ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY 0%7S A NOV 3 0 1927 "^S^AL MUSjA^ VOLUME IV DECEMBER 1923-1926 SEPTEMBER Numbers 1-5 Inclusive PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK 1927 New fork Zoological Swielg General Office: 101 Park Avenue , New York City (Pfltons President , Madison Grant; Vice-Presidents , Frank K. Sturgis; and Henry D. Whiton; Chairman , Executive Committee , Madison Grant; Treasurer , Cornelius R. Agnew. Secretary, William White Niles loarb of Managers Class of 1928 Percy R. Pyne, George Bird Grinnell, Anthony R. Kuser, Mortimer L. Schiff, Frederic C. Walcott, George C. Clark, Jr., W. Red- mond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler, George Gordon Battle, Bayard Dominick, Anson W. Hard Class of 1929 Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterjch George F. Baker, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, John E. Berwind, Irving K. Taylor, Harry Payne Bingham, Landon K. Thorne Class of 1930 Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, T. Coleman du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field ^rtrntifir i>taff W. Reid Blair, Director of the Zoological Park ; Charles H. Townsend, Director of the Aquarium ; Raymond L. Ditmars, Curator of Mammals and Reptiles ; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research ; Lee S. Crandall, Curator of Birds; ♦George S. Huntington, Prosector ; H. C. Raven, Associate Prosector; Charles V. Noback, Veterinarian ; Elwin R. Sanborn, Editor. iEbttnnal (Hmnmtttee Madison Grant, Chairman; Charles H. Townsend Elwin R. Sanborn, See’y. George Bird Grinnell ii W. Reid Blair William Beebe Zoologica, Vol. IV, Nos. 1-5 * TITLES OF PAPERS PAGE 1 — A List of Fishes, Amphibians and Reptiles Collected in Ashe County, North Carolina. . .Breder 3 2 — Problems and Facts About Frozen Siberian Mammoths ( Elephas primigenius) and Their Ivory. .................. .Lang 25 3 — The Galapagos Tortoises in Their Relation to the Whaling Industry Townsend 55 4 — Notes on Fishes from Three Panama Localities: Gatun Spill- way, Rio Tapia and Caledonia Bay. . Breder 137 5 — The Locomotion of Fishes Breder 159 in ' Zoologica, Vol. IV, Nos. 1-5. LIST OF ILLUSTRATIONS A LIST OF FISHES, AMPHIBIANS AND REPTILES COLLECTED IN ASHE COUNTY, NORTH CAROLINA Figures 1 to 8 inclusive PAGE Fig. 1. Mill Hill 5 Fig. 2. Beaver Creek in Front of the Post Office 5 Fig. 3. New River at the Mouth of Beaver Creek. 6 Fig. 4. Fyke Net Set in Beaver Creek . 6 Fig. 5. Semi-aquatic Habitat at the Base of French’s Knob .......... 7 Fig. 6. Terrestrial Habitat at the Base of French’s Knob 7 Fig. 7. Near the Summit of Bluff Mountain 8 Fig. 8. Beaver Creek 8 PROBLEMS AND FACTS ABOUT FROZEN SIBERIAN MAMMOTHS ( Elephas primigenius ) AND THEIR IVORY Figures 9 to 19 inclusive Fig. 9. Slope Where the Beresovka Mammoth Was Discovered. ...... 29 Fig. 10. The Beresovka Mammoth Ice-mummy in Process of Being Recovered 30 Fig. 11. Skull of the Adult Female Sangajurach Mammoth 32 Fig. 12. Portion of the Trunk of the Sangajurach Mammoth. ......... 32 Fig. 13. An Abnormally Spiraled Mammoth Tusk 34 Fig. 14. Outline Engraving of Wooly Mammoth Carved by Aurignacian Man 35 Fig. 15. A Piece of Frozen Mammoth Skin 36 Fig. 16. Various Stages in the Development of Upper and Lower Molars of the Mammoth 39 Fig. 17. The Mounted Mammoth Unearthed in 1901 as an Ice-mummy on the Banks of the Beresovka River 46 Fig. 18. The Beresovka Mammoth Skeleton as Mounted in the Petrograd Museum 48 Fig. 19. The Famous Frozen Mammoth of the Indigirka River, Siberia. 26 v V Illustrations THE GALAPAGOS TORTOISES IN THEIR RELATION TO THE WHALING INDUSTRY Figures 20 to 32 inclusive PAGE Fig. 20. Bark Morning Star of New Bedford. 56 Fig. 21. A Galapagos tortoise ( Testudo ephippium), on the U.S.S. Albatross 60 Fig. 22. Skeleton of Duncan Island tortoise ( Testudo ephippium ) as mounted in U. S. National Museum 62 Fig. 23. The South American tortoise ( Testudo tabulata ) 72 Fig. 24. A page from the logbook of the whaleship Isabella of New Bedford 74 Fig. 25. The Galapagos Islands. From U. S. Navy Chart No. 1798. . 76 Fig. 26. A Galapagos tortoise ( Testudo vicina ) from Albemarle Island. . 84 Fig. 27. A Galapagos tortoise ( Testudo vicina) drinking. From the New York Zoological Park 88 Fig. 28. Vegetation near Blackbeach, Charles Island, Galapagos 92 Fig. 29. A Galapagos tortoise ( Testudo abingdonii ) from Abingdon Island 94 Fig. 30. A Galapagos tortoise ( Testudo vicina ) from Albemarle Island. . 97 Fig. 31. A Galapagos tortoise ( Testudo vicina ) in a walking attitude. . . 100 Fig. 32. A Galapagos tortoise ( Testudo vicina ) in action 102 NOTES ON FISHES FROM THREE PANAMA LOCAL- ITIES: GATUN SPILLWAY, RIO TAPIA AND CALEDONIA BAY Figures 33 to 38 inclusive Fig. 33. Sketch map showing the location of the collecting sites at Gatun Spillway, Tapia, on the Rio Tapia, and Caledonia on Caledonia Bay 138 Fig. 34. The seaward end of a Caledonia Bay native fish trap 148 Fig. 35. A typical stretch of beach in Caledonia Bay 148 Fig. 36. A typical lane through a mangrove swamp at Caledonia 149 Fig. 37. Brackish pools just back of the beaches at Caledonia 149 Fig. 38. Doratonotus decoris 155 THE LOCOMOTION OF FISHES Figures 39 to 83 inclusive Fig. 39. Diagram of anguilliform locomotion 168 Fig. 40. The ostraciiform model. Perspective, side, and plan views. . . 170 Fig. 41. Diagram of the movements of the ostraciiform model. 172 Fig. 42. Diagram of a sculling oar, viewed from the rear 173 Fig. 43. Diagram of the path of particles of water about a rigid tail fin. 174 Fig. 44. Diagram of carangiform locomotion 175 Fig. 45. Caranx superimposed on Anguilla 176 Fig. 46. Diagram of the continual backward push of the caudal 178 Illustrations vii PAGE Fig. 47. Diagram of the movements of a carangiform fish. 181 Fig. 48. Undulation of the longitudinal structures 186 Fig. 49. The anguilliform model. Side and plan views 188 Fig. 50. Diagram of the pectoral movement as sometimes seen in Cich- lasoma 191 Fig. 51. Chilomycterus schoepfii ejecting water through the gill orifices. . 195 Fig. 52. Carcharias taurus swimming in shallow water 196 Fig. 53. Abramis chrysoleucas with pectorals removed 197 Fig. 54. Diagram of a fish turning by several methods 200 Fig. 55. Diagram of a tetradont such as Lagocephalus turning by several methods 201 Fig. 56. Pectoral control of elevation 203 Fig. 57. The effect of the relative positions of the fins on stopping 206 Fig. 58. Micropterus with the dorsal and anal lobes curved to one side and the caudal to the other so as to form a “sea anchor” 208 Fig. 59. Location of the center of gravity in fishes 212 Fig. 60. Diagram of the behavior of a Centrarchid ( Lepomis pallidus ) minus one pectoral 214 Fig. 61. Cross-section of Fundulus heteroclitus, indicating maximum roll of finless examples 215 Fig. 62. Maneuvering in a current 217 Fig. 63. Diagram of an epibatic tail as in Sphyrna 224 Fig. 64. Diagram of the action of a bilobed epibatic tail as in Mustelus . . 225 Fig. 65. Comparison of an isobatic and epibatic tail 226 Fig. 66. Comparison of action of model with a rigid and a flexible tail . . 229 Fig. 67. Behavior of model with tail fin insecurely attached to shaft. . . 230 Fig. 68. Progression of a leptocephalus. Conger sp 236 Fig. 69. Diagram illustrating the reason for Petromyzon marinus closing the mouth while swimming 240 Fig. 70. Diagram of the streamline contours of Rhinoptera bonasus . . . . 242 Fig. 71. The ultimate dorsal ray of Tarpon atlanticus 247 Fig. 72. The axillary scale of Elops saurus 247 Fig. 73. Pectorals modified for flight 249 Fig. 74. The trajectory of characins 250 Fig. 75. The motion of Atroblepus longifilis 251 Fig. 76. Hemiramphus leaping and skittering. 258 Fig. 77. Lophosetta maculata with dorsal fin rays in a resting position . . . 267 Fig. 78. Front view of Echeneis naucrates 268 Fig. 79. Pteraspis rostrata 274 Fig. 80. The ostraciiform model. Working drawing showing details of construction 291 Fig. 81. The anguilliform model. Working drawing, showing details of construction 292 Fig. 82. Dynamometer for determining the traction pull of^fishes 293 Fig. 83. Intermittent light device for determining the speeds^of un- dulating fins 295 r ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME IV. NUMBER 1 Papers from the New York Aquarium, Contribution Number 10 A LIST OF FISHES, AMPHIBIANS AND REPTILES COLLECTED IN ASHE COUNTY, NORTH CAROLINA By C. M. Bredee, Jr. The New York Aquarium , and Ruth B. Breder PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 27, 1923 Nm fork Xonlngiral ^nrirtg General Office: 111 Broadway, New York City GDffera President , Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee , Madison Grant; Treasurer, Cornelius R. Agnew Ifoarb of managers (Elans of 1924 Madison Grant, William White Niles, Frank K. Sturgis, Ogden Mills, Lewis Rutherfurd Morris, Archer M. Huntington, George D. Pratt, T. Coleman duPont, Henry D. Whiton, Edward Hatch, Jr., Cornelius R. Agnew, Harrison Williams (Ela00 of 1925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge C. Ledyard Blair, Anthony R. Kuser, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler Ollaoa nf 192fi Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor. i>taff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, D. V. S., Assistant to Director ; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of the Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector ; George A. MacCallum, Pathologist; El win R. Sanborn, Photographer and Editor. CMtorial (Eommttto Henry Fairfield Osborn, Chairman; William T. Hornaday, Charles H. Townsend. Corrected to October, 1923 ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME IV. NUMBER 1 Papers from the New York Aquarium, Contribution Number 10 A LIST OF FISHES, AMPHIBIANS AND REPTILES COLLECTED IN ASHE COUNTY, NORTH CAROLINA By C. M. Breder, Jr. The New York Aquarium, and Ruth B. Breder PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 27, 1923 First form on press December 27, 1923. (2) Zoologica Vol. IV, No. 1. A LIST OF FISHES, AMPHIBIANS AND REPTILES COLLECTED IN ASHE COUNTY NORTH CAROLINA By C. M. Breder, Jr. New York Aquarium, and Ruth B. Breder Introduction The County of Ashe, North Carolina, is little known from the biological point of view and the present paper stands as the first report on the ichthyological and herpetological fauna of it. Mr. C. S. Brimley of Raleigh, N. C., states that the only other reported collection previously made within its boundaries is a small one of insects. The area in which the specimens were collected on which this list is based is located in the central part of Ashe County and the time spent collecting in this territory extended from July 21 to August 17, 1922. This county is the most northwesterly one of the state, being bounded on the north by Virginia and on the west by Tennessee. The settlement at which headquarters were made is known to the postal authorities as Beaver Creek. It is located about fifteen miles southeast of the junction of the three states and is a typical hamlet of the southern Appalachians. All the collecting was carried on within a radius of five miles of the Beaver Creek post office, whilst by far the largest amount of it was done within less than two miles, and so represents a bit of concentrated collecting within a circumscribed area of small dimension. The land at this point is comparatively fertile and a considerable amount of it has been cleared, some quite recently, limiting the collecting grounds for terrestrial salamanders to rather well defined elevations which still in many cases are covered with the original stands of timber. Lumbering, however, is making rapid inroads on this type of territory. (3) 4 Zoologica: N. Y. Zoological Society [IV; I In the summer of 1915, the senior author spent two months at this same place engaged in other studies and it was then that the salamander fauna forced itself on the attention. Although no definite records were kept it is certain that there has been a con- siderable reduction in the abundance of these amphibians for which there seems to be no very apparent reason. It hardly seems possible that the lumbering above alluded to could have annihilated the animals in such great numbers for only in occasional places has the forest floor been disturbed to any great extent. Moreover, there seems to have been a considerable change in the fauna in general. Some species now quite or nearly absent were formerly abundant and vice versa. For example, only a single specimen of the common tumble bug, Canthon, was seen this year while previously they were ubiquitous. On the other hand, myriapods, arachnids, birds, and mammals were about as before, whilst terrestrial gastropods have noticeably increased in number. A partial explanation may be found in the fact that this summer was somewhat cooler than the one of 1915, and while numerous thunder squalls, which quickly dried, made a wet summer as far as agriculture was concerned, it lacked the drizzles which marked that of seven years previously which kept the soil continually damp. This territory exposes rocks of pre-cambrian formation and near springs at such outcroppings much of the Urodel collecting was carried on. The altitude varies from about 3,000 to somewhat over 5,000 feet above sea lq)vel. The fishes were collected by seine, fyke, dip net and angling gear. A small minnow seine of twenty feet was used in all waters in which its successful operation was possible, while the fyke (Fig. 4), which was likewise of small size and mesh, was set continually in the Beaver Creek, near headquarters. Small ditches, holes and backwaters were worked with a dip net. Angling was, as usual, least productive. The Beaver Creek, from which this place takes its name, runs within a few hundred feet of the post office and at this point an old time grist mill still flanks it. The section in which collecting was done wanders through tilled fields interspersed at various places by small patches of woodland. The Buffalo Creek, which drains the next valley to the northwest, a short distance on horse, empties into the north fork of the New River whilst the Beaver Creek empties into the south. These forks join at a distance of about twelve miles from the collecting site. The creeks are quite similar in general appearance, and while no great difference in the FIG. 2. BEAVER CREEK IN FRONT OF THE POST OFFICE Nigger mountain in the background. Baptist church in middle ground. 5 FIG. 1. M ILL HILL The unforested ground above the cottage was formerly inhabited by great numbers of Plethodon glulinosus. Zoologica Vol. IV, No. 1 FIG. 4. FYKE NET SET IN BEAVER CREEK Type of banks found in the open fields. 6 FIG. 3. NEW RIVER AT THE MOUTH OF BEAVER CREEK The water has little depth here. ZoQlogica Vol, IV, No. 1 FIG. 5. SEMI-AQUATIC HABITAT AT THE BASE OF FRENCH’S KNOB A typical haunt of Desmognathus. FIG. (>. TERRESTRIAL HABITAT AT THE BASE OF FRENCH’S KNOB A typical haunt of Plethodon. 7 Zoologica VoJ. IV, No. 1. 8 FIG. 7. NEAR THE SUMMIT OF BLUFF MOUNTAIN FIG. 8. BEAVER CREEK The type of rugged country encountered at the highest altitudes. View a short distance below the Post Office. French’s Knob in the Zoologica Vol. IV, No. 1. le^ background. 1923] Breder: Fishes — Amphibians — Reptiles 9 respective faunas could be noticed still there was a perceptible dif- ference in the relative frequencies of the various species as is in- dicated in Table No. 1. All amphibians and reptiles were collected by hand. Belts made similar to those used for cartridges, but instead fitted with pockets for holding tobacco tins, were found extremely useful and convenient as carriers, especially for living material of small size. Both the herpetological and ichthyological specimens were preserved in formalin in the field, and were transferred to alcohol on arrival at the laboratory. The collection has been deposited in the Amer- ican Museum of Natural History with the exception of some speci- mens that have been retained for reference. The locations of col- lecting sites mentioned throughout are defined under “ Distribution of fishes” or “ Distribution of salamanders” on pages 13 and 19 respectively. Dr. E. R. Dunn, of Smith College, gave very material aid in checking our identifications and in determining difficult specimens among the amphibia. Dr. G. K. Noble and Mr. J. T. Nichols permitted our use of the herpetological and ichthyological material of the American Museum, of which they have charge respectively. Mr. A. 1. Ortenberger, of the same institution, aided in the deter- mination of the snakes. To these gentlemen we are accordingly indebted and wish to here express our appreciation. Likewise we are grateful to Mr. and Mrs. R. A. Hamilton at whose home we were guests and who facilitated the collecting by their many kindnesses. The localities, sizes, et cetera, are given in some detail, even at the risk of being accused of undue verbosity, but on account of the violent changes which have taken place in the fauna of this region in the past seven years, as before noted, it is deemed advisable to have an accurate record of just what has been collected, for purposes of future comparison. Further, it is the writers’ belief that an error in this direction is preferable to one in the opposite. Such are only too often shown by regional lists of this nature. The metric system is used throughout and the measurements given for the fishes all refer to the standard lengths ; that is, the shortest distance from the tip of the snout to the base of the tail. The common name or names given for each species are those by which the natives know the animals. In several cases these are rather unique. In instances where more than one is given the first is the appellation most com- monly used. 10 Zoologica: N. Y. Zoological Society [IV; I Class PISCES Family CATOSTOMIDAE 1 — Catostomus nigricans LeSueur. Hogfish. Ten examples of from 51 to 240 mm. were taken between July 24 and August 3, all from the Beaver Creek from a short distance above the mill to a point about one mile further down stream. Family CYPRINIDAE 2 — Cyprinus carpio Linnaeus. Carp. Several of the more active natives have constructed carp ponds and have had sufficient success with them to supply their occasional demands for carp flesh. These ponds are of the simplest nature and are of small size. On account of the torrential rains common to this country the mud and silt brought down soon fills the ponds making it imperative to clean them out or build new ones. The two ponds visited were rather decadent, one appearing to be empty of fish and nearly filled with silt, whilst the other contained a fair crop of fry. Doubtless, many fishes escape from these ponds as very few safeguards have been taken to prevent losses in this manner. Such that do escape probably pass down to the relatively quiet waters of the New River in preference to the rapid mountain streams. Just why these people have taken to cultivating carp is not clear since their ponds are all fed by excellent trout streams. 3 — Campostoma anomalum (Rafinesque). Minnow. Of six examples ranging from 53 to 75 mm. only one was taken from the Beaver Creek, about a mile below the post office on July 28, while the remaining five were taken from the Buffalo Creek on August 12. Another of 57.5 mm. was taken from the Beaver Creek near the post office in the summer of 1915. 4 — Semotilus atromaculatus (Mitchill). Chub, Hornyhead. Eighty-one examples of from 35 to 154 mm. were taken. Sixteen of the smaller sized fishes were taken from the Buffalo 1923] Breder: Fishes — Amphibians — Reptiles 11 Creek on August 12. The remainder were taken from Beaver Creek, from above the mill to a point about one mile down stream, between July 24 and August 3. Only Rhinichthys exceeds this species in abundance in these waters. Many of the natives think the smaller ones are immature trout. 5 — Notropis photogenis photogenis (Cope). Minnow. Five examples of from 63 to 73 mm. were taken from the Beaver Creek within sight of the mill from uly 24 to August 3. This record stands as an addition to the fish fauna of North Carolina. The almost indistinguishable race, amoenus, well known from the coastal plain drainage, is replaced by photogenis in this territory which is Mississippian, as should be expected. In comparing these fishes with specimens in the American Museum of Natural History, there was found a somewhat larger individual, identical with the present material, which had been collected by Morton L. Church, at Marshall, N. C., that had not been recorded. Dr. Henry W. Fowler, of the Philadelphia Academy of Natural Sciences, kindly examined these specimens and considers them unquestionably photogenis. 6 — Rhinichthys atronasus (Mitchill). Minnow. One hundred and five examples of from 20 to 68 mm. were taken. Eighty-six were taken from the Beaver Creek, from above the mill to about a mile below it, from July 24 to August 3, whilst the re- mainder were taken from the Buffalo Creek on August 12. This species was particularly abundant, the smaller ones finding their way into the most tiny trickles whereas the larger examples were usually schooled up in shaded holes in the larger branches and creeks. Seen in the New River on August 13. 7 — Hybopsis kentuckyensis (Rafinesque). Hornyhead, Minnow. Three examples of from 109 to 110 mm. were taken from the New River at the mouth of the Beaver Creek on August 13 by hook and line. Family SALMONIDAE 8 — Salmo irideus Gibbons. Rainbow Trout. Reported by natives, who stated that a plant was made by the United States Bureau of Fisheries some years ago and that oc- casional examples have been taken since that time of under a foot in length. None seen by us. 12 Zoologica: N. Y. Zoological Society [IV ; I 9 — Salvelinus fontinalis (Mitchill) . Speckled or Brook Trout. Two immature examples of 119 and 163 mm. were taken on July 24 from the Beaver Creek above the mill. The stomach of the smaller example contained a crayfish about 12 mm. long and the larger one was full of insect fragments. Some years ago plantings of this species were made by the Bureau of Fisheries and these may be of that stock as lumber camp polutions practically wiped out the native trout some years back. Since then protective legislation has checked the contamination from them and the streams appear to be well recovered. Family CENTRARCHIDAE 10 — Ambloplites rupestris (Rafinesque). Redeye, Rock Bass. One example of 141 mm. was taken from the New River by angling, slightly above the mouth of the Beaver Creek (Figure 3), on August 13. Others of smaller size were seen at this time. Family PERCIDAE 11 — Etheostoma flabellare (Rafinesque). Young Hogfish. Ten examples of from 18 to 36 mm. were taken from Beaver Creek between a point slightly above the mill and one about a mile below it. The natives consider this the young of Catostomus. Family COTTIDAE 12 — Cottus ictalops (Rafinesque). Poisonhead. In the summer of 1915 this was one of the most abundant forms of fish life at this place, a simple seine made of a gunny sack often bringing up a dozen or more from a short haul. One specimen preserved at the time measured 61 mm. and was about the average size. A diligent search of all the waters in 1922 failed to reveal a single individual, and the crop of small boys that have sprung up in the intervening seven years do not know the name 'Poisonhead.’ The small boys of 1915, no longer such, aver that they do not re- member having seen any for a long time. The various small cypri- nids on the other hand seem to be more abundant. Distribution of Fishes at Beaver Creek Table No. 1 lists the species collected according to the three chief sites with reference to their relative abundance. The locality called Beaver Creek includes a stretch of that stream of about two 1923] Breder: Fishes — Amphibians — Reptiles 13 miles in extent, reaching from a point about three-quarters of a mile above the post office to one about one and one- quarter miles below it. It is here that most of the collecting was carried on and includes all the confluents of this stream between these two points. The other two localities were visited but once each. The Buffalo Creek was collected in, over a stretch of about one half mile, at the base of the northwest side of the Paddy Mountain. The south fork of the New River was angled in only, close to the mouth of the Beaver Creek. TABLE NO. 1 Distribution of Fishes Species 1 Catostomus nigricans 2 Cyprinus carpio 3 Campostoma anomalum 4 Semotilus atromaculatus 5 Rhinichthys atronasus 6 Notropis p. photogenis 7 Hybopsis kentukyensis 8 Salmo irideus 9 Salvelinus fontinalis 10 Ambloplites rupestris 11 Etheostoam flabellare 12 Cottus ictalops eaver Creek Buffalo Creek New River Common Reported Reported (large) Known from fish ponds only Occasional Common — Abundant Abundant — Abundant Abundant Common Occasional ■ — — — — Common Reported — — Not scarce Reported Reported — Common Rather common — — Absent, formerly abundant — — Class AMPHIBIA Order CAUDATA Family CRYPTOBRANCHIDAE 1 — Cryptobranchus allegheniensis (Daudin). Water Dog. Reported by natives from both the New River and Beaver Creek. The bed of the creek was recently shifted near West Jefferson for some road building operations and as a result several were noticed by the engineers who gave a fair description. No examples were seen by us. Family SALAMANDRIDAE 2 — Triturus viridescens viridescens Rafinesque. Water Lizard. Described by natives from a carp pond near Jefferson. On search we failed to find any, but very likely they occur here sparingly as the natives’ descriptions were reasonable. 14 [IV; I Zoologica: N. Y. Zoological Society Family PLETHODONTIDAE 3 — Plethodon cinereus (Green). Wood Puppy. Common in damp woods of not over 4,000 feet elevation. Taken on Mill Hill, near the Baptist church, and at the base of French’s Knob under damp litter, between July 22 and August 3. A curious feature is that out of the thirty-two individuals which varied from 34 to 76 mm. and numbers not collected, only a few with red backs were seen. This is especially interesting since Dunn1 in taking forty- eight specimens at Linville (thirty-three miles southwest of here) writes “only two had black backs.” Apparently this species holds the lower grounds as none were taken on the higher mountains. On the sides of Bluff Mountain it appears to be replaced by P. metcalfi. At least it is strange that the only place from which we took that species the present was absent. The point of greatest abundance for cinereus in our territory was the small patch of damp woods behind the Baptist church. Here large numbers of these supple little gray salamanders could be taken with slight effort. 4 — Plethodon glutinosus (Green). Wood Puppy. Abundant. Taken in a large variety of both damp and dry localities. Numbers were seen which were not collected, a few even in parched fields of stubble under stones or bits of wood of small size. Sixteen were collected ranging in size from 44 to 157 mm. between July 23 to August 2 from the following places: Mill Hill, damp woods; near Baptist church, damp woods; Base of French’s Knob, damp woods; Bluff Mountain, damp woods; Buck Mountain, near Mica Mine; Southwest slope of Nigger Mountain, both damp and dry localities. 5 — Plethodon metcalfi Brimley. Wood Puppy. Taken only on Bluff Mountain in damp woods as follows: 93, 98, 111 and 113 mm. The first varied from the typical metcalfi in that it possessed a few pigmentless areas on the sides which gave it a resemblance to P. glutinosus. Thus far all specimens of metcalfi have agreed in the uniformity of the gray coloration on the back and sides. This individual demonstrates that metcalfi may approach glutinosus in coloration by occasionally possessing these white spots 1 Dunn, Emmet R. Reptile and Amphibian Collections from the North Carolina Mountains with especial Reference to Salamanders. Bull. Amer. Muse. Nat. Hist. Vol. XXXVII, Art. XXIII, Oct. 13, 1917, N. Y. 1923] Breder: Fishes — Amphibians — Reptiles 15 even as the converse is true in that glutinosus sometimes lacks them. The presence or absence of these spots can therefore be used no longer by itself as a simple means of separating the two. 6 — Plethodon yonahlossee Dunn. Wood Puppy. One adult of 131 mm. was taken from under a rotten log on the damp southwestern slope of Mill Hill, July 26. Two young of 48 and 47 mm. on Nigger Mountain and two young of 64 and 56 mm. on Bluff Mountain. The young all showed the characteristic spots. These altitudes are slightly less than that of the type locality, they being from 3500 to 4000 feet whereas the latter was be- tween 4100 and 4400. This is the most northerly record for the species. 7 — Eurycea bislineata wilder ae Dunn. — This recently described race2 was taken in a variety of places, usually damp, although it was found further from water and in drier places, at times, than the writers had ever seen bislineata in the latitude of New York City or Washington, D. C. Five adults were collected from July 25 to August 8 varying from 28 to 71 mm. at these places: Mill Hill, both damp and relatively dry woods; near Baptist church, in spring; Bluff Mountain, in spring. The specimen of 28 mm. which was taken in a spring near the summit of Bluff Mountain presented an unusual type of coloration. The usual yellowish back, bordered with a black line on either side, was re- placed with a ground color of dusky brown. This color extended downwards to the insertion of the limbs, and only a faint suggestion of the black lines of typical individuals was present. About six lighter brown spots followed both these suggestions of lines at their dorsal edge between the fore and hind limbs. They showed a slight tendency towards occelation, as did other less conspicuous markings on the caudal region. These markings were not widely different from those of some of the larvae which seem to have a considerable range of individual variation. However, this specimen showed no larval fin fold or other immature characteristics and appeared to be fully metamorphosed, although it was smaller than some larvae we took, our largest being 32 mm. as against the 28 mm. of this example. Taxonomically, if otherwise checks well for this 2 Dunn, Emmet R. Some Reptiles and Amphibians from Virginia, North Carolina, Tennessee and Alabama. Proc. Biol. Soc. Wash., Vol. 33, pp. 129— 138, Dec. 30, 1920. 16 Zoologica: N. Y. Zoological Society [IV; I species and we believe it can be put down simply as an abnormality. The larval material was taken from the following places: Spring near summit of Bluff Mountain. One 32 mm. in company with the above described specimen and one adult of 30 mm., August 8. Spring near look off rock on Nigger Mountain. Temperature of water 54° F. Ten, 17 to 27 mm., August 9. McKeever’s spring, northeast base of Nigger Mountain. Temperature of water 48° F. Seven, 27 to 33 mm., August 9. 8 — Pseudotriton ruber niditus Dunn. Red Lizard. Four examples were seen, two of which subsequently escaped. The others represented the two most extreme types of variation of this race as yet seen by the describer. Both of these were in ex- ceptionally dry places and only about a mile apart. One near the Baptist church, on a hillock taken July 28, measuring 96 mm., was very lightly speckled with small black punctulations on the back, hardly any of which extended beyond the insertion of the hind limbs. The appearance in life was a bril- liant waxy red, the subspecific designation being especially appro- priate for this individual. The other on Mill Hill, at the edge of dry field, taken on July 31, measuring 85 mm., was covered with black spots of a heavy sort extending nearly to the tip of tail, although there was no tendency for the larger ones to fuse as in ruber. Neither was there any smaller stippling interspersed between the larger spots. Larvae were taken as follows: Mill Hill, July 25, 42 and 37 mm. ; Bluff Mountain, August 8, 29 mm. 9 — Desmognathus fuscus fuscus (Rafinesque). Wood Puppy. This widely distributed form was not hard to find but by com- parison with D. monticola was not abundant. Fifteen specimens were taken between July 25 and August 9 which varied from 46 to 75 mm. from the following places: Mill Hill, at the water’s edge and in springs; near Baptist church, in a nearly stagnant trickle; base of French’s Knob, in springs; Nigger Mountain, in spring near look off rock. In the two highest places in which taken (Mill Hill and Nigger Mountain) this species accompanied D. monticola whilst the latter was absent from its two lowest environments (near the Baptist church and at the base of French’s Knob) although the 1923] Breder: Fishes — Amphibians — Reptiles 17 forest floor in all four cases was generally similar. Not seen in places as dry as those in which monticola was at times. 10 — Desmognathus monticola Dunn. Wood Puppy. Abundant in damp and wet places up to 4,500 feet. Eighty-six were taken which varied from 23 to 125 mm. between July 25 and August 14 from the following places: Mill Hill, stream edge and damp places; Nigger Mountain, springs and damp woods; Bluff Mountain, damp woods; base of Paddy Mountain, spring; base of Mulatto Mountain, spring. While usually associated with damp- ness, they were frequently found considerable distances from water as compared with fuscus from here or other places. 11 — Desmognathus ochrophaeus carolinensis Dunn. Wood Puppy. This more terrestrial species of Desmognathus was taken in a wide variety of places. Twenty-nine individuals were taken between July 25 and August 9 which ranged from 19 to 90 mm. from the following localities: Mill Hill, dry woods; Nigger Mountain, dry woods, springs, and in a cave-like grotto; foot of French’s Knob, damp woods; Bluff Mountain, damp woods; Buck Mountain, near Mica Mine; Hayfield, near edge of spring. 12 — Desmognathus quadra-maculata (Holbrook). Taken in and at the edge of small streams, usually less than one half mile from their origin. Dunn states, “Around Brevard they come down to 2100 feet in large streams. ”3 As none of our territory was of much less elevation than 3000 feet we have no data on their environment below that level, but none that we saw were in streams over three feet wide, although we should have taken them, if present in the larger creeks on account of the ichthyological collecting. The locality of their greatest abundance was at a height of 4300 feet near the summit of Bluff Mountain. Here in a stream rising from a spring about a mile back from the bluff this species was particularly common. They were taken as near to the precipice as we dared approach with collecting in mind, at which point a picturesque streamlet with an average depth of about two inches and a width of about three feet plunged nearly vertically to the valley over eight hundred feet below. One of the individuals taken there was marbled all over with lighter, but is undoubtedly referable to this species nevertheless. One larval example of 55 mm. was taken in a 3See footnote 1, page 14. 18 Zoologica: N. Y. Zoological Society [IV; I spring on Buck Mountain near the Mica Mine. Another larva of 52 mm. was taken in the stream at the Bluff. Adults were taken in springs and streams on Mill Hill, Nigger Mountain and Bluff Moun- tain, as before noted, to the number of nine between July 25 and August 8 which varied from 55 to 110 mm. Distribution of Salamanders at Beaver Creek In Table No. 2 the species collected are arranged according to the chief localities. Those localities mentioned in this table that can not be found on the Cranberry or Wilksboro sheets of the United States Geological Survey map are explained below. Mill Hill — A wooded hillock of 3400 feet on one’s right if facing down stream at the Beaver Creek post office. See Figure 1. Baptist Church — A small glen behind a church of that de- nomination about a quarter mile down stream from Beaver Creek. See Figure 2. Rail Road Hill — A hill of 3200 feet, across the rail road from the post office, at a point about one quarter mile below the station. French's Knob — A densely tangled rise of land of 3600 feet, about one mile southeast of Beaver Creek. The northwest base was collected on very successfully. See Figures 5, 6 and 8. Mica Mine — A mine for high grade mica, now not in use, on the Buck Mountain is here referred to. It was in its near vicinity that the specimens were taken. Carp Pond — A disused carp pond near Jefferson on the property of Mrs. C. Neal. Order SALIENTIA Family BUFONIDAE 1 — Bufo americanus Holbrook. Toad Frog. Abundant. Many were seen, including practically all stages except the eggs. Eight adults measured as follows in head and body lengths, 105, 97, 86, 83, 73, 71, 65, 62 mm. The two smallest ones were decidedly reddish but lacked the dark throat patch character- istic of the male. One immature example of 8 mm. was nearly as red as the terrestrial form of Triturus viridescens. The natives consider these another species, calling them Bed frogs and tell wonderful tales of how they descend in the rain. The great numbers in which small toads of this size appear in these mountains after a Distribution of Salamanders 1923] Breder: Fishes — Amphibians — Reptiles Carp Pond Reportei 1 1 ' 1 1 1 I 1 1 1 Paddy Mount. I 1 1 1 1 1 1 ft i ft 'C =2 1 ft £5 ! | © 6 § 1 II 2 1 | 1 1 1 1 3 © ft NS 1 1 1 o 1 1 1 1 1 a Ul © JS £ - w S3 © © ift 32 2 2 •1 1 8 i 1 1 3 1 Ch ft 1 © <3 Sh 1 Sh rf £ W 1 1 |! 1 | 'd as 1 43 o 1 O ® o fc 5? S o 05 3 i nil 1 Mil 1 , | 1 , , =3 W ffS 05 ■£ 1 0 S3 i , § 2 , NS , 43 © Sh aj , a U cS 33 w o g a 1 |<3 73 ft 'd c3 4^ o 1 fc o_ 43 ? i © © 1 1 1 S S bO © C3 43 3 © ft s> © © , o o a a a 33 si 43 43 1 1 1 O O O >c O T3 ft © ft a a 1 o o 6 £ £ O O «1 © *-< 1 1 1 Si 4^ © © ft © © 43 © c be 5 b0 © ft © 2 ft S 1 fl 5 =3 3 2 ft 3 S3 bC q fc £ 43 1 1 1 o 1 6 O >» (M t a ft O J o 1 a § a wo O o 43 o fc ft a © oft © S3 . , o © , © o © O 3 1=5 1 § § 3 3 > a a g g £ W 1 § a 1 1 o o 1 33 ft ft -© ft © CO 4^> S 4^ O o o a o O O £ o £ O ^ © © > *3 © M I ! co - J p © .Si £ g co V. I »» S o : S co £ co co 2 cc? O © §; e O S rH -~ 'e © £ CC Oh' ^ ^ M ^ lO © N g -£ « I ? 3 i ' 2 | ;S S 9 © S *» * ? 5 . §, r* ■« ^ °' ■ © Q s* 'e g Is c5 o< e co Os 00 Ci 19 20 Zoologica : N. Y. Zoological Society [IV; I cloudburst is nevertheless extremely impressive. Tadpoles in various stages were seen as late as August 16. Family RAN I DAE 2 —Rana catesbeiana Shaw. Bull Frog. A single specimen of 91 mm. was taken in the minnow fyke in the Beaver Creek, near the post office. A number were heard along the New River on August 13 at mid-day in fair hot weather. 3 — Rana clamitans Latreille. Frog. Two tadpoles of this species 70 and 53 mm. long were seined in the Beaver Creek about one mile below the mill on July 28. No adults were seen. 4 — Rana palustris LeConte. Frog. Several examples were seen in the meadows about Beaver Creek. 5 — Rana sylvatica LeConte. Frog. One example of 30 mm. was taken on July 31 on the east side of Mill Hill at the edge of a dry woods. This individual departed from the typical sylvatica in that its back was markedly rugose, but in other respects was perfectly normal. Class REPTILIA Order SQUAMATA Suborder Sauria Family IGUANIDAE 1 — Sceloporus undulatus (Latreille). Scorpion, Fence Lizard (rare). This species which the natives hold in unnecessary awe was seen quite frequently, being perched as usual on old fence rails in the hot sun. Only two specimens were taken as there seemed to be no very good excuse for destroying numbers of these harmless and interesting lizards. One was a typical male of 135 mm. and the other a female of 134 mm. Both had their stomachs crammed full of fragmentary insects, most of which seemed to be coleopterous. July 27 and 28 are the respective dates of capture. On August 15 near the crest of Mill Hill in a clearing an individual was noted which was blotched with whitish. Partial albinism was thought of, 1923] Breder: Fishes — Amphibians — Reptiles 21 but when the animal allowed itself to be picked up and handled without attempting to escape it seemed likely that some disease afflicted the creature. It was allowed to perch freely on the finger, and strangely enough permitted a stroking of the white spots, responding only by craning its neck and blinking its beady eyes. This gentle stroking caused the scales to drop off in a shower of dandruff-like flakes and it was then obvious that the animal was merely shedding, as beautifully brilliant scales appeared wherever the old ones fell from. After a few minutes of this sort of treatment it decided it had stood for enough and with one powerful leap left the scene. Subsequent examination of the shed scales showed that they had completely sheathed the new formation simply tearing away all around the base. The whitish appearance is accounted for by the fact that air found its way between the loose old scales and the new. This contrasts strongly with such fine scaled forms as Anolis in which the epidermis is cast off in large patches, some- times the entire covering coming away almost as one piece. Suborder Serpentes Family COLUBRIDAE 1 — Carphophis amoena (Say). Snake. Two examples of 23 and 28 cm. were taken between July 27 and 31 and one of 24 cm. in 1915. A few others were seen, all in typical localities under stones and rubbish. The natives hardly know this unobtrusive form. 2 — Diadophis punctatus edwardsii (Merrem). Snake. Three examples from 29 to 35 cm. were taken in places similar to that of the above and usually close by between July 27 and 30. Numerous others were seen. 3 — Elaphe obsoleta obsoleta (Say). Black Snake. One example of 168 cm. was taken from the north side of the Mulatto Mountain on July 31. It is said to be fairly common by the natives, who hold it to be poisonous. 4 — Lampropeltis triangulum triangulum (Lacepede). Snake* One example of 49 cm. was taken at Beaver Creek on July 22* Not recognized as a distinct species by the inhabitants of that place. 22 Zoological N. Y. Zoological Society [IV; I 5 — Natrix septemvittata (Say). Water Snake, Water Moccasin. Three examples all in the vicinity of water, varying from 28 to 61 cm. in length, were taken between July 22 and August 11. The natives, are not sure as to whether these are dangerous or not. 6 — Natrix sipedon fasciata (Linnaeus). Water Snake, Water Moc- casin. Two examples, 71 and 79 cm., were taken near water on August 3 and 11. Many others were seen, it being a very common form in this vicinity. The natives confound this with the preceding species. 7 — Thamnophis sauritus (Linnaeus). Snake. One example from near the base of Buck Mountain was taken on August 11. It measured 54 cm. in length. 8 — Thamnophis sirtalis sirtalis (Linnaeus). Snake. Three examples from 56 to 105 cm. in length were taken between July 22 and August 13. The largest one was about to give birth to thirty-seven young which varied from 14.0 to 17.2 cm. and showed a mode of 16.0. A common species. Family CROTALIDAE 9 — Crotalus horridus Linnaeus. Rattlesnake, Rattler. The natives hold this snake in great fear and report that it is common and especially abundant at certain places. A careful though unsuccessful search was made both in 1915 and 1922 in the vicinity of reputed “dens.” Once did the junior writer believe a specimen was seen but it slipped away amid a pile of debris too rapidly for positive identification. Not one was definitely seen by us and it may be significant that not an individual questioned had a rattle to show as evidence. As boys living in areas infested with these reptiles usually have a few sets of such mementoes we infer that the species is not common in this vicinity. Order TESTUD1NATA Family CHELYDRIDAE 1 — Chelydra serpentina (Linnaeus). Terrapin. One specimen was seen with a carapace of 102 mm. It was typical and was taken in a small swamp near the north side of the Mulatto Mountain. This species is said not to be uncommon by the 1923] Breder: Fishes — Amphibians — Reptiles 23 natives, who relish it as food. Chelydra is known in this region simply as Terrapin while all others are known as Dry-land Terrapin whether they are aquatic in habit or not. The shorter and less cumbersome names of Turtle and Tortoise are unknown terms to most of these mountaineers. Family TESTUDINIDAE 2 — Clemmys muhlenbergi (Schoepf). Dry-land Terrapin. Two specimens were taken, one of which subsequently escaped. The measurements of the other were as follows. Carapace length 91.5 mm., width 69 mm., plastron length 80 mm., width 56 mm., length of tail (vent to tip) 17 mm., head width 17 mm. Carapace brownish black, no blotching whatsoever. Plastron with light brown and yellowish blotching, bridge similar. Nuchal plate 6 mm. Taken near the Baptist church on July 28 in a discarded horse trough. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME IV. NUMBER 2 i FEB^ni^ PROBLEMS AND FACTS ABOUT FROZEN SIBERIAN MAMMOTHS (. ELEPHAS PRIMIGENIUS) AND THEIR IVORY By Herbert Lang Associate Curator, African Mammals, American Museum of Natural History, Fellow, New York Zoological Society PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK January 8, 1925 Nm fork 2oologiral 8>ori?tg General Office: 101 Park Avenue, New York City (J&flfera Honorary President, Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer , Cornelius R. Agnew laard of iHattagrra Ollaaa of 1925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, C. Ledyard Bla r, Anthony R. Kuser, Watson B. Dickerman, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr. Ollaaa of 192fi Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich. George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor GHaaa of 192 7 Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field iririttiftr ^taff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant to the Director and Veterinarian; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector; Elwin R. Sanborn, Photographer and Editor. Editorial (Eammittor Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y. Zoologica Vol. IV, No. 2. PROBLEMS AND FACTS ABOUT FROZEN SIBERIAN MAMMOTHS (ELEPHAS PRIMIGENIUS ) AND THEIR IVORY1 By Herbert Lang Associate Curator, African Mammals, American Museum of Natural History, Fellow, New York Zoological Society (Figs 9 to 19 Incl.) Centuries ago rumors of the discovery in northeastern Asia of great curved tusks of ivory persistently drifted into Western Europe. Could it be true that animals apparently surpassing an elephant in size lived on the bleak tundra of northeastern Siberia? Did it not sound like a fairy story that such gigantic beasts bur- rowed and lived underground somewhat like our tiny moles? Ides, the famous Dutch traveler and ambassador to China, seems to have been the earliest to gather first-hand information. On traversing northern Siberia between the years 1692 and 1695 he learned that many of the Yakuts, Tunguses, and Ostyaks stead- fastly believed that these huge monsters spent their lives deep underground, moving about easily in spacious tunnels even though the earth was thoroughly frozen. Should they become particularly active the whole ground might rise above them, caving in later as they passed on. But let the “mamonts” or “ ground-dwellers0 come to the surface and breathe the warm air, they instantly died.2 This is not so strange a story when we consider that actual cir- cumstances helped strengthen native belief. At certain places in Siberia, after the melting of the snow, plenty of bones of what later came to be called the mammoth were lying about the surface or sticking up from the ground. Here and there after the thawing and slipping of portions of steep river banks the more or less complete remains of these proboscideans had been exposed to view in the very sites where inadvertently they might have reached the fateful daylight. At other times one of these frozen giants was discovered at a point the natives imagined to be the end of the mammoth’s diggings. 1 The photographs and some of the data in this article have been kindly contributed by Dr. E. W. Pflzenmayer, Curator, Natural History Museum, Stuttgart. Formerly: Assistant, Petrograd Zoological Museum; Member of the Beresovka Mammoth Expedition; Leader of the Sangajurach Mammoth Expedition. 2 Ides, Isbrand, 1704, ‘Dreyjarige Reize naar China.’ Amsterdam, p. 31. 25 First form on press January 5, 1925. Fig. 19. — The famous frozen mammoth of the Indigirka River, Siberia. From Beukendorf’s romantic story, Mrs. E. Rungius Fulda pictures the ice-mummy with hind limbs still anchored in the unthawed banks. Later, when the ground, together with the mammoth, was carried away by the force of the onrushing floods, the discoverers had a narrow escape. Reproduction from W. T. Hornaday’s “Tales From Nature’s Wonderland. ” Copyrighted by Charles Scribner’s Sons. 1924] Lang: Frozen Siberian Mammoths 27 It is with amusement we turn to the many heated, rather sprightly controversies that centered about the early finds in Europe of certain fossil bones of enormous proportions. Did they belong to giants? Certainly the few pieces available showed a more striking resemblance to those of man than to any other quadruped they could then be compared with. In Switzerland, after unearthing in 1577 some of these huge bones, the city elders of Lucerne desired to express pride in what they were pleased to consider their giant ancestor. After many enthusiastic comments they decided to figure him as bearer of the town escutcheon. More remunerative proved the resurrection of the supposedly nineteen foot tall Cimbrian king Teutobochus in 1613 near Montrigaud (Drome), in southeastern France. The astute surgeon Mazurier arranged a traveling show, making the curious crowds pay for the pleasure of viewing the relics. After parts of skulls, molars, and tusks had finally been obtained there was of course no question as to their belonging to some kind of elephant. Cuvier, who was the founder of the Department of Palaeontology in the Paris Museum, was the first to recognize that these gigantic bones exhumed in Western and Central Europe and the frozen remains in northeastern Siberia belonged to the same kind of animal, the extinct mammoth Elephas primigenius. There was naturally a keen desire on the part of the more enlightened to recover for scientific purposes at least one of the ice-mummies, soft parts and all. As early as 1722 Peter the Great of Russia gave orders to that effect to the governor of Siberia. From time to time quantities of bones collected apparently at random were sent in. Small portions of the coveted quarry occasionally reached interested centers and kept alive the yearning for real success. But in the hope of securing more complete remains the Petrograd Academy of Science before the close of the century sent several expeditions into Siberia to have exceptionally promising finds followed up, exhumed if possible, and transported to their zoological museum. During those early periods travel into such desolate, far-off regions was a slow and difficult process and rendered even the most hopeful of these enterprises uncertain. By the time adventurous men of science made their way over thousands of miles, the par- ticular frozen mammoth whose quest called them into the howling wilderness had literally melted away. Exposed soft parts rapidly decayed or were destroyed by carnivores that often scattered the 28 Zoologica: N. Y. Zoological Society [IV; 2 bones. Floods frequently carried whole portions away, or else the oncoming winter thwarted all further attempts at recovery. Not until 1806, however, came the really epoch-making find which solved many questions about this extinct form. A Tungusian fisherman in 1799 had located a complete, frozen mammoth on the banks of the Lena River at the threshold of the Polar Sea. Im- bedded in ice, as it had been for thousands of years, its meat was still in such condition as to be eagerly devoured by polar bears, wolves, and other carnivores attracted from great distances. As time went on every warm season bared more of the body; only the natives contested the booty by securing some of the meat for their dogs through the following years of exposure. It was then that the intrepid explorer and botanist Adams happened to arrive in the neighborhood and, hearing of the famed monster, lost no time in reaching it. Most of the soft parts were gone, one limb had been carried away, and a native had sawed off both tusks and sold them for about fifty rubles. Through Adams’ energy and foresight practically all remaining bones were collected. He also took to Petrograd a piece of the hide with the hair in place. It was from the still frozen side upon which the mammoth lay, and so heavy as to tax the strength of ten men to drag it along the shore. A large amount of loose, coarse hair, evidently trampled into the snow by feasting polar bears, was long enough to be considered as having formed a mane. This mounted “Adams” mammoth, to which some of the dried parts were left adhering, served Tilesius3 as a basis for the first figure of a complete skeleton, which by the way measures nine feet eleven inches at the shoulder and remains even to-day the largest ever recovered from Siberia. An Indian elephant from Ceylon but three inches less in height weighed 8,700 lbs. G. Cuvier soon after copied the figure of the Adams’ skeleton in his famous work on fossil bones.4 Subsequently the same illustration found its way into nearly every scientific text book and is still used in the eleventh edition of the Encyclopedia Britannica. It also was taken as a model for the setting up of practically all fossil mammoths found in Western Europe. Unfortunately it was far from satisfactory. The missing tusks of the mammoth had been replaced by com- 3 1815, Mem. Acad. Imp. Sci., St. Petersbourg, V, PI. X. 4 1821, ‘ Recherches sur les Ossemens fossiles.’ Nouv. ed., I, PI. XI, opp. p 204. Fig. 9. The steep slope where the Beresovka frozen mammoth was discovered. Masses of thawed ground slipped and uncovered the ice-mummy that reposed here perhaps anywhere from 12,000 to 25,000 years. 29 30 Fig. 10. The Beresovka mammoth ice-mummy in process of being recovered, after the debris had been cleared away and two-thirds of the frozen body exposed to view. The skull, in the upper left corner, has been cleaned of all flesh. 1924] Lang: Frozen Siberian Mammoths 31 bining several other pieces of ivory. According to later authorities5 these substitutes do not correspond in either size, length, direction, or curvature with those this huge bull originally seemed to have carried. For nearly a hundred years after Adams’ mammoth skeleton had reached Petrograd no important contributions were made in this line. Attempts to secure the entire frozen remains of some of the most promising of the twenty-one finds recorded during this period resulted practically in failure. They were too widely scat- tered over the bleakest of ice-bound solitudes, mostly in regions beyond the Arctic circle. Here nature seemed to be intent on holding on to one of its most fanciful creations — ice-mummies. Several Alaskan mammoths in a very much poorer and more fragmentary state were also investigated. The great credit for the rapid advance of our knowledge about frozen mammoths is due chiefly, however, to the extraordinary success of the three following expeditions: Herz-Pfizenmayer, on the Beresovka, a right tributary of the Kolyma River, Arctic Ocean drainage, Province of Jakutsk, 1901-1902; Pfizenmayer-Vollosovic, on the Sangajurach River, in the Arctic coast region opposite the New Siberia Islands, 1908; and Vollosovic, on the Liakhoff Islands, southernmost of the New Siberia Archipelago, Arctic Ocean, 1912-1913. The Beresovka Expedition was the first to profit by the rapid transportation facilities of the then new Trans-Siberian Railway. But even from Irkutsk, the last railroad station on their route, nearly 4,500 miles had to be covered on foot, horseback, and sleigh to the Beresovka River and back. In order to continue the work of salvage in the intense cold a hut had to be built over the partially exposed remains and stoves kept burning. After tremendous hardships and in the incredibly short time of ten months all that was worth while to be had of the mammoth was transferred to Petrograd. As it reached there the middle of February, most of it was still in frozen condition. This was the first time that the almost complete skin of any fossil mammal could be mounted for exhibition. Nearly all the hair had come off but some of it was put back later. For many reasons it was found advisable to represent the mammoth in the position in which it had met its untimely death. Careful study of the exceptionally perfect skeleton of this young bull, in which but one tusk was lacking, brought out many points of 6 Pflzenmayer, E. W.. 1907, Ann. Rept. Smithsonian Inst, for 1906, Washington, p. 332. Zoologica: N. Y. Zoological Society [IV; 2 Fig. 12. Portion of the trunk of the Sangajurach mammoth, showing essentially the same structure as those of living elephants. Near the lower part, to the left, a piece of the dense hair cover. Fig. 11. Skull of the adult female Sangajurach mammoth with soft parts removed except about upper portion of face. Tusks of female elephants being easily detached, es- pecially after a slight amount of decay, in this case they were not recovered. interest. Modern scientific methods of collecting made possible a number of unique results in the study of various parts, such as tongue, feet, tail, stomach, muscles, hide, fat, blood, as well as its food. (Figs. 9, 10, 17, and 18.) The Sangajurach Expedition, under the leadership of Dr. Pfizenmayer, seemed at first but little favored, for the greater part of the mammoth had been washed downstream or destroyed by Arctic foxes before the party arrived. But some lucky cause had 1924] Lang: Frozen Siberian Mammoths 33 preserved large pieces of the hide of the body and limbs with com- plete hair covering in place (Fig. 15). Even more fortunate was the recovery of some of the upper portions of the head and the nearly complete trunk (Figs. 11 and 12). In the case of the Beresovka ice-mummy the destruction of these particular parts as well as of the back by decay and carnivores had been a keen disappointment to all at the time. The Vollosovic Expedition was financed by Count Stenboek- Fermor, who presented the results to the Paris Museum. This mammoth proved to be in as good condition as the Beresovka speci- men and has helped to confirm and extend many of the researches made on the material from the two Russian expeditions. Following this the late Czar issued an imperial ukase prohibiting the exportation of any mammoth or parts thereof found in Russian territory, reinforcing a former order whereby all mammoth ivory and bone had to be submitted to a committee appointed by the Petrograd Academy of Science, that might retain any parts desired. The field observations and researches based upon the wonder- fully well preserved material from the Beresovka and Sangajurach mammoths settled a number of disputed questions. Different phases of the life history of the fabulous monsters of the frozen tundra were finally cleared up, such as appearance, structure, size, habits, and even relationship. No other fossil type has left such remarkably complete data as the Siberian mammoth and to a lesser extent its partner, the woolly rhinoceros. 6 Apart from its shaggy coat the main distinctions between the Siberian mammoth and living elephants were its much shorter, more massive body and above all its large, bulky head. The big skull had to furnish support to the enormous, spiraled tusks and weighty molars. As in recent elephants the tusks are variable in form and much smaller in the females. Their sockets run nearly parallel. At their point of emergence from the skull the tusks first diverge — sideward, forward, and upward — and then slightly converge in the general direction of the shoulder, with tips curved inward and downward. ■ The tremendous size and peculiar shape of mammoth tusks have aroused many discussions. Was so excellent a student as Adams7 right when he suggested that the hooked extremities thereof 6 Preserved parts of a mammoth and rhinoceros were also unearthed in 1907 in pits of mineral wax in Starunia, Galicia (Poland). 7 Adams, Andrew Leith, 1870, ‘Notes of a Naturalist in the Nile Valley and Malta.’ Edinburgh, p. 231. 34 Zoologica: N. Y. Zoological Society [IV; 2 Fig. 13. An abnormally spiraled mammoth tusk. Perhaps all the part rooted in the socket, and more, is missing. From the worn tip one might presume the ivory was a bothersome burden for its bearer. Much rubbing at that point somewhat reduced its thickness. may have been “used for pulling down and retaining branches of lofty coniferous and other trees”? Or is there reason to follow Pfizenmayer in his explanation that some apparently abnormal tusks with obliquely forward and downward directed tips served to break the crust of snow and scrape together food? Did these tusks grow to such gigantic proportions merely so the males might have a better chance to secure plenty to eat? Seldom would they care for the weaker among them. Nature would not treat in so step- motherly a fashion females and young, on whose welfare the con- tinuity of the race depends. In nearly all larger mammals the horns, antlers, and tusks serve essentially as weapons. In each case they are applied in the most suitable fashion. Among elephants the strongest bull of the herd enforces his right to perpetuate the race by battering every con- testant with his tusks. Just one wrong blow during the fury of a contest and these ivories snap off like glass. Not rarely have large African bull elephants left one of their tusks on such battlefields. 1924] Lang: Frozen Siberian Mammoths 35 Fig. 14. Outline engraving of woolly mammoth carved by Aurignacian man of early Magdalenian times on the rocky walls of the cavern at Combarelles, Dordogne, France. After Gapitan and Breuil, 1901. Of course the extremely large, recurved tusks of mammoths, de- scribing in many specimens fully three-quarters of a circle, un- doubtedly became useless even for such a purpose. Neuville may be correct in looking upon them as more embarrassing than useful and as showing degenerating influences at work. The largest Siberian mammoth tusk, preserved in the Petrograd Zoological Museum, measures along the outside curve thirteen feet seven and three-quarter inches, and weighs 186 lbs. The American Museum of Natural History possesses one from the Liakhoff Islands somewhat heavier, weighing 200 lbs., but only twelve feet eleven inches in length, with a greatest circumference of twenty-one inches. Lucas8 reports one from Alaska but slightly smaller, twelve feet ten inches. The trunk, as instanced by the Sangajurach specimen, from which only the tip was lacking, was dwarfed and weak in com- parison with those of recent elephants. With all proboscideans it is an important organ, the corner-stone of touch, scent, respiration, s Lucas, F. A., 1901, Ann. Rept. Smithsonian Inst, for 1899, Washington, p. 355. 36 Fig. 15. A piece of frozen mammoth skin from the upper part of the thigh with most of the hair still in place. Where longest this shaggy cpat measured about one foot six inches. From the Sangajurach specimen. 1924] Lang: Frozen Siberian Mammoths 37 and constantly used to secure food and drink, as well as for defensive and offensive purposes, and during swimming, when the body is submerged. In the mammoth, however, there is relatively little space reserved for the trunk between the huge, closely set tusks. Cor- respondingly small are the chief points for its support about the nasal and premaxillary bones. Evidently the principal function of this organ was to pluck grass from the forest meadows. Perhaps the Aurignacian cave man of Combarelles, Dordogne, France, whose rudely sketched outlines of the huge beast showed a two-fingered tip to the trunk, may still earn his fame as an observing naturalist. We might conclude from the very slight development of the trunk that, influenced by the boreal climate, the mammoth’s temper was of a milder sort. It seems not to have been used as an instru- ment of fury to devastate, break, and tear whatever may have been in its way, as is the case with the well-developed trunk of its African cousin. The ears were considerably smaller than those of the Indian elephant, measuring in the old Adams’ bull only about fifteen inches in length and six and three-quarter inches across their greatest breadth. They were densely covered with short, woolly, and longer, bristly hairs. The bony structure of the digits of the feet showed a pronounced tendency towards reduction. Some at least of these mammoths had already lost most of what in other mammals would correspond to the thumb and big toe and were four-toed (tetradactyl) and not, like living elephants, five-toed (pentadactyl) . The random numbers of toe-nails of the Paris mammoth were ascribed by Neuville9 to degeneration. Many of these supernumerary horny growths had striking resemblance to the normal nails, others were extra- ordinarily long and upturned, like those recorded from some menag- erie elephants. More decisive evidence of the mammoth’s truly boreal habitus was furnished by its heavy, shaggy coat. It covered the entire body, but even where longest it did not form a distinctive mane. In general appearance and arrangement it resembled that of the musk-ox. The dense, matted, woolly underfur, varying from fawn to golden brown, attained according to location up to two inches in length. A longer, coarser, yet fluffy hair had an average length on 9 1919, L’Anthropologie, XXIX, p. 207. 38 Zoologica : N. Y . Zoological Society [IV; 2 the body of a foot and a half; in color it was deep rusty brown, sometimes darker, sometimes lighter, according to peculiarities of preservation; its texture somewhat resembled the fibers in the hard outer covering of cocoanuts. Outstanding from this were the scarcer, flattened, considerably longer, black, but flexible bristles that ap- parently were evenly distributed over much of the body. Particu- larly graced with them were such parts as the chin, eyelashes, and ears. On the tip of the short tail they formed a long, fan-shaped tassel, but even there were only one mm. thick. The trunk was well covered with dense, short hair. On fore and hind limbs the longer coarse hair had an average length of one foot two inches; at the lower portions it was considerably shorter. As usual in mammals with dense underfur the epidermis in the mammoth was extremely thin and rather smooth, in that respect quite unlike the thick, horny, rugose, sparsely bristled skin of living tropical elephants. The leathery portion, however, according to various researches, proves to have been as thick as or thicker than that of present-day proboscideans. The histological character is essentially the same in both, neither of them possessing sudori- parous or sebaceous glands. Neuville suggests that the mammoth in evolving from ancestors living in a warmer climate and adapting itself to boreal conditions greatly reduced its epidermis. No more important factor could be cited indicating the coldness of the climate in which the mammoth lived, than the abundance of fatty tissue just below the hide. On the belly of the Beresovka male this layer was three and a half inches thick. Fat of any kind is practically absent in recent elephants, as is usual in game of tropical Africa except the hippopotamus. Its presence positively shows that at the time of death the mammoths preserved as ice-mummies were not on the verge of starvation. What better protection against the oncoming rigors of winter could be imagined than such an accumu- lation of fat, common in many boreal land and aquatic mammals and always in those that hibernate. Several lucky circumstances have contributed towards our fairly satisfactory knowledge about the feeding habits and food of mammoths. From what we know about living elephants the ex- perienced can tell from a glance at the molars that the mammoth secured its livelihood essentially by grazing and not by browsing. Its cheek-teeth present a densely crowded condition of the compo- nent transverse plates with comparatively even, yet characteris- 1924] Lang: Frozen Siberian Mammoths 39 Fig. 16. Various stages in the development of upper and lower molars of the mammoth C Elephas primigenius Blumenbach). Anterior portion pointing toward left. a. Third right lower molar at height of efficiency. Enclosed within the high rims of enamel of each of the numerous transverse lamellae or their parts lies the softer and lower dentine. The lamellae in turn are surrounded and united by cementum. Anteriorly this molar is worn low. b. Second right upper molar. Some of the anterior transverse lamellae have been shed, and the remaining ones are more worn than in Fig. a. c. Third right lower molar. Some of the posterior lamellae have not yet moved up from the jaw to the grinding surface. d. Second left upper milk molar before piercing the gum. The individual transverse lamellae, still crested, have not yet been united by cementum. e. Second left upper milk molar well worn, some of the transverse lamellae in front having been shed. /. First left upper milk molar with a grinding surface of only 12 mm. This cheek- tooth belonged to a calf probably not more than a year old. 40 Zoologica: N. Y. Zoological Society [IV; 2 tically rough, crown surfaces. Their peculiarity and efficiency are greatly influenced by the singular action of food-stuffs under the pressure and friction of mastication. It is the amount of sili- cate, grit, and other hard materials contained in the food that causes an unequal rate of wear of the three principal substances composing these grinders. Under these circumstances the ex- cessively hard enamel parts always remain as ridges, whereas the softer dentine and cementum form alternating depressions. The transverse plates of the cheek-teeth vary somewhat in complexity and number; from three in the first (milk molar) to as many as twenty-seven in the sixth or last molar. Osborn10 has shown that Jefferson’s mammoth ( E . jeffersoni) of the American Pleistocene, from Indiana, may have as many as thirty in the upper molar and from twenty-four to twenty-six in the lower. The masticating surface of the large, broadened molars* * of the mammoth forms an efficacious milling apparatus for the grinding to pieces of the rather tough, but very nourishing, boreal meadow plants. A similar arrangement would not answer so well for the bulky, succulent masses of tropical vegetation on which the living Indian and African elephants subsist. The relatively compact nature of the fodder of the mammoth may have helped lessen the need for accommodating immense digestive organs and finally have led to a general reduction in size of the animal’s body, back of the head, as mentioned above. Borodin11, was able to identify the plants found between the molars and in the stomach of the Beresovka mammoth. Above its ice mausoleum the flora of these Siberian forest meadows still showed essentially the same characteristics as thousands of years ago at the time of the victim’s entombment. The average temper- ature may have then been responsible for a more uniform, milder climate, somewhat lacking the intense severity of present winters 10 Osborn, Henry Fairfield, 1922, Amer. Mus. Novitates No. 41, pp. 1-16. 11 In: Salensky, W., 1905, Compt. Rend. Sean. VI Congres Internat. Zool., Geneve, (1904), p. 72. * The molars, as in other elephants, after emerging from the gum gradually move for- ward in the jaw. As the upper portion, the grinding surface, is worn down, the originally long, anteriormost roots are absorbed or “ eaten up ” within their sockets by special cells, the osteoclasts. Thus only a flat, thin, center portion of the anterior part of the molars remains. As this is pushed out beyond the forward part of the socket the thin pieces of the composing plates break off easily, the worn ones being replaced by succeeding new ones from the rear. In this way the rather voluminous and heavy molars are easily accommodated in so limited a space and retain their efficiency till a relatively advanced age. 1924] Lang: Frozen Siberian Mammoths 41 and allowing the forests to reach as far as about 74° North. The food gathered in abundance by the Beresovka mammoth just before death consisted of various kinds of grasses ( Alopecurus , Hordeum, Agrostis , Atropis, and Beckmannia) . Sedges were represented by two forms of Car ex. A mint {Thymus), pods of a leguminous plant {Oxytropis), wild poppies {Papaver), and seeds of the northern butter daisy ( Ranunculus ) made up the list. Some pine needles and bits of wood figure as incidental occurrences. From the above enumeration of characters it appears out of question, as formerly believed, that Indian elephants could be modified forms of the Siberian mammoth and had merely wan- dered southwards into the more luxuriant forests of tropical Asia, and in adapting themselves, had lost their heavy pelt and gradually changed otherwise. As shown above, the Siberian mammoth was in many ways too highly specialized to figure as an ancestor of the living Indian elephants, which must have evolved from some other form.12 As an argument against the boreal character of the mammoth there has been advanced the fact that in southern regions its remains were found mixed with those of such tropical types as the cave hyaena (■ Crocuta crocuta spelaea) and the cave lion {Leo leo spelaea ), that actually had gnawed its bones. The Pleistocene European hip- popotamus {Hippopotamus major) has been cited to the same purpose. It is not so uncommon a feature among various groups of recent mammals to travel about in regions having relatively great differences in temperature and presenting a variety of environ- ments. Our American bison once roamed from the plains of northern Mexico to the woodlands of Canada beyond Slave Lake. Another example is our puma, of which Theodore Roosevelt writes in his admirable account : “ It is found from the cold, desolate plains of Patagonia to north of the Canadian line, and lives alike among the snow-clad peaks of the Andes and in the steaming forests of the Amazon." Another instance is offered by a race of white-footed mice {Peromyscus) , whose footprints Dr. E. W. Nelson records having seen at 15,000 to 16,000 feet above sea-level on the volcanic ashes of Mt. Orizaba, Mexico. It thus furnishes the altitude record for North American mammals. In Africa the browsing elephant {Loxodonta africana), with 12 Osborn, Henry Fairfield, 1910, ‘The Age of Mammals,’ New York, p. 419. 42 Zoologica : N. Y. Zoological Society • [IV; 2 its preferred haunts in denser wooded parklands, roams also over trackless swamps and enters arid desert stretches. The grazing buffalo 0 Syncerus caffer radcliffei) leaves the plains and invades forests, making itself at home even at 10,000 feet. Both on Mounts Kenya and Kilimanjaro records of the two visiting snow-fields and glaciers are at hand. More surprising still are the giraffe ( Giraffa Camelopardalis tippelskirchi) , exploring the mountain forests of Kilimanjaro, and the eland ( Taurotragus oryx patter sonianus), going even beyond to the mountain meadows. The lion in East Africa ascends from the lowlands to above 7,500 feet as in the Rift Valley and within the range of the mountain gorilla {Gorilla beringeri) on Mount Sabinyo,13 reaching altitudes where the temperature during the night may drop below the freezing point. The lion has been bred with success freely exposed to the wintry rigor of the climate of Dublin. Leopard {Panther a pardus suahelica ) and hyaena {Crocuta crocuta germinans) , also typical animals of the lowland, go to over 9,000 feet on Mount Kenya. There, up to 15,000 feet, near the border of eternal snow, hyraxes {Procavia mackinderi mackinderi) too occur, differentiated only subspecifically from the lowland form. Colobus monkeys {Colobus abyssinicus kikuyuensis) are none the worse for icy-cold nights at 10,000 feet, though equally at home in the hot valleys far below. River-horses {Hippopotamus amphibius) even in captivity seem not to be so susceptible to cold as generally believed. In the zoological garden in London at least they were known to take their tubs in frosty weather. Africa, with its very restricted mountain areas, gives no fair basis as to what happens in Asia with its more extensive ranges and mountain plateaus, or to what might have taken place in this respect during the glacial periods of the Pleistocene. Even among recent mammals the list could be increased considerably. One need merely mention the hardy, long-haired Manchurian tiger and the well-furred snow-leopard with firmly established haunts in colder climes though their closest relatives inhabit the tropics. The camel {Camelus bactrianus ) and the yak {Poephagus grunniens), that sur- vive the icy blasts of the Tibetan plateaus, show not the slightest effect in their welfare or reduction in breeding on descending into more temperate zones. Certain it is that many of the high lands of the Pleistocene presented a wide, open expanse with an abundance of excellent i» Philipps, J. E. T., 1923, Geogr. Journ. London, LXI, p. 247. 1924] Lang: Frozen Siberian Mammoths 43 pasturage, as indicated by the large herds of gregarious mammals. There should be no surprise that some of the southern carnivores, like the lion, hyaena, and others, followed up such promising prey. Perhaps the borders of rivers in the summer offered also an abundance of choice fodder to the hippopotamus. All points considered, there is no reason why the hairy mammoth should not have wandered south. Its rambles may even have been undertaken during the colder season. Being great nomads, like most of their relatives, the mammoths unquestionably wandered back and forth through most of the northern countries of Europe, Asia, and America. During the moist, cool climate of the third glaciation they made their first ap- pearance in Western Europe, going as far west as the British Isles, at that time a peninsula, with the North Sea firm land; and even to Denmark and Scandinavia, where it was probably the remaining glaciers that stopped them at 62° North in Norway at Saejervaskter in Vaage; attaining, however, to 65° 30' North in several places in Finland bordering the extreme north of the Gulf of Bothnia.14 They also went southward to northern Spain and to Italy within the neighborhood of Rome. From northern Siberia they passed over to Alaska and America by way of Bering Strait or the Aleutian Islands before the separation of these continents took place, thence to California and across to North Carolina.15 On the American continent their evolution progressed into still more gigantic forms as they reached evidently more inviting regions farther south. Encouraged by slight fluctuations of temperature while glaciers were slowly advancing and retreating during the Pleistocene period the mammoths, like other mammals, shifted according to seasonal changes, either north or south, just as some of the African elephants nowadays accommodate themselves to dry and wet periods by travel- ing from the lower plateaus into the mountain forests and to escape from the annual grass-fires of the savanna into the safety of extensive swamp-lands. At the close of the glacial periods their haunts must have vitally changed. The mammoths apparently were not able to follow any more in the wake of the retreating ice and must have encountered conditions that sealed their fate. Most interesting is the evidence of what the gigantic beasts 14 Holst, Nils Olaf, 1913, L’Anthropologie, XXIV, pp. 363-364. “Matthew, W. D., 1915, ‘Mammoths and Mastodons,’ Amer. Mus. Nat. Hist., No. 43, Guide Leaflet Series, p. 6. 44 Zoologica: N. Y. Zoological Society [IV; 2 must have meant to the cave-dweller of Europe. The Cro-Magnon men of France16 were the first to leave for posterity authentic out- lines of these monsters they had hunted. One of the finest examples is the sketch of a huge, shaggy tusker cut into a slab of mammoth ivory found in “La Madeleine” cave. This and other equally characteristic pictures, such as a curious little figure of the mammoth found at Predmost, furnish proof of the absorbing interest early Paleolithic man evinced in glorifying the enormous beast his heroes succeeded in overpowering. These early artists evidently wished to commemorate the bearer of so bountiful a supply of meat. They undoubtedly used parts of it as talismans, as in the case of a child’s necklace of mammoth ivory beads found at Predmost. In all probability the extinction of the mammoths'was a gradual process and may have lasted hundreds of years or more. No single cataclysm, as Howorth17 believed, could have been widespread enough to account for their abundant, mostly scattered remains throughout the Holarctic regions preserved, as they are, in such different ways. Besides innumerable traces in northeastern Siberia, the neighboring Polar Sea, and the American continent, great accumulations of their fossil bones occur also in certain places in Europe. Predmost in Moravia, where some eight hundred or nine hundred individuals were counted, is particularly famous, but the mammoth deposits near Cannstadt in Wiirtemberg and Hofstade in Belgium illustrate similar instances. According to all authentic reports the mammoths preserved as ice-mummies, and found under various conditions of entomb- ment, perished singly. Some of them were in prime condition, as young and fat individuals prove, and had plenty of fodder in their stomachs. These facts strongly favor the view that they met with accident, as instanced by the Beresovka and other finds. For this reason they have hardly any direct bearing on the real causes of extinction of their race that is to be set at a much later period. In a way they might be compared with the frozen body and a skeleton of the African buffalo found by Ross18 and Mackinder, respectively, at about 14,000 feet on the glaciers of Mount Kenya. Wandering away from one of the many herds in the plains these rovers had perished on their unknown but curious excursion. is Osborn, Henry Fairfield, 1921, ‘Men of the Old Stone Age,’ 3rd ed., New York, pp. 397-398, figs. 197-199. 17 Howorth, Henry H., 1887, ‘The Mammoth and the Flood,’ London, p. xviii. 18 Ross, W. McGregor, 1911, Journ. East Africa and Uganda Nat. Hist. Soc., II, p. 63. 1924] Lang: Frozen Siberian Mammoths 45 It is significant that only the most gigantic mammals of this decidedly gregarious Pleistocene fauna have been transmitted to posterity in frozen condition. Their tremendous weight and re- lative clumsiness seem to have played an important role. Did they slip and fall, or were they precipitated to depths where cold would preserve them? Or were some of their bewildered troops devoid of the necessary agility and grit to extricate themselves from overwhelming storm and deep snowdrifts? Or did furious gales and blizzards cover them alive with icicles that quickly grew to encasing blocks of ice? As regards greater catastrophies, sub- siding land-masses may have brought their doom, or inundations engulfed them. A few may have found their final resting places in swamps and bogs. Considering the various finds, certainly some and perhaps all of the contingencies enumerated contributed their share to the final extinction of the mammoth. Every spring as a result of the setting in of warmer weather the important Siberian rivers move enormous masses of ice towards the Polar Sea. The clearing away of these obstructions is watched with intense excitement by the inhabitants of these ice-bound regions. A few months of river navigation means new freedom of traffic. They again can rove far and wide. It is then that hungry dogs may lead their masters to the masses of strongly smelling meat of the “mamonto” that has incidentally been uncovered along the thawing coves and banks. Mighty are the struggles oncoming spring leads against the wintry forces. For a while great portions of these streams are dammed up by mountains of constantly shifting ice-floes. When these finally break through, parts of a new channel are often enough quickly ploughed up. After the generally sudden subsidence of these temporary floods, the old, abandoned stream bed freezes over again, imprisoning huge blocks of ice, debris, and all. Some of the water for a while held below the icy cover drains off and here and there large scattered pits are left between the chaotic masses. After years have leveled out these sites, ponderous beasts like mammoths wandering over such' treacherous places might easily break through, be instantly killed, or become hopelessly mired. Later on erosive material piled over all may have formed the basis for the surface soil that practically furnished permanent protection. A few frozen bodies found in what was considered alluvial soil, mixed with pieces of ice, in a position as if ready to walk off, may have perished in this manner. Fig. 17. The mounted mammoth unearthed in 1901 as an ice-mummy on the banks of the Beresovka River, northeastern Siberia, as it appears on exhibition in the Zoological Museum in Petrograd. The animal has been partly restored. Much of the real skin could be used in the mounting. Some of the hair has been replaced. 1924] Lang: Frozen Siberian Mammoths 47 The cramped position, broken bones, large amount of clotted blood in the body cavity, as observed in the Beresovka mam- moth, point, as Salensky shows, towards instantaneous death by accident. The victim did not even have time to throw out or swallow the quantities of fodder between its molars and in process of mastication. Salensky gives a cause for such a tragedy. During extremely rigorous winters the formation of wide fissures in the ground is not rare in northeastern Siberia. With the oncoming warmer season these clefts rapidly fill with water which may cause extensive subterranean washouts. Later some colder spells may cover such basins with a sheet of ice, below which the percolation of the remaining water continues in other directions, thus giving rise to what really amounts to an underground cave. Subsequent strengthening of the surface ice and final covering with humus, until level again, would form a sufficiently strong cover for everything except the weight of so colossal a beast as the mammoth or rhino- ceros. Late in summer such places might be specially weakened and the unfortunate animal crashing through into the cavities would be instantly imbedded in the masses of ice and frozen debris loosened by the accident. Severe storms and periods of intensive freezing that usher in winter would soon remove all traces of the entombed. For thousands of years the victim might never be moved, except through the infinitely slow processes of floods and similar erosive actions. Without question all those mammoths discovered frozen in practically fresh condition were at the very moment of their de- struction surrounded by a temperature that completely excluded decomposition. The perpetually frozen ground of northern Siberia acted much like a modern refrigerator. For periods variously estimated at anywhere from 12,000 to 25,000 years, such ice- mummies may have reposed far below the protecting mantle of tundra vegetation. Here, as in the vast expanses of the “taiga,” the swampy Siberian forests, they were, so to speak, permanently protected, the ground being frozen generally at three feet below the surface even during the hottest of summers. That great numbers of these and other mammals perished without being pre- served in such perfect fashion is sure. In some places the earth and the shores of certain islands along the Polar Sea were literally crammed with their bones. The solid clay of higher sites inland preserved them pretty well, but elsewhere climatic conditions 48 Fig. 18. The Beresovka mammoth skeleton as mounted in the Petrograd Museum. One tusk and several ribs had to be replaced. To the top of the head the animal’s height is given as ten feet nine inches. 1924] Lang: Frozen Siberian Mammoths 49 fostered the more rapid decay of others, or without doubt there would be many more. Mammoth tusks have for many reasons aroused considerable interest. To the scientist they are the permanently growing second pair of upper incisors, composed mainly of a solid but peculiar den- tine, the “ivory” of commerce. To the poor Siberian hunter, with his pick and ax ready to chop into pieces whatever tusks he can discover on his migrations in the wilderness, they are “white gold.” To the Chinese artist the delicate, fine texture and peculiar pallor of the easily carved substance brings new incentive for his varied talents. Strongest is the claim of the superstitious. Small parts of mammoth ivory have meant to him the chance of his life. They have served even as relics of Christian saints. We are told that devout prayers addressed to them have given earnestly hoped for succor and success. Heathen with still stronger beliefs deeply implanted in their hearts as regards the occult powers of this mar- velous substance have had their satisfaction too. Equally many- sided were the supposed medicinal virtues of mere scrapings. The Chinese have been more beguiled by them than by their “dragon bones.” And western Europe did not free itself so very long ago from faith in such wonder cures. With their application, hemor- rhages, ulcers, broken bones, epilepsy, fevers, plague, and cholera19 would all vanish, according to the fancy of many. For that very reason they furnished princely revenues. As many as sixty tusks from the fossil mammoth deposits at Cannstadt were sent to the pharmacy of the Court, and became the precious powder of the “Licorne.” Gross credulity has been carried even into other channels. What of the warrior whose sword hilt, carved of mammoth ivory, is worth more to him than one wrought of gold or silver? In the matter of art Siberia itself has made little use of its great supply of ivory. Only a few figurines, animals, characteristic scenes of the native land, often in heavy relief or bold freedom, combs, vases, and boxes, are made in certain centers. Export of the crude tusks after all has been the mainspring of their efforts. A great impulse to the exploitation of Siberian mammoth ivory along the edge of the Polar Sea, its cliffs, and islands near the mouth 19 Kunz, George F., 191G, ‘Ivory and the Elephant,’ p. 239. 50 Zoologica : N. Y. Zoological Society [IV; 2 of the Lena, was given when Catherine II, Empress of Russia, took a personal interest in the matter. In October 1771 she wrote to Voltaire20: “But what proves, I think, that the world is a little older than our nurses tell us are the finds of bones of elephants long ago extinct in these regions and imbedded several fathoms below the surface of the ground in northern Siberia. Scientists . . . have said that it is fossil ivory, but, how is it possible? fossils do not grow in the form of very complete elephants.” Some time before, she had given orders to investigate the archipelago later known as the “New Siberia Islands,” whose highest point attains 1,200 feet. The southern two, low, and completely uninhabited Liakhoff Islands were named in honor of the fur mer- chant who, following the tracks of an enormous herd of reindeer coming from the north, discovered what later proved to be inex- haustible mines of mammoth ivory. The Czarina also had conferred on Liakhoff the exclusive right to hunt and to collect ivory on them. Many huge tusks were partly sticking out of the sand and others, together with bones, were constantly swept up on the shores by the waves. To what depth do these marvelous deposits cover the sea-bottom no one seems to know. Did these vast stores of wealth come from further inland, and were they carried out to sea with the crushing masses of ice in the spring? Here and there a frozen mammoth might have been moved thus along when whole sections of the partly thawed up river banks were undermined or torn out. Or did countless numbers of these huge beasts make their last desperate stand in these regions before the land was swallowed up by the sea? Evidence for subsidence of land masses is more certain, as apparently much of this expanse once formed part of the Asiatic mainland. On some of these islands Silurian coral and Devonian limestone, volcanic rocks, indicate that uplifts, as might be expected, had a part in the present physiographic configuration. On the northernmost, Hedenstrom found Tertiary strata with fossil bi- tuminous tree trunks in horizontal and upright positions, over 200 feet above sea-level. Other rich deposits of the same age with their interesting fauna and flora indicate a climate once very much warmer. But some of the lower islands off the coast show a few peculiar granite boulders and are covered with a deep mantle of drift formed chiefly 2° Boule, M., 1917, L’Anthropologie, XXVIII, p. 187. 1924] Lang: Frozen Siberian Mammoths 51 of sand, and buried ice in separate layers and incongruous blocks. These deposits were particularly rich in ivory tusks and masses of mammoth bones. Associated with them were the remains of other of the northern Pleistocene mammals, such as the woolly rhinoceros (Coelodonta antiquitatis) , Siberian bison ( Bison prisons), wild horse (, Equus caballus fossilis) , moose {Aloes latifrons), reindeer {Rangifer tarandus ), and musk-ox (Ovibos fossilis). Unquestionably the mammoth was boreal in habits and most abundant in the colder regions. In northern Siberia, it flourished in all the territories between the Ural Mountains, Obi, Yenisei, Lena, Indigirka, and Kolyma, and particularly in the adjacent islands of the Polar Sea. These, therefore, with their fabulous stores of ivory, are the greatest graveyard of mammoths known. Von Wrangell described some parts of this region as containing hecatombs of such remains before they were ransacked by those in search of the valuable tusks. Should we wonder that for over a hundred years organized ivory collecting flourished without any apparent diminution of the supply? The rigor of the climate imposed by far the greatest drawback to this greedy quest. At Liakhoff Island the open season lasts really but a few months. Bunge21 in 1882-1884 records 90° F. below freezing point in winter, with snow falling half through July. But even under such trying circumstances enormous quantities of ivory have been removed. In 1821 one trader alone sent off 20,000 lbs. Middendorff in 1841 estimated the annual output for the preceding twenty years as at least a hundred pairs of tusks. In 1881 Nordenskiold,22 basing his opinion partly upon the amounts still shipped, considered this figure as rather too low than too high. He arrived at the conclusion that since the conquest of Siberia select tusks from 20,000 mammoths had probably reached the markets of the world. From Westendarp we know that the fairly well stabilized imports to Europe of fossil ivory in 1872 — with London then the chief market — amounted to 1,635 mammoth tusks or about 245,250 lbs., granting an average weight of 150 lbs. apiece. The proportion of well preserved ivory among such lots is surprisingly small — only 14 per cent; and a slightly larger amount, 15 per cent, is absolutely 21 Bunge, 1893, Congres Internat. Zool., Moscou, Session II, (1892), pt. 2, p. 282. 22 Nordenskiold, A. E., 1881, ‘The Voyage of the Vega around Asia and Europe,’ London, I, p. 404, footnote. 52 Zoologica: N, Y. Zoological Society [IV; 2 useless. But even the really “bad,” amounting to 54 per cent, and the “still workable,” 17 per cent, when treated properly and fashioned into the plainer objects passed in the trade. For the most part these could not have been tusks of mammoths entombed in ice, the ivory of which compares well with that of re- cently killed elephants, but evidently had been subject to various disintegrating influences. So great was the demand for this re- munerative article that in Europe mammoth tusks of far inferior quality to the Siberian product were formerly dredged in quantities from the Doggerbank by the North Sea trawlers. In rare cases mammoth ivory is slowly impregnated with certain metallic salts and then known as odontolite or blue ivory. Used for jewelry it is highly prized for the delicate, yet vivid, blue, tur- quoise-like luster. The Eskimos of Alaska, according to Gilmore, are fond of a blue dye they secure from the phosphate of lime (vivianite) formed by the decomposition of mammoth tusks. Many hundreds of thousands of these enormous tusks must have completely decayed. What great herds of shaggy mammoths may have roamed during Pleistocene times in the proximity of the circumpolar area can be deduced from Darwin’s computations23 about the possible increase of the recent elephant, considered the slowest breeder of known animals. If, at a minimum rate of natural increase, between the ages of thirty and ninety years, only three pairs of young be raised, he comes to the conclusion that “at the end of the fifth century there would be alive fifteen million elephants, descended from the first pair.” The Pleistocene mammoths during several hundred thousands of years had totally adapted themselves to a life in a monotonous, frigid zone. Uniformity indeed is the hall-mark of boreal regions as much as diversity is that of the tropics. During so long a period they gradually became highly specialized, long-lived monsters. Being excessively slow breeders they entered a stage where further evolution or even slight adaptive changes were reduced to a minimum. This meant the death warrant of their race. Perhaps in the boreal climate the balance of endocrine functions had long before been disturbed so that undesirable specialization went on unchecked and possibilities of forming varieties of greater vitality were practically eliminated. But whatever the causes of their final 23 Darwin, Charles, 1860, ‘On the Origin of Species,’ 2nd ed., p. 63. 1924] Lang: Frozen Siberian Mammoths 53 extinction, here, at least, nature has preserved from the enormous numbers of these shaggy monsters a few victims of individual accidents as ice-mummies. They have now become a unique source of information. Still others rest in frozen Siberian ground waiting to disclose more secrets of bygone ages. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY THE GALAPAGOS TORTOISES IN THEIR RELATION TO THE WHALING INDUSTRY By Charles Haskins Townsend Director of the New York Aquarium PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK July 29, 1925 •Nfiu fork 2nnlogtral §>urU'ttT General Office: 101 Park Avenue, New York City ODfftrmi President, Madison Grant; Vice-Presidents, Frank K. Sturgis; and Henry D. Whiton; Chairman, Executive Committee, Madison Grant; Treasurer, Cornelius R. Agnew. Inarli of IHanagpra (Ela00 of 1023 Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor OIla00 of 102r Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field ^~ P P&sy/~~ fP^C &c7ot« r ^&^s4o^ £/0*0^o -csP'tP c-t+0 fipP~~ // t/tPt^ ^iPpT^Jc' 0 T/Ct SPtpyPT'&t^tP PP pi 'Pl Cy- s^x / 7 -^GTPP c^' -4^/C*Sp*7‘ P C*«ss7 tPj ^tppj£i i 2~ p,00*%7 ~P'>^iyP yc>*+~-7 &t/f ^ //'?(*? c > -*-«-i PP&-0 0^0 ^ &^y- 7 &Sss** pPr PPtPi ~P^£+S1 (p)c*sy £Ss7K0k^//rs<} d'yxs fPlsv J^4 &ooo-n ^^-<^S1 S~i J ' e'C*^6> y^y-ye^^s-/! yy Jfl^Pkt- ^>1 y~c7Pp7~ P'yJD-eo y O07t^,y, ox^sr 7%* P*. ~>^ 07—0, s^'Tyi y ^ y'y'er. ' /ypt^sjLfr 7 7^/00?/ yjP 7>T/y /^yp^pTy^ \/yp/ SP'*0*p-c~ & tP3> & 6>t7!} oT- Pppp-e ~or>x. y>0 +■ P-e-Ppyp TP" P? t>P? Pix Q?. sPPy - - . _ r Fig. 24. A page from the logbook of the whaleship Isabella of New Bedford. At Hood Island, Galapagos in 1831. Between December 4 and 8, this vessel captured 335 tortoises. 74 1925] Townsend: The Galapagos Tortoises 77 In' the following list of vessels there is shown a catch of 10,373 tortoises. The number of vessels participating is given as one hundred and fifty-one, but it should be understood that this number means visits when tortoises were secured; many vessels having called at the islands two or more times. The ship George and Susan ob- tained tortoises seven times between 1835 and 1856; the ship Hector seven times between 1832 and 1843, and the ship Congaree eight times between 1847 and 1853. Large catches were often made. Four vessels took over three hundred each, eight other vessels over two hundred each, thirty-three over one hundred each, and twenty- four over fifty each. The largest recorded catches of tortoises from a single island are 350, 335, 315 and 310: TORTOISES TAKEN FROM THE GALAPAGOS ISLANDS BY CERTAIN WHALESHIPS FROM 1831 TO 1868 8 Catch shown by Islands Year Vessel Island Tortoises 1831 Ship Isabella Hood 335 i ( “ Magnolia Charles 155 “ Hesper Hood 250 < 6 “ Fiances Charles 179 f Hood [ Abingdon 50 1832 “ Abigail 8 “ “ Hector Charles 226 pi 1833 “ Hector Albemarle 9 “ “ Pacific Indefatigable 44 1834 “ Abigail Indefatigable 140 “ “ Bengal Charles 100 “ Moss \ Chatham 8 [ Charles 350 “ Loper Hood 237 t ( “ Hector James 23 pi Bark Benezet j Charles 120 i i { Indefatigable 12 1835 Ship Barclay 9 Charles 50 “ Hector f James 124 { Albemarle 2 “ “ George & Susan James 68 8 For extracts from the logbooks of these vessels see appendix, where they are ar- ranged chronologically as in this list. 3 First voyage made in 1795. 78 Zoologica: N. Y. Zoological Society [IV; 3 1 [ Charles 40 1835 Bark Benezet j [ Abingdon 12 Ship Lima James 35 plus i i Albemarle 67 i i Phoenix Abingdon Hood 10 65 1836 Bark Pioneer Indefatigable 2 plus many << Ship Eliza Adams Albemarle 23 a Bark Hesper James 13 plus many Ship Lima f Chatham 20 i ( l ( James 118 1837 “ Abigail Abingdon 142 u “ Eliza Adams Charles 24 “ ‘ ‘ Lima James 224 a “ Omega Chatham 240 1838 “ Corinthian Hood 136 i i “ Charles Albemarle 8 u “ George & Susan Chatham 67 i i “ Phoenix James 12 plus 7 b loads 1839 “ George & Susan Barrington 22 i ( “ Charles Albemarle 20 \ Hood 12 il “ Robert Edwards i [ Albemarle 7 1840 “ Robert Edwards Chatham 59 “ “ Rousseau10 Hood 45 t: “ Mariner Chatham 115 1841 “ Elizabeth Chatham 102 | Albemarle (Crossman) 16 i L “ Chili ! James 93 [ Albemarle 10 6 6 “ Rousseau Albemarle 12 ( 6 “ Pocohontas Albemarle 47 “ “ Hector Albemarle 24 1842 “ James M unroe Albemarle 64 “ Ship Eagle Albemarle 36 a “ Rousseau Albemarle 10 “ Chili Chatham 118 a “ Lion Albemarle 5 i i “ Robert Edwards Chatham 107 i 6 “ Hector Hood 173 f Chatham 30 a “ Navigator i Hood 5 1843 Bark Garland Hood 100 < i Ship Robert Edwards Chatham 262 10 The oldest whaler built in 1801 for Stephen Girard of Philadelphia. Broken up at New Bedford in 1893. 1925] Townsend: The Galapagos Tortoises 79 1843 “ Hector Abingdon 67 f Hood 20 1844 “ Callao j [ Albemarle 4 plus [ Chatham 24 1844 Bark Equator < [ Albemarle 9 [ Hood 14 Ship Levi Starbuck j [ Chatham 130 ii “ Charles Chatham 100 1845 Bark Equator Albemarle 69 1 James 20 ii “ Alfred Tyler ] Abingdon 7 1 [ Indefatigable 45 1846 “ Equator j Albemarle 150 Chatham 14 i i (Name lost) Chatham 190 ii Ship Aurora f Albemarle 2 l Hood 7 ii “ Minerva Chatham 120 1847 “ Coral [ Albemarle 1 [ Abingdon 1 “ Susan Albemarle (Crossman) 30 “ Bark Alfred Tyler Abingdon 3 1847 Ship Charles Frederick Hood 67 i i “ Aurora Chatham 100 ii “ Congaree Chatham 4 “ . “ Elizabeth Chatham 100 1848 “ Susan | ' Albemarle 186 Abingdon 23 i i “ Corinthian Chatham 54 “ Roman j [ Duncan 50 ; Indefatigable 36 “ “ Congaree ’ j r Abingdon ; Chatham 10 70 “ “ Coral Chatham 200 1849 “ Susan Albemarle 2 i i (Name lost) Albemarle 63 i i Ship Congaree Chatham 130 ii “ Kingston Abingdon 6 f Hood 1 Brig Vesta | Abingdon 5 1850 Ship Susan Chatham 156 i i ‘ ‘ Peruvian Duncan 131 “ “ Martha Chatham 110 1851 “ Pocahontas Chatham 90 1852 , “ Congaree Abingdon 5 ( Albemarle 2 i i Bark Eugenia { Chatham 107 80 Zoologica: N. Y. Zoological Society [IV; 3 1853 Ship George & Susan Abingdon 3 i i Bark Henry H. Crapo Barrington 1 . ^ 1 f Chatham 315 i i Ship Congaree j [ Hood 7 “ Bark Peru Albemarle 150 “ “ Martha Chatham 13 1854 Bark Eugenia Abingdon 3 < i “ Superior Albemarle 1 ii Ship Potomac Chatham 43 1855 “ George & Susan Chatham 152 Chatham 28 i £ Bark Cornelia Albemarle 14 Chatham 4 6 ( Ship Mary Ann Duncan 17 6 ( Bark Superior Albemarle 11 “ “ Benjamin Cummings Chatham 310 1857 “ Bevis Albemarle 13 1858 “ Morning Star11 Albemarle 24 f Chatham 78 1859 “ Montgomery \ [ Albemarle 7 n Ship Lancer Chatham 70 1860 Bark Ohio Albemarle 81 i i “ Ospray Albemarle 122 “ “ Atkins Adams Albemarle 14 < ( Ship Edward Carey Albemarle 56 plus 1861 Bark Stella Albemarle 6 “ “ Ospray Albemarle 41 1 1 “ Morning Star Chatham 188 “ Ship Roscoe Chatham 50 < < “ Arnolda Chatham 42 “ Bark Atkins Adams Chatham 105 “ “ 0/w‘o Chatham 50 1862 “ Abingdon 4 i i Ship Edward Carey Albemarle 95 i i “ Roscoe Albemarle 63 , . , ^ 1 [ Chatham 1 1863 Ship Edward Carey [ Duncan 208 1867 Bark Osceola 2nd Abingdon 1 1868 Ship Roscoe Albemarle (Cowley Islet) 5 Total visits by above-named Total recorded 67 vessels — 151 catch 10,373 11 Built 1843, last voyage 1914, see frontispiece. 1925] Townsend: The Galapagos Tortoises Above Catch of Tortoises Arranged by Decades 81 Showing number of visits by vessels and average taken at each visit Decade Tortoises Visits by Vessels Average 1831-1839 3809 44 86 1840-1849 3567 61 58 1850-1859 1865 28 67 1860-1868 1132 18 63 Total Tortoises 10,373 Total visits 151 General Aver. 68 Same Catch Arranged by Islands 1. Chatham 4,326 6. Duncan 356 2. Albemarle 1,581 7. Abingdon 310 3. Hood 1,524 8. Indefatigable 279 4. Charles 1,244 9. Barrington 23 5. James 730 Total tortoises 10,373 In the following list of twenty-nine vessels making thirty-eight visits to the islands the numbers of tortoises taken are not recorded. Allowing these vessels the same average catch for each decade, as the sixty-seven vessels making one hundred and fifty-one visits, with definite records, their catch may be estimated conservatively as follows: Estimated Catch of Tortoises Based on the Recorded Catch — Arranged by Decades Decade Visits by Vessels Average Tortoises 1833-1838 12 87 1044 1840-1849 14 58 812 1850-1859 7 67 469 1860-1867 5 63 315 38 Estimated catch 2640 The recorded catch of 10,373 tortoises by vessels making one hundred and fifty-one visits, and the estimated catch of 2640 by vessels making thirty-eight visits, gives a total of 13,013 tor- toises taken during one hundred and eighty-nine visits. 82 Zoologica: N. Y. Zoological Society [IV; 3 List of Vessels Seeking or Obtaining Tortoises at Various Islands from 1833 to 1867, but Numbers Taken not Recorded Logbook Entries “At anchor at Abingdon” “went for terrapin” “the boat wint on shore” “employd giting turpin” “boats returned with turpin” “boats returned with turpin” “boats after turpin” “boats returned with turpin” “obtained Terapine” “imployed geting turpin” ‘stowing Terrapin” Year Vessel Island Date 1833 Ship Loper Abingdon Aug. 7-8 1834 “ Bengal Albemarle Apr. 1 i i “ L.C. Richmond Charles July 16 ii “ Ohio James Feb. 26- Mar. 1 1835 Bark Pioneer Charles May 7-14 “ Ship Lima Chatham June 23-25 1836 “ Ohio Charles Apr. 12-15 ii Bark Pioneer Charles July 16-17 1837 Ship Elizabeth Albemarle May 17-18 Indefatig- ii “ Eliza Adams j able James May 7-9 1838 “ Omega James June 26- July 2 Albemarle (Cross- 1842 “ Ocean man) Jan. 25-27 Chatham July 26- Aug. 3 “ George Wash- ington Hood Nov. 1 ii “ James Monroe Albemarle July 2-3 1843 “ Phebe Albemarle Jan. 12 1845 Bark Alfred Tyler Albemarle Nov. 11-12 ii (Name lost) James Aug. 18-23 1846 Ship Ann Alexander Hood (Records mis 1847 Bark Persia Hood Jan. 6-7 a “ Alfred Tyler Albemarle Apr. 24-25 a Ship Congaree < [ Abingdon [ Charles July Oct. 10 26-27 1848 “ Aurora Charles Jan. 12 1849 “ Phoenix Albemarle June 25-27 l r Chatham Mar. 19-22 1850 Ship Potomac j Albemarle Oct. 18-19 1854 “ Montreal Albemarle Jan. 19 1856 “ George & Susan Albemarle June 9-10 1858 Bark Stella Chatham June 10-12 ii Ship Fabius Charles Mar. 1 1859 Bark Ospray Chatham July 14-20 1860 Bark Montgomery Chatham July 8-13 “ Montgomery 1 Albemarle Feb. 9-10 1861 Chatham Mar. 15-19 “employed turpining” “getting turpin” “Terrapin very scars” “3 boats loded with turpin” “got a few Turrapin” “on shore for terphin” “got some turpin” slaid) “got but few tarapin” “on south head for terphin” “one Terrapin” “some Terrapin” “got a few Turrapin” “Several turpin” “3 boat loads of Terrapin” “after terrapin” “four boats go Turpin” “after Tarrapin” “few Turpin” “boats ashore after turtle” “employed Turpining” “the crew,after Terrapin” “some Terrapin” “crew after terepin” 1925] Townsend: The Galapagos Tortoises 83 1861 “ Alto Albemarle Feb. 13 “their boats after terrapin”12 1867 “ Osceola Albemarle Aug. 24-25 “after Turpin” Estimated Catch Arranged by Islands. Based on Recorded Catch 1. Albemarle 912 5. Hood 174 2. Charles 531 6. Abingdon 145 3. Chatham 472 7. Indefatigable 87 4. James 319 — Total 2640 The following table of the total catch shows that whaleships obtained a larger number of tortoises during the Thirties than in any subsequent decade. The catch from 1840 to 1849. was smaller, although more vessels visited the islands. The catch and the number of vessels both decreased during the two succeeding decades: Total Catch of Tortoises Arranged by Decades Decades Tortoises Visits by Vessels 1831-39 4853 56 1840-49 4379 75 1850-59 2334 35 1860-68 1447 23 Total tortoises 13,013 Total visits 189 Average per vessel 68 Arranging the total catch of tortoises by islands, we find Chatham Island, with 4,798, far in the lead in point of numbers taken. Albemarle is next, with 2,493; the position of other islands in respect of catch being in the following order: Charles, Hood, James, Abingdon, Indefatigable, Duncan, Barrington. The yield from the last four is small as compared with the others : Total Catch of Tortoises Arranged by Islands Chatham 4798 Abingdon 455 Albemarle 2493 Duncan 356 Charles 1775 Indefatigable 366 Hood 1698 Barrington 23 James 1049 Total 13,013 12 Reported in log of ship Edward Carey, February 13, 1861. 84 Fig. 26. A Galapagos tortoise ( Testudo vicina) from Albemarle Island, which has lived in the National Zoological Park at Wash- ington for twenty-seven years. Straight length of carapace, two feet. Photograph from the National Zoological Park. 1925] Townsend: The Galapagos Tortoises 85 Subsequent to the period under consideration — 1831 to 1868, the visits of whaleships had but little effect on the tortoises of the Galapagos Islands. The discovery of petroleum in 1859 affecting the price of whale oil, and the Civil War immediately after, with its heavy losses of vessels, marked the beginning of a decline from which the industry never recovered. Unpublished Notes on Galapagos Tortoises Contributed in 1924 Most of the later writers on the tortoises of the Galapagos have quoted extensively from the works of the navigators who saw them in the days of their greatest abundance. We need not here repeat these much- quoted accounts, but there are some works re- lative to the whaling industry that have not, so far as we know, been quoted in connection with the hunting of tortoises, which contain interesting information on this subject. It is possible to add also some unpublished accounts of tortoise hunting received from veteran whalemen and others while engaged in our search for logbooks of whaling vessels in 1924. Mrs. Johnson Whiting of West Tisbury, Martha’s Vineyard, contributes the following account of tortoise hunting, given her in August, 1924, by her neighbor, Mr. Russell Hancock, a vigorous man of eighty-two: “I went to the Galapagos Islands for terrapin. It was in 1865, on my first voyage to sea on the bark Samuel and Thomas of New Bedford, of which my brother-in-law, Captain Wm. Lewis, then of West Tisbury, was master. We landed on Albe- marle, uninhabited, of a volcanic mountainous construction, full of crevasses which made it hard and almost dangerous walking. We went ashore at daylight and walked until noon climbing up the mountain and crossing sort of bridges across the crevasses that former parties had made; then we left half our drinking water and went on capturing the terrapin which were feeding on the very sparse tufts of grass. We had hard work finding turtles small enough for two men to carry. When we did we rolled them over on their backs until we had enough, about 50 I think. Many huge turtles had carved on their backs the name of some whaling ship and a date of years before. I have often heard tell that a vessel went over from Panama with two donkeys and procured terrapin so large that two of them weighed 2,200 lbs. We had brought long iron 86 Zoologica: N. Y. Zoological Society [IV; 3 poles with us and we lashed the terrapins’ legs together, slung them on the poles and so carried them back to the ship, — one man on each end of the pole. We valued them very much for fresh meat. I don’t think anything ever tasted much better than fried terrapin liver. One thing we used to feed the turtles on board ship was bananas.” Mr. Lafayette Gifford of Westport Point, Massachusetts, makes the statement that his father, who was a whaling captain, “ visited the Galapagos in the ’fifties and saw there a renowned tortoise that was supposed to weigh over a thousand pounds. This tortoise had names carved all over its back. A British war ship finally took it away with the aid of tackles and a derrick. It was then but little longer than it was 150 years previously.” Perhaps the original dimensions were carved on its back. Mr. George A. Grant of Nantucket made a visit to the Gala- pagos in 1881 on the bark Alaska of New Bedford. At Perry Isth- mus on Albemarle Island he saw a tortoise of extraordinary size that was famous among the whalemen and was known as “Port Royal Tom.” There were dates and names carved on his back, the oldest date being 1791. Mr. Grant said that tortoises too heavy to lift were turned on their backs and dragged by means of ropes tied to their legs. An unpublished note on the Galapagos tortoises, written by Weston Howland, who visited the islands in a whaleship, “probably in 1830,” is contribtued by his daughter, Miss Rachel Howland, of Fairhaven, Massachusetts: “We proceeded to James Island, which is about sixty miles in circumference— and there anchored in a safe bay and sent a boat ashore. We found no inhabitants, but a pond of brackish water. Birds were plenty and so tame that a number of them were caught by hand, mostly teal, and the awkward flamingo with its beautiful plumage. In the morning a boat was fitted for an excursion in a bay or cove several miles away, for a load of ter- rapin. Arriving at the cove, a party consisting of sixteen officers and men, leaving we (two) boys to have a care of the boat, left for a cruise up the sharp volcanic side of the hills, to the feeding grounds of the hard shelled and long necked fellows. The principal food of the Gallipagos terrapin is the cactus tree and grasses which are found on the high lands and which get their moisture, from the near neighborhood of the clouds, as it seldom rains on the west 1925] Townsend : The Galapagos Tortoises 87 coast of South America so near the equator. The terrapin can and does, live six or more months without water, as he is provided with a vessel or bladder, in which he lays in a stock which he carries with him. This water is at any time as clear as amber and seems entirely pure and clean. The officers and men having returned from their mountain expedition, each with a large terrapin secured on his back, that would weigh seventy-five pounds or more, lunch and a good drink of water was served. They then returned to the feeding ground for another load, which they came back with ip time to load the boat and return to the ship before dark, with a supply of delicious food for a month or more.” The catch in this case appears to have been thirty-six tortoises, which, at seventy-five pounds each, weighed 2,700 pounds. Mr. Paul W. Ryder of New Bedford sends an account of tortoise hunting as related to him in December, 1924, by Captain Samuel Bumpus of Fairhaven, now in his eighty-third year: Captain Bumpus visited the Galapagos twice in 1860 in the ship Louisiana of New Bedford, John Kelly, master. Sixty tortoises were taken at Albemarle Island on the first visit and forty-eight on the second. He saw one tortoise of about 300 pounds lying in a moist spring hole which was too far up the mountain to be carried down. Most of the tortoises were secured about a mile inland. Some of those taken on board weighed as much as 300 pounds, but the average of most of them was from fifty to seventy-five pounds. While on board, the tortoises were fed on potato sprouts and peel- ings. At that time “turpin” as Captain Bumpus called them, were still plentiful. Fresh water was not found by the crew, which numbered thirty-two men. On the morning of the last day at Albe- marle the deck of the Louisiana was covered with volcanic ash. Captain Bumpus spoke with enthusiasm of tortoises as food, de- claring that the liver was the greatest of delicacies. The meat was usually prepared in the form of a stew called “ sea pie.” The fat made the finest of shortening. The following information was given by Captain Gilbert L. Smith of Vineyard Haven, in September 1924: Captain Smith was the master of the bark Northern Light of New Bedford on a voyage that took him to the vicinity of the Gala- pagos Islands about 1875. He never went ashore at the islands but was near enough to see them. He bought ten or twelve terrapin Fig. 27. A Galapagos tortoise ( Testudo vicina ) drinking. From the New York Zoological Park. 1925] Townsend: The Galapagos Tortoises 89 from a small Ecuadorian vessel that had been sealing at the islands. There were a few vessels from Ecuador that were engaged in sealing and terrapin hunting. The terrapin would be sold for various articles that the whalers might offer in exchange. Captain Smith said that terrapin were seldom fed anything when kept on board whaleships. Sometimes, however, they were given a few bananas. In his trading with the Ecuadorians, he bought also a barrel of terrapin oil, which he took home and dis- tributed among the housewives that he knew at Vineyard Haven. Captain Smith said that the terrapin on Charles Island were exterminated very early, and that Albemarle was the island where whalesmen of his day had the best luck. The terrapin were hard to locate, being usually high in the mountains. The smaller ones were carried on a man’s back by a strap arrangement, and the larger ones were carried on a pole, borne by two or more men. Captain Smith heard of one terrapin that the finders had to abandon. It was estimated to weigh about a thousand pounds, and the men were unable to get it down to the shore. It was found about four miles inland. Extracts Relative to Galapagos Tortoises from Works on Whaling The following passages relative to tortoises are from “ A nar- rative of the Sufferings and Adventures of Capt. Charles H. Bar- nard, in a Voyage Around the World during the years 1812 to 1816:” published in New York in 1829. The author’s first visit to the Galapagos, antedated that of Darwin by nearly twenty years. Captain Barnard, an American, was on board the British whaler Eliza when the latter visited the Galapagos Islands in company with another British whaler. They “came to at Charles’s Island,” . . . “Boats were dispatched from each ship on the 23d of June (1816) to the shore, to procure terrapin. They were so successful, that at night they returned loaded. . . . After a few days, the Indefatigable 13 (a British frigate there at the time) departed for England. The officers and crews of both ships had been actively engaged in pro- curing a full supply of terrapin.” Later, Captain Barnard was on 13 The vessel for which Indefatigable Island was named. 90 Zoologica: N. Y. Zoological Society [IV; 3 the ship Millwood (a merchant vessel) of New York, Captain Bailey, which called at Charles Island to procure tortoises: “At 4:00 A.M. of the 28th of October, we accompanied Mr. Cole and ten men, in the pinnace, to the black beach, about three miles distant, to procure terrapin: we arrived there at daylight, and proceeded to the spring, about two miles from the landing. We found a great many terrapin there. They were generally too large for a man to carry, and it was only by culling them that one could be obtained to convey down to the shore. While the men were gone to the boat, Mr. Cole and myself searched among the surrounding rocks and brambles for more terrapin, and by selecting the smallest, had procured one for each man on his return from the beach. “This spring of fresh water, the only one of living water on the island, is resorted to by the terrapin from the most distant parts of it, instinct being their pilot. They remain round the spring several days, occasionally drinking, until they have filled their five internal reservoirs, when having their twelve months’ stock on board, they return to their burrows. While we were here, there was a continual stir among them. Those that had obtained them stock were march- ing off, and others arriving to procure theirs. There was one re- markable for his size, as it was supposed he weighed six hundred pounds. Mr. Cole was desirous to get this mammoth on board, but to carry him to the pinnace was considered almost impracticable. I therefore instructed one of the boys how to manage and drive him, and calculated he would be able to reach the landing place by sunset; but he was one quarter of a mile distant from it, when we came up ; for his rogue of a driver, when he thought he was not observed, would get on his back, but the terrapin, not being well broken, would not proceed far without stopping. We turned him over, and lashed him to a tree to prevent his getting away, intending to terminate his land travels in the morning. On getting down to the beach, we found we had thirty-four fine terrapin there. On trial we perceived the boat would not carry them all at once; and accordingly five of them were left, four men remaining at the spring. We started for the ship, but the boat was so deep, and rowed so heavily, that we made slow headway, and it was ten o’clock before we got alongside. ... On the 29th we got out the long-boat before daylight, and when it was light, Mr. Cole and six men left for the Black Beach, to procure as many terrapin as they could. ... At 6:00 P.M. the long-boat not 1925] Townsend: The Galapagos Tortoises 91 appearing, I went in the pinnace, with a crew, to assist in getting her down, met and took her in tow, and got alongside about eight. Mr. Cole had forty-five terrapin in the boat, including the patriarch. Having now more than seventy on board, Captain Bailey considered that number sufficient.” Extracts from “A Narrative of the Life, Travels and Suf- ferings of Thomas W. Smith,” written by himself, and published in Boston in 1844. The following account concerns a part of his ex- periences on board a British whaling vessel, the ship Spring Grove of London, in the year 1821. “After this we proceeded to Woods’ [Hoods] Island, and came to anchor in a suitable harbor. Here we lay three days, during which time we collected 200 terrapins for the cruise and then pro- ceeded to Charles’ Island and from thence to Sea. . . . “Having watered our ship [coast of Peru] we sailed for the Gallipagos islands to take in a sufficient number of terrapins to last for the ensuing season on the coast. In two days we arrived at Chatham Island, where we took in 300 large terrapins. . . . “The terrapin resort to the low lands in the rainy season, drink- ing a sufficient quantity of water, at that time, to serve them during the dry season, which is six months. They then retreat to high ground, in consequence of which the labor of the ship’s crew, who go there to collect them, is great; as they have to pass through a thicket of bushes for a mile or two before they can fall in with any of them. “Individuals have strayed away in these thickets, in search of terrapins, and not being able to find their way out, have perished there for the want of water. My sufferings in this particular, as well as those of some of my ship-mates, were great; and we at times were under the extreme necessity of drinking the blood of the terra- pin, and even the water of the animal, with which they like the camel abundantly provide themselves for the season.” The following extracts, relative to tortoise hunting on the Galapagos Islands, are from “The Nimrod of the Sea” by William M. Davis, published in New York in 1874. This work is a composite account of the author’s whaling experiences. According to Star- buck’s History of the American Whale Fishery, Captain Davis was master of the ship Chelsea of New London, on two voyages to the Pacific Ocean — in 1827 and again in 1831. Chapter VIII begins after a landing had been made at Black Beach on Charles Island: 92 Fig. 28. Vegetation near Blackbeach, Charles Island, Galapagos. Photograph by U.S.S. Albatross Expedition 1891. 1925] Townsend: The Galapagos Tortoises 93 “We left two men to prepare camp, while the rest started for the back country to hunt terrapin Presently to my surprise, I saw our happy darkey ’Zekiel . . . sitting on the rear of an enormous terrapin about the size of a wheelbarrow, and much the shape of my mother’s forty-gallon apple-butter kettle . . . : Here was a 'baste’ that would weigh three hundred pounds at least. In the vicinity were numbers of others of more manageable size, and we selected two of perhaps fifty pounds weight. We tied the fore and hind legs of each so as to leave convenient loops through which to slip our arms, in- tending thus to carry our capture home, knapsack-fashion, on our backs. ... I have not a certain idea of the weight these creatures attain, but think I am within the mark in placing them at four or five hundred pounds. “The true way to carry a terrapin is to form a hand-barrow with deal clubs, or for the largest, of the steering oars, such a contrivance, manned by two or ten men, will bring down the capture with com- parative ease.” Penetrating into higher country the author con- tinues: “Great numbers of terrapin were about, some of them of immense size — very much larger than any seen on the shore plains here.” From Chapter IX: “we took the head off the largest terrapin we could find — one great enough to furnish a feast for a hundred men. — We were exceedingly thirsty, moreover, and had tried to satisfy our craving with the warm insipid juice obtained from the trunks of the giant cactuses, but in our capture, in our terrapin, we found the living spring of the wilderness. An ample supply of pure limpid water was discovered in the pearly sack placed at the base of the animals neck. There were some three gallons of water here, and, wonder of wonders, it was cool. The temperature of the animal is but 62°, but that of the country may reach 110° in the sun. . . . With one hundred and fifteen terrapin of all sizes secured, we then returned to the ship whose decks were covered with our sleeping captives and the cook’s galley steamed with a new and savory odor.” From Chapter XIX: “ A curious feature of the Galapagos is the novel post-office, established there by Commodore Porter, during the last war with England, while the Essex harbored in the island which bears the name of her worthy captain. He placed a large terrapin shell on a conspicuous point of Black-lava Rock. As round and white as a huge skull, it is a prominent landmark to vessels 94 Fig. 29. A Galapagos tortoise ( Testudo abingdonii ) from Abingdon Island. This specimen was among the first of these wonderful tortoises brought to the New York Zoological Park. 1925] Townsend: The Galapagos Tortoises 95 coasting among the islands. The enormous shell forms the roof of the letter-box, and it is the custom of ships to send a boat ashore and overhaul the mail for any letters that may have been left there for them, and to deposit any letters they may have directed to ships long out which may touch at the islands.” In “ Eighty Years Ashore and Afloat,” by E. C. Cornell, the following extract concerns an incident in the voyage of the ship Apollo . The account is probably true, as there was a ship Apollo of Edgartown, which sailed under Captain Daggett on a whaling voyage to the Pacific in 1816. The captain’s name, the vessel’s name and the year of sailing check with the book’s statements: “After many days reached Charles Island, where we fell in • with two Englishman whalers and a Nantucketer. We came to anchor close by them, and everything being secure went on shore after terrapin. Went far into the interior over to Black Beach, so called from its cinderry appearance. Trees called cabbage wood and prickly pears were scattered here and there; only one spring of water was found, and that on the extreme south end. We suc- ceeded in taking a good lot of terrapin, usually selecting those most convenient to carry on our backs, the usual way of transporting them. “Here we remained about one week, occupying ourselves daily in the same manner. Frequently it required some time to enable us to find the sized ones best suited to our ideas; they were all the way from as large as a silver dollar to the size of a Henry Clay cook stove. Some were so large that they could easily travel with four good-sized men on their backs. Their chief article of diet when on land is the cabbage-tree leaves, which are broken down by the force of the winds; but sometimes when no high winds lay their food on the ground for them, a large number will congregate, and with one accord gnaw into the bark of these trees, till, coming to the pith which is soft and tender, the tree falls before them. The trees grow to the size of half-barrel. I have often taken them from their work and pointed them in another direction, but if allowed they will return to complete their job, never leaving it until it is completed. Though they appear to enjoy eating as well as other animals, yet they will live and thrive on ship-board for months with nothing on which to subsist. “Three hundred were put on board our ship, stowed between / 96 Zoologica : N. Y. Zoological Society [IV; 3 decks or anywhere out of the way. They were a strange kind of birds; did not seem to care whether they stood on their head or heels. Their meat was most excellent; usually made it into stifles and soups. They were so fat that half a bucket full of grease could be taken from their upper shell when butchered. The fat was sometimes used to shorten those favorite “ duffs” previously alluded to. “The terrapin we had taken were stowed in different parts of the ship/some among the casks between decks, some on deck; it mattered little to us, and apparently less to them, what their ac- commodations were, so long as they kept out from under foot. With the food they afforded and that of the blackfish constantly on hand, we fared quite sumptously. Our cook used to parboil a sufficient quantity of terrapin over night for next morning’s break- fast, when not obliged to be in the boats.” There is a detailed account of tortoise hunting on the Galapagos in Captain Thomas Crapo’s “Strange but True” published in New Bedford in 1893, pp. 37-38. Captain Crapo visited Albemarle Island in 1858 in the bark Greyhound of Westport, Massachusetts, George G. Cathcart, Master; “On arriving there (Albemarle Island) we dropped anchor. After everything was put in shape, about two- thirds of the crew went ashore, taking with us boat sails to make tents of and water to drink and cook with, as fresh water cannot be found there. After fitting up our temporary camp we started for the mountains after turpin, which are very numerous, and are not found on any other islands. “Turpin are a species of turtle, the shell being in large checks like an alligators skin, and their flesh is unsurpassed as food for soups and stews: its equal cannot be found. The liver is far superior to any kind of meat I ever ate. It is as large as a beef critter’s (from a large one), and is many times superior to it in any way you choose to cook it. “ In order to get them we had to go high up in the mountains, as that seems to be their roaming ground. They are black in color and move very slow. We did not disturb the large ones, as we would have had to kill and cut them up and carry the pieces down on our backs, as many of them will weigh, I should think, nearly half of a ton. So w’e caught the smaller ones, none weighing over five or six hundred. “We went hunting them every day for a week, and as they are so Fig. 30. A Galapagos tortoise ( Testudo vicina) from Albemarle Island, which has lived twenty years in the New York Zoological Park. Weight when received 140 pounds. Present weight, 305 pounds. Photograph from the New York, Zoological Park. 97 98 Zoologica: N. Y. Zoological Society [IV; 3 clumsy and move so slow, made 'it an easy matter to capture them. We built a pen to put them in, and while on shore lived on them mostly, and used hard bread from the ship for soups and stews and other ways: the cook dished it out to us. The small ones we caught we carried down to camp on our shoulders, but we had to drag the larger ones. They are perfectly harmless and never known to bite. We caught about a hundred during the time. At the close of the week we took them aboard. Their weights would range from about five pounds to five hundred and over. We put them on deck and between decks, and let them crawl around as they chose. It was all of six months before they were all gone. I never knew one to eat or drink a drop while they were on board, and yet they looked as fat as a ball of butter when they were killed/ ' A record of tortoise hunting at the Galapagos from “ Whaling,” By Charles Boardman Howes, New York, 1924, p. 117, is as follows: Ship Sukey of Nantucket, John Macy, Master, at Charles Island, June 14, 1812: “I leave this port this Day with 250 Turpin.” In “ Wanderings and Adventures of Reuben Delano,” printed in 1846, the author having sailed on the ship Stanton of Fairhaven, in 1824, tells of a terrapin hunt he engaged in on Indefatigable Island, where 300 terrapin were taken. There is a brief reference to Galapagos tortoises in William Mariner’s “ Tonga,” published in Edinburg in 1827, which appears to have been overlooked by writers on this subject. Mariner ac- companied the ship Port au Prince , whaler and privateer, on a voyage to the Pacific in 1805. The record is as follows: “On Wednesday the 16th (October) made Chatham Island, one of the Galapagos. Whilst at this place, some turpin (land- tortoise) was procured from on shore.” There are some references to the tortoises of the Galapagos Islands in the journal of David G. Farragut,14 a young officer at- tached to the U.S.S. Essex in 1813: “At Charles Island we let some of the men go on shore daily to take a run. They amused them- selves by appointing one as a cook while the rest went in search of tortoises and water.” Among the captures made by the Essex were the British whale- ships Atlantic and Greenwich: “From these two vessels we secured water and tortoises euough to supply us for some time. On the 14 Life of Farragut, by his son Loyall Farragut, New York, 1879. 1925] Townsend: The Galapagos Tortoises 99 6th of June we saw a beautiful sight, a volcano in a state of eruption on the Island of Narboro.” Writing of the capture of the British whaleships Georgiana* and Policy, he says (p. 23): “In clearing their decks for action, they had thrown overboard several hundred Galapagos terrapins. The ap- pearance of these turtles in the water was very singular: they floated as light as corks, stretching their long necks as high as possible . . . many were picked up, which afforded us an abundant supply of fresh provisions for officers and crew. The meat, cooked in almost any manner, is delicious.” The Preservation of the Galapagos Tortoises The Executive Committee of the New York Zoological Society has signified its intention to procure if possible living specimens of Galapagos tortoises in the hope of averting the impending extinction of these animals. The edible value of the tortoises is sufficient reason for an attempt to locate a number in some southern locality where suitable climatic and food conditions would favor their in- crease. Their survival in the Galapagos where natives and wild dogs are equally destructive, seems to be quite hopeless. The last important collection of Galapagos tortoises was that made in 1905 under the direction of Mr. R. H. Beck for the Cali- fornia Academy of Sciences, when about 250 individuals were secured and preserved as museum specimens. More than one hundred of these were found in the high interior of Albemarle Island, where it is possible that survivors persist. This expedition spent nearly a year at the islands. The taking of such a number for scientific purposes appears to be justified in view of the heavy slaughter by Ecuadorian oil makers that had long been in progress on Albemarle, and was likely to be continued as long as there were profitable returns. While Galapagos tortoises have lived for many years in the New York Zoological Park, in the National Zoological Park at Washington, in England and elsewhere, they do not breed in northern climates. A fact to be noted in connection with their failure to reproduce is that the tortoises now living in captivity are nearly all males. Being hardy animals and easily cared for, a number of them 100 Fig. 31. A Galapagos tortoise ( Testudo vicina), in a walking attitude. This animal can easily carry a man weighing at least 150 pounds. It is ve'ry gentle and tractable, and follows the keepers like any domestic animal. From the New York Zoological Park. 1925] Townsend : The Galapagos Tortoises 101 should be transported to a carefully selected breeding base, where their protection would be assured. The present writer visited the Galapagos Islands in 1888 in the U. S. Fisheries Steamer Albatross, when eighteen tortoises were obtained from Duncan and Chatham islands; those from the latter having been taken there from Indefatigable Island. He revisited the Galapagos in the Albatross in 1891, when a single tortoise was obtained at Duncan Island, no search for tortoises being made on other islands. All of these reached Washington alive and in good condition but as a result of improper care did not survive the following winter. On board the Albatross they had the freedom of the deck and fed freely on such fruits and vegetables as were available. It is now known that they grow faster than was formerly supposed. According to Messrs. Daggett and Heller a twenty-nine pound speci- men taken to California in 1899 doubled its weight annually and in sixteen years attained a weight of 450 pounds. They are also long lived. There are authentic records of individuals that lived more than a century in captivity. It is known that the females lay numerous eggs. The species of giant tortoise ( Testudo elephantina) formerly abundant on Aldabra and other islands in the Indian Ocean, but later brought to the verge of extermination, is now increasing under Government protection. In 1906 Mr. M. J. Nicoll visited the Seychelles15 in the Indian Ocean, and described the giant tortoises now living there: “ In the grounds of Government House we saw a large number of Aldabra tortoises. Some of them were of large size and a great many were newly hatched. We were informed that they bred freely in confinement and that the young grew very quickly. These tortoises are used for food by the natives and on visiting the market we saw several tethered by the leg and exposed for sale. On all the islands and inhabited islets of this group, there were tortoise farms. In nearly all these farms the tortoises bear a number which is painted in white on the shell. Many farm-owners keep a record of all their stock, while at Government House a com- plete register is made with the dates of hatching and so forth. At the latter place we rode upon the largest tortoise. 15 Three Voyages of a Naturalist. Scribners, New York, 1908. 102 New York Zoological Park. 1925] Townsend: The Galapagos Tortoises 103 All these tortoises have been imported from Aldabra and there are now probably considerably more individuals on the Seychelles than on the former island.” Aldabra and the Seychelles lie north of Madagascar. There can be no doubt that the giant tortoises of the Galapagos are as well adapted to semi-domestication as the great tortoise of Aldabra. Having developed great size and high edible qualities on desert islands, the Galapagos tortoises might prove of great value among our food resources, especially in arid regions. With a steadily increasing population, the world’s food production must soon be increased. Three centuries of navigators feasted abundantly on the tortoises of the Galapagos. No other large land reptile ever figured so importantly in the food supply of mankind. The tortoise islands of the Indian Ocean, also visited by food seeking ships, could not apparently, have borne such numbers as the much larger islands of the Galapagos. According to the late Doctor Van Denburg16 who reported upon the numerous tortoises collected for the California Academy of Sciences in 1905-1906, the state of our knowledge respecting the numbers of tortoises remaining on the Galapagos Islands is_as follows: Island Species Status-1906 1. Abingdon Testudo abingdoni Rare 2. James “ darwini Rare 3. Jervis 66 wallacei Very rare 4. Duncan 66 ephippium Fairly abundant 5. Indefatigable 6 6 porteri Not rare 6. Barrington 66 7 Extinct 7. Chatham “ chathamensis Nearly extinct 8. Hood 6 6 hoodensis Very rare 9. Charles 6 6 elephantopus Extinct 10. Narborough 66 phantastica Very rare 11. Vilamil — Albemarle 6 6 guntheri Abundant 12. Iguana Cove — “ “ vicina Numerous 13. Tagus Cove — “ 6 6 microphyes Fairly numerous 14. Banks Bay — “ 6 6 becki Fairly numerous 15. Cowley Mt. — “ 66 Rare It is altogether improbable that the conditions of nearly twenty years ago obtain at the present time. The tortoises of the smaller 16 The Gigantic Land Tortoises of the Galapagos Archipelago. By John Van Denburg, Proc. Cal. Acad. Sci , 1914. 104 Zoologica: N. Y. Zoological Society [IV; 3 islands have doubtless suffered further depletion in numbers. Sur- vivors if sought for are most likely to be found in the large islands of Albemarle and Indefatigable, the interior regions of which are ex- ceptionally difficult to penetrate. A remarkable fact in connection with the giant tortoises is their amazing abundance at the time of the discovery of the islands. This continued for more than two centuries or until the first settle- ment in 1832, despite the inroads made upon them by food-seeking ships. All of the early navigators make mention of their abundance. Dampier who visited the Galapagos in 1684 says “It is incredible to report how numerous they are.” What a contribution could be made to the world’s food supply if the otherwise unimportant islands where, unknown to primitive man, the tortoises reached such an amazing development, could be cleared of the pests introduced by civilized man and the original conditions restored ! This is now unfortunately impossible on the Galapagos. The only remaining hope for the race is the establishment of survivors elsewhere. APPENDIX Logbook Records of Tortoises Taken from the Galapagos Islands by Certain Whaleships from 1831 to 1868 17 1831. Ship Isabella of New Bedford, Joseph Taber, Jr., Master. Hood Island 335 tortoises. Dec. If. — Harbor of Hood Island, anchored. Dec. 5 — “At 3 P. M. 2 boats went on shore after Terapin at 7 P. M. returned brought of about 30 at 4 A. M. 3 Boats went on Shore after Terapin at 7 P M Boats returned on Board Brought of about 100 Terapin Dec. 6 — at 4 A M 3 Boats went on Shore after Terapin at 5 P M the Boats came on board brought of about 90 Terapin Dec. 7 — At 5 A M 3 Boats went on Shore after Terapin at 4 P M Boats returned on Board Brought on Board 65 Terapin Dec. 8 — at 5 P. M. Boats Came on Board Brought of about 50 Terapin.” 1831. Ship Magnolia of New Bedford, Geo. B. Worth, Master. Charles Island. 155 tortoises. Dec. 11 — “At 4PM Came to Anchor at Charles Island. . . . Sent four Boats and Crews after Turpin and took about 110” Dec. 13—“. . . Sent three Boats after turpin and Returned with about 45” 17 Extracts from the logs are quoted verbatim. 1925] Townsend: The Galapagos Tortoises 105 1831. Ship Hesper of New Bedford, F. T. Brown, Master. Hoods Island 250 tortoises. Dec. 19— “came to anchor at Hoods Island sent 2 Boats ashore after turpine” Dec. 20 — “Employed in getting turpine” Dec. 23-24 — “These 2 days employed in getting turpine got about 250 alto- gether which cost us much trouble.” 1831. Ship Frances of New Bedford, Obed Alley, Master. Charles Island 179 tortoises. Nov. 12 — “ at dark lay our head yards aback to the Eastward of Charles Isle at daylight kepet off for the harbourd at ^ past 10 or there abouts come too in 7 fathoms of water” Nov. 13 — “two boats went a shore after terpin” Nov. 14 — “at night the boats come on board with 28 turpin” Nov. 15 — “the boats come on board with 44” Nov. 16 — “the boats come on board with 63” Nov. 17 — “we got 44 more terpin” 1832. Ship Abigail of New Bedford. Benjamin Clark, Master. Hood Island 50 tortoises. Abingdon Island 8 tortoises. April 1 — “5 A M Hoods Island distant about 10 miles sent 2 Boats Ashore for to get some Turpin” April 2 — “the Boats got 50 turpin ... all Hands employed Eating turpin” At sea. June 2— “all Hands employed about trying their strength by lifting an large turpin and looking for what we cant see that is sperm whales” June 6 — “tacking off and on Abington 7AM Captn Clark and second mate went on shore with two Boats to get some turpin” June 7 — “6 P M Boats came on board with 8 turpin” 1832-33. Ship Hector of New Bedford. John O. Morse, Master. Charles Island 226 tortoises. Dec. 27 — “kept her off for Charleses Island ... at 10 A. M. the starboard boat went on shore” Dec. 28— “at 5 P. M. the boat came on board brought some turpin” Dec. 29 — “at 11 P. M. came to anchor [Post Office Bay, Charles I.] at 3 A. M. called all hands to go after tarpin . . . the governer came on board” Dec. 30 — “at 7PM the boats came on board & brought 80 tarpin” Dec. 31 — “the boats went after Tarpin 14 miles from the ship” Jan. 1 — “at 6 P. M. the boats came on board & brought 90 tarpain ... at 2 A. M. the boats weant after turpin” Jan. 2 — “a sundown down the boats came alongside with 6 turpin Later the waist boat went on shore at the landing for 50 turpin that govner cent down the beech” 1833. Ship Hector of New Bedford. John O. Morse, Master. Albemarle Island 9 tortoises. 106 Zoologica: N. Y. Zoological Society [IV; 3 1833. Ship Hector ( cont .) March H — “standing off & onn from Albermar the Larboard & waist boats went on shore after Turpin wee found eight vary large ones” March 15 — “the boats went on shore after turpin could not find but one” 1833. Ship Pacific of New Bedford; Paul Chase, Master. Indefatigable Island 44 tortoises. Oct. 27 — “came to an anchor at Indefatagable Island sent 3 boats 21 men on shore after tirpen at dark retimed on board and never saw one” Oct. 28 — “the boats retimed on board with 8 tirpen ... at daylight got under way and beet up to the weather harbour” Oct. 29 — “sent 3 boats after tirpin” Oct. 30 — “the boats retimed with 27 tirpin” Oct. 31 — “the boats retimed with 9 tirpin” 1834. Ship Abigail of New Bedford. Benjamin Clark, Master. Porter’s Island 140 tortoises* May 15 — “steering for Porters Island [Indefatigable] 4PM Came to Anchor in Downes Bay in 7 fathoms” May 16 — “Boats came on Board with 21 large turpin” May 17 — “Boats came on Board with 31 turpin” May 19 — “Boats Came on Board with 40 live turpin” May 22 — “the Boats Returned with 40 live terpin ... we got here 140 Terepin and 10 Boats Load of wood” 1834. Ship Bengal of Salem, Russell, Master. Charles Island 100 tortoises. Mar. 22 — [Visited the settlement and] “had through the hospitality of one of the residents a good dinner of terrapin at sunset arrived on board the other boates got 50 terrapin larg an small.” Mar. 21 \ — “at 4 A. M. all the Boates whent a boute 12 miles to Swains landing on the N. W. point of the Island after anchoring the Boates we landed on the rocks and by the help of a rope we succeeded in getting up the precipice we found a plain with some large terrapin on of which & Backet to the Boate.” ... “we got 50 terrapin 5 of the large ones died on the rocks at 8 P. M. on board tired oute” 1834. Ship Moss of New Bedford. Shubael* Clark, Master. Chatham Island 8 tortoises. Charles Island 350 tortoises. Feb. 15 — “at 9 A. M. came to anchor at Charles’s island . . . sent all hands A shore after terrapin” Feb. 16 — “the boats came on board with 8 terrapin” Feb. 1 7 — “the boats came on board with 6 terrapin . . . took the anchor and stood off and onn sent A boat on shore we found the island to be, Chatham island by obs” Feb. 18 — “A boat on shore at sunset the boat came of with 8 terrapin” Feb. 21 — “at 4PM came to anchor in post-office bay at Charleses island . . . all hands A shore after terrapin” 1925] Townsend: The Galapagos Tortoises 107 1834. Ship Moss (cont.) Feb. 2 If — “150 terrapin on board.” [All hands getting terrapin from Feb. 21 to March 1] March 1 — “finished giting terrapin — got on board the No of 350” [from Mar. 2 to Mar. 13, the men went wooding] 1834. Ship Lover of Nantucket. John Cotton, Master. Woods [Hoods] Island 237 tortoises. Sept. 13 — “At 6 P. M. the Boats Returned With But 21 Tortoises.” ■ Sept. Ilf — “One Boat Came on Board With 60 Tortoises & the other 2 Boats Stoped all Night at Day Light the Boat Went on Shore Again” Sept. 15 — “At 5 P. M. the 3 Boats Came on Board With 30 Tortoises” Sept. 16 — “At 6 P. M. the Boats Came on Board with 50 Tortoises” Sept. 17 — “At 6 P. M. the Boats Returned With 35” Sept. 18 — “At 6 P. M. the Boats Returned for the Last Time With 41 Tor- toises” 1834. Ship Hector of New Bedford. John O. Morse, Master. James Island 23 + tortoises. April 12 — “standing of & onn from James Isle came to anchor at 8 A. M. two boats went after turpin” April 13 — “the boats came on board turpin was vary scarce we got 3” April 14 — “loard three boats & wint after turpin” April 15 — “the boats came on board caught twenty turpin Latter part the boats went after turpin” April 16 — “the boats came again on board did not do vary well” 1834. Bark Benezet of New Bedford. Chas. Pitman, Jr., Master. Charles Island 120 tortoises. Indefatigable Island 12 tortoises. Feb. 26 — “concluded to go in at Charles Island . . . at 9 A M came to anchor in 12 fathoms water” Feb. 27 — “one Boat after terpin Capt gone to town” Feb. 28 — “at 8PM Boats all on board with 50 terpin” March 1 — “at 8PM Boats abord with 30 tirpin” March 2 — “at 5PM came aboard with 40 tirpin” April 13 — “at 5 killed some tirpin” June If — “at 2AM came too Anchor at Portors Isle in 8 fathoms water at daylite 2 Boats went a tirpining at sunrise the 3rd Boat” June 5 — “got 8 tirpen” June 6 — “came aboard with 4 large tirpen” 1835. Ship Barclay of New Bedford, Henry Cottle, Master. Charles Island 50 tortoises. July Ilf — “anchored Charles Island Harbor” July 15— “at daylight sent 2 Boats for Turpin” July 16 — “at 6 the Boats returned with 20 Turpin” 108 Zoologicai N. Y. Zoological Society [IV; 3 1835. Ship Barclay ( cont .) July 17 — “2 boats a turpining, one Boat to Town for potatoes three of the men Deserted Caleb Halsted Alfred Overtwin Ronald Blanchard the boat returned without them at 7 oclock the Boats Returned with 30 turpin” July 18 — “employed in Wooding Caught the three runaways put the Ring- leader Caleb Halstead In Irons and Kept him Below the Gideon Basto [Barstow] Left and the Ships Washington and Baleaner of New Bedford Arrived” Note. — This vessel made her first voyage in 1795. 1835. Ship Hector of New Bedford. Thomas A. Norton, Master. James Island. 124 tortoises Albermarle Island 2 tortoises. June 3 — “steering in for Breakfast Isle at 2 P M let go the anchor in 18 fathoms of water in Compy with stanton of Fairhaven. . . . At 4 A M lowered 3 Boats and landed at James Isle to procure turrapin” June U — “At Breakfast Isle ... at 6 P M Boats Come off with 34 Turapin ... at Daylight 3 Boats went on shore for Turapin” June 5 — “at 6PM Boats came off having procured 26 Turapin” June 6 — “at 6PM Boat Come off with 40 Turupin” June 7 — “at 6PM the Boats Come off with 24 Turupin” Nov. 25 — “at Daylight kept off for South head [Albemarle] at 7 A M 2 Boats went on shore for turapin at south head lying off and on” Nov. 26 — :“at 2PM the Boats Came off having procured 2 turapin and some fresh fish” 1835. Ship George and Susan. D. E. Wight, Master. James Island 68 tortoises. Oct. 2)+— “at six the boats returned with ten terapen. Could not find them anywhere handy had a long ways to go for to get them but we got 68 all told grate and small” 1835. Bark Benezet of New Bedford. Charles Pitman, Jr., Master. Abing- don Island 12 tortoises. Jan. 1 — “at daylite kept of for Abingtons Island at 7 started 2 Boats for tirpen” Jan. 2 — “at 6 P M all of with 10 live and 2 dead ones” 1835. Bark Benezet of New Bedford. Charles Pitman, Jr., Master. Charles Island 40 + tortoises. April 13 — “at 5 a boat went in for Charles Island for tirpen” April 15 — “at ancher . . . the Boat got 40 tirpen” April 18 — “at 6 P M all abord with a few tirpen” 1835. Ship Lima of Nantucket. William Wyer, Master. James Island 35 + tortoises. Albemarle Island 67 tortoises. June 27 — “at noon came to anchor at James Hand in 12 fathoms water” June 28 — “went on shore 2 boats returned at 5 with some terrappin” 1925] Townsend: The Galapagos Tortoises 109 1835. Ship Lima ( cont .) June 29 — “returned with some terrappin” June 30 — “returned with 20 terrappin” July 1 — “returned with about 15 Terrappin” Nov. 21 — “ at 8 A M two boats wemt on shore at south head” [Albemarle] Nov. 22 — “at 3PM the boats returned with one dozen good Terrappin” Nov. 25 — “at 6 P M came to anchor In Elizabeths Bay . . . at 9 A M 2 boats went on shore after terrappin but saw none” Dec. 1 — “at 4 A M 2 boats went after terrappin at South head” Dec. 2 — “at 5PM came of with 25 terrappin” Dec. 3 — “at 4PM went on shore to be there early in the morning” Dec. ^ — “at 3PM the boats came off with 30 terrappin” Dec. 7 — “killed a terrappin weighing 250 lbs.” 1835. Ship Phoenix of Nantucket; Isaac B. Hussey, Master. Abingdon 10 tortoises. Hood Island 65 tortoises. James Island 7 boat loads tortoises. Feb. 6 — “Steering for Abingtons Island. . . . Sent 2 boats on Shore and and Cat [caught] Sum turping” Feb. 7 — “Got 10 turping” July 18 — “At 5 Came to Anchor At hoods Island . . . Employed in giting of turpin” July 19 — “At TPM got to the Ship with 45 turpin” July 20 — “At 6PM got Along side with 20 turpin it being hard to git them July 23 — “At 8 A. M. Came to Anchor At Jamess Island” July 25 — “Git on board with 2 boatload of turpin” July 26 — “Came on board with 2 boat lode of turpin” July 28 — “Came on board with 3 boat lode of turpin” 1836. Bark Pioneer of New Bedford. Reuben Russell, 2nd, Master . Porters Island. 2 plus many more tortoises. July 22 — “at 5 P M came too an anker at porters Island. . . . At 4 A M sent two boats for Terpin” July 23 — “at 4PM The boats returned with two tirpen” July 21+ — “at 6PM one Boat returned with terpin” July 25 — “at 8PM wone Boat returned with terpin” July 26— “at 7PM wone Boat returned with terpin” July 27— “at 6PM the Boats returned with terpin” Note. — This vessel is celebrated for having made the most profitable of all whaling voyages. In 1856 the value of her cargo was $151,060. 1836. Ship Eliza Adams of Fairhaven, John O. Morse, Master. Albemarle Island 23 tortoises. Aug. 21+ — “Off S head ... at 5 a.m. 3 boats went on shore for terpins” Aug. 25 — “got altogether 23 terpins” 110 Zoologica : N. Y. Zoological Society [IV; 3 1836. Bark Hesper of Fairhaven. Obed Fosdick, Master. James Island 13 plus several boat-loads of tortoises. Feb. 19 — “lying off and on James Island at 8 A M Come to Anchor in 8 fathoms water sent 2 Boats wooding one boat Turrapin” Feb. 20 — “1 Boat Turapining” Feb. 21 — “at 6PM got off 1 Boat load Turrapin. Last part 2 Boats Tur- rapining” Feb. 22 — “at 6PM the Boats returned to the Ship loded with Turrapin Last part 2 Boats Turrapin” Feb. 23 — “at 6PM the Boats returned to the Ship loaded with Turrapin Last part all hands employed Turrapining” Feb . 2k — “at 6PM the Boats returned to the Ship loded with Turrpin Last part 2 Boats Turrapining” Feb. 25 — “at 6PM the Boats returned to the Ship loded with Turrapin Last part 11 men Turrapining” Feb. 26 — “at 4PM the Boat returned to the Ship with 13 turrapin” 1836. Ship Lima of Nantucket; William Wyer, Master. Chatham Island 20 tortoises. James Island 118 tortoises. July 1 — “at 4PM came to anchor at Stephens Bay in 9 fathoms water, . . . went on shore 2 boats after terrappin” July 2 — “came off with about 20 terrapin” July 3 — “at 8AM came to Anchor at James Hand and went on shore after terrapin” July k — “Come off with about 18 terrapin” July 5 — -“employed getting terrappin 30” July 6 — “came on bord with 40” July 7 — “brougt on board about 30” 1837. Ship Abigail. Wm. Raynard, Master. Abingdon Island 142 tortoises. Jan. 21 — “Went ashore and got two terpen” Jan. 25 — “All hands employed in making belts to go after terpen at Abington island got ashore one oclock sixteen men got 30 terpen” Note. — Another log of this voyage continues the record as follows: Jan. 26 — “laying off and on 2 Boats on shore after Teripin at 7 P M returned with 53 Turin pin” Jan. 27 — “at 9PM the Boats returned from the shore with 40 Turpin” Jan. 28 — “the Boats on shore for Turipin at 9 P M they returned with 17 Turpin” 1837. Ship Eliza Adams of Fairhaven, John O. Morse, Master. James Island several tortoises. Charles Island 24 tortoises. Oct. 22 — “at 2 came to an anchor under Jamses Island and went on shore for turpin at night came of with a few turpin and wood ... at noon the Omega anchored here.” (See Omega, 1837, Chatham Id.) Oct. 23 — “a turpining and wooding” 1925] Townsend: The Galapagos Tortoises 111 1837. Ship Eliza Adams ( cont .) Oct. 21 \ — “imployed in wooding and backing turpin at night came of with a number of turpin” Oct. 27 — “at 2AM tuck our anchor for the harbor” Oct. 28— “two boats on Shore at noon came of with 3 Spanyards and 24 turpin the inhabitants of the Island” [The “harbor” mentioned is doubtless at Charles Island because of the reference to “inhabitants”.] 1837. Ship Lima of Nantucket; William Wyer, Master. James Island 224 tortoises. May 28 — “at 6PM James Hand bore West 20 miles dist. ... at 11 A M came to anchor in 10 fathoms” May 29 — “Employed getting terrappin got 17” May 30 — “got 26” May 31 — “got 17” June 1 — “got 46” June 2 — “got 53” June 4 — -“got 65” 1837. Ship Omega of Nantucket; Albert C. Gardner, Master. James Island ? tortoises. Chatham Island 240 tortoises. Oct. 23 — “came to Ancor on the N E side of James Island” Oct. 2 1> — “imploied turpining” Oct. 25 — “tuck the Ancor and but to sea” Oct. 28— ‘ “Came to Ancor in Chatham Island” Oct. 29 — “All hands on shore for turpin.” Nov. 5 — “Boats returned from turpin and maid up the number of 240” 1838. Ship Corinthian of New Bedford. Leonard Crowell, Master. Hood Island 136 tortoises. June 21 — “At 7AM came to Anchor at hood Island in 193^> fathoms. 3 Boats A shoar after Teripen” June 22 — “at 6PM the Boats came off got 36 Small Teripen” June 23 — “at 7PM the Boats came off with 22 Teripen” June 2J+ — “at 7PM the Boats came off with 33” June 25 — “at 6PM the Boats came off With 45” 1838. Ship Charles of New Bedford. Morselander, Master. Albemarle Island 8 tortoises. May 16 — “at 4AM went in at the South Head of Alber Marl Island with 2 boats after Terapin” May 17 — “at 9PM Came on board with 8 large Terapin” 1838. Ship George and Susan of New Bedford, H. C. Cushman, Master. Chatham Island 67 tortoises. 112 Zoologica : N. Y. Zoological Society [IV; 3 1838. Ship George and Susan ( cont .) Sept. 2 — “P. M. boats came on board with 27 larg tarapin” Sept. 3 — “fitting the boats to go after turapin at 3 P M Called all hands at 4PM lowred 3 boats Capt. Second and 3d Mates went after tarapin” Sept. 5 — “at 8 oclock P M 2 Boats came on board at 10 P M the other arived brought 40 turapin” 1838. Ship Phoenix of Nantucket; Isaac B. Hussey, Master. James Island 12 tortoises. May h—r' “At 7AM one Boat went on Shore to James Isle For turpin” May 5 — “At 5PM the Boat Return with 12 turpin” Sept. 25 — “At 3 P M A Boat went on Shore [at James I.] For turpen and stopt on Shore all night. . . . Sent another Boat on Shore for turpen” Sept. 26 — “At 3PM the boats Returned with a few Turpen” 1839. Ship George and Susan of New Bedford. H. C. Cushman, Master. Barrington Island 22 tortoises. Dec. 21 — “3 boats on shore at barington island at 7 P M came on board with 12 turpin” Dec. 22 — “of the E. end of Barington Island .... 3 boats on shore at 6 P M came on board with 10 tarpin” 1839. Ship Charles of New Bedford. Morselander, Master. Albemarle Island 20 tortoises. Feb. H — “Steering in for the South Head of Alber Marl at 3 P M went on Shore with one boat at 7 P M the boat came on board with some wood and 2 Terapin” Feb. 15 — “lying of and on at Albermarle at 6 P M the boats Come on board with 18 Terapin” 1839. Ship Robert Edwards of New Bedford. — Howland, Master. Hood Island 10- 12 tortoises. March 21+ — “at 7 too boats went on shoar for Terrapins on the South side of Hoods Island” March 25 — “at 4 the boats come off found it to ruged to land with safety 4 of us howeve got on shoar and got 10 or 12” 1839. Ship Robert Edwards of New Bedford. — Howland, Master. Albemarle Island 6-7 tortoises. March 26 — “at 7 the cap* went on shoar for Terapin. . . . South head of Albemarl N 12m” March 27 — “one boat on shoar at 4 she come of with 6 or 7 terapin” 1840. Ship Robert Edwards of New Bedford. Howland, Master. Chatham Island 59 tortoises. March 31 — “sent a boat for terapin at 4 came of with 4 or 5 wee anchored 1925] Townsend : The Galapagos Tortoises 113 1840. Ship Robert Edwards ( cont .) under the W point of Chatham Island in 9 fathoms water . . . two boats on shoar Turpining” April 1 — “the boats came of with 14 Terapin ... at 4 A M two boats went after Terapin” April 2 — “at 6 one boat came of with 29 terapin lost one boat on the rocks Left 5 men on the shoar and 12 terapin on the bech . . . at 4 A M went on shoar took of the boat and the terapin” 1840. Ship Rousseau of New Bedford, Luce, Master. Hoods Island 45 tortoises. Feb. 2— '‘saw Hoods Island ... at 4 o’clock P M came to anchor in 15 fathoms of water” Feb. 3 — “two Boats went after turpin” Feb. 1 !+ — “got 45 turpin” 1840. Ship Mariner of Nantucket. George Palmer, Master. Chatham Island 115 + tortoises. May 18 — “at 9 [A. M.] came too [at Chatham I.] with the larboard anchor in 15 fathoms water and sent the boat on shore with 2 gangs for terrapin . . . 1 at night the boats came off without much success” May 21— “At 5 A M 2 boats went to the east end of the island at 6 P M the came back with 45 terrapin” May 22 — “came back with 30 terrapin” May 21+ — “came back with 40 terrapin” 1841. Ship Elizabeth of New Bedford, H. F. Eastham, Master. Chatham Island. 102 tortoises. July 23 — “got 51 Terrapin at night all boats on board” July 21+ — “All Boats away, at night returned with 44 Terrapin . . . the Boats went and got 7 Large Terrapin” 1841. Ship Chili of New Bedford. D. B. Delano, Master. Crossman Island 16 tortoises. James Island 93 tortoises. Albemarle Island 10 tortoises. Sept. 19 — “laying off and on at Crossmans Isle [Albemarle] 4PM the boats came off bringing 16 terrapin” Sept 20 — “heady N W for James Isle 2PM sent in two boats for terrapin 4 came too in 10 fathoms 8PM boats came off with two terrapin . . . 234 A M sent off two boats to Albemarle and one here for terrapin” Sept. 21 — “3 P M boat came off with 7 . • • sent in one boat for terrapin” Sept. 22 — “4 P M boat came off with 8 . . . daylight sent in the boat 11 A M 2 boats came from Albemarle with 10” Sept. 23 — “ came off with 8 ” Sept. 21+ — “two boats came off with 26” Sept. 25 — “4 P M boat off with 20 6 P M two boats came with 22 have lost a man . . . daylight sent all hands ashore to look for the lost man” 114 Zoologica: N. Y. Zoological Society [IV; 3 184L Ship Chili ( cont .) Sept. 27 — “could not find him left bread and water and directions in a bottle if any one should ever find him” 1841. Ship Rousseau of New Bedford. John E. Brayton, Master. Albe- marle Island 12 tortoises. Oct. 23 — “South head bareing S E 3 Ships in sight at 7 A M the Capt Went on shore with 2 boats to look for Tarrapin” Oct. 2. 4 — “at 9AM the boats returned with 12 Terrapin” Note. — The oldest whaler. Built 1801 for Stephen Girard of Philadelphia. Broken up at New Bedford 1893. 1841. Ship Pocahontas of Holmes' Hole, Smith, Master. Albemarle Island 47 tortoises. May lb — “saw the island of Albemarl and run for it sent 2 boats on shore for tarpens” May 15 — “the boats came off from shoar with 7 tarpens . . . sent 3 boats on shore for turpens caught a seal” May 16 — “the boats came off with 23 turpens” May 17 — “sent 3 boats on shore to the south of head of albemarl for turpen” May 18 — “the boats on shore for turpen came off and left them on shore with 5 men . . . sent 2 boats on shore for the turpen and men and got them off 17 in number” 1841. Ship Hector of New Bedford. James Gray, Master. Albemarle Island 24 tortoises June 8 — off South Head, Cape Christopher, [Albemarle] “At dark the boats returned with some half dozen Terrapins” Nov. 1 — “landed two boats on South Head” [Albemarle] “spoke the Rodman” Nov. 2 — “at9PM the boats returned with 10 Terrapin. Rodman about the same. Stood off and on all night: at daylight 7 ships in sight” 1841. Ship James Munroe of Fairhaven. Benjamin Cushman, Master. Albemarle Island 64 + tortoises. Sept. 15 — “Ship laying off and on . . . sent 3 Boats on shore for Turpin South hade [Albemarle] Bairing E N E 10 miles” Sept. 17 — “at 9PM the Boats came on Board with Turpin” Sept. 18 — “at 9PM the Boats Came on Board with 9 Turpin” Sept. 19 — “at 6PM the Boats Came on Board with 8 Turpin” Sept. 29 — “at 7AM sent 3 Boats on Shore for Turpin at South Hade” Sept. 30 — “at 4PM the Boats Came on Board with 17 Turpin” Oct. 1 — “at 5PM the Boat Came on Board with 30 Turpin” 1842. Ship Eagle of Fairhaven. Samuel Perry, Master. Albemarle Island 36 tortoises. March 1 4 — “at 3AM made the Island of Arlbemarl south Head at 9 lowered all the boats and went on shore and at night came on board with 36 Terpine” 1925] Townsend: The Galapagos Tortoises 115 1842. Ship Rousseau of New Bedford. John E. Brayton, Master . Albe- marle Island. 10 tortoises. April 12 — “at 5AM went On shore at South head with 2 boats after Ter- repin” April 13 — “at 5PM returned with 10 large Terrapin” 1842. Ship Chili of New Bedford. D. B. Delano, Master. Chatham Island 118 tortoises. Aug. 15 — “1 P M spoke Ship Robt Edwards . . . Chathams Isle bearing N. dist 10 leagues . . . latter . . . running for the Anchorage . . . prepared to anchor but seeing a reef ahead on which we had like to run kept off again the R. E. anchored” Aug. 16 — “at 2PM came too in 17 fathoms found here Ships N Bedford. . . . Aurora. . . . boats terrapining. the ships laying at the lee anchorage sunset the boat came off with three terrapin” Aug. 17 — “sunset came off bringing 9 ... all hands ashore terrapining brot off 34 + 2 = 36” Aug. 18 — “brot off 35” Aug. 19 — “sunset came off bringing 8 . . . latter came off bringing 27” 1842. Ship Lion of Providence, R. I. Chas. F. Howland, Master. Albe- marle Island 5 tortoises. Mar. 19 — “Albemarl in sight” Mar. 21 — “the Nantucket and Awashonks in company ... at 6 went on shore after terapin” Mar. 22 — “in company with Awashonks a teraping. At 6 came off with two and three dead ones” 1842. Ship Robert Edwards of New Bedford. Burgess, Master. Chatham Island 107 tortoises. Aug. 15 — “steering in for Chathams Island at Meridian came to anchor 3^ mile from the shore in 10 fathoms” Aug. 16 — “the boats went in and got 7 Terapin ... all hand Terapining” Aug. 17 — “got off 9 Terapin . . . got off 21 Terapin” Aug. 18 — “got off 49 Terapin” Aug. 19 — “took off 21 Terapin” 1842. Ship Hector of New Bedford. James Gray, Master. Hoods Island 173 tortoises. Abingdon Island 8 tortoises. Aug. 25 — [Abingdon Island] “at dark the boats returned with some wood and eight Terrapin” Sept. 21+ — [Hood Island] “Sent three boats in for Terrapins. They returned at 7 p. m. having taken 25 rather small” (2 lost men) Sept. 25 — “went on shore, found the two lost men and 42 Terrapins” Sept. 26 — “obtained with hard labor 55 terrapins” Sept. 27 — “got 45 terrapins alive and 6 dressed” 116 Zoologica: N. Y. Zoological Society [IV; 3 1842. Ship Navigator of Nantucket; Elihu Fisher, Master. Chatham Island 30 tortoises. Hood Island 5 tortoises. May H — “at 2PM went a shore to Chatham Island the Island barring N N W at 5 come of with 5 tirapen . . . laying of and on” May 15 — “2 boats a tirapining at 5 Com of with 25” Oct. 20 — “at 10 Saw hoods Island at 11 went on Shore a turipening” Oct. 21 — “come of with 5 turpin” 1843. Bark Garland of New Bedford. Albert Scranton, Master. Hood Island 100 + tortoises. Sept. 30 — “At 4PM Came to Anchor at Hoods Island in 9 fathoms water . . . sent 2 boats to git Tarrapin” Oct. 1 — “the boats Came on Board with a few Tarrapin they were scarce” Oct. 2 — “Sent the Boats to git Tarrapin” Oct. 3 — “the Boats Came on Board with 34 Tarrapin . . . busey gitting Tarrapin” Oct. It — “the Boats Came on Board with 26 Tarrapin . . . busey gitting Tarrapin” Oct. 5 — “the Boats Came on Board with 40 Tarrapin” 1843. Ship Robert Edwards of New Bedford. Burgess, Master. Chatham Island 262 tortoises Dec. 19 — “At 4 Ship came to anchor in 17 fathoms of watter % of miles from the shore. Kicker Rock N. W. by N. 2^ Miles Chathams Island” Dec. 20 — “Bought 72 Terapin” (“ Terapining” until the 24th) Dec. 2 It — “At 9 P. M. Boats all came off and brought the remainder of their Terapin, makeing 190 in all, and 72 that the Capt. Bought” 1843. Ship Hector of New Bedford. George Manter, Master. Abington Island 67 tortoises. Mar. 1 — “10 live terrapins” “some more terrapins — 32” Mar. 2 — “obtained about 25 terrapins” 1844. Ship Callao of New Bedford. James A. Norton, Master. Hood Island 20 + tortoises. Albemarle Island 4 tortoises. Also “a load” of tor- toises. July 9 — “at 4PM came to anchor in the Roads of Hoods island in 15 fathoms of watt ... all hands went on shoare in persute of turpin Gut about 20” Note. — During the next four days, all hands went ashore “aturping” each day, but the number of tortoises caught is not recorded. Nov. 27 — “Loard 2 Boats to Goe on shore for Turpin at sonset came of with 4 So ends this day of South Head [Albemarle]” Nov. 31 — “Laying off & on South Head ... 2 [Dec. 1] boats on shore at 5PM came off with load” [of terrapin] 1844. Bark Equator of New Bedford. Thos. H. Mathews, Master. Chatham Island 24 tortoises. Albemarle Island 9 tortoises. 1925] Townsend: The Galapagos Tortoises 117 1844. Bark Equator ( cont .) April 2 4 — “Came to anchor at Chatham island in 7 fathoms of water Ship frances hear also too boats from each ship went after terpen thear is a Mr gerney lives hear geting out terpen oil his wife & 8 spanyards” April 25 — “at 7PM the boats retur with 8 terpen to each ship” April 26 — “went again with 8 men & got 8 terpen” May lb — “cloas in to Charlees islan at 6 P M loard the boats & towed in to 14 fathoms of water one mile N W of black beach” May 16 — “this day we got of 17 barrels of potatoss 30 pumpkins 15 bunches of benaners 50 pound of choacklet paid in trade & som money they com aboard as to a groacery stoar for small stoars at 6 P M got through & set them a shoar” May 19 — “ship being of South head went a shoar with 2 boats fro terpen landed at a crick to weather of goana cove [Iguana Cove, Albemarle] but found no terpen went to leward and landed our boats” May 21 — “got of clear through a verey hevey serf with 7 terpen the ship Daniel webster had two boats ashoar got 2 terpen” 1844. Ship Levi Starbuck of Nantucket; Jos. P. N ye, Master. Hood Island 14 tortoises. Chatham Island 130 tortoises. Jan. 31 — “at 2 P M we came to anchor in Woods [Hood] Island in 20 fathoms of watter, went on shore to look for tarraping at sun set return to the Ship with one tarraping” Feb. 1 — “the Boats on shore tarraping at night they return to the ship with 13 tarraping” Feb. 2 — “at daylight we got under way from Woods [Hood] Islands and went to Chatham Island, and at sun set we came to anchor in 22 fathoms of watter” Feb. 10 — “at 5 P. M. the bots came for the last time, we have got 130 tar- raping large & smal” 1844. Ship Charles of New Bedford. Gardner, Master. Chatham Island 100 tortoises. Feb. 9 — “at 4PM made the Land the East End of Chatham Island bearing W by S Dist 30 miles . . . when within 8 or 10 miles of the Land dispatch’d 2 boats in quest of tarapin” Feb. 10 — “at 7PM the boats return’d with tarapin” Feb. 11 — “at 5PM the boats return’d with tarapin” Feb. 12 — “at 5PM the boat return’d Onboard with Tarapin . . . beat up to the anchorage at daylight anchored in 10 fathoms water” [On Feb. 13, 14, 15, the men were “employ’d at getting Tarapin”] Feb. 16 — “got under way . . . after Obtaining 100 tarapin” 1845. Bark Equator of New Bedford. Thos. H. Mathews, Master. Albe- marle Island 69 tortoises. May 1 — “. . . ship heading to the E . . . south head [Albemarle] bearing N W distant 10 milds at 6 P M tacked ship & hauled up the courses” 118 Zoologica: N. Y . Zoological Society [IV; 3 1845. Bark Equator ( cont ) May 2 — “at 8AM went in with labor & waist boats for terpen & got 11 large ones” May 3 — “went in with two boats to goana cove [Iguana Cove, Albemarle] in company with ship James Allen & got terpen to both ships” May 17 — “at 8AM being close in to cape Roas [Cape Rose, Albemarle] about 3 miles to the W we went in with two boats for terpen & found nise boat cove & got 27 terpen” Aug. 20 — “sent a boat in for terpen on the third hill from south head [Albe- marle] but got non” Aug. 21 — “being up in the head of weather bay sent two boats in for wod at 4PM got of 8 boat load” Sept. 17 — “at 7AM went in with two boats fror terpen at Albemar abreast of brattle island at 7 P M got aboard with 23 terpen” 1845. Bark Alfred Tyler of Edgartown, Luce, Master. Indefatigable Island 45 tortoises. Abingdon Island 7 tortoises. James Island 20 tortoises. May 5 — “running down for Porters Ileand” [Indefatigable] May 6 — “at 2 P.M. came to under the lee of the Ileand” May 7 — “sent one boats crew after terphin” May 8 — “the boat returned with 12 terphin” May 9 — “the boat returned with 20 terphin” May 10 — “at 7 P. M. the boat returned with 13 terphin” Sept. 7 — “laying of and on at Jameses Ileand at 4 P M the Boat Returned with 8 terphin” Oct. 2 — “run in to Abbington and sent 2 boats for terphin” Oct. 3 — “at 5PM the boats returned with 7 dead terphin” Oct. 11 — “sent two boats with 13 men to Jameses Ileand for terphin and suckceded in getting 12” 1846. Bark Equator of New Bedford. Thos. H. Mathews, Master. Albe- marle Island 150 tortoises. Chatham Island 14 tortoises. Feb. I,. — “at 8AM went in with two boat & got 12 large terpen goana cove bearing N” [the vessel was “off South head, Albemarle”] Feb. 5 — “captain & went in with two boats & sent off 11 verey large terpen captain & 4 men stayed ashoar goana cove bearing N E” April 11,. — “off point Essex” [Albemarle] April 16 — “all this day employed beating up weather bay in company with 3 other sail for terpen” April 17 — “went in & got one live terpen & two dedons” [dead ones] April 18 — “cam to anchor in weather bay tagers cove [Tagus Cove, Albe- marle] bearing W N W distant 8 milds in 25 fathoms of water one mild from shoar” April 20 — “got off 14 terpen” April 21 — “got off 80 terpen” April 22— “took the anchor & towed out in companey with Roussau & Aurora got 26 good terpen” 1925] Townsend: The Galapagos Tortoises 119 1846. Bark Equator ( cont ) Oct. 10 — “at 7AM went into fresh water bay ]Chatham Island] & got 6 cask of water & 5 terpen” Nov. 20 — “being off freshwater bay took a raft of 9 cask ashoar & filled them & got 9 terpen” Dec. 7— “being of Essex point [Albemarle] at 11 A. M. went in with two boats for terpen & got 4” 1846. Whaleship — No name found. Chatham Island 190 tortoises. July 20 — “anchored at Chatham Island in 10 fathoms water and all hands after Turpin” July 27 — “went to sea 190 turpin on Board” 1846. Ship Aurora of Nantucket; Frederick W. Coffin, Master. Albemarle Island (at least 2 tortoises). Hood Island 7 tortoises. April 16 — “standing in to the Bay [Elizabeth Bay, Albemarle] . . . Lat. 00”45 S” April 17 — “trying to get up into the Bay going in pursuit of Turrapin in company with bark’s Equator, Franklin & ship Rousseau” April 19 — “came to an anchor in 30 fathoms of water” April 20 — “all hands ashore after Turpin” April 21 — “all hands ashore after Turrapin” Sept. 1 — “off South Head . . . went ashore with one boat & got off the meat of 2 large Turrapin” Oct. 21 — “came to an anchor at Hoods Isld in 12 fathoms of water” Oct. 22 — “One watch ashore got 7 Turrapin” 1846. Ship Minerva of New Bedford. J. S. Macomber, Master. Chatham Island. 120 tortoises. Nov. 2 — “At 10 A M Come to anchor in fifteen fathoms water of Chatham Island” Nov. 7 — “finish Turepning got One hundred and twenty” 1847. Ship Coral of New Bedford. Humphrey W. Seabury, Master. Albe- marle Island 1 tortoise. Abingdon Island 1 tortoise. May 28 — “steering for the S W part of Albamarle at 6 P M Cape Rose bore W by N dist 10 miles . . . Ship off & on” May 29 — “at 10 all on board met with poor success got one Turpin Teripin” Nov. 2 If — “At daylight off the S W part of Abington. Went on shore with 2 boats & 20 men for terrapins” Nov. 25 — “At 4PM boats returned from the shore. Landed at 3 different places on the S side of the island & only found 1 terrapin — By all appearances they had retreated into the mountains as everything appeared to be dried up with the sun. Caught plenty of fish with which the shores abound & killed 1 fur seal” 120 Zoologica: N. Y. Zoological Society [IV; 3 1847. Ship Susan of Nantucket; Charles B. Ray, Master . Crossman Island 30 tortoises. June 10 — “lying off and on the Island of Abington, boats on shore after terrapin” June 11 — “at 5 P. M. boats came off with terrapin and fish” June 17 — “came to anchor att Portors Island in 7^ fathoms water” June 18 — “Went with 3 boats to Crossmans Island [off Albemarle] after terrapin” June 19 — “At Crossmans Island” June 20 — “Came on board with 30 terrapin . . . took our anchor and run over to James Island and 3 boats went on shore for terrapin” June 21— “at 4 P. M. came on board with the boats” 1847. Bark Alfred Tyler of Edgartown, Luce, Master. Abingdon Island 3 tortoises. May 2 If — “Saw Abbington bearing W • • . at 5 A M loured two boats and went on shore for terphin and fish” May 25 — “at 4PM the boats returned with 3 terphin and plenty of fish” 1847. Ship Charles Frederick of New Bedford. H. P. Barnes, Master. Hood Island 67 tortoises. July 30 — “came to anchor at hoods island at daylight sent two boats on shore for turpin” July 31 — “at dark the boats came off with 27 turpin at daylight sent on shore again” Aug. 1 — “at dark the boats came off with 40 turpin” 1847. Ship Aurora of Nantucket; Frederick W. Coffin, Master. Chatham Island 100 tortoises. Mar. 27 — “came to an anchor at Chatham Isd.” Mar. 28 — “a part of the crew ashore after Turrapin got 30” Mar. 29 — “got off 36” Mar. 31 — “got under way & went to sea 100 Turrapin on board” Oct. 26 — “the Capt ashore on Charles Isld got a few Turrapin” 1847. Ship Congaree of New Bedford. Aaron C. Cushman, Master. Chatham Island 4 + tortoises. July 1 — “ship lying off and on at Wreck bay [Chatham] at 2 P M the Capt came off with a few Terapin” July 6 — “at 3AM hove too off Kickor rock [Chatham] at 8. 3 boats started for Terrapin. Saw 3 ships at anchor at Terrapin Road” July 7 — “at 8PM the boats came on board with 44 Terrapin” 1847. Ship Elizabeth of New Bedford. M. Baker, Master. Chatham Island 100 tortoises. Aug. 21f—[ at anchor off Chatham Id.] “3 boats went in a tearpaning” Aug. 29 — -[still at Chatham] “took on board 100 Terrapin” 1925] Townsend: The Galapagos Tortoises 121 1848. Ship Susan of Nantucket; Charles B. Ray, Master. Abingdon Island 23 tortoises. Albemarle Island 186 tortoises. June 6 — “lying off and on the Island of Abingdon, boats after terrapin” June 7 — “at 4 P. M. boats came on board clean” June 8 — “lying off and on Abingdon at 4 P. M. boats came on board one terrapin” July 5 — “off Abingdon’s Island . . . boats shore after terrapin” July 6 — “lying off and on Abingdon at sunset boats came off with 8 large terrapn” July 9 — “boats came off with 14 large terrapin” Sept. 12 — “off south head 2 boats on shore after terrapin at sunset returned with 25” Sept. 14— “lying off and [on] South head boats on shore at sunset returned with 150 terrapin” Oct. 19 — “Off Albemarle 2 boats on shore after terrapin” Oct. 20 — “came off with 11 terrapin” 1848. Ship Corinthian of New Bedford. Armington, Master. Chatham Island 54 tortoises. June 5 — “at 8 A. M. three boats went on shore on the S. E. part of Chatham Island for turapin. . . . June 6 — “at 9PM the boats returned from the shore with 14 turapen” Oct. 12 — “at 10 A. M. took on board 40 large turapin lying at Anchor at Chatham Island Stephens bay” 1848. Ship Roman of New Bedford. Sanford Wilbur, Master. Duncan Island 50 tortoises. Indefatigable Island 36 tortoises. June 6 — “At 7AM Came to anchor the E side of Albemar i a bay duncans island bearing E by S went off with 6 boats for terpen but found non terpen gon back in the mountain” June 8 — “At 4 A M 3 boat from the Margrate Scot with three of ours went over to duncans island” June 9 — “at 9PM returned with about 50 terpen” June 10 — “stood across to Porters island [Indefatigable] & came to anchor in conway bay in 7 fathom of water” June 11— “the captain returned from the setlement reports the potatos full of woms got 30 terpen” June 15 — “At 6AM got under way & stood down the Jameses island chanel got 36 terpen 14 bunches of benanars” 1848. Ship Congaree of New Bedford. Aaron C. Cushman, Master. Abing- don Island 10 tortoises. Chatham Island 70 tortoises. July 6 — “at 11AM lowerd 3 boats and went on shore at Abingdon” July 7 — “at 7PM the boats came off with 7 Terapin 3 live ones and 4 dead ones . . . at 9 A M Went on shore with 3 Boats” July 8 — “at sunset the Boats came off with 3 Terapin one alive” 122 Zoologica : N. Y. Zoological Society [IV; 3 1848. Ship Congaree ( coni .) Sept. 2 h — “at 8AM went on shore at wreck bay [Chatham! and took off 70 Terapin” 1848. Ship Coral of New Bedford. Humphrey W. Searury, Master. Cha- tham Island 200 tortoises. Feb. 27 — “At 11 came to anchor on the S side of Chatham island at fresh water bay in 28 fathoms water — }/& mile from the shore. At 12 the Hope came to anchor near by. We have the cascade, a stream of water which can be seen running from the bank at the dist. of 4 miles bearing N. N. E.” [Feb. 28, to March 2 were spent getting terrapin and water] March 3 — “Weighed anchor made sail & steered out to the S in company with the ship Hope. . . . We are 12 miles from our anchorage with 200 terrapin on board most of them large which we obtained very easy as we found them near the shore & but a short dist. from the ship. We also filled up our empty casks with water & could have obtained any amount had we wished. As we found it good watering at the foot of a deep ravine that makes down from the top of the island. A stream running therein which forms a pond back of the beach which is in a fine cave at the foot of the ravine. A ship in my opinion may anchor within % mile from this place in about 20 fathoms water with perfect safety — from the months of December to April ships in coming to anchor should always keep well to the E. as there is a strong current setting to the W. although we found tides near the shore but the current sets but a very short time to the E.” 1849. Ship Susan of Nantucket; Charles B. Ray, Master. Albemarle Island 2 tortoises. Aug. 15 — “off South head . . . sent one boat ashore on south head after a fresh mess of terrapin” Aug. 16 — “bot returned from shore with 2 terrapin” 1849. Whaleship — no name found. Albemarle Island 63 tortoises. July 17— “went on shore with two boats found a plenty of terepin about two or 2 miles from the landing. We Succeeded in backing down 8 noble fellows at nine in the evg.” July 18 — “Went on shore early in the morn with three boats we made out to back down a bout twenty terepin about 200 cwt (some of them) each, reached the ship about eight in the eve.” July 1 9 — “ Started early after terepin to a new place With two boats. Cooper & four others to the old place after two terepin that was left on the road & then made sail for the other place — when we had reached we found that they had all all gone to the mountain & left us to haul the boat up ... we all returned at dark with a bout twenty terepin” July 20 — “started as usual for the shore at day light & brought down fifteen terepin two that would weigh from 200 to 250 each” 123 1925] Townsend: The Galapagos Tortoises 1849. Ship Congaree of New Bedford. Aaron C. Cushman, Master. Cha- tham Island. 130 tortoises. July 25 — “went into Stephens bay came to an anchor at 6 P M in 17 fathoms of water” July 26 — “took off 98 Terrapin” July 27 — “3 Boats on shore after Terrapin at 4 P M one boat came off with 8 Terrapin” July 28 — “At 10 A M the other boat came off with 7 Terrapin . . . finished getting off 130 Terrapin” 1849. Ship Kingston of Fairhaven, Leonard Luscomb, Master. Abingdon Island. 6 tortoises. Nov. 5 — “ At 7 A M went on Shoare with two boats after Terpin. . . . Laying off and on at Abbington” Nov. 6 — “At 6PM returned to the Ship with six Terpin” 1849. Brig Vesta, Osander Mayhew, Master, sailed from Edgartown April 10, 1849 bound to California (not a whaling voyage). Hood Island 1 tor- toise. Abingdon Island 5 tortoises. Oct. 19 — “at daylight saw the Island called Hoods Island run down hove to at 10 A M sent 2 boats on shore Looked all day for turpin at night came on board . . . with only one turpin that weighed 41 lbs.” Oct. 21 — “at 8AM lowered two Boats and went on shore at Abingdon Island Cruised all day and took 5 Turpin.” 1850. Ship Susan of Nantucket; Charles B. Ray, Master. Chatham Island 156 tortoises. June 23 — “At 4AM made Chathams Island. ... 12 M came to an anchor in 19 fathoms water off N. E. point” June 21 \ — “went on shore with 3 boats at sunset returned with 14 terrapin” June 30 — “at 4PM boats came off with the last terrapin making all together 156 terrapin” 1850. Ship Peruvian of Nantucket; George B. Folger, Master. Duncan Island 131 tortoises. Sept. 17 — “at 2PM ran in and came to an anchor in Grand Harbour Porters Isl in 6 fathoms water . . . at 4 P M 3 boats went across to an island about 10 miles dist [Duncan] for terrapin” Sept. 19 — “at 6PM the boats returned with 48 terrapin ... at 4 A M 3 boats started on another cruise across to the island for terrapin” Sept. 22 — “at 3PM the boats returned from the other island with 83 ter- rapin” 1850. Ship Martha of Fairhaven; Island 110 tortoises Skinner, Master. Chatham 124 Zoologica : N. Y. Zoological Society [IV; 3 1850. Ship Martha ( coni .) May 7 — “heading in for Chatham Is’d. . . . came to an anchor in 9 fathoms water the Catawba & Empire at anchor Turpapining” May 16 — “got under way . . . with 110 Turpen” 1851. Ship Pocahontas of Holmes Hole, J. Dias, Jr., Master. Chatham Island 90 tortoises. Aug. 20 — “sent two boats ashore (for terapin) with provisions and water for 3 days” Aug. 21+ — “one boat came off to bring a few turpin” Aug. 25 — “at 1 a boat came off with more Turpin — a small sloop from San Francisco after a cargo of Turpin. I should think she would carry as many as 20 good large ones” Aug. 26— “one boat came off with a load of Turpin, at 3 the other boats came off in each a load of Turpin making in all about 90 and I think pretty good ones” 1852. Ship Congaree of New Bedford. Martin Malloy, Master. Abingdon Island 5 tortoises. Aug. 15 — “at 7AM went on shore at Abington for Terrapin” Aug. 16 — “At 9PM the boats came on board with 5 Terrapin” 1852. Bark Eugenia of New Bedford. Wm. Wood, Master. Albemarle Island 2 tortoises. Chatham Island 107 tortoises. June 2 — “at 9AM went on shore with two Boats at South head [Albemarle] and got two small Terrapin. ***** Aug. 10 — “At 9.30 A M steered for S Bay [Chatham] at noon Came to an Ancor” Aug. 11 — “Took on Board 107 Terrapin” 1853. Ship George and Susan of New Bedford, J. S. Jenckes, Master. Abing- don Island. 3 tortoises. Sept. 21+—' “2 Boats returned from the Shore with the meet of 3 Terrapin and one nice one” 1853. Bark Henry H. Crapo of South Dartmouth. Spooner Jenkins, Master. Barrington Island 1 tortoise. April 12 — “Afternoon went on shore to Barington island With one boat for turpin and gut one” Note. — The log mentions the names of six other whaling vessels in sight of the Henry H. Crapo at that time. 1853. Ship Congaree of New Bedford. Martin Malloy, Master. Chatham Island 315 tortoises. Hood Island 7 tortoises. Jan. 3 — “run off for the East end of Chatham” 1925] Townsend: The Galapagos Tortoises 125 1853. Ship Congaree ( cont ) Jan. 4 — “At 4PM came too an anchor in 12 fathoms in Hobbs Bay . . . the hands after terrapin” Jan. 5 to Jan. 11 — “After terrapin” Jan. 11 — “took off 175 Terrapin” ***** Sept. 26 — “At 3PM Came to an anchor in Gardners Bay, Hoods Isle” Note. — The next few days were spent coopering oil and stowing Oct. 2 — “Took on board 7 terrapin” Dec. 13 — “at 8AM came to an anchor in 14 fathom of water in Hobbs Bay” [Chatham] Note. — No entry in the log-book between Dec. 13th and 18th Dec. 18 — “finished getting 140 Terrapin” 1853. Bark Peru of Nantucket; Charles E. Starbuck, Master. Albemarle Island 150 tortoises. July 8 — “boats off after turpin . . . the place where we are is on the east side of Albemarle abrest of Cowleys Island” [Two other ships in company, the Sea Queen and the Clifford Wayne.] July 9 — “one off the boats has been onshore abrest of the ship got 5 terapin have not heard from the other boats, they all stop ashore nights Latter part boats returned with 9 terrapin” July 18 — “our terrapin cruise is at an end. we obtained about 150” 1853. Ship Martha of Fairhaven. Meader, Master. Chatham Island 13 tortoises. July 29 — “At one oclock P. M. sent two boats on Shore at Chatham island for terpin” July 30 — “at two P M Came On Board with four turpin. . . . Came to an anchor at Chatham Island in fourteen fathoms water and veared out fifty fathoms cable . . . lowered three boats and went on Shore to see if we Could not get Some turpin” Note. — July 31 to Aug. 2, boats were after tortoises. Aug. 1+. — “the Boats returned with 9 terpin” 1854. Bark Eugenia of New Bedford. William Cattle, Master. Abingdon Island 3 tortoises. May 30 — “At 3PM sent Two Boats on shore at Abington Island, Contrary to Orders they remained on Shore all night” May 31 — “At 1PM one Boat returned with fish, At 10 P M the other one returned with Two Terrapin that they Butchered” July 4 — “At 7.30 A M sent Two Boats on shore at Abington Island, after Terrapin” July 5— “At 1PM the Boats returned with one Terrapin” 1854. Bark Superior of New Bedford. Charles L. Norton, Master. Albe- marle Island 1 tortoise. 126 Zoological N. Y. Zoological Society [IV; 3 1854 Bark Superior ( cont .) Feb. 19 — “Steering in for South head [Albemarle] at 1 oc P. M. Scent 2 boats on Shore for turbin got one” 1854. Ship Potomac of Nantucket; Enoch Ackley, Master. Chatham Island 43 + tortoises. June 12- — “steering N along the E side of Chatam Isl. ... at daylight steered off N. for the N. E. point of the Isl sent in two boat for turpin” June H — “got a few” Aug. 25 — “under the lea of Chatam Isl” Aug. 26 — “At anchor in Terrapin Rroads at 4 P. M. took off 10” Aug. 27 — “took on board 16” Aug. 28— “got off 9” Aug. 29 — “got on board 8” 1855. Ship George & Susan of New Bedford. J. S. Jenckes, Master. Cha- tham Island. 152 tortoises. June 29 — “At 6PM Boats returned Bring 6 Tarrapine” June 30 — “at 6 P. M. Boats returned Bringing 5 Turrapine” July 1— “at 7PM Boats returned Bringing 6 dead ones & 2 live ones” July 2 — “at 10 Boats returned Bringing 19 Tarrapin” July .4 — “At 10 A. M. Boats returned Bring 25 Tarrapine leaving one man behind who got lost one boat in search of the lost man” July 5 — “At 5 P. M. the two Boats returned Bringing 65 Tarrapin and at 7 the other Boat returned Bringing the lost man and one Tarrapin.” July 11 — “at 3 Boat went on shore at Stephans Bay. Chatham Island, to finish trading for Terrapin got off 23 more” Aug. 30 — Off Charles Island, “spoke a Brig from the Coast wanted to sell Tarrapin” 1855. Bark Cornelia of New Bedford, Reuben W. Crapo, Master. Chatham Island 28 tortoises. Albemarle Island 14 tortoises. Oct. 21 — “at daylight . . . running in for Chatham Island at 8 A M 2 boats went on Shore for Turpins Ship laying off and on” Oct. 22 — “at Sundown boats Came on board with 18 Turpins and one Turtle” Oct. 23 — “at 5 P M.Came to Ancor on the North Side of Chatam Island in 9 Farth of Water . . . got 5 Turpins” Oct. 2i \ — “at Sundown boats Came on board with 5 Turpins” Oct. 26 — “at 9 A M 2 boats went on Shore at South head [Albemarle] for Turpine Ship laying off and on” Oct. 27 — “at dark boats Came on board with 14 Turpins” 1855. Ship Mary Ann of Fairhaven. Thomas Dallman, Master. Chatham Island 4 tortoises. Duncan Island 17 tortoises. Oct. 2 — “kept away North for Chatham island at 10 A M the mate & second 1925] Townsend: The Galapagos Tortoises 127 1855. Ship Mary Ann ( cont .) mate went ashore for turpin & stoped untill 8 PM, found seven but did suc- ceed in getting off but four” Oct. 3 — “at 8AM the first, second & third mates with their boats went ashore & stoped untill 3PM but could not find any turpin” Oct. I \ — “at noon the first, second & third mates went ashore at Duncan’s island & stoped untill 5 PM. they got 17 small terpin” 1855. Bark Superior of New Bedford. Charles L. Norton, Master. Albe- marle Island 11 tortoises. May 4 — “lying of & on South head [Albemarle] to Boats of on shore After turpin At Sunset returned to the ship and Brought 11 turpin” 1856. Bark Benj. Cummings of Dartmouth. S. Jenkins, Master. Chatham Island. 310 tortoises. July 22 — “Came to anchor at Hobbs Bay Chatham Island for Terrapin 23 men Terapining Got a few” July 23 — “These days Terpining [24 to 31] 150 live Terpin” Nov. 25 [same voyage] — “ Came to Anchor at Chatham Island for Terpin all hands ashore” Nov. 26 to Dec. 4 — “all hands came on board with about 160 Turpin” 1857. Bark Bevis of New Bedford. David G. Peirce, Master. Albemarle Island 13 tortoises. Feb. 5 — “At 7 A. M. the Capt. took a boat and pulled in to Iguanno Cove [Albemarle] after terapin” Feb. 6 — “at 43^ P. M. Capt returned with 6 terapin” March 4 — “Off and on at Iguanna Cove, at 7 A. M. sent two boats in after terapin” March 5 — “At 8 P. M. the boats returned with one large and six small terapin” 1858. Bark Morning Star. H. D. Norton, Master. Albemarle Island 24 tortoises. July 27 Elizabeth Bay — “three boates went ashore after Tirpin today found none” July 28 — “Three boates went ashore after tirpin found two or three” Aug. 5 — “at 5 A M all Started up [mountain] again today we got down about 22” 1859. Bark Montgomery of New Bedford. R. N. Crapo, Master. Chatham Island 78 tortoises. Albemarle Island 7 tortoises May 4 — “at 9AM two Boats went in to South head [Albemarle] after Turerpin” May 5 — “at 7PM the Boats came off with Seven Turerpin” July H — “at 3PM Saw Chatham Ilseland ... at daylight Steered in for 128 Zoologica: N. Y. Zoological Society [IV; 3 . 1859. Bark Montgomery ( cont .) the Anchorage at 9 A M came to Anchor in twelve fathoms in company with the Ospra three Boats from each bound off after Tererpin” July 15 — “two Boats came down to the Barks with 14 Tererpin” July 16 — “the two Boats went up to the other Boats with Water for the rest and after Tererpin” July 17 & 18 — “after Tererpin” July 19 — “at daylight the Boats started from their landing for the ships with 51 Turerpin” July 20 — “got 13 Turerpin” 1859. Ship Lancer of New Bedford. 0. Fisher, Master. Chatham Island 70 tortoises. May 21+ — “Steering S W for Terpin road Chatham Is.” May 25 — “all hands on shore for Terpin at 4 P M took twenty on board.” May 26— “at 5PM returned with 8. this day found the Terpin farther off or about 6 miles from the beach found it very hard to back them” May 27 — “at 4PM returned with 14” May 28 — “at 5PM returned with 17 large ones and 3 small ones, makeing in all 65 live ones and 5 that were to large we killed and brought down there meat . . . one man shipped on board he had been there 10 mts all Alone at 7 A M sent A boat for his Clothing” & 1860. Bark Ohio of New Bedford. David Baker, Master. Albemarle Island 81 tortoises. July 6 — “ankered again at Cowlys Inlet” [Albemarle] Note. — From July 7 to 9 — “all hands on shore geting terpin” July 10 — “All hands employed geting terpin hav got 81 on board” 1860. Bark Ospray of New Bedford. J. E. Stanton, Master. Albemarle Island 122 tortoises. July 2 — “Bound to Perrys Ismuss [Albemarle] for wood in company with Bark Ohio and Vigilant” Note. — From the third to the eleventh, the crew was employed getting wood and tortoises. July 12 — “all hands came on Board with 122 Terpin” 1860. Bark Atkins Adams of Fairhaven. William Wilson, Master. Albe- marle Island 14 tortoises Ayg. 1+ — “at 8PM came to an anchor in weather bay [Albemarle] in 25 fathoms water one eighth of a mile from the shore ... at 7 A M three boats started in pursuit of Tarrepin” Aug. 5 — “at 7PM the boats returned without success, found only one Tarrepin in the mountains” Aug. 7 — “at 5PM Tarrepin cruisers returned with three Tarrepin” Aug. 8 — “at 6PM Tarrepin cruisers all down from the mountains but four, 1925] Townsend : The Galapagos Tortoises 129 1860. Bark Atkins Adams ( coni .) got ten Tarrepins, found them plenty about fifteen miles from the landing ... at daylight went ashore and took on board two of the absentees, at TAM started in pursuit of Tarrepin at another landing at 10 A M went on shore and took onboard another of the absentees.” 1860. Ship Edward Carey of Nantucket; Francis M. Gardner, Master. Albemarle Island 56 + tortoises. Mar . 19 — “came to anchor at Albemarle in 18 fathoms of water brattle Hand bearing E by South Sent three boats on shore to look for terrapin” Mar. 21 — “at sunset two boats came off with eleven terrapin the other boat remained waiting for two men which lost their way” Mar. 22 — “at dark the boats came off with a load of wood and twelve ter- rapin ... at 3 A. M. got under way and [went] further to the Westward say about five miles, four boats after terrapin at different beaches” Mar. 23 — “at 5P.M. two boats came off with 26 terrapin & left some penned up on Shore found terrapin quite plentiful the other two boats got 4” Mar. 21+ — “boats all on shore all hands seeking terrapin at 8 P. M. boats all got off heavy surff on shore left three terrapin on shore experienced very strong westerly current . . . fresh breeze at daylight sent two boats in after the above said terrapin.” 1861. Bark Stella of New Bedford. Frederick Hussey, Master. Albemarle Island. 6 tortoises. June 25 — “at 7 A. M. 2 boats went ashore on S. head [Albemarle] & got 2 large Terapin” Dec 18 — “at 9 A. M. two boats went in to the S. head [Albemarle] affter terrapin . . . laying off & on” Dec. 19 — “at 4 P. M. the boats came on board & brought off 4 scutteled Terrapins” 1861. Bark Ospray of New Bedford. J. E. Stanton, Master. Albemarle Island 41 tortoises. April 30 — “the Bark working up to Elizabeth Bay [Albemarle] at 5 p m came to Anchor Sent a Boat to finde a place to get wood” May 2 — “Latter part Sent 2 Boats in company with 2 from the Levy Star- buck to get Tarapan” Note. — For the next two days, it is written “much the Same” May 5 — “took the Anchor & ran to the North 12. m. & came to Anchor again all hands after terapin” “Monday May 6th to Saturday May 11th Employed bringing terapin out of the mountains & geting them on board 41 larg ons” 1861. Bark Morning Star. H. D. Norton, Master. Chatham Island 188 tortoises. Note. — Boats were ashore daily after terrapin from June 27 until July 9. 130 Zoologica: N. Y. Zoological Society [IV; 3 1861. Bark Morning Star ( cont ) July 10 — “all three Boats came on Board each one brought 20 Turpin” July 11 — “Brought the rest of the Turpin which made 188 alltold” 1861. Ship Roscoe of New Bedford. G. H. Macomber, Master. Chatham Island 50 tortoises. Nov. 30 — “At 10 A. M. the boats came on board brought 50 turpin” Note. — As the boats were ashore daily “after turpin” from Nov. 23 to 30, the above number may or may not have been the total. 1861. Ship Arnolda of New Bedford. J. A. Crowell, Master. Chatham Island. 42 tortoises. June ^ — “at 10 A M came to anchor at Chatham Island” Note. — From June 5 to 10 there were “boats on shore for terapin” daily. June 11 — “at 7AM took the anchor and went to sea ... we bring out 42 live terrapins” 1861. Bark Atkins Adams of Fairhaven. William Wilson, Master. Chatham Island 105 tortoises. July 5 — “Employed in geting straps ready to fetch tearpin with” July 18 — “headed in for N E point Hobs Bay Chatham island . . . anchored in 10 fathom of water” July IP— “Employed in tearpining” Note.— From July 19 to 31 “all hands Employed in tearpining” Aug. 1— “all hands Employed in receiveing tearpings on Board, we have on Board 105” 1861. Bark Ohio of New Bedford. David Baker, Master. Chatham Island 50 tortoises. March 10 — “at 2PM ankered at Hobes Bay” [Chatham] [Another vessel in company] March 15 — “From the last date up to the presant hav been at anker in Hobes Bay hav had three Boats crews on shore geting Terpin hav got fifty on board” 1862. Bark Stella of New Bedford. Frederick Hussey, Master. Abingdon Island. 4 tortoises. June Ilf — “off & on at Abingdon two boats crews went on shore after Ter- rapin” June 15 — “at 4 P. M. the boats came on board & brought two live terrapin & two scutled ones” 1862. Ship Edward Carey of Nantucket; Francis M. Gardner, Master. Albemarle Island 95 tortoises. Nov. 10 — “Ship heading in for the weather bay [Albemarle] . . . 11 P. M. came to anchor in Thirteen fathoms of water and close to the Clara Bell, Hector and Luisianna” 1925] Townsend : The Galapagos Tortoises 131 1862. Ship Edward Carey ( cont .) [During the next few days the crew was employed cutting wood.] Nov. Ik — “send two boats with seventeen of the hands after terrapin” Nov. 17 — “at 8 P. M. one boat returned with five Terrapin and intirely ex- hoisted” Nov. 19 — “working up for Iguana Cove where each ship have sent one boat with nine men apeace after terrapin lowered another boat and send up to the place” Nov. 20 — “returned with eleven terrapin and reported seventy more on the beach” Nov. 21 — “at 3 P. M. got all the boats off all hands and about 90 Terrapins in all” 1862. Ship Roscoe of New Bedford. G. H. Macomber, Master . Albemarle Island 63 tortoises. July 20 — Eighteen men on shore after terapin and got eight. July 22 — two boats came on board with terapin. July 23— the terapin gang came on board at 6 P.M. with fifteen terapin. July 2k — the terapin gang came on board at 6 P.M. with 17 terapin. July 25 — boats came on board at six P. M. with seven terapin. July 26 — boats came on board at 6 P. M. with six terapin. 1863. Ship Edward Carey of Nantucket; Francis M. Gardner, Master. Chatham Island 1 tortoise. Duncan Island 208 tortoises. Dec. 23 — “at sunset saw Chatam Island . . . daylight kept away for the Island at 10 A M sent in 2 Boats for Terripen” Dec. 2k — “dropt ancor in 19 faths dist mile from the shore at 11 P M the Boats came off bringing 1 Terripan” Dec. 26 — “stood over to Porters Island at 5 P M dropped ancor in 13 fathoms mile from shore . . . at 4 A M lowered 3 Boats with 21 men & stood over to Duncan Island for Terripan” Dec. 28 — “at 10 A M a small schooner Anchored close to us the Capt went on board bought of them 78 Terripan. . . . Also rec’d a Boat load of Terripan from the Islands” Dec. 30 — “at sunset rec’d the Boats from the Island with a cargo of Terripan” Dec. 31— “at sunset the [boats] came again with a Cargo” Jan. 1 — “at 4AM the Boats went back to the Islands at sunset all hands came on board having been gone a week & got 130 Terripan” 1867. Bark Osceola, 2nd, of New Bedford, John M. Shaw, Master. Abingdon Island. 1 tortoise. June 10 — “at 11 A M raised Abingdon steering towards it” June il — “went ashore got a mess of fish and one Turpin” 1868. Ship Roscoe of New Bedford. Geo. H. Macomber, Master. Cowley Island [Albemarle] 5 tortoises. 132 Zoologica: N. Y. Zoological Society [IV; 3 1868. Ship Roscoe ( cont .) Nov. 20 — “at 12 M anchored at Cowleys Islet in 6 fathom W” Nov. 21 — “one boats crew ashore cutting wood” Nov. 23 — “ 17 men ashore looking for Terrapin at sundown all came off brought 5 small Terrapin . . . employed cuting wood” Record of Vessels Seeking or Obtaining Tortoises, but Numbers Taken not Recorded 1833. Ship Loper of Nantucket, John Cotton, Master. Abingdon Aug. 7-8 At anchor at Abington 1834. Ship, Bengal of Salem. Russell, Master. Albemarle Island. Tues. Aprill 1 “went for terrapin to the south head” April 3 “not a drop of water where we shall go next or what we shall do is known to god only.” 1834. Ship, L. C. Richmond of New Bedford, John Tucker, Master. (Stone Fleet — see ship Potomac p. 134) Charles Island. July 16 — “the boat wint On Shore at the Island” 1834. Ship Ohio of Nantucket, Charles W. Coffin, Master. Feb. 25 — “at 9 A.M. anchored in Jameses island harbour in 16 fathoms water three boats went after turpin” Feb. 26 to Mar. 1 — “employd giting of turpin” 1835. Bark Pioneer of New Bedford. Reuben Russell, 2nd, Master. May 7 — “At 10 P.M. came too an anker at Charles Island” May 8 — “At 4 A.M. two boats went after terpin” May 9 to May 1\ \ — “the boats returned with terpin” every day. 1835. Ship Lima of Nantucket; William Wyer, Master. June 23 — “at 3PM came to anchor at Chatham Hand in 8 fathoms water went on shore two boats returned at 7 a few small terrappin” June 21+ — “at 5PM the boats returned with some terrappin” June 25 — “at 4PM the boats returned with a few terrapin” 1836. Ship Ohio of Nantucket, Charles W. Coffin, Master. April 12 — “At 5 P.M. anchored in blackbeach bay [Charles I] in 13 fathoms of water . . . two boats after turpin” April 13 — “two boats after turpin” April 11+ — “two boats after turpin & giting of potatoes” April 15 — “three boats after turpin” 1836. Bark Pioneer of New Bedford, Reuben Russell, 2nd, Master. July 16 — “At 9 A.M. came too an anker at Charles Island. At 10 A.M. sent one boat for terpin and two boats for wood” 1925] Townsend: The Galapagos Tortoises 133 1836. Bark Pioneer ( cont .) July 17 — “At 2 P.M. the boats came off with wood. At 6 P.M. the boat returned with Turpin” 1837. Ship Elizabeth of Salem, Isaac G. Hedge, Master. May 17 — “A.M. Stood in toward Albemarle & boats over for fish & Terapin” 1837. Ship Eliza Adams of Fairhaven, John 0. Morse, Master. May 7 — “at 3PM came to an anchor at Porters island” [Indefatigable] May 8 — “imployed in geting turpin” May 9 — “imployed in geting turpin . . . went over to James island for turpin” 1838. Ship Omega of Nantucket; Albert C. Gardner, Master. June 26 — “standing in for James’s Island at 3 P.M. came to anchor in 12 fathoms . . . sent 3 boats in after Terrapin” [To July 1 the crew was busy hunting for terrapin, but the numbers found are not recorded.] July 2 — “at 10 A. M. Took the Anchor and steered W. N. W. Empd. stow- ing the Anchors & Terrapin” 1842. Ship Ocean of Nantucket; Elijah Parker, Master. Jan. 25 — “came to anchor to Poters island in 7 fathoms of water . . . three boats went on Crosmans isl for turpin” Jan. 26 — “employed turpining” Jan. 27 — “employed turpining . . . took the anchor and stood to sea” July 26 — “Came to anchor to Chatham island in 13 fathoms of water the Henry of Nantucket lying there” [From July 27 to August 3, the crew was employed “getting turpin,” but the results are not recorded.] 1842. Ship George Washington of New Bedford. Taylor, Master. Nov. 1 — Hoods Ih “At 5 P.M. the boats returned found Terrapin very scars ” 1842. Ship James Munroe of Fairhaven, Benjamin Cushman, Master. July 2 — “At 6 A.M. 3 boats went on shore for Turpin at noon, Albamaral S Bore E By S dis 5 miles” July 3 — “At 4 P.M. the Boats came on board Loaded with Turpin” 1843. Ship Phebe of Nantucket. Samuel W. Harris, Master. Albemarle Island. Jan. 12 — “got a few Turrapin” 1845. Bark Alfred Tyler of Edgartown, Luce, Master. Albemarle — ? Nov. 11 — “at 7AM spoke the President run in to South Head” [Albemarle] “together and the two Captains went on shore for terphin” 134 Zoologica: N. Y. Zoological Society [IV; 3 1845. Whaleship — (No name found). Aug. 18 — “came to anchor at James Island after turpin and wood” Aug. 19 — “got some turpin” Aug. 20 — “got some turpin” Aug. 21 — “turpining” Aug. 22 — “turpining and wooding” Aug. 23 — “turpining and wooding” 1847. Bark Persia of New Bedford, Richmond Manchester, Master. Jan. 6 — “the first of this day Ship heading in to hoods island harbour at 2.30 P.M. came to anchor in fathoms water coral bottom at 5 A.M. went on shoar for tarapin found but verry few” Jan. 7 — “all hands imployed after tarapin . . . got but few tarapin” 1847. Bark Alfred Tyler of Edgartown, Luce, Master. Albemarle Apr. 2^ — “sent two boats ashore on south head for terphin” [Albemarle]. Apr. 25 — “at 5PM the boats returned with small luck” 1847. Ship Congaree of New Bedford, Aaron C. Cushman, Master. July 10 — “at 2 P.M. lowered 2 Boats and went on shore at Abington 6 miles off, at 9 P.M. got back with part of a Terrapin and some Earth to cure the scurvy” Oct. 26 — “At 7 A.M. went on shore with one boat at Black Beach” [Charles I] Oct. 27 — “At 5 P.M. boat came off with some hogs Terapin and fowls” 1848. Ship Aurora of Nantucket; Frederick W. Coffin, Master. Jan. 12 — “off and on at Charles Island got a few Turrapin” 1849. Ship Phoenix of Nantucket; Perry Winslow, Master. June 25 — “Steering for South Head” June 26~‘ “Steering for the land At 2 P M went on shore At dark came on board with several terepin. ... At daylight went on Shore with 3 Boats after turpin” June 27 — “At Sunset the Boats came of to the Ship with Several turpin” 1850. Ship Potomac of Nantucket; Charles Grant, Master. Mar. 19 — “at 2PM Anchored off Wreck Bay. Chatham. ... 3 Boats went around on the weather side after Terrapin” Mar. 20 — “1 Boat returned with Terrapin” Mar. 22 — “at 6AM [boats] started for Stephens Bay at 11 returned with 3 Boat Loads of Terrapin” Oct. 18 — “at 10 A M 2 Boats on Shore after Terrapin . . . Off S. Head” [Albemarle] Oct. 19 — “at 6 Boats came off” Note. — This vessel was one of the “Stone Fleet” of 40 whaleships loaded with stone and sunk in southern harbors during the Civil War to prevent blockade running. 1925] Townsend: The Galapagos Tortoises 135 1854. Ship Montreal of New Bedford, S. L. Gray, Master. Jan. 19 — “ship standing in for south head at 7 A .M. lourd four Boats to go shore Turpin. . . . Latter part ... at 11 A.M. boats com on” 1856. Ship George & Susan of New Bedford, J. S. Jenckes, Master. June 9 — “off south Head of Albemarle Isl. at 5 P.M. lowered down two Boats went in shore after Tarrapin” June 10 — “At 4 P.M. Boats returned to the ship” 1858. Bark Stella of New Bedford, R. W. Hathaway, Master. June 10 — “Send in two boats for Turpin and so ended off & on Chatham Island” June 11 — “two Boats shore on Chatham Island looking for Turpin at 5 P.M. returned to the Ship with none” June 12 — “2 boats shore for Turpin at 8 P.M. returned with few” 1858. Ship Fabius of New Bedford, Geo. A. Smith, Master. March 1 — Off Charles Island “the boats a shore after turtle” 1859. Bark Ospray of New Bedford, J. E. Stanton, Master. July 14— “At 9 A.M. Anchored in Hobbs Bay [Chatham I.] 8 fathoms watter 45 chain” From the 15th through the 20th “employed Turpining” every day. The number caught is not given. 1860. Bark Montgomery of New Bedford, R. N. Crapo, Master. July 8 — “came to an Anchor in Hobses Bay [Chatham I.} at 11 o’clock” July 9 — “one Boats crew a fishin the rest of the Ships Company a terupaning” July 10, 11, 12 & 13, the crew went after terrapin but no numbers are given. 1861. Bark Montgomery of New Bedford, R. N. Crapo, Master. Feb. 9 — “sent in two boats in to Guano Cove after Terapin” Feb. 10 — “the Boats returned to the ship with some Terapin” . March 15 — “Latter at 9 o’clock came to an Anchor in Hobses Bay [Chatham I.] “seven fathoms watter in company with the Bark Columbas” March 16 — “sent three Boats from each ship around on the East side of the Island after Terapin about 4 O’clock the boats landed and hailed up for the night Latter sent the crews after terepin found a few they ware very scarse” March 17 — “the men returned to the Boats had seen few Terapin latter returned to the Ship” March 18 — “Latter all but 4 men went on shore to look for Terapin” March 19 — “At Sundown returned to the ship” 1867. Bark Osceola, 2nd, of New Bedford, John M. Shaw, Master. Albe- marle Aug. 21+ — [anchored at Albemarle, the crew wooding] The Cap gone after Turpin got back at 11 P M” Aug. 25 — “nearly all hands ashore fishing catching turtle pipies and guanoes” ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY y VOLUME IV. NUMBER 4 NOTES ON FISHES FROM THREE PANAMA LOCALITIES GATUN SPILLWAY, RIO TAPIA AND CALEDONIA BAY By C. M. Breder, Jr. New York Aquarium PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK September 18, 1925 Nftu flnrk Zoologtrariwirtti General Office: 101 Park Avenue, New York City G&fltora President, Madison Grant; Vice-Presidents, Frank K. Sturgis; and Henry D. Whiton; Chairman, Executive Committee, Madison Grant; Treasurer, Cornelius R. Agnew. Utaarlftrf JUattagmi (ElaaH of 192fi Henry Fairfield Osborn, Lisfenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor (Ela00 of 19 27 Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field QUa00 of 192H Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, Anthony R. Kuser, Mortimer L. Schiff, Frederic C. Walcott, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler, George Gordon Battle, Bayard Dommick William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant Director ; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds ; George S. Huntington, Prosector ; H. C. Raven, Associate Prosector; Elwin R. Sanborn, Asst, to Sec’y. and Editor. £Mtm*ial (Enmmittrr Madison Grant, Chairman; William T. Hornaday William Beebe Charles H. Townsend Elwin R. Sanborn, Sec’y- Zoolog ica Vol. IV. No. // NOTES ON FISHES FROM THREE PANAMA LOCALITIES: GATUN SPILLWAY, RIO TAPIA AND CALEDONIA BAY. By C. M. Bredek, Jr.1 New York Aquarium (Figs. 33-38 incl.) Introduction The notes comprising this paper are based on three small col- lections of fishes which were made while attached to the Marsh- Darien Expedition in 1924 as the representative of the American Museum of Natural History. They were acquired quite incidentally to the main ichthyological work of the trip which was the surveying of the fish fauna of the Rio Chucunaque drainage and is to be re- ported elsewhere. As these collections have no especial bearing on that work, they are recorded here separately and with it complete the list of fishes taken on the trip, excepting those purchased in the markets simply as museum specimens. Each collection is treated as a unit, since they have no particular connection with each other, and should be considered as independent, being issued together for purposes of convenience. As with all other papers based on this expedition I am of course primarily indebted to Mr. R. 0. Marsh whose financial and other aid made the work possible. Regarding the section on the fishes of Caledonia Bay I am indebted to Mr. J. T. Nichols of the American Museum and to Mr. L. L. Mowbray of the New York Aquarium for assistance in diagnosing some of the smaller examples of the more difficult species. All measurements refer to the standard lengths of the specimens, without caudal. The material is deposited in the American Museum. Fishes from Gatun Spillway, Canal Zone While equipping at Colon preparatory to our plunge into the jungles of Darien the opportunity was taken to make a collection at Gatun Spillway (Fig. 33.). The kindness and aid of Mr. 1 Illustrations from drawings and photographs by the author.* 137 138 Zoologica: N. Y. Zoological Society [IV: 4 Fig. 33. Sketch map showing the location of the collecting sites at Gatun Spillway, Tapia, on the Rio Tapia, and Caledonia on Caledonia Bay. William Markham, founder of the Panama Canal Tarpon Club, greatly facilitated the gathering of numerous small fishes at the foot of the spillway and the obtaining of information concerning many of the fishes that reach here only at a large size and of ones not collected during my short stay at the well appointed club house. Mr. Markham remarked, nevertheless, that the present collection was unusually varied and he is in a position to know as he habitually takes his bait fishes at this point. The collection was made on January 26 and 27. Most of the small fishes were taken in a small minnow seine although a dip net was also used. A few gobies were taken in the confluents of the lower Rio Chagres about two miles below the dam and in an overflow from the bait tanks located under the club house. Otherwise all others were taken at the foot of the spillway unless special mention is made to the contrary under the specific headings. At this point there is a most interesting mixture of the fresh and salt water faunas. The sea fishes working up the stream are of course stopped here where they seem to stay in considerable numbers making the place one of particular favor with both local and visiting anglers. Apparently these marine fishes subsist largely on the many smaller fresh water fishes rushed over the spillway. Nu- merous larger ones come over also and it is odd that so many sur- vive the swift and turbulent torrent presumably uninjured, but that they do, in part at least, is evidenced by the large number of strictly fresh water forms taken on the hook here. The Rio Chagres 1925] Breder: Notes on Panama Fishes 139 below the spillway is practically at sea level and the slight Atlantic tide can be detected to the very foot of it. This is most prominent when all gates are closed and water is discharging only through the huge hydro-electric plant which operates at all times. The water is fresh, except for the minute and practically negligible amount of salt water that climbs through the locks and comes over by way of the dam, for 'the sea merely pushes back the river flow during tidal periods. Usually one or two gates, at least, in the great Gatun dam are left open, supplying water to the lower Rio Chagres, in order to prevent Gatun Lake from rising too high. However, during the dry season these are sometimes left closed for during this period efforts are directed to maintain the necessary depth. Such action causes a rapid lowering of the stream below and leaves many small pools in pockets and depressions on either bank. Formerly these were always found to contain great numbers of small fishes. Sometimes even fishes of a larger size were so ensnared, it not being uncommon to find some running up to ten 'or fifteen pounds in weight. These were usually snook or jack. For some unknown reason, for the last eight years, according to Mr. Markham, who has carefully studied the fishes of this region from the anglers’ point of view, no fishes have been so entrapped by the receding waters, although there has been no marked decrease in River’s fauna. Mr. Markham further stated that at no time in his experience has a tarpon been so cut off from the stream. He explains it on the basis that, accord- ing to his observations, on the slightest slackening of flow all the tarpon at once rush into mid-stream, the point of maximum speed of current, for at least the tarpon observable from the apron of the spillway react in this manner to the slightest tampering with one of the gates. It would further seem probable that if one were at least partially entrapped it would leap as a reaction to the constraining influence of a small body of water and would soon find its way back to the main stream. Even this short list of the more prominent species existing at the Gatun Spillway seems especially appropriate at this time for in less than a year after the collection was made the United States Bureau of Fisheries released a number of North American species in Gatun Lake and its confluents. That is, on December 3, 1924, the following species were released (Fisheries Service Bulletin No. 116)., 140 Zoological N. Y. Zoological Society [IV : 4 Micropterus salmoides (Lacepede) 4 inch fish 2,250 Pomoxis annularis Rafinesque 2 inch fish 500 Lepomis pallidus (Mitchill) Fingerlings 500 It will be exceedingly interesting to watch the progress of these fishes, if any, accustomed to a comparatively large seasonal fluctu- ation of temperature, under tropical conditions, especially as there already exist in the lake various cichlids which appear superficially, at least, to occupy rather similar ecological niches. However, Gatun Lake covers much territory and shows numerous different types of habitat. At the foot of the dam one of the most striking absences was the complete lack of the smaller characins, for example, such as Astyanax which was found to swarm in practically all other places. The common names given in the list which follows are those used by the local English speaking anglers. ANNOTATED LIST OF SPECIES Family I. SILURIDAE 1 — Rhamdia wagneri (Gunther). Catfish. Fairly common. One example of 179 mm. Family II. CHARACIDAE 2 — Brycon chagrensis (Kner). Salmon. Said to be common by the local anglers. Regarded as a fair game fish and sometimes eaten, but highly relished only by the natives. One example, a green male of 325 mm., stomach empty. Family III. ANGUILLIDAE 3 — Anguilla rostrata (Le Sueur). Eel. Young ones from 54 to 68 mm. common in small pools under flat stones at the foot of the spillway where their further ascent, is prevented. Family IV. ELOPIDAE 4 — Tarpon atlanticus (Cuvier and Valenciennes). Tarpon. Common. The principal object of the spillway fishermen. The largest example taken here had a total length of 188 cm. and a weight of 38.5 kilograms, whilst the smallest had a length of 254 mm. Although none were taken during my stay, they were seen constantly leaping near the foot of the dam. On June 22 I again visited this place but did no collecting. The rainy season was then under way and while a thunder storm confined us to the club house, hundreds of tarpon were seen disporting themselves. So numerous were they that they literally wallowed and rolled with their backs out of water. It is 1925] Breder: Notes on Panama Fishes 141 interesting in this connection to note that some San Bias Indians that were with us, taken to view the Gatun locks, were much more impressed, judging from their gesticulations by these tarpon, their favorite food fish, than by the mechanical marvels of the white men. See the general discussion preceding this list for further data on the tarpon. 5 — Elops saurus Linnaeus. Bonyfish. Common but not valued by the anglers. One example of 355 mm. In- dividuals larger than this are rare at the spillway. Family V. ENGRAULIDAE 6 — Anchovia elongata Meek and Hildebrand — Common. Five examples ranging from 68 to 74 mm. In two the pec- torals reach the base of the ventrals but in the rest fall a trifle short of it. Family VI. POECILIIDAE 7 — Poeciliopsis isthmensis Regan. One example, a young female is questionably referred to this species. Taken in company with small specimens of Dormitator in a very foul drainage ditch a short distance below the spillway. Family VII. BELONIDAE 8 ^—Tylosurus timucu (Walbaum). Not common. One example of 272 mm. Family VIII. ATHERINIDAE 9 — Thyrina chagresi (Meek and Hildebrand). Needle fish. Common and much used for bait. Seven examples ranging from 45 to 62 mm. Family IX. MUGILIDAE 10 —Agonostomus monticola (Bancroft). Needle fish. Common and used for bait. Fifty-three examples, ranging from 27 to 54 mm., showed a mode of 47 mm. Apparently confused with Thyrina by the fishermen. 11 — Joturus pichardi Poey. — — Not especially common at the foot of the spillway, the local anglers having no name for it, not knowing it as a food fish. However, it is much valued by the natives. One example, a green male of 355 mm. The stomach was filled with an evil smelling black paste. Family X. CARANGIDAE 12 — Caranx hippos (Linnaeus). Jack. Apparently this species appears periodically at the spillway. Its identity coujld not be established with certainty from the descriptions as no examples were seen. 142 Zoologica: N. Y. Zoological Society [IV: 4 Family XI. CENTROPOMIDAE 13 — Centropomis pectinatus Poey. Snook. Two small examples of 31 and 34 mm. are somewhat reservedly referred to this species. The anglers rate snook as second only to the tarpon at this place but probably refer to another species, likely C. parallelus Poey, that attains greater proportions, specimens up to thirty pounds being spoken of. Family XII. LUTIANIDAE 14 — Lutianus ay a (Bloch). Red snapper. Apparently this species is occasionally taken here, judging from descriptions of the anglers although not taken in Panama by Meek and Hildebrand ’25. Examples reaching up to ten pounds were mentioned but none were seen. Family XIII. HAEMULIDAE 15 — Pomadasys crocro (Cuvier and Valenciennes).' Bass. Small examples common. Thirteen ranged from 25 to 65 mm. with modes at 25 and 55 mm. Those in the smaller group have the soft anal rays slightly longer than the enlarged second anal spine, but as size increases this ratio becomes inverted, the transition coming between 28 a^nd 35 mm. This dif- ference is to be seen with the fin erect. Larger specimens are valued by the fishermen. Family XIV. GERRIDAE 16 — Eucirtostomus calif or niensis (Gill). Snook. Small examples common. Forty examples ranging from 14 to 49 mm. showed a mode at 25 mm. while the three larger 63, 70 and 84 mm. fell without the curve. These young fish are apparently confused with young Centropomis by the anglers. Family. XV. CICHLIDAE 17 — Cichlasoma maculicauda Regan. Black perch. Common. Sometimes used for food. Three examples, 74, 88 and 102 mm. Family XVI. TETRAODONTIDAE 18 — Spheroides testudineus (Linnaeus). Blow-fish. Not very common. One example of 94 mm. Family XVII. GOBIIDAE 19 — Dormitator maculatus (Bloch). Doby. Common, but not in the main stream. Two examples of 39 and 41 mm. from a puddle under a leaky spigot outside the club house, and many seen in the overflow from the bait tanks. Thirteen examples from a filthy, hot and stinking drainage ditch below the spillway. These ranged from 18 to 35 mm. and showed a mode of 22 mm. 20 — Eleotris pisonis (Gmelin). Three were found at the foot of the spillway and six in a tiny spring about 1925] Breder: Notes on Panama Fishes 143 a mile below. This spring forms a small, but crystal clear, pool near the edge of a banana plantation about which a small stand of native plants clusters, oasis-like, on the baked and cracked flood plain of the lower Chagres. Many small birds and one spiny tailed rat came to drink in the short time I was there. The only species of fish in the pool appeared to be the present, but there were a few individuals of a slightly larger size that I was unable to capture. Those collected ranged from 34 to 80 mm. 21 — Sicydium salvini Grant. Common at the foot of the spillway. Nine examples ranged from 24 to 30 mm. and showed a mode at 25 mm. Fishes from the Rio Tapia While continuing our outfitting, at Panama City, the oppor- tunity was taken to make a small collection of fishes from the Rio Tapia at a point where the new Tapia Road crosses the river (Fig. 33). Most of the collecting was done nearly under the bridge and all within sight of it. The collection was made on February 1 and 2, mostly with a twTenty foot seine, although a few specimens were taken by dip net. It is believed to represent nearly the entire fish fauna at this point. It being the dry season, the water was crystal clear and shallow. This extremely interesting section is now easily reached by automobile over the excellent new road. The list is chiefly of interest because the locality is one that has not been col- lected in previously and therefore is an extension of local records. The Rio Tapia is one of the few short independent streams of the Pacific slope east of the Canal, not connected with any master drainage system. ANNOTATED LIST OF SPECIES. Family I. CHARACIDAE 1 — Curimatus magdalenae Steindachner. One example of 145 mm. 2 — Compsura gorgonae (Evermann and Goldsborough). Ten examples of from 20 to 25 mm. In company with the more numerous Pseudochierodon. 3 — Pseudocheirodon affinis Meek and Hildebrand. Forty-three examples of from 20 to 28 mm. A specimen of either this or possibly Compsura was seen to be attacked by a large spider in the very swift water in which both species were disporting themselves. The attack was made while I was at extremely close range, the spider being well observed. It was 144 Zoological N. Y. Zoological Society [IV: 4 noted not to relinquish its hold on the bank with its hinder legs, simply dipping its forward parts enough to effect the capture. An inadvertent move caused it to drop its prey and allowed it to escape while the fish was rapidly swept away by the swift water, apparently quite dead, although it had only been in the spider’s palps a very short time. See Gudger ’25, page 263, for the full account. 4 — Gephyrocharax atricaudata (Meek and Hildebrand). Seven examples of from 23 to 31 mm. 5 — Astyanax ruberrimus Eigenmann. Eighty-nine examples of from 25 to 98 mm. Some of the larger individuals were ripe or nearly so. The dark shoulder spots so conspicuous in most of the examples subsequently taken in the Rio Chucunaque were practically absent in the present series, showing faintly in only a few. Possibly some of these should be referred to A. fasciatus (Cuvier) but all showed some evidences of a more or less distinct caudal spot. 6 — Roeboides occidentalis Meek and Hildebrand. Sixteen examples of from 62 to 139 mm. This species was seen in large schools which appeared to be made up exclusively of this form and which were not seen to mix with the equally common Astyanax. Numerous specimens were ripe. 7 — Ctenolucius beani (Fowler). One example of 175 mm. Small schools composed of individuals of about this size constantly cruised about near the surface but were exceedingly difficult to seine in the clear water on account of their leaping ability which compares not unfavorably with that of Mugil. Family II. SYNBRANCHIDAE^ 8 — Synbranchus marmoratus Bloch. Three examples ranging from 88 immature to 248 mm., a spent female. Taken by seine amid the bottom debris. Family III. POECILIIDAE 9 — Poeciliopsis elongatus (Gunther). Two females of 21 mm. are probably referable to this species. Family IV. CICHLIDAE 10 — Aequidens coeruleopunctatus (Kner and St'eindachner). Three examples of from 88 to 118 mm. Family V. GOBIIDAE 11 — Philypnus maculatus (Gunther). Two examples of 67 and 180 mm. 1925] Breder: Notes on Panama Fishes 145 Fishes from Caledonia Bay The material and data upon which these notes are based was collected at Caledonia Bay, a small arm of the Caribbean Sea, that slightly indents the Atlantic coast of Panama near the Colombian border. See Fig. 33. As a collecting site it is of interest chiefly because of its comparative remoteness from other Atlantic coast collecting points. Porto Bello is the nearest place from which a collection has been reported (Meek and Hildebrand ’23). In the other direction a much greater distance is spanned before another site of collection is found. My presence here was in a certain sense accidental, having just emerged from the Chucunaque drainage by force of necessity, after crossing the continental divide over an un- speakable trail. I was stationed at the Indian village of Caledonia for slightly more than two weeks* although all the actual collecting was done within a period of four days, May 4 to 8. Many obstacles made even this collecting of extreme difficulty, amongst which may be mentioned a dearth of preservatives and containers, most of my supplies being nearly exhausted, a lack of proper collecting gear, not being equipped for marine collecting and numerous other ac- tivities which occupied much of my time and finally the inroads of illness into my heretofore unbroken health which eventually made my withdrawal from the field imperative. It is hoped that this fragmentary data will be of use to those who must eventually take up the much needed study of the life histories of the Caribbean fish fauna. In all cases where it was available the local appellation given the species by the Indians at Caledonia is indicated with syllabication, the spelling being purely phoenetic. Other names, representing forms not taken but of which we had fair descriptions from the natives, follow. Most of the collection was taken in a twenty foot seine operated over weedy patches, just off the beaches between coral snags. Calcium carbide, intended for use in acetylene lamps was found to be excellent as a poison for rock pools in the absence of any less Shark Sawfish Ray Nali Su-coo Ne-der-di-be Nali-ooru Oo-coo-ma-dera Oo-new-su Remora Flounder Any small fry 146 Zoologica: N. Y . Zoological Society [IV: 4 expensive substance. The efficiency of this gas generating substance was found to be high if strewn rather evenly over the bottom of such pools. The effect was rapid and not infrequently the doomed fishes would leap clear of the water landing several inches back from its edge. A small amount of angling was done whilst most of the larger fishes recorded, were collected for food by the natives of the village. The methods these Indians used in capturing fishes, which I saw during my stay at their village, have been considerably influenced by their contact with small trading sloops and schooners that barter with these people for the splendid cocoanuts that grow here in pro- fusion and the valuable shell of the hawksbill turtle. Of course the Indians receive a ridiculously small amount of merchandise in return, but to them the machine made artifacts of civilization are indeed of great value. In this way by degrees, many of them have ac- cumulated a considerable store of very presentable fishing tackle, that is, at least, good hooks and stout lines. As a result, much of their fishing is simply angling after our own manner, but always by hand line. A primitive sort of surf casting is occasionally indulged in. That is, a piece of coral is tied below usually two hooks and thrown bodily from shore over sandy stretches where it will not snag and is then slowly retrieved. In doing this a coil of line is thrown seaward much as sailors use a heaving line. Live bait is generally employed, most usually Hepsetia, Sardinella or Anchovia although they sometimes troll from their dug-outs for jack and barracuda with a piece of white or colored rag as a lure. What appear to be the more truly native methods may be tabulated as follows although these also show the effects of contact with civilization. Spear — Spearing is a common practice and the natives are naturally expert. The spear of this region generally consists of a point made from a square iron rod, obtained from some trader, along the edges of which notches have been filed to serve as barbs, a head of black palm into which the point is affixed and a shaft of some light hollow reed. No doubt originally the black palm head was shaped into a point in lieu of the iron spear. Bow and arrow — A certain amount of fishing is done by means of a small bow and a multi-pointed arrow, with black palm points and a reed shaft. This however is more of a fluvitile implement and not especially characteristic of the coast. Harpoon — Harpoons are constructed somewhat similar to the 1925] Breder: Notes on Panama Fishes 147 spears but with the characteristically removable point to which a stout line is affixed. This point is made of a three-cornered file from which the temper has been drawn so that it may be shaped by a second one. The three flat sides are deeply grooved in such a way that the cross section is a three pointed star. Then deep notches are filled into each of the three vanes to form barbs. I do not know if the finished point is retempered. Sharks, tarpon and other large fish are the quarry of these devices. Trap — A device which seems to be the primitive forerunner of a fish trap was found in considerable numbers near the village of Sasardi Nuevo, just south of Point Sasardi. Such a trap (Fig. 34) consists of a vertical wall of slats lashed together with vines and running out at right angles to the shore line for 200 or 300 feet, here . curving around abruptly for about half a circle with a radius of about five feet. Stationed at this point in a “Cayuka" or dug-out the fisherman stands with his spear poised ready to strike at the first edible fish deflected toward him by the wall. Some of these “ traps" are improved by a platform on which to stand and I was informed that they are frequently used at night by lantern light. No true traps, however, were seen in the possession of these Indians, but one of the negro cocoanut traders had a typical West Indian basket trap or “fish pot" on board. Seine — The closest approach to a seine which I saw was a small affair used for bait catching. A piece of coarse cloth about three by four feet strung on two long poles, much like a seine but without corks or leads was operated by two men standing in a cayuka holding the extending ends of either stick, with the net placed diagonally in the water. When a school of small fish came about it was sud- denly raised bringing some of them up. Its success resides chiefly in the abundance of these small bait fishes which were mostly Anchovia. However, the Indians that helped me operate my seine took hold with real skill, giving me the impression that they had used such gear before. This was in decided contrast to the dif- ficulty encountered with both our own negroes and the Indians of the Pacific drainage who were very unsatisfactory as seine men. Caledonia Bay is shored for long stretches by gentle sloping beaches of coral sand (Fig. 35) here and there interspersed with old weathered aeolian rock formations or dense stands of mangrove (Fig. 36) about which fine flocculent silt accumulates. The beaches 148 Zoologica: N. Y. Zoological Society [IV: 4 Fig. 35. A typical stretch of beach in Caledonia Bay. Along such shores small fishes find shelter in weeds growing just below the tide line. Fig. 34. The seaward end of a Caledonia Bay native fish deflector or trap. 1925] Breder: Notes on Panama Fishes 149 Fig. 36. A typical lane through a mangrove swamp at Caledonia. Here small Tetraodonts swarm and Anchovia come for shelter. Fig. 37. Brackish pools just back of the beaches at Caledonia, such as this one, generally are quite barren of fish life. 150 Zoologica: N. Y. Zoological Society [IV: 4 are narrow, dense shrubbery and coco palms growing almost to the water edge (Fig. 37), under which are brackish pools quite barren of fishes, for the tide here is so slight as to be practically negligible. Below the tide line in many places patches of Zoster a grows in pro- fusion. Here the young of many species seek shelter together with the adults of the more diminutive forms. Small tide pools, formed in the aeolian rock by the slight rise and fall, harbor a variety of invertebrates and small fishes, gobies and blennies being especially abundant in such places. Just off the edges of the mangrove swamps, schools of Engraulids and Clupeids surge back and forth retreating into its shelter when danger approaches, whilst Tetraodonts of small size teem in places under these upthrust tangled mats. Between May 4 and 16 about fifty feet from shore the water temperature ranged from 84° to 89° at the surface, whilst closer it reached 91° frequently and even higher in the rock pools. The accompanying air temperature ranged from 80° to 85°. A short distance off shore the gradual beaches take a rather abrupt drop, meeting the fantastic coral beds of the typical Carib- bean variety. Here the larger fishes commonly associated with such places disported themselves. Still further off large numbers of small islands arranged in a single or multiple series parallel the coast. Some consist of a purely coral origin whilst others attaining greater heights clearly have had at least a nucleus of other genesis. These islands are much more numerous than any map shows, mul- titudes of tiny ones, some not over one hundred feet in diameter being scattered all about. They are variously fringed by shores similar to those found on the mainland. Fending off the force of the breakers still further seaward a typical fringing reef, just about awash, marks the shoreward approach of the open ocean. In the lanes, often of considerable depth, between the islands and between them and the reef the larger fishes roam, sharks, barracudas and so forth as well as schools of various Scombrid-like fishes. The fish fauna, as would be expected, is typical of the Caribbean region and probably differs in no essential respect from that at Colon. As the collection is extremely small and fragmentary its variety indicates the richness of the region. In all only two rock pools were poisoned, two seining parties, one dip net party, four trolling and four angling parties, were undertaken and four native catches were examined. In addition to those species taken numerous others were seen but owing to circumstances were impossible of 1925] Breder: Notes on Panama Fishes 151 capture, such as large sharks, probably Carcharhinus, Scombrids, probably S comber omorus, Hemiramphids and numerous reef fishes more or less familiar. A considerable paucity of small species was apparent although the shores abounded with little fishes but these were mostly the young of larger species, the only small fishes found in abundance being Anchovia and Hepsetia. The complete absence of Poeciliids was striking and in such places as they are usually found, in man- grove swamps and similar locations, young Tetraodonts swarmed. However, near the head of brackish water and above in the fresh water of the exceedingly short streams of this slope they were not uncommon although none were seen which could be considered marine. ANNOTATED LIST OF SPECIES. Family I. SYNBRANCHIDAE 1 — Synbranchus marmoratus Bloch. Yar-be. Contrary to the findings of Meek and Hildebrand ’23 this species was noted to be fairly common in the fresh waters of Panama, being taken in numbers on several occasions. It is included in this list of marine fishes as a single example of 248 mm. was seined out of the sea at Caledonia. Gilbert and Starks ’04 who found it common at Miraflores suggested that it burrows in the mud as they took none with their seine. Although three specimens were taken by seine in the Rio Tapia on the present expedition this secretive habit was found to be characteristic for it was only necessary to drop some effervescent substance in a sluggish creek or pond that superficially appeared to be barren of them to see them come wriggling out of their places of refuge which were usually hollows under rocks. The example taken in salt water at Caledonia showed a very evident emaciation and it is quite possible that it was washed down to the sea in a weakened condition after one of the cloud bursts of the early rainy season, since the species appears to be a typical inhabitant of the head waters, as all the other material was taken well inland, chiefly in small tributaries. Further- more ripe examples were taken far from the sea, the species probably not being catadromus. Family II. MURAENIDAE 2 — Gymnothorax funebris Ranzani. Yar-be. A single example of 234 mm. Its stomach contained the carapace of a crab 16 mm. across. The coloration was of as bright a green as I have ever seen this species show. The natives evidently confuse Synbranchus with this as probably they are not very familiar with the former. 3 — Echnida catenata (Bloch). Ti Naipe. Not uncommon, but greatly feared by the natives. Their name literally 152 Zoologica: N. Y . Zoological Society [IV: 4 translates to water snake. This they also apply to serpents which dwell along the w^ater courses. One specimen of 370 mm. This example had its digestive tract crammed with the remains of small crabs, the carapaces of which averaged about 25 mm. across. These were intertwined with numerous parasitic worms. It was undeveloped sexually although Meek and Hildebrand ’23 took a ripe female of not more than 120 mm. longer in April at Colon. Family III. ELOPIDAE 4 — Tarpon atlanticus (Cuvier and Valenciennes). Me-la. Fairly common. Appears periodically in large schools. Valued by the Indians to the extent that it is apparently their most highly prized food fish. One large school which appeared on May 6 caused the bulk of the male popu- lation to give chase, which however was unsuccessful. Family IV. CLUPEIDAE 5 — Sardinella macrophthalmus (Ranzani). Sardina. Common near the edges of mangrove swamps, in small schools of individuals up to about 230 mm., or as scattered individuals in company with the more numerous and smaller Anchovia. Two typical examples of 44 and 46 mm. In weedy places along sandy shores smaller examples were common in company with juvenile Anchovia. Four examples from 23 to 26 mm. just out of the larval stage. With these were five specimens ranging from 22 to 24 mm. which appear to be of this species but still show numerous larval characteristics. Most prominently they still have a somewhat rounded ventral outline and are more elongate. The name used by the Indians is obviously borrowed from the Spanish. Family V. ENGRAULIDAE 6 — Anchovia brownii (Gmelin). Oo-new-su. Found in tremendous shoals of thousands of individuals skirting along the edges of mangrove swamps, into which they retreat on the slightest provocation only to reappear a few moments after quiet is restored. The precision and una- nimity with which these schools wheel and turn is striking, reminding one of the actions of a wandering flock of European starlings. Twenty specimens of from 41 to 51 mm. with a mode of 43 mm. In grassy patches along sandy beaches, smaller sized individuals were found in more scattering schools. These were just passing out of the larval Stage and ranged from 22 to 30 mm. showing a mode of 24 mm. (fifty two specimens). Family VI. BELONIDAE 7 — Tylosurus timucu (Walbaum). Tabu-garraty. Small examples were common close to shore. Larger examples about 300 mm., likely of this species, were seen over greater depths. Three examples, 100 to 106 mm. Family VII. SYNGNATHIDAE 8 — Syngnathus mackayi (Swain and Meek). Time-mass-su. Found amid patches of marine growth near shore. Previously not re- 1925] Breder: Notes on Panama Fishes 153 corded south of Porto Bello. Two examples, 132 and 134 mm., both males. Family VIII. ANTHERINIDAE 9 — Hepsetia stipes (Muller and Troschel). Oo-new-su. Abundant, fringing the shores all over. Previously Porto Bello marked the southernmost limit of its known range. The Indians apparently do not distinguish this from Anchovia. Twenty-three examples ranging from 29 to 46 mm. with a mode of 40 mm. Family IX. SPHYRAENID A E 10 — Sphyraena barracuda (Walbaum). Da-bu. Common along sandy beaches, especially the very young, which roamed around near the edges of weedy patches generally. Very large examples of this and possibly other species cruised about in the deeper water between the fringing reefs and keys more offshore. Whilst we were not equipped for col- lecting such material, trolling spoons were contrived by Mr. Marsh to the number of six or eight. Although supplied with heavy wire leaders they were all lost at the first strike of these fish, such was their force, some of which were much over 125-cm. The most numerous ranged between about 30 and 90 cm. Nine examples ranging from 47 to 71 mm. with a mode of 55 mm. Family X. HOLOCENTRIDAE 11 — Holocentrus ascensionis (Osbeck). Oo-ah-dar-see. Common, the very small often in company with Upeneus, young Haemulids, et cetera. Five examples, one of 55 mm. and four of about 153 mm. each, the former from a weedy patch and the latter by hook and line about coral heads. Family XI. MULLIDAE 12 — Upeneus maculatus (Bloch). Se-no-oo-ah. Very common about weedy patches. Nine examples ranging from 48 to 75 mm. with a mode of 55 mm. Family XII. CARANGIDAE 13 — Caranx ruber (Bloch). One example of about 300 mm. was taken by Mr. Charles Charlton on a trolling spoon intended for barracuda. Many of this or closely similar species were seen, all well offshore, mostly in the lanes between the islands. This species appears not to have been recorded south of Cozumel. 14 — Caranx crysos (Mitchill). Ca-lu. Fourteen small carangins, taken near weed patches, which range from 38 to 65 mm. and show a mode of 42 mm. are somewhat questionably referred to this species, as they show numerous small differences from the adult de- scriptions of this well known form. Family XIII. SERRANIDAE 15 — Cephalopholis fulvus (Linnaeus) Marga-too-go-willy. One example of 103 mm. 154 Zoologica: N. Y. Zoological Society [IV: 4 16 — Epinephelus morio (Cuvier and Valenciennes). Not uncommon, but not nearly as abundant as E. striatus. 17 — Epinephelus adscensionis (Osbeck). Marga-too-go-willy. Not uncommon. Apparently not distinguished from Cephalopholis by the natives. 18 — Epinephelus striatus (Bloch). Thirteen examples ranging from 150 to 610 mm. 19 — Epinephelus guttatus (Linnaeus). Too-goo. Eight examples ranging from 125 to 254 mm. Generally in company with E. striatus near reef formations. Family XIV. LUTIANIDAE 20 — Lutianus synagris (Linnaeus). Oo-ah-nalu. One example of 110 mm. 21 — Lutianus jocu (Bloch and Schneider). Nalu-oo-sele. Not uncommon. One example of 235 mm. was apparently immature and contained a single small crab in its stomach. 22 — Lutianus apodus (Walbaum). Nalu-oo-sele. Fairly common. Two examples of 150 and 343 mm. The latter, a nearly ripe female had a fish sound in its stomach whilst the former was immature. Naturally this is not distinguished from L. jocu by the natives. 23 — Lutianus ambiguus (Poey). One specimen of 100 mm. which considering its small size agrees well with the descriptions of this little known form. 24 — Ocyurus chrysurus (Bloch). Yala-tail-a. Not uncommon. The Indian name is obviously a corruption of the English West Indian name “Yellow tail.” A favorite food fish of the natives. Family XV. HAEMULIDAE 25 — Bathysoma rimator (Jordan and Swain). Oo-ah-su. One example of 103 mm. 26 — Haemulon flavolineatum (Desmarest). Seven examples of 101 to 152 mm. Twenty-three small Haemulids ranging from 22 to 35 mm. with a mode of 27 mm. are questionably referred to this species. The agreement is fair but the coloration and pattern is considerably different. These were taken in weedy places close to shore. 27 — Haemulon sciurus (Shaw). \ Bu-too. Not uncommon. Generally found in company with H. flavolineatum. 28 — Anisotremus surinamensis (Bloch). Wati-car-see-che. A single medium sized specimen was seen. 1925] Breder: Notes on Panama Fishes 155 Family XVI. GERRIDAE 29 — Eucinostomus calif or niensis (Gill). Small examples were found chiefly in weedy places and occasionally off the coral island opposite Caledonia village. One group of eight specimens ranged from 25 to 39 mm. and showed a mode of 25 mm. and another of seventeen specimens ranged from 12 to 20 mm. and showed a mode of 14 mm. Family XVII. POMACENTRIDAE 30 — Abudefduf saxatilis (Linnaeus). Canis-ah-oo-wa. Young Qua-beb. Fairly common about snags and coral heads. Young common in rock pools. Here they tend to school and keep more to the open places than the Pomacentrus there associated with them. The coloration of these was very pale in life, much less so than examples of similar size seen in Bermuda. They were somewhat reminiscent of Cyprinodon variegatus Lacepede in appearance and action. Twelve examples ranging from 12 to 29 mm. showed a mode of 17 mm. 31 — Pomacentrus fuscus Cuvier and Valenciennes. Was-che-che. Common in tide pools. Three examples of from 15 to 45 mm. Family XVIII. LABRIDAE 32 — Doratonotus decoris Evermann and Marsh. Ah-bu. Two specimens (32 and 40 mm.) of this exquisite little bright green labrid were seined from a patch of weed on a sandy beach. The apparent irregular boundary of the dorsal, the completely transparent nature of the distal margins of the vertical fins, the entire transparence of the pectorals and the full color- ation of the ventrals gave them in life, an extremely broken outline (Fig. 38), Fig. 38. Doratonotus decoris. A sketch of the larger example, showing the unusu- ally broken and abrupt outline of the coloration extending on the vertical fins, beyond which they are nearly invisible in life. 156 Zoologica: N. Y. Zoological Society [IV: 4 so that they greatly resembled torn fragments of sea weed. As they swam in the collecting pail, after the manner of their family, slowly winging their way along by means of their pectorals alone, which were perfectly invisible, and steering by their nearly equally invisible caudal, the illusion was greatly en- hanced, and were it not for their irregular and erratic gyrations I might not have recognized them as fish. Even when lifted in the hand the sense of touch was called in to supplement that of vision in definitely establishing the presence of the tail fin. Unfortunately much of this transparence has vanished in spirits. Examples from other localities do not always show this extremely irregular outline of color, there being apparently much variation in this character. Both of the present examples were ripe females. The ovarian eggs average 0.30 mm. in diameter. The abdomens were considerably distended. This is the southern- most record for the species. Family XIX. SCARIDAE 33 — Sparisoma hoplomystax (Cope). Ah-oo. Two examples of 70 and 350 mm. The former may possibly be S. niphobles Jordan and Bollman which closely resembles this species and is almost im- possible to differentiate at this size owing to the inconstancy of the differential characters. 34 — Scarus croicensis (Bloch). Ah-oo. Three examples ranging from 14 to 35 mm. They lack the characteristic light abdominal streak but otherwise the agreement is good considering their small size. Family XX. CHAETODONTIDAE 35 — Pomacanthus arcuatus (Linnaeus). Oo-ha-sigel-de. Common about coral heads in company with what appeared to be Ange- lichthys as near as could be told without the aid of a water glass. One example of 80 mm. Family XXI. TEUTHIDAE 36 — Teuthis bahianus (Castelnau). Oo-ah-nali. Seven examples ranging from 31 to 46 mm. with a mode of 33 mm. All near weed patches. Although showing numerous immature non-diagnostic characters they are referred to this species because of the shape of the tail even though it lacks the filament often found in the adults, the light margining of that fin which was blue in life and the body depth which increases from 1.7 in the smallest to nearly 2.0 in the largest. Family XXII. BALISTIDAE 37 — Batistes vetula Linnaeus. Old-wipe. Three examples of from 101 to 203 mm. Common about coral heads. The Indian name is clearly a corruption of the English West Indian name “Old wife.” 1925] Breder: Notes on Panama Fishes 157 Family XXIII. MONACANTHIDAE 38 — Monacanthus ciliatus (Mitchill). Two examples of 31 and 42 mm. In weed patches near shore. In life * numerous dermal flaps were prominently scattered over the sides which to- gether with the general greenish coloration appears to attain nearly the same degree of deceptiveness that Doratonotus does by other means. Both these species living side by side and normally of slow movements greatly resemble fragments of sea weed to the human eye at least. This seems to be a case of parallel development induced by similar needs of protective resemblance for such it certainly appears to be, one meeting the requirements of the adults of a small species and the other those of the young of a larger one that loses it later. 39 — Alutera scripta (Osbeck). One example of 301 mm. Family XXIV. OSTRACIIDAE 40 — Lactophrys bicaudalis (Linnaeus). Cala-pa-too. One example of 381 mm. Family XXV. TETRAODONTIDAE 41 — Spheroides testudineus (Linnaeus). No-sardele. Common in mangrove swamps, particularly where fresh waters are empty- ing out, -but all small. See Fig. 36. Here they “poke” about slowly, apparently feeding on small invertebrates attached to the roots and stalks of the mangrove trees. Occasionally about the shores of coral islands. Two examples 14 and 51 mm. Family XXVI. GOBIIDAE 42 — Gobius soporator Cuvier and Valenciennes. Too-goo. Common in tide pools. Twenty examples of from 16 to 71 mm. showed modes at 20, 45 and 65 mm. Family XXVII. BLENNIIDAE 43 — Labrisomus nuchipinnis (Quoy and Gaimard). One example of 21 mm. from a rock pool. Only one opercule showed the characteristic spot. 44 — Blennius cristatus Linnaeus. Too-wala-lady. One example of 36 mm. In company with Gobius. 45 — Salariichthys textilis (Quoy and Gaimard). Too-wala-lady. One example of 31 mm. In company with Gobius and Blennius. Natur- ally the Indians do not have separate names for these small fishes that are not of use to them. 158 Zoologica: N. Y. Zoological Society BIBLIOGRAPHY [IV: 4 Fisheries Service Bulletin 1925 January 2, No. 116, U. S. Bureau Fisheries p. 116. Gilbert, C. H. and Starks, E. C. 1904 The Fishes of Panama Bay. Mem. Cal. Ac. Sci., Vol. IV. Cont. Hopkins Seaside Lab. XXXII. Gudger, E. W. 1925 Spiders as Fishermen and Hunters. Natural History, Vol. 25, No. 3, May-June, pp. 261-275. Meek, S. E. and Hildebrand, S. F. 1923 The Marine Fishes of Panama. Field Mus. Nat. Hist. Pub. 215. Zoo. Series, Vol. XV, Part I., Dec. 20. 1925 Do. Part II., April 15. i ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME IV. NUMBER 5 THE LOCOMOTION OF FISHES By C. M. Breder New York Aquarium PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK September 28, 1926 Ncut fork Zoological 8>nrictu General Office: 101 Park Avenue , New York City GDflfens President, Madison Grant; Vice-Presidents, Frank K. Sturgis; and Henry D. Whiton; Chairman, Executive Committee, Madison Grant; Treasurer, Cornelius R. Agnew. Secretary, William White Niles $nar b of JHanagpns Ollaaa of iaar Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, T. Coleman du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field CGlaaa of 192B Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, Anthony R. Kuser, Mortimer L. SChiff, Frederic C. Walcott, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler, George Gordon Battle, Bayard Dominick (Elaaa of 1929 Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, John E. Berwind, Irving K. Taylor, Harry Payne Bingham, Landon K. Thorne W. Reid Blair, Director of the Zoological Park; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Mammals and Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds ; George S. Huntington, Prosector; H. C. Raven, Associate Prosector; Elwin R. Sanborn, Asst, to Sec’ y. and Editor. iE&itmial (EtmtmUto Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y- Zoologica, Vol. IV, No. 5. THE LOCOMOTION OF FISHES1 By C. M. Breder, Jr. Research Associate, New York Aquarium Introduction (Figs. 39-83 incl.) The locomotion of animal forms has been studied in more or less detail from time immemorial, certain ancient mechanical devices, such as the oar, battering ram, et cetera, having been obviously based on the observation of the performances of organic structures. In historic times biologists, physicists and engineers have investi- gated the problem of animal locomotion from both the philosophical and the utilitarian standpoints, striving for various reasons to attain a better understanding of the physical laws underlying the multiform locomotor efforts of the various animate objects moving about them. Many of these laws, when understood, have been applied to useful mechanical devices. Frequently the methods of the animals studied have been surpassed, for in many cases it has been possible to go beyond the limitations of animal structure in the employment of the inert materials used in machine construction. The manifold significance to biologists of the manner in which changes in position are effected by animals from the osteological, myological, phylogenetic and the purely philosophical standpoints are too patent to require elaboration. In spite of the large amount of work done, there still remain many poorly understood, and even some misunderstood, methods of producing changes in spatial relationships that have even been employed by animals existing long before the advent of man. Progress through the water as exemplified by fishes is prominent among those methods that are comparatively slightly understood even though phases of it appeared early in geologic history. The problem is complicated, as is any study involving observations on living fishes, by the fact that the subjects inhabit a different medium 1 Awarded the A. Cressy Morrison Prize in 1925, by the New York Academy of Sciences. 159 160 Zoologica: N. Y. Zoological Society [IV; 5 than that which is necessary for the life of the observer. Further, they often fail to act in a normal manner when confined in a glass- sided vessel, and in a considerable number of cases it is even im- possible to successfully transfer them to such a container in a healthy state. The accompanying bibliography testifies to the widespread in- terest this field of study has stimulated, although it is surprising to note that so much of the work has been in the nature of abstract philosophical discussion and that relatively so little has been based upon actual experiment and study of living fishes. While there is an extensive literature on the locomotion of fishes, as just noted, much has been left unanalyzed and unexplained. It is the purpose of the present paper, therefore, to amplify our existing incomplete knowledge of the subject as much as possible at this time; to refute that which is untrue in its literature, and finally to unify and systematize the whole. The hope is entertained that it will be of use to both the philosophical biologist and the practical student who may seek in natural phenomena an inspiration for his inventive genius. In developing the present paper the author has carefully confined himself to a consideration of the physical forces and the external body forms involved in the movements of fishes. All anatomical details concerning the manner in which a given movement is ac- complished have been avoided as much as possible, as it is felt that such details might better be the subject of a separate disserta- tion. Thus for the present purposes, fishes are simply considered as mechanisms capable of reacting to their environment in various ways, so as to effect their spatial relationships. How their acts produce the known results is the only question under consideration. It is evident that such data are a prerequisite to a thoroughly satis- factory explanation of the various types of locomotor organization among fishes. It was found advisable to consult a considerable number of people concerning the various points herein discussed. The points of view so obtained varied greatly and as a consequence much profit was derived in the way of suggestions, corrections and criti- cisms. To these gentlemen who gave generously of their thoughts and time the author is greatly indebted, and is especially beholden to Prof. W. K. Gregory, of Columbia University and the American 1926] 161 Breder: Locomotion of Fishes Museum of Natural History, for advice and general encouragement; to Mr. J. T. Nichols, for translating certain papers and for philo- sophic criticism, to Dr. E. W. Gudger, for bibliographic aid, both of the American Museum; to Dr. C. H. Townsend, of the New York Aquarium, for various facilities; and to Mr. L. L. Mowbray, also of the Aquarium, for descriptions of the behavior of certain fishes afield not seen personally; to Dr. G. B. Pegram, Dean of the Depart- ment of Physics Columbia University; Mr. C. M. Paxton, inventor, Mr. E. C. Bennett, naval architect, and the Blakeman-Hartshorn Co., consulting engineers, for mechanical and mathematical data; to Mr. Will Simmons, artist and student of animal life, and to his father, for criticism and editorial assistance. The paper has been divided into two chief parts: “Part I— Physical,” which treats of the locomotor efforts of fishes, grouped according to various characteristic types of movement, and “Part II — Systematic,” which treats of the way in which typical repre- sentatives of the larger taxonomic groups use their locomotor apparatus. Following these are the conclusions drawn, and various infor- mation of an appended nature. The various divisions and subdivisions are indicated in the Outline below. As such a subject is naturally intricate, many cross- references have been inserted, for nearly every part bears in some way on every other. OUTLINE Part I — Physical Introduction . 163 Forward Rectilinear Movement 166 Body Movement. 166 Anguilliform Movement 167 Ostraciiform Movement 169 Carangiform Movement 175 Comparison and Discussion 177 Movements of the Appendages 185 Longitudinal Structures 186 Caudal Undulation 189 Pectoral Propulsion 190 Jet Propulsion 190 Actions in a Current 198 Comparison, Interrelation and Discussion 198 162 Zoologica: N. Y. Zoological Society [IV ; 5 Maneuvering . 200 Turning 200 Rising and Falling 202 Fin Movements 202 Hydrostatic Elements 203 Starting and Stopping 205 Maintaining a Stationary Position 209 Effects of Current on Maneuvering 216 Comparison, Interrelation and Discussion 218 Movements other than Swimming 218 Burrowing 219 Creeping 219 Leaping 220 Flying 221 Comparison, Interrelation and Discussion 222 Reomorphism 222 Relationship to Habits and Development 233 Part II — Systematic Introduction 237 Treatment by Orders 238 Class — Cyclostomata 239 Order — Myxinoides 239 Order — Petromyzontes 239 Class — Pisces 239 Subclass — Elasmobranchii 239 Order — Plagiostomi 239 Sub-order — Selachii 239 Sub-order — Batoidei 241 Order — Holocephali 244 Subclass — Teleostomi 245 Order — Crossopterygii 245 Order — Chondrostei 245 Order — Holostei 245 Order — Teleostei 246 Suborder — Malacopterygii 246 Suborder — Ostariophysi 249 Suborder — Synbranchii . 253 Suborder — Apodes * 253 Suborder— Haplomi 254 Suborder — Heteromi 256 Suborder — Catosteomi 256 Suborder — Percesoces 257 Suborder — Anacanthini 259 Suborder — Acanthopterygii 260 Division — Perciformes 260 Division — Scombriformes 264 1926J Breder: Locomotion of Fish3s 163 Division — Zeorhombi 266 Division — Kurtiformes 267 Division — Gobiiformes 267 Division — Discocephali 268 Division — rScleroparei 269 Division — Jugulares 270 Division — Taeniosomi 270 Division — Atelaxia 270 Suborder — Opisthomi 271 Suborder — Pediculati 271 Suborder — Plectognathi 271 Division — Sclerodermi 271 Division — Gymnodontes 272 Subclass — Dipneusti 273 Locomotion of Fossil Fishes 273 Part III — Conclusion Recapitulation and Discussion 277 Definitions of New Terms 282 Summary _ 284 Bibliography Appendix Descriptions of Mechanical Devices 291 Models 291 Apparatus other than Models 293 Suggested Lines of Research 294 PART I— PHYSICAL Introduction Although the term “fishes” is used to designate an extremely diversified assemblage of vertebrates, there is, nevertheless, a basic- similarity in all their multiform movements, for with the exception of a few highly specialized off-shoots, they retain a primitive meta- meral arrangement of the lateral body muscles such as is not found in the tetrapods. The paired appendages are not ordinarily con- trolled by such huge muscle masses as in the tetrapods, in which the conditions are rever*sed, for there the emphasis is commonly placed on the paired limbs. Thus all the fishes may be considered under one rather natural locomotor grouping. On account of the density of water, it is necessary that an extremely fine streamline form be attained if any considerable 164 Zoologica: N. Y. Zoological Society [IV; 5 speed is requisite. This fact at once limits fishes requiring con- siderable mobility to a symmetrical, evenly rounded, unangulated and somewhat fusiform outline. With such a form, made necessary by evident mechanical circumstances, the primitive metamerism, together with other generally varying though often restricted features, gives ample means for the production of all necessary speed and agility. In forms that have departed radically from the basic bio-physical necessity of a streamline form, a decided restric- tion of speed is at once apparent, which is proportional to the amount of departure from the typical ichthyized form. In fact, only where mobility ceases to be important can these frequently bizarre forms, survive. Tetrapods that have returned to an aquatic habitat, such as Cetaceans and the extinct Ichthyosaurs, have revived the old streamline forms wherever speed was necessary. The musculature, while differing in detail on account of widely different ancestry, has revived the old body movements, especially in those with elongate fish-like bodies, the only ones that have acquired speed in swimming.2 The average vertebrate animal, excluding the heavily armored ones, has a specific gravity approximating that of sea water, if the buoyant effect of the lung cavity in pulmonate forms be discounted. This may be associated with a remote ancestry adapted to an aquatic existence. Fishes, at least, seem to have been able to approximate closely the various specific gravities of the different densities in which they are found without any profound specializa- tion. This has been done principally by means of the air bladder, which effects a great muscular economy entirely unknown to ter- restrial animals which leave terra firma in locomotor .flight. It is clear that for insects or birds to develop a sufficiently large sack for containing some substance lighter than air, they would require an immense and unwieldy apparatus for lifting their bulk in such a tenuous medium. Gliding animals, such as flying squirrels, are, of course, out of the discussion as are the ballooning spiders which simply drift about with air currents. The latter are more comparable to planctonic forms with extended surfaces to assist in flotation. In this connection a typical fish might better be likened to a dirigible 2 The body movements may even be in another plane as seen commonly in the cetaceans. 1926] Breder : Locomotion of Fishes 165 balloon than to any truly flying animal, which is fundamentally more nearly similar to an aeroplane. On account of the much greater density of water, the reaction of forces through or in it is much more direct than in such a tenuous medium as air. Therefore in comparisons of apparently similar creatures inhabiting either sphere respectively, due allowance must be made for this difference. It might be said that movement in water is slowed as compared with that in air, as every swimmer knows. A submarine motion picture bears a considerable resem- blance to a slowed study of a certain speed. This simply means that a much greater resistance is encountered in water than in air, and that in the interest of economy a much closer approach must be made to the ideal streamline form. Partly offsetting this is the greater reactance of any stroke under water, which is also due to its density. In water, certain differentials may become reversed. For example, in forms with a specific gravity of less than that of their habitat, the inverse of gravitational effect is felt, proportional to the difference in the two specific gravities. However, in most fishes the specific gravity is so nearly that of their native waters that the effect is practically nil. In a study of the motor efforts of fishes the following factors and conditions are always to be kept in mind. For a reaction to be possible, it must conform to these conditions: 1. A given set of motional phenomena must be attributed to an appropriate set of forces applied by the fish to the water and the resulting reaction of the water on the fish. 2. To produce the appropriate set of forces the fish in question must be physically able to move the various members in a manner that the occasion demands. 3. The fish must be observed to actually perform in the given manner Obvious as these three statements are, it seems necessary to give them, as there exist, in the literature of the subject, numerous instances where students have boldly neglected such considera- tions. As it is possible that the same result may be obtained in any one of several ways, it takes no little patience to observe clearly and note the motions of the various elements involved, and to resolve the resultant into its often numerous components. In this connec- tion the study of motion picture film prints and numerous still pictures has proved invaluable. 166 Zoologica: N. Y. Zoological Society [IV; 5 For purposes of convenience the present section is first divided under three main heading groups as follows: (1) Forward Rectilinear Movement, (2) Maneuvering, and (3) Movements Other Than Swimming, such as creeping, leaping and flying. The first two topics include not only movements commonly referred to as swim- ming but also all other acts of special adjustment on the part of a fish that is free from any contact with solid surfaces. To simplify matters, in all cases except where mention is made to the contrary, it is understood that the fish studied is in perfectly still water, because, unless some special use is made of a current, a fish is either carried along with it, the actual course being the resultant of the current and of the fish's own direction of motion, or some special act is performed to offset the effect of the current. Following these sections are two more special ones: (4) “Reomorphism, ” which deals in a general way with the external form of fishes, and (5) “ Relations to habits and development," which considers briefly the influence of these factors on locomotion. Forward Rectilinear Movement. Under this heading are considered all basic methods employed to produce a straight forward horizontal motion in still water while free from contact with any solid. This section covers those move- ments here referred to as swimming (in the restricted sense). Body Movements. Nearly all fishes make some use of flexures of the trunk and tail portions of the body for the purpose of inducing forward pro- gression. In only a few aberrant forms is this not done, for even most fishes that use the fins as the chief locomotor organs, will, at times of extreme hurry, resort to this more primitive method as an auxiliary, if their bodies are not so highly specialized as to prevent it. What at first appear to be two distinct types of oscillation are mechanically possible in the bodies of fishes in which a muscular metamerism is persistent. These are : (1) a somewhat serpentine motion as seen in the waving of a flag; which is induced by differential alternate contractions of the myomeres and is subsequently referred to as the anguilliform motion; (2) a wig- wag motion somewhat as seen in a fan, which is induced by a nearly simultaneous contraction of all the myomeres on each side, alternately, this is subsequently referred to as ostraciiform. These two movements are not as distinct 1926] Breder: Locomotion of Fishes 167 as would first appear and are usually seen as elements of a single motion. It is only in the extremes that one stands out clearly through the lessened complication of the other. With this thought in mind these twx> motions may be described and analyzed separately as follows: Anguilliform Movement: One extreme type may be so termed for it is illustrated almost perfectly in the Anguillidae and similar long-bodied forms. Primarily, this type of movement consists in throwing the anterior portion of the body into what is practically a sine curve and in passing this curve backwards by differential alternate contractions and relaxations of the serial myomeres. A mechanically analogous movement may be effected by grasping a long rope at one end and giving it a quick jerk at right angles to its axis. A wave will be seen to pass down it, dying out eventually because the one action alone is the impelling agent, whereas in the case of the living body an added impetus is imparted by each successive myomere. As soon as the first wave is completely formed and started rearward, a second follows, but on the opposite side, and so on alternately, giving the animal its typical waving motion. The initial wave is started by contracting the first few myomeres on one side of the body in such a manner as to slightly flex the head and to a less degree the posterior part of the body and form the first curve, or the wave may start further back, leaving the head practically still. The wave is then carried back by con- tracting the next posterior myomere and relaxing the first con- tracted one progressively. Maurey (1895) in studying animal locomotion in general, published two illustrations of a swimming eel which show this character of the anguilliform movement admirably, although he made no mention of the forces involved. Dean (1895) repeats one of these illustrations and makes the following statement concerning such movement. “It is the pressure of the fish’s body against the water enclosed in these incurved places which causes the for- ward movement.” The forward motion is certainly attained, as Dean wrote, by the pressure of the fish’s body against the water and in the following manner. The mechanical forces brought to bear on the water are diagonally backward as indicated by the small arrows in the diagram (Fig. 39 A). As these are distributed sym- metrically3 about the line of progression, a forward resultant of re- 3 This does not mean that the planes of pressure are at all times paired, from side to side. There may be an excess of not more than one on either side, but as soon as that one passes off the tip of the tail, the excess appears on the other side, so that, considering time as an element, these sine curves may be considered symmetrical about the axis of motion. 168 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 39. Diagram of anguilliform locomotion. The feathered arrow indicates the direction of progression. The short dark arrows indicate the direction of pressure from the backwardly moving waves. The dotted arrows indicate the vector quantities. A, the simple basic action of anguilliform locomotion. Note the relative sizes of the vectors. B, the increase in amplitude and decrease in wave length as generally employed. Note the relative sizes of the vectors as compared with the effective vector of A. The transverse dashed lines indicate the posterior decrease in distance between the crests, and the longitudi- nal ones the accompanying increase in amplitude. action follows as is indicated by the feathered arrow, for pressure from a moving plane is always at right angles to its surface. It might be objected that as the eel is moving ahead there is likewise adverse pressure diagonally forward from the anterior sides of these backwardly moving waves. The truth of this is evident and it simply makes it necessary for the fish to pass these waves posteriorly at a rate considerably faster than it expects to move forward. Cinematography has proven in certain cases, at least, that in comparatively slow movement the waves travel at over twice the speed attained by an eel in its forward progression. 1926] Breder: Locomotion of Fishes 169 The speed of the waves moving backward must exceed that of the forward motion of the animal as a whole. If the two speeds just equalled each other, it would mean that any point on a wave, such as its crest, would be stationary with reference to the sea bottom; but as one is dependent on the other this is obviously impossible. The difference in the two speeds is greater than that theoretically necessary on account of other factors, due to what in machines would be called mechanical imperfection. Maurey (1895) dismisses a certain interesting point with this statement, “ ... the undulations of the tail are .... more pronounced than those of the rest of the body.” By “tail” it is taken that he means that portion of the fish posterior to the vent. This is perfectly true but prompts further analysis. The change that actually takes place in the waves in progressing back- ward is that their amplitude increases and the distance between the consecutive crests decreases, as diagrammed (Fig. 39 B). Both these changes operate to effect movement more efficiently, for as indicated (Fig. 39 A and B), the vector quantities of force shown by the small dotted arrows vary as these two aforesaid factors become greater and less respectively, the effective vector force becoming relatively greater. There is a further advantage in this on account of the anatomical structure of eels in that they are nearly cylindrical anteriorly and well flattened posteriorly; for, because of internal stresses and strains, a ribbon shaped piece bends more readily (transversely) than does a cylindrical one. Furthermore the viscera are all contained in the anterior rounded portion whereas the caudal portion is devoted mainly to muscula- ture. Also more thrust may be had from a blade-like structure than from a cylindrical one (the diameter of which equals the width), in that it presents its surface more effectively to the water, so if for no other reason the emphasis is placed on the posterior portion. As the myomeres carry on the waves, each one, as noted on page 167, gives its little added impulse. These added impulses appear to be in excess of that needed to maintain the original curve to the end of the tail and so they keep increasing the amplitude and decreasing the spaces between the crests of the waves until the tail’s tip is reached. The diagram of this (Fig. 39 B) is purposely exaggerated for the sake of clarity, for actually, the changes noted are often slight, an instantaneous photograph sometimes being necessary to establish their presence. Ostraciiform Movement: The type of locomotion here dis- Zoologica: N. Y. Zoological Society [ft; 5 Fig. 40. The ostraciiform model. Perspective, side, and plan views. 1926] Breder: Locomotion of Fishes 171 cussed is illustrated well by the Ostraciidae and represents the opposite extreme from that of the anguilliform. It has been the center of considerable controversy, for it has been argued to be physically impossible to obtain a continuous forward thrust from this sort of movement. Since the Ostraciidae are incased in hard and unflexible tests it is obviously impossible for them to pass waves posteriorly as do the more flexible fishes. The tail protrudes from an opening in the test and is supported on a peduncle too short to be thrown into waves and is therefore insufficient to give any efficacious movement of the anguilliform type. In these fishes the locomotor emphasis is placed on the pectoral fins and other parts while the tail is used primarily as a rudder. However, at times when more than ordinary speed is required, the tail is given the only motion possible, a lashing from side to side. Borelli (1680) argued that such movement in itself was sufficient for locomotion and stated it as the basis of general fish progression. Pettigrew (1874) claimed this view to be erroneous, and stated that some flexure or turning over of the body was necessary. His logic would seem to be correct but that he was in error has been positively demonstrated by the construction of a model (Fig. 40). Here only a side to side swing was possible for the “tail fin,” but neverthe- less this mechanical contrivance moved forward with a sure and steady gait. The failure to include a certain elusive factor appar- ently led him into this error. Although the fact that such a device will move forward is generally recognized by physicists, a discussion of the behavior of the model will serve to illustrate better why motion of this kind will produce a forward component. Fig. 41 shows diagrammatically how this model starts off and picks up speed until its maximum is reached. The first swing of the tail does not appear to produce a backward component as might be thought and as Pettigrew reasoned, but being a simple bending at the pivotal point, swings the nose as is indicated from position 1 to position 2. That this is true may be shown by grasping a sculling oar, located centrally, and giving it a swing to one side. The boat will show no tendency to back up but will swing around to a position similar to “2” in the figure. It may be here pointed out that this is the elemental basis of sculling and only because of this fact is it possible to move a boat forward in such a manner. In fact a skiff may be sculled 172 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 41. Diagram of the movements of the ostraciiform model. Eight successive positions are shown. The dashed line indicates the path of the tail’s tip and the dot and dashed line that of the nose. Position 1 is that of the model at rest and 5 the return to the initial position after passing through one complete cycle, 2, 3 and 4 being intermediates while 6, 7 and 8 are repetitions of them but separated more widely on account of the in- creasing speed as the model gets under way. by simply pressing the blade from side to side in a manner identical to that of the model's “tail/' the other motions of a skilled sculler making for efficiency and ease on account of the structure of the boat and its position on top of the water. Pettigrew tried to explain this away in a roundabout and confusing fashion. The twisting of the wrist which tips the blade from side to side with each stroke serves the simple function of holding the oar in the oar-lock. Fig. 42 shows this diagrammatically. If the relation of the stroke and the angle of the blade is reversed, as in the dotted position, the oar jumps out of the oar-lock, as all beginners at sculling can testify, for the pressure of the water is diagonally upward instead of down- ward at the oar's tip. Other “fancy" movements of skilled scullers may make for efficiency in various ways but the fundamental principle remains the same. The following continued description of the action of the model serves to elucidate further the action of the sculling oar as well as the tail of rigid bodied fishes. The model now in position 2 (Fig. 41) is sent forward by a return stroke, as the tail presses diagonally backward. By the time the nose has reached the axis of progression the tip of the tail has also, and the model is as in the initial position, but advanced to 3, and is now mov- ing forward by inertia so that by the time 4 is reached it is fur- ther along but is gradually slowing down. From 4 it shifts to 5 and from 5 to 6 and 7 and so on, with the nose wig-wagging and the tail beating. The factor which Pettigrew misunderstood or failed to recognize is this oscillation of the anterior part. It is hardly necessary to point out in this connection that a freely sup- ported body such as a boat or a fish if jointed as described has no rigidly fixed pivot (in space) to work on but either part may be considered as bending about the other. Breder: Locomotion of Fishes 173 / 1926] Fig. 42. Diagram of a sculling oar, viewed from the rear. The horizontal dashed arrows indicate the path of travel of the blade. The angle the blade makes with the surface of the water is indicated on both sides of a stroke. The pressure is indicated by the arrows pointing diagonally upwards. The dotted blade indicates the position which causes it to rise out of the oar-lock and the dotted arrow indicates this pressure. The movement thus obtained is very close to that of the trunk fishes as in such forms a sharp contraction of the lateral muscles inserted in the peduncle first on one side and then on the other suffices to produce this action. The greatest difference be- Y' tween such a fish and the mechanical model described is that in the latter there is a single joint about which the fore and aft portions turn whereas in the former a series of a few joints (in the vertebral column) cause a broad curve to be formed as each vertebra bends on the next adjacent one. This, of course, is merely a quantitative difference as pointed out on page 171. In addition to the explanation given above, a rearward flow (or a forward motion) is automatically induced by the back and forth oscillation of even a rigid tail piece. A mechanical analogy may be made by placing a cue flat on a billiard table, with a ball in contact near the handle end. If a swinging movement be im- parted to the cue with the butt end as a pivot the ball in addition to its lateral path will travel relatively rapidly along the length of the cue away from the center of rotation. This is true irrespective of the speed at which the cue is moved. An important point not considered as yet bears on the impinge- ment of a flowing stream of water on a moving blade, in that on striking a surface it spreads out and follows it as a sheet flow. However, the alternate striking and drawing of a particle of water causes it to travel backward in a zigzag line progressively more and 174 Zoologica : N. Y. Zoological Society [IV; 5 Fig. 43. Diagram of the path of particles of water about a rigid tail fin. The dotted lines indicate the paths of three particles of water starting at various points along the fin. The original thrust is indicated as though at right angles to the plane of pressure. The initial path would not be quite at right angles to the plane because of the circular motion of the blade but is so shown here for the sake of charity. Note that the particles starting near the pivotal point attain a more nearly straight backward thrust than do those starting further back. more nearly parallel to the axis of progression. Fig. 43 diagrams this. Under this action the particles follow the paths in the dia- gram under the constant urging of the blade only. For example, if any point be taken on one of the paths and connected with the pivot, it will give the position of the tail when the water particle is at that point. The breeze from the edge of a fan will demonstrate the truth of this, for otherwise a fan would fail in its purpose. This action coupled with the fact that the fin is curved, gives a nearly backward thrust, greatly increasing the effective power and reducing the swinging of the head to a minimum. The relative size of the body and tail of course, influence the speed, as do the amplitude of oscillation of the tail, the shape of the body and many other similar factors. The exact mechanical relations of these various elements are outside the province of the present discussion, but doubtlessly they could be reduced to and expressed as formulae, varying with each form. A piece of apparatus made so that the two parts (body and tail) were exactly equal in size, shape and weight would fail to move forward at all, simply remaining in one place while bending and re-bending upon itself. This may be observed sometimes in terrestrial insect larvae. Occasionally a caterpillar, very similar fore and aft, in its efforts to extricate itself from a puddle, will violently bend from side to side so that the arch occurs centrally with no progressional movement at all. 1926] Breder: Locomotion of Fishes 175 The greater inertia and resistance to motion sidewise through water of the larger part of the model, referred to above as the body, causes it to move through less of an arc than the lighter and more mobile tail portion, which fact tends to keep the model or fish on a true course and causes the force to be expended in a back- wardly direction, as forward is the path of least resistance. Carangiform Movement: The movements displayed by most typical fishes are intermediate between the two extremes just examined and the more generalized members of the family Carangi- dae, such as Caranx, occupy about the middle position in this respect. Many fishes do obviously never throw their bodies into a series of reverse curves as described under Anguilliform Move- ment nor are their bodies hard and completely inflexible as in the case of the Ostraciidae, but the entire body flexes as from a pivot (or fulcrum) at the atlas. As in Anguilla, the first act preparatory to locomotor effort is the contraction of the first few anterior myomeres on one side. The head being less in bulk and offering slighter resistance to the water than the rest of the body, moves through fy\ t \ \ I \ I \ V \ V \ %\-| V\ 1 A Fig. 44. Diagram of carangiform locomotion. The positions are serial from left to right, with the preceding indicated by dotted lines for comparison. A, the initial or resting position. B, the contraction of a few anterior myomeres on one side. Note the great deflection of the head and the slight deflection of the body and tail. C, the continuing backwards of the metameral contractions with relaxations immediately following swings the tail as shown. Here two intermediate positions are indicated. The fish is now in a similar but opposite position to that of A. D, Identical but opposite to B. E, Identical but opposite to C, completing one cycle and returning the fish to A. F, a composite of the movements of the preceding five positions indicating the amplitude of the caudal and cephalic oscillations. The lower dotted lines indicate the head’s oscillation, projected. The feathered arrow indicates the axis of progression. 176 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 45. Carartx superimposed on Anguilla, indicating the similarity in movement and the abbreviation of the one over the other. the greater arc and may be said, relatively, to be thrown to one side, as a result of this primary contraction. The myomeres then succes- sively contract and relax caudad, bringing the tail to the axis of the head with a powerful sweep. Fig. 44, A, B and C, diagram this in a series of positions viewed dorsally. Because of the pivotal point being at the base of the skull a short swing of the head is all that is necessary to place the tail in a position for a long telling stroke. Fig. 44, D and E, complete the cycle to the other side * and thus the fish returns to the original position. F is a composite of the movements showing the amplitude of the caudal and cephalic oscillations. While both the caudal and cephalic axes cross the axis of progression, the former is always moving toward the axis of the head, while the latter is always moving away from the axis of the bocjy. This is possible because of the constant shifting of these two axes, which, as we have seen, reaches its maximum prominence in the Ostraciidae. Fig. 44 C shows two intermediate positions of the body and tail in making a stroke, and clearly indicates its relationship to the anguilliform type of movement, while the other parts of Fig. 44 show equally well the relationship of the carangiform to the ostraciiform. Fig. 45 in which Caranx is superimposed upon Anguilla , serves to illustrate that the move- ment of the body of the former is simply an abbreviated form of the latter, in which never more than one half of a sine curve is formed. It should be noted here that while the pivot is at first at the atlas, it moves backward (as the point of greatest curvature) with the sweep of the body and tail so that near the end of the stroke it is nearer to the tail than to the head. This is especially advantage- ous for at first the anterior part is relatively short while later the posterior part is. From this it should be evident that an advan- tage is had over the ostraciiform type in which the pivot is fixed well behind the middle of the body. Likewise an advantage is had over the anguilliform type, which possesses what may be considered 177 1926] Breder: Locomotion of Fishes multiple pivots, a condition in which this advantage is practically lost. Comparison and Discussion: The preceding outline of the action in the body movements of the two extremes (Anguilliform and Ostraciiform) and the norm between them (Carangiform) makes it now possible to discuss in some detail the various characteristics of the results of progressive metameral contractions as seen in fishes generally. There is such a complete gradation through these three rather arbitrarily chosen types just described that it is impos- sible to say where one leaves off and the other begins. In fact, the anguilliform type might be considered, in a sense, as a great number of ostraciiform units arranged serially, or the latter as an abbreviated form of the former. A fan in oscillating is analogous to the ostraciiform movement if rigid, but if sufficiently elongate and flexible the same driving power produces an anguilliform (flag waving) movement. It is obvious that these differ only as do their proportions and the flexibility of the oscillating members. This series includes all fishes that have a functional tail, that is, one which is capable of being swung from side to side in loco- motor effort whether it is habitually so used or not, and it excludes only such unorthodox forms as Hippocampus. Further, it might be shown by means of a statistical curve of variation that the carangiform type of movement is the one most frequently encount- ered. That is to say, fishes of that type would roughly form the mode while the anguilliform and ostraciiform would fall at either end of such a curve. All others would come between these in in- creasing frequency as the mode is approached. Furthermore, the carangiform method is pre-eminently fitted for speed and appears to be of the highest efficiency, since it always accompanies fishes of high speed, such> as Pomatomus , Scomber , and fast oceanic forms generally. The other two types are both principally methods of comparatively sluggish littoral forms. The ostraciiform motion is usually used only as an auxiliary to other locomotor efforts not concerning the trunk proper, while the anguilliform is typical of fishes such as the thigmotactic eel and moray, although these are capable of speed utterly unknown to the passive trunk fishes and are even occasionally to be found among the pelagic fauna as in the case of the Ophidiidae. Doubtless locomotor characteristics in general have a con- 178 Zoologica: N. Y. Zoological Society [IV; 5 Pig. 46. Diagram of the continual backward push of the caudal. Each curved line represents the vertebral axis of an anguilliform fish. Ten successive positions are shown numbered from O through 1 to 9 and back to O. The diagram has been broken in two simply to avoid a confusion of lines, 5 in the left, being identical with 5 in the right. Note the dotted arrows indicating that the push of the tail’s tip is always diagonally backward. There is no “non-effective” stroke. The dark disc on line O in the left half is the same as that on O in the right in reference to the wave motion and indicates the distance that one wave has traveled in passing through the ten successive positions. The direction of motion is opposite to the travel of the waves. siderable phylogenetic significance. However, the great frequency and extent of parallelism and convergence among widely different groups due to the direct contact of such plastic animals as fishes with such a powerful environmental factor as similar locomotor needs, shows that no great importance should be attached to the grosser movements of the trunk and tail as indicative of close phylogenetic relationship. That the tip of the tail, no matter which direction it is moving in, is continually “pushing” water diagonally behind is made clear in 1926] Breder: Locomotion of Fishes 179 Fig. 46 wherein is represented a series of positions of the tip of the tail showing how it is always presenting a backwardly moving plane. An anguilliform type is represented, but it is obvious that it is equally true for the others. In the case of the ostraciiform type the difference is more of quantity than of kind, as no matter how short the projecting peduncle is, there still remains a few myomeres to operate in the conventional manner as far as interfering cir- cumstances permit, and the tail fin is always flexible. As pointed out in the diagram of the ostraciiform model in motion, in this device the second half of any stroke is entirely concerned with swinging the nose and does not have a forward thrust on account of the mechanically rigid tail piece. This clearly never obtains completely in nature on account of the flexibility of caudals but doubtless is Pettigrew's “less effective stroke." The preceding paragraphs show that these three designated types of body movement are in reality nothing but varying forms of the same rather involved motion, differing only on account of the differences of shape of the body in which the metameral con- tractions occur. They are broken up here only for purposes of analysis. Pettigrew (1874) states that fish always throw their bodies into two opposite and complementary curves, which is necessary to his explanation. This is at least conceivable for many forms, but is never done by Caranx and a host of others. Simple observa- tion will suffice to prove this. He further intimates that as the tip of the tail moves away from the axis of motion it is impeding prog- ress, and drags in such absurdities as having the fish roll over slightly so as to present an oblique blade to the water. Large sturgeon when swimming slowly, especially when starting off, have a noticeable roll, which is partly due to the fact that the small vertical fins have only slight powers of steadying; the stabilizing effects of which are explained under “Reomorphism.” The reason for this roll is purely a mechanical one and the action is seen in the ostraciiform model when starting, especially before the nose has attained its full swing. It is apparently this movement that he construed to be a voluntary turning over, although how it is supposed to be voluntarily accomplished is not indicated. Pettigrew further writes that the mackerel twists its tail to such a degree as to be “very much after the manner of a screw in 180 Zoologica: N. Y. Zoological Society [IV; 5 a steamship” and thus “to drill the water.” Observation con- firms this in no way whatsoever, and a great many more objections could be raised to other parts of his explanation, which, however, may be passed over here. Probably the most remarkable part of it all is the lack of criticism that he has had. Du Bois-Reymond (1914) touches on Pettigrew’s work, showing some knowledge of the ostraciiform effect, but leaves the subject essentially as it was. Bridge, in the Cambridge Natural History (1904), in dealing with locomotion, simply paraphrases Pettigrew, thus giving the latter’s views a very wide circulation. The movements have thus far been considered only in reference to the fish itself and have not been referred to the water through which the fish is moving, except in the case of the ostraciiform model. Fig. 47 diagrammatically represents the motion of a caran- giform fish relative to the surrounding medium. It is to be noted that the path is truly rectilinear. The boundaries of the single hatching represent the limit of oscillation of the tail’s tip induced by the metameral undulations. Those of the double hatching repre- sent the limit of the snout’s swing. The track or path through which the fish moves is rectangular in cross section, if viewed end on. That the head makes a much smaller sweep is indicated by the double and single hatching. The initial position in which the fish is shown is the same as C (Fig. 44), and the head is ready to be flexed to the left. Assuming the fish to be already in motion, the nose is brought to cross line 2 and the tail to 2'. The body is then in position for a really effective stroke which runs the fish forward to position 3 and 3', after which the other half of the cycle is com- pleted, bringing the fish to 5 and 5'. In this diagram momentum has been omitted for the sake of clarity as it is quite obvious that the thrust from the body and tail would carry the fish along at scarcely diminished speed while it wagged its head in order to give a stroke from the other side. Also it is clear that the stroke of the body and tail must tend to deflect the head toward the midline, the direction to and beyond which it is to go at the finish of the stroke. It is to be noted that most of the time the head is either travelling on the right or left limiting boundary, the path being indicated by the dot and dashed line, while the tail is occupied for the most part in passing from side to side, except for momentary touchings of its limiting lines on either side, indicated by the dashed line, 1926] Breder: Locomotion of Fishes 181 Fig. 47. Diagram of the movements of a carangiform fish. Four positions are shown. The dashed line indicates the path of the tail’s tip, and the dot and dashed line that of the nose. Position 1-1' is identical with C of Fig. 44, 2-2' with D and 3-3' with E. This represents one-half a cycle, the remaining positions not being shown except the one at the extreme left which is 4-4' (B of Fig. 44) of the next cycle. Momentum and increasing speed have been omitted for the sake of clarity. The single hatching indicates the extent of the amplitude of the tail’s swing and the double hatching that of the nose. The black area indicates the orthokinetic part. while the head is switched to the same side with which the tail is in contact. This is in agreement with observation, as it means that the head is comparatively quiet in its oscillation while the caudal portion appears in violent motion and actually is, for, in addition to the above relative movement, it swings through a wider arc. A mackerel swimming rapidly shows the posterior portion of the body simply as a blur, while the head can be seen simply weaving from side to side ever so slightly. While most of the fish's body is oscillating over a shaded area in one way or another, there is a point about at the atlas which travels directly forward in a straight line. That portion of the body that does not leave the boundaries of the double hatching at any time, indicated in black, represents a solid core of the fish's body which is continuously concentric about the axis of motion. The remaining parts are occupied first on one side and then on the other part of the track, which is shaded. It is plain that it is not always the same exact materials which occupy the central core, but this variable core, always the same in bulk as long as the fish swims by means of similar oscillatory movement, is the only part of the fish which actually goes continually forward without making a waved track through the water leaving alternate unoccupied areas on either side. This section may be called the orthokinetic part. The parts which oscillate about this section, anteriorly and posteriorly may be called respectively the cephalic amphikinetic , and the caudal amphikinetic parts. It is unnecessary to figure the anguilliform movement in this manner for it follows that if a long produced form occupied the place of the fish in the diagram, similar paths of motion could be 182 Zoological N. Y. Zoological Society [IV; 5 traced about it. The orthokinetic part would bear the same relation to the fish and there would be no second or third as might be sup- posed, because of the numerous crossings of the axis of motion made by the sinuous body. As the undulations are passed backward, a series of sections of the fish alternating with interspaces would follow this path but would produce a no more continuous section than would the sidewise movement of the head and tail. However, if the undulations were so slight that the crest of any wave never caused the body to pass completely to one side of the axial line, the orthoki- netic part would be considerably attenuated. This probably seldom happens in active swimming. On the other hand an eel violently agitated may thrust the head from side to side in such amplitude that no orthokinetic part at all is formed. When this takes place, which seems only to be at times of mortal danger, except in greatly compressed forms, the progress made is not proportional to the apparent energy expended and the fish usually soon resorts to the more normal method of slipping along. Therefore it is believed that this is an abnormal effect caused by the over stimulation of great danger. It is doubtful if such locomotion is nearly as effective as when the orthokinetic part is allowed to remain intact. In the ostraciiform model the point that moves forward continually is, of course, at the joint (Fig. 41), for this point in each position is on the axis of progression. It would therefore seem that the anterior placement of the orthokinetic part is associated with the efficiency of locomotion through water by means of body undulations. The foregoing analysis of the differences and similarities of the three phases of the body movements of fishes induced by metameral contractions may now be contrasted in the following table: Anguilliform 1. Cephalic amphikinetic part moves imperceptibly or very slightly from side to side, normally. 2. More than one-half a sine wave is formed by the body, typically several en- tire waves. TABLE I Carangiform 1. Cephalic amphikinetic part moves through a con- siderable arc, and its alter- nation with the body and tail movement is generally observable. 2. Not more than one-half a sine wave is formed by the body and frequently it is anatomically impossible to form even that much. Ostraciiform 1. Cephalic amphikinetic part moves through a large arc, but there is no alter- nation, the nose shifting with the tail. 2. Hardly any curve is formed at all by the small peduncle, simply a swing from side to side being ob- tained. 1926] Breder: Locomotion of Fishes 183 3. The pivot is at the base of the skull. 4. The orthokinetic part is concentric about the axis vertebra but is followed by a number of backwardly mov- ing sections. It may even be absent. 3. The pivot is at the base of the skull at the beginning of a stroke, but migrates backward with it. 4. The orthokinetic part is concentric about the axis vertebra, but is not followed by any sections. 3. The pivot is at the base of the caudal peduncle. 4. The orthokinetic part is concentric about the ver- tebrae at the base of the peduncle. This table may be taken as indicative of the characters by which the three forms of body movement are defined and by which the dynamic metameral characteristics of a given species may be recognized, for it is to be thoroughly understood that they simply represent phases of the same thing that have been forced on fishes in response to their changes in bodily form, often by evolutionary factors not directly concerned with locomotion. Special acts of the tail fin relative to rectilinear movement but not the result of the cumulative metameral contractions are dis- cussed under the head of “ Movements of the appendages,” and their form under “ Reomorphism, ” but the importance of the tail to generalized fishes may be considered here in relation to its motion induced by the cumulative effects of the metameral body muscles. This cumulative effect of the lateral musculature gives the tail a wide swing and in so far is effective in much the same way that it is in the ostraciiform type; moreover the tail, being flexible, carries out the movement of the rearward-traveling waves by momentum; but this movement is modified by the differential forces at work on t the tail incident to its motion. The functional significance of the tail is indicated by the following experiments made on Scardineus erythrophthalmus: Experiment 1. The tail fin from one example was carefully amputated. The fish was then placed in a trough eight feet long together with an entire example of equal size. As usual, these two fish rested side by side at the far end of the trough. Carefully approaching this end, a smart blow with the flat of the hand was delivered against it, which sent them scurrying to the other end where they took up similar positions. Each time that this was tried, they arrived at the distant end apparently simultaneously after traversing the intervening distance side by side. The only obvious difference was that the example minus the tail oscillated the body faster and through a greater arc. This might be taken as a measure of the efficiency of the tail, making a greater 184 Zoologica: N. Y. Zoological Society [IV; 5 effort incumbent on the mutilated fish in order to keep abreast of the normal one. However, it is doubtful if the normal example was travelling at much less than top speed since none of this species has been seen to travel measurably faster in an eight foot run under any circumstances. For simple mechanical reasons the change in oscillations would be expected to take place. The greater resistance of the tail to movement from side to side being removed, with similar exertion it would follow that a wider arc or more abrupt curve would be passed through by the stump, and at a higher rate of speed. This increased speed of the lessened area of pressure moving by means of the same muscle mass appears to ‘offset to some extent the difference between it and that of a slower motion with a greater functional pressure area. However, in a longer run there is no doubt, that a marked difference would appear in the time required to cover a given distance. Experiment II. The tail of another fish was immersed in liquid air, which instantly killed and made rigid the portion so treated. The film of ice formed about the tail tended to make the fish float up, but aside from this inconvenience it was obvious that the wig-wag of this now rigid fin was extremely inefficient. A series of individuals were so frozen as to make each successive one rigid a little more anteriorly. A regular decrease in speed was noticeable, until all myomeres were frozen and motion ceased. To demonstrate that it was the mechanical effect of a rigid tail, and not nervous inhibition incident to freezing that caused the above result, the fore part of the head of one fish was frozen, with ice solid in the gill and mouth cavities. Strangely enough this fish showed no prominent reduction in speed. Further, in thawing out, fishes partially frozen from the posterior end, resumed some of their former speed although the frozen section was lifeless and flexed only because of the movement transmitted to them by the live anterior myomeres. Experiment III. In other examples the nerve cord was severed just past the viscera in such a manner as to paralyze the posterior portion. This caused a slight slowing down which is believed to be proportional to the number of myomeres cut off. The deadened portion oscillated as before, however, on account of the continuing effect of the ripple started anteriorly, although of course the efficiency was less. These results were practically identical with those obtained by thawing tails after freezing them in liquid air, as in Experi- ment II. Experiment IV. In still another example, the body was made rigid by passing a fine steel wire parallel to and above the vertebral column to a point just behind the brain case. This succeeded in inhibiting the undulations and locomotion almost ceased. A very slight forward movement was produced by wagging the head and tail fin, which the wire did not prevent. On removal of the wire, locomotion proceeded as before. The inference drawn from these experiments is that the meta- meral contractions and the resulting flexures of the body act in- dependently and directly on the surrounding medium as well as by their cumulative effect on the tail fin, the importance of which 1926] Breder: Locomotion of Fishes 185 is entirely based on its terminal position. The significance of this seems never to have been stated, although it is clear from the above four experiments that by far the greater effect is due to the con- tracting myomeres themselves, rather than to the tail fin; this is especially evident by experiments I and IV. See Breder (1925b) for other similar data. The relative importance of the tail fin varies with its shape and size as well as the amplitude and wave length of the body oscillation. Regnard (1893) found that the pull on a tow line of a Cyprinus carpio was reduced by one-half by removal of the tail fin. This is questioned, for similar experiments failed to yield the same results, this being found a very unsatisfactory method because of induced nervous inhibitions. Also, the subjects suffered from exhaustion very rapidly, usually giving a few vigorous tugs and then sulking. It would seem that after such an operation, a measured pull could be easily half as great on account of many causes aside from the simple loss of the tail fin. Movements Of The Appendages. Movements of the appendages are more generally concerned with maneuvering than with locomotion in a straight line. How- ever, there are numerous forms representing different locomotor types in which all fins (except the pelvics) have been so modified as to be the primary locomotor organs. Further, most species commonly using the more primitive locomotor method, body undulation, also use the fins at times for inducing forward move- ment, especially for slow, short travels. In some, two or even more methods appear side by side so as to provide multiple locomotor systems to be used either together or separately. In fishes which seldom, if ever, use body undulations, the same general body form is retained, unless they are of exceptionally slow movement, for it is primarily the necessity of conforming to streamline effect that shapes them. Various fins of numerous fishes are capable of being undulated for purposes of progression. These movements are comparable to the undulations of the body examined under the heading “Body Movements.” Certain of these are nearly the same in application (those involving the longitudinal fins), while others (those involving the caudal and pectoral fins) introduce other elements. That is, 186 Zoologica : N. Y. Zoological Society [IV; 5 Fig. 48. Undulation of the longitudinal structures. A, Amiatus, dorsal undulation. (Amiiform). B, Gymnotus, anal undulation. (Gymnotiform) . C, Monacanthus, inclined dorsal and anal undulation (Balistiform) . D, Raja, pectoral undulation, (Rajiform). in many forms the undulations of certain fins may be considered as localized parallels to the body movements of other forms. Longitudinal Structures: The dorsal and anal fins in certain fishes carry on wave movements which affest motion in a manner similar to that caused by anguilliform movement, although of a gentler sort. Usually in such forms undulations of the body are resorted to when a higher speed is desired. Amiatus is an excellent example of the type which shows undulations being carried on in the 1926] Breder: Locomotion of Fishes 187 dorsal fin. This may be called Amiiform (Fig. 48 A). However, as above noted, this species undulates the entire body for rapid movement in a manner intermediate between anguilliform and carangiform locomotion. Gymnotus, on the other hand, has the seat of similar undulations in the anal fin. This may be called the movement gymnotiform (Fig. 48 B). Batistes shows this form of movement in both dorsal and anal fins, each being inclined a little from the horizontal, but oppositely, so that their resultant causes a forward movement. This may be called the balistiform movement, (Fig. 48, C). In these longitudinal fins the waves are propagated by simply moving the rays from side to side serially in a method analogous to the anguilliform movement. This has been demon- strated by a model (Fig. 49), which incidentally may serve as a mechanical representation of the anguilliform motion as well. Instead of myomeres contracting to bend the body, muscles attached to the sides of the fin rays at their bases contract, so deflecting them alternately from a vertical position in the appropriate manner. Raja represents a slightly different sort of longitudinal undulation. Here the greatly expanded pectorals effect a similar end but operate in a vertical plane. Maurey (1895) illustrated these movements beautifully by serial photographs. This may be called rajiform movement (Fig. 48, D) . It is to be noted that longitudinal structures used in this manner are practically without anterior stiffening. That is, fishes with spines in the dorsal or anal fins do not undulate those fins. In such examples where spines are present they are entirely free from the rayed and undulating part of the dorsal, as in Batistes. However, in this case there is a slight thickening of the anterior rays which is discussed below. In addition to fishes using longitudinal structures in an anguil- liform manner, others, chiefly with short fins, use them in what we may consider an ostraciiform manner. Lagocephalus and Spheroides often so use the dorsal and anal as a unit, simply flapping them from side to side. This may be called the tetraodontiform movement. (Fig. 55 A). These fins may be thought of as an ostraciiform tail in two parts moved slightly forward dorsally and ventrally. The slight divergence from straight ahead of the reaction, owing to the positions of these fins, is, of course, compensating. In those used in this manner there is a tendency to a thickening or stiffening of the anterior part of the fins. 188 Zoological N. Y. Zoological Society [IV; 5 Fig. 49. The anguilliform model. Side and plan views. Between the anguilliform and the ostraciiform extremes there are intermediates, although it would serve no useful purpose at this point to consider any as carangiform. The thickening of the anterior ray of the dorsal and anal of Batistes shows a tendency toward an ostraciiform fin movement, and indeed not infrequently does it flap these fins as units instead of undulating them. This thickening of the anterior part of such fins seems also to be associated with angular divergence from the axis of motion, whereas in such forms as Amiatus and Gymnotus , as previously noted, in which the longitudinal propellor is practically parallel to the axis of motion there is no such thickening. 1926] Breder: Locomotion of Fishes 189 Caudal Undulation: While the caudal is operated primarily by the action of the body muscles, many fishes at times may be seen to undulate the tail fin vertically and move forward slowly without any other apparent effort. These waves travel at right angles to the undulations of the longitudinal structures and might be expected to raise or lower the posterior part of the fish. Observa- tion shows that such is not the case, other factors entering which translate the thrust into a forward one. One difference is that the rays are convergent instead of parallel. Considering each ray separately, in waving from side to side, it naturally has a forward reaction of the ostraciiform type. The fact that about one half the number operates in an opposed manner to the other half causes a double effect which obviates the necessity of a head oscillation. This may be roughly likened to the squeezing of an orange pip between the fore finger and thumb, the considerable force attained by pressure on its inclined sides being not altogether different from the pressure of the alternate rays. Of course, in the latter the rays are not directly opposite and a solid and a fluid are considered in- stead of two solids, the efficiency being much less, but this may help to visualize the effect. What little thrust of a similar nature acts. to press Amiatus downward and Gymnotus upward on account of their dorsal and anal fins (with parallel1 rays) respectively, is inconsequential because of the difficulty of moving a streamline form at right angles to its intended direction, just as the little vertical thrust derived from the present method dissipates itself. A slight twist of a fin, practi- cally imperceptible to the eye, would readily offset any such action. Du Bois-Reymond (1914) states that fishes often obtain a thrust from the tail by moving the inferior border to one side and the superior to the other. This is not quite the same as the above described undulation. The central ray of the tail remains still and simply twists on its axis in the latter, while in the former all rays pass through a complete cycle of oscillation. In the method described by Du Bois-Reymond, the upper part of the tail is given a backward thrust diagonally to one side, while the other simulta- neously is given a similar one to the opposite side. This has been observed well in Roccus , Epinephelus, and a few others, but appears to be much more rare than the previous method. By a slight stretch, these might be thought of as anguilliform and ostraciiform 190 Zoologica: N. Y. Zoological Society [IV ; 5 respectively, and intermediates found, but it would seem to serve no useful purpose to so designate them or to create special terms for them at this time. Pectoral propulsion: The pectoral fins in many fishes are used partly, and in some almost exclusively, for propulsive purposes. Such forms that have comparatively narrow pectorals used to this end, as Abudefduf and Teuthus, may be, in one sense, almost thought of as rowing their way along, in that they bring the fin forward almost edgewise and force it back broadside. However, a simple synchronous flapping of the pectorals produces a forward component similar to that of the ostraciiform tail. This is often seen in such broader finned forms as Scams and Tautoga, and may be called the labriform movement. If such a pectoral be thought of as an ostraciiform tail moved forward, it is clear that its flapping would drive the fish forward and curve it to the opposite side. The other pectoral would deflect it oppositely and equally with the resultant reaction directly forward, as observed. The proof of this is that by turning the tail of the ostraciiform model so that it flaps against one side of the hull, thus approximating a pectoral of this sort, that end moves forward and curves as above described. Other species, as Spheroides, undulate their pectorals in a manner similar to the way in which the tail is sometimes employed, as described under “ Caudal undulation.” In this case also these fins may be thought of as two forwardly displaced caudals whose angular divergence is complementary. This may be called the diodontiform movement. In actual practise, the movement is complicated by the fact that often the two above described methods are used simultaneously, combining the force of the first with the smoothness of the second, or an intermediate movement may obtain. Many other minor movements are made by the pectorals of various fishes, but all have the preceding elements fundamentally. The following may serve as an illustration. Cichlasoma has been seen to produce a forward reaction by bending over both edges of the fin and beating them up and down together as indicated in Fig. 50. This causes two diagonal thrusts with a horizontal resultant. Jet Propulsion. That the force of the water exhaled from the gill orifices of fishes has a reactance which of necessity must be of some aid in 1926] Breder: Locomotion of Fishes 191 Fig. 50. Diagram of the pectoral movement as sometimes seen in Cichlasoma. A, side view. B, section. Both dorsal and ventral edges are turned over and beaten in unison producing divergent components with a forward resultant as indicated by the arrows. sending them forward there can be no doubt. The amount of this force and its importance as a locomotor asset varies greatly, be- coming large in certain forms and falling to nearly nothing in others. The underlying physical principles that are of advantage in placing slits in such a position as commonly seen has been applied to practical purposes by an engineer. These have been described by Breder (1924)4, parts of which may be restated here as follows: Mr. Clifford M. Paxton has invented and claimed patent rights on a strikingly novel method of propelling ships which he calls an “induced streamline system. ” The following brief con- sideration of this propulsion method is necessary to a proper under- standing of some of these factors of fish propulsion. The movement of a vessel is chiefly impeded by three obstacles, as described below: (1) Inertia resistance. When a ship is propelled through the water all the submerged surfaces of the entrance section are subject to the adverse pressure of the relatively stationary water which has to be forced out of the way to make room for the advancing hull. This water is projected away from the ship and represents lost energy. As the ship moves, other water has to flow in by gravity to fill up the space the hull moves out of. At low speed this is not a serious matter, as the water is moved slowly and has ample opportunity to readjust its level by gravity flow. At higher speeds, however, this is not the case and the water is “pushed and piled up” in the form of a bow-wave about the entrance portion of the ship. (2) Cavitation. Likewise, in reference to the after portion of the ship, water can not flow fast enough in response to gravity to fill in instantly the space vacated by the ship, and there result hollows or “low pressure areas” about the run or after portion of the hull. When a ship is propelled by the rearward pro- 4 This paper was the first published account of Mr. Paxton’s invention. 192 Zoologica: N. Y , Zoological Society [IV; 5 jection of water from the stern the adverse effect of this cavetation is consider- ably increased. The rear low pressures augment the retarding effect of the high pressure created so that there results a pressure differential with a large rearward component, which is the dominant limiting factor in the speed of ships. (3) Skin Friction. The frictional resistance between the ship’s surface and the water through which it is forced, is not a limiting factor, and does not increase in the same ratio to speed as the pressure differential does; it is, never- theless, an important item of resistance. The first two of these resistance factors are as great or greater in the case of a fish, although the adverse pressures are not ordinarily visible as waves on the surface. With an apparently simple but nevertheless very ingenious arrangement of “developed” jets, Mr. Paxton has greatly reduced the retarding effect of these factors and at the same time has been able to develop sufficient reactive thrust to overcome the remnants of resistance that still remain. The invention has progressed beyond the theoretical stage, so the inventor's actual model will be described in lieu of a necessarily longer exposition of the abstract principles. This model is nearly an exact reproduction of a modern destroyer, Class 186, reduced to an overall length of thirty-four and one-half feet on a scale of 1 to 9. On each side of the hull, midway between the waterline and the keel, a short distance aft of the bow a rearwardly directed nozzle is located, so placed as to cause water expelled therefrom to sheathe the under-water hull a short distance aft of them, this sheath completely surrounding the hull at midship. The position of the intake orifice is of slight importance, as long as it is always submerged, it usually being placed where most convenient and presented forward. At this point certain principles concerning the behavior of jets may be mentioned. Contrary to popular belief, the water set in motion surrounding a stationary submerged jet moves slowly in at right angles to the edge of the moving stream and then on contact passes along with it at a velocity only slightly inferior to that of the jet. This induced flow causes more water to move with it in a similar manner, and so on, thus spreading out the stream rapidly. The initial jet increases in sectional area by its deceleration, to which is added the constantly increasing induced flow. Paxton finds that with jets of high velocity the cumulative stream as thus built up may be more than a thousand times the cross section of the initial jet stream. The truth of the foregoing has been satisfactorily demonstrated by experiment. 1926] Breder: Locomotion of Fishes 193 Another fact to be here noted is that a stream ejected along the side of a curved form will follow the contour presented, even if the curve is convex to the axis of the stream, provided it is not too abrupt. The stream does not veer off at a tangent as might be supposed, but closely follows the bending of the curve. With these considerations in mind the manner in which Paxton overcomes the three obstacles to the speed of vessels by his method may be con- sidered. Through the nozzles described he pumps a small quantity of water at a high velocity and neutralizes the three impediments to progress, as will be shortly described. This system of propulsion is not to be confused with many which have appeared from time to time that were based on nozzle reaction and used a large volume of water at comparatively low velocity with the orifice located else- where. All such have been proven to be less efficient than the modern screw propeller. One important difference is that Paxton adjusts, within limits, the lengths of the pressure-reducing and pressure-increasing portions of the jet stream to the respective lengths of the bow and stern sections of the hull. (1) Inertia resistance. This may be lowered slightly by a small amount of water passing into the propelling system through the intake orifice; but this is wholly inconsequential, since possibly less than one-half of one per cent of the water moved away from the high pressure region forward is taken into the ship. The discharge nozzle slits are' so located as to be in advance of the maximum pressure region and by virtue of the water movement induced by the sheet dis- charged through these slits the bow-wave is in practice actually eliminated. (2) Cavitation. The posterior depression is filled in part by the water ejected from the jets, but principally by the flow induced by them, so that the wake trends somewhat rearward instead of forward, and due to the water ex- cavated from between the sheet jets and the hull it has a forward instead of a rearward component. (3) Skin Friction. Considered as resistance to the ship’s motion, this is largely overcome in that a considerable part of the frictional surface is trans- ferred to the surface of the jet stream that sheathes the hull, for this may be considered practically as part of the vessel while it still follows the contour of it'. There is an important addition to friction, however, on account of the high velocity streams being in contact with the hull. The thrust of reaction of the jets on the nozzles is and must be equal to the remnant of resistance which remains after the net resistance reduction has been deducted. The reason for giving the jets a long narrow section instead of a circular one is for the purpose of placing a larger surface area in contact with the adjacent water and sheathing the hull satisfactorily; as well as concentrating 194 Zoologica : N. Y. Zoological Society [IV; 5 the movement, reducing the time required for it, and adjusting the jet stream to the form of the hull. According to Paxton the modern screw propeller is more efficient than his new method at very low speeds, but at relatively high speeds the relationship is reversed.5 Returning to the fishes, it should now be obvious that the water ejected through the gill clefts of typical acanthopterygians is extremely similar in its effect to that of this new mechanical device that has actually propelled a model successfully. It seems, however, almost impossible to obtain an accurate measure of just what locomotor advantage this exhalation of water may be to most species, but it is readily demonstrated to be a real item in others. Brugmans (1812) seems to have been the only naturalist hitherto to have considered the possible effect of exhalations and of course knew nothing of Paxton’s factors. An examination of over 300 diverse species of free swimming fishes taken at random (including both Teleosts and Elasmobranchs), most of which move at a considerable rate of speed, shows that over 90 per cent possess gill clefts at a place which Paxton pronounces to be the theoretically correct position for the most efficient use of this method, as far as he could tell from available material, con- sidering the varying forms. The remainder consist of a few slight variations, none of which are large. Furthermore, even in sluggish forms the gill slits hold these positions fairly closely. It is only in such fishes as have progressed far from the typical ichthyized form that any wide variation is seen. Prominent among the latter are such highly specialized fishes as Hippocampus, Histrio and Lophius. In the fast moving forms long, narrow, gill clefts are found, as typified by Seriola, Scomber and Pomatomus, while in the more sluggish forms small and often nearly circular exhalant pores are present as in Spheroides, Balistes and Lactophrys. Practically all intergrades are found between the two extremes which are beautifully correlated with other locomotor structures and known habits. It is naturally difficult to obtain a measure of the force of exhalant water from living fishes moving at their higher rates of speed and at the lower rates perforce used in confinement, the body 5 All statements concerning his invention have been personally checked and approved as correct in a general way by Mr. Paxton, although naturally sketchy and inadequate for a full understanding of his invention as applied to ship propulsion. They are, however, suf- ficient for our purposes here. 1926] Breder: Locomotion of Fishes 195 Fig. 51. Chilomycterus schoepfii ejecting water through the gill orifices while being held in the hand at the surface. A, the start of an exhalation. B, an exhalation in progress. Some drops that have reached a considerable height are falling at the right. C, the end of an exhalation with the last drops falling at the right. The jets have not been retouched in any way. and fin movements which may be roughly analogized to a screw propeller have a great advantage. However, on the sluggish forms a definite demonstration of this force of the exhaled water is a simple matter, the Tetraodonts demonstrating it most clearly, al- though it must be admitted that here, on account of the rather wide divergence of the exhalant apertures, “nozzle reaction” plays a relatively larger part. Of these, Chilomycterus schoepfii shows its ability in this direction better than any others so far examined. It is simply necessary to hold an individual of this species with its mouth immersed to observe this. Having little flexibility of body, it is unable to squirm about and necessarily confines its attempts to escape to violently lashing the caudal, anal and ventral from side to side and waving the pectorals about in addition to squirting powerful jets of water through the gill orifices. In a fish six inches long these jets may attain a height of considerably over two feet above the surface of the water, (Fig. 51). That these jets are of great use in locomotion there can be no doubt. In fact, specimens of this species have been seen to impel themselves forward through the water by this means alone at not much less than top speed. Other direct evidence was obtained as follows: A six-foot Carcharias taurus was permitted to swim in shallow water so that its back protruded. This caused a “bow wave” and a “posterior cavitation” to form as indicated by the solid line (Fig. 52). Every time the fish exhaled, the bow wave was considerably lowered and 196 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 52. C archarias taurus swimming in shallow water. The solid line represents the bow wave and cavitation formed and the dotted line their reduction on each exhalation. the cavitation more than filled as indicated by the dotted line in the same figure. On removing the pectorals of an Abramis crycoleucas it was found to be unable to rest without some forward motion unless touching some solid surface (Fig. 53) which shows the ex- pedient it usually resorted to. More detailed evidence is given under the section “ Maintaining a stationary position.” As submerged jets of pure water are perfectly invisible, it is only when a suspended particle is acted upon by the jet that its force may be noted. Fishes that were held perfectly rigid so that there was no fin or body movement whatever appeared to be unable to eject jets with any force, simply respiring lightly, but as soon as the slightest tremor was permitted in the body the water was expelled violently. This suggests the possibility of a sympathetic nervous connection between the trunk and tail movements and respiration. Fishes of high speed, such as Caranx and Seriola, which could only be held with difficulty in a manner similar to that described for Chilomycterus, failed to respond appropriately, either flapping violently, or not respiring or if so only feebly, in such a manner that nothing could be deduced therefrom. The powerful adductor operculi, together with the branchiostegal rays and other compressible parts of the head, must make it possible for these species to eject water with a considerable velocity, and it may be mentioned that Paxton main- tains that there is ample water ejected from fish held under such conditions to effect the purpose, the difficulty apparently being one of the velocity of ejection, which the musculature of the opercular and mandibular region could easily effect, were it not for some nervous inhibition incident to restraining the fish. Regarding the considerable strength with which fish can eject water through the mouth under experimental conditions, see Breder 1925 a. This is taken to indicate the great ejaculatory strength of the oral appara- tus, the mandibular and opercular valves controlling only the place of the water’s exit. 1926] Breder: Locomotion of Fishes § 197 Fig. 53. Abramis chrysoleucas with pectorals removed. Illustrating the method by which the effect of the exhaled jets may be overcome, i. e. by dragging the caudal fin. The reduction of skin friction is probably negligible in fishes on account of their effective mucous coat, but the other two and more important obstacles to speed must be overcome by muscular action. Looking at the question from a phylogenetic standpoint there seems to be no very good reason why the port of exhalation in such diverse animals as elasmobranchs and teleosts should have such a community of placement, unless there is a positive advantage to be attained by so placing them or a definite disadvantage in having them placed anywhere else. This might be analogized to the necessarily similar streamline forms seen throughout fishes of high speed. If there was not some sound advantage in ejecting water forcibly it would seem a useless expenditure of energy on the part of many fishes while swimming to pump water in and out when by simply opening the mouth a greater amount would flow over the gill membranes as long as the fish moved forward, because the flow would not be intermittent. Actually, this has been observed in both Carcharias taurus and Anisotremus surinamensis when swimming leisurely. Furthermore, fishes which might be expected to make excellent use of this simplified manner of breathing, such as many members of the Carangidae and Scombridae, have a particularly well-developed opercular apparatus. As the respiration of fishes, however, is an intermittent process it is clear that their mechanism would not be as efficient as a machine giving continuous flow. The exhalation of fishes is not to be con- founded with the simple reactive jets of the so-called syringograde 198 Zoologica: N. Y. Zoological Society [IV; 5 animals which suck up water slowly and expel it with violence through the same or a nearby aperture, such as the cephalopods, medusae and certain Odonata nymphs. This notoriously inefficient method may be compared to the discredited jet propulsion systems of the past. Actions in a Current. The preceding sections, for the sake of simplicity, deal only with forward motion through static bodies of water. If a fish, in progressing in any of the previously described manners, is in a straight flowing current instead of a still body of water, it is clear that in the main, the path traversed would be the resultant of the force and direction of the flow and that of the energy expended by the fish and its direction. However, a complication enters here which may modify the above somewhat, as the action of the current is de- pendent on the shape of the fins and body presented to it and their angle of presentation, as w^ell as the eddy currents consequently pro- duced. These generally tend to deflect a fish from its course and are considered under ‘'Effects of current on maneuvering.” In pursuing a straight line under such conditions, appropriate compensating acts are consequently required to offset these effects for, as the same angles are not always presented to the flow, due to the movements of the fish’s body necessary to locomotion, the force of the current does not continually tend to press the fish in a single direction. Many fishes, however, will suffer themselves to be carried along by the current, especially if of a pelagic habit. In irregular currents, eddies, et cetera, straight line locomotion is more difficult and not infrequently impossible owing to the constant adjustments neces- sary to hold the course. This may be true even if the speed and force of the flowing water is comparatively slight, it being not simply a case of the fish’s muscular efforts being overpowered. Comparison, Interrelation and Discussion. That the three primary methods employed by fishes to induce forward motion, (body movements, appendage movements, and respiration) are all advantageously interrelated is patent. In the majority of fishes, certain phases of each are present and are working harmoniously to the common end of driving the fish forward accord- ing to its requirements. No definite measure of the relative im- 1926] Breder: Locomotion of Fishes 199 portance of each factor has been obtained for any given form although it can be said with little, fear of contradiction that the metameral contractions of the lateral muscles are the most important to the majority of typical fishes, and that most frequently the other two are auxiliary. The appendage movements are especially brought into play for slow progression, and the respiratory jets (according to Paxton) for high speed travelling. There appears to be no timing of the respiration to make it synchronize with the body movements, commonly several swings of the tail being made to one respiratory cycle. However the exhal- tion in fishes of carangiform locomotor tendencies seems to reach its maximum between flexures when the fish is straight forward, which would be clearly advantageous for mechanical reasons. In forms where jet and pectoral fin propulsion are found together, these fins are necessarily kept out of the way of the jets, for efficiency in straight forward swimming. After having analyzed the various expressions of muscular energy directed toward propulsion, it may be well to consider the same from a synthetic point of view, for, after emphasizing the differences, it should be borne in mind how closely similar these different modes of propulsion are in reality. First we considered the body undulations, split up in various ways for purposes of discussion, but actually representing only abbreviate or elongate expressions of the same thing. Secondly we considered the movements of the appendages, which are simply localizations of the same phenomenon and subject to the same gamut of elongation or abbreviation. Thirdly the effects of respira- tion were considered. It thus becomes evident that the means used by fishes to effect movement from place to place are among the most direct to be found anywhere and of great fundamental uni- formity. The simplicity of “blowing” themselves along, coupled with the direct reaction of the primitive metameral lateral muscles, is not to be compared with the comparatively elaborate apparatus of the tetrapods. Even in the most highly specialized of fishes the same elemental movements are to be found, hardly beclouded by the extreme specializations under which they operate. The great expression of undulation, reciprocation and wave motion throughout the swimming of fishes brings to mind Spencer’s “ Synthetic Philosophy” (1892) and his views of the general signif- 200 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 54. Diagram of a fish turning by several methods. Myomeres on one side con- tracted, pectoral fin held out, tail held to one side and exhalation through only one gill orifice. icance of undulation. The philosophic concept of artists, which regards a circle as indicative of place and a straight line as indicative of direction and a sine curve (their combination) as indicating motion, also would seem to find expression in the swimming of fishes. Maneuvering. In general, maneuvering in fishes consists largely of differential application of the locomotor efforts which would otherwise produce forward motion, together with a variety of special acts, most of which are limited to certain forms, rather than being generally applied. Turning. Turns may be made while moving forward by any of the following means: Body movements. (1) Metameral waves may be propagated in greater number or be of larger size on one side than the other, instead of simply alternating uniformly. (2) Several myomeres on one side may be held tightly contracted so as to hold the body flexed while some other effort drives the fish forward (Fig. 54) . Fin movements. (1) A pectoral fin may be held out at an appropriate angle to the body. The further forward the fin is thrust the sharper will be the turn (Fig. 54). (2) The tail may be held rigidly to one side, acting like a rudder (Fig. 54) . This usually accompanies “2” of “Body movements,” as a posterior continu- ation. (3) The dorsal and anal may be held over to one side. (4) Fishes using pectoral propulsion may cease the operation of one fin while the other continues as before. 1926] Breder: Locomotion of Fishes 201 A B Fig. 55. Diagram of a tetraodont such as Lagocephalus turning by several methods. Dorsal and posterior views. A, straight line swimming. (Tetraodontiform. ) B, turning to the left. Pectoral extended, caudal held to one side and dorsal and anal oscillated one side. (5) Fishes using a simple flapping movement of the dorsal and anal may oscillate them both over to one side (Fig. 55). Respiratory action. (1) The operculum of one side may be held down tightly, forcing all the water out through the opposite one. The gill cleft on the outside more or less automatically opens and the inside one closes if the body is flexed to any great extent because of its bending away from the head on the convex side (Fig. 54). These comprise the movements on which turning depends when swimming actively. The side to which the turn is made is obvious and needs no elaboration. Usually more than one method is used at a time and in such sharp turns as is seen in Salmo after having risen to a fly it is likely that as many as possible are used in combination. Fishes which wheel about in great circular paths 202 Zoologica: N. Y. Zoological Society [IV; 5 often use a pectoral alone. Lagocephalus and others may turn very abruptly by backing with one pectoral and oppositely moving the other, with all vertical fins “hard over.” Schlesinger (1911, b) points out with a diagram that in turning by means of extending a single pectoral the pressure of the water on the extended dorsal and anal swings the fish around with the pectoral somewhat as a pivot. This is especially noticeable in long bodied forms, such as Esox. Fishes at rest may face about without perceptibly moving forward by opposite fin movements on either side of the body. That is, such pectoral fin movements mentioned above for Lago- cephalus will practically turn the fish on a point if no forward motion is being made. In such long bodied forms as Esox and Lepi- sosteus , the pelvics may be brought into play also to aid in this action. Many methods are thus employed by various species. Probably the most common method is a single flap of the tail to one side. While this, of course, gives the fish some forward motion, the pectorals are frequently held out as breaks so that no long glide results. In this action the tail is expanded to its full extent in making the stroke and contracted on its return to the median position in order to prevent a reverse turning action or any more forward motion than necessary. Rising and Falling. In rising and falling both fin movements and hydrostatic elements enter, the former as direct acts and the latter rather passively. Fin Movements: As an appropriate movement of the pectorals can deflect a moving fish from a straight line to the right or left so may they by proper twisting cause it to move diagonally upwards or downwards as diagrammed (Fig. 56). This method is effective only if the fish is actively swimming as the fins are held passively up or down, their position as planes, and not any motion of theirs, determining the course taken. The resultant is the product of the inclination of the plane and the horizontal thrust. Fishes not actively swimming may use the pectorals somewhat as active paddles and obtain a similar result by twisting them appropriately and employing methods described under “Pectoral propulsion.” In the case of fishes whose undulations are in the vertical plane as 1926] Breder: Locomotion of Fishes 203 Fig. 56. Pectoral control of elevation. A, pectoral turned to effect an upward move- ment. B, pectoral turned to effect a downward movement. The angled arrows indicate the manner of impingement of the fins. The curved arrows indicate the directions of travel. with skates and flounders, certain of the methods of body undulation used for turning in other fishes cause ascent or descent. If a temporary tendency to rise or fall is not desirable and not readily obviated by the hydrostatic apparatus, the fins may be used to offset it and maintain a definite horizontal plane until an adjustment is made, by using any of the various muscular efforts, that will effect the desired result. Hydrostatic Elements: In most fishes lacking a swim bladder, the specific gravity is slightly greater than that of the waters they inhabit, and immediate although generally slow sinking follows on a cessation of active swimming. In some forms this may be greatly retarded by the expansion of a large horizontal surface as in the case of the triglids with their long plane-like pectorals (heavy although possessing a swim bladder). Others are so close to the specific gravity of the waters they inhabit, due to the presence of sufficient fat, that only the slightest movement suffices to keep them from sinking or rising, as in the case of Poronotus. Prominent among the various fishes lacking a swim bladder may be mentioned Cyclostomata, most Selachii, Scopelidae, Scomber scombrus, Menti- cirrhus, Alepocephalidae, Cyclopteridae, and Pleuronectidae. Both free swimming and bottom dwelling forms are here represented. 204 Zoologica : N. Y. Zoological Society [IV; 5 This lack of a swim bladder in other than bottom forms is found only in marine fishes, the quantity of fat or other buoyant substance necessary to float a fish in the less dense fresh water being practically prohibitive as suggested by the work of Taylor (1922). Obviously any fish permanently or temporarily heavier than water may descend at will in any direction and at nearly any gradient by inclining their fins appropriately, depending on their structures. This is very commonly seen in the triglids although nearly all fishes of high specific gravity appear to use it at times. The vast majority of fishes, however, possess a definite hydro- static organ, the swim bladder. Much controversy has centered about this organ for in many forms it is undoubtedly' used for numerous other purposes, such as a means for sound production or as an auxiliary breathing apparatus. The fact remains, however, that it must of necessity affect the flotation of the possessor. It has been proven to be adjustable to various depths as Du Bois-Reymond (1914) points out and makes a stationary position, with regard to vertical movement, readily possible within certain limits of depth. Various methods by which the gas in the swim bladder is probably controlled have been suggested from time to time and the probability is that nearly all have some truth in them. Although a detailed discussion of the action of the swim bladder is not within the province of the present paper, the following brief consideration may be given it. Even if the bladder be con- sidered as a passive reservoir, the pressure of the superimposed column of water above would tend to compress it to a certain extent, proportional to the depth, but as it is well protected by its place in the fish’s body, this effect is probably comparatively slight. In accordance with this is Du Bois-Reymond’s corollary that a fish with a specific gravity of unity at a given point will descend with increasing rapidity if once started, on account of the slight com- pressibility of water and the rapidly increasing superimposed weight of water (one gram per sq. cm. for each cm. of depth) causing compression of the gas and a consequent increasing specific gravity differential. The opposite is the case in rising. It is conceivable that voluntary muscular control of the size of this organ and conse- quent variation in the bulk of its contents might be used by some species to cause a rising or falling of the fish. In changes of con- siderable depth the secretion or reabsorption of gas by the blood 1926] Breder: Locomotion of Fishes 205 no doubt plays a prominent part in stabilizing the animal and moreover is acting reflectively at all times with the purpose of keeping the fish at the desired depth. That these various factors are always working to maintain the proper specific gravity is clear if it be considered that in the normal life-processes of food taking, digestion and excretion, the bulk of any fish is practically incessantly changing slightly, and small adjustments must be made continually. The following experiments demonstrate that in Fundulus heteroclitus and majalis, at least, both of which possess well developed swim bladders, the adjustment in flotation to different densities is not instantaneous. One dozen of each species was transferred as follows from salt to fresh-water and vice versa: TABLE II Exp. No. Transferred from 1 Salt water sp.g. 1.024 2 Fresh water sp.g. 1.000 3 (No. 1 back to salt water) 4 (No. 2 back to fresh water) Transferred to Fresh water sp.g. 1.000 Salt water sp.g. 1.024 Result Sank Rose Rose slightly. Sank slightly. From this table it is evident that the reactions are the result of an adjust- ment that is not immediate. In Nos. 1 and 2, all the specimens occupied the bottom and top halves of the jars respectively, and it was clear from the pectoral movements that in the first case these were directed to drive them upward and in the second downward. In between five and ten minutes, they began to distribute themselves more equitably, but there was still a slight difference between the two jars. By fifteen minutes, the two sets were indistinguishable. On transferring back (No. 3 and 4) a similar lack of immediate adjustment was evident, but not nearly so great, from which it is judged that the first adjust- ment was not as complete as it first appeared. Starting and Stopping. In starting ordinarily the movements of swimming are simply initiated, as pointed out under descriptions of the various types of locomotor efforts. However, the pectorals, usually held out from the body when resting, are quickly brought back flush with the body and remain there except in the pectoral type of locomotion. A violent exhalation may be made at the same time also. These actions, of course, give an added impetus to the initial effort. Fishes may come to a stop gradually by simply ceasing to operate the locomotor musculature, in which case they glide for a 206 Zoological N. Y. Zoological Society [IV; 5 Fig. 57. The effect of the relative positions of the fins on stopping. A, manner in which the hind end of Esox rises if pectorals only are used as brakes, unless checked by the pelvics as shown. B, manner in which an abrupt stop is made by Esox with both pectorals and pelvics used as brakes. Note that approximately A: B : : X : Y. C, Comparison with Vomer . Note that here too approximately A : B : : X : Y, but the pectoral is much nearer to a central location, obviating the necessity for the large posteriorly placed pelvics. considerable distance before coming to absolute rest. Most fre- quently, however, they employ some muscular activity to neutralize the no longer desirable motion initiated by other contractions of muscle fibers. Anguilliform fishes may suddenly reverse the direc- tion of the motion of the undulations and by this means quickly check forward movement or they may suddenly hold themselves rigid in any position and check the motion more gradually. Fishes with such undulations localized in the fins accomplish the same 1926] Breder: Locomotion of Fishes 207 result with their particular locomotor structures in a similar way. This can be readily observed in Amiatus and Gymnotus, but Raja seems never to employ a reverse movement although it may rigidity itself. All fishes which have been noted to reverse their primary locomotor “gear” have also been noted to deliberately travel, at least short distances, backward by this means. Gymnotus and related genera are the most expert in this performance of any of the forms studied. There are good mechanical reasons why Raja could not do it very effectively, judging from its shape. Fishes of a carangiform tendency in locomotion are not known to show a reversal of the primary locomotor movements, the pector- als playing an important part instead. These are usually stuck out equally on both sides, thus forming an effective resistance to forward movement. In such long bodied forms as Esox it is clear that if the low hung pectorals were simply dropped, while they would stop the fish, they would likewise “trip it up,” i. e., cause the caudal portion to raise up because the point of pressure is anterior and low (Fig. 57 A). In order to avoid this the pelvics which in these forms are placed well aft are held as indicated in the figure which, because of the close similarity in size and shape of the two sets of fins, counteract the tendency of the tail to rise. This expedient is resorted to in an attempt to make a slow stop. If an abrupt stop is desired both pectorals and pelvics are thrown down simultaneously at approximately the same angle, thus producing two points of resistance, one forward and the other aft, which causes no tendency for either end to raise. (Fig. 57 B). In short bodied fishes this tendency to tip up is materially lessened on account of their short fore and aft dimension and a corollary to this is the fact that the pectorals are closer to the middle of the body, from front to back as well as from top to bottom (Fig. 57 C). With this may come a reduction of the pelvics as in Vomer, or the pelvics may remain large as in Lepomis. The former condition is found typically in forms which swim continuously for long periods, as pelagic fishes, e. g. Poronotus, while the latter condition is typical of lacustrine, fluvitile and reef fishes which habitually stop frequently in their locomotor efforts, e. g. Lepomis, C}iaetodipterus and Pomacanthus. Roughly, the tendency connected with a shortening, deepening and narrowing of the body is for the pectorals to become more nearly centrally located, antero- 208 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 58. Micropterus with the dorsal and anal lobes curved to one side and the caudal to the other so as to form a “sea anchor. ” posteriorly and dorso-ventrally, while with a lengthening and widening of the body the pectorals tend to move forward and down- ward and the pelvics backward and downward and to approach them in size. Generally speaking, approximately A : B :: X : Y (Fig. 57 B and C). As in nearly all of the modifications mentioned in this paper, a complete series of variations is found grading from one extreme to the other. Acanthopterygians that have the bases of the pectorals and pelvics usually placed rather close together and are comparatively long bodied also usually have the tips of the dorsal and anal lobate. These are frequently brought “hard over” to check movement. The genera Micropterus, Epinephelus and Micteroperca quite commonly employ the tail in this manner, virtually forming a sea anchor. Any tendency caused by this to bring about undesirable deflection to one side is counterbalanced by an appropriate pectoral movement. Lepomis and Micropterus at times curve their tails to one side and the dorsal and anal lobes to the other, thus checking movement and avoiding deflection (Fig. 58). As many fishes may eject water through the mouth, it follows that there is some slight reactive effect which probably in most cases, however, is negligible and rarely used. The only direct evidence we have of its effect is that to be noted by watching Batistes. This genus, especially B. carolinensis, with its mouth a few inches away from sandy bottoms will repeatedly blow water out excavating a small hole in the sand in its search for small worms and other burrowing forms, habitually seeking food in this manner (Breder 1925 a). The force of this jet, directed against the bottom is indicated by the way the fish braces its pectorals, undulates its 1926] Breder: Locomotion of Fishes 209 dorsal and anal and so on, although even then, at times, a slight backward recoil is noticeable. In stopping, the dorsal and anal of most spiny rayed fishes are usually erected to their fullest extent. That these fins are chiefly concerned with the maintenance of an even keel may be proved by the removal of them. Fishes of this type so deprived are able to swim practically as fast as before, but roll slightly from side to side with each half cycle of undulation, especially when starting or coming to a stop. Once well under way, this twisting largely disappears. Rapidity of motion continuously made, without accel- eration or deceleration and absolute rest obviates to a certain extent the necessity of these keels which are then ordinarily folded back. See under “Reomorphism. ” The pelvic fins further this same maintenance of position by functioning after the manner of bilge keels. This is especially prominent in many of the sharks with the long and evenly outstanding pelvics. Maintaining a Stationary Position. The maintenance of stationary position is by no means a simple matter with fishes which do not merely rest on the bottom or hold fast to some object by means of prehensile or suctorial specialization. Allowing the hydrostatic apparatus to be perfectly adjusted so that there is no tendency to rise or fall, which condition is frequently approached, and that the fish is in perfectly still water there are still other factors to be reckoned with. It has been argued by Osburn (1906) and others that the continuous movements of the paired fins while a fish is at rest are for the purpose of maintaining equilibrium. The experiments of Osburn were performed on Fundulus heteroclitus from which he claimed to have found the following: “. . . When a single pectoral fin was removed the fish tended to turn partly on one side, due probably to the action of the pectoral of the opposite side. This, however, the fish soon learned to regulate. After the removal of both pectorals the fish when swimming slowly apparently moved as usual, but when forced to turn quickly it was unable to accurately balance or otherwise undergo movements requiring nice adjustment. ... A study of the movements of many species of fishes in the New York Aquarium is entirely confirmatory of the view that one function of the pectoral is to balance and accurately adjust the fish in swimming. . . . Fishes with the pectorals removed would at first frequently run 210 Zoologica : N. Y. Zoological Society [IV; 5 against the side or bottom of the tank, but later they learned to avoid this by a strong movement of the tail. . . . , it is a point of observation without a single exception in my experi- ence that the ordinary, actively swimming type of fish when resting on the bottom does not move the fins at all. ... On the other hand, all fishes that I have observed use the pectorals when they are suspended in the water. Moreover, other fins are often brought into use at the same time. Thus the elongate pike ( Lucius ) and gar ( Lepi - sosteus ) are seen to move the pelvic fins slowly, coordinately with the pectorals, and short bodied forms such as the butterfly fish ( Chaetodon ) move the pectorals and caudal, while in species intermediate in form the caudal, anal and dorsal may, any or all, be used in addition to the paired fins when suspended in the water. This array of facts makes it quite clear that the function of the pectorals when the fish is stationary is that of equilibration and not the removal of water charged with carbon dioxide.” This latter statement refers to Duges (1905) who contended that the motion of the fins was to cause a current supplying fresh and removing vitiated water from the gill region. How the fish managed to remain in a stationary position if freely suspended while doing this appeared not to concern him. Osburn summarized the functions of the pectorals as follows: “Guiding and balancing the body in swimming; To act as a brake in arresting the progress; Equilibration when suspended stationary in the water and Locomotion, either forward or backward.” While Osburn’s work is here accepted in general, it is objected that the balancing and equilibrating function upon which he places so much stress is of minor importance and almost negligible as the subsequent evidence will serve to indicate. In spite of the various functions ascribed by other students6 to the movements of the fins while at rest, ranging from that of supply- ing new water to the gills to the more rational one quoted in part above the following is offered in substitution. Fishes in respiring eject water through their gill orifices in a backward direction as pointed out under “Jet propulsion.” It is axiomatic to state that if a jet of any sort whatsoever be ejected at a velocity however low from a body freely suspended, there will be a proportional reaction in the opposite direction. Therefore, if a fish is not to move forward at a speed dependent upon the rate 6 Since this manuscript has been prepared, a paper “On the Functions of the Fins of Fishes” Schmalhausen (1916) has been received. It is in essential agreement with the con- clusions presented here and discusses certain factors in more elaborate detail accompanied by six interesting diagrams. The effects of respiration are not considered. 1926] Breder: Locomotion of Fishes 211 and force of respiration when resting, some counteracting effort must be made. The following is given in support of this view and is taken in part from Breder (1924). Over a dozen diverse species were experimented upon by the removal of some or all of the fins; the species including those used by Osburn and others and varying in form from such extremes as the log shaped Esox reticulatus to the deep, thin bodied Vomer setapinnis. The results obtained were an embarrassment of move- ment, depending on what fins or combinations of fins were removed, but in no case was a distinct disturbance of equilibrium obtained. As long as the individuals remained at rest and attempted no turning or other maneuvering they retained the normal horizontal position. The present experiments demonstrate that the stability of the equilibrium of fishes is not controlled by fin action, general statements notwithstanding. True, dead or very sick fish often float belly or side up, but this is no doubt attributable to other more or less obscure physiological causes: derangement of the digestive tract, lack of control of the gases of the swim bladder, et cetera. The centers of gravity of numerous free swimming species were found to fall within the air bladder. This was found by balancing them in two planes on a knife edge.7 Various comparisons made in and out of water showed that taken as a whole these fishes could be considered as being of virtually uniform specific gravity, even though constructed of many different and diverse substances. This is not in accord with the generally accepted belief that fishes are in an unstable equilibrium normally, which view has been based on injured or diseased examples and their clearly unstable equilibrium. Fig. 59 illustrates the position of the center of gravity with reference to the swim bladder in four species. An inspection of this will show that if the body be considered as of uniform specific gravity, the center of gravity of the entire fish and that of* the swim bladder itself practically coincide. This clearly makes for an ease of motion from side to side as there is no great pendulum action to rapidly erect (or turn over) a fish, and partly explains the ease with which certain labrids, scarids, and others swim on their sides at times. Mr. E. C. Bennett points out in this connection that it is his belief that the center of gravity and the center of buoyancy would be found to coincide in most fishes. 7 See Appendix, page 293 for a description of the apparatus used in this connection. 212 Zoologica : N, Y. Zoological Society [IV; 5 Fig. 59. Location of the center of gravity in fishes. The cross lines indicate the posi- tion of the center of gravity. The shaded area indicates the position of the swim bladder. A, Chipea harengus. B, Fundulus majalis. C, Aplodinotus grunniens. D, Eupomcentrus leucosiictus. 1926] Breder: Locomotion of Fishes 213 A superficial glance at nearly any typical fish poised quietly in the water should convince anyone that the pectorals are engaged in backing water. That is, the effective thrust is forward, which would tend to move the specimen in a backward direction, or just the reverse of what is described under “Pectoral propulsion.” However in connection with the effect of the exhaled water it is conceived that the function of this movement of the pectorals is to neutralize its reactive force. This backwardly moving inter- mittent stream of respiration is not to be understood, however, to have any considerable force, while the fish is at rest, as naturally the respiration is slower and furthermore the gill clefts are observedly opened wider at such times, thus reducing the velocity of the emerging streams as well as increasing the cross-sectional area of them and consequently reducing both their velocity and surface area per unit of volume. The pectorals being usually placed directly behind the gills are enabled to intercept the stream and check the original direction of the thrust. The truth of these assertions is by no means simple to demonstrate, owing to the large number of locomotor organs that generalized fishes employ either singly or in numerous combinations. In fact, it is rather seldom that fishes are seen free in the water with no apparent motion other than the pectorals. At such times they are seen to back water rhythmically and usually in perfect synchronism with the respiratory movements. That is, as the pectorals come forward the operculum lowers and forces the vitiated water out, the inhalation accompanying the return stroke. Usually, however, there are some other movements such as undulation of the dorsal or caudal as well as various others which complicate matters so that it is a matter of patient waiting for a proper opportunity to see these two factors working together alone in direct opposition to each other. The significance of these other motions is treated under the heading “Relationship to habits and development. ” In very cold water when Chaenobryttus is in a state of semi-hibernation they may sometimes be observed with no fin movement at all and with both sets of paired fins securely pressed against the sides. At such times the respiration is extremely superficial and appears to have little reactive effect. However, after a long period of time it was observed that they do move forward ever so slightly and that very occasionally the pectorals are brought into play to regain the previous position. However, they generally allow themselves to come in contact with some piece 214 Zoologica : N. Y. Zoological Society [IV; 5 Fig. 60. Diagram of the behavior of a Centrarchid ( Lepomis pallidus ) minus one pectoral. In attempting to maintain a stationary position the backing of the pectoral was insufficient and the fish curved forward toward that side. An increased pectoral action caused it to curve backward to the opposite side. Later the dorsal lobe was brought into play and a stationary position was maintained. of brush or other object in which case the friction is sufficient to stop motion. The removal of a single pectoral from a specimen of Lepomis pallidus demonstrated this still further. On composure after release it backed water as usual with the remaining pectoral fin but as the force applied was only one-half of that previously used and on one side only, the fish moved forward, slowly curving toward the side possessing the fin. This motion appeared to disturb the specimen, causing it to speed up the number of oscillations. As now the force of the fin overcame that of the jets the fish moved slowly backward and curved slightly to the opposite side. In a short time the fish learned to compensate for the missing member by waving the posterior tip of the soft dorsal which it bent towards the side of the missing pectoral, and from then on had no difficulty in maintaining any position desired (Fig. 60). Most of the Centrar- chidae use either or both median fins in this manner occasionally, making the learning of this accomplishment no new feat. On this account it is usually done by specimens practically immediately on coming to rest, the particular individual mentioned above probably representing a slight abnormality in nervous adjustment. The removal of fins from Fundulus heteroclitus yielded the 1926] Breder: Locomotion of Fishes 215 Fig. 61. Cross-section of Fundulus heleroclitus, indicating maximum roll of Unless examples. The solid lines indicate the normal horizontal and vertical axes. The dot and dash lines indicate the .extreme angular displacements of the vertical one. following results. Compare with those of Osburn quoted on pages 209 and 210. These observations are given with reference to equi- librium only, as at no time were the specimens sufficiently quiet to observe the effect of their exhalations. One pectoral removed. — Very slightly inconvenienced, movement almost as in the normal fish. Turns toward the side of the missing member were made with difficulty. Two pectorals removed. — A slight tendency to roll while swimming straight ahead was noticeable. Any turning difficult. One pelvic removed. — Even less inconvenience experienced than with the removal of one pectoral. Two pelvics removed. — A slight tendency to roll was noticed, but much less than with both pectorals missing. One pectoral and one pelvic (same side) removed. — A slight tendency to roll was again noticeable. Turns were negotiated with difficulty as were changes in level. Two pectorals and one pelvic removed.— A greater tendency to roll was noticed. On release the fish dove to the bottom and at first had considerable difficulty in rising. The following day all were up and active having accommodated themselves to their various handicaps. Fig. 61 shows in cross section the greatest amount of angular swing noted in any. This could be hardly construed as a turning ‘‘belly side upwards” as mentioned by Bridge in the Cambridge Natural History (1904). Further, this diagram is somewhat exaggerated, and the rolling became progressively less as the specimens became adapted to circumstances and learned better to control themselves with dex- terous twists of the remaining appendages. Further experiments on other species are given below: Lepomis pallidus. — One pectoral and one pelvic (same side) removed. The r 216 Zoologica: N. Y. Zoological Society [IV ; 5 action was indistinguishable from that described previously with a single pec- toral missing. Two pectorals removed. This fish failed to stop forward motion except when resting with the pelvics in contact with something, rolled slightly in swim- ming and could not turn well. Two pelvics removed. Did not stop as abruptly as a normal fish. Not inconvenienced while at rest unless in a slight current, when a wabbling motion was noticeable. Abramis chrysoleucas. — Two pectorals removed. Could not stop forward motion except by dragging the tail along the bottom, thus resting in a diagonal position. See Fig. 53. Two pelvics removed. Only very slightly inconvenienced in turning and stopping. Esox reticulatus.— Two pectorals and two pelvics removed. Swam and turned more awkwardly than a normal fish. Bent dorsal and anal and so maneuvered while nearly at rest. Rose and fell fairly well. Two pectorals, two pelvics, dorsal and anal removed. Swam with dif- ficulty. The median fins seem to have a distinct propulsive function, being placed so far aft and automatically sharing in the oscillation of the body, as pointed out by Schlesinger (1909). Maneuvering power greatly restricted. Did not remain perfectly still as is their habit. Dagodon rhomboides and Eupomotis gibbosus. — One pectoral removed. Dorsal lobe bent around and compensated generally at once. Vomer setipinnis. — Two pectorals removed. Unusually little inconveni- ence noted. In a tank of normal examples it could not be distinguished for any abnormal behavior, although this species constantly waves its pectorals about. Scardineus erythropthalmus. — Pectorals held down tightly to body and operculums held closed by a band of very thin rubber. Short stops not made successfully, the ventrals being insufficient brakes. Turning not so gracefully or accurately done. Effects of Current on Maneuvering. Maneuvering in a current has much greater complication than straight forward swimming under such circumstances. That is, a current may generally either help or hinder maneuvering in a more marked manner than it does rectilinear locomotion. Turning, rising and falling are affected either advantageously or adversely depending on the angle of the flow in relation to the direction of the changing axis of progression. That is, in a turn produced by any of the aforesaid methods the current may so impinge on the curves of the body, the projecting fins etc., as to hasten the turn or retard it. This, of course, is also true of altitudi- nal changes. 1926] Breder: Locomotion of Fishes 217 Fig. 62. Maneuvering in a current. The direction of flow is indicated by the arrows. A, normal mean position in swimming actively against a current. B, and C, either side of a carangiform oscillation from which active swimming is started to regain a lost position. Starting, stopping and the maintenance of a stationary position is somewhat more complicated however, and may be considered briefly below. In holding a position in a current, fishes head up stream as this minimizes the frictional resistance of the flow against their bodies by reason of their streamline shape; and in this position they are properly orientated for swimming against it. Here the effect of respiration aids in holding their position and need not further be considered in this connection. A nearly stationary position may be held simply by actively swimming against the current just fast enough to balance the effect of the water in carrying the fish down stream, i. e. by swimming just as fast in one direction as the water flows in the opposite. This method may be observed in a fish culturist’s trough of salmonid fingerlings. A fish so maintaining a stationary position in a current may desire to drop back to a position further down stream. This is generally done, not by turning, but by simply stopping the swim- ming movements and drifting passively back. The drifting back is generally not done in a position with the body axis parallel to the current, but at an angle to it equal to the amplitude of the propulsive contortions (Fig. 62), in which A represents the mean position, parallel to the direction of flow, and B and C the fish at either side of an oscillation (similar to A and C of Fig. 44). These latter two are the positions held for drifting down stream. Starting from one of these, as is frequently done to regain the original position, the fish is already in the best possible position to initiate motion against the stream flow, instead of having to 218 Zoologica: N. Y. Zoological Society [IV; 5 make that first flexure from the less effectual position directly in line with the flow. See under “ Body movements. ” In the ordinary distances so drifted there appears to be very little change of this angle due to the action of the current. The speed that a fish facing a current can sometimes attain is surprising, as its actual motion is, of course, the sum of its observed travel plus that of the stream flow. See Stringham (1924). Comparison, Interrelation and Discussion. It should now be evident that all the various elements that go to make forward progression possible are likewise involved in maneuvering, together with numerous others not directly concerned with straight line locomotion. The “propelling devices” and the “steering mechanisms” are so inseparable that it is generally impossible to say where one leaves off and the other commences. Maneuvering in the main is effected by differential applications of the motor system generally; body movements, fin movements and exhalations, all being capable of use in maneuvering as well as in driving the fish forward. However, in more generalized fishes the metameral muscles of the trunk may be roughly compared to the driving engine and the appendages to the steering apparatus, in a very broad sense. The innumerable interrelations of the many factors concerned with orientation and nicety of movement makes the isolation and description of any one of them particularly difficult for it is sel- dom indeed that one is to be seen operating alone. The removal of members while giving collateral evidence is not completely satisfactory in itself. Observation of many individuals of varied species under normal and abnormal physical conditions acting in a manner slow enough for the eye to follow has been found in many cases, after much experimentation to furnish the most reliable data, when studied with the axioms listed on page 165 in mind. Movements Other Than Swimming. Nearly all fishes are primarily adapted to locomotion while freely suspended in water and all factors treated under the present head may be considered as secondary adaptations to special environ- ments and habits. 1926] Breder: Locomotion of Fishes 219 Burrowing. One of the common habits of many fishes not properly to be considered as swimming is burrowing. This is generally associated with the strict anguilliform type, but not necessarily, e. g. many labrids, such as Iridio, are capable burrowers. Burrowing is gener- ally effected by an active swimming with the nose pointed into the sand. This is continued until a sufficient length is covered to allow the various muscular contractions to obtain a grip on the sand, after which progress is more rapid, the tail portion then frequently trailing into the burrow passively. Other forms such as skates and flounders generally throw the sand over themselves, not bur- rowing in the sense described above, while still others employ somewhat intermediate methods. Creeping. Many various groups are modified for motion along a solid surface, generally only in a nearly horizontal plane. The triglids, for example, are able to creep slowly over the bottom by means of the free pectoral rays. The motion is produced by placing the tips of the rays in contact with the bottom and pushing backwards. This movement may be simulated by running the hand over a table top with three finger tips touching it and moving somewhat as legs. There seems to be no especial sequence in which the rays are used. Fishes with a suctorial disc such as the Petromyzontes, Gobio- formes and Loricariidae, all seem to be able to adhere to any surface almost independently of its angle of inclination, providing it is of the proper texture. By slight movements of these discs they can inch their way along slowly for short distances, in various degrees of effectiveness depending on the structure of the disc. Flounders may push themselves forward by means of the fringing rays and skates can kick back with their modified ventral fins. See “Part II — Systematic” for further details of this sort. Anguilliform fishes of nearly cylindrical cross section can move over solid surfaces out of water by applying their particular locomotor movements. In such a case the contact with the sup- porting surface is the only place of effective pressure, the air being so tenuous as to be entirely ineffective as a resisting medium to these movements. Therefore the oscillations are of wider amplitude, as it is natural that they would be and indeed need to be, for this 220 Zoologica: N. Y. Zoological Society [IV; 5 manner of movement is very inefficient. Advantages not dissimilar to those to be had in the case of burrowing are found with eels moving through grass or over rocky places on account of the in- creased contacts. A fish of carangiform locomotor apparatus out of water flops from side to side because in contracting the myomeres of one side as it normally does, the tail hits down smartly and the reaction throws the fish upward. Incidentally in this connection experiments were made upon Fundulus heteroclitus in an effort to determine whether they could direct their flipping movements out of water and so find their way seaward if left stranded on a beach as is claimed, Mast (1915). The results of the present experiments were as follows: If the surface (in full view of and near to open water) is perfectly smooth and level, the fish simply hop up and down “progressing” indifferently in any direction and often coming back to the same point. If it is slightly tipped in any direction, the vast majority move down the incline as would be expected for mechanical reasons. Similarly if a strong wind is blowing they tend to move with it. If a small pool is made in the sand and a fish placed in it, it stays there until the water sinks through the sand and leaves the fish dry, after which it starts to flip about apparently aimlessly. However, if a slight trickle is allowed to enter the pool, it heads up stream and tries to wriggle out of the pool if even the stream is too small to float it. In its efforts to make progress, it often makes short leaps, usually falling back in the stream again, and continues to show this simple positive rheotaxis as the only directional influence until open water is reached. In the case of a stream flowing out of a shrinking puddle a negative rheotaxis is sometimes evident. As these experiments were not carried on in an identical manner to those of Mast’s, nor as extensive, the only real divergence in results appears to be in the matter of going uphill which none of the present fish did at all. However in the related Rivulus directional locomotion over land is common. Leaping. The leaping of fishes may be considered simply as a rapid swimming up through the surface of the water, momentum alone carrying the fish forward after the tail has left the water entirely. In leaping, the passing into air, a less dense medium, comparatively accelerates the speed and makes possible leaps that otherwise would appear too great for what seems to be slight effort. The active propulsion is obtained in water, a dense medium, with its patent advantages of comparative solidity whilst the glide pro- duced by momentum is in a light medium which has comparatively little resistance. The course the fish takes after leaving the surface is dependent on the manner in which the body is held and is modified 1926] Breder: Locomotion of Fishes 221 by external forces, such as wind velocity and the angle of its direction to the fish. If the body is held rigidly in a straight line the path will be straight, barring external factors. Mugil usually leaps in this manner. If the body is flexed, the fish follows the curve and falls to the concave side. Salmo and Tarpon usually show this. The final stroke before leaving the water is often of great amplitude in these fishes and sends them upward in a great curving path. Flying. Much of a controversial nature has been written concerning the flight of fishes, that is, as to whether they fly in the strict sense or simply soar, and as Clark (1925) suggests there is probably some truth in both views, That is, probably it is largely a soaring flight, although there is a strong probability that distinct wing movements of muscular origin aid them at times, especially in the case of the very young in which the wings vibrate through a relatively large arc with a distinct “hum.” However, the pectoral muscles of the exocoetids are approximately equal to those of the hemiramphids (Ridewood, 1913). At least some of the vibration of the pectorals of large examples is due to the forcing of them, as planes edgewise through the air, and some according to J. T. Nichols, is likely a muscular quiver incident to holding the “wings” out under tension. Ridewood regards the flying characins, Gastropelecus , as more likely true flyers on anatomical grounds, although the observed behavior of Thoracocharax would hardly lead to this conclusion (see page 250 and Fig. 73, A). The wings of the exocoetids are ideally suited for soaring, as indeed is the entire fish, from an aeronautical point of view, as Dowd (1921) points out. Other flying fishes, Pantodon and the Dactylopteridae, are less well equipped and their flights are of shorter duration and comparatively clumsy. Allied to this type of flight is the skimming or skittering over the surface, of hemiramphids, etc. Here the pectorals are only sufficient to raise the weight of the head and forepart of the body into the air, leaving the tail submerged. The bulk of the fish being in the air, reduces head resistance considerably, while the tail still has the advantages of operating in the denser medium. Flying fishes pass through this stage as a transitory one as they leave the water at the beginning of a flight and lapse back into it if their momentum ceases and they do not wish to entirely submerge again. 222 Zoologica: N. Y. Zoological Society [IV; 5 The latter they generally accomplish by a short leap and a head first plunge. When the tail is so submerged it is violently oscillated. This naturally vibrates the entire animal. Being supported in such a precarious manner this is especially noticeable at the tips of the “ wings” on account of their position. This may have been contributary to the belief that the wings are actually flapped in locomotor effort. Such aerial excursions start as a simple leap. Comparison, Interrelation and Discussion. The chief locomotor efforts of fishes other than swimming are represented as specializations, none of which is the common property of a large number of fishes. Further, these various specializations are more or less mutually exclusive. That is to say that burrowing fishes do not fly and leaping forms do not creep as a rule. On account of the narrow limitations of most of these specializations and their, for the most part, evident operation they are only touched on here, the lesser details being discussed in their proper places in “Part II — Systematic.” Reomorphism.8 For the most efficient rapid motion through water a torpedo- like form is essential, varying somewhat in detail with the actual speed. That is, it should be circular in cross section throughout and somewhat cigar shaped. Of course, this ideal is never attained in fishes as there must always be some apparatus for applying the driving force and changing the course when necessary as well as other structures not at all concerned with mobility, such as the defensive and offensive requirements call for. The closest approach to the ideal is seen in oceanic pelagic forms not concerned directly with either surface or bottom, for in animals living close to a plane of division and under the positive or negative influence of gravity a marked dorso-ventral differentiation is present. Du Bois-Rey- mond (1914) believes a sharply pointed anterior end on a fish to be desirable for high speed but points out that it is physiologically impossible, all of which is clearly untrue. He was apparently unfamiliar with the modern studies of streamline forms and the experiments proving that a comparatively blunt nose is the ideal and did not realize the existence of such spear-like forms as Tylosurus and Lepisosleus, not to mention a host of less exaggerated types. 8 First used by Karrer (1924). Defined as designating “the structural adjustment in organism toward streamline contours.” 1926] Breder: Locomotion of Fishes 223 It seems likely, judging from the existing diversity of fish heads, that any desirable entering section could be evolved, and probably those existing on high speed fishes are nearly theoretically correct. See Parsons (1888) for a discussion of the forms of fishes from a mathematical standpoint. The maximum speed that various fishes can attain is largely undetermined, but is considerable in some, especially oceanic forms. Stringham (1924) gives the following estimates in miles per hour for some freshwater species: Salmon, 6.75 to 7.0; Pike, 8.0 to 10.0; Alewife, 6.8. None of these approach the speed of some pelagic forms. The normal speed of a fish, its maximum velocity, the length of time it can maintain it, its locomotor classification, its normal bathometric habitat, its agility, maneuvering, leaping ability, et cet- era, are all reflected in every external structure. Even to the most untrained eye such reflections serve as a fairly accurate gauge of the mobility of an example. With some study and experimentation, a really accurate measure of the method of movement may be visual- ized and even an average illustration will reveal unmistakable indications of locomotor ability. The shape of the body is no doubt truly streamline for the normal speed at which a given fish is built to move, modified accord- ing to the mechanical necessities of locomotor efforts or other requirements (Houssay, 1911, 12 and 14), except where motion is normally so slow that the pressure differential due to movement is entirely inconsequential. Here streamline shape is usually sacri- ficed for armature or some adaptation not directly concerned with locomotion. Some torpedo-like form is absolutely necessary if any relatively high speed is to be attained. Reference to “Part II — Systematic” will reveal instances where, what are apparently anything but streamline shapes, in reality are excellent ones, con- sidering the other necessary factors, and are simply disguised by the peculiar locomotor apparatus of the possessor, e. g. some of the Batoidei. The investment of the body, usually imbricated scales, forms another reflection of the locomotor prowess of the possessor. Primitively they are metameral, being grouped in the same number of vertical rows as are the myomeres. Specialization has changed this in various ways, in some cases removing them completely, as in the Siluridae, where a tough flexible skin replaces them, and increasing them in others as in the Pleuronectidae, where 224 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 63. Diagram of an epibatic tail as in Sphyrna. The diagonal dashed arrows indicate the manner of impingement of the caudal and pectorals in horizontal swimming. The section A-A indicates the swinging of the dependent part of the caudal. there are less myomeres than rows of scales. That is, the scales are placed according to the lines of strain incident to the body flexures (Ryder, 1892). Primarily they were designed as a flexible armor and still generally retain that function although in some cases, such as the Ostraciidae, flexibility has been completely sacrificed for armature strength (See also Woodward, 1893). The mucous cover- ing of the body in most cases doubtless reduces skin friction con- siderably, not only because of its inherent slipperiness, but also because it fills up any small irregularities, such as spaces between the teeth of ctenoid scales, and makes the fish actually smoother than it feels to the hand, because the pressure of the latter passes through the mucus so readily. The position, shape, prominence and sculpturing of the scales all reflect the natatorial nature of the possessor. This is likewise true of outstanding armor of any kind, which is always sacrificed by fast swimming fishes, the slower ones only being able to afford it. The fins as well as the body form, especially the caudal fin, form a very fine index of speed and agility, the same types being repeated over and over again in the most unrelated fishes of similar habits of swimming, being in many cases beautiful illustrations of homodynamic adaptations. Various types of tails are generally considered by taxonomists, such as heterocercal, homocercal, dyphycercal, isocercal and proto- cercal. The fact that homocercal tails are heterocercal internally need not concern us here, for it is the fin as a whole that presses against the water, and its exterior which measures its reaction and not the details of the internal osteology. We may better use the terms epibatic for externally heterocercal tails; isobatic for tails externally symmetrical, no matter what may be the internal structure, and hypobatic for tails with an enlarged and thickened lower edge as in Exocoetus. Whether all tail fins are strictly homol- ogous or in some cases merely analogous need not concern us here either. 1926] Breder: Locomotion of Fishes 225 Fig. 64. Diagram of the action of a bilobed epibatic tail as in Mustelus. A, position of upper and lower lobe to one side of body when they are both on the same side of the crest of one wave. B, position of upper and lower lobe on alternate sides of the body when they are on either side of the crest of the same wave a little later. The primitive epibatic tail (heterocercal) is unsymmetrical about a horizontal plane passing through the axis of the body in that the vertebrae margin it above. As the rest of the tail depends from this it follows that being flexible, the tail would wave from side to side in its passage from right to left (Fig. 63). The pressure would be alternately downward from side to side as indicated in the enlarged section A-A. This would tend to raise the tail and pitch the nose downward, were it not for the pectorals being held out at an appropriate angle. Indeed, the fins of sharks are almost permanently fixed at this angle. Daniel (1922) has shown by pinning the pectorals down that fish with epibatic tails under such conditions always head downward and are then unable to rise. Fig. 64 shows such an epibatic tail from above in two successive positions. In A both the upper and lower lobe is pressed to one side, that is, when they are both on the same side of the crest of one wave. When this has travelled back as in B to where it splits between them, the lower lobe switches to the other side and is later followed by the upper so that they are again on one, but the opposite, side of a crest. While the crest is between them it is clear that the two lines of pressure are diagonally opposed, thus forcing the tail forward in a line more nearly parallel and closer to the axis of progression. This is obviously an economy which tends to reduce the oscillation of the nose. In the isobatic (homocercal) tail, there is no tendency to lift or depress the tail, the force all being in a horizontal plane. The various forms this type of tail may take are discussed on pages 226 to 229 together with the special contours of the epibatic tail. Fig. 65 compares the two types. The acuminate isobatic (diphy- cercal) type appears only where all locomotor efforts have been removed from that organ and have been transferred, for example, to the anal, as in Gymnotus (Fig. 48 B). The hypobatic tail is a comparatively rare structure associated 226 Zoologica : N. Y. Zoological Society [IV; 5 Pig. 65. Comparison of an isobatic and epibatic tail. A, isobatic tail as in Pomatomus. B, epibatic tail as in Mustelus. with flying fish and their kind. The action is naturally the reverse of the epibatic and apparently has some bearing on the frequency with which they ascend rapidly in preparation to flight or skittering over the surface. The many and various shapes of the tails of fishes are indicative of the speed and type of movement enjoyed by the different forms that use movements of the body for propulsive effect. Fishes with large squarish or spatulate tails, as Promicrops, Micropterus and Ephinephalus are comparatively slow but are capable of extremely sudden short spurts of speed, but never of long continued high speed efforts. Fishes with deeply forked or lunate tails are capable of long continued swimming at high velocity, the more lunate the tail the faster being the fish as in Coryphaena , Pomatomus and Sarda. These tails from tip to tip span a distance equal to from 18 to 25 per cent, of the total length, and frequently the accompany- ing narrow peduncle is strengthened with keels. Twelve high speed fish averaged 21 per cent, in this proportion. Extremely slow speed fishes, in which the tail functions as a propellor very slightly, are unforked and proportionally large, as in Alutera. Heterocercal tails while generally forked, at least in the Salachii, subscribe to the same generalization as is evidenced by the fast I sums of open waters with its crescentic tail and by the sluggish Ginglymostoma of littoral environs in which the tail is very slightly notched and asymmetrical. Again in inland waters, Polyodon, fast as compared to Amiatus and Lepisosteus, exhibit the same contrast although the large awkward rostrum of the former to some extent interferes with swift motion. In the Selachii, an evenly oval cross section is associated with a nearly symmetrical tail, while a subtriangular section is associated with an asymmetrical tail. This condition again reflects a bottom and pelagic habit. Certain fishes with forked caudals show a very slight prolongation of one lobe over the other as in Rachycentron in which the upper 1926] Breder: Locomotion of Fishes 227 lobe is slightly larger, and in many characins the lower is somewhat produced. It hardly seems possible that these slight differences could have locomotor significance, or at least in the present state of our knowledge to attribute such would be mere speculation. Nichols (1915 and 20) gave the substance of the following as an explanation of the fork in fishes’ tails. As the bodies of speedy fishes are streamline forms, the water that is displaced forward moves around the fish from front to rear more rapidly than does the fishes’ motion relative to the water outside the limit of its influence. For a maximum efficiency this displaced water should meet immediately on leaving the converging curves of the body. A square tail would prevent this and thereby offer some impedance as it causes a parallel run of the water after, in the interests of economy, the two streams should have merged. Also, by placing the functional tips of the caudal fin well above and below this point of confluence, a much better purchase is had on the water, which at these points is not moving away from the fish so rapidly. While this hypothesis is believed to be true, there are other probably equally important factors. Furthermore, Nichol’s idea does not give a positive reason as to why some tails are squarish, it being left as an assumption that a fish moves more rapidly if its tail happens to be forked. If an ordinarily spatulate tailed individual has a prominent fork cut in the tail, no greater speed is attained nor is it visibly reduced, but the motion of the body is different for reasons explained on pages 228 and 229. On the other hand, if a “comet” goldfish ( Carassius auritus) with its cumbersome and over-developed tail, produced by artificial selection, has it so trimmed as to be similar to that of a common or “plain-tailed” goldfish, it will attain the normal locomotion of the species,9 al- though these often seem to be considered as fishes of speed. An explanation of what the changes in the body movement accompany- ing a trimming of the tail indicates is superfluous, because the fol- lowing description of the ostraciiform model treated in a similar manner serves better in its stead. Both a forked tail and a square tail, otherwise similar, were constructed for the model (see Appendix) so as to be interchangeable. 9 In performing this experiment care must be exercised in the selection of an example. One should be taken with a normal body, as frequently a change in shape in the body ac- companies this excessive size of the tail. A young example is preferable, for generally old fish, even if normally bodied at first, are so modified by being forced to manipulate this excessive growth for a long time as to be worthless in this experiment. 228 Zoologica : N. Y. Zoological Society [IV ; 5 The difference in the model's action was studied comparatively, from which the following was found: With a square tail. The highest speed attained with this tail, less than the following, was obtained by adjusting it to move through an arc of about 35°. A larger swing of the tail with such a large surface caused so great a swing of the nose that the motion lost in that manner impeded forward progress to a con- siderable extent. With a forked tail. The highest speed attained with this type of tail, higher than the above, was obtained by movement through an arc of about 70°. The amplitude of the swing of the nose was not increased as rapidly as it was in the case of the former with the increased caudal amplitude, for there was less resistance to the passage of this tail of lessened area (about one-fourth less) through the water. This, on the other hand, formed a less effective blade on account of the reduced surface of pressure. It is evident that a forked tail swinging through a wide arc is more suited to fast movement than a square one of larger area, and also more efficient than a square tail given less amplitude. In these experiments the driving mechanism was set to move at a definite speed with no load applied. As various loads were then placed on the clockwork motor, it slowed down proportionally. As the greater the resistance offered to the tail the slower its movement became, the speed of progression stood as an index of the thrust of the various tail forms. The connecting mechanism was so designed as to give one complete cycle of the tail’s oscillation for each revo- lution of the driving shaft. As this was true for any amplitude, it follows by the laws of leverage, that the short strokes were more powerful than the longer ones, it taking the same force to produce each. From this, it is inferred that for a continuous high speed a deeply forked tail oscillating through a large arc is more easily manipulated than a large one making short strokes and eating up more energy on account of the accompanying great swing from side to side of the nose, which is lost motion. The fact that water particles impinging on the tail fin near its base attain a more nearly backward path may have something to do with shaping the tail in fast swimming forms (Fig. 43). That is, the center part of the tail is cut away reducing the amount of area that gives more of a sidewise and less of a directly backward thrust, with its accompanying large deflection of the head. Coupled with Nichol's factor, this effect is probably of considerable importance. Other experiments with the model concerning caudal fins demonstrated the following: the more flexible the tail, the less the nose oscillates. This would naturally follow for a flexible tail gives a more nearly backward thrust than a rigid one, as explained on pages 171 to 173. If such a flexible tail be made long enough, a complete sine curve or more will be formed i. e., the anguilliform type carried on by momentum. This calls to mind the motion of tadpoles and spermatozoa. As fish tails are all more or less flexible, 1926] Breder: Locomotion of Fishes 229 Fig. 66. Comparison of action of model with a rigid and a flexible tail. A, with rigid tail. B, with flexible tail. The dashed arrows indicate the angles of pressure. The dashed curves at the nose indicate the amount of its swing. it follows that the really extreme ostraciiform type as seen with such a rigid tail as in the model is never actually encountered in life. Fig. 66 compares the motion of a rigid and flexible tail on the model and the accompanying oscillation of the nose. As a further demonstration of the differential pressure of such a tail movement due to its position at the end of the body, the following is given. If the tail is not secured perfectly rigid to the turning support and ac- curately centered, it will eventually work itself around to one side causing the model to take an inward spiral path. (Fig. 67) first three positions. After this has gone on for a time and the tail oscillates equally about a line drawn at right angles to the axis of the boat through the tail post, it simply turns on a point as in the fourth position (Fig. 67). After the tail passes this position and the major part of each oscillation is anterior to this line, the model starts to back up and spirals outward until halted by the tail slapping against the side of the hull, as in the final positions (Fig. 67). This forcing of the rudder around on its not too tightly secured axle can only be accounted for by the presence of less resistance to the second half of a stroke than to the first. That is, the part of the stroke which causes the nose to deflect most sharply is less in resistance than the part which chiefly moves the boat ahead. In the moving boat the passage of it through the water with the consequent currents flowing aft must have nothing to do with it for if they controlled such a loosely fastened tail, their effect would be to keep it centered and even if started off center they would return it. It seems to be referable to the oscillation of the anterior part. This would indicate that after once being started the nose swings largely by momen- tum on the last half of a stroke with speed nearly as great as that of the caudal causing little pressure to be felt from the flowing water relative to the tail fin. In backing out of the spiral the tail acts like one pectoral fin as described under Pectoral Propulsion. Pectorals of a spatulate form accompany fishes of slow or moderate speed, while long falcate, or at least not spatulate ones, accompany rapid swimmers which use them largely for wheeling and seldom if ever for either propulsoin or backing water. The actual form of the ventrals appears to be of little significance in rapid swimming, they being for the most part auxiliary maneuver- ing instruments or at best “ bilge keels. ” They are uniformly kept tucked close to the body in rapid swimming. Generally 230 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 67. Behavior of model with tail fin insecurely attached to shaft. The arrows indicate direction of travel. Note that the tail oscillates progressively more and more to one side. considered, they are the fins that fishes can best dispense with, excepting the dorsal or anal in certain forms. They are the only fins on which the primary locomotor significance is not placed in some forms. Examples are numerous of cases of the predominant importance of all other fins, e. g. caudal, Micropterus; pectoral, Raja; dorsal, Gymnarchus , anal, Gymnotus. Also they are the fins most frequently modified to such an extent as to lose all loco- motor function, as seen in Phycis and Colisa, and not infrequently they are entirely missing. These facts coupled with the experiments described on pages 215 and 216, lead to the belief that the pelvic fins are of comparatively minor locomotor importance. Their form generally follows that of the pectorals quite closely. See also “ Starting and stopping.” The dorsal and anal fins in fast fishes are generally somewhat similar in size and shape and it is only in relatively slow forms that any great developments of one over the other is seen. However, 1926] Breder: Locomotion of Fishes 231 there seems to be a general tendency for the anal to be somewhat shorter than the dorsal. The reasons for this appear to be two fold. The anal cannot very well extend forward of the vent with economy for several anatomical reasons, which by reference to the preceding parts should be clear, as well as probably for phylogenetic reasons. The pelvics appear to compensate for lack of length of the anal, acting as “ bilge keels” as pointed out above, for, primarily, the dorsal and anal are keels. Comparatively elongate forms that have the pelvics greatly reduced or wanting, generally have the anal unusually long with the vent often displaced forward. In the fastest forms the vertical fins can be depressed so completely as to mar the perfection of the streamline form in no way at all. In fishes with their greatest bulk above a line from the tip of the snout to the middle of the peduncle, the dorsal is generally larger than the anal and conversely, when the bulk is below that line, the anal is generally larger; whereas in fishes nearly symmetrical about such a line, these fins are practically equal in size as noted by Abel 1925. This appears to be merely a placing of these steadying keels at the point of greatest vantage. A fish if inclined to roll will do so about this line, giving the greater bulk the largest swing on which a steadying keel would be most effective. The remaining parts not mentioned specifically including axillary scales, produced fin rays, erectile processes, either employed for locomotor or other purposes or both, are formed according to streamline necessity in proportion to the speed desirable to the possessor. As these are specializations usually of a limited character they are treated under their respective heads in Part II — Syste- matic. The chief external characters of fishes affected by or affecting locomotion may be tabulated according to their variations. Such a table follows, its purpose being to make more clear the broad gamut of variations through which fishes range, although it should be borne in mind that there is no attempt to go into details, simply the main elements being indicated, with their extreme modifications and the norm between them. This table lists the principal elements of fishes’ external topog- raphy and gives their chief ordinary range of variation. It is believed that the list is complete in a broad way, i. e.f that any species could have its various elements checked off according to the eleven items, which with their various descriptive divisions 232 Zoologica: N. Y. Zoological Society [IV; 5 Elements Center of gravity Shape of body Surface of body Snout Mouth Gill opening Dorsal and Anal Caudal Pectorals and Ventrals TABLE III Variation Vertical position ( High — M edian — Low ) Horizontal position (Anterior — Median — Posterior) r Longitudinal Elongate — Fusiform — Truncate) Transverse ( D epr essed — C y lindrical — C ompressed ) Special (Bizarre — Rotated through 90° — etc.) r Protective Covering * (Naked — Scaled — Rugose and armored) • f Relative Length * (Attenuate — Medium — Blunt) [ Size I (Large — Medium — Small) Position (Superior — Terminal — Inferior) Size ( Large — M edium — S mall ) Shape (Slit-like — Oval — Circular) Vertical position (Superior — M edian — Inferior) Special (Multiple — United below — etc.) Length ( Long — M edium — Absen t ) Height (High — Medium — Absent) Shape (Lobate — Rectangular — Falcate) Special [ (Multiple — Produced rays — etc.) Length (Long — M edium — Absent) Spread ( Great — M edium — Absent) Shape (Spatulate — Truncate — Forked) Type (Epibatic — Isobatic — Hypobatic) Special ( (Prehensile — Produced rays — etc.) Length (Long — Medium — Absent) Height (High — Medium — -Absent) Shape (Spatulate — M edium — Falcate) Vertical position (High — Medium — Low) Horizontal position (Anterior — Medium — Posterior) Special (Sensory rays — Intromittent organs — etc.) 1926] Breder: Locomotion of Fishes 233 could be placed in one category or another. It would be found if a large number of species were tabulated according to this list that they would fall into certain rather constant groups. That is, certain characters are definitely associated with others. For example, fish with spatulate pectorals do not have forked tails, fishes with falcate pectorals are never exceedingly elongate, and so on. This brings us to what is probably the most important consideration, one not shown by the tabulation itself. That is the relationship of these factors, as above noted and not their absolute intrinsic quantities alone, deter- mines the locomotor characteristics of a given species. It explains how fishes of different appearance may have nearly similar locomotor functions, equal speed and so on, for not alone does the absolute response determine these conditions, as the co-relation of the involved elements must be of a given sort to produce a specific result. Relationship to habits and Development. Habits and development are both determined by locomotor requirements and determine locomotor abilities. A few examples may be considered here. The habit that many smaller fishes have of following larger ones, ships or almost any moving body of considerable size, may among other advantages, enable them to slip along with less effort than they would otherwise have to make. For example, a small shark following a large vessel is swimming in a flowing stream of water running toward the stern of the boat to fill up the cavitation created in moving forward. As this is running in the desired direction, the effect is the same as though the fish were being carried along by a tide. In modern screw propelled vessels the wake trends clearly forward. A small fish following a larger one receives a similar pull, but to a lesser degree because the jets, as pointed out under the section on Jet Propulsion, tend to make the wake trend back- ward. That they fail to do so completely, at the lower speeds at least, may be observed by watching suspended particles. The undulations of the body and other factors offset and overcome the tendency of the exhaled water to reverse the direction of the wake. In a similar manner it would seem probable the lagging members of a school dropping behind from fatigue or exhaustion would be unconsciously helped along by their more sturdy brothers swimming in the van. 234 Zoologica: N. Y. Zoological Society [IV; 5 A Berlin engineer, H. Schieferstein, working on the effects of tuned oscillating parts on reciprocating machines, noticed an increase in power output in machines that were made up of parts with a definite periodicity, (Gradenwitz 1923). This efficiency led him to suggest that flocking birds might gain a mutual advantage through the intervening air spaces. If this be proven true, it is more than likely that fishes in schools receive a similar mutual benefit on account of the fact that mechanical shock and vibrations are much better transmitted through water than air. Then too, it is a matter of simple observation to note how often several ad- jacent members of such a company are moving with perfect syn- chronism. More data on the entire subject of the oscillatory effects on power machines is essential before a thorough understanding of the subject relative to the schooling of fishes may be had. At least it is probable that an individual fish commonly vibrates its body in accordance with the proper frequency, considering the animal or its oscillating part as a vibrating element. In the section “Maintaining a stationary position/’ mention was frequently made of movements of the various fins. Certain of these have been explained as counteracting the disturbing in- fluence of respiration, but a considerable percentage of the move- ments that make it difficult to appreciate the real effect of respiration is not explainable on that basis. As these movements are, neces- sarily, most often neutralized by others in order to maintain a resting position, it is evident that they serve no equilibrating function. Here then is an apparent waste of effort on the part of the muscular system, from which no useful result is obtained. An illustration may often be seen in the Centrarchidae when a slight caudal undulation is accompanied by an increase in the backing efforts of the pectorals far above that necessary to offset the thrust of the exhalations. Another may be seen in Amiatus which at times will start a series of waves in the dorsal at its anterior end preparatory to moving forward, but before the inertia is over- come, some negative impulse causes the initiation of a similar series from the posterior end, effectively checking the reaction of the first; these opposing series meet at about half way between, and then die out. The explanation of such muscular displays is referred to the nervous system, as it is conceived that the muscular activity ob- served is the visible effect of nervous impulses not altogether 1926] Breder: Locomotion of Fishes 235 inhibited, as fishes in a tank or natural body of water are constantly receiving various stimuli of an opposing nature. For example, if some object be dropped into a tank containing fishes, they usually rush toward or away from it, depending on the amount of commotion it makes in breaking the surface, its size, etc. If the object be something inert, like a stone, the fishes soon quiet down again, often coming back to their previous positions. This is an illustration of a case where the reversal of the impulse follows some time after the original one. If however, some less violent stimulus, as, for instance, another fish too large to be suitable for food and too small to be dangerous, comes within the sensory range of an individual, the first stimulus will be to move, but, lagging behind this by only a fraction of a second, will come the details of size, and in this hypothetical case no change of position being necessary or desirable, the reversal of the fin movements will be nearly instantaneous. Fishes dependent upon flight both for safety and for pursuit of food, show this characteristic of quick reversals or counter movements most strongly, if indeed they do not keep in almost constant motion. On the other hand, bottom forms such as the Rajidae, Pleuronectidae, Cottidae and Batrachidae, which depend primarily on coloration and shelter for safety, fail to show this pass- ing over of impulses at all. In one sense the rapid alternation or starting and checking of movements might also be likened to the “dancing” of a boxer, in search of an opening for a thrust. In some forms, as Umbra, this activity is so definite that it may be considered a fixed and constantly present habit. It seems possible at least that this keeping open of the nervous paths may serve a useful purpose when it is necessary to make an immediate depar- ture. Vertebrate animals accelerate their rate of respiration on fright. This has usually been attributed to a primitive reflex preparatory to precipitous flight. In the case of many fishes whose practically only reaction to a fearsome object, under most circum- stances, is flight, this increased respiration caused by fear acts directly in the actual effort of moving away from the disturbing influence, thus being a mechanical advantage as well as a purely physiological one. The larval forms of fishes which are free to move about begin life with a sub-carangiform or even anguilliform movement, even if they develop into something entirely different subsequently, e. g. 236 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 68. Progression of a leptocephalus. Conger sp. After Dean (1912). Spheroides (Welsh and Breder, 1922). This is always found to be the case, as indeed would be expected, except in very specialized incubation or developmental habits. In these cases the free swim- ming stage is possibly abbreviated or obliterated, and may be passed through either within the egg capsule, or in the female’s body, or in some special environment; or the locomotor apparatus is functionless as a means of propulsion because of the possession of some special structure making locomotion undesirable or impossible. An example of the first case of reduction in swimming ability in the larval stage is seen in Fundulus heteroclitus, where the young fish do not leave the egg until they have become post-larvae with a full complement of fins, when they can swim substantially as do their parents. Any ovoviviparous fish, such as Lebistes reticulatus , illustrates the second case. The third is exemplified in the larvae of practically any nest building fish that stands guard over its offspring, as Macropodus, Gasterosteus, or in the larvae of any form showing oral gestation, as Felichthys or Paratilapia. The larvae of the Salmonidae are so encumbered by the great yolk sac which does not disappear until after the larval state has been well passed, that they simply lie on the bottom in passing through this stage, awaiting the yolk’s absorption. Many of the long bodied larvae, such as leptocephali, show the most extreme form of anguilliform motion to be seen anywhere (Fig. 68). The curves resemble, to a certain extent, those of a snake progressing over a very smooth surface. This movement is no doubt correlated with the extreme ribbon shape of the leptocephali, making this excessive bending relatively easy to perform. Dean (1912) mentions that he could not notice any “slip” in the progres- sion of a larval Conger and that a pencil placed in one of the loops 1926] • Breder: Locomotion of Fishes 237 was not touched as the animal moved forward. However, Dean’s Leptocephalus while apparently having a high coefficient of efficiency would have eventually touched his pencil if it had been sufficiently long. It is nevertheless remarkable that such a great amount of effective energy could be delivered. In this type of movement no orthokinetic port is formed. See page 182. As metamorphosis proceeds, this method is supplanted by the more familiar anguilli- form type. The primitive pectorals are generally spatulate in shape and early function as paddles. As they increase in prominence, the anguilliform motion decreases in those species destined to lose it, until the post-larval, and finally the adult, form of locomotion is attained. The more special details of habit are treated under their respective heads in “Part II — Systematic.” PART II— SYSTEMATIC Introduction. Having considered the physical forces applied by the various types of fishes in moving about in “Part I — Physical,” it now remains to examine the various major groups of living forms in order to understand how their individual specializations agree with the more general statements set forth and developed previously. In the actual work upon which this paper is based, the opposite method was naturally employed. That is, a great number of species, each usually represented by many individuals, was studied and experimented upon before the generalizations of the previous part were hazarded. It is obviously impossible for any one person to examine all extant fishes in the flesh, and it is altogether reasonable to make inferences on the belief that like locomotive structures within one group are applied in closely similar manners. No attempt has been made to describe in minute detail the very small differences between closely related fishes for such varia- tions are referable in most part to causes other than locomotor requirements. Such changes as a slightly larger or smaller number of fin rays are so bound up in the sum total of phylogeny that it is impossible to ascribe a locomotor advantage of one over the other. 238 Zoologica : N. Y. Zoological Society [IV; 5 If any such advantage actually exists, our knowledge of the habits of fishes is manifestly too inadequate to enable a correlation to be made that is more than grossly speculative. On the other hand, decidedly different fishes often accomplish similar results in slightly different ways because, for non-locomotor reasons, members pri- marily concerned with motion have secondarily acquired added and different functions, e. g., the change of the anal fin in male ovoviviparous Poeciliidae to an intromittent organ. Treatment by Orders. Under this section the great groups of recent fishes are con- sidered in taxonomic order as arranged by Bridge and Boulenger in the Cambridge Natural History (1904). 10 That is, the arrangement given in that work is followed completely down to and including families. The species and genera mentioned, however, follow more recent taxonomic practice for obvious reasons. Under each group- ing a very general statement is given explaining the locomotor classification or classifications into which its various members fall.* 11 In many cases the general types of locomotion are so similar to others that for the present purposes a few words are sufficient and in most cases the correlation with the propelling structures is evident by referring to the previous part. To avoid repetition, page references are given to Part I, in which members of the group are discussed. Following this, specific instances and side lights on those species most closely worked with are mentioned. These are placed in smaller type as their bearing on the main theme is slight, but they serve to illustrate various points in detail and to show what sort of differences might be expected in any group not so intimately known. At the end of each group is tabulated a list of the species studied in life to enable other workers to check the present paper intelligently and to indicate the breadth of the base upon which the generalizations were based. Species in light faced type were studied by observation only, either in a state of nature, in captivity, or, in most cases, both. Those in heavy type were experimented upon in various ways as partially indicated in other parts of the paper. i° This classification is used simply because of its widespread use, and its adoption here should not be taken as an endorsement of it. 11 Acraniate forms are omitted from the present discussion as being too far removed from the chief forms under consideration to warrant inclusion for the present purposes. 1926] 239 Breder: Locomotion of Fishes CLASS— CYCLOSTOMATA Orders— MYXINOIDES and PETROMYZONTES These elongate jawless forms use the simple anguilliform method of necessity, as they possess no appendages of a locomotor nature. The continuous fin fold can only function if the body is used in an anguilliform manner. They are skilled at burrowing. The Petromyzontes are also able creepers, accomplishing this by aid of the suctorial mouth. See pages 203 and 219 for other data. Petromyzon marinus, and probably all others, when swimming or forcing a passage through weeds and debris, usually folds the suctorial disc together upon itself from side to side. This makes the entrance portion of the animal regu- larly conic in shape, thus reducing the hindrance that would otherwise be formed by the open buccal cavity. If open, it would tend to deflect the head upwards and as these animals possess no paired fins at all, difficulty might be encountered in offsetting this deflection, as it would have to be done by exertion of the body muscles alone. Fig. 69 shows the passage of the stream lines about the head of an example both with the mouth open and closed (Breder 1923). The ability of this animal to inch its way up smooth, damp surfaces is well known and may be observed at any dam or stream frequented by them. Species studied in life — 1. Petromyzon marinus Linnaeus. 2 — Lampetra wilderi Jordan and Evermann. CLASS— PISCES Subclass — Elasmobranchii Order— PLAGIOSTOMI Suborder — Selachii The typical sharks display swimming movements intermediate between the anguilliform and carangiform. Mustelus approaches the former and Carcharodon the latter. Some of the depressed forms approach the movements displayed by the Batoidei (rajiform) in certain respects. The family Rhinidae is fairly intermediate, but is closer to the skates in its movements than to the generalized sharks. Correlated with the transition from a practically anguilliform shark to a sub-carangiform one is associated an accompanying change from an epibatic to a sub-isobatic tail and from a sub- triangular cross section to a nearly elliptical one. 240 Zoologica: N. Y. Zoological Society [IV; 5 A CD Fig. 69. Diagram illustrating the reason for Petromyzon marinus closing the mouth while swimming. A, head of a swimming lamprey with mouth open indicating how water below the axis of the body would be deflected downward from the mouth, thereby forcing the head up. B, head of a swimming lamprey with the mouth closed indicating how the water both below as well as that above the axis of the body would be parted without any deflecting effect. C, ventral view of the head of a lamprey with the mouth open. D, ventral view of the head of a lamprey with the mouth closed. The function of the notch in the upper lobe of the tail is un- known, and there is a possibility that there is none today, its presence being accounted for phylogenetically, for sharks deprived of the tab certainly appear to swim as well as those with it. As the tail waves from side to side this pendant piece simply flaps from one side to the other, trailing along after the tail proper. Abel (1912) believes that the tab was originally the whole caudal fin and that a second anal migrated backward and forced the tip out to where it still remains today, while the anal became the functional caudal. This theory lacks conviction and a function may yet be ascribed to the notch. At least it is certain that this tab increases the tendency to raise the tail because of its greater flexibility. The fins of sharks are very limited in their movements practi- cally their whole function being that of keels and rudders. On this account, sharks are unable to make an abrupt stop, swerving to one side of an obstacle instead, for they are unable to use the pectorals or any other members as “brakes.” Many sharks, such as Mustelus, Carcharhinus and Squalus flex the dorsal and anal at times to aid in turning. That is, they bring the posterior parts of these fins toward the concave side of the turn. The under surface of the lobe of these fins is slightly concave and quite possibly forms a suction to the side to help hold the deflexed fins in place during the turn. See under Tarpon, page 247, for other data concerning the use of fins in this manner. The turning over of sharks when feeding from the surface is 1926] Breder: Locomotion of Fishes 241 not a necessary act, for often they will deliberately swim up, pro- truding the snout, so as to place the inferior mouth at the surface. However, when such turns are made, they are controlled entirely by warping the guiding keel-like fins. See pages 194-195, 203, 226, 233, and Figs. 52, 63, 64, 65 B, for other data. Mustelus canis when chafed by confinement will often swim up vertically through the surface of the water and hold a position with the fore part of the body exposed for some time by the continued action of the submerged hind portion. Over one-third of the body is often so lifted, and for shorter times even as much as one-half has been noted. This gives a measure of the force exerted, for it is not a matter of momentum as in leaping, but simply a direct and continued “push.” Sphyrna zygaena is believed to use the lateral expansions of the cephalic region as a bow rudder acting in a vertical plane, enabling it to rise and dive with unusual alacrity. If Vulpecula marina Valmont possesses a locomotor advantage in its elon- gated tail, this is unknown. Allen (1923) believes that it is used as a “whip” to maim smaller fish in order to facilitate their capture. Rhineodon typus Smith, although little known, seems to gain some advan- tage from its exhalations. Mr. L. L. Mowbray, in speaking of a thirty-five foot specimen taken at Long Key, Florida in June 1923, stated that in its struggles to free itself while tied by the tail, it ejected powerful jets of water through the gill slits causing a great current to flow back so strongly that a miniature breaker was formed at about the first dorsal. Also, when the fish was at rest a piece of seaweed inadvertently taken in at the mouth would be ejected violently through the gill clefts by an extra strong jet. Species studied in life — Family— SCYLLIIDAE 3. Ginglymostoma cirratum (Bonnaterre). Family— CARCHARIIDAE 4. Mustelus canis (Mitchill). 5. Carcharhinus milberti (Muller and Henle). 6. Carcharias taurus Rafinesque. Family— SPHYRNIDAE 7. Sphyrna zygaena (Linnaeus). Family— SPINACIDAE 8. Squalus acanthias Linnaeus. Suborder — B atoidei The primary locomotor organs of most members of this sub- order are the tremendously enlarged pectorals in which are localized 242 Zoologica: N. Y. Zoological Society [IV ; 5 A Section B'B Fig. 70. Diagram of the streamline contours of Rhinoptera bonasus. The sections of the actual photograph A-A and B-B were made by slicing the animal shown and tracing the outlines. anguilliform undulations in a vertical plane (rajiform Fig. 48 D). Certain examples approach the movements of the Selachii, Pristis, for example, being more shark-like than skate-like in its movements. These two orders intergrade completely in locomotor methods, following of necessity the changes in body form, especially regarding the degree of depression. The Pristidae have a close parallel in the Pristiophoridae among the Selachii, whereas the Rhinobatidae some- what resemble the Rhinidae. In the family Rajidae, the tail steers to a certain extent, being^ held out rigidly, although at times it may be lashed about with some trivial locomotor effect, chiefly because of the position of the reduced dorsals near its tip, which form a small but functional tail fin. The family Torpedinidae has a less expanded disc and a functional caudal fin. The pelvics have a special function when the skate is resting on the bottom; they operate to push the animal 1926] Breder: Locomotion of Fishes 243 forward by kicking back much as though they were feet. Frequently in rising the members of this family give a single violent backward thrust, so launching themselves on their way. While the fish rests on the bottom, most of the excurrent water passes out through the spiracles, and on rising, if it is physically possible, the water is likely to be directed through the gill slits so as to react forcibly on the solid substratum. The families Trygonidae and Myliobatidae have carried the rajiform undulations a step further. The tail has become a mere trailing wisp, which, while it may have some slight course-deter- mining action, appears to trail passively after the animal. Steering is accomplished for the most part by differential action of the pectorals. That is, in curving, the outside “wing” is speeded up or the inside one retarded which swings the animal accordingly. These forms, being broader than long, do not appear superficially, to be especially well fitted for cleaving the water, although they are notably faster than the skates. One evident adaptation is the frequent loss of all rugosities which generally ornament the skates. It is clear that this apparently unhandy shape is necessary for a high development of this type of locomotion. On close examination, however, it will be seen that for a body of their peculiar shape a maximum streamline form has been developed. The longitudinal sections (Fig. 70) indicate the nature of the wedge shape of the body. See pages 187, 207, 223, 230, and Fig. 48D, for further data. Pristis microdon in ordinary swimming swings along with the gait of the shorter bodied sharks, such as Carcharias, although in addition, the edges of the large pectorals flutter. Most of this fluttering is probably incident to their motion through the water although some of it is doubtless muscular and of locomotor function. At times when frightened they will flatten themselves against the sand in a manner similar to Raja. Then and when very slow motion is made, the movements of the pectorals are prominent. They appear to reverse these ripples as a gentle braking action. The armed rostrum weaves back and forth somewhat as would a bowsprit on the ostraciiform model. Raja eglanteria sometimes adheres to smooth vertical surfaces. At such times most of the exhaled water escapes through the spiracles, which may serve as a slight reactive function to hold the skate against the surface. That all does not so escape is demonstrated by the slight ripples at the posterior part of the disc caused by the water flowing from the gill slits. As they invariably head up, this must tend somewhat to offset the effect of gravity. As the fish slightly arches its body, it would seem that some suctorial effect was attempted although this would be destroyed at each exhalation allowing water to pas« through the gill clefts. 244 Zoological N. Y. Zoological Society [IV; 5 Species studied in life — Family— PRISTIDAE 9. Pristis microdon Latham. Family — RAJIDAE 10. Raja erinacea Mitchill. 11. Raja diaphanes Mitchill. 12. Raja eglanteria Bose. 13. Raja stabuliforis Garman. Family— TRYGONIDAE 14. Urolophus jamaicensis (Cuvier). 15. Dasybatus centrura (Mitchill). 16. Dasybatus say (Le Sueur). 17. Pteroplatea micrura (Bloch and Schneider). Family— MYLIOBATIDAE 18. Rhinoptera bonasus (Mitchill). Order— HOLOCEPHALI Nothing can serve better here than a paraphrased summary of Dean's (1906) excellent description of the behavior of Chimaera colliei Bennett. As the caudal is reduced to a diphycercal wisp the force derived from metameral undulations is dependent upon the expanse of the long low dorsal and anal which are produced to the posterior ex- tremity of the body. The large delicate pectorals appear to have two types of movement which may be used together or separately They may be used somewhat after the manner in which Abudefduf wings its way through the water as described under “Pectoral locomotion," or they may be undulated when held out at right angles to the body in a rajiform manner. When moving slowly these fish rock from side to side slowly showing an imperfect stabiliz- ing effect of the fins acting as keels. Normally they are rather sluggish but at times of great excitement may show sudden spurts of speed. They sometimes rise considerably above the bottom and then sink slowly, probably after the manner of the triglids. Callorhynchus with its decidedly epibatic tail probably ap- proaches the sharks more in its manner of locomotion, that is, the tail is relatively more important as a locomotor organ, while Har- riota with relatively large pectorals and an attenuate caudal appears to have the emphasis placed on the former, and doubtless some, such as Chimaera, have anguilliform flexures localized in the dorsal fin. 1926] Breder: Locomotion of Fishes 245 Subclass Teleostomi Order— CROSSOPTERYGII Evidently most of these fishes swim in a manner approaching the anguilliform, although they present a certain stiffness, some- what as in Tylosurus or Esox. Harrington (1899) writes, “Peculiar in the swimming movements of Polypterus is the manner in which the head moves freely from side to side. This produces the appear- ance of a progression more or less snake or eel-like, although in general the powerful sweeps of the strong tail characterize the progression as fish-like.” Its lobate pectorals are, no doubt, used for progressional purposes at times. Calamoichthys is probably more similar to Anguilla in its mode of locomotion and little aid can be expected from the small pectorals. The purpose of the interrupted dorsal is not clear although it may serve in a capacity somewhat similar to that of the finlets of the Scombridae. However, neither the general build of the fishes nor the anterior position of the “finlets” would suggest that such a device would be of any particular service. Order— CHONDROSTEI The fishes of this group swim in a manner intermediate be- tween that of the anguilliform and carangiform type. The epibatic tail makes the warping of the pectorals imperative as with the Selachii. Likewise abrupt stops are not possible on account of the lack of an efficient “brake.” The small vertical and pelvic fins allow of a considerable rolling motion which their small area does not overcome. See page 179 for other data. Species studied in life — Family— ACIPENSERIDAE 19. Acipenser sturio Linnaeus. 20. Acipenser rubicundus Le Sueur. 21. Acipenser brevirostrum Le Sueur. 22. Scaphirhynchus platorynchus (Rafinesque). Order— HOLOSTEI The locomotor movements of this order are sub-carangiform. That is, their movements embody hardly more than one-half a sine curve, for, although they are elongate the rigidity of their 246 Zoologica: N. Y. Zoological Society [IV; 5 bodies prevents anguilliform curves from being formed. This is no doubt associated with their heavy armature. See pages 186-189, 202, 207, 222, 226, 234, and Fig. 48 A, for other data. Amiatus, in addition, in much of its movement employs localized undu- lations situated in the long dorsal (amiiform) (Fig. 48A). The pectorals are employed largely in backing water for which their size and shape are beauti- fully adapted. Lepisosteus osseus and others of the genus, on account of their long and rel- atively rigid bodies turn awkwardly, employing both the pectorals and pelvics in a manner which Dr. W. K. Gregory aptly likened to that of a “hook and - ladder truck,” there being a steering device at either end. Their stopping is made in a manner similar to that described on page 69 and illustrated by Esox, Fig. 57, A and B. Species studied in life — Family — AM 1 1 DAE 23. Amiatus calva (Linnaeus). Family— LEPIDOSTEIDAE 24. Lepisosteus osseus (Linnaeus). 25. Lepisosteus platostomus Rafinesque. 26. Lepisosteus tristoechus (Bloch and Schneider). Order— TELEOSTEI Suborder — M alacopterygii Fishes of this sub-order while showing a great range of loco- motor variation generally approach the carangiform type of loco- motion closely and consequently are generally strong and active swimmers. The pectoral fins, while capable of being folded back do not have the flexibility to be seen in many of the higher teleosts and are not often capable of being used effectively to back water with. Their use is more nearly like that of the sharks’ pectorals, except that they are tucked away in active swimming as the tails are isobatic and consequently there is no tendency to depress the fish that requires offsetting. Their prime function is wheeling and turning. While capable of some braking action, short stops are not the rule with these fishes. A turn to avoid an obstacle is the more usual method (Clupeidae). The tails vary from truncate to forked, which is somewhat associated with a fluvitile and marine habitat respectively, although in the more specialized forms it may disappear entirely as an active fin ( Gymnarchus ). If the adipose fin, common 1926] Breder: Locomotion of Fishes 247 Fig. 71. The ultimate dorsal ray of Tarpon atlanticus. A, cross section. B, the tip showing the groove below. in this group, has a locomotor significance it is unknown. As this is true of the adipose fin in other groups as well, no further mention will be made of it. Some of the members of this suborder are capable of tremendous leaps ( Tarpon and Salmo), and leaping amounts to almost a group characteristic. The numerous specialized fishes in this suborder that do not conform to the above general statements include the highly specialized Mormyridae with their elon- gate snouts, and the Pantodontidae with their powers of flight. Gym- narchus (Dahlgren 1914) and Notopterus (Schlesinger 1910), other specialized offshoots, propel themselves by localized undulations of the long dorsal and anal respectively. The Stomiatidae with their obvious adaptations to a very special environment, and others might be mentioned, but the above will serve here for the purpose of illustration. See pages 201, 216, 221, 223, 230, and Fig. 59A, for other data. Tarpon atlanticus possesses a long wisp at the posterior margin of the dorsal. This is concave below and tapers to a slim point (Fig. 71). Mr. L. L. Mowbray ascribes a special and definite function to this with reference to the famed leaping habits of the species. Preparatory to making a leap, according to Mowbray, the tarpon lashes this whip around to one side of the body and clamps it tight to its side. Here it adheres because of suction as the grooved piece lifts slightly from the body at its center. The dorsal is thus held rigidly to one side, so aiding in determining the course. Even in a dead fish the adhesion is considerable when the ray is applied in the manner described. The turn is made toward the side to which the ray is adhering. A Section A ~A Fig. 72. The axillary scale of Elops saurus. Side view with fin pressed to body and section with fin in same position. 248 Zoologica: N. Y. Zoological Society [IV; 5 Elops saurus shows a very delicate adjustment to streamline contours in the axillary scale of the pectoral. When that fin is held flat to the body, the scale rises and occupies the triangular area immediately above it (Fig. 72), thus giving a very smooth section to this region. When the fin is extended as in wheeling, the scale drops down behind it forming a sort of chute for the water to pass over behind the fin. This is of a very gently curving form, no doubt af- fording a minimum of frictional resistance. Ophisthonema oglinum possesses a similar ultimate produced dorsal ray to that of Tarpon . It would seem that this would serve a similar function, for even in very small examples a perceptible suction could be detected. Thymallus ontariensis appears not to use the large dorsal at all for propulsive purposes, although its size might suggest it even if its shape and proportions do not. Species studied in life — F amily — ELOPI D AE 27. Tarpon atlanticus (Cuvier and Valenciennes). 28. Elops saurus Linnaeus. Family— ALBULIDAE 29. Albula vulpes (Linnaeus). Family— CLUPEIDAE 30. Clupea harengus Linnaeus. 31. Pomolobus mediocris (Mitchill). 32. Pomolobus pseudoharengus (Wilson). 33. Pomolobus aestivalis (Mitchill). 34. Alosa sapidissima (Wilson). 35. Opisthonema oglinum (Le Sueur). 36. Brevoortia tyrannus (Latrobe). 37. Ilisha fiirthii (Steindachner). 38. Anchovia elongata Meek and Hildebrand. 39. Anchovia brownii (Gmelin). 40. Anchovia mitchilli (Cuvier and Valenciennes). Family— SALMONIDAE 41. Coregonus clupeiformis (Mitchill). 42. Oncorhynchus tschawytscha (Walbaum). 43. Salmo fario Linnaeus. 44. Salmo mykiss Walbaum. 45. Salmo irideus Gibbons. 46. Salvelinus fontinalis (Mitchill). 47. Cristivomer namaycush (Walbaum). 48. Thymallus ontariensis Cuvier and Valenciennes. 1926] Breder: Locomotion of Fishes 249 Suborder — Ostariophysi This order divides itself into two very differently appearing groups, the Eventognathi and the Nematognathi. The former includes fishes that in the main agree closely with the general locomotor habits of the suborder Malacopterygii. There is, how- ever, a general increase in the flexibility and use of the pectoral fins. Many cases of parallelism exist in the two groups. For example, certain characins bear more than a passing resemblance to Salmo, and a functional similarity exists between Gymnotus and Gymnarchus , and so on. So many of the South American Characini- dae and oriental Cyprinidae are narrowly and peculiarly specialized that no one broad statement can take them all in. Generally their locomotor habits resemble more closely some type in another group, entirely unrelated, which they happen to resemble, rather than some close relative. Thus Ctenolucius behaves more like Esox than its immediate relatives. Leaping is a fairly well developed character- istic of many forms, especially those resembling the generalized Malacopterygii in locomotor habit. Misgurnus shows nearly anguilliform movements and in many ways approaches the Nema- tognathi in locomotor habits. Their relatively large pectorals are Fig. 73. Pectorals modified for flight. Dorsal and lateral views. A, Thoracocharax. B, Cypselurus. 250 Zoologica: N. Y. Zoological Society [IV; 5 e t i AMD \ GASrl Fig. 74. The trajectory of characins. Astyanax, and other short-pectoraled forms. Thoracocharax and Gasteropelecus, long-pectoraled forms. largely in the nature of planes controlling the angle of elevation. The Nematognathi propel themselves by methods reaching from the nearly anguilliform to the carangiform and even beyond. The pectorals are largely planes as in Misgurnus. They are typically sluggish bottom fishes and the form of the body movements is directly correlated with their length and with the extent of their dermal armature. Thus some of the elongate naked Siluridae approach the anguilliform movements and the short, tightly incased plecostomids reach beyond the carangiform and extend toward the ostraciiform type. Some are adept at burrowing; others at creeping and still others at navigating over land. See pages 183-185, 187-189, 196, 207, 216, 219, 221, 223, 227, 230, 236, and Figs. 48B, 53, 56, for other data. Thoracocharax maculatus, owing to its long falcate pectorals is capable of soaring for considerable distances. Compared with the exocoetids there seems at first glance little similarity, but if viewed dorsally there is a pronounced one (Fig. 73). The similarity is even greater between these characins and the “two-winged” flying fish, Halocypselurus. This is destroyed in the lateral view by the greatly dilated ventral region devoted to pectoral musculature. Except for housing this musculature it probably has little to do directly with their flight. This flight seems to be the simple trajectory of a characin leap flattened by the presence of these pectorals (Fig. 74), although at times they seem to be able to sustain themselves for some little distance, and there may be some fin movement as Ridewood (1913) suggests for Gastropelecus. Carassius auratus developed into monsters by artificial selection hardly need any elaboration as most of the excess finnage is clearly an encumbrance to activity which is plainly evident on inspection. Further details are given on page 99. Synodontis membranaceus Geoffroy and some others of this genus are per- fectly accommodated to swimming in an inverted position. This suggests an adjustment to a displacement of the normal center of gravity. The perfect ac- commodation to this habit is indicated by the complete reversal of the color dif- ferential, these species being dark ventrally (the upper half) and light dorsally (the lower half). Astroblepus longifilis inhabiting mountain torrents is able to hold tightly 1926] Breder: Locomotion of Fishes 251 Fig. 75. The motion of Atroblepus longifilis overland. The portions of the caudal swing during which the mouth alternately releases and adheres is indicated. fast to smooth surfaces by use of its suctorial mouth, both under water and out of it. The method of progressing overland as seen by the author is essentially a fish-like flapping coupled with an appropriate action of the mouth, it holding fast on the flexure of the caudal portion, and releasing on the extension (Fig. 75). The depressed body and the extended pectorals and pelvics cause it to rest easily on its ventral surface. The inching along by means of the muscles at- tached to the pelvic girdle as given in Evermann and Kendall (1905) for an allied species was not observed, although no doubt the rough pelvic spines with their backwardly directed spinules act as a resistance to a possible backward drag. See also Johnson (1912). Loricaria altipinnis and other elongate loricariates with greatly depressed peduncles seem to approach the anguilliform type of locomotion, but as the peduncle is constructed to cleave the water and not react to it, virtually all of the effective thrust is centered in the caudal fin. As this then swings from side to side its action more closely approaches that of the ostraciiform type. That is, if the tail of the model be extended by an equally non-resisting rod, the effect would be nearly similar. Species studied in life — Family— CHARACINIDAE 49. Curimatus magdalenae Steindachner. 50. Characidium marshi Breder. 51. Piabucina festae Boulenger. 52. Compsura gorgonae (Evermann and Goldsborough). 53. Pseudocheirodon affinis Meek and Hildebrand. 54. Brycon striatulus (Kner). 55. Aphyocharax rubropinnis Pappenheim. 56. Hyphessobrycon bifasciatus Ellis. 252 Zoologica: N. Y. Zoological Society [IV; 5 57. Astyanax ruberrimus Eigenmann. 58. Creagrutus affinis Steindachner. 59. Bryconamericus emperador (Eigenmann and Ogle). 60. Hemibrycon dariensis Meek and Hildebrand. 61. Gephyrocharax atricaudata Meek and Hildebrand. 62. Thoracocharax maculatus (Steindachner). 63. Roeboides occidental is Meek and Hildebrand. 64. Ctenolucius beani (Fowler). 65. Hoplias malabaricus (Bloch). Family— GYMNOTIDAE 66. Gymnotus carapo Linnaeus. 67. Sternopygus dariensis Meek and Hildebrand. 68. Hypopomus brevirostris (Steindachner). Family— CYPRINIDAE 69. Carpiodes velifer (Rafinesque). 70. Catostomus commersonii (Lacepede). 71. Catostomus nigricans Le Sueur. 72. Erimyzon sucetta oblongus (Mitchill). 73. Moxostoma macrolepidotum (Le Sueur). 74. Cyprinus carpio Linnaeus. 75. Carassius auratus (Linnaeus). 76. Tinea tinea Linnaeus. 77. Scardineus erythrophthalmus (Linnaeus). 78. Campostoma anomalum (Rafinesque). 79. Chrosomus erythrogaster Rafinesque. 80. Semotilus bullaris (Rafinesque). 81. Abramis crysoleucas (Mitchill). 82. Notropis procne (Cope). 83. Notropis cornutus (Mitchill). 84. Rhinichthys atronasus (Mitchill). 85. Exoglossum maxillingua (Le Sueur). 86. Brachydanio rerio (Hamilton, Buchanan). 87. Brachydanio albolineatus (Blyth). 88. Danio malabaricus (Jerdon). 89. Puntius semifasciolatus Gunther. 90. Puntius conchonius (Hamilton, Buchanan). 91. Misgurnus fossilis fossilis (Linnaeus). Family— SILURIDAE 92. Felichthys felis (Linnaeus). 93. Ictalurus punctatus (Rafinesque). 94. Ictalurus fur catus (Le Sueur). 95. Ameiurus natalis (Le Sueur). 96. Ameiurus nebulosus (Le Sueur). 97. Schilbeodes gyrinus (Mitchill). 98. Clarias anguillaris (Linnaeus). 1926] Breder: Locomotion of Fishes 253 99. Rhamdia wagneri (Gunther). 100. Pimelodella chagresi (Steindachner). 101. Pimelodus clarias punctatus (Meek and Hildebrand). 102. Ageneiosus caucanus Steindachner. 103. Astroblepus longifilis (Steindachner). 104. Cory dor as paleatus Steindachner. 105. Hoplosternum punctatum Meek and Hildebrand. Family— LORICARIIDAE 106. Plecostomus plecostomus panamensis Eigenmann. 107. Lasiancistrus planiceps (Meek and Hildebrand). 108. Chaetostomus fischeri Steindachner. 109. Ancistrus spinosus Meek and Hildebrand. 110. Loricaria altipinnis Breder. 111. Loricaria variegata Steindachner. 112. Sturisoma panamense (Eigenmann and Eigenmann). 113. Stirisoma citurense (Meek and Hildebrand). Suborder— Synbranchii The swimming of this group is typically anguilliform as it must be on account of the shape and lack of locomotor appendages. Burrowing is a prominent characteristic and the speed with which these fish can enter loose soil is remarkable. Their speed through open water is also comparatively great as they habitually simply dash from cover to cover never remaining in an exposed position for long. Species studied in life — Family— SYNBRANCHIDAE 114. Synbranchus marmoratus Bloch. Suborder — Apodes These forms all show the typical anguilliform movements; those with pectoral fins using them simply in very small adjust- ments of position, chiefly when resting on the bottom. Their use is especially prominent when backing the cephalic part while the rest is in contact with some surface. Water expelled through the nearly circular gill openings is, of course, of use only as straight reaction, as well as on account of the shape of the body. Their very positive thigmo taxis causes the ejection of water to be of no consequence when resting. In some forms such as Myrichthys , in addition to the more common general body movement, undulation may be localized in the dorsal and anal. 254 Zoologica: N. Y. Zoological Society [IV; 5 It is interesting to note that many of these start life as ribbon- shaped larvae, the leptocephali (Fig. 68), which are pelagic in habit. As soon as the ribbon shape disappears and they become more nearly circular in cross section the pelagic habit is lost and life is taken up along shores, much of which time is spent in contact with solids. See pages 166-169, 175-183, 219, 114, 235, 237, and Figs. 39, 45, 46, 49, 68, for other data. Anguilla rostrata not infrequently arches its body and “coasts” downward by gravity alone, controlling the direction by turning the head and the radius of the arc which the body describes. Species studied in life — Family— ANGUILLIDAE 115. Anguilla rostrata (Le Sueur). 116. Myrichthys oculatus (Kaup). Family— MURAENIDAE 117. Gymnothorax funebris Ranzani. 118. Gymnothorax moringa (Cuvier). 119. Echidna catenata (Bloch). Suborder —Haplomi The propulsive methods of this group are about equally divided between body movements which are sub-carangiform and pectoral efforts, although the group includes a great variety of forms. Here, for the first time, species are found in which the pectorals take a considerable part in the production of a forward thrust (barring, of course, the rajiform movements). In the long bodied forms, with the pelvics placed well behind the pectorals, the movements are somewhat as in Lepisosteus. In comparatively many of the short bodied forms, the pectorals take on a large share of the progressional effort. As would be expected, the more compressed forms display a greater bending of the body than do the forms more nearly cir- cular in cross section. See pages 202, 203, 205, 207, 210, 211, 214-215, 220, 222, 235, and Figs. 57A, B, 57, 59B, 61, for other data. Umbra pygmaea habitually uses the pectorals to a great extent as pointed out on page 235. Their characteristic position of rest is diagonal to the surface, with the caudal portion drooping. At such times the pectorals and dorsal may show only as a blur as they move with great rapidity. What effect there may be from exhaled water is apparently taken care of by an excess of the 1926] Breder: Locomotion of Fishes 255 pectoral over the dorsal action. The cycles of the pectorals and dorsal are syn- chronized and are about twice as fast as those of respiration. Any sudden sound causes a pronounced increase in the rate of these with no change in position. Slow forward motion is effected by the pectorals, but rapid movement by a sub-carangiform body movement. Mollienesia latipinna at times slightly undulates the large dorsal fin (the male only) for progressional purposes. Xiphophorus helleri in its “love play” performs its backing movements by means of the pectorals in such alteration that a hesitant movement results. Other ovoviviparous species, such as Lebistes reticulatus, perform in a manner closely similar in their amorous maneuvers. Platypoecilia maculatus, being short-bodied, has a practically carangiform movement. Rivulus uropthalmus and chucunaque, being longer bodied, tend to approach the anguilliform method when not employing the pectorals alone for propulsive purposes. Species studied in life — Family— ESOCIDAE 120. Esox americanus (Gmelin). 121. Esox reticulatus (Le Sueur). 122. Esox lucius Linnaeus. 123. Esox masquinongy Mitchill. 124. Umbra pygmaea (De Kay). Family— CYPRINODONTIDAE 125. Gambusia affinis (Baird and Girard). 126. Priapichthys tridentiger cana (Meek and Hildebrand). 127. Priapichthys darienensis (Meek and Hildebrand). 128. Lebistes reticulatus (Peters).' 129. Heterandria formosa (Girard). 130. Phalloceros caudomaculatus (Hensel). 131. Mollienesia latipinna Le Sueur. 132. Mollienesia sphenops (Cuvier and Valenciennes). 133. Mollienesia caucana (Steindachner). 134. Xiphophorus helleri Heckel. 135. Platypoecilus maculatus pulcher Boulenger. 136. Platypoecilus maculatus ruber Stausch. 137. Cyprinodon variegatus Lacepede. 138. Jordanella floridae Goode and Bean. 139. Fundulus majalis (Walbaum). 140. Fundulus heteroclitus macrolepidotus (Walbaum). 141. Fundulus diaphanus (Le Sueur). 142. Fundulus luciae (Baird). 143. Fundulus bermudae Gunther. 144. Fundulus chrysotus Holbrook. 145. Panchax panchax (Hamilton, Buchanan). 256 Zoologica: N. Y. Zoological Society [IV; 5 146. Panchax lineatus Cuvier and Valenciennes. 147. Panchax chaperi (Sauvage). 148. Aphyosemion australe (Rachow). 149. Rivulus uropthalmus Gunther. 150. Rivulus chucunaque chucunaque Breder. 151. Rivulus chucunaque sucubti Breder. Suborder — Heteromi Apparently these fishes propel themselves by anguilliform or near anguilliform methods, as well as by using the long anal fin after the manner employed by Gymnotus. The pectorals in some forms doubtless have some locomotor function. Suborder — C atosteomi The locomotor efforts of the majority of fishes comprising this sub-order are confined to movements of the fins. In many, the encasing armament precludes any extensive body movement. Extreme and unusual specializations of various sorts are character- istic of the group allowing of no very general statement concerning their locomotor habits beyond the above, for they range from such large pelagic forms as Lampris luna (Gmelin) to tiny gasterosteids and syngnathids living in very special littoral environments. See pages 177, 194, for other data. Apeltes quadracus and other gasterosteids have pectoral movements not dissimilar to those of Umbra. See page 235. Aeoliscus strigatus (Gunther) and other members of this group swim with the axis practically vertical for most part although they may assume any other position. See Willey (1902) for an illustration of this species in action. Fistularia tabacaria Linnaeus appears not to have any locomotor function connected with the great prolongation of the central caudal ray. Syngnathus fuscus when highly excited may lash its body into anguilliform curves but makes slight progress, the pectorals and dorsal being the chief loco- motor organs. The latter is undulated as in Amiatus. Hippocampus hudsonius which usually moves with the backbone in a nearly vertical position is necessarily modified accordingly. In rising or falling in the water appropriate localized undulations in the dorsal give the desired motion on account of its position. A violent flapping from side to side, coupled with the action of the pectorals drives the fish slowly forward, ventral side foremost. However, when high speed, for a seahorse, is desired it leans forward with the vertebral column nearly horizontal and undulates the dorsal in localized anguilliform fashion. The much reduced anal may also be vibrated. With these various combinations of fins, any nicety of .spatial adjustment may be at- tained. Generally speaking, the pectorals and dorsal are used in conjunction for the fastest movements, and the higher the speed attained the nearer the 1926] Breder: Locomotion of Fishes 257 vertebral axis approaches the horizontal, thus drawing the divergent forces closer together and more nearly parallel to the axis of motion, so allowing for the efficiency necessary for a given speed. See Ryder (1881) for further dis- cussion of the locomotion of Hippocampus. Species studied in life — Family— GASTROSTEIDAE 152. Pygosteus pungitius (Linnaeus). 153. Gasterosteus bispinosus Walbaum. 154. Apeltes quadracus (Mitchill). Family— SYNGNATHIDAE 155. Syngnathus fuscus Storer. 156. Syngnathus mackayi (Swain and Meek). 157. Syngnathus elcapitanense (Meek and Hildebrand). 158. Hippocampus hudsonius De Kay. Suborder — Percesoces The locomotion of the members of this group is typically sub-carangiform, except in the short deep forms where pectoral pro- pulsion is well developed, as in the Stromatidae. In most elongate forms there is not as close an approach to the anguilliform type of motion as might be expected judging from the form alone. These, such as the tylosurids, are rather rigid fishes and exhibit no great flexibility, somewhat resembling Esox in simple locomotor or maneuvering efforts. Many limited specializations are found in this group, skittering over the surface and flying, finding its greatest development here. Other members are skilled leapers. See pages 203, 221, 222, for other data. Tylosurus marinus in maintaining a stationary position frequently fans its tail from side to side and compensates with the pectorals. In shifting its position when so resting, which is frequently done, a single flexure of the long body accomplishes this. The forward glide so initiated is nullified by the action of the pectorals, practically turning the fish on a point. This genus and its relatives, more especially the hemiramphids, are expert at leaping and skittering over the surface of the sea. Their bodies are held at an angular elevation from the surface of about 30° or even more in an extremely rigid manner and their submerged tail is vibrated rapidly. This is usually alternated with short leaps from which they alight tail first for further activity, or may sometimes plunge head first for submersion (Fig. 76). Cypselurus sp. and related^ “four-winged” forms doubtless have better control of their flight than the “two-winged” species, Halocypselurus, for the reason that the pelvics, which serve as posterior warping planes, give them more 258 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 76. Hemiramphus leaping and skittering. Note the functioning of the hypobatic tail in skittering. purchase for their limited amount of maneuvering. The latter more closely resemble the flying characins, Gasteroplecus, etc. See Fig. 73, pages 221 and 250. Menidia menidia, despite its somewhat elongate appearance, is an extremely stiff form. The pectorals are rather stiffly articulated and do little more than aid in turning and wheeling. They are capable of extremely sudden and rapid darts which are entirely dependent on the operation of the lateral muscles. Mugil curema displays somewhat more flexibility than do the Atherinidae although the movements are of the same order. This genus is marked by unusual leaping ability. A straight path is followed in the leap in all cases ob- served by the author. Sphyraena barracuda and related forms partake of pike-like movements except that they are inclined to show greater flexibility, in continuous swimming. Poronotus triacanthus and related forms, such as Peprilus paru, are il- lustrative of the suppression of ventrals in short deep bodied forms. On the other hand, Nomeus has extremely large ones, the exact use of which is not clear. They swim both by carangiform movement and pectoral operation. However, on removal of the pectorals, they seem to be able to navigate quite well and re- main in an upright position. Channa fasciata, with its long dorsal fin has not been observed to operate it after the manner of Amiatus, although structurally it might well be expected to. The locomotor efforts are mainly concerned with the body muscles, which tend toward the anguilliform type. The pectorals are used as active paddles and in backing, in the conventional manner. Anabas testudineus, being a short bodied fish living in weedy places, natur- ally uses the paired fins in moving about to considerable extent, and in obtaining a nicety of position. Its famed scansorial powers are apparently exaggerated or are inhibited except under very favorable circumstances. Captives removed from an aquarium appear extremely distressed, and while the operculi are thrown outward and the spines prominent, no attempt has been seen to use them except when the fish was placed in a sloping groove. Then, by the simple wriggling and flapping of the tail, the opercular spines acted rachet-like, and inched the fish slowly along. Species studied in life — Family— SCOMBRESOCIDAE 159. Tylosurus marinus (Walbaum). 160. Tylosurus timucu (Walbaum). 161. Hemiramphus sp. 162. Cypselurus sp. 1926] Breder: Locomotion of Fishes 259 Family— AMMODYTIDAE 163. Ammodytes americanus De Kay. Family— ANTHERINIDAE 164. Hepsetia stipes (Muller and Troschel). 165. Thyrina chagresi (Meek and Hildebrand). 166. Menidia menidia notata (Mitchill). Family— MUGILIDAE 167. Mugil curema Cuvier and Valenciennes. 168. Mugil cephalus Linnaeus. 169. Agonostomus monticola (Bancroft). Family— POLYNEMIDAE 170. Polydactylus octonemus (Girard). Family— SPHYRAENIDAE 171. Sphyraena barracuda (Walbaum). 172. Sphyraena borealis De Kay. Family— STROMATEIDAE 173. Peprilus paru (Linnaeus). 174. Poronotus triacanthus (Peck). Family— OPHIOCEPHALIDAE 175. Channa fasciata Steindachner. Family— ANABANTIDAE 176. Anabas testudineus (Bloch). Suborder — Anacanthini This group is marked by a tendency toward elongation and in contrast to the preceding by a considerable flexibility. Thus the movements are typically of a type approaching the anguilliform, the comparative slenderness of form being an index of the approach to this type of movement. The Macruridae, possessing an attenuate caudal however, doubtless make use of their long* anal by the application of localized undulations, and possibly also the pectorals to a considerable extent. See page 230 for other data. Microgadus tomcod often uses the pectorals to wing its way along in a some- what labriform manner. Phycis regius has been observed to bury itself in sand and reclining on its side with the head protruding, watch the movements of other fishes with apparent interest. 260 Zoologica: N. Y. Zoological Society [IV; 5 Phycis chuss in turning sometimes has its low long dorsal and anal flexed far to the outside of the curve. This is just the opposite of that which would cause the turn. Therefore it is inferred that the controlling muscles are simply relaxed and the water pressure causes this outward bend, it being only seen to occur in comparatively slow motion, and when other turning efforts were being obviously applied. Species studied in life — Family— GADIDAE 177. Merluccius bilinearis (Mitchill). 178. Microgadus tomcod (Walbaum). 179. Gadus callarias Linnaeus. 180. Lota maculosa (Le Sueur). 181. Phycis regius (Walbaum). 182. Phycis tenuis (Mitchill). 183. Phycis chuss (Walbaum). Suborder— Acanthopterygii Division— PERCIFORMES This great division while showing numerous specializations is marked by a carangiform or sub-carangiform type of body movement. As the body is short generally, frequently more so than that of Caranx, this necessarily follows, and even in some a faint suggestion of the ostraciiform movement is present. This group contains fishes of the greatest body depth known. Pectoral locomotion is common and used to a large extent in certain families, such as the Labridae and Scaridae, and a few forms, such as Malacanthusf make use of dorsal and anal undulations. See pages 175, 189, 194, 197, 207, 208, 215, 216, 226, 230, 236, and Figs. 50, 58, 59C and D, 60, 65A for other data. Aphredoderus sayanus with its large fins is sluggish and almost clumsy in its use of them in moving slowly about amid the weedy places it inhabits. Pomoxis annularis and the other generalized centrarchids are covered in the preceding section and need no especial treatment here. Thulia arge reminds one more of a characin in general swimming than its closer relatives. Boleosoma nigrum and the other darters use the pectorals in making their characteristic darts, usually a single stroke of them being all that is made before coming to rest again. Malacanthus plumeri, an unusually elongate form for this group, according to Mowbray uses both the anal and dorsal for locomotor purposes by means of localized undulations. 1926] Breder:' Locomotion of Fishes 261 Menticirrhus saxatilus, Eques acuminatum and others of these two genera make considerable use of a pectoral type of propulsion, although it is not other- wise especially characteristic of the Sciaenidae. Species studied in life — Family— BERYCIDAE 184. Aphreoderus sayanus (Gilliams). 185. Holocentrus ascensionis (Osbeck). Family— CENTRARCHIDAE 186. Pomoxis annularis Rafinesque. 187. Pomoxis sparoides (Lacep&de). 188. Ambloplites rupestris (Rafinesque). 189. Chaenobryttus gulosus (Cuvier and Valenciennes). 190. Enneacanthus obesus (Baird). 191. Mesogonistius chaetodon (Baird). 192. Lepomis auritus (Linnaeus). 193. Lepomis pallidus (Mitchill). 194. Eupomotis gibbosus (Linnaeus). 195. Micropterus dolomieu Lacepede. 196. Micropterus salmoides (Lacepede). 197. Kulnlia arge Jordan and Bollman. Family— CYPHOSIDAE 198. Kyphosus sectatrix (Linnaeus). Family— LOBOTIDAE 199. Lobotes surinamensis (Bloch). . Family— NANDIDAE 200. Badis badis Hamilton, Buchanan. 201. Polycentrus schombergki Muller and Troschel. Family— PERCIDAE 202. Stizostedion vitreum (Mitchill). 203. Stizostedion canadense (Smith). 204. Perea flavescens (Mitchill). 205. Boleosoma nigrum olmstedi (Storer). 206. Etheostoma flabellare Rafinesque. Family— SERRANIDAE 207. Roccus lineatus (Bloch). 208. Roccus chrysops Rafinesque. 209. Morone americana (Gmelin). 210. Cephalopholis fulvus (Linnaeus). 211. Epinephelus adscensionis (Osbeck). 212. Epinephelus striatus (Bloch). 262 Zoological N. Y. Zoological Society [IV; 5 213. Epinephelus guttatus (Linnaeus). 214. Epinephelus morio (Cuvier and Valenciennes). 215. Hypoplectrus puella (Cuvier and Valenciennes) 216. Promicrops itaiara (Lichtenstein). 217. Mycteroperca venenosa (Linnaeus). 218. Mycteroperca bonaci (Poey). 219. Myctoperca olfax (Jenyns). 220. Mycteroperca phenax Jordan and Swain. 221. Centropristes striatus (Linnaeus). 222. Diplectrum formosum (Linnaeus). 223. Pseudopriacanthus altus (Gill). 224. Neomaenis griseus (Linnaeus). 225. Neomaenis jocu (Bloch and Schneider). 226. Neomaenis apodus (Walbaum). 227. Neomaenis ay a (Bloch). 228. Neomaenis analis (Cuvier and Valenciennes). 229. Neomaenis buccanella (Cuvier and Valenciennes). 230. Neomaenis synagris (Linnaeus). 231. Ocyurus chrysurus (Bloch). 232. Theropon jarbua (Froskal). 233. Centropomis unionensis Bocourt. 234. Pomatomus saltatrix (Linnaeus). Family— SCIAENIDAE 235. Cynoscion regalis (Bloch and Schneider). 236. Bairdiella chrysura (Lacepede). 237. Sciaenops ocellatus (Linnaeus). 238. Leiostomus xanthurus Lacepede. 239. Micropogon undulatus (Linnaeus). 240. Menticirrhus americanus (Linnaeus). 241. Menticirrhus saxatilis (Bloch and Schneider). 242. Aplodinotus grunniens Rafinesque. 243. Pogonias cromis (Linnaeus). 244. Eques acuminatus (Bloch and Schneider). Family— GERRIDAE 245. Eucinostomus calif or niensis (Gill). Family— PRISTIPOMATIDAE 246. Haemulon parra (Desmarest). 247. Haemulon sciurus (Shaw). 248. Haemulon plumeri (Lacepede). 249. Haemulon flavolineatum (Desmarest). 250. Brachygenys chrysargyneus (Gunther). 251. Bathystoma striatum (Linnaeus). 252. Bathystoma rimator (Jordan and Swain). 253. Anisotremus surinamensis (Bloch). 1926] Breder: Locomotion of Fishes 263 254. Anisotremus viriginicus (Linnaeus). 255. Orthopristis chrysopterus (Linnaeus). Family— SPARIDAE 256. Stenotomus chrysops (Linnaeus). 257. Calamus bajonado (Bloch and Schneider). 258. Calamus arctifrons Goode and Bean. 259. Lagodon rhomboides (Linnaeus). 260. Archosargus unimaculatus (Bloch). 261. Archosargus probatocephalus (Walbaum). Family— MULLIDAE 262. Upeneus maculatus (Bloch). Family— CHAETODONTIDAE 263. Chaetodipterus faber (Broussonet). 264. Chaetodon ocellatus Bloch. 265. Scatophagus argus (Bloch) 266. Pomacanthus arcuatus (Linnaeus). 267. Pomacanthus paru (Bloch). 268. Angelichthys isabelita Jordan and Rutter. 269. Angelichthys ciliaris (Linnaeus). 270. Angelichthys townsendi Nichols and Mowbray. Family— TEUTHI D IDAE 271. Teuthis caeruleus (Bloch and Schneider). 272. Teuthis hepatus Linnaeus. 273. Teuthis bahianus (Castlenau). Family— OSPHROMENIDAE 274. Macropodus cupanus cupanus (Cuvier and Valenciennes). 275. Macropodus cupanus dayi Kohler. 276. Macropodus chinensis (Bloch). 277. Macropodus opercularis (Linnaeus). 278. Betta splendens Regan. 279. Colisa fusciata (Bloch and Schneider). 280. Colisa lalia (Hamilton, Buchanan). Family— CICHLIDAE 281. Haplochromis multicolor Hilgendorf. 282. Hemichromis bimaculatus Gill. 283. Cichlasoma nigrofasciatum (Gunther). 284. Cichlasoma faceatum (Jenyns). 285. Cichlasoma calobrense Meek and Hildebrand. 286. Cichlasoma umbriferum Meek and Hildebrand. 287. Cichlasoma tuyrense Meek and Hildebrand. 264 Zoologica: N. Y. Zoological Society [IV; 5 288. Cichlasoma maculicauda Regan. 289. Aequidens coeruleopunctatus (Kner and Steindachner). 290. Neetroplus panamensis Meek and Hildebrand. 291. Geophagus crassilabris Steindachner. 292. Pterophyllum sclare (Cuvier and Valenciennes). Family— POMOCENTRIDAE 293. Eupomocentrus leucostictus (Muller and Troschel). 294. Abudefduf saxatilis (Linnaeus). Family — LABRIDAE 295. Lachnolaimus maximus (Walbaum). 296. Harpe rufa (Linnaeus). 297. Tautoga onitis (Linnaeus). 298. Tautogolabrus adspersus (Walbaum). 299. Iridio radiatus (Linnaeus). 300. Iridio bivittatus (Bloch). 301. Thalassoma bifasciatus (Bloch). 302. Doratonotus decoris Evermann and Marsh. Family— SCARIDAE Pseudoscarus quacamaia (Cuvier). Pseudoscarus plumbaeus Bean. Scarus caeruleus (Bloch). Sparisoma abligardi (Bloch). Sparisoma viride (Bonnaterre). Sparisoma hoplomystax (Cope). Division— SCOMBRIFORMES In this group are found those forms after which the carangi- form type of locomotion was named. They are a rather stiff bodied aggregation, and even the longer forms represent movements close to the above named type. The pectorals are typically stiff and with- out much mobility and are used almost entirely for wheeling and turning. Abrupt stops are unknown. They are mostly fast swim- mers of great endurance. The relatively great amount of muscular tissue speaks eloquently of the extraordinary natatorial powers, as does the beautiful response of body and appendage form to stream- line conditions. Considering these fishes as machines their relatively small “ furnaces” and large propelling “engines” imply that they must consume their “fuel” rapidly, if only to be able to provide room for more, that is, they run their “boilers” at a high rate and on readily “combustible” materials of high grade. The fact that they have a high body temperature for fishes is noteworthy in this 303. 304. 305. 306. 307. 308. 1926] Breder: Locomotion of Fishes 265 connection. This is contrasted with many sluggish fishes that live on relatively indigestible foods and work it through long digestive tracts while quietly poking around, as for example, Catostomus. The following percentages are based on data gathered by Corson (1923) and illustrate this proportion roughly. TABLE IV Species Percentage of viscera to entire weight Gymnosarda alleterata (Rafinesque) .05 + Thunnus thynnus (Linnaeus) . 05 — See pages 175, 183, 194, 203, 207, 211, 216, 226, 227, and Figs. 44, 45, 47, 57C for other data. Caranx crysos in common with the other slimmer peduncled forms, has a prominent strong keel flanking it which doubtlessly functions chiefly as a strengthening brace. Rachycentron canadus when adult has pectorals that are shark-like in shape and use. There is, accompanying this, a slight approach to an epibatic con- dition of the tail, the upper margin being very slightly longer than the lower and somewhat stiffer, which may possibly be connected in some way with the manner in which the pectorals are used. In this connection it might be men- tioned that the young (about 80 mm. long) have large, fan-shaped isobatic tails and use the pectorals in a labriform manner. Sarda sarda and other forms with finlets apparently use them as multiple rudders. Their proximate margins are slightly concave and it seems that they adhere to the body as do the dorsals of Tarpon and some sharks. The exact mechanical advantage of these finlets is not clear although it may be that it allows an unhampered continual use of the caudal and body movements. Here again in the Scombridae, the peduncle is strengthened with a keel or keels if slender. Istiophorus nigricans (Lacepede) must possess a powerful keel in the en- larged dorsal fin but its exact use is not evident, although it is reputed to project the fin above the surface and actually sail by that means. Species studied in life — - Family— CARANGIDAE 309. Seriola zonata (Mitchill). 310. Selar crumenopthalmus (Bloch). 311. Caranx hippos (Linnaeus). 312. Caranx crysos (Mitchill). 313. Caranx ruber (Bloch). 314. Alectis ciliaris (Bloch). 315. Vomer setapinnis (Mitchill), 316. Selene vomer (Linnaeus). 317. Trachinotus falcatus (Linnaeus). 266 Zoologica: N. Y. Zoological Society [IV; 5 318. Trachinotus goodei Jordan and Evermann. 319. Trachinotus carolinus (Linnaeus). Family— RHACHICENTRIDAE 320. Rachycentron canadus (Linnaeus). Family — SCOMBRIDiE 321. Scomber scombrus Linnaeus. 322. Scomber colzas Gmelin. 323. Sarda sarda (Bloch). 324. Scomberomorus maculatus (Mitchill). Division — ZEORHOMBI This division contains greatly compressed forms with short bodies, whose locomotion must be typical for fishes of such propor- tions, excepting those that have developed an asymmetry con- comitantly with their habit of lying on one side. These, the Pleuronectidse, even when swimming retain the reclined position, although the undulations of the body go on as before. This places the plane of undulation in a vertical position from which fact the family name is derived. The great compression of the body makes comparatively short undulations possible, and while viewed in profile, they must be considered short and deep, their movement approaches that of the anguilliform because of the above mentioned thinness. The exhalation from the under gill slit no doubt aids in the send-off, by its impact on the bottom, as do both those of the skates. Turning is effected partly by the paired fins and partly by movements of the body. See page 203, for other data. Pseudopleuronectes americanus as well as others, in slow movement over the bottom undulate the marginal dorsal and anal with the tips of the rays in contact with the solid substratum thereby securing greater thrust. Lophopsetta maculata and probably all pleuronectids in resting on the bottom often draw the anterior rays backward and the posterior ones forward so bracing themselves against any strong current that might tend to dislodge even so flat a form. In inaugurating a sally t6 the open water, they hasten their start by pushing off by means of the fringing rays. Fig. 77 shows this and indicates how the rays are thrust back after such a start. It is clear that the caudal portion of the fin and the first few rays have the most effect because of the larger arc through which they swing. Achirus fasciatus attaches itself to smooth surfaces by pressing the fringing fins downward and raising the central part of the body thus producing con- siderable suction. In fact, most of the members of the present family possess 1926] Breder: Locomotion of Fishes 267 Fig. 77. Lophopsetta maculata with dorsal fin rays in a resting position with the manner in which they are thrown back in starting indicated. Four rays are shown in the new posi- tion with the path their tips traversed, as well as the margin of the fin. The anal operates in a similar way. this faculty to a slight degree, but it finds its greatest expression in this genus. Vertical surfaces are even commonly so clung to. Species studied in life — Family— PLEURONECTIDAE 325. Paralichthys dentatus (Linnaeus). 326. Paralichthys oblongus (Mitchill). 327. Pseudopleuronectes americanus (Walbaum). 328. Lophopsetta maculata (Mitchill). 329. Etropis microstomus (Gill). 330. Citharichthys gilberti Jenkins and Evermann. 331. Achirus fasciatus Lacepede. 332. Achirus panamensis (Steindachner). Division— KURTIFORMES The single species of this division necessarily must confine its body movements to the carangiform, partly on account of its general shape, but more especially because of the anterior stiffening effect that the ossification of the outer membrane of the swim-bladder implies. Very likely the long anal is the seat of localized undulations and the pectorals of flapping movements somewhat like those of the Labridae. Division— GOBIIFORMES This group is marked by a quiescent nature. Many of the species behave in a manner quite similar to the darters, swimming from place to place simply by single strokes of the pectorals between rests. A sub-carangiform method of propulsion is found in others. See page 219 for other data. 268 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 78. Front view of Echeneis naucrates. Note positions of the paired jins. Philypnus maculatus t a comparatively well rounded form; has rather small pectorals and leads a more active life than do such species that are prominently flattened above and possess large pectorals. Sicydium salvini and other forms with united ventrals by employing them as a sucker are enabled to remain in one place in a strong flow without active muscular effort. Species studied in life — Family— GOBIIDAE 333. Dormitator maculatus (Bloch). 334. Dormitator latifrons Richardson. 335. Eleotris pisonis (Gmelin). 336. Eleotris picta Kner and Steindachner. 337. Philypnus maculatus (Gunther). 338. Sicydium salvini Grant. 339. Gobius soparator Cuvier and Valenciennes. 340. Gobius daguae Eigenmann. 341. Microgobius miraflorensis Gilbert and Starks. Division— DISCOCEPHALI The swimming movements of this group vary from a sub- carangiform to an anguilliform type depending on the length of the species considered. The paired fins, of nearly equal size, are placed one pair over the other, so that when viewed end on, as in Fig. 78, they almost divide the fish into quadrants. This placing of these fins appears to be associated with the ability of the fish to rotate on its longitudinal axis in swimming, apparently turning to any such position with ease. This is clearly necessary to insure attachment to sharks or other bodies. That fishes of such natatorial powers possess such a highly specialized organ as the cephalic disc is interesting, for they have apparently suffered no particular degeneration of their power to care for themselves in a locomotor sense, for of course they must first catch their shark. In this 1926] Breder: Locomotion of Fishes 269 connection, their square tail is of special significance as short spurts of speed are naturally requisite. Echeneis naucrates, an elongate form frequently attached itself to the glass front of its tank and indulged in what appeared to be the regular swimming motions. A careful measurement (traced on tissue held against the glass) showed the tail’s tip to describe an arc, the chord of which equaled 29% of the entire length of the fish. While the head was of course restricted in its move- ment on account of the attaching disc, the tail’s tip swung an arc of just 15° or 7.5° to either side of the axis as measured from a point over the axis vertebra. Remora remora is skilled at leaping for a fish of its build, whereas the former being much longer bodied is not. Species studied in life — Family— ECHENEIDIDAE 342. Echeneis naucrates Linnaeus. 343. Remora remora Linnaeus. Division— SCLEROPAREI The body movements of this group are sub-carangiform in the main. Being bottom fishes, the pectorals of many are developed as propelling organs which they use much as do the darters and some gobies. In others, such as the Triglidae, the pectorals are used as gliding planes and still in others, the Dactylopteridae, they are carried a step further and the fishes rise from the surface of the water by means of their large “wings, ” somewhat after the manner of the exocoetids, but less successfully. However, the wings vibrate rapidly and their flight has been compared to that of grasshoppers. It seems likely, however, that the vibration is a mechanical effect caused by the plane-like fins being forced edgewise through the air. The Cyclop teridae are said to be able to creep over rocks by manipulating their ventral sucker. See pages 203, 221, for other data. Prionotus evolans and carolinus when ceasing active swimming generally extend their large pectorals and the weight of the fish (although it possesses a swim-bladder) which is heavier than the water generally, enables them to glide forward at will, wheeling about by “warping” the planes. Sometimes they will “volplane” through the water in wide circles by this means alone. Species studied in life — Family— COTTIDAE 344. Uranidea gracilis (Heckel). 345. Coitus ictalops (Rafinesque). 346. Myoxocephalus octodecimspinosus (Mitchill). 347. Myoxocephalus aeneus (Mitchill). 270 Zoologica: N. Y. Zoological Society [IV; 5 Family— TRIGLIDAE 348. Prionotus carolinus (Linnaeus). 349. Prionotus evolans strigatus (Cuvier and Valenciennes). Division— JUGULARES This division shows typically an intermediate position between the anguilliform and carangiform types of movement, although some approach the former closely. The ventrals may be reduced to a few rays in bottom forms or united to form a sucking disc somewhat similar to that of some of the gobies. The movements of such forms are generally quick darts made in a manner not very dissimilar to those made by the darters, some gobies and the Cottidae. Species studied in life — Family— BLENNIIDAE 350. Labrisomus nuchipinnis (Quoy and Gaimard). 351. Blennius pholis Linnaeus. 352. Blennius cristatus Linnaeus. 353. Salariichthys textilis (Quoy and Gaimard). Family— BATRACHIDAE 354. Opsanus tau (Linnaeus). Division—' TAENIOSOMI These deep water forms, Du Bois-Reymond (1914) states, move forward slowly by means of undulating the long dorsal. He doubts that they can effect an anguilliform movement on account of their extreme thinness and delicacy, but apparently fails to realize that at the depth at which they normally live, this delicacy disappears as in all deep water forms in their natural habitat. Schlesinger (1911, a) has shown that they undulate the body in curves of large amplitude, not unlike those of the leptocephali of apodal fishes, as well as employing the method mentioned by Du Bois-Reymond. The peculiar caudal doubtless forms an efficient rudder, and may possess a somewhat epibatic effect. Division — ATELAXIA The locomotion in this group is probably highly similar to that of the preceding. The body is doubtless convoluted in an anguilliform manner and similar movements are probably localized 1926] Breder: Locomotion of Fishes 271 in the long low dorsal. The extremely attenuate caudal wisp is, however, probably without any particular course-determining significance. Suborder — Opisthomi The movement of Mastacembelus appears to be somewhat after the manner of Gymnotus and Gymnarchus combined, and not simply anguilliform as sometimes illustrated, although it is likely that they resort to such a method when high speed is requisite. That is, in general locomotion both the dorsal and anal are seats of localized undulation. * Suborder — Pediculati These highly specialized fishes show a sub-carangiform move- ment when body contortions are attempted. The angulated paired fins are used as paddles and in many cases resemble feet more than they do fins, the mechanics of which are evident. * See page 194, for other4 data. Lophius piscatorius pushes the body along the bottom at times by kicking back with the ventrals. Swimming is accomplished by undulating the caudal portion of the body, the large head having small sidewise motions. Ogcocephalus vespertilio (Linnaeus) and other similar forms are reputed to use the fore and hind “limbs” in a manner reminiscent of frogs, the latter aiding in launching them on their way. Species studied in life — Family— LOPHIIDAE 355. Lophius piscatorius Linnaeus. Suborder — Plectognathi Division — SCLERODERMI This division is marked by an emphasis of the locomotor efforts being placed on the fins and a suppression of them in the trunk. In the rigidly encased forms, such as the Ostraciidae, the typical ostraciiform movement is seen, from which group it derives its name. The dorsal and anal are typically a seat of localized an- guilliform undulations, especially if rather lengthy as in Batistes (Fig. 48C) or of localized ostraciiform movements if short as in Lactophrys. See pages 166, 169-175, 177-183, 187, 194, and Figs. 40, 41, 42, 43, 48C, for other data. 272 Zoologica: N. Y. Zoological Society [IV; 5 Batistes vetula and others possess a long movable pelvis, which when erected seems to act as a keel. This is usually done on coming to a stop and at times when other fishes would erect their anal. Lactophrys triqueter and others of the same genus make comparatively slight efforts to escape when held in the hand. Even the jets which may be quite powerful at times are not ordinarily forced into action. The dorsal and anal, being very short are most frequently simply waved from side to side al- though at times a few small waves may form at their tips. The nearly spheroidal shape of the young (10 mm.) make their movements very sluggish and irregular as would be expected. Species studied in life — Family— BALISTIDAE 356. Balistes carolinensis Gmelin. 357. Balistes vetula Linnaeus. 358. Canthidermis maculatus (Bloch). 359. Melichthys piceus (Poey). 360. Monacanthus hispidus (Linnaeus). 361. Monacanthus ciliatus (Mitchill). 362. Alutera schoepfii (Walbaum). 363. Alutera scripta (Osbeck). Family— OSTRACIONTIDAE 364. Lactophrys triqueter (Linnaeus). 365. Lactophrys trigonus (Linnaeus). 366. Lactophrys tricornis (Linnaeus). Division— GYMNODONTES The extreme shortness of body of most of the members of this division limit them to an ostraciiform or sub -ostraciiform move- ment when the lateral myomeres are employed. However, the emphasis is placed on the pectorals, dorsal and anal in all forms known to the author. The force of the exhaled water is of con- siderable significance if from simple reaction alone. See pages 187, 190, 194, 202, 236, and Figs. 51, 55, for other data. Lagocephalus laevigatus most frequently fans its dorsal and anal from side to side (localized ostraciiform motion) in unison, but occasionally alternates them. The significance of this is not clear. Species studied in life — Family— TETRODONTIDAE 367. Spheroides maculatus (Schneider). 368. Spheroides harperi Nichols. 369. Spheroides sphengerli (Bloch). 1926] Breder: Locomotion of Fishes 273 370. Spheroides testudineus (Linnaeus). 371. Lagocephalus laevigatus (Linnaeus). Family— DIODONTIDAE 372. Chilomycterus schoepfii (Walbaum). 373. Diodon hystrix Linnaeus. Subclass — Dipneusti The appendages of Lepidosiren are nearly useless as locomotor organs on account of their size and shape but doubtless aid in maneuvering, while in Neoceratodus they must be of considerable importance. An intermediate between the anguilliform and car- angiform types of movement is here obtained, with a close approach to the former in some cases. Locomotion of Fossil Fishes. Although our knowledge of the earlier groups is fragmentary and that of their methods of locomotion is necessarily in the nature of inferences based on anatomical similarities to living forms, the physical forces at play in moulding these ancient fish bodies were the same in early geological ages as now, so by comparison a reasonable degree of certainty as to their methods of progression may be obtained, for it has usually been the external locomotor parts which have been preserved. Probably the most interesting fact to be brought out in this connection is that fishes have not been forced or able to make any great fundamental improvement in their methods of propulsion since the earliest pre-gnathous forms swarmed in Silurian and Devonian seas. True it is that a great number of highly specialized forms have arisen, but practically all these are kinds that have not given rise to great groups, since the specializations are of such natures as to confine them to more or less restricted areas and make them utterly unfit for wide dispersion or differentiation inter se. Examples are seen in Hippocampus, Histrio , etc., but as before mentioned, fishes which are still distinctly recogniz- able as such, that is, are still somewhat ichthyized, all use some form of undulation, generally in the body itself, or at least localized in certain members for purposes of progression. Even the examples used as exceptions have carried these undulatory methods along with them in little altered form to their remote positions, e. g. the dorsal and pectorals of Hippocampus. It may be truly said that no known fish, living or fossil, whatever be its modification for 274 Zoologica: N. Y. Zoological Society [IV; 5 Fig. 79. Pteraspis rostrata. Note the manner in which the curve of the rostrum deflects the water so as to offset the action of the epibatic tail. progression over solid surfaces, has ever been able or found it desirable to completely shake off the heritage of some form of undulatory movement for general progression while free in the water. This would seem to indicate that the optimum in method of animal locomotion through water had been reached long ago, as far as the limits of biological structures permit. It is equally true that for simple mechanical reasons, if nothing else, the effect of the exhaled water has had to be reckoned with, either as an advantage or disadvantage dependent on whether forward motion was desirable or not, from about the same early period. Examining such early forms of pre-gnathous fish-like animals as the Ostracodermata, we find among the Heterostraci such forms as Pteraspis rostrata, which, if the current restorations are correct, must have progressed somewhat after the manner of modern teleosts which are nearly rigid anteriorly. The anterior region was corseletted in a calcified test, from which extended a posterior tapering portion covered with small rhomboidal scales. Of this much we are certain and it is believed that the tail was heterocercal (epibatic). Only one inference can be drawn from the fact that the animal was covered with a scaly protection posteriorly, and that is that it was to gain flexibility. The restored length almost precludes the throwing of this section into a complete sine curve. Therefore it is believed that these fish had a motion not unlike that previously described as carangiform. The caudal could not have been decidedly epibatic, or horizontal swimming would have been impossible unless the shape of the snout acted somewhat after the manner of the pectorals in the sharks of today, as suggested in Fig. 79, for as far as known, this creature lacked any other appendages except the dorsal spike, which could hardly have even any function as a keel on account of its small blade area, unless it supported a membrane. The fact that the paired fins were wanting is of small consequence, as the tadpoles of frogs, for example, manage to swim rather effectively today without them, although of course all such forms have isobatic tails, to say nothing of the 1926] Breder: Locomotion of Fishes 275 surprising degree of success attained by teleosts deprived of their accustomed fins. The only loss seems to be one of a nicety of positional adjustment. It would seem that the pectorals were originally developed to offset the deflection of an epibatic or hypo- batic tail, and when once well established took on other significance. That Pteraspis was a free swimmer can hardly be doubted on account of its fusiform outline. Others of the same order seemingly propelled themselves similarly. Drepanaspis was probably a slow moving bottom form and probably could lash its tail into about one half a sine curve. Little can be said of Thelodus and Lanarkia as not much more than a crushed outline of them has been preserved. In the Osteostraci a similar manner of locomotion was doubt- less employed, but in the case of Cephalaspis, the test covered a short anterior portion and we know there was a metameral arrange- ment from there on posteriorly which no doubt corresponds to and is possibly homologus with the myomeres of the modern fishes. In this case, it may have been possible for the animal to throw itself into a full sine curve, although it is likely that it used less than this usually, probably swimming after the manner of the shorter bodied loricariates of today. The well developed dorsal “fin” formed a satisfactory keel and the pectoral lappets doubtless offset the action of the epibatic tail in driving the head downward, even though they appear to be much less flexible than pectoral fins. Passing on to the order Anaspida, we find a great group of fusiform animals which were doubtless somewhat free swimming in habit. The metameral arrangements of the body segments and the well developed caudal fin make it almost certain that these primitive forms swam in the manner roughly similar to the modern generalized cyprinids or characins. The anal fin functioned as a keel, while the dorsal row of spines probably aided it in that function to some extent. The pectoral appendages established by Kiaer (1924) might have been predicted on account of the tail form, for, as previously pointed out, only animals with isobatic tails can dis- pense with them, unless there is some other substituting factor. In the Antiarchi, we have bottom forms encased anteriorly and flexible posteriorly as the scale-like covering indicates. These forms so far as known drove themselves forward in a manner similar to the pteraspids, but with this difference, — that they had in addition well formed dorsal and pectoral appendages. Just what function these structures had is not clear, but in swimming they 276 Zoologica: N. Y. Zoological Society [IV; 5 were probably used somewhat after the manner of pectoral fins in turning and wheeling whilst they no doubt functioned also, while the animal was on the bottom, as ambulatory organs. In the Arthrodira, we find more conventional forms which evidently swam much after the fashion of modern fishes of similar general shape, and it is hard to imagine them lacking pectorals, on account of the tail form. The anterior armor extended nearly , half way back, thus delimiting the effective undulation to the posterior part of the body which no doubt in this case means that they were limited to not more than half a sine curve. Their denti- tion indicates a raptorial habit and as a consequence they were necessarily powerful and comparatively fast swimmers, as their form indicates. Among the first true jaw-bearing fishes, the Acanthodii are prominent. These forms had a full complement of fins and the gills opened externally by separate clefts as in modern sharks. They varied from short bodied fishes to those of rather elongate shape and judging from the squamation, were apparently flexible from the skull backward. The short bodied ones no doubt propelled them- selves in a nearly carangiform manner while the longer forms probably threw their bodies into a greater number of curves. The single spine supporting each fin allowed of no localized undulations as the posterior margins were adnate to the body. The probability is that the paired fins were used exclusively for wheeling and turning and offsetting the epibatic depression, while the median ones were simply keels. Here, for the first time, we find the gill clefts definitely marked as such and as they are in the conventional position and appear typical, we must assume, in the absence of evidence to the contrary, that these Devonian fishes must have gained some forward thrust from the exhaled water, as has been shown for recent fishes. This, coupled with the structure of the pectorals indicates that they must have moved somewhat after the manner of sharks and were unable to make abrupt stops, turning to avoid obstacles. From here on in all groups we find similar thoracic exits for the excurrent water, and further specific mention of them is not neces- sary. The Cladoselachii, while phylogenetically rather different from the above, have many points in common with them from a locomotor standpoint, but as all were elongate, they probably formed more 1926] Breder: Locomotion of Fishes 277 than one wave in moving forward. The better and more highly developed caudal suggests a greater speed than the acanthodians were capable of and the cartilage rays of the fin suggest more flexibility in these members and a consequent greater use than was possessed by the former. That is, they were probably able to “warp” the pectorals in a manner which slightly approached these movements in modern sharks. It seems reasonable to suppose that the Ichthyotomi undulated the long dorsal after the manner of Amiatus or Gymnarchus, holding the body rigid, and they may have paddled with the pectorals and pelvics as well, for these seem fitted for this method of movement. They represent the first forms with what appears to be really flexible paired appendages, such as are not again found until we come to the less primitive Teleostomi. Being swamp-inhabiting forms, it is probable that these latter two methods were employed for nice adjustments of position, as in the above mentioned Amiatus of today. The fossil dipnoans so nearly resemble the recent forms in general external formation, that what has been written concerning them applies equally well to Dipterus, the most primitive of them, while fossils of higher groups (Chondrostei, et cetera) more closely re- semble recent forms and need no separate treatment, being strictly comparable to recent forms. That is, the fossil sharks, actinop- terygians and crossopterygians are comparable with their various recent representatives, or, by parallel development, to other groups modified only by slight or obvious changes in form and structure. Various parallelisms are apparent, for example, Chirothrix being apparently a flying member of the Haplomi. PART III— CONCLUSION Recapitulation and Discussion. After analyzing the movements of fishes in the preceding two parts, we may review them from a broader aspect and consider the fundamental characteristics that are concerned with locomotion. Following is a tabulation of the larger divisions into which the swimming movements of fishes are divided, with an example showing- each characteristic. 278 Zoologica: N. Y. Zoological Society [IV; 5 Type op Flexures Anguilliform Carangiform13 Ostraciiform TABLE Y Body Anguilla Caranx Lactophrys Location op Flexures Median Members12 Gymnotus Batistes Lagocephalus Pectorals Chilomycterus Lepomis Scarus The only other swimming efforts of fishes, the exhalations of jets, may be considered in the following tabulation, using the same examples as those of Table V. TABLE VI Effect of Jets Type of Gill Orifice Slit Oval Circular Induced Streamline Caranx Intermediate Lactophrys Simple Jet Reaction Chilomycterus These two tables (V and VI) include all the basic efforts which fishes employ in swimming and trace them through their various modifications. This at once brings out forcibly the truth of the similarity of these undulatory movements. The only one not included in Table V is the “ Caudal undulation'' of page 189, not of great importance to any fish. This, at times, is similar to the dorsal movements of Hippocampus, and with it completes the list of every known fundamental motion. It will be noted that metamerism of the activating muscles is basically present in all, only in the ostraciiform pectoral oscillation is it reduced to a practically single muscular act; in all others, at least two or more units act serially. Even in the oldest fossil forms this same condition obtains. There is thus a great uniformity underlying this superficial diversity. Regarding Table VI little need be added, for it simply indicates that with a change from a slit-like opercular aperture to a circular one, whatever the significance of the exhaled water may be it changes from the induced streamline effect of Paxton to that of simple jet reaction. Note also that slit-like opercular ports are accompaniments of swift fishes with fusiform bodies and that other shaped exhalant vents not ejecting the water in a sheet are the property of fishes not especially fast nor of a shape directly respon- sive to streamline contours. The body forms of fishes unless so modified as to be of no streamline value at all, such as Ogcocaphalus, vary from the elongate 13 “ Carangiform ” is used here in a comparative sense, as an intermediate between “Anguilliform” and “Ostraciiform” in reference to “ Median members ” and “Pectorals.” See pages 189-190. 1926] Breder: Locomotion of Fishes 279 through the fusiform to the truncate. The central condition, “fish-like,” is that of high speed, whilst either extreme is slower. The continuation of carangiform movements in elongate types (anguilliform) gains for the fish greater pressure areas from its sides directly, but loses the terminal effect of the tail fin. In fact, the caudal fin is generally entirely lost, as it would be practically valueless in the extreme elongated forms. Here the direct thrust of the insinuating body is the sole propeller. The truncate fishes, being unable to flex their bodies effectively on account of their shortness, have the locomotor emphasis placed on the appendages. In these fishes armature can be afforded as the body flexures cease to be important and speed is impossible. Thus other protection is desirable. The fins, paired or median may be various but they run a similar gamut as far as their positions and forms allow. That is, they may partake of either anguilliform or ostraciiform movements. The carangiform is hardly represented in its true sense, but the intermediate type of movement may be considered as such, as indicated in Table V. The median fins if long and low have anguilli- form undulations localized in them; if high and short, they have ostraciiform movements. The pectorals if wide have the former, and if narrow the latter. The only movements which remain to be considered are those not to be considered as swimming, i. e., creeping, flying, et cetera. These are all narrow specializations which have no special bearing on the present connection, as they represent in each case a divergent off-shoot of rather limited possibilities. With this resume, the major groups of fishes may be listed according to their chief locomotor characteristics. Table VII thus tabulates twelve factors, each with three divisions, either extreme and the norm between them. While it does not pretend to be absolute or entirely complete, the belief is entertained that it gives a true representation of these twelve characteristics in all of the groups of fishes down to Divisions. X in a column indicates the presence of the condition mentioned at its head. The placing of an X not directly in a column, but between two, indicates an inter- mediate condition. Dashes indicate entire absence of the condition, or, in the case of certain fossil forms, incomplete information. It is evident in such a tabulation of intergrading factors and elements, that in many cases only an approximation can be given TABLE YII 280 Zoologica : N. Y. Zoological Society [IV; 5 j'einojio XX X Gil Ori flee IEAO 1TIS XXX xxx xxxx. X XX © ft e^'eoanjia X xxx XX XX cS .d eq.'eounjj, X X X XX CO G^'euTiunoy XX X XX XX xxx a O © oi^eqod^n Pi oi^'eqosi XX X X X X X X X XX xxx £h OT^-eqTda xxx •i >» QiqxxGia 1 1 X IX X X 1 © s G^'eipeuiao^ux 1 i 1 XX XX 1 ft -ft BPS 1 1 X X X 1 X XX 1 Sh B Avoaa'ej^; 1 1 1 X 1 o G^tpQUIJG^UI 1 1 X X X 1 X XX 1 © ft ft p^ojh 1 1 X 1 X XX X XX 1 A M raaojip'eaxso 1 X 1 1 x 1 1 1 X XX 1 1 a £ GX'BTpQUIJGXUI 1 1 X 1 X XX 1 ft 1:1 uuojnimSuy 1 1 1 1 X 1 1 "cS „ B qSiq pm3 poqs 1 1 X XX XX 1 .3 | o GX'eipGUIJGXUJ 1 1 X X XX XX XX 1 I 1 ft moi pui3 Suoq 1 1 X X xxx XX 1 ’So o , uuojiio'eaxso 1 1 1 1 1 1 1 1 1 1 d d O Sh © 2 GX'eipGUIJGXUI 1 1 1 1 1 1 1 ft 00 ft ^ raaojnnnSuy 1 1 1 X 1 1 X 1 1 1 X XX 1 •pH >» GiqtxGM XX XXX xxx xxx « -*2 © a GX'eipQtujQXux XX xxxx XX ft 2 BttS XX XX 1 d pGSSGJdGQ XX X J>» o 9 © .2 X^oupuii^O , XX xxx XX XX XX X X xxx xs o GO pGSSGJdraoo X X XX ft s QX'BOUtUX X X X' XX Sh O raaojisna XX X X xxx X XX XX ft GXeSuoig; XX X X xxx xxx Flex- ures uuojip'eaxso ' XX XX X X X ra.iojT8ui3.reo taaojnimSuy XX xxx XX X XX XX xxx d h Tj N ce >> ■s § g I ‘El i s r. © © X* -3 -d •” O u u Pl, HI O O HI 'fctO .pH b I -s 5 S 3 O >> 53 3-S g E “I Ph m < i-i ?-» i ® © o ! 3 'd Td f-i *h "■ _a 1 s 2 © w so .2 n So© 3 £ ft O Sh -Q © d V oa M S-4 W o o o o a *a cS d jb o PS +=> ft W © O §,2 ^H © w m Sh W © c3 xi 'o Sh i °-§ co t>> o ftOO M © © o xs xi ^ *h u O o o w « i £h .ft fd Sh Sh Sh © £ £ ® © © TJ CC CO 'd XJ Xj O 1 1 o 5 rd o O ft b0 s|g ||,s 8-2 <2 ^ >> CO £ « £ O Eh O O O &h X2 pO pO d d d CO CO CO O O O 1926] Breder: Locomotion of Fishes 281 X X X X 1 1 1 X 1 1 X X XX X XX 1 II II xxxxx xxxxxxxxxxx X 1 II II X XXX XX X XX X XX X XXX XXX XXXX X XXX X X X X XX X XXX X xxxx X X X X xxxxxx xxxxxxxxxxxx XXX X X XX XX xxxxx X X XX X XXX 1 1 1 XXX xxxxxxxxxxx 1 1 X 1 X X XX 1 1 XX 1 X XXX X 1 1 1 xxxxxx XXXXXXXXXX X XXX 1 1 XX 1 X XX X X XX X XX 1 1 X 1 XX XXX xxxx XX 1 1 1 1 X 1 xxxxxx xxxxxxxxxxxx XXX 1 III 1 X XX X X XX 1 III 1 XX X 1 1 X XXX XX XXXX X XX 1 1 xxxx X xxxx xxxx xxxx X X 1 1 i III 1 XX 1 1 1 1 1 1 1 X III 1 XX 1 IMII X 1 xxxx XXXX 1 1 1 XXXX 1 X X 1 1 1 II 1 xxxx XX XX xxxx X X X XX XX XX XX X X XX x X X X X XX XX XX XXX X XX XX XXX X XXXX X XX X x xxx xxxxx XXXX XXX XXX XX XXX X XX XX X X X X v XXX XAXXXX XXXXXXXX X XXX XX XX xxxx X X X X XX X X X X XX X XX XX X X X XXXXXXXX X X XXX XX xxxx X X a a § J . O Sh 0C on&g ft & ® ce M +* ® ft ft O P ft o a ^ S « -S H 5 $ ©HO ft <1 <1 H H H H H © o © o o TJ 'd ’d T3 tj H H H H H o o o o o .O -ft X2 X3 d d 25 IS 00 02 CO C/2 £2 £2 £> 3 3 3 00 00 00 H, M ■ » B 3 ® H ft ® © B as-gsa-sBy §1^11 § I'll § S o o ^ B ® 6(1 ® " flflflflf3fl?3f333 .2 .2 .2 .2 .2 .2 .2 .° .2 ’3 ‘3 ’3 ‘3 '3 "3 ‘3 '3 ‘3 > > '£ > ’£ > '£ '£ > > § § SqSSS5SpQQ32,q oo no £3 ’d » & O * 2 & rH ® 43 > -d ® © Jm aii M I! *.§ £ o § § 3 3 » o © p3 O :P oo H TABLE VII Class Cyclostomata1* Order Myxinoides .... Order Petromyzontes . Class Pisces HI si 111 312 11 ill 1 s lit llll!? der Plagiostoml . Sub-order Batoldei Order HolocophaU ub-class Teleostomi Order Crossopterygll .... Sub-order Ostoolepida16 Sub-order Cladlstia. . . . Order ( Order Teleostoi Sub-order Malacoptorygii . Sub-order Ostariophysl . . . Sub-order Synbranchii17 . . . Sub-ordor Apodes ' * . Sub-order Haploml . . Sub-order Heteromi . Sub-ordor Percesocos Sub-order Anacanthini Sub-ordor Acanthopterygii . , Division Perciformes .... Division Scombriformes . . Division Zeorhombi1*. . . . Division Kurtiformos .... Division Gobiiformes. . . . Division Discocepbali .... Division Scleroparoi Division Jugulares Division Taeniosomi'* Division Atolaxia19 Sub-order Opisthomi Sub-order Podiculati Sub-order Plectognatlii Division Sclerodormi Division Gymnodontes. . . Sub-class Dipneustl Fossil forms of uncertain affinities Palaoospondylidao Ostracodormi Order Heterostraci Order Osteostraci Order Anasplda11 Antiarchi Arthrodira xxx : xxx : xx i includes the longitudinal pectorals of the Batoidei. In other groups the dorsal and anal only, s multiple. s of this order have distinctly trilobed tails which are isobatlc. s united ventrally, median. group are rotated through 90° — the Pleuronectidae. of these Ashes if ever used in a locomotor sense is doubtless somewhat epibatic on account of their upturned i 282 Zoologica: N. Y. Zoological Society [IV ; 5 and a considerable part must rest on the judgment of the compiler. In cases where definite information or first hand knowledge was not available the positions of the check marks are based on a study of the extensive literature on fish locomotion, inferences from anatomi- cal data, et cetera. The data concerning fossil forms is based on similarities with existing fishes as noted under “Locomotion of fossil fishes.’' The maneuvering abilities of fishes are accomplished, in the main, by differential actions of the locomotor apparatus combined with the presenting of planes to the water when in motion. There are therefore few external structures devoted solely to maneuvering in fishes. On the other hand, the organs concerned with locomotion other than swimming are generally of some definite and highly specialized development. The fins primitively were broad based and relatively fixed in position, (Gregory, 1914) and later took on distinct movements with the necessary accompaniment of a com- paratively restricted base. The habits of fishes are naturally closely connected with their locomotion. Just which is the cause and which the effect is impossible to say in most cases. This connection is especially evident in the movements of fishes other than true swimming, such as burrowing and leaping, for here, as above noted, there are generally limited and peculiar structures to guide one to a correct inference as to their function. The simple metameral contractions are probably the original primitive vertebrate progressional efforts. They are clearly inti- mately associated with the reasons for and the functions of the primeval vertebral column. The following definitions of new terms and the summary bring together in brief form the chief factors entering into the present discussion of fish locomotion and point to the significance of them. Definitions of New Terms. It has been found necessary and desirable to coin some new terms for the present purposes. The more important types of muscular propulsion and certain body zones have been so designated, using in the former the names of characteristic and well known forms as the word stems, and in the latter definitive terms. It should be noted that the terms ending in “form” refer to types of movement and not to body forms and are therefore not parallel to 1926] Breder: Locomotion of Fishes 283 others such as disciform, fusiform and more recently sagittiform (Schlesinger 1909) which are used to describe the body shapes indicated by the first part of the word. In the present series that part refers to the name of a family exemplifying the characteristic. These may be defined as follows. Movements of the Body. Anguilliform Movement.20 That form of locomotion exemplified by the Anguillidae {Anguilla). It consists of a series of sine waves (as in the waving of a flag) which are passed caudad by successive contractions and relaxations of the metameral body muscles. More than one-half a sine curve is formed at least. The pivot is at the base of the skull. See pages 167-169, and Table I. Carangiform. That form of locomotion exemplified by the typical Carangidae ( Caranx ). It consists of not more than one-half of a sine wave, which is passed caudad by successive contractions and relaxations of the metameral body muscles. The pivot at the beginning of a stroke is at the base of the skull, but moves caudad terminating with the end of the stroke in the peduncle. See pages 175-177, and Table I. Ostraciiform. That form of locomotion exemplified by the Ostraciidae (Lactophrys) . It consists of hardly any curve forming at all, the few peduncular myomeres acting almost as a unit in sweep- ing the flexible tail from side to side. The pivot is in the caudal peduncle. See pages 169-175, and Table I. Movements of the Longitudinal Structures. Amiiform. That form of localized anguilliform undulation found in the dorsal fin of the Amiidae ( Amiatus ). See pages 186- 190. Gymnotiform. That form of localized anguilliform undulation found in the anal fin of the Gymnotidae {Gymnotus). See page 187. Balistiform. That form of localized anguilliform and inter- mediate21 undulation found in the dorsal and anal of the Balistidae ( Balistes ). The angular divergence of the two fins from the hori- zontal is complementary, reducing the reactance to a horizontal resultant. See page 187. 20 This movement should not be confused with that of serpents (serpentine) moving over solid ground as there are numerous differences which, however, need not be gone into here. 21 “Carangiform” is not used here for reasons explained on pages 189-190. 284 Zoologica: N. Y. Zoological Society [IV ; 5 Rajiform. That form of localized anguilliform or intermediate undulation found in the expanded lateral pectorals of the Rajidse (Raja). See page 187. Tetraodontiform. That form of localized ostraciiform undula- tion found in the dorsal and anal of the Tetraodontidae ( Lago - cephalus). See page 187. Movements of the Pectorals. Labriform. That form of localized ostraciiform motion found in the pectorals of the Labridae (Tautoga). See page 190. Diodontiform. That form of localized anguilliform undulation found in the pectorals of the Diodontidae (Diodon). See page 190. Body Sections. Orthokinetic Part.22 That portion of a fish’s body which moves forward in a rectilinear path through the center of which passes the axis of progression. See page 181. Cephalic Amphikinetic Part.22 That anterior portion of a fish’s body which does not move forward in a rectilinear path, but oscillates to either side of the orthokinetic part. See page 181. Caudal Amphikinetic Part.22 * That posterior portion of a fish’s body which does not move forward in a rectilinear path, but oscillates to either side of the orthokinetic part. See page 181. Summary. (1) The locomotion of fishes has received the attention of students for a long time but there are still many unexplained factors. (2) The methods basically used by fishes in effecting locomo- tion are extremely ancient in a geological sense. (3) The aquatic habitat is of great potency in producing a direct response to streamline requirements, both as regards form and propelling mechanism. (4) The locomotion of fishes is distinct from that of the tetrapods and shows a certain similarity throughout its various modifications in that the emphasis is placed on the lateral muscles instead of on the paired limbs. (5) The primary locomotor apparatus of fishes centered in the lateral muscles to induce propulsive contortions in the body itself, has clearly conditioned the origin of the vertebral column. 22 The etymology of this term has been suggested by Dr. W. K. Gregory. 1926] Breder: Locomotion of Fishes 285 (6) The length and degree of flexibility of fishes using such methods determines the form which these movements attain. (7) A series of graduations is obtained from the waving move- ments of the eel (anguilliform) through the typical movements of generalized fishes (carangiform) to the wig-wag of rigidly incased fishes (ostraciiform) . Although differing superficially, on account of body form, these movements are fundamentally similar. (8) The thrust effects of the muscular contortions upon the water are direct and not only significant on account of the terminal position of the caudal fin. (9) . Movements similar to those of the body may be localized in various fins and range through a somewhat similar gamut. When localized in various longitudinal structures, the effect of the thrusts is direct, as in the body flexures, (amiiform, gymnotiform, balistiform, tetraodontiform, rajiform), but when in the caudal or pectorals and rarely the pel vies, forward motion is produced by various modifications of the elements entirely dependent, upon position and mechanical circumstances (labriform, diodontiform). (10) All the movements of fishes when swimming (except exhalation) are fundamentally of an undulatory muscular nature even though obscured by various specializations, and are induced by the serial action of metameral muscles. (11) A certain point in a fish’s body moves straight forward, about which a certain core of material does likewise (orthokinetic part). Anterior and posterior to this are two oscillating parts (cephalic amphikinetic and caudal amphikinetic parts). (12) The water exhaled through the gill clefts has an effect useful in driving fishes forward, which varies greatly with different forms, being of considerable importance to some and of little or none to others. It may operate mainly by simple reaction (if the gill clefts are circular) or may partake of the induced streamline effect of Paxton (if the gill clefts are long narrow slits). (13) In the earliest forms in which either single or multiple gill clefts are traceable, they are in the normal position of modern fishes, a fact which indicates the great antiquity of jet propulsion. (14) Maneuvering in fishes consists mainly of differential application of locomotor efforts that would otherwise produce for- ward motion, and the presenting of passive surfaces or planes when in motion. There are few external structures in fishes solely de- voted to maneuvering. 286 Zoologica: N. Y. Zoological Society [IV ; 5 (15) Turning is accomplished by a variety of methods includ- ing body movements, the use of fins and exhalations. (16) Rising and falling is controlled both by fin movements and the swim-bladder, if present, which is primarily a hydrostatic organ. (17) Stops are generally made by a braking action of the pectorals if the organization allows of such. (18) Fishes which reverse their primary locomotor apparatus in coming to a stop, generally also make a practice of travelling backward, for short distances at least. Fishes that show a carangi- form or ostraciiform type of body movement are not able to do this, the pectorals playing a relatively large part. (19) The exhalant respiratory streams of freely suspended fishes make it necessary for them to neutralize these thrusts by various fin actions in order to maintain a stationary position. (20) A current may either hinder or help locomotion or a given maneuvering act. (21) While the propelling and steering mechanism of fishes is intimately connected, there is a general tendency to place the burden of the first on the body muscles and that of the latter on the fins. (22) Locomotor efforts other than swimming, such as flying, creeping, et cetera, are specialized developments and not the common property of any great number of fishes, and are often accompanied by elaborate apparatus developed for that end. (28) Leaping may be considered a swimming up through the surface of the water at high speed and flying a continuation of it chiefly by planing, whereas burrowing may be considered at the start as a swimming into the soil after which other muscular con- tractions continue the progress underground. (24) The entire external, character of fishes , from the most general aspect down to minute details, reflects their locomotor qualifications, and indicates the power of their environment in moulding them. (25) The habits of fishes both are modified by and modify the locomotor qualifications of any given species, these commonly change in the life of an individual, as different habits are acquired during its development. (26) A great amount of homodynamic adaptation is found throughout fish life in response to certain constant factors found in the aquatic environment, such as the density, pressure, et cetera. 1926] Breder: Locomotion of Fishes 287 (27) It is the combination of locomotor characteristics that determines a fish's natatorial powers and not the intrinsic qualities of each alone. (28) Even in fossil fishes the same fundamental character- istics that are found in recent fishes obtain throughout. (29) The pectoral fins first appear as guiding planes and to offset the depressing action of the epibatic tails. Later they take on definite locomotor movements demanding greater flexibility. (30) The external factors involved in the swimming of fishes in a straight forward horizontal path through open still water may be tabulated thus. (A) Effects of planes of pressure (1) Body flexures (Anguilliform to Ostraciiform) (2) Longitudinal fin flexures (Localized* Anguilliform to Ostraciiform) , (3) Paired fin flexures (Localized Anguilliform to Ostraciiform) (B) Effects of jets (1) Exhalations (Paxton’s factor to Nozzle reaction) This table is designed to include all the efforts of fishes to move directly forward under the conditions mentioned above. BIBLIOGRAPHY Only titles actually referred to in the text are listed here, for it is felt that with the Dean bibliography (1916) available23, others would be superfluous. Du Bois-Reymond (1914) also gives an extensive list of references. Abel, O. 1912. Grundziige der Palaeobiologie der Wirbeltiere. Stuttgart, pp. 708, figs. 470. 1922. Ueber den wiederholten Wechsel der korperformen in Laufe der Stammesgeschichte der Teleostomen. Bijdr. Dierk. Amster- dam. Vol. 22, pp. 73-87. Allen, W. E. 1923. Behavior of the thresher shark. Science, n.s., vol. LVIII, No. 1489, July 3, pp. 31-32. Borelli, G. A. 1680. Philosophia de motu animalium ex principio mechanico-statico. 2 vols. Romae, 1680-82. pp. 168. 23 See especially under the following headings: Adaptations,. Adhesive disk, Air bladder, Dermal skeleton, Temperature and number of fin rays, Flying fishes and their flight, Form of fishes, Hearing and the static sense, Integument or skin, Locomotion, Metamerism, Sleep of fishes, Technique and methods, Teratology. 288 Zoologica: N. Y. Zoological Society [IV; 5 Boulenger, G. A. 1904. Fishes, systematic account of teleostei. (In Cambridge Natural History, Vol. VII, pp. 541-727.) London. Breder, C. M. Jr. 1923. Spring notes on lampreys. Bull. N. Y. Zool. Soc., July, vol. XXVI, No. 4, pp. 86-89. 1924. Respiration as a factor in locomotion of fishes. American Natur- alist, March-April, Vol. LVIII, pp. 145-155. 1925a. Fishes squirting water. Bull. N. Y. Zool. Soc., May, vol. XXVIII, No. 3, pp. 69-72. 1925b. Tailless pearl roach. Ibid., pp. 72-74. Bridge, T. W. 1904. Fishes, exclusive of the systematic account of Teleostei. (In Cambridge Natural History, Vol. VII, pp. 139-537.) London. Brugmans, S. J. 1812. Aanmerkingen over de middelen door welke de visschen zich bewegen in het algemeen en over het vermogen der Uitademing tot dat einde in het bijzonder. Verh. I Kl. Nederl. Instit., pp. 185-217. Clark, Austin H. 1925. Animal flight. The Scientific Monthly, January, 1925, pp. 5-26. Corson, R. H. 1923. Fire Island fish notes. Copeia, October 15, No. 123, p. 108. Dahlgren, U. 1914. Origin of the electric tissues of Gymnarchus niloticus. Carnegie Instit. Washington, Pub. no. 183, pp. 159-194. Daniel, J. F. 1922. The Elasmobranch fishes. Univ. Calif. Dean, Bashford. 1895. Fishes, living and fossil; an outline of their forms and probable relationships. Col. Univ. Biol. Ser., Ill, pp. 1-300. 1906. Chimaeroid fishes and their development. Carnegie Instit., Washington, Pub. no. 32, pp. 1-195. 1912. Changes in the behavior of the eel during transformation. Ann. N. Y. Acad. Sci., December, Vol. XXII, pp. 321-326. 1916. A bibliography of fishes. Amer. Mus. Nat. Hist., N. Y. Vols. I, II, III. Dowd, R. E. 1921. The aeronautics of the flying fish. Aerial Age Weekly, January 10, pp. 464-465, figs. 3. Du Bois-Reymond, R. 1914. Physiologie der Bewegung (In Winterstein, 1914), pp. 166-185. Duges, A. 1905. Note on the functions of the fins of fishes. Science, n.s., Vol. 22, pp. 798-800. Evermann & Kendall. 1905. An interesting species of fish from the high Andes of central Ecuador. Proc. Biol. Soc. Washington, Vol. 18, pp. 91-105. Breder: Locomotion of Fishes 289 1926] Gradenwitz, A. 1923. Oscillatory power. Science and Invention. February, Vol. X, No. 10, whole No. 118, pp. 950-951. Gregory, W. K. 1914. Adaptations illustrating “habitus” and “heritage.” Locomotor adaptations in fishes. Ann. N. Y. Acad. Sci., Vol. 23, pp. 267- 268. Harrington, N. R. 1899. The life habits of Polypterus. Amer. Naturalist, Vol. 33, pp. 721-728. Houssay, F. 1911. The body shape of fishes. Water pressure as the determining cause. Sci. Amer. Suppl., Vol. 74, p. 12. 1912. Sur la stability des poissons en mouvement. C. R. Assoc. Franc. Avanc. Sci., 40. sess., pp. 34-35. 1914. The effect of water pressure upon the form of fishes. A study of evolution of form resulting from conditions of life and habits. Sci. Amer. Suppl., Vol. 78, pp. 376-378. Johnson, R. D. O. 1912. Notes on the habits of a climbing catfish ( Arges marmoratus) from The Republic of Colombia Ann. N. Y. Acad. Sci., Vol. 22 pp. 327-33. Karrer, Enoch. 1924. The terminology of certain physical and biological effects of light. Science, n.s., Vol. LX, No. 1541, July 11, pp. 30-33. Kiaer, J. 1924. The Downtonian fauna of Norway. Videnskapsselskapets Skrifter I, Mat.-Natur. Klasse, No. 6. Mast, S. 0. 1915. The behavior of Fundulus, with especial reference to overland escape from tide-pools and locomotion on land. Jour. Animal Behavior. Vol. V, No. 5, pp. 341-350. Maurey, E. J. 1895. Movement. Translated by Eric Pritchard. The Internat. Sci. Ser. New York, No. 73, pp. 323, figs. 200. Nichols, J. T. 1915. On one or two common structural adaptations in fishes. Copeia, July 27, No. 20, pp. 19-21. 1920. The Phylogeny of Carangin fishes. Ibid., July 31, No. 84, pp. 57-63. OSBURN, R. C. 1906a. The functions of the fins of fishes. Science, n.s., Vol. XXIII, No. 589, pp. 585-587. 1906b. The uses of the fins of fishes. Bull. N. Y. Zool. Soc., No. 25, pp. 347-348. Parsons, H. de B. 1888. The displacements and area-curves of fish. Trans. Amer. Soc. Mechanical Engineers, Vol. IX, 17 meet., pp. 1-17. 290 Zoologica: N. Y. Zoological Society [IV; 5 Pettigrew, J. B. 1874. Animal locomotion; or walking, swimming, and flying, with a dis- sertation on aeronautics. The Internat. Sci. Ser., New York, No. 8, pp. 1-264. Regnard, P. 1893. Dynamometre permettant de mesurer la puissance musculaire de l’appareil caudal du poisson. C. R. Mem. Soc. Biol. Paris, 9. ser., Vol. 5, pp. 1-80. Ridewood, W, G. 1913. Notes on the South American fresh water flying fish, Gastropele- cus and the common flying fish Exocoetus. Ann. Mag. Nat. Hist., Vol. 8, ser. 12, pp. 544-548. Ryder, John A. 1882. A contribution to the development and morphology of the lopho- branchiates (Hippocampus antiquorium ) the sea horse. Bull. U. S. Fish Comm., Vol. I, (1881), pp. 191-199. 1892. On the mechanical genesis of the scales of fishes. Ann. and Mag. Nat. Hist. 6. ser. 11, pp. 243-248. SCHLESINGER, G. 1909. Der sagittiforme Anpossungstypus nektonischer Fishe. Ver. Zool.-Bot. Ges. Wien. Vol. 59, pp. 140-156. 1910. Die Locomotion der Notopteriden. Zool. Jahrb. ( Abth . Syst .) Vol. 29, pp. 681-688. 1911a. Die Locomotion der taenioformen Fische. Ibid., Vol. 31, pp. 469-490. 1911b. Schwimmen und Schwimmtypen flinker Meerefischen. Kosmos. Jahrg., pp. 284-287. SCHMALHAUSEN, I. 1916. On the Functions of the Fins of Fish. Rev. Zool. Russe, Moscow. Pp. 185-214. Resume in English. Spencer, H. 1892. A system of synthetic philosophy. Vols. I, 2, 3. London. Stringham, E. 1924. The maximum speed of fresh water fishes. Amer. Nat. Vol. Vol. LVIII, March-April, No. 655, pp. 156-161. Taylor, H. F. 1922. Deductions concerning the air bladder and the specific gravity of fishes. Bull. U. S. Bur. Fisheries, April 24, Vol. XXXVII, doc. No. 921, pp. 121-126. Welsh, W. W. & Breder, C. M., Jr. 1922. A contribution to the life history of the puffer, Spheroides macu- latus (Schneider). Zoologica, Vol. II, No. 12, pp. 261-276. Willey, A. 1902. Contribution to the natural history of the pearly nautilus. I. Personal narrative (In his Zoological results based on material from New Britain, New Guinea, Loyalty Islands and else- where. Collected 1895-97. Cambridge, pt. vi.). 1926] Breder: Locomotion of Fishes 291 WlNTERSTEIN, H. 1914. Handbuch der vergleichenden Physiologie, vol. 3, Halfte 1, Theil 1, pp. 1-248. Jena. (Schwimmen der Fische, by Du Bois- Reymond, R.) Woodward, A. S. 1893. Note on the evolution of the scales of fishes. Nat. Sci., Vol. Ill, pp. 448-450. APPENDIX Descriptions of Mechanical Devices The following descriptions and the diagrams accompanying them are here- with appended in order to make more clear the methods used in the phases of the work requiring aids of this nature. They are sufficiently detailed to enable others to construct identical devices if so inclined and allow of a complete check on all such data. Models. Two models were constructed in the analysis of the general body movement of fishes. They represent either extreme as discussed under Body Movements. Ostraciiform Model. The construction of the ostraciiform model (Fig. 40) is shown (Fig 80) and needs scant elaboration in writing. The hull was carved from soft wood and extended upward by a sheathing of sheet metal. The motive power was supplied by a small spring motor, such as is used in phono- graphs using disc records. No modifications were made on it, the spindle in- tended to receive the record plate being used as a driving shaft. To this was attached a stiff crank arm supporting a pin bearing a roller. Details of this are shown in the central part of Fig. 80. This grooved roller rested between two 292 Zoologica: N. Y. Zoological Society [IV; 5 PL AM VIEW FROM BELOV wiring; diagram Fig. 81. The anguilliform model. Working drawing, showing details of construction. parallel rods attached to a travelling carriage to which it imparted a reciprocal motion. A chain attached to either end of the cross arm on the tail piece and passing around a pulley in the bow was secured to one side of the travelling carriage. This imparted a wig-wag motion to the tail piece. The speed of this could be adjusted by regulating the governor on the motor and its amplitude of oscillation could be modified coarsely by slipping the hooks on the ends of the chain in various holes in the tail piece cross arm, as seen in the “Plan” of Fig. 80. Fine adjustment could be obtained by moving the driving pin on the motor shaft in the slot of the crank arm as indicated in the central details. A hole was bored in the hull to accommodate the extended winding crank. A cork pre- vented any water from being shipped here. Other details of structure are evi- dent on the drawing. Anguilliform Model. The construction of the anguilliform model (Fig. 49) is illustrated in the working drawing (Fig. 81). The construction of this was somewhat more difficult and as it was desirable to reverse the movements a small electric motor was used for supplying the power. A flat board, the deck, was sheathed above and below with sheet metal, as indicated. A chain of clock work gears was attached to the motor for reducing the speed, the last shaft which supported a sprocket wheel connected with a chain to a similar one at the end of a crank shaft running nearly the full length of the craft. This chain ran 1926] Breder: Locomotion of Fishes 293 through a hole in the deck around which was built a water-tight housing extend- ing above the water line to prevent water reaching the deck and motor. Each crank of the crank shaft was advanced 36° beyond the last, so that every tenth one was in an identical position. To each of these was attached a small con- necting rod, linking them with long stiff rods pivoted on another and parallel shaft. Rotation of the crank shaft imparted a wig-wag motion to each of these rods as indicated in the central details of Fig. 81. On account of the placing of the cranks each such rod followed its leader in wig-wagging. Considering them as a whole, a wave was thus caused to pass down them. A piece of thin cloth, doubled, with spacing stitched for the reception of each rod completed the analogy to a longitudinal fin. Power applied by the motor would cause move- ment of the craft in a direction opposite to the travel of the wave along the series of webbed rods, this being closely analogous to the anal of a gymnotid. Long, flexible insulated wires allowed of freedom of motion. A complete reversal of direction could be obtained by changing the direction of rotation of the motor. This was accomplished by operating a double-poled reversing switch held in the hand and wired as shown in the wiring diagram. Other structural details are evident on the drawing. Apparatus Other Than Models. Three pieces of apparatus found to be generally useful were constructed as follows: Their uses were various and are either specifically mentioned in the text or are inferred at various places throughout the body of the paper. Balance. A balance for determining the position of the center of gravity of fishes was constructed as follows: To a rigid base a vertical metal plate with a Fig. 82. Dynamometer for determining the tractive pull of fishes. 294 Zoologica: N. Y. Zoological Society [IV; 5 knife edge was attached. On this was balanced a glass plate to which cross ruled tissue had been glued so that one of the sets of ruling was parallel to the knife edge. Then a fish was laid on the plate with its long axis parallel to the rulings perpendicular to the knife edge by trial until it balanced. Two pins were used as markers fixing the axis through which it balanced. Then the fish was swung through 90° and the process repeated. The intersection of the two lines was over the center of gravity which was half way through the fish (sym- metrical from right to left). In this way the data of Fig. 59 was obtained. Dynamometer. Owing to the fright incident to handling, and the rapid fatigue of fishes the desired data was not obtained by the following described device, but much corollary information was incidentally uncovered. For this reason its description is given. Fig. 82 illustrates it and obviates detailed ex- planation. A line attached to the long arm and passing over the two pulleys was attached by a bridle to the specimen under experiment. The spring balance then indicated the amount of pull on the line when properly calibrated. This balance could be moved lengthwise of the arm for purposes of leverage depending on the strength of the fish in hand, each position requiring new calibration. However most examples failed to perform in a satisfactory manner, although one Centropristis momentarily registered considerably more than its own weight out of water. Intermittent Light Device. This device aided in the study of fin move- ments in small fishes. Its external appearance is shown in Fig. 83. A powerful electric lamp burned continuously within the housing and the circular shutter was rotated by a toy electric motor above it. Control was obtained as follows. The length of the periods of light could be varied in relation to the periods of darkness by the adjustable sector in the heavy cardboard shutter attached to the motor shaft by small nuts. The speed of the flashes could be varied to any desired time by a rheostat in the motor circuit. It could be driven at such a speed that the intermittent effect could hardly be detected. As sixteen images per second are about as many as the normal human retina can interpret satis- factorily due to the lag of the sensory nerves, it is clear that a speed of much over 1,000 r.p.m. would be of little value. The device was first tested by rotating a spoked wheel in an otherwise darkened room. By regulating the aperture, and speed it could be made to apparently stand still or rotate in either direction at speeds other than its own. Similar to the effect sometimes seen in motion pictures of wagon wheels. Thus fish fins could often be reduced to apparently slow motion although others, such as the dorsal of Hippocampus probably on account of its high speed could not be studied with satisfaction in this way. It seems possible that an accurate measure of its speed might be gotten in some such way by further study. Suggested Lines of Research. It is particularly patent to the author that the preceding pages are hardly more than a start, and are open to much expansion. Many fascinating di- gressions from the main theme on subjects which are only touched on here have tempted him consderably in the course of the work and various problems have been noted down. Some of those which seem to promise to be the most prof* 1926] Breder: Locomotion of Fishes 295 Fig. 83. Intermittent light device for determining the speeds of undulating fins. itable for future investigation are suggested in the following list. Possibly none of them will be undertaken by the author personally and it is highly de- sirable that they be investigated by others, if only for the sake of freshness of viewpoint. 296 Zoologica: N. Y. Zoological Society [IV; 5 (1) The maximum and normal speeds of various fishes should be accurately- determined. (2) The exact relationship between the speed of the body movements and the speed of progression should be determined. (3) The tractive strength of fishes should be determined after a suitable technique is devised. (Improve on the dynamometer described in this paper.) (4) An accurate measure of the jet effects on high speed swimming should be obtained, the only condition under which the effect of Paxton could be proper- ly determined. (5) An exact measure of the relation of the shape and size of the caudal to its use should be reduced to an index if possible. Just how is the fork of a tail connected with speed? (6) A study regarding the hydrostatics of fishes with and without a swim- bladder should be carefully made and the position of the center of buoyancy, the metacenter and their effects on the stability of various forms should be deter- mined. Fishes that habitually turn over, should be studied particularly. (7) The relationship of the size, shape and placement of the fins and other external parts in connection with the body form should be measured in detail and their various coefficients found, if possible. (8) The internal structures, especially the formation of various bones an- choring the natatorial muscles, and the size and functioning of the swim-bladder should be studied in relation to the swimming of the subjects. (9) A careful cataloging of the various fishes should be made according to locomotor characteristics such as is suggested by Tables III and VII, but in much greater detail and study. (10) The flight of flying fishes is still open to discussion and should have first hand study. (11) The change of locomotor characteristics in the ontogeny of any group should be useful. Which is “habitus” and which “heritage”? (12) The possibility of using the finer details of locomotor efforts as in- dicative of relationship should be considered. (13) In connection with the above, the possibility of different groups at- taining the same locomotor ends, and having sufficiently similar locomotor characteristics to conceal their different origins, should be considered. In other words, are some of the homodynamic adaptations so close as to be in- distinguishable? (14) The effects of the exhaled jets of various forms in comparison with other locomotor efforts at various speeds should be studied. (15) The swimming of any fossil group, as revealed by remains, should be studied in detail. This should be of particular value in making restorations and interpreting various data as suggested under “Locomotion of fossil fishes.” (16) The details of the locomotor habits and their causes in any group of recent fishes as well as their ancestors should be studied. (17) The number of myomeres in reference to the accompanying number of vertebrae, scales, etc. should be studied. This might give an index of the flexibility of the body. 1926] Breder: Locomotion of Fishes 297 (18) The possibility of definite correlation between the position of the gill slits and the greatest mid-section or other proportion in reference to the ex- halations should be considered and reduced to formulae. (19) The various specializations of fishes concerned with locomotor efforts other than swimming, such as creeping, flying, etc. should be compared between different groups in all respects. (20) The manner in which the habits of fishes relate to their movements should be studied in detail. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V, NUMBER 1 Department of Tropical Research, Contribution Number 151 WILLIAMS GALAPAGOS EXPEDITION By William Beebe Director, Department of Tropical Research and Honorary Curator of Birds New York Zoological Society PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 31, 1923 N?uj fork Xonlogtral Swirly General Office: 111 Broadway, New York City GDffkmi President , Henry Fairfield Osborn; Vice-Presidents , Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer, Cornelius R. Agnew lfoarb of managers QHaas of 1924 Madison Grant, William White Niles, Frank K. Sturgis, Ogden Mills, Lewis Rutherfurd Morris, Archer M. Huntington, George D. Pratt, T. Coleman duPont, Henry D. Whiton, Edward Hatch, Jr., Cornelius R. Agnew, Harrison Williams (Class of 1 925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge C. Ledyard Blair, Anthony R. Kuser, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler Class of I92fi Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor. £>rirntiftr £>taff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, D. V. S., Assistant to Director; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of the Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector ; George A. MacCallum, Pathologist; El win R. Sanborn, Photographer and Editor. C&itnrial (Eummitter Henry Fairfield Osborn, Chairman; William T. Hornaday, Charles H. Townsend. Corrected to December, 1923 ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V, NUMBER 1 Department of Tropical Research, Contribution Number 151 WILLIAMS GALAPAGOS EXPEDITION By William Beebe Director, Department of Tropical Research and Honorary Curator of Birds New York Zoological Society PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 31, 1923 Plate A. SKETCH MAP OF GALAPAGOS ISLANDS Route of the Noma, and details and location of the Archipelago. 2 First for m on press December 3.1 102, Zoologica Vol. V, No. 1 V)t3 WILLIAMS GALAPAGOS EXPEDITION By William Beebe Director, Department of Tropical Research, and Honorary Curator of Birds .New York Zoological Society Photographs by the Author and John Tee-Van 1. Resume This expedition, to one of the least visited corners of the earth, was conceived and achieved in record time, every hope was con- summated, every expectation realized. First and last, the credit belongs to Harrison Williams, Esq., who initiated and financed the whole trip, and then to the twelve members of my party, who made possible all that we accomplished during the limited time at our disposal. We left New York on the steam yacht Noma on March 1, and returned on May 16. This was just in time to rush the collections of live mammals, birds and reptiles to the Zoological Park, and to frame and hang for exhibition the one hundred and thirty oil paint- ings and water colors made during the trip, — in readiness for the Annual Garden Party of the Zoological Society on May 17. During the trip we steamed a total distance of nine thousand miles, and crossed the equator eight times. Twenty-one memorable days were spent on the Galapagos Islands, and we touched besides at Charleston, Key West, Havana, Colon and Panama. To the living collections of the Zoological Park were added the following, most of which were new to the collections, some being exhibited for the first time anywhere in the world: Mammals. — 5 monkeys, 3 opossums. Birds. — 3 penguins, 2 flightless cormorants, 3 gulls, 3 doves, 1 hawk, 10 parrakeets, 2 jays, 3 mockingbirds. Reptiles. — 42 lizards. For the American Museum there was collected material for two lizard groups, including vegetation, rocks, shells, sand and many photographs, together with a giant tortoise, eighteen lizards and a family of sea-lions. 3 4 Photograph by William Beebe, 1923] Beebe: Williams Galapagos Expedition 5 \ \ f vl ' FIG. 2. FTSHING FROM A BOWSPRIT Strange fish, worms, jellies and even insects can be caught from this swinging seat, at full speed and in rough weather. For study by the department of Tropical Research of the Zoological Society: 90 water color plates by Miss Cooper. 40 oil paintings by Harry Hoffman. 46 pen and ink drawings by Mr. Broking. 400 photographs and 11,000 feet of moving picture film by Mr. Tee-Van. 160 bird skins. Many nests and eggs. 150 reptiles. 200 fish. 3,000 insects. 40 jars of specimens. 60 vials and jars of plankton. 200 microscopic slides of plankton. 100 specimens of plants. 300 pages of narrative, records, notes and catalogues by Miss Rose. 6 FIG. 3. DARWIN BAY, TOWER ISLAND A sheltered, mile- wide bay which seems never to have been described or mapped. Every niche of cliff, every bush and shrub holds nests of gulls, teros, boobies, doves or frigate-birds. 1923] Beebe: Williams Galapagos Expedition 7 This material is remarkable both for its rarity, excellent preser- vation and for the fact that it was almost all collected within a period of three weeks. The various groups of organisms will be studied by members of the expedition or sent to specialists, and the results published in the following numbers of Zoologica, while the more general matter has appeared in a volume by William Beebe, called “Galapagos; World’s End,” published under the auspices of the Zoological Society, by George P. Putnam’s Sons. II. Brief Narrative The Noma, with all the members of the Williams Galapagos Expedition, steamed from her berth in Brooklyn at noon on March 1, but swinging the compass and engine adjustment kept us in the lower bay for thirty hours. This gave opportunity for unpacking and storing our vast quantity of paraphernalia, and in fitting up a laboratory, a fortunate interlude as it proved, for the passage down the coast was rough and stormy. Scientific work on a yacht under way was a new experience to me, and we ran the gamut from comfort to absolute cessation of work. Until we learned to fasten everything down, a sudden terrific heave would sweep the laboratory tables quite clean, and on unusually rough days we would continue our work seated on the floor, as chairs were useless. This was the exception, however, and in the usual calm weather, the twenty-five hundred horse-power, twin-screw engines gave forth not a tremor or vibration so that even high power microscopic research could be carried on. By the time we were off the Florida coast the sea permitted me to occupy my usual perch in a boatswain ’s seat, over the bow, close to the water, where, with a long-handled net, I secured sufficient fish and sea- weed fauna for days of study. We put into Key West to pick up Dr. James Mitchell and to obtain additional supplies of coal and water, and then crossed to Havana for a supply of 95 per cent, alcohol for the preservation of our specimens. While in port we dredged sand bars, and caught sharks and various tropical fish from the rail. From the Windward Passage between Cuba and Haiti, to Colon, we were in the trough of a heavy sea, and rolled steadily, occasionally as much as 34 degrees. At Colon we were overhauled and coaled, giving time for collecting trips to the jungle and coast beyond Fort Sherman, for horseback rides to the Chagres River, 8 Zoologica: N. Y. Zoological Society [V; 1 FIG. 4. LAND LIZARDS OF THE GALAPAGOS On the sandy upland veldt of Seymour Island we caught these great iguanas, red and green, yellow and ivory white, — as eager to bite as their fellows of the surf were innocuous. tarpon fishing at the Gatun spillway, and trips back and forth across the Isthmus. In Colon we met with the greatest disappoint- ment of the voyage as Mr. Williams was compelled, for business reasons, to return to New York. It was with the deepest regret that we saw him go, for his enthusiasm in the expedition had been great, and now the real excitement of exploration was just ahead. For a supply of fresh water beyond that which the Noma carried, we depended upon the islands, as a supply was marked on the chart in several places, and the pilot book mentioned even a pipeline on the dock at Chatham. We passed through the Canal without special incident, arriving at the Panama end in time to see the final searchlight display of the combined Atlantic and Pacific fleets. On the night of March 24 we steamed into the Pacific, which was as smooth as a lake during the four days it took us to reach the Archipelago. Indeed during our time there and on our voyages to and from Panama this ocean lived up to its name, and we experienced only perfect weather and summer calm, a welcome change from our Atlantic memories. 1923] Beebe: Williams Galapagos Expedition 9 FIG. 5. A FEARLESS SEA-LION Like the birds in the background, this big male seal had probably never seen a human being, and could conceive no harm as coming from such a strange creature. At dawn on March 28 we sighted the islands, and steaming slowly among their misty shapes, recognized Indefatigable, James, Seymour, Daphne, Jervis and Duncan, and dropped anchor about 10:00 A. M. in Conway Bay on the north-west side of Indefatigable. This anchorage, chosen more or less at hazard because of our in- complete information concerning the islands, proved to be a fortunate choice. Sheltered on the west by Eden, an isolated volcanic peak of an island, a sandy beach at the back of a natural lava breakwater provided an easy and safe landing for our small boats. Here we pitched two tents for temporary laboratories, though most of our work in arranging and studying specimens continued to be done on board ship. During our entire stay at the islands we lived on the Noma, thus eliminating the extra work of transporting supplies; also we had the benefit of the evening breezes and escaped the mosquitos which on seme of the islands appeared at dusk in innumer- able swarms. 10 Indefatigable in the distance. 1923] Beebe: Williams Galapagos Expedition 11 The shore life at this landing, which we named Harrison Cove, was plentiful and most interesting. The instantaneously arresting feature was the astounding tameness of all the creatures. Having never seen human beings they had little fear, the birds and sea-lions being particularly indifferent to us. Perhaps indifference is hardly the word, since in many cases they showed great curiosity about us. Mockingbirds would follow us along, hopping from branch to branch within arm’s reach; little flycatchers would perch a foot from our faces, in close inspection of our mystifying presences. It was found almost impossible to alarm some of the big pelicans or gulls and even among the crabs some individuals would stand as quietly as the lava while we touched or pushed them about. During our first hour ashore a wild duck flew down and alighted at our very feet and a short-eared owl perched on my helmet as I walked through the low scrubby undergrowth. Our first day at Harrison Cove was rich in interest and no one of our succeeding days fell below its high standard. The fact that a large percentage of the fauna and flora of the Galapagos is peculiar to this Archipelago, and the presence of such rare forms as Ambly- rhynchus, the only marine lizard in the world, and Conolophus, an extraordinary land lizard whose numbers are rapidly decreasing, makes the study of these islands of particular interest. At Conway Bay we had a wide field from which to choose. Eden, in spite of its small size, yielded a great quantity and variety of specimens. It was here in one small cove that I obtained our collection of living Amblyrhynchus, and a host of interesting facts concerning their life history. Insects on the Galapagos are very limited as to numbers as well as species, but some unusual ones were collected here, while tide-pools among the lava shore were inexhaust- ible mines of beauty and value. Guy Fawkes Rocks to the north- east of our anchorage were favorite haunts of sea-lions and many memorable hours were spent under the over-hanging cliffs in photo- graphing these animals and in delightful tests of their tameness. Later, specimens were secured here. We had not found fresh water at Conway Bay and our supply was rapidly diminishing. Even now we were on rations, shaving in Whiterock or Poland Water and bathing only in salt water. Accord- ing to the chart, there was fresh water at James Bay on James Island, not far to the north of Indefatigable, and four of the party went off in one of the larger motor-boats to investigate. They 12 The yacht Noma in the haunts of the Flightless Cormorant. The bird sits on its nest without fear, seeing for the first time human beings. This individual is at present living in the New Tork Zoological Park, 1923] Beebe: Williams Galapagos Expedition 13 returned late the same night, reporting that they had found no water but giving such glowing accounts of the island that we determined to stop there if only for a short time on our way to some other spot in search of water. At noon on April 4 we anchored at James Island where eighty- eight years ago Charles Darwin had spent a week. In James Bay we hurried ashore in small boats. The landing was a difficult one in spite of a long sandy beach, for the surf was very heavy and a bad undertow combined with swirling cross currents made it a risky spot. This island differed from Indefatigable in that trees of considerable size grew close to the shore, which made it possible with slight effort to reach the forested slopes of the crater. On Indefatig- able we had not been able, in the limited time at our disposal, to penetrate the miles of country, covered with jagged broken lava, cactus and thorny scrub, which separated the semi-arid coast from the forested high country of the interior. On James the going was also made comparatively easy by the well-defined donkey trails. Each of the larger islands that we visited seemed to have some sort of animal, once domestic, which had bred and multiplied and reverted to a wild state. Indefatigable, for instance, has its wild dogs; South Seymour its flocks of wild goats; Albemarle its cattle, and James, judging from the number and well-worn condition of the trails, is the home of large numbers of wild donkeys. Here we also found the skeletons and tracks of wild pig and one of our number shot a large sow. Whether these animals were left here by buccaneers or whalers as a future food supply, or whether they are the only survivors of ship-wrecks of long ago, no one knows. It is an interesting fact that these imported forms, all of which we are accustomed to consider as thoroughly tame, should here, after a few generations of non-domestication, be the only really wild animals. They have reverted to a completely feral state, that is to say, of fear of man, while such creatures as birds or reptiles from which we expect no confidence, are, in these islands, tamer than barnyard fowls. On Indefatigable one glimpse of wild dogs was vouchsafed to me, wolfish looking animals who, on sight, snarled and slunk away. During our few hours at James Bay we saw only two donkeys, one of which was pure white, though the hills often reverberated to their hearty braying, and the one wild pig was secured only after a stalking as cautious as though a deer had been the object of the 14 Zoologica: N. Y. Zoological Society [V; 1 chase. The contrast is great between this sort of pursuit and our experiences in lifting up frigate birds and cormorants from their nests, and patting sea-lions on the head. We found no fresh water on James, only brackish pools close to the sea, where ducks and herons were plentiful. Here, too, we saw flamingos passing overhead, but there were few seabirds, as the closely wooded shores and absence of islets offered no attraction to them. The water question was sufficiently pressing to prevent us from spending more than one day here, and it was decided to steam for Tagus Cove on Albemarle, which was marked on the chart as a good anchorage, with two places on the shore where fresh water could be obtained. In returning to the Noma that evening, three of our party had a narrow escape from what might have been serious injury. In launching the small motor boat, it was overturned by a big breaker and they had a bad few minutes in the surf. Luckily they escaped with nothing worse than a few cuts and bruises. The boat was smashed and rifles and personal belongings were lost. Later in the evening when their predicament was discovered they were brought off in a lifeboat. Early next morning we left for Tagus Cove, steaming around the north end of Albemarle and passing between it and Narborough. On these two islands we saw what seemed like the most recent evidences of volcanic activity, great black swathes of lava slashing across the green of trees and undergrowth. It became noticeably colder in passing to the west of Albemarle on the open ocean side, so much so that sweaters were comfortable for an hour or two. At first we were doubtful of the identity of Tagus Cove, it seemed so small and unlike in shape to that anchorage shown on the chart. But once inside, a more perfect shelter would be hard to conceive. Long and narrow, between straight towering cliffs, with deep water up to within a few feet of land, it was a satisfactory and a wonderfully picturesque anchorage. The landing facilities left much to be de- sired , but that was of small moment compared to our disappointment when the chart was once more proved to be over-optimistic on the subject of water. Not a drop of the precious fluid was to be found, although this was the height of the rainy season, and our only hope now was to go to Chatham, in search of that pipeline of which the pilot book spoke so glibly. 1923] Beebe: Williams Galapagos Expedition 15 We calculated that with our shortage of supplies it would not be advisable to stay long at Tagus Cove, but our few hours there yielded a rich harvest. Some of the party explored the slopes adjacent to the Cove, finding quantities of nests and eggs of the black finches (Geospiza), and other indigenous birds, besides insects, lizards and botanical specimens. Others climbed the steep cliffs around the Cove, carrying with them, by enormous exertion, motion- picture and other cameras, plates and equipment up the almost perpendicular slopes. In this Cove we secured live penguins and flightless cormorants, as well as the nests and eggs of the latter. Boobies, pelicans and terns were abundant and nesting. We left Tagus and steamed toward Chatham, crossing the equator four times in twenty-six hours. Early the next morning we anchored at Wreck Bay which boasts the only lighthouse in the Archipelago, visible for four miles, which is not bad for a gasoline light on a long pole. Nothing else is to be seen of human occupancy in this Bay except a square white shack where the lighthouse keeper lives, and a very shaky pier. The pipeline of the pilot book did not exist. The lighthouse keeper, an Ecuadorian who said he was also Captain of the Port, came aboard with an old Englishman, and we were told that the only way to obtain fresh water was to have it brought in casks on the back of oxen from a distance of five miles up in the mountains. As we needed forty tons of water, this was an impossible way of obtaining it, and the prospect was very gloomy. The old Englishman who told us he was “ Johnson of London” and who had lived so long in Wreck Bay that he had almost forgotten his native tongue, volunteered to pilot us around the island to Fresh Water Bay where he was sure we could get a sufficient supply. So, having stopped hardly long enough to anchor, we got under way again, and cruised around to the Bay with the promising name. Here we found two cascades of fresh water, one of good size, which plunged over high cliffs and poured into the sea. Against the foot of the cliffs surged a tremendous surf, which kept all small boats a hundred feet off shore. The Bay was such only by courtesy, for there was almost no incurve to the forbidding coast line and it was on the weather side of the island . There was no bottom a quarter mile off shore, and the Captain dared approach no closer. So we watched the tantalizing spectacle of quantities of fresh water running to waste in a spot which for us was utterly inaccessible. 16 The site of a submerged crater. Out of the crevices of the tortured, dead cinders spring grotesque cacti, on which perch tamest of mockingbirds, singing their hearts out. 1923] Beebe: Williams Galapagos Expedition 17 However, with three others I made an attempt upon a bit of low pebbly beach with a mass of green back of it. I leaped over- board and let the roller wash me up on the piled pebbles, and there, immediately behind, was a broad stream of pure water rushing down into the sea. We rigged up the long rubber deck hose, one end being in a lifeboat a hundred feet from shore, and the other on top of the pebbly beach. This end I held as high as possible in the air, while the rest poured buckets of water into a funnel. In an hour we had four tons of sweet water in the life-boat, and we towed this off to the yacht. There we found that, unable to anchor, we were consuming coal at a rate which would still further curtail our stay. So we had to give up our hard-earned plan of filling the tanks on the next morning. After landing “ Johnson of London” who had given us so little in return for the enormous quantities of food and cocktails which he had consumed, we steamed back to Conway Bay for a last clean-up of the specimens which we needed. Then on to Panama which we reached on the last ton of coal in the bunkers and the last gallon of fresh water. At Panama we added to our party Prof. William Morton Wheeler of the Bussey Institution, Harvard, who had been with me at Kartabo in British Guiana, and whose philosophical grasp of evolution and life on the earth makes his presence on any expedition a tremen- dous asset and pleasure. We coaled again, covering the lower decks with great sacks and besides laid in quantities of bottled water. When we again sighted the Galapagos it was decided to try another anchorage near Indefatigable, farther to the east, in the shelter formed by the two Seymour Islands. During the first day ashore two attempts were made to penetrate to the crater, but both were defeated by the terrific going. We realized that two or three camps must be established to accomplish this feat, and in the limited time which we had., it seemed infinitely wiser to concentrate on the vast mass of material ready to hand along the shore, rather than give up precious days to the mere performing of a stunt. Here we again found the small islands in the vicinity to be far more interesting than the large one. South Seymour, to the east, was geographically quite unlike any other island, as back from the shore it consisted of open veldt-like country. This was covered with grass and dotted sparsely with cactus and fair-sized trees, where moving flocks of spiral-horned goats took the place of ante- 18 FIG. 9. SEYMOUR ISLAND 1923] Beebe: Williams Galapagos Expedition 19 lope in corresponding places in Africa. This, too, was the only place where we found Conolophus, the giant land lizards. Daphne Major, five miles to the north, was visited twice. It is a perfect island crater, and after landing on its most inhospitable cliffs, we climbed its precipitous sides covered with loose, easily- sliding shale and looked down into the deep crater. The floor covered with white sand was dotted everywhere with hundreds of nesting boobies, all of the blue-footed species. We went down and walked about among them, collecting a chick, or an egg or an adult here and there, and taking photographs at close range without causing more disturbance among them than an occasional gurgling protest. Except for one dead pelican we saw no other kind of sea- bird on the floor of the crater, though on the outside slopes were numbers of nesting tropic-birds, terns, Creagrus and Galapagos gulls. In all our wanderings we had seen no tortoise nor traces of one anywhere, although not so many years ago they were probably the most usual sight on the islands. The whaling ships used to carry them away by the hundred to provide a welcome change of diet on long voyages. Oil hunters from the mainland have made great inroads on their numbers and wild dogs and pigs have probably accounted for numberless eggs and newly-hatched young. Where the tortoises are not actually extinct, the survivors have evidently betaken themselves to the craters of the interior. In 1907 it was reported that these reptiles were most numerous on Duncan, so five members of the expedition went to Duncan in a large motor boat, thirty-six miles away, hoping to verify this report. They beat over the land near the shore and much of the interior of the lesser crater and found only one moderately large tortoise, which, after the most exhausting labor, they managed to carry back to the boat. It seems certain that another unique form of life is well on the road to extinction, thanks to the efforts of man. Our last anchorage in the Archipelago was at Tower Island, in Darwin Bay, a hitherto unmapped bay which we discovered and named. The bay is over a mile square, with deep water up to the very foot of the high cliffs with which it is surrounded. Our one landing beach was extraordinarily beautiful and interesting, fronting a nesting place for hundreds of frigate-birds, boobies, gulls, doves and other native birds, as tame as we had come to expect all the creatures of these islands to be. Here were also deep pools and 20 Zoologica : N. Y . Zoological Society [V ; 1 FIG. 10. THE ISLAND OF INDEFATIGABLE Twenty-five miles across, reaching an altitude of 2300 feet, with dozens of craters, the center of this island is wholly unexplored. Where buccaneers once buried their treasures, wild dogs now roam — waifs from many wrecks. wave-made wells in the rocky lava coast where inconceivably brilliant parrot and angel-fish swam in crystal clear water, and tiny sheltered sandy coves where families of sea-lions basked and played . Here we spent four unforgettable days, working from dawn to dusk to learn all we could of the life of this no-man ’s-land. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V. NUMBER 2 Department of Tropical Research. Contribution Number 152 GALAPAGOS HETEROCERA WITH DESCRIPTIONS OF NEW SPECIES By W. Schaus Honorary Assistant Curator, United States National Museum PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 31, 1923 Nrtu fork Znnlngtral ^ortrty General Office: 111 Broadway , New York City (itiftrrni President, Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer, Cornelius R. Agnew luarb of IHanagrra QHaaa of 1924 Madison Grant, William White Niles, Frank K. Sturgis, Ogden Mills, Lewis Rutherfurd Morris, Archer M. Huntington, George D. Pratt, T. Coleman duPont, Henry D. Whiton, Edward Hatch, Jr., Cornelius R. Agnew, Harrison Williams (Elaaa nf 1925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge C. Ledyard Blair, Anthony R. Kuser, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler (Claaa of 192fi Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor. William T. Hornaday, Director of the Zoological Park; W. Reid Blair, D. V. S., Assistant to Director; Charles H. Townsend, Director of the Aquarium ; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of the Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector; George A. MacCallum, Pathologist; El win R. Sanborn, Photographer and Editor. CMtnrial (ftummitto Henry Fairfield Osborn, Chairman; William T. Hornaday, Charles H. Townsend. Corrected to December, 1923 ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY WITH DESCRIPTIONS OF NEW SPECIES By W. Schaus Honorary Assistant Curator, United States National Museum PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V. NUMBER 2 Department of Tropical Research, Contribution Number 152 GALAPAGOS HETEROCERA December 31, 1923 Route of the Noma, and details and location of the Archipelago. 22 First form on press December 31, 1923. Zoologica Vol. V, N c. GALAPAGOS HETEROCERA WITH DESCRIPTION OF NEW SPECIES By W. Schaus Honorary Assistant Curator, United States National Museum This collection of Heterocera was made by William Beebe in the Galapagos Archipelago, during the period of March 28th to April 9th, and April 19th to 29th, 1923. The opportunity was afforded by the Williams Galapagos Ex- pedition of the New York Zoological Society. The types of new species have been deposited in the collections of the United States National Museum in Washington, D.C. Family ARCTIIDAE Utetheisa ornatrix Linn. Syst. Nat. I. p. 511 (1758) Conway Bay, Indefatigable 13 April 1st. Tagus Cove, Albemarle 1 April 6th. Utetheisa galapagensis Wallgrn. Wien. Ent. Mon. IV. p. 161 (1860) South Seymour 7 Tagus Cove, Albemarle 1 Family NOCTUIDAE Subfamily Agrotinae Chloridea cystiphora Wallgrn. Anthoecia cystiphora Wallgrn. Wien. Ent. Mon. IV. p. 172 (1860) Anthoecia inflata Wallgrn. Wien. Ent. Mon. IV. p. 172 (1860) Anthoecia onca Wallgrn. Wien. Ent. Mon. IV. p. 172 (1860) Conway Bay, Indefatigable 15 c? 13 $ April 1st. James Island 3 c? 5 $ April 7th. Tagus Cove, Albemarle 5'c? 1 $ April 6th. April 23rd. April 6th. 23 24 Zoologica: N. Y. Zoological Society [V ; 2 Chatham Island 3d1 1 $ South Seymour 14 cP 17 $ Chloridea virescens Fabr. Spec. Ins. II. p. 216 (1781) Phalaena rhexia Smith and Abb. Ins. PL 100 (1792) Chatham Island 1 Euxoa williamsi sp. nov. Conway Bay, Indefatigable 8 9 James Island 1 9 Tagus Cove, Albemarle 1 cP South Seymour 1 cP 6.9 Lycophotia oceanica sp. nov. Conway Bay, Indefatigable 1 9 Tagus Cove, Albemarle 1 cP South Seymour 1 cP Subfamily HadeNiNae Cirphis cooperi sp. nov. Conway Bay, Indefatigable 1 9 Subfamily Acronyctinae Magusa orbifera Walk. Cat. Lep. B. M. XI. p. 761 (1857) Tagus Cove, Albemarle 1 9 Trachea roseae sp. nov. Conway Bay, Indefatigable 1 cP Perigea apameoides Guen. Noct. I. p. 229 (1852) James Island Perigea ruthae sp. nov. Conway Bay, Indefatigable 1 cP 1 cP 2 9 April 7th. April 23rd Georgia, II. p. 199 April 7th. April 1st. April 5th. April 6th. April 23rd. April 1st. April 6th. April 23rd April 1st. April 6th. April 1st. April 5th. April 1st. 1923] Schaus: Galapagos Heterocera 25 James Island 1 cf 5 9 April 5th. Tagus Cove, Albemarle 3 rirnttfir &taff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant to the Director and Veterinarian; Charles H. Townsend, Director of the Aquarium ; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosed r; Elwin R. Sanborn, Photographer and Editor. editorial (Eammittre Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V. NUMBERS 4-16. Department of Tropical Research Contribution Numbers 154-162 and 165-169. 4. ICHTHYOLOGY 5. APTERYGOTA 6. HOMOPTERA 7. MALLOPHAGA 8. DIPTERA 9. ARACHNIDA 10. FORMICIDAE 11. TRIUNGULIN LARVAE 12. CHILOPODS 13. COCCIDAE 14. BRACHYURAN CRABS .J. T. Nichols . J. W. Folsom .Herbert Osborn . H. E. Ewing .Charles W. Johnson .Nathan Banks .William Morten Wheeler .Charles T. Brues .Ralph V. Chamberlin ..Harold Morrison .Mary J. Rathbun 15. MACRURA AND ANOMURA ... Waldo L. Schmitt 16. HYMENOPTERA S. A. Rohwer Collected by The Williams Galapagos Expedition PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK February 27, 1924 Route of the Noma, and details and location of the Archipelago. 62 First form on press February 27, 192k Zoologica, VoL V, No. k . A CONTRIBUTION TO THE ICHTHYOLOGY OF THE GALAPAGOS. By J. T. Nichols. (Figure 11). The American Museum of Natural History has recently re- ceived, through Mr. William Beebe, Director of the Harrison Wil- liams Galapagos Expedition of the Department of Tropical Re- search, New York Zoological Society, an interesting collection of marine fishes from the Galapagos Islands, made in 1923. Various fish collections have been made in these islands in past years, but these have been of a fragmentary nature, and the fishes of the islands are still imperfectly known. Two undescribed forms have been found in the present material, and two or three others, known from neighboring seas, deserve special mention, as follows: Cypselurus nigricans (Bennett). Bennett, 1840, described this flying fish from “both the Atlantic and Pacific Oceans, in lat. 5° N.” It is close to Cypselurus furcatus (Mitchill) 1815, New York. Recent authors have recognized furcatus and nigricans as distinct species both occurring in the Atlantic, but it is probable that Atlantic material referred to each is identical; and as the fish to hand agrees with Bennett’s description, and is almost certainly what he had from the Pacific, his name is available for it. It is 215 mm. in length to base of caudal, longest dorsal ray Vz head. Dorsal, 14; anal, 11. Pectorals blackish with a diagonal white stripe and white edging; dorsal pale with a black blotch between the 5th and 11th rays; ventrals pale with slight indication of dusky shading; anal pale; caudal dusky with a pale margin. Haemulon scudderi (Gill). A specimen about 13 cm. long, from South Sey- mour I., of this common grunt of the west coast from Mexico to Panama, seems to be the first definite Galapagos record. On a priori grounds it was to have been expected there, but not necessarily so, as grunts are not very strong swim- mers. Eupomacentrus beebei1 sp. nov. The type, our only specimen, No. 8270, American Museum of Natural History, was collected in a rock pool at Eden, Indefatigable Island, April 1, by the Williams Galapagos Expedition, 1923. It is illustrated in a colored plate in the narrative volume of the Williams Expedition; “Galapagos; World’s End,” by William Beebe, published by Putnam. 1 Named for William Beebe in view of his appreciation of the interest and beauty of a tropical reef and its brilliant fishes. First form on press February 27, 192k- 63 64 Zoologica: N. Y. Zoological Society [V; 4 It has a very striking color pattern as* follows: in the alcoholic specimen dusky all round including pectoral and lower fins; center of top of head, top of back and spinous dorsal, orange (later fading to whitish) ; soft dorsal grey with a dusky base. A large black blotch surrounded by a blue ring on last two spines and anterior soft rays of dorsal, and extending onto the back below. Peduncle pale; caudal grey. Length to base of caudal, 15 mm. Depth in this length, 1.8; head, 2.3. Eye in head, 2.4; interorbital, 3.0; maxillary, 4.5; depth of peduncle, 2.5; pectoral, 1.4; ventral, 1.3; longest dorsal spine, 2.7; longest dorsal ray, 2.1; longest anal ray, 2.2. Upper and lower profiles of body about equally arched, forehead moderately steep and rounded. Tips of ventrals filamentous. Caudal with a shallow fork. Dorsal XII, 15, anal II, 14. Scales, 29. There are light blue marks on the head, including a mark from snout to over front of eyes, thence, after a short break, continued back to nape. A spot in the mid-line between the eyes opposite the break in the above-mentioned mark. A blue spot on the base of the soft dorsal, and a blue spot bordered be- hind with dusky in its axil. Two blue spots followed by a blue stripe on the base of the anal. Some of the scales on sides faintly barred with blue. A color sketch from life has electric blue sides and a scarlet back. There is nothing like this little fish described in the genus Eupomacentrus, where it seems to belong. Young Pomocentrids are at times very deceptive, and it should be noted that we have not seen so small a specimen of Hypsypops rubicundus. Teuthis crestonis (Jordan and Starks). A large surgeon fish from Indefatigable' I. falls within probable individual variation for this species, found from Mexico to Panama. Its caudal is well forked, the upper lobe longest and narrow; color all round dusky, except a yellow margin to the pectoral fin. Runula albolinea sp. nov. Differs strikingly in color from Runula azalea with which it has been directly compared (specimens from Cape San Lucas). No alternate black and white cross marking on the fins as in that species. A narrow white line from over eye to upper caudal origin. The type, No. 8271, American Museum of Natural History, was collected on Indefatigable Island by the Williams Galapagos Expedition, 1923. It is 43 mm. long to base of caudal. Depth contained 6.5 times in this length; head 4.5; snout and eye equal, 3.5 in length of head; width of mouth, 3.0; pectoral, 1.7 ; ventral, 2.6; caudal, 1.2; forked for about half its length. Dorsal beginning on the nape; its rays, 41; anal, 28. Color in alcohol pale greyish to the level of the lower margin of the pupil, white below that line. A narrow white line passing over eye is continued back to the upper caudal origin, and there is another similar white line on the center of the head before dorsal. Dorsal, with a narrow pale edge subtended by a dusky stripe which broadens to tinge all but the base of the central half of the fin; other fins pale. 1924] Nichols: Ichthyology of the Galapagos 65 FIG. 11. RUNULA ALBOLINEA sp. nov. A second example is of the same size and similar in every respect. Four other specimens measure from 35 to 46 mm. in standard length. The largest of these, which has been figured by Miss Isabel Cooper, is somewhat deeper. The color pattern of all is the same. All are from Conway Bay, on the northwest coast of Indefatigable Island, which island is also the type locality of R. azalea. It would seem more probable that the Runula from the coast would differ from that in the islands, than that a second species should occur in the islands where R. azalea was originally described. However, Cape San Lucas material examined agrees with the type description of azalea, and albolinea neither with that description nor with Cape San Lucas material. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” .. . .. - - — Zoologica, Vol. V, No. 5 APTERYGOTA OF THE WILLIAMS GALA- PAGOS EXPEDITION By J. W. Folsom. (Plates 11I-V incl.). The Apterygota collected by the Williams Galapagos Expedition sent out by the Department of Tropical Research, of the New York Zoological Society, and forming the subject of the present report, are as follows: Heterolepisma intermedia sp. nov. Acrotelsa galapagoensis Banks. Lepidocyrtus inter mixtus sp. nov. The two species of Thysanura were collected by Professor W. M. Wheeler, and the single species of Collembola by Mr. William Beebe. Types and other specimens are in the collections of the Depart- ment of Tropical Research of the New York Zoological Society. Five species of Apterygota are now known from the Galapagos Islands, the Hopkins Stanford Expedition, 1898-1899, having taken three species, which were described by Mr. Nathan Banks. One of these three (. Acrotelsa galapagoensis Banks) was present in the Williams collection, but the two others ( Lepisma insularis Banks and Machilis mutica Banks) were not. Heterolepisma intermedia sp. nov. (Plate III, figs. 1-8; Plate IV, figs. 9-12). Dorsally grayish or brownish with the scales; ventrally whitish; body color yellowish white. Legs whitish or spotted with brown or purplish, with yellow setae. Head short and broad. Thorax in width to height as 10:7; thoracic nota subequal; pronotum not strongly narrowing anteriorly, subquadrate with rounded angles. Body elongate, almost parallel-sided throughout, sub-cylindri- cal, dorsally strongly convex. Abdomen narrowing but slightly posteriorly, almost twice as long as the thorax (as 13:7). Tenth urotergite (figs. 1, 2) four times as long as the ninth, trapezoidal, with posterior angles rounded and pos- terior margin slightly concave. Antennae white, spotted with brown or purplish, the spots forming annulations in large specimens; length of antennae unknown. First form on press February 27, 192 4. 67 Plate III. 1, Heterolepisma intermedia, telson of female, X 80. 2, Heterolepisma inter- media, extremity of telson of male, X 80. 3, Heterolepisma intermedia, right maxillary pal- pus, X 100. 4, Heterolepisma intermedia, left labial palpus, X 100. 5, Heterolepisma inter- media, ventral aspect of extremity of abdomen of female, X 53. 6, Heterolepisma inter- media, ventral aspect of extremity of abdomen of male, X 53. 7, Heterolepisma inter- media, seta from front, X 370. 8, Heterolepisma intermedia, setal cluster of left postero-lateral angle of pronotum, X 175. 68 1924] Folsom: Apterygota of the Galapagos 69 Palpi whitish, tinged with brown or purplish, with yellow setae. Maxillary palpi (fig. 3) with last four segments pigmented apically. Labial palpi (fig. 4) with last three segments pigmented faintly; last segment stout, subclavate, with five rounded papillate sense organs (fig. 4). Cerci of unknown length. Styli (fig. 5) two pairs, on eighth and ninth abdominal segments, respectively; white with yellow setae; styli of ninth segment long and large; those of the eighth small. Ovipositor (fig. 5) rod-like, tapering apically, extending beyond the paramedian processes a distance equal to four times the length of the latter; dorsal and ventral valves pseudosegmented. Paramedian processes (fig. 5) each extending not quite to the middle of the adjacent stylus; shorter and broader in the male (fig. 6) than in the female. Eighth urosternite undivided in the male (fig. 6). All the setae are naked. Most of the setae of the head and body are bilobed apically (fig. 7), straight or slightly curving, and of different sizes. They are numerous on each side of the front, on the sides of the head, -above the bases of the antennae, and on the posterior part of the abdomen. On the lateral margins of the thorax are sparse short setae, with occasional long setae. Each thoracic notum has 2+2 setal clusters or combs: at each postero-lateral angle a cluster (fig. 8), and on each side of the posterior margin a comb consisting of one macro- chaeta between two small setae. Abdominal segments one to seven, inclusive, bear 3+3 setal combs: inner dorsal, outer dorsal and lateral (figs. 9-11); abd. 8, 2 + 2, dorsal and lateral; abd. 9 and 10, none. The macrochaetae of the combs are reduced in number as compared with those of most other species. Thus the inner dorsal comb has but one large seta between two small ones (fig. 9); the outer dorsal, two or three macrochaetae (fig. 10); and the lateral comb, two (fig. 11). Ventrally, abdominal segments two to seven, inclusive, bear 1+1 combs, each comb consisting of two (rarely three) macrochaetae between two small setae (fig. 12). On the eighth abdominal segment there is one (sometimes two) macrochaeta on the mesal side of the articulation of the stylus (figs. 5, 6). In the specimens most of the macrochaetae of the clusters and combs had fallen off, but were represented by their sockets, as shown in the figures. Scales oval, orbicular, suboblong, subelliptical, etc., being very variable in form; very finely striate; pale brown or colorless. Length, 5.5 mm.; width, 1 mm. Eleven syntypes. Indefatigable, April 19th, under roots of Bursera, five males and four females, W. M. Wheeler (No. 2262). South Seymour, April 22nd, sifted from soil near beach, one male and one female, W. M. Wheeler (No. 2297). In its generic characters this species is intermediate between Isolepisma Esch. and Heterolepisma Esch. It agrees with the former genus in its arched dorsum and long telson, and with the latter in its subquadrate pronotum, scarcely narrowed anteriorly, and the general character of its setal clusters and combs. It seems preferable to refer this species to Heterolepisma, rather than to erect a new genus for it. H. intermedia is nearly related to the neotropical species H. pampeana Silv., recorded from Buenos Aires, Rio S. Cruz and Porto Piramides, and reported to be abundant on the pampas of Patagonia. Plate IV. 9, Heterolepisma intermedia , right inner dorsal setal cluster of the third abdominal segment, X 370. 10, Heterolepisma intermedia, right outer dorsal setal comb of the fifth abdominal segment. The macrochaetae are represented only by their sockets, X 370. 11, Heterolepisma intermedia, right lateral setal comb of the third abdominal segment, X 370. 12, Heterolepisma intermedia, right ventral setal comb of the sixth abdominal segment, showing one of the two macrochaetae, X 175. 13, Acrotelsa galapagoensis, telson of female, X 25. 14, Acrotelsa galapagoensis, left maxillary palpus of female, X 25. 15, Acrotelsa galapagoensis, left labial palpus of female, X 20. 16, Acrotelsa gala- pagoensis, ventral aspect of extremity of abdomen of female, X 25. 17, Acrotelsa galapa- goensis, ventral aspect of extremity of abdomen of male, X 34. 70 1924] Folsom: Apterygota of the Galapagos 71 Acrotelsa galapagoensis Banks. Lepisma galapagoensis Banks, 1901. Proc. Wash. Acad. Sc., Vol. 3, pp. 541-543, figs. 47-50. (Plate IV, figs. 13-17; Plate V, figs. 18-20). This giant lepismid described by Banks was represented in the collection by five specimens, the study of which has enabled me to add certain details to the original description. Dorsally dark brown; mottled with dark brown, pale brown or grayish scales; body color white. Ventrally silvery white; or with yellowish white thorax and brownish abdomen. Legs golden brown proximally, pale yellow distally, with brown scales and yellow or yellowish brown setae. Head short and broad, rounded anteriorly; eyes not prominent. Thorax broader than the abdomen; pronotum rounded laterally, narrowing anteriorly. Abdomen narrowing slightly posteriorly. Tenth urotergite (fig. 13) elongate-subtriangular, apically acute, fringed with stiff yellow setae. Antennae longer than the head and body by one third; annulated with dark brown and white. Maxillary palpi (fig. 14) white with brown setae (female) or brownish with last three seg- ments darker (male); with segments in relative lengths as 3:5:7. 5:7:8 (female) or as 3:5:7 :5:13 (male). The fifth segment in Bank’s figure is probably abnor- mally short. Labial palpi (fig. 15) white with brown setae (female) or pale brown (male); last segment foot-shaped. Cerci and pseudocercus dark brown, annu- lated with white or yellowish white; the former three fourths as long as the head and body; the latter somewhat longer than the cerci. Styli (figs. 16, 17) seven pairs, on third to ninth abdominal segments; slender, white with brown setae (female) or brownish with golden brown setae (male). Dorsal and ventral valves of ovipositor subequal in length. Ventral valves (fig. 16) finger-shaped in ventral aspect, obliquely pseudosegmented, extending almost as far as the ad- jacent paramedian process of the ninth gonocoxite; this process (figs. 16, 17) tapering uniformly to an acute apex, and extending two fifths as far as the ad- jacent (last) stylus in the female, and one third as far in the male. Eighth ventral segment divided in the female, forming a pair of gonocoxites (fig. 16), undivided in the male (fig. 17). Seventh ventral segment (fig. 18) undivided, rounded posteriorly, without a median fold on the posterior margin. Parameres of male (fig. 17) subclavate, extending as far as the adjacent paramedian pro- cesses of the ninth abdominal segment. Stiff, minutely fringed setae, apically bilobed (fig. 19) occur abundantly; most of them being yellow, though some are brown. On the head are “fourteen tufts of hair seen from above as follows: one in front of each eye, one behind each antenna, two above base of each antenna, one in front of each antenna, and two each side on anterior margin; below on clypeus there are other tufts” (Banks). Marginal clusters of these bilobed setae occur on the thoracic terga, occupying the yellowish pores mentioned by Banks. Each thoracic notum bears seven or eight lateral clusters of setae, the last cluster being at the postero- lateral angle, and bears in addition, near the posterior margin, 1+1 dorsal combs. The combs are simply clusters in which the setae are in alignment, as in figure 17. On the abdomen, in dorsal aspect, the following setal combs or clusters are visible: abd. 1, 2 + 2; abd. 2-7, 3+3; abd. 8, 2 + 2; abd. 9, Plate V. 18, Acrotelsa galapagoensis, ventral plate of seventh abdominal segment of fe- male, X 14. 19, Acrotelsa galapagoensis, seta from front, X 240. 20, Acrotelsa galapagoensis, setal sockets of left outer dorsal comb of fourth abdominal segment, X 110. 21, Lepismid, outline of egg, X 15. 22, Lepismid, egg shell, X 15. 23, Lepismid, egg shell, X 15. 24, Lepismid, nymph, reconstructed from exuviae, X 25. 25, Lepismid, right antenna of nymph, X 100. 26, Lepismid, telson, cerci and pseudocercus of nymph, X 53. 27, Lepidocyrtus intermixtus, eyes of left side, X 320. 28, Lepidocyrtus intermixtus, right hind foot, X 1260. 29, Lepidocyrtus intermixtus, left mucro, X 1260. 72 1924] Folsom: Apterygota of the Galapagos 73 1+1; abd. 10, 3 + 3 large marginal clusters and sometimes a small fourth distal pair (fig. 13). Of these just enumerated, lateral clusters occur on abd. 1-9 inclusive. Most of the clusters and combs contain many macrochaetae (fig. 20). The thoracic sternites are fringed with stiff yellow setae; the proster- num has three combs on each side, the meso- and metasternum each two. On the abdomen, ventrally, there is a setal cluster on each side of the articulation of each stylus on abd. 3-7, inclusive (fig. 18), the mesal clusters forming combs; and a mesal comb on abd. 8 (figs. 16, 17). Scales simple, varying greatly in form: obovate, ovate, suboval to orbi- cular, etc., very minutely striate. The scales are sometimes colorless but usually pigmented with brown, especially apically; the fine close striae give interference colors of violet or blue. Length of largest female, ‘18 mm.; width of mesonotum, 5 mm.; length of antennae, 25 mm.; of a lateral cercus, 15 mm. Length of largest male at hand, 9 mm. Six specimens, collected by Prof. W. M. Wheeler. Two females, Indefatig- able, April 8th, 22nd, under stones on beach (Nos. 2188, 2306). One female, Daphne Major, April 24th (No. 2333). One male, South Seymour, April 25th, (No. 2423). Two males, James Island, April 5th (pinned). The types described by Banks were taken on Hood Island in May. A. galapagoensis is closely related to the West Indian species A. gigantea Esch., as regards the number of styli, the form of the labial palpus, etc., but differs from the latter species in regard to the color of the legs, styli and palpi, and in some other respects, particularly in having no median fold on the posterior border of the seventh ventral plate of the female. The genus Acrotelsa contains the largest known lepismids. A. gigantea Esch. attains a length of 21 mm., and A. galapagoensis Banks, 20 mm. Lepismid Eggs and Nymphs. (Plate V, figs. 21-26). Mr. Beebe collected, on South Seymour Island, April 22nd, hundreds of egg shells, which were at the time correctly identified by Professor Wheeler as lepismid. These were found under a slab of lava rock upon a small amount of earth . They were not attached to the stone but were loose, on and in the thin layer of soil. These were shells of eggs that had hatched. Among them was rarely an unhatched egg, like that represented in figure 21. The shells are brownish yellow, the eggs broadly elliptical to sub- ovate, and smooth externally. At one end, the narrower end of the subovate egg, the nymph escapes by pushing off a cap, leaving usually an irregular torn margin (fig. 22), though occasionally a straight edge (fig. 23), indicating a pre-existing line of weakness. The eggs average 1.45 mm. in length by 0.9 mm. in width. Fre- quently several egg shells are held together in a cluster by means of a gelatinous substance. 74 Zoologica: N. Y. Zoological Society [V; 5 Among the egg shells were hundreds of minute lepismid. skins, and occasionally a skin of the same kind could be seen within an egg shell. These skins had all belonged to nymphs of practically the same size and were probably exuviae of the first molt. The more complete skins showed a median dorsal split extending always the length of the head and thorax and frequently to the end of the abdomen. Though all the skins were more or less distorted and fragmentary, it was not difficult to reconstruct from them the form of the nymph, shown in figure 24. As little is known about the early stages of lepismids, except what Heymons has recorded, it is worth while to describe this im- mature form, in comparison with adult lepismids. The head is vei y large in proportion to the body. The thoracic segments are equal and simple, and the thorax but slightly broader than the abdomen ; the latter being only one fourth longer than the thorax, with parallel sides scarcely narrowing posteriorly, and with segments mostly sube qual . There is no imbrication of the abdominal sclerites, which simply meet, edge to edge. In the exuviae, at least, there are no traces of external organs of reproduction, with the exception of a median slit dividing the ninth ventral segment into halves almost to the posterior border of the eighth. The mouth parts are peculiar, forming a cone ; there being an enclosing ventral sheath (probably the labium, as in Hemiptera) which is split along the median dorsal line — a condition that 1 have not as yet encoun- tered in any adult lepismids. The antennae (fig. 25), inserted close together, are short and stout, only three fifths longer than the head, with twenty -four segments (the adults would have very many), the segments being mostly subequal in length. The cerci and pseudo- cercus (fig. 26) are slightly longer than the head, stout, and elongate- conical; the cerci having thirteen segments (rarely twelve) and the pseudocercus fifteen (the adults would have many more). There are no traces of styli. The legs have short stout segments, like those of an embryo; the tarsus bearing three claws, as in all lepismids. The setae of the body and legs are sparse, short and stiff. No scales are present, though the cuticula is elevated into minute, closely-set papillae. Characters of generic value are practically absent in these nymphs; though they might possibly be found in the mouth parts. The form of the telson is one common to many genera of Lepismidae. 1924] Folsom: Apterygota of the Galapagos 75 Clues to the identity of these nymphs were found, however, in two cast skins of well-grown individuals that occurred among the egg shells and small exuviae. These two larger skins were far from perfect, but showed the same peculiar kind of mouth parts and the same form of telson found in the smaller skins; and exhibited, more- over, well developed setal combs, in which the macrochaetae were still in place. These large setae were exceptionally long, bilobed. apically, and strongly fringed. The large skins showed lepismid eyes, of which I did not detect any traces in the small exuviae; corneae may have been present in the latter, but if so, were obscured by foreign matter and by the distortion of the cuticula. There were seven pairs of styli. The eggs and nymphs do not belong to either of the two species of lepismids described here; for the form of the telson excludes them from Acrotelsa and the fringed setae from Heterolepisma. The num- ber of dorsal setal combs, the number of macrochaetae in each comb (inner dorsal, 2; outer dorsal, 4 or 5; lateral, 4 or 5) and the number of styli, indicate that the species belongs in or near the genus Ctenolepisma. Lepidocyrtus intermixtus sp. nov. (Plate V, figs. 27-29). White throughout (excepting the black eye-spots), or white with a faint pigmentation of irregular spots of blue, frequently surrounding colorless round or oval spots made by the hypodermal nuclei. The pigment consists of minute round separate granules of blue, and when present occurs scatteringly on the following regions: head, dorsally, laterally and ventrally; an imperfect V-shaped line connecting the ocular spots anteriorly; mesonotum, antero-dorsally and laterally; metanotum, dorsally and laterally; abd. 1-3, a feeble pigmentation dorsally and laterally; abd. 4, dorsally except on anterior fourth, laterally, also ventrally on each side of the middle region; abd. 5, a little pigmentation dor- sally. Antennae white throughout, or all segments spotted with blue but color- less apically. Legs slightly pigmented basally, otherwise colorless. Furcula white throughout. Eyes (fig. 27) three on each side, equal, not on separate pigment spots, but on a common black spot, which is small and roughly elongate-triangular. Anten- nae slightly longer than the head, with segments in relative lengths about as 10:16:17:33; first two segments subcylindrical, third subclavate, fourth narrowly elliptical. Mesonotum covering the prothorax but not projecting over the head; with a dense anterior fringe of divergent setae, feebly clavate and naked. Fourth abdominal segment more than twice as long as the third (from 2.1 to 2.6 times as long). Fifth and sixth abdominal segments with curving fringed setae. Claws minute. Unguis (fig. 28) almost straight, without lateral teeth, with inner margin subequally bidentate, there being one tooth in the middle and an- 76 Zoologica: N. Y. Zoological Society [V; 5 other one fourth from the apex. Unguiculus extending as far as the proximal tooth of the unguis, straight, sublanceolate. Tenent hair feebly knobbed. Furcula attaining the ventral tube. Manubrium and dentes subequal in length, with short curved fringed setae dorsally, and scales ventrally. Dentes crenulate dorsally on the proximal three fourths, the distal fourth bearing minute serra- tions, which are continued beyond the dens on the dorsal side of the mucro (fig. 29). Mucro minute, tridentate; apical and anteapical teeth subequal; proximal tooth oblique as usual, spine-like, in the middle of the dorsal margin. A few long fringed setae extend from the dens nearly or quite to the end of the mucro. A dorsal pair of erect subclavate fringed setae was seen on the second ab- dominal segment of one specimen, and such sensory setae doubtless occur nor- mally on other segments also. Very few scales had remained on the specimens; these scales were oval or elliptical, with obscure, extremely minute and close, short, irregular, longitudinal striae. Maximum length of specimens, 1 mm. Twenty-two syntypes. South Seymour Island, April 22nd. This little col- lembolan was taken incidentally by Dr. William Beebe; it appeared among the hundreds of lepismid egg shells that he collected. This form, like a few other species of Lepidocyrtus, approaches the genus Sira as regards the shortness of the mesonotum. As the furcula bears ventral scales, however, the species is referred to Lepidocyrtus. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica Vol. V, No. 6 HOMOPTERA OF THE WILLIAMS GALA- PAGOS EXPEDITION. By Herbert Osborn. The few species of Homoptera submitted for examination were collected by William Beebe in the Galapagos Islands, and are interesting as bringing to light a species long ago described by Stal with some closely related forms, and also a species of Jassus not hitherto known from these islands. The Williams Galapagos Expedition was sent out by the Department of Tropical Research of the New York Zoological Society. Jassus galapagoensis sp. nov. Head broad, blunt; vertex, female, nearly twice as wide as long, narrower in male, faintly carinate; ocelli close to border, not sunken in pits; front narrow, rather faintly carinate; clypeus twice as long as width at base, expanding toward tip. Pronotum a little longer than vertex, minutely granulate. Genitalia: Female, last ventral segment three times as long as preceding; hind border scarcely produced, faintly sinuate, strongly carinate. Male, plates very slender, widened a little toward tip, extended beyond tip of pygofer. Color: Yellow, tinged with orange; border of front and middle of clypeus, orange or reddish. Pronotum and scutellum blackish, in the female with spots at anterior border of pronotum and borders of scutellum, yellowish; elytra mostly black in the male, but with broad discal stripe and claval nervures, blackish in the female; costa yellowish transparent — this including also most of outer apical areole; wings smoky; veins blackish. Beneath yellowish with black areas on the thorax and coxae; legs yellowish; tarsal claws blackish. In some individuals, especially in the males, there is a dense coppery or bronzy pruinose covering which gives the insect a greenish coppery appearance. Length, female 8 mm.; male 6 mm. Seven specimens, two females (type and paratype), four males (allotype and paratype), and one nymph, from Conway Bay, Indefatigable, Galapagos, April 1st, 1923, and James Islan, Galapagos, April 5th, 1923. The females and males differ considerably in size, but agree closely in marking, except the extent of blackish on the elytra. There is however little question as to their belonging to the same species. The nymph has the vertex elongate, considerably longer than width at base, the borders of the front cinnabar red, extended on basal half of clypeus. The prothorax and mesothorax with large fuscous spots; anterior wing pads mostly fuscous; abdominal seg- First form on press February 27 , 192k 77 78 Zoologica: N. Y. Zoological Society [V; 6 ment greenish yellow with fuscous patches at the sides. Beneath greenish; borders of abdomen tinged with reddish; legs greenish; femora with fuscous patches and a dark annulus near tip. The wing pads extend to base of second abdominal segment which would indicate a third or fourth instar. This species seems most closely related to some of the South American forms, especially auratus Fab., but does not agree with any species known to me, and it seems probable that it was derived from a South American source at so remote a period as to have formed a distinct species. Genus Philatis Still. Rio Jan. Hmip. p. 68, 1858. Philatis productus Stal. Mycterodus productus Stal. Eugenies Resa, Omkring, Jorden, p. 278, 1853. Stal described this species as from “Insulae Galapagenses, Callao, et Panama.” Specimens from Tower Island agree very perfectly with his des- cription. The female has the last complete ventral segment deeply indented each side of the middle and emarginate toward the border with the succeeding segment either hidden at the middle or divided by the base of ovipositor. The pygofer is short, compressed, upturned, the sub-anal plate flattened, narrowed posteriorly and bluntly rounded at tip. Male terminal segment twice as long as preceding; hind border concave; lateral angles reaching above base of the plates; plates closely appressed or fused, deeply spoon-shaped, narrowed to blunt tips extending beyond tip of pygofer. Length 5 mm. Two specimens, male and female, Tower Island, April 28th, 1923. Philatis cinerea sp. nov. Smaller than productus and distinctly grayish in color with minute fuscous punctations, a light gray patch on the disk of elytra and a series of whitish cross veins next the claval suture. Head produced, but not distinctly carinate; margins of vertex slightly elevated; margins of front thin, slightly reflexed. Pronotum short, a little more than half as long as vertex, narrowed at sides; elytra broad, convex; hind border obliquely truncate, nearly right-angled at tip of clavus; wings rudimentary. Genitalia: Female, last complete segment indented each side near the middle, sinuate at the sides; succeeding segment with sides separated by ovi- positor; pygofer short, compressed, scarcely produced upward at tip; sub-anal plate rather long; margins reflexed; tip bluntly rounded. Male, last ventral segment twice as long as the preceding; hind border truncate; plates fused; broadly carinate; tip contracted, blunt, exceeding pygofer. Color: Dark gray; vertex and front tinged with green; body sprinkled with minute fuscous dots; a broad whitish patch, on disk of elytra and a light area 1924] Osborn: Homoptera of the Galapagos 79 bordering the clavus; outer border between veins distinctly fuscous. Abdomen above smoky, beneath somewhat greenish. Length, to tip of elytra, female 5 mm.; male 4.5 mm. Five specimens, three females, two males, Tower Island, April 28th, 1923. Philatis major sp. nov. Larger than productus, with carina less distinct, and color more uniformly pale testaceous. Head triangular; vertex depressed, without distinct carina at the middle; borders slightly raised; front without distinct carina; lateral margins thin, slightly expanded. Pronotum about half as long as vertex, angularly emargi- nate behind; elytral veins conspicuous, reticulate; costa broadly rounded; wings rudimentary. Genitalia: Female, last complete segment shallowly indented near the middle, deeply sinuate each side; pygofer compressed, curved. Male, last ventral segment equal to preceding, these about three times as long as basal segments; plates narrowed, closely appressed, broadly carinate each side; tips blunt, extended beyond tip of pygofer. Color: Pale testaceous; veins in the female faintly margined with fuscous; the costal and apical margin minutely dotted with fuscous. Abdomen of male below faintly tinged with greenish. Length, to tip of elytra, female 6.5 mm.; male 5.5 mm. Four specimens, two females and two males from Conway Bay, Indefatig- able, Galapagos, April 1st, 1923. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica Vol. V. No. 7 BIRD-INFESTING MALLOPHAGA COL- LECTED BY THE WILLIAMS GALA- PAGOS EXPEDITION. By H. E. Ewing. Bureau of Entomology, United States Department of Agriculture. (Figure 12). Our knowledge of the Mallophaga of the Galapagos Islands dates from 1902 when Professor V. L. Kellogg and S. 1. Kuwana published their extensive paper ( Proc . Wash. Acad. Sci., Vol. vi, pp. 457-499, pis. xxviii-xxxi) on the biting lice collected by the Hopkins Stanford Galapagos Expedition, 1898-1899. The louse specimens were collected by R. E. Snodgrass, at present of the Bureau of Entomology, whose researches on the morphology of the biting lice and several other orders of insects are well known. Snodgrass obtained no less than 43 different species from the islands. Later (1906) Professor Kellogg published another paper (Trans. Am. Ent. Soc., Vol. xxxii, pp. 315-324) dealing with the Mallophaga of these islands and the Revillagigedo Islands. This paper was based on a collection made by Rollo Beck in 1901, which included fifty-four species. During the summer of 1923, Mr. William Beebe sent to the writer a small collection of bird-infesting Mallophaga obtained by the Williams Galapagos Expedition of the New York Zoological Society during the same year. Although the number of species sent in by Mr. Beebe is small yet the collection is of considerable interest as it contains a record (new species) from the flightless cormorant ( Nannopterum harrisi). First form on press February 27, 192 4 82 Zoologica: N. Y. Zoological Society [V; 7 LIST OF SPECIES, WITH DESCRIPTIONS OF A NEW SPECIES AND A NEW VARIETY. Family MENOPONIDAE Mjoberg. Genus Menopon Nitzsch. M. auri-fasciatum Kellogg. New Mallophaga, Pt. Ill, p. 43, pi. iv, fig. 5 (1899). From Fregata aquila. Three females and one nymph. M. navigans Kellogg. New Mallophaga, Pt. I, p. 156, pi. xiv, figs. 4 and 5 (1896). From Sula nebouxii, two females and one male; and from the same host (second lot), four females and one male. Genus Colpocephalum Nitzsch. C. unciferum Kellogg. New Mallophaga, Pt. I, p. 140, pi. xii, figs. 1-3 (1896). From Pelecanus sp. Two females. Family PHILOPTERIDAE Burmeister. Genus Philopterus Nitzsch. P. breviformis (Kellogg and Kuwana). Proc. Wash. Acad. Sci., Yol. iv, p. 463, pi. xxviii, fig. 3 (1902). From Progne modesta. One female specimen. Genus Esthiopterum Harrison. E. helleri (Kellogg and Kuwana). Proc. Wash. Acad. Sci., Yol. iv, p. 479, pi. xxx, fig. 3 (1902). From Sula piscator. Three male specimens. E. nannopteri sp. nov. Fig. 12 Female: Head stout, subtriangular; forehead subconical, sides very slightly concave; postantennal region longer and broader than forehead; temporal lobes large, evenly rounded; posterior margin of head slightly convex. Clypeus broader than long, signature shield-shaped, slightly longer than broad and clearly outlined. Trabeculae minute. Mandibles stout, bifid at tip. Eyes small but with projecting corners and jet black pigment spots. Lateral margins of fore- head each with six small setae three of which are situated on clypeal region. Temples each with a very long seta at the posterior angle and three minute prickles on lateral margin. Antennae about half as long as head; second segment longest; last segment tipped with minute setae. Thorax about as long as head but not as wide. Prothorax fully twice as broad as long and bare above except for a pair of long and a pair of small setae at each of the posterior angles. Meso-metathorax twice as long as prothorax and broader, sides divergent posteriorly; each posterior angle with a tuft of four 1924] Ewing: Mallophaga of the Galapagos 83 FIG. 12.— ESTHIOPTEBUM MANN OP TERI sp. nov., female, X 30. long, unequal setae, some of which are longer than the segment itself. Legs large and conspicuous; the posterior pair, which is slightly the largest, reaches to the middle of abdomen. Claws unequal, those of the second legs are nearly equal. Femora slightly swollen and subequal in length to the tibiae. Abdomen swollen and broadest at the fourth or fifth segment; pleural bands dark brown, almost black; tergal bands broadly interrupted at the middle and not quite as broad as the segments, brown in color. Above, the abdomen bears four submedian rows of long setae and on the posterior corners of each segment is one or more long setae, the number being greatest on the next to the last segment which bears a tuft of five such setae at each posterior corner. The first segment of the abdomen is the shortest, the second is longer than either I, III or IV, the last is almost as long as broad. Length, 2.3 mm.; greatest width, 0.8 mm. Nymphs: Similar in shape to adult, but smaller and paler. Smaller nymphs without abdominal bands. Last nymph with dark brown pleural markings. Length of last nymph, 2.0 mm.; greatest width, 0.6 mm. Type host and type locality: From Nannopterum harrisi, Galapagos Islands. Type: Cat. No. 23764, U. S. N. M. Described from one female and four nymphs. Holotype a female. This is the first record of an Esthiopterum species from the flightless cor- morant, and as far as the writer has been able to determine is the only Mallo- phagan record from this remarkable and rather recently discovered bird. Snod- grass obtained some of these cormorants in the Galapagos but no lice from them. 84 Zoologica: N. Y. Zoological Society [V; 7 Beck in his trip to the Galapagos in 1901 took some of these flightless cormorants, two of which are now mounted specimens in the United States National Museum. The Mallophaga which he collected this year were worked up by Professor Kellogg, yet in Kellogg’s paper I find no mention of any lice from the flightless cormorant. Also this host is not mentioned in the list of hosts of the Mallophaga known from the birds of the Galapagos Islands published earlier by Kellogg and Kuwana. The species is distinctive and does not resemble any of those previously listed from the Galapagos. However, it is closely related to E. farallonii (Kellogg), described from the Farallone Cormorant, Phalacrocorax dilophus albociliatus, taken at Monterey Bay, California. It differs from Kellogg’s species in being smaller, having a decidedly wider head, longer setae on the abdomen and much larger legs, and, probably most important of all, in not having the median abdominal blotches. Although of about the same dimen- sions as E. acutifrons (Rudow), taken from Phalacrocorax sulcirostris, the speci- mens from Nannopterum differ from Rudow ’s species decidedly in the shape of the head and in various body dimensions. E. pelagicum (Denny). British Anopleura, p. 173, pi. xiv, fig. 2 (1842). From a petrel. One male specimen. E. potens var. minor, var. nov. Similar to the type form taken from Sula piscator in the Galapagos and described by Kellogg and Kuwana. It differs from the species described by these authors chiefly in size. Males of var. minor are but little over 3 mm. in length, while the length given by Kellogg and Kuwana for potens proper is 4 mm. The females also of minor are somewhat smaller than those of the type form. Speci- mens of minor are more strongly banded than represented in the drawing of potens given in Kellogg’s and Kuwana ’s paper. Type host and type locality: From Sula nebouxii taken in the Galapagos Islands: Type slide: Cat. No. 23765, U. S. N. M. Described from a male, two females and a nymph in one lot and from two males, two females and two nymphs of another lot. All specimens from Sula nebouxii taken in the Galapagos. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica, Vol. V, No. 8. DIPTERA OF THE WILLIAMS GALAPAGOS EXPEDITION. By Charles W. Johnson. (Figures 13-14). This report is concerned with a collection of Diptera made by William Beebe on the Williams Galapagos Expedition. This was sent out under the auspices of the Department of Tropical Research of the New York Zoological Society. The first Diptera described from the Galapagos Islands were collected by Charles Darwin, while on the memorial cruise of H. M. S. Beagle around the world. The specimens were collected from September 15th, to October 20th, 1835, and nine species were de- scribed by Francis Walker in his “ List of Diptera” in 1849. During the cruise of the Swedish frigate Eugenies, the Galapagos were again visited, and the Diptera collected, — numbering twelve species, — were described by C. G. Thomson in 1868. The British S. S. Petrel visited the islands in 1875, and among the insects were two Diptera recorded by F. Smith in the “ Proceedings of Zoological Society of London for 1877, page 84.” During the cruise of the U. S. S. Albatross in 1887 and 1888, these islands were again visited, but only one undetermined species of Culex was reported. In 1898-1899 there was an expedition known as the Hopkins Stanford Galapagos Expedition. Among the insects collected by Mr. R. E. Snodgrass, the entomologist, were thirty-five species of Diptera, that were recorded by D . W. Co quillett . 1 Of these, twenty- four had not previously been reported from the islands and of these ten were described as new. The present paper deals with twenty- eight species, of which thirteen have not been before recorded. Deducting synonyms it makes a total of fifty species of Diptera recorded from the Galapagos Including three undetermined species, the number apparently peculiar to the islands is thirty-eight. 1 Proceedings Washington Academy of Sciences. Vol. Ill, pp. 371-379, 1901. First form on press February 27, 192k 85 86 Zoologica: N. Y. Zoological Society [V; 8 FIGS. 13-14. GALAPAGOMYIA LONGIPES gen. et sp. nov. Fig. 13. Wing. Fig. 14. Front leg. Family CHIRONOMIDAE (The Midges). Chironomus sp. One imperfect specimen in alcohol, South Seymour, April 22nd. Ceratopogon gaiapagensis Coquillett. C. gaiapagensis Coquillett, Proc. Wash. Acad. Sciences, Vol. Ill, p . 372, 1901. Four specimens in alcohol, South Seymour, April 20th. T any pus sp. One imperfect specimen in alcohol, South Seymour, April 22nd. Galapagomyia longipes gen. et sp. nov. Male: Head yellowish, the prominent facial protuberance bearing long black hairs, the proboscis nearly as long as the face, palpi large, yellow with black hairs, antennae yellow, scape about three times the diameter of the first joint of the flagellum, the joints of the latter six in number, are rounded and each bear three verticilli, the terminal joint, which is about double the length of the preceding joint, tapers to a point. Thorax brown, with three dorsal rows of hairs, pleura yellow, with a large, brown, central spot, scutellum brown. Ab- domen with both the dorsal and ventral segments brown, margined posteriorly with yellow, hypopygium comparatively small, in form similar to a Diamesa. Legs long, femora yellow, thickened at the basal half, tibiae brown, the front tibiae about one-fourth longer than the femora, the others but slightly longer, tarsi brown, the metatarsi about one-half the length of the tibiae, the other joints of the tarsi together not quite as long as the metatarsi, fourth joint less than one-half the length of the third, legs covered with fine black hairs, halterers yellow, wings brownish hyaline, the costa, first and second veins hairy. Length, 4 mm. Female: similar to the male but only 3 mm. in length. Ovipositor short. The eggs show distinctly through the thin distended sides of the abdomen. Two specimens in alcohol, Seymour Bay, Indefatigable, April 26th. The italicized portions of the above description represent the generic characters. Family TIPULIDAE (The Crane-flies). Dicranomyia sp. One specimen, Conway Bay, Indefatigable, April 1st. The discal cell is open, the venation resembling that of D. floridana O. S. The abdomen is missing and it would be inadvisable to describe it without the male genitalia. 1924] Johnson: Diptera of the Galapagos 87 Family CULICIDAE (The Mosquitoes). Aedes taeniorhynchus portoricensis (Ludlow). Taeniorhynchus niger Gibs ( non Theobald) Jour. Troy. Med., Yol. 7, p. 382, 1904. Culex portoricensis Ludlow, Can. Ent., Yol. 37, p. 386, 1905. Aedes ( Taeniorhynchus ) portoricensis Dyar. Proc. U. S. Nat. Mus., Vol. 62, p. 88, 1922. Two mounted specimens, Conway Bay, Indefatigable, April 1st, and some forty-eight specimens in alcohol, from Seymour Bay, Indefatig- able, South Seymour and Tower, April 18th to 28th. This is undoubtedly what Coquillett recorded from the Galapagos as Culex taeniorhynchus. Differs from taeniorhynchus only in having the last hind tarsal joint strongly blackish tipped. The habits are the same as those of taeniorhyn- chus of which this is to be considered a local race.” (Dyar). The larva were „ in “salt water tide pool,” at Eden, April 2nd. Family TABANIDAE (The Horseflies). Tabanus vittiger Thomson. T. vittiger Thom., Eugenies Resa, p. 451, 1868. Four specimens, Conway and Seymour Bays, Indefatigable, April 1st and 22nd. This species resembles the common T. lineola Fabr. Family BOMBYLIIDAE (The Bee-flies). Villa primitiva (Walker). Anthrax primitiva Walker, List Diptera, II, 257. 1849. Anthrax lateralis Thomson, Eugenies Resa, p. 482, 1868. non Say 1823. Anthrax nudinscula ? Coquillett, Proc. Wash. Acad. Sciences, III, 373, 1901. non Thomson. One specimen, Conway Bay, Indefatigable, April 1st. The description by Walker seems to agree with this species, and as Thom- son’s name is pre-occupied I have adopted it. Coquillett placed A. lateralis Thomson doubtfully as a synonym of A. nudinscula Thomson, described from Panama, but the description does not substantiate this. Villa tincta (Thomson). Anthrax tincta Thomson, Eugenies Resa, p. 483, 1868. One specimen, Seymour Bay, Indefatigable, April 22nd. The specimen is rubbed, but from the form of the head and color of the wings I can only refer it to this species. Family ASILIDAE (The Robber-flies). Ommatius marginellus Fabricius. One specimen, Conway Bay, Indefatigable, April 1st. This species was recorded from James by Coquillett. The specimen before me is a female and therefore the determination is somewhat doubtful, as there are species in which the costa of the male is not thickened, as for example* O. saccas Walker of Jamaica. 88 Zoologica: N. Y. Zoological Society [V; 8 Family DOLICHOPODIDAE (The Long-legged Flies). Asyndetus versicolor sp. nov. Male: Face with whitish pubescence, front metallic green, palpi black, antennae black, first joint about twice the length of the second, third joint rounded and partly covered by the second, arista about as long as all of the three joints together, thickened at the base. Thorax and scutellum metallic green, covered with a thin yellowish pollen. Abdomen bronze black, with a strong purplish reflection, the first segment and a wide posterior margin on the other segments greenish, which in certain lights show as whitish pollinose spots on the sides of the second, third and fourth segments. Legs greenish black, the front coxae each bearing a pair of long bristles projecting forward. Halter- ers yellow, wings hyaline, veins dark brown. Length, 3 mm. One specimen, South Seymour, April 23rd. Family PHORIDAE (The Hump-backed Flies). Aphiochaeta scalaris (Loew). Phora scalaris Loew Cent., VII, p. 100, 1869. One specimen, Tower, April 23rd. Described from Cuba, it seems to be a widely distributed tropical species. Family SYRPHIDAE (The Flower-flies). Baccha clavata (Fabricius). Syrphus clavata Fabr., Ent. Syst., IV, 298, 1775. Baccha fascialis Thomson, Eugenies Resa, p. 504, 1868. Two specimens, South Seymour, April 23rd. A widely distributed tropical and subtropical species. Family SARCOPHAGIDAE (The Flesh-flies). Wohlfahrtia inoa (Walker). Sarcophaga inoa Walker, List Diptera, IV, 832, 1849. Three specimens, Conway and Seymour Bays, Indefatigable, April 1st, and 22nd. This species is readily recognized by its prominent epistoma and pubescent aristae, the abdomen has four rows of uniformly pollinose spots on a permanent black ground, third vein of the wings hairy at the base, almost to the cross vein. This is referable to the genus Wohlfahrtia, although there are some authors who would erect a new genus for it without wincing. Sarcophagula occidua (Fabricius). Musca occidua Fabr., Ent. Syst., IV, 315, 1794. Sarcophagula occidua Aldrich, Sarcophaga and Allies, 40, 1916. One specimen, Daphne Major, April 22nd. A widely distributed tropical species. Sarcophaga violenta Walker. S. violenta Walker, List Diptera, IV, 826, 1849. One specimen, South Seymour, April 23rd. The specimen is a male a little larger (14 mm.) than the measurement 1924] Johnson: Diptera of the Galapagos 89 given by Walker, but otherwise agreeing with the description. It was recorded by Coquillett from Albemarle. Sarcophaga reversa Aldrich. S. reversa Aldrich, Sarcophaga and Allies, 127, 1916. Two male specimens, South Seymour, April 23rd. This has a wide distribution and the specimens agree so well with the description and figure of the hypopygium, that there is little doubt that they represent this species. Sarcophaga obtusifrons Thomson. »S. obtusifrons Thomson, Eugenies Resa, 536, 1868. One specimen in alcohol, Seymour Bay, Indefatigable, April 23rd. Family MUSCIDAE. Codiliomyia macellaria (Fabricius). Musca macellaria Fabr., Syst. Ent., p. 776, 1775. Musca phanda Walker, List Diptera, IV, 869, 1849. Lucilia quadrisignata Thomson, Eugenies Resa, p. 544, 1868. Compsomyia macellaria, E. L. Arribalzaga, Anales Soc., Cien. Argentina, X, p. 70, 1880. Williston, Proc. U. S. Nat. Mus., XII, 203, 1889. Chrysomyia quadrisignata Coq. Proc. Wash. Acad. Sci., Ill, 375, 1901. Cochliomyia macellaria Towns. Jour. Wash. Acad. Sci., V, 646, 1915. Callitroga macellaria Johns, Bull. Amer. Mus. Nat. Hist., XLI, 439. 1919. One specimen, Conway Bay, Indefatigable, April 1st. The specimen agrees with the descriptions of both Walker and Thomson and I find no character to separate it from the common and widely distributed “Screw-worm fly” C. macellaria Fabr. The synonymy of this species is large, although some of the twenty-six species placed there by Arribalzaga and Willis- ton will ultimately prove to be good species when more thoroughly studied. The Musca ochricornis Wied., recorded from the Galapagos by F. Smith, may also represent this and not the Brazilian species, which is placed in the genus Lucilia. Family ORTALIDAE. Pareuxesta latifasciata Coquillett. P. latifasciata Coq., Proc. Wash. Acad. Sciences, III, 376, 1901. P. intermedia Coq., 1. c. p. 377. Seven males and seven females mounted and thirty-nine in alcohol. Tower, April 28th and 29th. “Hundreds of these flies clustered on gull excrement on the beach.” The color of the last two abdominal segments of the female varies from yellow to black, thus eliminating one of the characters used by Coquillett to separate P. intermedia from P. latifasciata. The width of the apical band on the wing is also variable, and it is impossible to draw the line between two- thirds as wide as the preceding hyaline interval in one, to one-half to three- fifths as wide in the other. I am therefore making P. intermedia a synonym. The larvae were also collected. They are whitish, cylindrical, tapering anteriorly to a point. The hook-like mouth parts can be seen through the some- what transparent tegument, the posterior end is truncated with two prominent 90 Zoological N . Y . Zoological Society [V;8 anal tubercles, ventral surface with low transverse ridges armed with rows of small chitinous spines. Length, 4 mm. Pareuxesta obscura Coquillett. P. obscura Coq., Proc. Wash. Acad. Sci., Ill, 377, 1901. Seven, specimens, Daphne Major, April 22nd. All the specimens are males, but the narrow bands of the wings readily separates it from P. latifasciata. One specimen has the bands obsolete, suggest- ing that possibly P. hyalinata Coq., with unmarked wings, may represent an extreme variation. Family CHLOROPIDAE. Hippelates pusio Loew. H. pusio Loew, Cent., X, 87, 1872. Eight specimens (four in alcohol). Tower, April 28th, 29th. “These flies were attracted to small abrasions on our hands or arms.” This, and several other species, are often prevalent in the southern States and West Indies and largely instrumental in spreading an infectious disease known as “sore eye” or pink eye, see Scharz, “Hippelates plague.” Insect Life, Vol. 7, p. 137, 1895. Family DROSOPHILIDAE (The Pomace-flies). Drosophila willistoni Sturtenant. D. pallida Williston, Trans. Ent. Soc. London 1896, p. 415 (non Zetter- stedi 1847). D. willistoni Sturt., Ann. Ent. Soc. Amer., IX, 327, 1916. N. Amer. Drosophila, p. 89, 1921. One specimen, a female, in alcohol, South Seymour, April 22nd. This seems to belong to this widely distributed species of tropical America. Family AGROMYZIDAE. Odinia williamsi sp. nov. Male and female: Front, vertex and occiput grayish white, the front and vertex together forming a quadrangle, with three pairs of frontal orbital bristles, the two upper pairs reclinate, the lower pair convergent, the two pairs of vertical bristles reclinate, the lower pairs of ocellar bristles proclinate and the upper pair reclinate, a fine curved line extends from above the base of each antennae, upper half of the face black, the lower part and cheeks silvery white, vibrissae black, curved, with the basal half thickened, proboscis and palpi yellow, anten- nae with the second joint white above and black below, third joint about One and a half times as long as the second, yellow, with a broad longitudinal black stripe, arista black, base yellow. Thorax grayish with very small black spots at the base of each hair and bristle, one pair of acrostichals and four dorso- centrals, scutellum gray margined with brown and with four marginal bristles, abdomen gray, sparsely covered with black hairs, second, third and fourth seg- ments each with pairs of brown subdorsal and smaller lateral spots. Femora black, tips of the femora and the tibiae and tarsi yellow, the posterior tibiae with a basal and subapical band of brown, halterers yellow, wings grayish, 1924] Johnson: Diptera of the Galapagos 91 with some thirty irregular black spots, that are surrounded by a narrow hyaline margin, tegulae white. Length, 3 mm. Two specimens, South Seymour, April 22nd, 23rd. This beautiful little fly is dedicated to the promoter of the expedition. Family HIPPOBOSCIDAE (The Tick-flies). Olfersia spinifera (Leach). Feronia spinifera Leach, Mem. Wernerian Nat. Hist. Soc., II, 557, Tab. 24, /. 1-3, 1817. Eleven specimens, Tower, April 26th-28th. Living upon the Frigate-bird ( Fregata aquila). Olfersia fossulata Macquart. O. fossulata Macq., Dipt. Exot., II, part 3, p. 434, 1843. One specimen, Daphne Major, April 22nd. From the Brown Pelican, ( Pelecanus fuscus calif or nicus) . Recorded by Coquillett from Wenman. In the absence of specimens of the true O. spinifer I cqnfused this with that species in my paper on the Diptera of the Bahamas (Psyche XV, 80, 1908). The species there recorded from the Cormorant and Booby are really this species, which measures only about 5-6 mm., while O. spinifer is from 8-9 mm. in length. Ornithoponus americanus (Leach). Feronia americana Leach, l. c. 557, Tab. 27, f. 1-3, 1817. Ornithoponus americana Aldrich, Ins. Ins. Menst. , XI, 77, 1923. Seven specimens, Seymour Bay, Indefatigable, April 22nd. From a hawk, Buteo galapagensis. The specimens agree with this common and widely distributed species, which has been recorded from three species of Buteo and several of the owls. Ornithoponus intertropicus (Walker). Or nithomyia inter tropica Walker, List Diptera, IV, 1144, 1849. Olfersia intertropica Austen, Annul, Mag. Nat. Hist., ser. 7, XII, 264, 1903. Three specimens, Seymour Bay, Indefatigable, April 22nd. From a heron, Butorides sundevalli. This species is closely allied to 0. albi- pennis Say, which frequents the various herons of North America. The types described in this paper are in the Laboratory of the Department of Tropical Research, New York Zoological Society. ADDENDA. In preparing the above paper I did not discover until after I had returned the galley proof that Dr. C. H. T. Townsend had described four new species, — the genotypes of four new genera, from the Galapagos. The descriptions are based on species previously recorded by Coquillett under other names in his report on the Diptera collected by the Hopkins Stanford Galapagos Expedition, but this fact is not stated by Dr. Townsend. In the Insecutor Inscitiae Menstruus, vol. 5, p. 163, 1917, under the title “New Genera of Amobiinae” he describes Opsophytopsis insularis, Albemarle, 92 Zoologica: N. Y. Zoological Society [V ; 8 Jan. 18, 1899. Under “Genera of the Dipterous tribe Sarcophagini” (Proc. Biol. Soc. Washington, vol. 30, p. 196, 1917), he described Prosthetocirca cana, Narborough, Jan. 13-29, 1899, Albemarle, Jan. 1-18, 1899, Gigantotheca galapagensis, Albemarle, Jan. 18, 1899, and Sarothromyiops cinctus, Culpepper* Dec. 10, 1898. The types are in the U. S. National Museum. The only comment, by Dr. Townsend relative to previously described species is, — “I can identify none of the above Galapagos forms with Sarcophaga inoa Walker.” It seems likewise difficult from the description to identify Dr. Townsend species. Therefore, a study of his types is just as essential as a study of those of Walker. In referring this complex to Dr. J. M. Aldrich he says: — “It is. too bad you have not been enjoying these new genera for the past six years. There are a few good genera among them and I think Prosthetocirca is one, as. the front rows do not diverge below, which is a pretty good character. Coquil- lett’s specimen with his label Sarcophaga inoa was included by Townsend in the material of his Prosthetocirca cana but he did not mention that helpful item. I do not believe that Coquillett was correct because Walker’s statement about the ‘four hoary spots,’ on each segment does not seem to agree. Town- send’s Sarothromyiops cinctus is a synonym of P. cana as he could easily have ascertained if he had spread the genitalia. He separates the genera on anterior acrostichals which are in this case not of specific importance. We have several other specimens and one or two show a single acrostichal developed.” This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica, Vol. V, No. 9 ARACHNIDA OF THE WILLIAMS GALA- PAGOS EXPEDITION. By Nathan Banks. (Figures 15-18 incl.). This expedition of the Department of Tropical Research under the auspices of the New York Zoological Society visited the Gala- pagos in April, 1923. The collection of Arachnida was made by William Beebe. The Arachnida taken by the expedition are mostly those pre- viously recorded from the islands. Four species are described as new. There are twenty-four species of spiders, five of mites, and two other Arachnids, thirty-one in all; nine of these are new to the fauna of the islands. Several of the additional species are common in the West Indies or Central America. Order ACARINA. Family ERYTHRAEIDAE. Erythraeus remotus sp. nov. (Fig. 15). Body about one and three-fourths times as long as broad, broadest in front of the third legs, but not much narrowed behind, in front with a broad, rounded lobe. Dorsum clothed with many short, spine-like hairs, some near the eyes First form on press February 27, 192it 93 94 Zoologica: N. Y. Zoological Society [V; 9 and crista longer. Cephalothorax not marked behind, two eyes each side, about diameter apart; crista short, enlarged at tip, and in front, where there are five long bristles. Legs very slender, front pair longer than the body, hind pair twice as long as body, second and third pairs about as long as body. Tarsus I is about two-thirds as long as the metatarsus, tarsus IV about two-fifths as long as the metatarsus, latter a little longer than the tibia. Palpus rather slender, penultimate joint stout, pointed, last narrowed at base, with several bristles at tip. Length, 1.7 mm. Under leaves on beach, Indefatigable, April 19th. Atomus sp. One specimen, Tower, April 27th. FIG. 16. AMBLYOMMA WILLIAMSI sp. nov. Dorsum of male. Shield and stigmal plate of female. Family IXODIDAE. Amblyomma williamsi sp. nov. (Fig. 16). Male : red brown; a large pseudoscutum, and two radiate spots each side more yellowish; legs pale yellowish; venter reddish or yellowish. Body about as broad as long, in front rather shouldered, making the front broader than usual; pseudoscutum truncate behind, two raised spots each side and a median streak deeply and heavily punctured, except the median streak; the punctures separate; hind margin with a few hairs. Coxal tubercles very small, hardly noticeable on the hind coxae; hind tarsus tapering, without a spur below. Stig- mata more slender than in female, the smooth area a little larger. 1924] Banks: Arachnida of the Galapagos 95 Female: red brown, more reddish beneath; legs pale yellowish; scutum with large pale spot on the hind lobe, and extending forward each side on the lateral lobes. Body with many short, stout hairs, not arranged in rows; no long hairs. Scutum rather broader than long, cordate, posterior sides evenly convex, finely, evenly and deeply punctured, the punctures separate; eyes small, hardly convex; porose areas rather large, nearly circular and about diameter apart; coxae with only very small tubercles. Stigmata subtriangular, with a long smooth area in front above, the stigmata proper being cone-shaped. Length of male, 2.5 mm.; engorged female, 8 mm.; the scutum; 1.6 mm. From Conolophus lizards, South Seymour, April 20th. Differs from darwini by hairs not arranged in rows, and from hirtum by absence of long hairs. Amblyomma darwini Hirst. From Amblyrhynchus lizards on South Seymour, April 20th, and Indefatigable, March 28th and from Tropidurus lizards on Eden, April 2nd. Ornithodoros talaje Guerin. From Eden Island, April 2nd,; occurs in Central America. Order SOLPUGIDA. Ammotrecha solitaria Banks. Several from Tower, April 27th, and Daphne, April 23rd. Order SCORPIONIDA. Hadruroides lunatus Koch. Specimens from Eden, April 1st; South Seymour, April 20th and 22nd.; and Daphne, April 23rd. Order ARANEIDA. Family FILISTATIDAE. Filistata fasciata Banks. Immature from Tower, April 28th. Family SCYTODIDAE. Scytodes fusca Walck. One from Indefatigable, April 4th; known from the West Indies. Scytodes hebraica Simon. One from Tower, April 28th; known from Central ^America and West Indies. Sicaroides ultriformis Butler. Immature from Eden, April 2nd, under stones. Family DYSDERIDAE. Ariadne tarsalis Banks. One from South Seymour. 96 Zoologica: N. Y. Zoological Society [V; 9 FIG. 17. DICTYNA REMOTA sp. nov. Palpus from side and beneath; side view of mandible of male and epigynum. Family DRASSIDAE. Zelotes galapagoensis Banks? An immature specimen from the beach of Indefatigable, April 19th, agrees with the description, but one cannot be certain without a mature specimen. Family CLUBIONIDAE. An immature specimen of a Clubiona, or allied genus, is from Tower, April 27th. 1924] 97 Banks: Arachnida of the Galapagos Family DICTYNIDAE. Dictyna remota sp. nov. (Fig. 17) Cephalothorax yellow-brown, with white hairs; legs pale, unmarked; abdomen whitish to greyish, with brown basal median stripe and two lateral stripes behind; venter pale. Related to D. parietalis of Central America, with which I formerly identified it. It differs in several respects; the female shows no bands on the legs the epigynum shows two smaller dark spots, farther apart, and the dark lines from them run mostly forward. The male has the same shaped mandibles as in D. parietalis, with the spine near outer base; the palpi are similar, the patella has a swelling at inner tip, the tibia has a hook at base similar to that of D. parietalis, but the subapical process is very different, a slender pointed process, whereas D. parietalis has a broad plate. Length, 1.6 to 2 mm. Daphne, April 23rd. Family THERIDIIDAE. Lathrodectes apicalis Butler. From South Seymour, April 20th. Argyrodes jucundus Cambr. From Albemarle, April 6th. Family EPEIRIDAE. Gasteracantha insulana Thorell. From Albemarle, April 6th. Argiope argentata Fabr. From Indefatigable, April 8th, Duncan, April 25th, Tower, April 27th, Daphne, April 23rd, and South Seymour, April 20th. Epeira oaxensis Keys. From Indefatigable, April 4th and 8th, Duncan, April 25th, Tower, April 27th and 28th, James, April 4th, Eden and Tagus, April 6th, South Seymour, April 25th. Epeira labyrinthea Htz. From Daphne, April 23rd, and Tower, April 27th, 28th and 29th. Epeira prompta Htz. From Indefatigable April 4th, and South Seymour, April 20th. Cyclosa conica Clerck. From Daphne, April 23rd. Nephila clavipes Linn. From Eden, April 6th, common in tropical America. 98 Zoologica: N. Y. Zoological Society [V ; 9 Family SPARASSIDAE. Heteropoda venatoria Linn. One without locality. Olios galapagoensis Banks. From Indefatigable, April 8th, Eden, April 6th, and South Seymour, April 20th'. Family CTENIDAE. Odo insular is Banks. From Eden, under stones, April 6th. FIG. 18. PHANIAS DISTANS sp. nov. Palpus, tibia and metatarsus I, and fang and grooves of mandible. Family ATTIDAE. Plexippus paykulli Aud. et Sav. Immature from South Seymour, April 20th. Philaeus pacificus Banks. From Tower, April 27th. Metacyrba insularis Banks. From Daphne, April 23rd. Phanias distans sp. nov. (Fig. 18). Cephalothorax brown, eye area nearly black, mandibles reddish, legs yellow- ish brown, darker at bases and tips of many joints above; black haired, no fringes. 1924] Banks: Arachnida of the Galapagos 99 some short yellow hair on femur I. Abdomen yellow brown, with much black hair, no distinct markings, faint traces of median stripe in a few narrow chevrons; sternum and venter yellow brown with black hairs. Cephalothorax moderately flattened, not much broadened in middle. Abdomen slender, depressed. Each mandible with a bicuspid tooth on inner groove; sternum tapering in front and behind, the coxae I, however, separa e by nearly width of lip; legs slender, femora I and II more swollen, these with two long bristles above near base; tibia I wi h four spines on inner row, three on outer, tibia II with three in each row, metatarsi I and II with two in each row, tibia IV below with one at base and one near tip, metatarsi IV with one at basal third, and a pair at tip below; patellae III and IV unspined. Male palpus has a process from tibia, at first stout, then very slender and forked before tip, alongside of bulb is a rather stout style from base to beyond tip. Length, 9 mm. From South Seymour. Evidently belongs to Simon’s group of Maevia and Peckham’s Marptusa group. Differs from Marptusa in more separated anterior coxae and rather fewer spines on legs, a though this is doubtless variable. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” ZooloQica, Vol. V, No. 10. THE FORMICIDAE OF THE HARRISON WILLIAMS GALAPAGOS EXPEDITION. By William Morton Wheeler. (Figures 19-27 incl.). Four years ago I published an account of the Galapagos ants collected by the Expedition of the California Academy of Sciences in 1905 and 1906 and of those previously taken on some of the voy- ages of the “Albatross/’1 The fact that the description of these materials revealed almost nothing in regard to the habits of the insects and the prospect of finding additional species led me to accept the very generous invitation of Mr. Harrison Williams and Mr. William Beebe to join their expedition to the archipelago. Although the region that could be covered in the time at our disposal was limited, we were able to secure nineteen different forms, eight of which (two species and six varieties) are new to science and one (M onomorium floricola) a well-known tropicopolitan ant not hitherto recorded from the islands. The collection of so small a number of forms in any spot on the American mainland, except its arctic and antarctic ends, would have very little or no significance, but con- sidering the meagerness of the Galapagos ant-fauna and the time and effort required in securing even a small number of specimens, those obtained and the observations made on their habits are well worth recording. Many other groups of insects, such as the butter- flies, bees, wasps, termites, many families of Diptera, Coleoptera, etc., are even less abundantly represented in the Galapagos fauna. To the general statements on the Formicidae in the introduction to my paper of 1919 I have nothing to add. Family FORMICIDAE. Subfamily Ponerinae. Cylindromyrmex williamsi sp. nov. (Fig. 19, b and c ). V Worker. Length 4.5-6 mm. Closely related to C. striatus Mayr. Head longer than in that species, oblong, about one and a fourth times as long as broad, with nearly straight, parallel sides and feebly excavated posterior border. Eyes rather large, but 1 The Ants of the Galapagos Islands, Proc. Calif. Acad. Sci. (4) 2, 1919, pp. 259-310, First form on press February 27, 1924 101 102 Zoologica: N. Y. Zoological Society [V; 10 a c b FIG. 19. CYLINDROMYRMEX STRIATUS MAYR; C. WILLI AMSI sp. nov. a , head of female C. striatus; b, head of female and c, bead of worker C. wi'lli amsi. not very convex, their anterior orbits at the median transverse diameter of the head. Small workers with distinct ocellar pits, large workers with a small an- terior ocellus. Mandibles rather flat, with nearly straight external borders, the apical tooth large and blunt, the apical border somewhat undulating but not distinctly denticulate. Clypeus short, depressed, flat in the middle, more convex on the sides, which are produced and rounded anteriorly while the median border, though broadly rounded, recedes. Frontal area elongate ellipti- cal, deeply impressed. Frontal carinae moderately prominent, extending back to the posterior orbits and thence continuous with rugae to the posterior corners of the head as in striatus. Antennal scapes reaching to the eyes, about four times as long as broad; funicular joints 2-6 much broader than long, 7-8 also transverse but proportionally longer, terminal joint about one and one-half times as long as broad. Thorax as in striatus but somewhat longer and more rectangular, about twice as long as broad; epinotal declivity sharply marginate laterally and above. Petiole in the smallest worker as broad as long, in larger individuals distinctly longer than broad; its anterior surface marginate on the sides and above, its anteroventral surface with a large, blunt, compressed tooth. Postpetiole from above trapezoidal, one and one-third times as long as its anter- ior border, the sides much less convex than in striatus. First gastric segment about one-fourth broader than the postpetiole; pygidium subtruncate, with a row of acute denticles on each side of its posterior half. Legs very distinctly more slender than in striatus, the anterior tibia fully three times as long as broad (in striatus scarcely two and one-half times). Middle and hind tibiae each with a single pectinated spur. * Shining; mandibles subopaque, sharply longitudinally striate. Middle of clypeus smooth, its sides, the scapes and the large antennal foveae finely longitudinally striate; cheeks and gular surface of head coarsely and somewhat irregularly, longitudinally striate; head above, thorax, petiole and postpetiole sculptured much as in striatus, with coarse longitudinal striae separated by rounded rugae, those on the thorax, petiole and postpetiole very regular, on the 1924] Wheeler: Formicidae of the Galapagos 103 head somewhat divergent posteriorly and dividing, thus increasing their number at the occipital border. There are also some coarse interrugal punctures in this region. The number of rugae on the vertex between the posterior continuations of the frontal carinae is about 15, there are about 10 on the thoracic dorsum and petiole and about twice that number on the postpetiole. Epinotal declivity and anterior surface of petiole shining and very finely shagreened. Gaster and legs very smooth and shining, with minute, scattered, piligerous punctures. Hairs whitish, very sparse, conspicuous only on the clypeus, tip of gaster and venter; gaster with sparse, rather long pubescence; hairs on the tibiae very sparse, subappressed. Black; tip of last funicular joint and terminal tarsal joints testaceous. Female (dealated). Length nearly 7 mm. Differing from the worker in having the head fully one and one-half times as long as broad. The eyes are larger and there are three rather small ocelli. Thorax somewhat depressed above, the pronotum with concave sides, broader behind than in front and broader than long, its lateral borders straight. Meso- notum and scutellum small, the former as broad as long, arcuately rounded in front. Epinotum with subequal base and declivity, the former of nearly the same shape and size as the pronotum. Sculpture and color similar to that of the worker but the striae on the sides of the thorax are finer. Hairs on the femora and tibiae much more numerous. This species, which I dedicate to Mr. Harrison Williams, is described from two workers and a female taken near the south end of South Seymour Island, April 20th, 1923. In my paper of 1919 I erroneously identified this ant from speci- mens taken by Dr. F. X. Williams at Academy Bay, Indefatigable Island, as Mayr’s C. striatus. On comparing one of the workers collected by Dr. Wil- liams and the three specimens described above with a female of the true striatus (fig. 19 a) taken by Prof. C. T. Brues at Guayaquill, Ecuador and Cameron’s figure of the worker, which he erroneously described as Holcoponera whymperi, I find that the Galapagos species is quite distinct. The legs of striatus are shorter, much more robust and of a different color, the head is shorter, the eyes smaller and less convex and the median portion of the clypeus longer and more produced, the antennal scapes are broader, the rugae on the head, thoracic dorsum, petiole and postpetiole are somewhat stronger and more even, the pubescence on the gaster is shorter and the hairs on the legs much less numerous. The striae on the mandibles, on the contrary, are more superficial and the man- dibles, clypeus, cheeks and antennae are dark red. The three specimens which I took on South Seymour Island, together with several young larvae, formed the entire personnel of a colony which was nesting in the dead branch of a Celastraceous shrub ( Maytenus obovata Hook. fil.) growing near the beach. Many of the dead branches of the same shrub contained flourishing colonies of Calotermes pacificus Banks, but the ants did not actually live among them though evidently occupying galleries which they had once inhabited. This fact is of interest, because colonies of the allied genus Simopone of the Ethiopian and Malagasy regions have been taken in dead branches. Arnold records S. marleyi Arnold as occurring in hollow stems of the castor oil plant at Durban, Natal. That the ants of the two genera, Simo- pone of the Old, and Cylindromyrmex of the New World, which together 104 Zoologica: N. Y. Zoological Society [V;10 constitute Emery’s Ponerine tribe Cylindromyrmicini, prey on termites and tend to establish their colonies near these insects is also indicated by Mayr’s statement that Hetschko found C. brasiliensis in wood in the galleries of a ter- mite at Santa Catharina, Brazil. The larvae of C. williamsi, to which I have referred, were unfortunately lost through breaking of the vial in which they were contained so that I am unable to figure them. They were very long and slender, with narrow, curved neck and small head. The body was smooth, i. e., nontuberculate and covered with numerous, short, even hairs, as in Stigmatomma and allied Ponerine genera. Among the specimens in my collection I find a winged female of a peculiar Bolivian Cylindromyrmex, of which I insert a description: Cylindromyrmex (Met a cylindromyrmex boliviae sp. nov. (Fig. 20). Female. Length 10 mm. Closely related to C. godmani Forel. Head subrectangular, fully one and one-half times as long as broad, as broad in front as behind but somewhat nar- rowed in the middle; its posterior border rather deeply and subangularly excised. Eyes moderately convex, but not as large as in the preceding species, their anterior orbits at the median transverse diameter of the head. Mandibles large, convex, with rounded external borders, a blunt apical tooth and broad, toothless apical border. Clypeus very short, abrupt in the middle, overarched by the very large and projecting frontal carinae, which extend back to the middle of the eyes. Frontal area elongate-elliptical, deeply impressed. Antennal scapes about two and one-half times as long as broad, abruptly narrowed at the base, with straight posterior and convex anterior border as in the preceding species; funicular joints 2-6 extremely short and transverse, the two penultimate joints nearly as long as broad. Thorax depressed above; pronotum transversely sub- oblong, only slightly broader behind than in front, its sides concave, its lateral borders subparallel and marginate; mesonotum as broad as long, its anterior border semicircular, its lateral borders concave; base of epinotum longer than the abrupt declivity which is concave in the middle and submarginate on the sides and above. Petiole subcylindrical, about one and one-fourth times as long as broad, concave on the sides, its anterior surface submarginate on the sides and above, its anteroventral surface with a large, blunt, compressed tooth. Post- petiole as long as broad, somewhat broader behind than in front, with straight sides, anterior and posterior borders, its ventral portion very convex and pro- jecting in front. Pygidium slightly truncated behind, armed with a row of acute denticles on each side of its posterior half. Legs stout as in C. striatus, the fore tibiae somewhat less than two and one-half times as long as broad. Median and hind tibiae each with two well-developed pectinated spurs. Shining; mandibles with fine, interrupted, superficial striae; anterior half of gula with coarse longitudinal striae, posterior half coarsely and sparsely punctate. Cheeks with a long, pronounced longitudinal stria or groove. Anten- nal foveae rather coarsely striate, remainder of head with longitudinal striae separated by rugae which on the front and occiput become coarser, more regular and very slightly divergent. Scapes very finely striolate. Pronotum with about 1924] Wheeler: Formicidae of the Galapagos 105 a FIG. 20. CYLINDROMYRMEX ( METACYLINDROMYRMEX ) BOLIVIAE sp. nov. Female: a, profile, wings removed; b, head. 20 strong longitudinal rugae, separated by deep striae, mesonotum smooth behind, longitudinally striate in front, the striae diverging posteriorly. Scutel- lum smooth and shining, with a few punctures near its posterior border. Epi- notum and petiole longitudinally striate, the striae rather irregular or branching. Sides of epinotum and lower mesosterna rather finely and sharply striate, upper mesosterna and sides of pronotum very smooth and shining. Postpetiole like the gaster, very smooth and shining, with small, scattered, piligerous punc- tures. Hairs yellow, short, erect or suberect, more numerous than in williamsi on the head, thorax and petiole, on the gaster less numerous, longer and more erect. Tibiae and tarsi with long, uneven, suberect, somewhat bristly hairs. Black; mandibles, funiculi, tips and bases of scapes, coxae and tip of gaster deep castaneous red; legs brownish yellow, knees and tips of tibiae darker. Wings distinctly infuscated, with dark brown veins and pterostigma. A single specimen from Mapiri, Bolivia (Staudinger). This species is very closely related to C. godmani of Panama and Ecuador, also known only from female specimens, but the latter species is larger (12.5 mm.), its antennal scapes and tibiae are shorter (scarcely twice as long as broad), the apical borders of the mandibles are bluntly denticulate, the petiole is shorter and has traces of striae, the femora are darker and the wings are described as brownish violaceous as in the bee Xylocopa violacea. The workers and females of the seven known species of Cylindromyrmex may be separated by means of the following table: 106 Zoologica : N. Y. Zoological Society [V; 10 1. Workers 2. Females. 6. 2. Eyes very small and flat; anterior border of petiole emarginate in middle; first gastric segment striate 3. Eyes large and convex, anterior border of petiole entire; first gastric segment smooth and shining 4 . 3. Head nearly twice as long as broad; apical borders of mandibles edendulate. Length 8 mm. (Brazil) longiceps Ern. Andre. Head one and one-half times as long as broad; apical borders of mandibles bluntly denticulate. Length 5. 5-6. 5 mm. (Venezuela) meinerti Forel. 4. Striae of head, thorax and pedicel coarse and regular; coxae trochanters and femora black or dark brown 5 . Striae of head, thorax and pedicel finer and less regular; legs yellow throughout. Length 6-7.3 mm. (Brazil, Paraguay) brasiliensis Emery. 5. Legs slender, entirely black, except the terminal tarsal joints. Length 4.5-6 mm. (Galapagos Islands) .. williamsi sp. nov. Legs stouter, tibiae except their ends, pale ivory yellow (Suri- nam, Peru, Ecuador) striatus Mayr. 6. Frontal carinae moderately large; head not narrowed at the median transverse diameter; postpetiole coarsely longitu- dinally striate; middle and hind tibiae each with a single well-developed pectinated spur 7 . Frontal carinae larger and more projecting; head appreciably narrowed at the median transverse diameter; middle and hind tibiae each with two well-developed pectinated spurs . . 8 . 7. Head one and one-half times as long as broad; legs black, except the terminal tarsal joints. Legs slender. Length 7 mm williamsi sp. nov. Head shorter; legs stouter; tibiae, except their tips, pale ivory yellow. Length 7 mm striatus Mayr. 8. Postpetiole nonstriate; fore tibiae more than twice as long as broad; femora yellow; wings brownish. Length 10 mm. (Bolivia) boliviae sp. nov. Postpetiole somewhat striate; tibiae scarcely twice as long as broad; femora reddish black; wings brownish violaceous. Length 12.5 mm. (Panama, Ecuador) godmani Forel. The foregoing table shows that the species of Cylindromyrmex may be arranged in three distinct groups or subgenera, namely, longiceps and meinerti , which have very small, flat eyes in the worker and for which I suggest the name Hypocylindromyrmex subgen. nov., with longiceps as the type; striatus , brasiliensis and williamsi, which have large, rather convex eyes, moderate frontal carinae and only a single well-developed pectinated spur on the posterior tibiae ( Cylindromyrmex sens, str., with striatus as the type), and godmani and boliviae with large, rather convex eyes, very large frontal carinae and two well- developed pectinated spurs on the posterior tibiae (Metacylindromyrmex subgen. nov., with godmani as the type). The brownish color and minute eyes of the 1924] Wheeler: Formicidae of the Galapagos 107 two species of Hypocylindromyrmex indicate that they do not inhabit twigs like the species of the two other subgenera but that they lead a hypogaeic life, probably in the abandoned galleries of terrestrial termites. Ponera beebei sp. nov. (Fig. 21). Female (dealated). Length nearly 2 mm. Head subrectangular, about one-sixth longer than broad, with distinctly concave posterior border, the sides behind the eyes straight and parallel, con- tracted somewhat in front of the eyes, which are small, rather fiat and scarcely longer than their distance from the anterior corners of the clypeus. Ocelli small and widely separated. Mandibles moderately large, their broad apical FIG. 21. PONERA BEEBEI sp. nov. Head of female. borders with four rather widely separated, acute teeth. Clypeus short, abrupt, strongly carinate, its anterior border advanced and rounded in the middle, sinuate on each side. Frontal carinae rounded, with ciliated borders; frontal groove distinct, extending to the anterior ocellus. Antennal scapes not reaching the posterior border of the head; funicular joints 2-6 subequal, very transverse, more than twice as broad as long; remaining joints forming a 5-jointed club, all the joints of which, except the last, are broader than long and gradually increase in length apically, the terminal joint somewhat shorter than the three preceding joints together. Thorax slightly more than twice as long as broad, as broad behind as in front, rounded anteriorly and posteriorly, its dorsal outline straight in profile, the mesonotum and scutellum small and flat, the former broader than long, with semicircular anterior border; epinotum with straight base and declivity, the latter longer than the former, its sides rounded, not marginate. Petiolar node thick, about one and one-half times as high as long, much narrower than the epinotum, truncated anteriorly and posteriorly and with a rounded summit, which is distinctly thinner than the base. Seen from behind the node is subcircular. The ventral surface in profile is feebly convex, not swollen, and without an anterior tooth. Postpetiole trapezoidal from above, broader than long, narrowed in front, with straight anterior and 108 Zoologica: N. Y. Zoological Society [V; 10 lateral borders; strongly truncated in front. Constriction behind the post- petiole well-developed. First gastric segment distinctly broader than long, from above transversely rectangular. Legs rather slender. Shining; mandibles smooth, with minute scattered piligerous punctures; remainder of body finely and distinctly but not very densely nor deeply punctu- late; sides of thorax finely rugulose; antennal scapes and legs more finely and densely punctulate and less shining. Hairs and pubescence white, abundant, the former short and uneven, most distinct on the clypeus, summit of petiole and tip of gaster, the pubescence long, subappressed, grading into the pilosity, conspicuous on the thorax, petiole and gaster, somewhat shorter on the head, especially on its dorsal surface. An- tennae and legs with fine, dense, subappressed pubescence. Dark brown, almost black, mandibles and legs brownish yellow, or testa- ceous; clypeus, antennae and a large round spot on the front brownish red. A single specimen which I found under a large stone embedded in the sand of the small beach of Tower Island. I have described this form as new because I am unable to refer it to any of the species known to me from specimens or descriptions. The antennae seem to be most like those of P. gleadowi Forel of India and Borneo, but the color, sculpture, shape of petiole, etc., are very different. Subfamily Myrmicinae. Pheidole williamsi Wheeler var. seymourensis var. nov. Soldier and Worker. Differing from the typical williamsi of Indefatigable Island in color, the body being yellowish brown, with the posterior portion of the head, the dorsal surface of the thorax and nodes darker, the gaster almost castaneous brown, and the middle portions of the femora slightly infuscated. Two very small colonies of this variety were found, one in a nest in the sand just beyond the beach on South Seymour Island, the other under a large stone in the bottom of the crater of Daphne Island. Ph. williamsi seems to be closely related to the tropicopolitan Ph. megacephala Fabr., but is smaller, the soldier has a smaller, more rounded head, with longer antennal scapes and there are differences in the structure of the thorax and pedicel, the humeri being less prominent and more rounded and the postpetiole shorter and more trans- versely elliptical. Monomorium floricola Jerdon. Several colonies of this minute tropicopolitan ant, hitherto unrecorded from the Galapagos Islands, were found nesting in dead twigs of Bursera graveolens Trian. and Planch, in the thickets on Tower Island. The workers were assidu- ously visiting the nectaries of the flowers of Cordia lutea Lam. and of the large cactus, Opuntia helleri, which is peculiar to the island. This ant must have reached Tower Island in floating vegetation since the females are wingless. Solenopsis globularia F. Smith subsp. pacifica Wheeler. (Fig. 22). Female (dealated), undescribed. Length nearly 4 mm. Resembling the worker. Head proportionally shorter and more rectangular; eyes much larger. Thorax from above elongate elliptical, narrower than the 1924] Wheeler: Formicidae of the Galapagos 109 head, about two and one-half times as long as broad; epinotum in profile rounded and steeply sloping, without distinct base and declivity; petiole higher and more compressed anteroposteriorly than in the worker; postpetiole short, trans- verse, about two and one-half times as broad as long. Gaster large and elongate. Sculpture, pilosity and color as in the worker, but each of the gastric segments with a broad, poorly defined, brown transverse band. Numerous workers and a single female from several colonies found at Sey- mour Bay, Indefatigable Island, Daphne Island and Tower Island. On Daphne FIG. 22. SOLENOPSIS GLOB XJLARIA F. Smith subsp. PACIFICA Wheeler. a, head of worker; b, thorax and pedicel of same in profile; c, pedicel, dorsal view. Island these ants were nesting under stones at the bottom of the crater, in the other localities under small logs lying on the sandy beach or among the thickets above high water mark. This subspecies was first taken on Albemarle and Tower Islands by F. X. Williams and the Albatross Expedition of 1899. Tetramorium guineense Fabr. In my previous paper I recorded this widely distributed tropicopolitan ant from South Albemarle (F. X. Williams). Emery mentioned it from Chatham Island (G. Baur) and the Albatross Expedition of 1899 secured it from the islands but failed to note the precise locality. I found it only on Tower Island, confined 110 Zoologica: N. Y. Zoological Society [V; 10 to the small beach and nesting under stones and the bark of bushes. The workers were exploring the foliage of the bushes on which great numbers of frigate birds and gannets were nesting. Subfamily Dolichoderinae. Dorymyrmex ( Conomyrma ) pyramicus Roger subsp. albemarlensis Wheeler. (Fig. 23). This ant, previously known only from Albermarle Island, is common at Seymour Bay, Indefatigable Island and on South Seymour Island. It was FIG. 23. DORYMYRMEX ( CONOMYRMA ) PYRAMICUS Roger subsp. ALBEMAR- LENSIS Wheeler. a, head cf worker; b, thorax and petiole of same in profile. also taken by members of the expedition at Tagus Cove, Albermarle and on Eden Rock. It nests near sea-level in flat, crater nests in the sandy soil, or decomposed lava in the hot open spaces among the vegetation. The behavior of the very active and enterprising workers is like that of other forms of pyrami- cus which ranges from our Southern and Southwestern States to Argentina. Tapinoma melanocephalum Fabr. A widely distributed tropicopolitan ant, the “hormiga bottegaria” of the Latin Americans, previously taken on the east side of Indefatigable Island by Dr. F. X. Williams and on the boat at Chatham Island by Dr. G. Baur. I found several populous colonies on Tower Island, nesting in abandoned Calo- termes galleries in dead branches of Bursera graveolens. 1924] Wheeler: Formicidae of the Galapagos 111 Subfamily Formicinae. Prenolepis (Nylanderia) vividula Nyl. subsp. guatemalensis Forel var. eden- ensis var. nov. Eight workers and a dealated female taken by members of the expedition from a single colony on Eden Rock agree closely with the var. cocoensis Forel from Cocos Island, except in color, the gaster being dark brown, in the eyes of the worker which are smaller and more convex than in Forel ’s variety and the pubescence which, especially on the head and gaster, is distinctly longer, so that these regions are less shining. Prenolepis ( Nylanderia ) fulva Mayr subsp. nesiotis Wheeler. A number of workers taken from a single colony nesting in the sand above high water mark at Seymour Bay, Indefatigable Island agree closely with the cotypes of nesiotis from James Island in my collection, except in the coloration of the gaster, which is somewhat darker. The differences do not seem to be sufficient to justify the introduction of a new varietal name. Genus Camponotus Mayr. Perhaps no other group shows the poverty of the ant-fauna of the Galapagos so clearly as the genus Camponotus, which is repre- sented by such an enormous number of forms in continental tropical America. Only three small species have been recorded from the archipelago: senex, macilentus and planus , all by Frederick Smith. He recorded the first as having been taken by W. E. Cookson, commander of the Petrel while on its voyage to the islands in 1875. As this species has since been taken only in Central and South America and as Smith was notoriously careless in making identifica- tions even of species which he had himself described, I believe that we are justified in dropping it from the Galapagos faunal list. In British Guiana during the summer of 1920 and again in Panama during the summer of 1923 I had an opportunity to study C. senex. As Forel was the first to ascertain,2 this ant, like Oecopbylla and many species of Polyrhachis in the Old World and C. formiciformis Forel in Central America3 makes a peculiar nest by using its larvae for spinning leaves together with layers of silk in the form of a ball. The nest may vary from the size of an orange to that of a football and is firmly attached to the twigs of a tree, sometimes at a con- siderable distance from the ground. Several nests may occur on a 1 Biol. Centr. Amer. Formicidae, 1899-1900, p. 139, pi. 2, fig. 5. 3 According to observations recorded in my paper “On the Presence and Absence of Cocoons among Ants, the Nest Spinning Habits of the Larvae, etc. Ann. Ent. Soc. Amer. 8, 1915, p. 323-342, fig. 1. 112 Zcologica: N. Y. Zoological Society [V; 10 single tree and may, perhaps, belong to a single polycladic colony. So far as 1 have been able to observe, senex lives only in the low jungle and almost always on trees or bushes growing along water- courses. This is certainly not an environment to be encountered in the Galapagos Islands. There remain therefore only two species of Camponotus, macilentus and planus , as occurring on the archipelago. They have been taken on nearly all the larger and on several of the smaller islands and are, in fact, the most abundant and therefore the dominant components of the ant-fauna in the low xerothermal zone, the only part of the islands which has been carefully investi- gated. Both are very timid species which rarely attempt to bite even when their nests are violently disturbed. Although they coexist in the same localities, their habits are so different that they do not encroach on each other's activities. The pale yellow C. macilentus is a nocturnal ant, which lives in rather small colonies in dead branches of trees and shrubs (Bur sera graveolens and Mayte - nus obovata) that have been hollowed out by Calotermes and wood- boring beetles. The workers and soldiers were never seen abroad in the day-time. At the time of my visit to the islands in the latter part of April, the nests contained considerable brood and many males and winged females. These phases were also taken in August by Dr. F. X. Williams, so that there must be either two brief periods or, more probably, a single protracted period during which the sexual forms are produced. The black C. planus, on the other hand, is a strictly diurnal ant which forms much more populous communities and nests in the soil about the roots of trees, shrubs, the large arbores- cent Opuntias or in old logs. The entrances are small and concealed so that the nests are found only by accident or by following foraging workers. The latter are most frequently seen on the foliage of the thickets in search of insect prey or visiting the flowers for their nectar. The beautiful clustered yellow flowers of Cordia lutea attract them as well as many other insects, Xylocopa, Chrysopa, beetles, etc., in considerable numbers. On Indefatigable and South Seymour I failed to find any males or winged females in the nests during late April, but these phases were secured by members of the expedition April 1 to 7 on Chatham, Albemarle, James and at Conway Bay, Indefatigable Island. Dr. Williams collected males and winged females on Duncan, Charles and Indefatigable during August, October and November. It would seem, therefore, that 1924] Wheeler: Formicidae of the Galapagos 113 this species either sends off its sexual phases at very different times of the year on different islands or that they are developed continu- ously over a long period from August to the beginning of April. 1 maintained in my former paper that both macilentus and planus have produced a number of varieties, one or more of which occur on each of the islands of the archipelago. The materials col- lected by the Harrison Williams expedition confirm this statement and fill some of the gaps in the previously known distribution of the species. No forms of macilentus were previously recorded from Indefatigable, but the expedition took a new dark variety and a paler undescribed form on that island and South Seymour. A hitherto unnamed variety was also secured on Tower Island. Pre- vious expeditions took no forms of planus on James, but members of our party captured specimens of a new variety on that island and also the hitherto unknown male and female of the var. peregrinus Emery on Chatham. The two species of Camponotus are so variable and the differences between their varieties often so feeble and illusive that much more material will have to be studied before a satisfactory conspectus of their characteristics and distribution can be given. Future collectors in the islands should therefore make a strenuous effort to obtain as many specimens as possible of these ants. In my former paper I placed C. macilentus in the subgenus Myrmamblys but Emery has recently removed it to his subgenus Pseudocolobopsis4 and I have followed him in this paper. He agrees with me in placing planus in the subgenus Myrmorhachis Forel, but I have recently given the American species of this group the name Myrmocladoecus5 and this name will have to be substituted for Myrmorhachis (Emery 1920). But all the known neotropical species of Myrmocladoecus ( bidens Mayr, bispinosus Mayr, latangu- lus Rog., mucronatus Emery, quadrilaterus Mayr, etc.) nest in plant cavities, whereas planus nests in the ground. The structure of the head and thorax, the pilosity and sculpture would justify one in placing it in Orthonotomyrmex Ahsmead, were it not that this subgenus is now restricted to Old World species. The precise position of planus must therefore be left to be determined by the future student who can make a detailed study of the various sub- 4 Le Genre “ Camponotus ” Mayr, Nouvel Essai de la Subdivision en Sousgenres. Rev. Zool. Afr. 8, 1920, p. 229-260. ‘ Professor Emery ’s Subgenera of the Genus Camponotus Mayr. Psyche 28, 1921, p. 19. 114 Zoologica: N. Y. Zoological Society [V; 10 genera of Camponotus and of their limits. For the present 1 retain the species from the Galapagos in the subgenus Myrmocladoecus. Camponotus (Pseudocolobopsis) macilentus F. Smith var. sapphirinus var. nov. (Fig. 24). Worker major. Length 6-7 mm. Head about one-fifth longer than broad, distinctly broader behind than in front, with nearly straight sides and posterior border; anterior borders of cheeks not very convex nor projecting. Clypeus subrectangular, scarcely broader in front than behind, somewhat longer than broad, its posterior half distinctly FIG. 24. CAMPONOTUS ( PSEUDOCOLOBOPSIS > MACILENTUS F. Smithvar. SAPPHIRINUS var. nov. a , head of worker major; b, thorax and petiole of same in profile; c, head of worker minor. carinate in the middle, its anterior border very feebly and broadly rounded, rather receding. Scapes reaching nearly twice the width of their tips beyond the posterior corners of the head. Thorax strongly compressed behind, the epinotum in profile not evenly rounded but made up of three subequal planes meeting at rounded obtuse angles. Petiolar node low and thick, its upper border blunt, entire and when seen from behind broadly rounded. Shining; mandibles and anterior portion of head less so than the remainder of the body, which is very finely shagreened. Pilosity much as in the other varieties, the color considerably darker. Body 1924] Wheeler: Formicidae of the Galapagos 115 brownish yellow; mandibles and anterior border of head brownish red; antennae, a large spot on the front and vertex, the whole meso- and epinotum, the tibiae, tarsi and a broad transverse band on each gastric segment, brown; the cephalic spot, apical half of the scapes and gastric bands darker, more castaneous brown. Owing to its fine transverse shagreening the upper surface of the gaster in direct sunlight has a pronounced blue iridescence, which is absent or very faint in the other varieties. The top of the head and sides of the thorax display a similar but feebler iridescence. Worker minor. Length 4. 5-5. 5 mm. Closely resembling the major worker, except in the shape of the head, which in the minima is nearly one and one-third times as long as broad, with more rounded posterior corners, straight, anteriorly somewhat converging cheeks and broader and less distinctly carinate clypeus, The scapes extend about one- third their length beyond the posterior corners of the head. Color similar to that of the worker major, but there are differences in the head, which is whitish or ivory yellow in front and diffusely brownish behind; the mandibles are brown- ish yellow, the antennae, especially the apical halves of their scapes, paler than in the major. The pronotum may be somewhat brownish posteriorly, but this is also the case in some of the larger workers. Female (dealated). Length 7-8 mm. Like the major worker, but with larger eyes, shorter clypeus and the anterior portion of the head more sharply sculptured. Petiolar node very thick and broad, its superior border entire, very blunt, straight and transverse. The pilosity and color are also similar but in some specimens the dark brown area on the front and vertex is more extensive and less sharply defined, in others restricted to the ocellar triangle. Each ocellus has a black spot at its inner border. Mesonotum with a large rectangular brown spot on the middle of its posterior three-fourths in some specimens, in others there is merely a brownish cloud. Male. Length 5-5.5 mm. Pale ivory yellow, head and gaster brownish, the latter with the posterior borders of the segments yellow; the former with the ocellar triangle dark brown, almost blackish, and each ocellus with a black spot at its inner border; the front pale brown but somewhat variable in different specimens. Wings pale yellowish hyaline, the pterostigma scarcely darker than the veins, which are pale yellow. There is a small black spot at the base of each of the anterior wings. Described from numerous specimens nesting in hollow twigs of Maytenus obovata, previously inhabited by Calotermes pacificus colonies, near sea-level at Seymour Bay, Indefatigable Island and on South Seymour Island. This variety of macilentus differs markedly from all those previously des- cribed in the deeper and more extensive infuscation of the head, thorax and gaster and the distinct iridescence of the gaster in the worker and female. Camponotus (Pseudocolobopsis) macilentus var. pervious var. nov. Worker major. Length 4.5-5 mm. Very similar to the preceding variety but smaller, more thickset, with proportionally larger head and paler, posteriorly less compressed thorax. Thorax and petiole immaculate, the head with a pale brown, sometimes very faint, spot on the vertex. Bands on the gaster much narrower, each produced anter- 116 Zoologica: N. Y. Zoological Society [V; 10 iorly as a point in the middle line. Apical half of scapes in some specimens with a black streak on its anterior and posterior border. The ground color of the body, especially of the thorax, petiole and gaster is paler than in the var. sapphirinus and there is scarcely a trace of iridescence. Worker minor. Length 4 mm. Very similar to the major worker in color; the head as in the var. sapphirinus but the body smaller. Female. Length 6-6.5 mm. Smaller than sapphirinus and colored like the major worker, but with a faint brownish cloud on the vertex and posterior portion of the mesonotum. Each ocellus has the usual black spot at its inner border. Wings 6.5 mm. long, distinctly brownish yellow, with brownish yellow veins and pterostigma. Male. Length 4.5 mm. Also smaller than the corresponding sex of sapphirinus, but the gaster is as pale as the head and thorax and the suture between the mesonotum and scutel- lum is black. The vertex is brownish, the ocelli as usual with a black spot at their inner borders. Several specimens from a single small colony taken in a Maytenus obovata twig at Seymour Bay, Indefatigable Island. The small dimensions of all four phases show that we are dealing with a distinct variety or subspecies of macilentus. Camponotus ( Pseudocolobopsis ) macilentus var. jacobensis Wheeler. Single winged female and male specimens taken by members of the expedi- tion on James Island April 5th, though somewhat darker, agree well with my topotypes collected by Dr. F. X. Williams. Camponotus (Pseudocolobopsis) macilentus var. castellanus var nov. (Fig. 25). Worker major. Length 6-7 mm. Head fully one and one-fourth times as long as broad, subrectangular, scarcely narrowed in front except at the extreme anterior end of the cheeks, the posterior border and sides straight. Cheeks in front convex and projecting on either side of the clypeus, which is subelliptical and notched behind at the indistinct frontal area. Scapes extending beyond the posterior corners of the head a distance equal to the width of their tips. Posterior portion of thorax moderately compressed, the base of the epinotum in profile convex and rounded, decidedly longer than the abrupt, concave declivity. Petiolar node smaller and thinner than in the other varieties, its superior border rather sharp, trans- verse and feebly concave in the middle. Sculpture of the mandibles, anterior and dorsal surface of the head rather sharp so that these regions are distinctly less shining than the remainder of the body. Brownish yellow; coxae and femora paler, thorax and petiole immaculate; mandibles, anterior border of clypeus and cheeks, apical half of scapes, a large spot on the front and vertex, tibiae and tarsi reddish brown. Gaster dark brown, with the posterior border and two transverse spots at the base of each segment, brownish yellow. 1924] Wheeler: Formicidae of the Galapagos 117 Worker minor. Length 4-5.5 mm. Head very similar to that of the major worker, but with sides feebly rounded, scarcely narrower in front than behind, clypeus shorter, distinctly carinate. Antennal scapes extending about one-third their length beyond the posterior border of the head. Color similar to that of the major but mesonotum, epinotum and petiole brownish. Tibiae only slightly darker than the femora, which FIG. 25. CAMPONOTUS ( PSEUDOCOLOBOPSIS ) MACILENTUS F. Smith -var. CASTELLANUS var. nov. a, head of worker major; b, thorax and petiole of same in profile; c, head of worker minor. like the coxae, anterior portion of the head and paler portions of the gaster, are whitish or ivory- yellow; knees brownish. The epinotum and petiolar node have the same shape as in the major worker. Female. Length 7-7.5 mm. Head large, like that of the major worker, not narrowed in front, but the scapes extend about twice their greatest diameter beyond the posterior corners. Color as in the worker major but the scutellum and a large rectangular spot on the posterior portion of the mesonotum are brown. Wings nearly 8 mm. long, distinctly brownish yellow, with pale brownish pterostigma and pale yellow veins. Male. Length 5.5-6 mm. Brownish yellow, the gaster a shade darker, the mandibles, anterior portion of head, pleurae, coxae and femora paler; a small black spot at the base of each fore wing and the usual black spot at the inner border of each ocellus 118 Zoologica: N. Y. Zoological Society [V; 10 very distinct. Ocellar triangle more or less brownish. Wings paler than in the female, the pterostigma not darker than the veins, which are pale yellow. •Described from numerous specimens taken from dead twigs and branches of Bursera graveolens, abandoned by colonies of Calotermes pacificus, on Tower Island. In my paper of 1919 (p. 287) I recorded a major and two minor workers of this variety, among the material taken in 1899 by the “Albatross” on the same island, but the specimens were so greasy and defective that I did not give them a name. a, head of worker major; b, thorax and petiole of same in profile; c, head of worker minor. This variety is distinct in the shape of the head in the worker and female and in that of the petiolar node of the worker. In these phases there is a faint trace of iridescence on the gaster when it is viewed in direct sunlight. The same is also true of the var. jacobensis. Camponotus ( Myrmocladoecus ) planus F. Smith var. peregrinus Emery. Female (undescribed). Length 7.5 mm. Compared with the typical planus of Charles Island, the differences are mainly in color and pubescence. The body is black, the red areas of planus 1924] Wheeler: Formicidae of the Galapagos 119 being more restricted and the red shade of the appendages deeper. On the gaster the appressed pubescence is distinctly shorter, less dense and less silky, so that this region has a different, faintly grayish luster. The pterostigma of the wings is pale yellow, like the veins. Male (undescribed). Length 5.5 mm. Also darker than the male of the typical planus, the body being jet black; the mandibles, funiculi and trochanters reddish, and the legs and antennal scapes black, slightly tinged with red. Pterostigma and veins of wings pale brown. Single specimens taken by members of the expedition on Chatham Island, April 7th, 1923. Camponotus ( Myrmocladoecus ) planus var. isabelensis Wheeler. I refer two males taken April 6th, at Tagus Cove, Albemarle Island to this variety. Camponotus ( Myrmocladoecus ) planus F. Smith? var. santacruzensis Wheeler. (Fig. 26). Specimens from several large colonies were collected at Seymour Bay, Indefatigable Island and on South Seymour Island during the latter part of April. They were nesting in the ground about the roots of trees and shrubs and the large arborescent Opuntias peculiar to the islands. The largest workers attain a length of 7 mm. which is a millimeter more than I recorded for the speci- mens collected by Dr. Williams. The body has little pilosity and the head is entirely black, not red anteriorly as in the following variety: Camponotus ( Myrmocladoecus ) planus var. indefessus Wheeler. A number of small workers and males were taken by members of the expedition April 1st, 1923 at Conway Bay, Indefatigable Island. Camponotus (Myrmocladoecus) planus F. Smith? var. sansalvadorensis var. nov. (Fig. 27). Worker minor. Length 4. 5-5. 5 mm. Head trapezoidal, slightly longer than broad, with straight anteriorly converging sides, the eyes at the posterior corners and apparently somewhat more projecting and convex than in the other varieties. Antennal scapes slender and terete at the base, even in the media (in other forms of the species broader and at least slightly flattened), extending somewhat less than half their length beyond the posterior border of the head. Thorax with the base of the epinotum distinctly concave in profile, its posterior angles somewhat dentate, much as in the var. indefessus. Pilosity as abundant as in that form but slightly shorter and paler. The appressed pubescence on the gaster is distinctly paler and sparser so that it has only a faint, silvery luster. Body subopaque and of the usual sculpture and color, only the mandibles, anterior edges of the cheeks and the appendages being red. Male. Length 4-5.3 mm. Indistinguishable from the male of the var. peregrinus Emery. Six workers and five males, taken April 5th, 1923, on James Island by some of the members of the expedition. 120 Zoologica: N. Y. Zoological Society [V; 10 The distinguishing character of this variety seems to be the terete, or cylindrical base of the antennal scapes, which are broader and distinctly more or less flattened in the other forms of the species. Probably the unknown worker major has flattened but very narrow scapes. The known distribution of the various forms of C. macilentus and planus is given in the following table, which is merely an amplification of the one pub- lished in my paper of 1919. It will be seen that while some form of macilentus is known to occur on each of the islands on the list except Chatham, planus is unknown from Hood, Bindloe and Tower. I feel confident that it really does not exist on the outlying Tower and that the same statement is probably true of Bindloe, but Hood, judging from its geographical position, should yield a distinct variety. The occurrence of a peculiar variety of macilentus may also be pre- dicted for Chatham. FIG. 27. CAMPONOTUS ( MYRMOCLADOECUS ) PLANUS F. Smith var. SANSAL- VADORENSIS var. nov. a, head of worker media; b, thorax and petiole ol same in profile. ISLANDS Macilentus, typical Charles planus, typical (? an undescribed variety) . . Chatham .... var. peregrinus var. narboroensis Narborough . var. fernandinensis var. albemarlensis ) Albemarle . . . var. isabelensis var. vulcanalis. . . ) var. duncanensis .......... Duncan var. pinzonensis var. pervicus ) Indefatigable ( var. indefessus var. sapp innus } ) South Seymour j var. santacruzensis 1924] Wheeler: Formicidae of the Galapagos 121 var. hoodensis Hood (? an undescribed variety) var. barringtonensis Barrington . . var. fidelis var. jacobensis James var. sansalvadorensis var. bindlofnsis Bindloe (? absent) var. castellanus Tower (absent) In conclusion I append a complete list of the Galapagos Formi- cidae with their distribution, so far as it has been ascertained from the materials collected by the Harrison Williams and previous expeditions. For reasons given above I have omitted Camponotus senex and have substituted Cylindromyrmex williamsi for C. striatus. As revised the list shows that forty-two different forms have so far been taken in the islands, that there are only five species, but twenty-eight varieties or subspecies, peculiar to their fauna and that some nine species are common tropicopolitan vagrants, or tramps. The list also shows that nearly twice as many ants (12) are recorded from Indefatigable as from any of the other islands. This merely means, of course, that more numerous collections have been made in that one locality. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” 122 Zoologica: N. Y. Zoological Society [V;10 Names of Species, Subspecies and Varieties Narborough Albemarle Duncan Charles ■d o o W Chatham | Barrington Indefatigable Eden Rock Sh d o a 03 m A +3 d o . GQ Daphne James Tower Bindloe Abingdon Cylindromyrmex williamsi — X X — — — — — Ponera beebei — — — — — — — — — — — — X ■ — Odontomachus haematoda subsp bauri. — — — X — , X — — — — — — — — — Pheidole williamsi — — — — — — X — — ■ j§|§ — var. seymourensis — — — — — — — — — X X - Pheidole sp. — X — — — — — — — — — — — Solenopsis saevissima — — — — — — X — — — — Solenopsis geminata Solenopsis globularia var. • — X — III — — — — — — galapagaea var. pacifica — X — — — X ■ — X — X — var. rubida — — — — X — 1 — — — — — — — Monomorium pharaonis — — — — — — — X — — — — — X Monomorium floricola — ~ Tetramorium guineense — X — Hj X — — — — — X ■ — Tetramorium simillimum — — — X — — — — ' — — — — — — Dorymyrmex pyramicus sub- sp. albemarlensis — X — — — — — ' X X X — — — — Tapinoma melanocephalum — — — X X — X — — — — X — — Prenolepis longicornis — — — X — — — — — 'W — — — — • — Prenolepis vividula var itin- erans — _ — — — — ' X . — X — — j — — — — : — var. edenensis 11 1 1 Prenolepis fulva subsp. nesiotis — — — — — — — X — — — X — ■ — — Camponotus macilentus — var. jacobensis — — — — — — V — X — jj§ — var. albemarlensis — X — — — • — — — — — — — var. vulcanalis — X — — — — — — — — — — — — var. duncanensis — — X — — — — — — — — — — — — var. narboroensis X — — var. hoodensis X var. barringtonensis — — — X — — — - ^ ■ — var. bindloensis — X — var. sapphirinus 1 II — " 1 1 (1 — X x_ — II — | — var. pervicus X var. castellanus X Camponotus planus X var. peregrinus 0 1 X var. isabelensis — X — var. indefessus — — — — — — — X — — — — — — var. santacruzensis — — — — — — X — X — — — — — var. fldelis X var. fernandinensis X var. pinzonensis X Ill var. sansalvadorensis X Number of forms on each island 2 7 2 6 3 6 2 13 2 5 2 3 6 1 1 Zoologica, Vol. V, No. 11. TRIUNGULIN LARVAE FROM THE WILLIAMS GALAPAGOS EXPEDITION TR1UNGULINS OF A MELOID BEETLE BORNE BY XYLOCOPA, WITH REMARKS ON THIS TYPE OF LARVA IN THE COLEOPTERA AND STREPSIPTERA.1 By Charles T. Brues. (Figures 28-32 incl.). Among the numerous specimens of the bee Xylocopa transitoria Perez collected by the members of the Williams Expedition to the Galapagos, there is one female from South Seymour Island, taken by William Beebe, which is of particular interest. This specimen bears attached to the posterior part of the thorax a large number of Coleopterous triungulins which undoubtedly belong to some beetle parasitic upon this large bee. The Xylocopa in question was preserved in a vial of alcohol and after its long voyage to New York and subsequent shipment to Boston still retains most of the triungulins attached to its body (Fig. 28). These now number approximately 100, in addition to some twenty-five which have become detached during transit. With the exception of two on the basal concavity of the abdomen, all are confined to the scutellum and to one side of the posterior declivity of the propodeum where they form a dense yellow mass. Each triungulin rests with the posterior end of its body directed away from the body of the bee. Examination of the individual triungulins shows that they maintain this position by closing the mandibles about a single body-hair of the bee, or by grasping several together in such a way that the branched hairs pass through the space between the closed mandibles and are pressed between these and the interior surface of the head, while the apical portion of the hair passes backward along the ventral surface of the body. Thus attached, the triungulin is further securely anchored by a peculiar modification of the mandibles which are not toothed along the inner 1 Contribution from the Entomological Laboratory of the Bussey Institution, Harvard University, No. 231. First form on press February 27, 1921+ 125 FIG. 28. XYLOCOPA TRANSI TORI A Perez Attached to the thorax is a group of triungulins of Horia maculata Swed. FIG. 29. Mandibles of a triungulin of HORIA MACULATA Swed. In microphotograph -A- the lens has been focussed at a higher level than in -B- to show the narrow' spaces between the +ee+h r.t their bases. 126 1924] Brues: Triungulin Larvae of the Galapagos 127 FIG. 30. Triungulin of HORIA MACULATA Sued. A, lateral view; B, dorsal view; C, mandibles; D, antenna; E, palpus; F, tibia and tarsus of hind leg; G, body-hair taken from propodeum of Xylocopa transitoria Perez. cutting edge as is usual among insects. They are instead provided each with a series of tubercular projections or teeth on the upper surface, that is to say on the side which faces the under surface of the head. Thus when the mandibles are closed (Fig. 30, C; Fig. 29) the projections form a transverse line, each extending backward and serving to catch the hair (Fig. 30, G) so firmly in the notches between them that the triungulin is able to retain its hold continu- ously in spite of the active movements of its conveyer. 128 Zoologica: N. Y. Zoological Society [V ; 11 As will appear later, these triungulins are undoubtedly larvae of a species of Horia, most probably H. maculata Swed. They may be described as follows: Length 1.3-1. 7 mm. Uniformly pale brownish yellow, the eyes intensely black. Head much flattened; thorax less so, and abdomen nearly cylindrical in cross section. Head gradually narrowed anteriorly, the front border acutely rounded. Eyes set just back of the middle, small, oval, appearing in specimens that have been cleared in potash as composed of two ommatidia, a larger dorsal and a smaller ventral one. Antennae three jointed, very small; basal joint short, second and third cylindrical, the third more slender and slightly the longer; apical seta long and so extremely thin apically that its tip is difficult to see even with an oil-immersion lens; fully as long as the head. Palpi longer than the antennae; basal joint short; second and third longer; tip of third joint truncate and bearing a few very minute setae. Mandibles large, concealed beneath the head when closed; acutely pointed at tip; upper edge (7. e., the surface next to the lower surface of the head) bearing four tooth-like ridges that extend back- ward. Thoracic segments longer than the abdominal ones; prothorax the longest and the metathorax shortest; all three entirely destitute of bristles. Abdomen long and tapering, composed of nine segments in addition to an apical membran- ous process (so-called anal sucker; not shown in the figure) which is sometimes extruded to nearly the length of the last segment; all the segments bear along the sides and posterior edge a number of bristly hairs. Legs moderately long, all three pairs of about the same size and form; femora somewhat thickened; tibiae more slender; tarsus (fig. 30, F) reduced to a single curved claw on each leg. The femora and tibiae are sparsely clothed with bristly hairs and each coxa bears one or two bristles. Having noted the remarkable structure of the mandibles in this species, I was led to examine other triungulins for similar modifications and find that such exist in at least one other form. A female Andrena collected by Prof. W. M. Wheeler at Colebrook, Connecticut, bears attached to the body-hairs, several triungulins quite similar to those on the Xylocopa from the Galapagos, although differing in a number of characters. This larva is probably a Meloid of the tribe Zonitini represented in this region by at least one species, Zonitis bilineata Say. According to the studies of Cros (’13; ’20) the larvae of this group and of the tribe Sitarini are distinguishable from the triungulins of other Meloidae and of the Rhipiphoridae by the presence of a pair of dorsal projections arising in the membrane between the eighth and ninth abdominal segments, formerly thought to be suckers or attachment organs, but shown by Cros to be res- piratory organs. These are present in this triungulin, which further has bi-ocellate eyes and tarsal claws with a long basal bristle. The mandibles are strikingly similar to those of the Horia larvae from 1924] Brues: Triungulin Larvae of the Galapagos 129 the Galapagos, but are less highly modified. The teeth are on the upper surface of the mandible, but are closer to the edge and actually project over the edge in optical section. This type of dentition is therefore intermediate between that with the usual dentate inner edge and the modification described above where the teeth have shifted away from the edge and have come to lie entirely on the dorsal surface. Unfortunately no Meloidae or Rhipiph.oridae are known from the Galapagos and it is impossible to determine positively the syste- matic position of the larvae. There are no records which I can find of Xylocopa bearing triungulins in other places where these bees are abundant, but as I hope to show from their structure in con- junction with published records of Meloid beetles reared from the nests of Xylocopa in other parts of the world, the presumptive evidence is very strong that the Galapagos triungulins are those of Horia maculata Swed. In general form, tarsal structure and by the presence of bi- ocellate eyes they are very similar to the larvae of Sitaris, Hornia and other Sitarini and Zonitini as described by Cros (’20), but the respiratory horns near the apex of the abdomen are entirely lacking and they are present in all the members of these tribes in the first larval stage, so far as is definitely known.2 As mentioned above the form of the mandibles is also different although easily to be derived from a type which occurs in these tribes. The triungulins of several Meloini are well known and were familiar objects to several of the earlier entomologists. They were observed by Goedart, DeGeer, Kirby and Dufour (who first called them triungulins in 1828), and as early as 1845 Newport was able to explain their presence attached to the bodies of bees on the basis of their larval habits. Later the larva of Meloe was studied by Riley (’ll), by Beauregard (’90) and notably by Cros in several recent papers. The legs terminate in three similar curved claws, of equal thickness or sometimes with the median one stouter and straight, two types very aptly termed by Cros “en fourche” and “ en trident de Neptune/’ The head bears a single eye on each side and the abdomen is furnished with long caudal bristles. Obviously the present larva does not belong to this group, although some species 2 Williams and Hungerford (’14) figure a triungulin which they suspected was that of Hornia, but this is quite likely referable to some other genus of Meloidae. 130 Zoologica: N. Y. Zoological Society [V; 11 are bee-parasites and are regularly found attached to the bees. All do not cling to the hairs, however, and one remarkable species, Meloe cavensis, actually perforates the intersegmental membrane of the bee by forcing its flat and pointed spiny head between the abdominal plates of its unhappy mount. Triungulins of a number of genera of the tribe Lyttini are known but none have been found attached to Hymenoptera as they appear to depend entirely upon their own powers of locomotion to locate their hosts. The claws are bifid or tripartite and the eyes are placed far forward, on the anterior half of the head. Our larvae could hardly be referred to this group. The early larva of the small tribe Horiini is known through a careful study of Bugnion (’09) on the larva of the Ceylonese Cissites testaceus and by supplementary details given later by Cros (’20). This larva resembles that of the Zonitini-Sitarini, but lacks the apical abdominal horns, although it is supplied with two caudal setae. The eyes are composed of a single ocellus according to Cros, but there is a suggestion of the eye being double in Bugnion ’s figure. The leg claws are bifid, consisting of a stout claw with a smaller appendage at the base. In all, the larva is very similar to the Xylocopa-parasite which I have described, and furthermore Cissites inhabits the nests of Xylocopa. Bugnion obtained his material from the galleries of the common Indian X. tenuiscapa Westw., with which it has been known to occur since 1833 when Westermann found it in Xylocopa nests where he mistook it for the primary agent excavating the wood. The main differences be- tween Cissites and the form in question from the Galapagos lies in the bi-ocellate eye of the latter, as well as the smaller bristle at the base of the tarsal claw (c/. Fig. 31, c.) and in the dention of the mandibles, which are said to be feebly denticulate on the edge in Cissites by Cros, although Bugnion failed to detect any teeth and described them as smooth. According to Gahan (’08) the genus Cissites is restricted to the Old World, and the Neotropical species which are very closely related are to be placed in Horia. Horia was found associated with Xylocopa nearly a century ago by that well known naturalist and keen observer, the Rev. Landsdown Guilding (’25) on the island of Barbados. He described the later larva of a Horia which he referred to H. maculata Swed., but which according to Champion ('92) was undoubtedly that of H. auriculata Duges. Guilding also shows one 1924] Brues : Triungulin Larvae of the Galapagos 131 of the large brightly marked beetles on the colored plate which accompanies his paper. It is now evident that Guilding was correct in his belief that the Horia is parasitic in the nests of the New World Xylocopas, just as Cissites is in those of Africa and Asia. Champion was not inclined to accept the accuracy of Guilding’s conclusions, for he says, in referring to Horia auriculata ( loc . cit. p. 372: “ Speci- mens of this species were chiefly obtained by me in the open veran- dahs of houses, and on more than one occasion I have observed the insect crawling on the wood-work in the close vicinity of the FIG. 31. Triungulin of CISSITES TESTACEUS Fabr. A, dorsal view; B, ventral view of head; C, tibia and tarsus of hind leg. (after Bugnion) nests of a large blue wasp, these nests being commonly found in such places. It is probable, therefore, that H. auriculata really preys upon this wasp, and not upon Xylocopa.” There can be little question that the triungulins which I have described belong to H. maculata or possibly to some as yet unde- scribed species, for maculata occurs from Mexico southward to Ecua- dor, while auriculata is not known south of Costa Rica. One other record of a Meloid associated with Xylocopa has come to my attention. This is an observation by Davidson (’07) and relates to the North American Nemognatha scutellaris Lee. which he reared from a cell of Xylocopa, probably X. orpifex, as well as from another bee of the genus Alcidamea. The first stage larvae of the family Rhipiphoridae are not so well known as those of the Meloidae, but those of several genera have been described with considerable care. Like some of the Mel- oids, Rhipiphorus ( Myodites auct.) is parasitic on various bees 132 Zoologica: N. Y . Zoological Society [V ; 11 (Melander & Brues ’03; Pierce ’04). The triungulin of R. soli- daginis Pierce has been figured by Silvestri (’06) whose drawing is reproduced in the accompanying figure (Fig. 32, B). It will be seen that this larva is extremely similar in nearly all details to the sup- posed Horia from Xylocopa described above. Silvestri does not state from whence his specimens came, but as he does not question the identification there is presumably no reason to doubt its accuracy. In his figure of the head three ocelli are shown on one side and two on the other, the larger number being characteristic of Rhipiphoridae FIG. 32. A, triungulin of RIIYZOSTYLOPS IN QU1 REND US Silv. Ventral view (.after Silvestri) B, triungulin of RHIPIPHORUS SOLID AGIN1S Pierce. Dorsal view (after Silvestri) and two being characteristic of certain Meloids. The mandibles do not appear to bear any teeth. Pierce’s figures (’04) are not suf- ficiently detailed to show the form of the eyes, nor of the tarsal claws in this species, but he describes the leg as “ terminated by a curved claw, almost concealed by a large semi-transparent elliptical pulvillus of twice its length, ” which is certainly very different from the tarsus in the Meloidae. The first larva of Rhipid.ius has been several times observed and R. denisi Chobaut is well figured by Chobaut (’19). It may be considered together with Rhyzostylops inquirendum Silvestri (’06). The latter is a very aberrant Rhipiphorid known 1924] Brues: Triungulin Larvae of the Galapagos 133 only from the extremely degenerate female and the first-stage larva. Silvestri considered this insect as more or less intermediate between Rhipidius and the Strepsiptera. The triungulins of these two genera are remarkably similar (fig. 32, A, Rhyzostylops) . The legs are more slender than those of the Meloidae, with long thin tibia, tipped by a pair of extremely minute claws, the eyes are multi-ocellate and the body is heavily bristled. In at least some species of Rhipidius (Murray ’70), there is a large pulvillus between the tarsal claws. The mandibles are simple, without teeth. In another Rhipiphorid, Macrosiagon ( =Emenadia ) which occurs in the nests of certain solitary wasps (Odynerus, Eumenes, etc.) there is a pulvillus also, much as in Rhipiphorus. The structure and arrangement of the claws of the legs of Meloid and Rhipiphorid triungulins is of particular interest in connection with the possible relationship of these beetles to the Strepsiptera. The triungulins of the latter are usually destitute of claws on the legs, which are tipped with flattened sucker-like pads. In some Strepsiptera, however, two pairs of legs bear the terminal sucker while the other is provided with a single claw. This is the case in Xenos nigrescens Brues as figured in a former paper ( ’05) and a recent re-examination of these triungulins leaves no doubt as to the actual dissimilarity of the legs, the hind pair each with a claw and the four anterior legs each with a terminal disc. The same arrangement occurs in an undetermined genus “ allied to Xenos’ ’ figured by Per- kins (’05) and in Stylops. Terminal pads on all three pairs of legs occur also in some Strep- siptera, at least they have been so figured in Stichotrema (Pierce ’18). Single claws on all the legs are present in several genera, e. g., Elenchus (Perkins ’05), Pentozocera ( = Bruesia Perk.) (Per- kins ’05) and in Stylops calif ornica Pierce (Pierce ’18) although the presence of discs on the four anterior legs of some other species of Stylops would make it seem probable that S. calif ornica must repre- sent another genus. Triple claws on each leg have been found by Pierce (’18) in Callipharixenos which thus approaches the typical Meloid triungulin in tarsal structure, although the claws are very delicate and appear as three slender filaments attached to a distinct elongate tarsal joint. Thus it will be seen that the Strepsipteran triungulins show a great variation in regard to the terminal armature of the legs and 134 Zoologica: N. Y. Zoological Society [V; 11 exhibit no constant character which will serve to distinguish them from those of either the Meloidae or Rhipiphoridae. There appears to be no distinguishing characteristic in the presence of long caudal setae as these while present in all Strepsiptera are as well developed in many Meloids and Rhipiphorids. Whether the difficulty of dis- tinguishing the larvae of the two groups indicates genetic relation- ship is of course not altogether clear, but the many similarities strengthen the opinion held by many entomologists that the Strep- siptera are more closely related to the Coleoptera than to any other group of insects. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” BIBLIOGRAPHY. Audouin, J. V. ’35 Observations sur la maniere de vivre des larves de Sitaris humeralis Ann. Soc. Entom. France, vol. iv, bull. p. lxxvii. Barber, H. S. ’15 Macrosiagon flavipenne in Cocoons of Bernbex spinolae. Proc. Entom. Soc. Washington, vol. xvii, pp. 187-188. Beauregard, H. ’90 Les Insectes Vesicants. Paris. 1 vol. 8vo. Brues, C. T. ’05 Notes on the Life History of the Stylopidae. Biol. Bull., vol. viii, pp. 290-295, 2 figs. Bugnion, E. ’09 Le Cissites testaceus Fabr. des Indes et de Ceylon; metamorphoses, appareil genital. Bull. Soc. Entom. Egypt, 1909, fasc. 4, pp. 182-200, 3 pis. Champion, G. C. ’92 Biologia centrali-americana, Coleoptera, vol. 4, pt. 2, p. 372. Chobaut, A. ’91 Moeurs et metamorphoses de Emenadia flabellata F. Ann. Entom. Soc. France, vol. ix, pp. 447-456, 1 fig. ’19 Description des deux sexes, de 1’oeuf et de la larve primaire d’un nouveau Rhipidius de Provence. Bull. Soc. Entom. France, 1919, p. 200-206, 2 figs. 1924] Brues : Triungulin Larvae of the Galapagos 135 Cros, Aug. ’10 Contribution a 1’etude des especes algeriennes du genre Meloe. Bull. Soc. Hist. Nat. Afr. Nord, vol. ii. ’17 Forme des ongles des larves primaires des Meloidae et valeur du terme “Triongulin.” Ann. Ent. Soc. France, vol. lxxxvi, pp. 159-164, 5 figs. (Several earlier papers of the author are listed on p. 160 of the above.) ’19 Nemognatha chrysomelina F. La larve primaire. Bull. Soc. Hist. Nat. Afrique du Nord, vol. x, pp. 55-61. ’20 Contribution a 1’etude des Rhipiphorides Algeriens. Macrosiagon tricuspidata (Emenadia bimaculata) Bull. Soc. Hist. Nat. Afrique du Nord, vol. xi, pp. 56-68, 70-75. ’20 Notes sur les larves primaires des Meloidae avec indication de larves nouvelles. Ann. Ent. Soc. France, vol. lxxxviii, pp. 261-279. Davidson, A. C. ’07 Beetles from Bee Cells. Entom. News, vol. xviii, p. 446. Dufour, L. ’28 Description d’un genre nouveau d’ insectes de 1’ ordre des Para- sites. Ann. Sci. Nat., vol. xiii, pp. 62-66, 4 figs, on pi. 9. Fabre, M. ’57 Memoire sur 1’ hypermetamorphose et les moeurs des Meloides. Ann. Sci. Nat. Zool., (4) vol. vii, pp. 299-365, 1 pi. ’58 Nouvelles observations sur 1’ hypermetamorphose et les moeurs des Meloides. Ann. Sci. Nat. Zool., (4) vol. ix, pp. 265-276. Gahan, C. J. ’08 Notes on the Coleopterous Genera Horia and Cissites and a List of the Described Species. Ann. Mag. Nat. Hist., (8) vol. ii, pp. 199-204. Guilding, Lansdowne. ’25 The Natural History of Xylocopa teredo and Horia maculata . Trans. Linn. Soc. London, vol. xiv, pp. 313-317, 1 pi. Mayet, V. ’75 Moeurs et metamorphoses du Sitaris colletis. Ann. Soc. Entom. France, (5) vol. v, pp. 65-92, 1 pi. Melander, A. L. & C. T. Brues. ’03 Guests and Parasites of the Burrowing Bee Halictus. Biol Bull.,, vol. v, pp. 1-27, 7 figs. Murray, M. ’70 Some Facts Toward a Life History of Rhipidius paradoxus. Ann. Mag. Nat. Hist., (4) vol. vi, pp. 314-328, 1 pi. Perkins, R. C. L. ’05 Leaf-hoppers and their Natural Enemies. Bull. Div. Entom. Hawaiian Sugar Planters’ Assoc., No. 1, pt. 3, pp. 90-111, 4 pis. Pierce, W. D. ’04 Some Hypermetamorphic Beetles and their Hymenopterous Hosts. Univ. Nebraska Studies, vol. iv, pp. 153-190, 2 pis. 136 Zoologica: N. Y. Zoological Society [V ; 11 ’18 Comparative Morphology of the Order Strepsiptera, together with Records and Descriptions of Insects. Proc. U. S. Nat. Mus., vol. liv, pp. 391-501, 15 pis. Riley, C. V. ’77 On the Larval Characters and Habits of the Blister-beetles be- longing to the Genera Macrobasis Lee. and Epicauta Fabr., with Remarks on other Species of the Family Meloidae. Trans. St. Louis Acad. Sci.,vol. 3, pp. 544-562. (Reprinted in First Rept. U. S. Entom. Comm., pp. 293-302 (1878) 1 pi. 5 figs. ’79 Notes on the Life History of the Blister-beetles, and on the Structure and Development of Hornia. Proc. Amer. Assoc. Adv. Sci. for 1878, vol. xxvii, pp. 284-285. SlLVESTRI, F. ’06 Descrizione di un nuovo genere di Rhipiphoridae. Redia, vol. iii, pp. 315-324, 1 pi. Westermann, B. W. ’33 Ueber die Lebensweise der Insekten in Ostindien und am Kap; Schreiben an Wiedermann. Germar Magaz. Entom., pp. 411- 427, pi. 4. Westwood, J. 0. ’39 Introduction to the Modern Classification of Insects. Vol. i, pp. 298-299. Williams, F. X. & H. B. PIungerford. ’14 Notes on Coleoptera from western Kansas. Entom. News., vol. xxv, pp. 1-9, 2 pis. Zanon, V. ’22 La larva triungulina di Meloe cavensis (Petagna) dannosa alle api in Cirenaica. L’Agric. Colon., Florence, vol. xvi, pp. 345- 354, 5 figs. ZoolOQica Vol. V, No. 12. CHILOPODS OF THE WILLIAMS GALAPAGOS EXPEDITION By Ralph V. Chamberlin (Plate VI, figs. 1-4) This expedition, initiated and financed by Mr. Harrison Williams, was sent out under the auspices of the Department of Tropical Research of the New York Zoological Society. While but few centipedes were secured by Mr. William Beebe and his associates on this expedition, these represent two of the three species pre- viously recorded from this archipelago, and, in addition, two new species, one of which typifies a new genus in the family Schendylidae. Thus the number of species now known from these islands is raised to five. Four of these species are indigenous. In a previous paper (Psyche, 1914, 21, p. 85) the writer erron- eously listed, among species from the Galapagos Islands, Cryptops navigans Chamberlin and Mecistocephalus parvus Chamberlin; but these forms were, in fact, taken on Clipperton Island, which lies fifteen hundred miles north-west of the Galapagos group. The only one of the five species not represented in the present collection is Orphnaeus brevilabiatus (Newport). This is a very common geophiloid throughout the warmer parts of both hemi- spheres. One specimen of it was taken on Hood Island by the Stanford Expedition of 1898-99. The types of new species are in the Department of Tropical Research of the New York Zoological Society. Family CRYPTOPIDAE Cryptops beebei, sp. nov. . General color yellowish. Head with sides subparallel over the middle portion. Without paired sulci. First dorsal plate with a cervical sulcus which is angularly bent back at the middle. Paired longitudinal sulci extend from caudal margin to the cervical sulcus. Each longitudinal sulcus is furcate at its anterior end. Paired sulci present on the second and subsequent tergites. Last plate with a median longitudinal depression which does not extend to the caudal angle. Prosternum not punctate or distinctly furrowed. Anterior margin convex; bearing 4+4 setae of which the ectal on each side is reduced or may First form on press April 28, 19 2^. 137 PLATE VI. NANNOPODELLUS PURPURASCENS, sp. nov. 1, anterior end, dorsal view; 2, prehensors, ventral view; 3, claw of right prehensor, ventral view, more highly enlarged; 4, caudal end, ventral view. 138 1924] Chamberlin: Chilopods of the Galapagos 139 be indicated only by the basal nodule. A little caudad of the anterior margin are in addition 2+2 bristles. Elsewhere the setae are short and sparse. Ventral plates smooth, not punctate. Last ventral plate trapeziform, the sides convex, more strongly rounded toward and about the posterior corners; caudal margin a little convex. Coxopleurae caudally subtruncate. Spiracles moderate, in part a little longitudinally elliptic. Tarsi of anterior legs uniarticulate. Third joint of last legs armed ventrally and laterally with numerous spines and spinescent setae of which the ventral ones are stouter; ventral surface with a median longitudinal space free from spines. Fourth joint with similar but fewer spines and with a similar spine-free median space. Fifth joint (tibia) with a series of four ventral teeth. The sixth joint with two ventral teeth. Ultimate joint bearing only setae. Length, 16 mm. Locality. — Tower Id. Two specimens taken 28 April, 1923. In characters of the anal legs this species resembles Cryptops navigans Chamberlin of Clipperton Id. From that species, however, it is readily dis- tinguished in having paired sulci on the first as well as on the second tergites, in having setae on the anterior margin of the prosternum, etc. The present species is named for Mr. William Beebe, leader of the expedition, by which the types were secured. Family SCOLOPENDRIDAE Scolopendra galapagoensis Bollman Proc. U. S. Nat. Mus., 1889, 12, p. 214. Scolopendra galapagoensis Chamberlin Psyche, 1914, p. 86. In the stomach of a Galapagos hawk, Buteo galapagensis, shot on South Seymour, April 19th, 1923, were found the well-preserved posterior portions of five specimens of this large centiped. Heads and anterior segments were missing. From a second hawk, killed at the same place and on the same date, fragments of two other individuals of this centiped were removed. In this case the head end of one specimen is present and entire, while of the second specimen the head proper is missing although the poison-jaws are present.* Fragments, including head of another specimen of the same species, were found in the stomach of a short-eared owl, Asio galapagensis, taken April 28th, 1923 on Tower Id. This species would seem to be the most abundant, or at least certainly the most conspicuous, chilopod of the Galapagos Is. It was previously recorded' from Hood, Chatham, Bindloe, Narborough, and Albemarle Islands. Family SCHENDYLIDAE Pectiniunguis albemarlensis Chamberlin Pectiniunguis americanus Chamberlin (nec Bollman) Ent. News, 1913, p. 122. 140 Zoologica: N. Y. Zoological Society [V; 12 Pectiniunguis albemarlensis Chamberlin Psyche, 1914, 21, p. 86; Proc. Cal. Acad. Sci., 1923, ser. 4, 13, p. 394, f. 3. This species was based upon a single female specimen with 61 pairs of legs which was taken upon Albemarle Id. by R. E. Snodgrass while a member of the Stanford Galapagos Expedition of 1898-’99. Two additional specimens were taken on Tower Id. by the Williams Expedition on the 18 and 27 April, 1923, respectively. These are likewise females. One, with sixty-one pairs of legs, has the prebasal plate exposed and the coloration normal. It is 62 mm. long. The other individual has 65 pairs of legs and has the prebasal plate covered, apparently by artificial retraction of the cephalic plate in the alcohol. The cephalic plate is somewhat differently shaped and the antennae are shorter. The geminate black dorsal stripe is absent or only vaguely indicated in the posterior region. It is 45 mm. long. Nannopodellus, gen. nov. Mandibles with a dentate lamella which is typically tripartite. Labrum pectinate at sides, truly dentate (?) at middle. Claw of the palpi of the second maxillae with margins pectinate. Sternites without pore-areas. Coxopleurae of last legs each with two homogeneous glands. Anal legs six-jointed; without claws. Genotype. — N. purpurascens sp. nov. Related to Nannophilus in general features, but different in the absence of ventral pores on the sternites. It is at present impossible to say just what relationship the present form bears to the several South American species placed by Silvestri in the genus Nannophilus, though it is quite possible they are con- generic. The species clearly referable to Nannophilus have been found in the Mediterranean region. Nannopodellus purpurascens, sp. nov. (Plate VI, figs. 1-4) The general ground color is yellowish, but the body is conspicuously 'marked throughout its length above, below and laterally with numerous purplish marks and dots which tend to form longitudinal stripes. The head shows no frontal suture. It is longer than wide in the ratio 8 : 7, and is widest at middle of its length. Caudal margin incurved. Anterior margin obtusely angular, (fig. 1.) Antennae 1.8 times as long as the cephalic plate. The ultimate article as long as, or longer than, the three preceding articles taken together, (fig. 1.) Prebasal plate exposed. Basal plate very short, its exposed area rather more than five times as wide as long. (fig. 1.) Claws of prehensors when closed not attaining front margin of head; unarmed at base but serrate along mesal side of middle portion as shown in figs. 2 and 3. Proximal joints of prehensors unarmed. Prosternum as exposed a little wider than long, without definite chitinous lines. 1924] Chamberlin : Chilopods of the Galapagos 141 The last ventral plate very wide; covering the coxal pores on each side. Pores evident only after special clearing of the specimen. (See further, fig. 4.) Anal legs of male very stout, (fig. 4.) Pairs of legs in the male, fifty-three. Length, about 16 mm. Locality. — South Seymour Id., 20 April, 1923. One male which is in poorly preserved condition. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a Volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica Vol. V,No. IS. THE COCCIDAE OF THE WILLIAMS GALAPAGOS EXPEDITION By Harold Morrison Bureau of Entomology, U. S. Department of Agriculture, Washington, D.C. (Figs. 33 to 37 incl.) This expedition was sent out by the Department of Tropical Research of the New York Zoological Society. The Coccidae re- ported on below were very kindly offered to the writer for study by their collector, Dr. William Beebe, who has also furnished all the collection data quoted for the different species represented. The writer is indebted to Miss Amalia Shoemaker for the ac- companying figures showing the structural details of the newly described species. In spite of its very obvious deficiencies, the scheme of classi- fication given in the Fernald Catalogue of the Coccidae of the World, 1903, has been followed in assigning the species to genera and sub- families. Subfamily Margarodinae Genus Margarodes Guilding Margarodes similis sp. nov. Adult Female. — (Description drawn up from fragments of one specimen only.) Nothing available regarding the normal external appearance, actual size or shape of body; the single antenna available 5-segmented, with the 4th segment incompletely divided twice, the antennae therefore probably normally 7-segmented; measurements of the segments in microns as follows: I, 71; II, 61; III, 22; IV, 100; V, 64; antennae beset with the usual stout spines and setae, not displaying any apparent peculiarities; legs characteristic for the genus, the thickened claw of the posterior pairs somewhat more slender than the same part in the legs of the nearest known relative, rileyi; mouthparts presumably wanting; thoracic spiracles very considerably larger than the abdominal, with a distinct lateral chitinous bar and four or perhaps more bilocular center, multi- locular disk pores within the external opening, and two tiny pores externally immediately adjacent to the opening; abdominal spiracles much smaller, tubular, each with a pore collar of about 4, similar, bilocular center pores, spiracles pre- sumably present in seven pairs, although only a single spiracle available for examination; derm pores nearly circular, with a large quinquilocular center surrounded by a band of smaller loculi, this in turn surrounded by a heavy First form on press April 28, 192 k. 143 144 Zoologica: N. Y . Zoological Society [V; 13 FIG. 33. MARGARODES SIMILIS, sp. nov. A, antenna, X 120; B, fore leg, X 60; C, thoracic spiracle, X 230; D, abcu.mina, spiracle, X 530; E, middle leg, X 60; F, body spines of Margarodes rileyi, for comparison X 530; G, body spines, X 530; H, derm, showing papillae, X 530; I, detail of disk pore, X 1500; J, detail of body seta, X 530; all adult female. collar; no other sorts of derm pores except those described in connection with the spiracles located; all of the derm available for examination bearing many, rather crowded, uniformly distributed, slightly convex to nearly hemispherical papillae; derm with numerous, moderately stout spines with apices varying, according to the stoutness, from acutely to almost obtusely pointed, and with numerous rather stout setae; the arrangement of these spines and setae not ascertainable from the fragments available for study; anal tube not available and no ventral cicatrices evident. Immature Female Tests. — Of moderate size, made up of more or less distinctly overlapping layers, golden to reddish-brown in color, not differing in any evident particulars from the tests of M. rileyi and M. formicarum. This spBcies has been described from a single, very fragmenta y, adult female and from a considerable number of tests of the immature stage female collected by Dr. William Beebe with the following data: “from rootlets of Maytena, Seymour Bay, Indefatigable, April 25” (Invert, no. 2263) (holotype adult female and intermediate female tests); “from roots of yellow-plumed ground plant, Eden Island, N. W. Coast of Indefatigable, April 8th” (Invert., no. 2219); “from debris under Bursera close to beach, South Seymour, north- east of Indefatigable, April 19th” (Invert, no. 2262); “Seymour Bay, Inde- fatigable,” no date, host nor number; (all of these last represented by female tests only). The type female and a number of the tests are in the U. S. National Collection of Coccidae. 1924] Morrison: Coccidae of the Galapagos 145 This species, on the basis of the material available for study, is extremely closely related to M. rileyi Giard, as this species is identified from the Florida Keys, differing positively, so far as can be observed, only in having the body spines normally pointed at apices and without the nipple-like continuation of each spine apex which appears to be characteristic of M. rileyi. Subfamily Ortheziinae Genus Orthezia Bose. Orthezia galapagoensis Kuw. This species of Orthezia is represented in the collection by two lots of material as follows: “from stems of Heliotropium parviflorum, Seymour Bay, Indefatigable, April 25” (Invert, no. 2369) and on “Bursera, Duncan, April 25” (Invert, no. 2390). Subfamily Dactylopiinae Genus Eriococcus Targ. Eriococcus papillosus sp. nov. Sac of Female. — Broad oval, moderately convex, the single sac available appearing, after immersion in alcohol, thin and fragile, white; length about 3 mm., width about 2 mm. Adult Female. — Length as mounted 2.5 mm., width 1.7 mm.; oval, tapering more or less distinctly at apices; derm clearing completely on treatment with caustic potash, except for anal lobes; antennae normally 6-segmented, the measurements in microns of the segments of those available for examination as follows: I, uncertain, due to distortion; II, 36-46; III, 104-118; IV, 22-25; V, 18-25; VI, 36-40; beak and spiracles not unusual for the genus, the former about 100 u wide and 140 m long; legs not unusual for the genus, the measure- ments of a middle leg in microns as follows: coxa about 114, trochanter 75, femur 157, tibia 132, tarsus 118, claw 36, tarsal digitules 54, claw digitules 39; tarsal digitules longer and stouter than those of claw, all distinctly knobbed at apices, tarsal claw with the usual denticle; anal lobes not unusually large nor prominent for the genus, chitinized, with the usual three dorsal spines, two ventral setae, and an apical seta, the latter about 286 m long; anal ring with the usual single complete row of pores and eight setae, the longest of these about 121^; preapical setae of anal lobes about 96m long; body bearing rather numerous large spines over the dorsum as well as at the margin, these, on the abdomen at least, arranged in more or less distinct transverse segmental bands, with spines of smaller size interspersed between the larger, all these spines stout and only slightly constricted at base, tapering almost uniformly to a slender but bluntly rounded tip; those of anal lobes proportionately distinctly more slender than the remainder; longest anal lobe spine about 40 m in length, largest marginal spine about 54 m in length, largest dorsal spines about 50 m in length, smallest dorsal spine about 22m in length; large tubular ducts with the tube of average length, the inverted cup wide and fairly deep, these ducts scattered in segmental ar- rangement over the dorsal surface, and at the margins ventrally; body also 146 Zoologica: N. Y. Zoological Society [V; 13 FIG. 34. EBIOCOCCUS PAPILLOSUS, sp. nov. A, outline of body, optical section, showing appendages, spines and larger setae, X 40; B, antenna, X 120; C, posterior leg, X 120; D, large tubular duct, X 1500; E, small tubular duct, X 1500; F, marginal spine, X 530; G, large dorsal spine, X 530; H, small dorsal spine, X 530; I, apical anal lobe spine, X 530; J, outer anal lobe spine, X 530; K, inner anal lobe spine, X 530; L, portion of derm showing papillae, X 530; M, disk pore, X 1500; all adult female. with minute symmetrical tubular ducts dorsally, but in much fewer numbers than the preceding; ventrally with smaller, quinquilocular disk pores; derm dorsally and at margins bearing numerous, but scattered, tiny papillae or tubercles rather uniformly distributed over the whole area; largest ventral abdominal seta about 50 /x. No other stages available for examination. This species has been described from two specimens mounted on slides, collected in the Galapagos Islands in company with a large number of specimens of Orthezia galapagoensis and having the following data: “From stems of Heliotropium parviflorum, Seymour Bay, Indefatigable, April 25.” (Invert, no. 2369.) The types are in the U. S. National Collection of Coccidae. Of the species of the genus Eriococcus with which the writer is familiar, 1924] Morrison: Coccidae of the Galapagos 147 FIG. 35. PHENACOCCUS PARVUS, sp. nov. A, outline of body, optical section showing position and relation of parts, X 21; B, apex of abdomen, dorsal and ventral, X 88; C, apical cerarian spine, X 1500; D, penultimate cerarian spine, X 1500; E, lateral abdominal cerarius from unexpanded adult, X 530; F, multilocular disk pore, X 1500; G, triangular pore, X 1500; H, middle leg, X 120; I, an- tenna, X 120; all adult female. this appears to approach most closely Eriococcus palmeri Ckll., described from Lower California, from which species it differs most obviously in the larger size and more elongate and more slender antennae as well as in the possession of the cuticular tubercles, these not being in evidence in E. palmeri. Genus Phenacoccus Ckll. Phenacoccus parvus sp. nov. Adult Female. — Only alcoholic specimens available for examination, ^so no description of normal external appearances can be given; length of fully dis- tended specimens as mounted on slide 1.8 to 2 mm.; width 1.2 to 1.3mm.; normal shape either uniformly oval or very slightly broader behind; derm clearing 148 Zoologica : N. Y. Zoological Society [V; 13 completely on treatment with potassium hydroxide in old specimens, but re- taining faintly suggested indistinct disks below each cerarius in undistended adult females; of the usual Pseudococcine type, antennae normally 9-segmented, the range of measurements in microns of those available for study as follows: I, about 36; II, 46-61; III, 46-53; IV, 32-39; V, 29-36; VI, 29-36; VII, 29-32; VIII, 32-36; IX, 54-61; (segments 4 and 5 incompletely separated in one an- tenna); legs not unusual, hind coxae without pores, measurement of a middle leg in microns as follows: coxa 125, trochanter 79, femur 186, tibia 190, tarsus 89, claw 29; claw with a distinctly developed denticle about one-third of length from apex, digitules slender, those of claw knobbed, those of tarsus acute at apices; beak elongate, triangular, more or less distinctly 2-segmented, length about 114 m; with the usual anterior and posterior pairs of slit-like dorsal ostioles; with eighteen pairs of cerarii along the body margin, each of these composed of two slender lanceolate spines and a few (3 to 9) triangular pores, and each underlaid, in newly emerged and undistended adult females only, by a fairly distinct, but only slightly chitinized, circular to oval disk; rarely with one or two smaller dorsal spines approximating the cerarian area so closely as to appear to be a part of the cerarius, average length of cerarian spines 11 n, spines of apical cerarii about 17 m long, correspondingly larger than the others on the body and accompanied by more triangular pores than any other cerarii; apical setae of anal lobes about 196 /x long; without ventral chitinized thickenings on anal lobes, but with several setae of various sizes ventrally just anterior to the apical setae; anal ring bearing six setae averaging about 107 n. in length and each half composed of an inner scalloped band of irregularly shaped larger pores and an outer band of nearly uniformly rounded, smaller, but less heavily chitinized, pores; dorsally, at least in the abdominal region, with transverse segmental rows of scattered triangular pores and slender lanceolate spines, the former much more abundant than the latter; with tubular ducts of moderate size surrounding the cerarii dorsally and ventrally at the body margin and extending in transverse segmental bands of scattered ducts across the ventral abdominal segments; with five transverse segmental rows of scattered large circular disk pores ventrally in the abdominal region; also ventrally with transverse seg- mental single rows of setae of different sizes; with a single, transversely oval to quadrate, cicatrix. No other stages of this species have been available for examination. This species has been described from four mounted specimens collected “on bush near shore, Tover, Apr. 28” (Invert, no. 2413). The types are in the U. S. National Collection of Coccidae. The present state of the classification of Phenacoccus and related genera is such that the writer can make no precise suggestions regarding the relation- ships of the species described above. Its structural characters appear to be quite commonplace, leaving its comparatively small size the most evident feature of the species. Pseudococcus galapagoensis sp. nov. Adult Female. — Only alcoholic specimens available for examination, so nothing regarding the external appearance can be given; length 3.5 mm., width 2 mm.; elongate oval, apices tending to be pointed; antennae normally 1924] Morrison: Coccidae of the Galapagos 149 FIG. 36. PSEUDOCOCCUS GALAPAGOENSIS, sp. nov. A, outline of body, optical section, X 21; B, posterior apex of abdomen, dorsal and ventral, X 60; C, apical cerarian spine, X 1500; D, penultimate cerarian spine, X 1500; E, multilocular disk pore, X 1500; F, small tubular duct, X 1500; G, triangular pore, X 1500; H, body seta, X 1500; all adult female. 8-segmented, lengths of segments in microns as follows: I, 64; II,. 78; III, 68; IV, 43-46; V, 46-57; VI, 39-43; VII, 39; VIII, 93-96; legs not unusual, lengths of parts of middle leg in microns as follows: coxa 196, trochanter 107, femur 268, tibia 278, tarsus 103, claw 36; claw without denticle, both tarsal and claw 150 Zoologica: N. Y. Zoological Society [V; 13 digitules slightly knobbed at apices, those of claw distinctly stouter near base, hind coxa without distinctly developed, enlarged pores, although both coxa and femur with some such enlarged pores or aerolations faintly suggested; beak elongate triangular, distinctly 2-segmented, length 171 width at base 114 n) with the usual two pairs of dorsal ostioles, the anterior pair relatively inconspicuous; with 17 pairs of cerarii, the typical arrangement of each con- sisting of two spines surrounded by a cluster of pores and, at outer margin of this, 4 setae, this arrangement varying however, as few as three pores and one seta being present; spines of posterior cerarii distinctly larger and stouter than any of the others, the size of these gradually reduced anteriorly; a tabulation of the cerarian spines showing the following count: I (anterior), 3; II, 2; III, 2-3; IV, 1-2; V, 2; VI, 2-3; VII to XVII inclusive, 2 each; pores in the anal lobe cerarii fairly closely crowded, those in the other cerarii more scattered; apical seta of anal lobes about 121 n long, ventral surface of anal lobes with a small, not very distinctly chitinized, thickening; anal ring with the usual inner and outer rows of pores and with six setae, these averaging about 130 a* in length; derm dorsally, at least in the abdominal region, with triangular pores and some small setae, and, on some abdominal segments, with an enlarged tubular duct just within each cerarius and another similar duct about half way between the center line and each of the submarginal ducts; ventrally with larger setae and large multilocular disk pores in definite transverse rows in the abdominal region, clusters of small short tubular ducts around and between the cerarii, mostly on the ventral surface, and similar ducts and tri- angular pores scattered over the mid- ventral area; the single ventral cicatrix quite large and more or less distinctly quadrate with rounded corners. This species has been described from a single mounted specimen having the following data: “from roots of a yellow-plumed ground plant, Eden Island, N.W. coast of Indefatigable, April 8” (Invert, no. 2204). The type is in the U. S. National Collection of Coccidae. Pseudococcus insularis sp. nov. Adult Female. — Only alcoholic specimens available for examination, so nothing regarding normal external appearances can be given; length as mounted on slide 3.4 mm., width 1.9 mm.; body elongate oval, tapering somewhat posteriorly; antennae of the normal Pseudococcine type, 8-segmented, measure- ments in microns of the segments available for study as follows: I, 64-68; II, 89; III, 86-89; IV, 57; V, 57-64; VI, 50; VII, 46; VIII, 107; legs of the usual Pseudococcine type, hind coxae without pores, length in microns of a middle leg as follows: coxa about 160, trochanter 125, femur 300, tibia 300, tarsus 107, claw 36; claw without denticle, claw digitules fairly distinctly knobbed at apices, tarsal digitules very slightly knobbed; beak fairly stout-conical, indistinctly 2-segmented; with the usual anterior and posterior pairs of slit-like dorsal ostioles; differing from the normal Pseudococcus type in the possession of only fifteen pairs of cerarii, each of these composed of triangular pores, accessory setae and spines, the latter running as follows in the cerarii examined: I, 3; II, 3; III, 3; IV, 2; V, 2; VI, 2-3; VII-XV, 2 each; number of accessory setae varying from 1 to 4 in the anterior and lateral cerarii, increasing to 5 in the 1924] Morrison: Coccidae of the Galapagos 151 A, outline of body, optical section, X 21; B, apex of abdomen, dorsal and ventral, X 60; C, multilocular disk pore, X 1500; D, triangular pore, X 1500; E, body seta, X 1500; all adult female. anal lobe cerarii; spines of anal lobe cerarii conspicuously larger than any of the others, those of the preapical cerarii intermediate in size, the remaining cerarian spines much smaller and more slender; triangular pores of apical cerarii closely crowded to form a dense oval cluster surrounding the spines, those of the re- maining cerarii distinctly but not widely separated from one another; apical setae of anal lobes broken, but probably, from diameter of socket, longer than 152 Zoologica : N. Y. Zoological Society [V ; 13 those of anal ring; anal ring setae about 160 ^ long; no ventral chitinous thicken- ings on the anal lobes; dorsally, at least in the abdominal region, with transverse segmental rows of widely scattered slender setae and much more numerous but scattered segmental bands of triangular pores, also, at least on some segments, with one or two median and one or two submarginal dorsal enlarged tubular ducts; ventrally, at least in the abdominal region, with seven transverse rows or clusters of large circular disk pores, with indistinct clusters of widely separated small tubular ducts beneath each cerarius and with more or less distinctly developed, transverse rows of widely scattered triangular pores, smaller tubular ducts and slender setae; anal ring rather narrow, with six setae and the usual inner and a single outer row of pores; ventral cicatrix large, more or less distinctly quadrate, with rounded corners, somewhat constricted transversely about the middle. This species has been described from a single mounted specimen collected “under stone near brackish water pool, South Seymour, Apr. 20“ (Invert, no. 2272). The holotype is in the U. S. National Collection of Coccidae. The insect above described rather closely resembles the preceding species, differing, however, in presumably important characteristics, in that it has only 15 pairs of definitely developed cerarii and has no visible ventral chitinized thickening on the anal lobes. There are also apparent differences in the relative and actual numbers of the various types of pores present but the constancy of such differences has not thus far been established. Subfamily Coccinae Ceroplastes sp. Two lots of specimens of a species of Ceroplastes were included with the material examined. The outer waxy covering of a few of the specimens was in fairly good condition, but only fragmentary portions of the female body could be obtained for study purposes. While the species does not appear to agree with any of the described forms with which the writer is familiar and is quite probably undescribed, the material available does not permit the preparation of an adequate and satisfactory description. The following data accompanied the two lots of material: “from bag of earth and leaves taken from under Maytena bushes on Eden Island, N.W. of Indefatigable, Apr. 8, 1923“ (Invert, no. 2183) and “Under lava, Seymour Bay, Indefatigable, April 25“ (Invert, no. 2552). This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica Vol. V,No. Ilf. BRACHYURAN CRABS COLLECTED BY THE WILLIAMS GALAPAGOS EXPEDITION, 1923 By Mary J. Rathbun Division of Marine Invertebrates, U. S. National Museum, Washington, D. C. (Plate VIL Fig. 38) The collection numbers 23 species, 12 of these being from the Galapagos and the remainder from the Atlantic side of Panama and the vicinity of Key West. There are several interesting occurrences. Two very small crabs are referable to Xanthias insculpta, a species not reported since the type from Lower California was described by Stimpson. The author had in hand but one individual, which is no longer extant, having probably been destroyed with extensive collections by the Chicago fire of 1871. The grey box crab, Calappa convexa, had not up to this time been taken at the Galapagos. Its occurrence there was to be ex- pected, as the crustacean fauna of the archipelago is in great part identical with that of the adjacent mainland. Colon is a new locality for the seemingly rare fiddler crab, Uca heterochelos, although this is not an extension of its range. It is a satisfaction to report a new form of megalops, referable probably to the Galapagos sand crab, thus adding another bit of knowledge to that little known subject, the development of the crab from the egg to the mature crab form. Mithrax bellii Gerstaecker. (Moss-back Crab). Mithrax ursus Bell, Proc. Zool. Soc. London , vol. 3, 1835 (1836), p. 171 (not Cancer ursus Herbst, 1788); Trans. Zool. Soc. London, vol. 2, 1836, p. 52, pi. 10, Jigs. 2, 2c, 2d, 2e and 3. Mithrax bellii Gerstaecker, Arch. f. Naturg., vol. 22, part 1, 1856, p. 112. Eden Island, Galapagos Islands, in rock pools, April 6; 1 young male, with carapace 11.7 mm. long, 10.3 wide. The young of this species are almost con- cealed by a dense mosslike covering of setae, which largely disappears with age; the carapace also grows broader until its breadth exceeds the length. The young have only five marginal spines. Range. — Galapagos Islands; Chile. First form on press April 28, 192Jf. 153 PLATE VII. MEGALOPS OF (?) OCYPODE GAUDICHAUDII . From surface of a puffer in a pool at Eden Island. Carapace 4 6 mm. long. 1, front view; 2, dorsal view; 3, ventral view. Drawn by C. J. Fish. 154 1924] Rathbun: Galapagos Brachyuran Crabs 155 Ocypode albicans Bose. (Ghost Crab or Sand Crab). Ocypoda albicans Bose, Hist. Nat. Crust., vol. 1, an X (1801-1802), p. 196, not pi. 4, fig. 1. Ocypode arenarius Say, Journ. Acad. Nat. Sci. Philadelphia, vol. 1, 1817, p. 69. Ocypode albicans Rathbun, Bull. 97, U. S. Nat. Mus., 1918, p. 367, pis. 127 and 128. Colon, Panama; 2 males (1 young). This is the common sand crab of, the eastern coast of America, ranging from Block Island to Brazil. Ocypode gaudichaudii Milne Edwards and Lucas. Ocypode gaudichaudii Milne Edwards and Lucas, d’Orbigny's Voyage dans VAmer. Merid., vol. 6, 1843, Crust., p. 26: vol. 9, atlas, 1847, p'l. 11, figs. Jt-kb. (Plate VII, figs. 1-3). A small crab was taken from the surface of a puffer, Spheroides annulatus, in a pool at Eden Island. It proved to be a megalops, or one of the later develop- mental stages of a crab; it is similar to a known megalops of Ocypode albicans , and for that reason I have ventured to give it the name of the only sand crab occurring at the Galapagos, viz.: O. gaudichaudii. In the adult of this species the eyes are remarkable in having a slender style projecting from them; this style may be as long as the eye and its stalk. The carapace of the megalops is 4.6 mm. long, 4 mm. wide. Its sides are high and are crossed obliquely by three furrows into which as many ambulatories may fit; between the first (or anterior) and the second groove, there is a promi- nent, rectangular, hepatic tubercle; on the branchial region on the posterior margin of the second groove there is a sharp oblique ridge. The mesogastric and cardiac regions are each set off by deep grooves; the mesogastric is partially divided into three parts. The front is deeply cut into three narrow, deflexed lobes, of which the lateral are only half as long as the median lobe. A deep median groove extends forward from the gastric region. The body is covered with pigment spots which are larger anteriorly and diminish in size and number posteriorly, being very few on the sixth abdominal somite and absent from the telson. The raised portions of the carapace are covered with a short pubescence, while a transverse line of hairs crosses the branchial and anterior cardiac regions. The posterior border of the first six abdominal somites is fringed with short hair, and of the pleopods with long hair. The ambulatories are sparingly dotted with fine pigment spots which thin out distally, being absent from the dactyls and upper half of the propodites. Five spines below each dactyl, the second spine from the tip being the longest. No hairs between bases of second and third ambulatories. Uca galapagensis Rathbun. (Broad-fronted Fiddler Crab). Uca galapagensis Rathbun, Proc. Washington Acad. Sci:, vol. 4, 1902, p. 275, pi. 12, figs. 1 and 2; Bull. 97, U. S. Nat. Mus., 1918, p. 403, pi. 142, text- fig. 167. James Island, common about salt ponds, April 4; 3 males, 1 female, all very small. i 156 Zoologica: N. Y. Zoological Society [V ; 14 South Seymour Island, April 21, 1 male from a salt pool, 1 female from the beach. Indefatigable Island, April 25; 1 male. Eden Island; 1 male. This is the larger of the two fiddlers that inhabit the Galapagos. Uca heterochelos (Lamarck). (Narrow-fronted Fiddler Crab). Ocypoda heterochelos Lamarck, Syst. Anim. sans Vert., 1801, p. 150. Gelasimus heterocheles, Kingsley, Proc. Acad. Nat. Sci. Philadelphia, 1880, p. 137, pi. 9, fig. 2 (part). U ca heterochelos Rathbun, Bull. 97, U. S. Nat. Mus., 1918. p. 381, pi. 131, figs. 1 and 2. Colon; 1 male, half grown. This is one of the groups of fiddlers in which the front between the eyes is very narrow, tongue-like. While the species ranges from the Bahamas to Brazil, it has not before been noted from Panama. Cardisoma guanhumi Latreille. (Large Land Crab). Cardisoma guanhumi Latreille, Encyc. Meth., Hist. Nat., Entom., vol. 10, 1825, p. 685. Rathbun, Bull. 97, U. S. Nat. Mus., 1918, p. 341, pis. 106 and 107, text-fig. 155. Colon; 1 male. Found in great numbers, all having the same light blue color. Widely distributed on the Atlantic coast of America, from Bahamas to Brazil. Gecarcinus lateralis (Freminville). (Small Land Crab). Ocypoda lateralis Freminville, Ann. Sci. Nat., ser. 2, Zool., vol. 3, 1835, p 224. Gecarcinus lateralis Guerin, Icon. Regne Anim., pi. 5. fig .1. Rathbun, Bull. 97, U. S. Nat. Mus., 1918, p. 355, pis. 119 and 120, text-fig. 161. Colon; 1 female. This species is considerably smaller than the Cardisoma and is always of a deeper, richer color. It ranges on the Atlantic coast from the Bahamas to Guiana. Grapsus grapsus (Linnaeus). (Rock Crab). Cancer grapsus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 630. Grapsus grapsus Ives, Proc. Acad. Nat. Sci. Philadelphia, 1891, p. 190, Rathbun, Bull. 97, U.S. Nat. Mus., 1918, p. 227, pis. 53 and 54, text-fig. 135. Eden Island, off Indefatigable Island; 1 male, 1 female. Indefatigable Island; 1 young female. Common in the tropics on rocky shores on both sides of the American continent. Geograpsus lividus (Milne Edwards). (Small Rock Crab). Grapsus lividus Milne Edwards, Hist. Nat. Crust., vol. 2, 1837, p. 85. Geograpsus lividus Stimpson, Ann. Lyc. Nat. Hist. New York, vol. 7, I860, p. 230. Rathbun, Bull. 97, U. S. Nat. Mus., 1918, p. 232, pi. 55. Tower Island, Galapagos; 1 young. A smaller, and duller-colored form than the preceding; it has acutely pointed fingers instead of spoon-shaped ones. Found on both sides of the continent. i 1924] Rathbun: Galapagos Brachyuran Crabs 157 Goniopsis cruentata (Latreille). (Mangrove Crab). Grapsus cruentatus Latreille, Hist. Nat. Crust., vol. 6, 1803, p. 70. Goniopsis cruentatus Rathbun, Bull. U. S. Fish. Comm., vol. 20, for 1900, pt. 2 (1901), p. 15, pi. 1 {colored)-, Bull. 97, U. S. Nat. Mus., 1918, p. 237, pi. 57. text-fig. 136. Colon, Panama; 1 young specimen. This crab can be told by its brilliant coloring in connection with the flashing white outer surface of its palms. Planes minutus (Linnaeus). (Sargassum Crab). Cancer minutus Linnaeus. Syst. Nat., ed. 10, vol. 1, 1758, p. 625. Planes minutus White, List Crust. Brit. Mus., 1847, p. 42. Off eastern Cuba, in Sargassum, 13 specimens from four different stations. For variations in this widely distributed species, see Verrill, Trans. Connecticut Acad. Sci., vol. 13, 1908, p. 325, pi. 13. FIG. 38. XANTHIAS INSCULPTA STIMPSON. Immature male, carapace 4 mm. wide. Off Eden Island. Xanihias insculpta (Stimpson). (Sculptured Crab). Xanthodes insculpta Stimpson, Ann. Lyc. Nat. Hist. N. Y., vol. 10, 1871, p. 105. Xanthias insculptus Rathbun, Bull. Labor. Nat. Hist. State Univ. Iowa, vol. 4, 1898, p. 271. Off Eden Island, 5 fathoms; 1 immature male, 1 young. This species was known hitherto only from the type from Cape St. Lucas. The larger of the Galapagos specimens is about the same size as the type. The species probably grows much larger. The gastric and frontal regions are deeply lobulated, the antero-lateral regions and also the dorsal aspect of the wrist and palm are lumpy. There are four equal, small, but well-marked teeth on the antero-lateral margin of the carapace behind the orbital angle, and a 158 Zoologica: N. Y. Zoological Society [V ; 14 small tooth at the outer end of each of the frontal lobes. There is a longitudinal ridge through the middle of the palm and above it a few obsolescent transverse ranges of minute granules. The dark color of the immovable finger is con- tinued back on the palm for a ways. Extreme length of carapace 3 mm., width 4 mm. Xanthodius lobatus (A. Milne Edwards). (Oblong Crab). Leptodius lobatus A. Milne Edwards, Crust. Reg. Mex., 1880, p. 271, pi. 49, fig. 4. Eden Island, rock pools; 1 male. Inhabits Clarion Island and Chile as well as the Galapagos. Menippe nodifrons Stimpson. (Stone Crab). Menippe nodifrons Stimpson, Ann. Lyc. Nat. Hist. N. Y , vol. 7 1859 p. 53 Rathbun, Bull. U. S. Fish Comm., vol. 20, for 1900, pt. 2 (1901), p. 37. A young specimen of this species was taken from the dock piles at Colon Harbor. Ozius verreauxii Saussure. Ozius verreauxii Saussure, Rev. et Mag. de Zool., ser. 2. vol. 5, 1853, p. 359. pi. 12, fig. 1. Indefatigable Island, April 25; 1 male. Is found also on the mainland from Lower California to Ecuador. Eriphia granulosa A. Milne Edwards. Eriphia granulosa A. Milne Edwards, Crust. Reg. Mex., 1880, p. 339, pi. 56. fig. 2. Eden Island, rock pools, April 6; 1 young. This species has been recorded from Chile as well as the Galapagos Islands. Eriphides hispida (Stimpson). (Red-Bristle Crab). Eriphia hispida Stimpson, Ann. Lyc. Nat. Hist. N. Y., vol. 7, 1860, p. 218. Pseuderiphia hispida A. Milne Edwards, Crust . Reg. Mex., 1880, p. 340, pi. 56, fig. 1. Eden Island; 1 female. Known from Central America, Panama and the Galapagos. Portunus sayi (Gibbes). (Sargassum Crab). Lupa sayi Gibbes, Proc. Amer. Assoc. Adv. Sci., vol. 3, 1850, p. 178 [141. Neptunus sayi A. Milne Edwards, Arch. Mus. Hist. Nat., Paris, vol. 10, 1861, p. 317, pi. 29, fig. 2. This is the common pelagic swimming crab of the Atlantic; 5 specimens were taken at 3 stations off eastern Cuba, in Sargassum. Callinedes danae Smith. (Brown Swimming Crab). Callinectes danae Smith, Trans. Conn. Acad. Sci., vol. 2, 1869, p 7. Rath- bun, Proc. U. S. Nat. Mus., vol. 18. 1895, p. 357, pi. 16; 26, fig. 4: 25, fig. 3; 26, fig. 3; 27, fig. 3. 1924] Rathbun : Galapagos Brachyuran Crabs 159 Colon; 1 male. Extends from Florida to Brazil. Differs from the common edible or “blue” crab of our coast, by its dull color, four teeth instead of two on its front margin between the antennae, by the greater width of the circumscribed area in the middle of the carapace, and by other less evident characters. Cronius ruber (Lamarck). (Red Swimming Crab). Portunus ruber Lamarck, Hist. Nat. Anim. sans Vert., vol. 5, 1818, p. 260. Cronius ruber Stimpson, Ann. Lyc. Nat. Hist. N. Y., vol. 7, 1860, p. 225. Achelous ruber A. Milne Edwards, Arch. Mus. Hist. Nat., Paris, vol. 10, 1861, p. 345, pi. 23, fig. 1. Off Eden Island, 5 fathoms, April 1; 1 young specimen. This form appears to be indistinguishable on opposite sides of the con- tinent. It ranges from South Carolina to Brazil and from Lower California to Ecuador. Cronius tumidulus (Stimpson). (Small Swimming Crab). Achelous tumidulus StMpson, Bull. Mus. Comp. Zool., vol. 2, 1871, p. 149. Cronius bispinosus Miers, Challenger Bept., Zool., vol. 17, 1886, p. 188, pi. 15, fig. 2. Off eastern Cuba, in Sargassum; 4 young specimens. This is not one of the two so-called “Sargassum crabs,” but a much more uncommon swimming- crab than Portunus sayi. C. tumidu'.us has been found sparingly from the Bahamas to Bahia. 4 Calappa convexa Saussure. (Grey Box Crab). Calappa convexa Saussure, Rev. et Mag. de Zool., ser. 2, vol. 4, 1853, p. 362, pi. 13, fig. 3. Eden Island; 1 male. This is known from Cape St. Lucas to Ecuador. Calappa flammea (Herbst), (Purple-striped Box Crab). Cancer flammeus Herbst, Natur. Krabben u. Krebse, vol. 2, 1794, p. 161, pi . 40, fig. 2; vol. 3, part 3, 1803, p. 19. Calappa flammea Bose, Hist. Nat. Crust., vol. 1, 1802, p. 185. Rathbun, Bull. U. S: Fish Comm., vol. 20, for 1900, pt. 2 (1901), p. 84, pi. 2. Key West; 1 male. This is the common box crab of our Atlantic coast, from North Carolina to Brazil. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica Vol. V, No. 15 THE MACRURA AND ANOMURA COLLECTED BY THE WILLIAMS GALAPAGOS EXPEDITION, 1923 By Waldo L. Schmitt Curator of Marine Invertebrates, U. S. National Museum, Washington, D. C. (Figs, 39-41 incl.) This collection, though not large in point of numbers, is of great interest in its extension of our knowledge of the carcinological fauna of the Galapagos Islands. In the crustacean collections made by the Hopkins Stanford Galapagos Expedition of 1898-99, and reported on by Dr. Mary J. Rathbun1 there were five macrurous forms not represented in the present one. On the other hand it gives us five new records for the Islands, two of which, Hippolyte williamsi and Lysmata galapagensis are apparently undescribed species, in addition to two already known from the Galapagos, but not returned by the Hopkins Expedition. En route three well- known species of Sargas^um shrimps were taken at several stations in West Indian waters, and a familiar fresh- water crayfish, or rather shrimp, from the Canal Zone. Penaeopsis kishinouyei (Rathbun). Parapenacus kishinouyei Rathbun, Proc. Washington , Acad. Sci., vol. 4, 1902, p. 288, pi. 12, figs. 13-15. Penaeopsis kishinouyei de Man, “ Siboga ” Exped., Monog. 39 a, Decapoda, pt. 1, Penaeidae, 1911, pp. 8, 55. Eden, off Indefatigable Island; dredged in five fathoms, April 1; 2 im- mature females. Exopodites are present on all the legs, and epipodites on all but the last two pairs. The meri of the fifth legs of the males of the type lot are not notched, and the telson bears three pairs of lateral marginal spines. The Eden specimens have respectively nine, and eight rostral teeth, counting the gastric one; the antero-lateral angle of the carapace is spinous. This species is known only from the Galapagos Islands, “the types were taken at Tagus Cove, on the reef north of Tagus Hill, Albemarle Island,” while four other specimens collected about the same time were “taken in 2 fathoms in Tagus Cove.” 1 Proc. Washington Acad. Sci., vol. 4, 1902, pp. 275-292, pi. 12, text-figs. 1-4. First form on press April 28, 192'f. 161 162 Zoologica: N. Y. Zoological Society [V; 15 Crangon bouvieri var. chilensis (Coutiere). Alpheus bouvieri var. chilensis Coutiere, Lenz, Zool. Jahrb., Suppl. vol. 5, 1902, p. 732. Eden; rock pools, April 6; 1 specimen. This species hitherto was known only from the three specimens of the type lot from Calbuco, Chile. Synalpheus nobilii Coutiere. Synalpheus nobilii Coutiere, Proc. U. S. Nat. Mus., vol. 36, 1909, p. 40, text- fig. 22. P Synalpheus neptunus Rathbun, Proc. Washington Acad . Sci., vol. 4, 1902, p. 289. a, second leg of right side; b, third leg; c, dactyl of same, more enlarged. Eden, rock pools; April 6; 1 specimen. The large hand of this specimen has much the appearance of that of S. sanlucasi Coutiere (op. cit., p. 41) with which Miss Rathbun had later iden- tified her material from Albemarle Island. I have been unable to examine those specimens, as they are still in the hands of Prof. Coutiere, who is mono- graphing the National Museum Alpheids. 1924] Schmitt: Galapagos Macrura and Anomura 163 The proportions of the second legs of the specimen before me definitely- set it apart from S. sanlucasi; and the antennules, antennae, spines, scales, and rostrum have the slender form as in S. fritzmulleri and S. nobilii. On the basis of the stouter third legs it is identifie4 with the latter rather than the former species. The only other known specimen of the species is the type in the Paris Museum, from St. Helena, Ecuador. Hippolyte williamsi sp. nov.? Type Locality. — Eden; rock pools, April 6; 1 9 with carapace and rostrum together, but slightly more than 3.5 mm. in length. Description. — Rostrum reaching four sevenths the length of the antennal scale, exceeding the antennular peduncle by about the length of the first two segments of the thicker flagellum, equalling about three fourths the length of the carapace; above the rostrum has two subequal teeth, the anterior of which is situated just behind the middle of the rostrum, and the first at about the distal third of the interval between the second tooth and the orbital margin; below there are three teeth of which the first is situated just before the distal margin of the first segment of the antennular peduncle, being about as far in advance of the second dorsal as that tooth is in front of the first dorsal, the second ventral is not quite half the distance between the first and third in advance of the former, the last tooth is immediately behind the acute tip, giving it a bifid appearance; on the dorsal margin a little in advance of the first ventral tooth, there is an inconspicuous tiny notch or tooth which might possibly be con- sidered as a third dorsal. The last two joints of the antennular peduncle are subequal; taken together they are shorter than the first joint; antennal peduncle reaching to or just beyond the distal margin of the second antennular segment. Supra-orbital and branchiostegal spines well developed. Distally, the outer, lateral face of the merus of the third legs is armed with three conspicuous spines, increasing in size from behind forwards, the largest, the anterior spine, is placed lower than the others; fourth leg with two such rneral spines, of which the anterior is almost ventral; fifth leg with merus unarmed; carpal joints of last three pairs with a spine on outer face at about one third the length of the joint from the posterior margin; propodus of third leg short and stout, less than three times as long as the dactyl; the dactyls of the last three pairs of legs are quite powerful, stout and well spined, spines terminally quite strong, and set close together; posteriorly, the dactyls are produced, forming a peculiar, unarmed “heel.” The third legs fall short of the end of the antennal scale, though they exceed its spine by about the length of the dactyl, the second legs and third maxillipeds reach about as far forward as the distal margin of the first joint of the antennular peduncle. Fifth abdominal somite about two thirds the length of the sixth, sixth nearly as long as the telson which is slightly shorter than the inner branches of the uropods. 2 Named in honor of Harrison Williams, Patron and Curator of Ichthyology of the Department of Tropical Research. New York Zoological Society, whose interest and liberality rendered possible the collection of the material upon which this report is based. 164 Zoological N. Y. Zoological Society [V; 15 FIG. 40. HIPPOLYTE WILLI AMSI, sp. nov. a, lateral view of carapace; b, antennule; c, antennal scale; d, third maxilliped; e, first leg: f, second leg; g, third leg; h, dactyl of same, more enlarged; i, telson. Remarks. — Our species seems to be quite distinct from the other known Hippolytids that may have a similar rostral formula, three teeth below and two or three above. Hippolyte pleuracantha (Stimpson)3 has two to three teeth above and three below, but the legs are much more slender and the dactyls have a different shape, being long and narrow instead of short and broad, they 3 Virbius pleur acanthus Stimpson, Ann. Lyc. Nat. Hist. N. Y., vol. 10, 1871, p. 127. Stimpson described this species as having but one tooth on the inferior margin of the rostrum, as a matter of fact, it has quite uniformly three teeth below. 1924] Schmitt: Galapagos Macrura and Anomura 165 have slender spines on the ventral margin and lack the “heel” found in H. wil- liamsi; the propodus of the third leg of.H. pleuracantha is about six and one third times as long as its greatest width; in H. williamsi it is about four and one half times as long as wide, and the dactyl is armed with short stout spines. In H. obliqui-manus Dana4 from “Rio Janeiro,” the dactyl and propodus of the third leg are proportioned as in H. pleuracantha; the rostrum has three teeth below if we count the ventral tooth of the bifid tip as figured in with the two described as constituting the armature of the lower margin; above there are four teeth; the rostrum is half again as long as the antennular peduncle. In H. pleuracantha, though the rostrum has been generally described as “about half as long as the carapace,” “scarcely more than half as long as the acicle,” and “reaching to extremity of antennular peduncle,” in about half of the specimens I have examined, it is about as long as the scale, exceeding the an- tennular peduncle by about one half the length of the peduncle. Other than in the possession of a fourth tooth on the upper margin of the rostrum, I can detect but one other feature in Dana’s figure of obliqui-manus, distinguishing it from H. pleuracantha. What might have been intended to represent the bran- chiostegal spine in the former species is situated on or at the anterior margin of the carapace, while in the latter, the tip of that spine is removed by distance at least half, and usually nearly the length of the spine from the anterior margin. H. williamsi has the branchiostegal spine placed so close to the anterior margin of the carapace that in part it extends beyond the margin. Heller’s Mediter- ranean H. gracilis ,5 has two to three teeth beneath, and three or four on the upper margin, but of the dorsal teeth, two were described as being “behind the eyes,” at least one is on the carapace and the second about over the orbital margin, more or less; the branchiostegal spine is figured as being behind the margin, and the rostrum is about half again as long as the antennular peduncle. Hippolyte calif or niensis Holmes,6 though it may have as few as three teeth below and the same number above, need not be considered here, because of its vastly different form; the tip of the rostrum is usually trifid, the second joint of the antennular peduncle is two to three times as long as the third and more than twice as long as wide, and the sixth abdominal somite is twice as long as the fifth and longer than the telson. Lysmata galapagensis sp. nov. Type Locality. — Northeast of Eden; seven fathoms, dredged, April 6; 56 specimens. The carapace and rostrum of the figured female together are about 6 mm. long. Description. — Rostrum about four ninths the length of the carapace, as long as, or a little longer than the eyestalks, and reaching the distal margin of the first segment of the antennular peduncle; rostral crest is continued backward onto the anterior third of the carapace; above it is usually armed with five or six. more rarely, seven teeth, of which one, less often two are on the carapace; as in the figured specimen there is always a decided hump between the first and 4 Crust. U. S. Expl. Exped., pt. 1, 1852, p. 564; atlas, 1855, pi. 36, fig. 3a-/. 5 Virbius gracilis Heller, Sitzb. Wiener Akad. Wissen., vol. 45, 1862, p. 397, pi. 1, figs. 19, 20. 6 Proc. Calif. Acad. Sci., ser. 2, vol. 4, 1895, p. 576, text-figs. 21-26. 166 Zoologica: N. Y. Zoological Society [V; 15 a, lateral view of carapace; b, antennule; c, antennal scale; d, third maxilliped; e, first leg: f, second Jeg; g, third leg; h, dactyl of third leg of another specimen; i, dactyl of third speci- men; j, telson. second teeth from which arises a conspicuous long hair and maybe one or two smaller ones; similarly hairs occur at about the mid-points of the intervals, between the other teeth and the last tooth and the rostral tip as well, but the humps marking their insertion are much less pronounced; the interval between the first and second teeth is as great as between the second and fourth; below 1924] Schmitt: Galapagos Macrura and Anomura 167 there is sometimes, but not always, a single tiny tooth, close behind the tip of the rostrum, giving it a bifid appearance; posterior dorsal region of carapace with a scattering of short hairs. The outer antennular flagella are biramus; the free portion of the shorter ramus is composed of three, four or five articles, the fused portion of from six to nine; the second and third articles of the peduncle are subequal and taken together shorter than the basal article; the blade of the antennal scale is broadly rounded and slightly exceeds the spine; mandible without palp or incisor process; third maxillipeds with exopodite, and reaching by nearly half their length beyond the antennal scale; the first legs fall short of antennal scale, the second legs exceed the third maxillipeds and reach about as far forward as the third, exceeding the antennal scale by two thirds of their carpal joints, the third legs extend beyond the scale by the length of the propodus; the carpus of the second legs appears to be divided rather uniformly into seventeen articles, the merus also is annulated, as well as the distal portion of the ischium. The fifth abdominal somite is not quite three fourths the length of the sixth, and the sixth slightly more than three fourths the length of the telson, the telson is shorter than the uropods. Remarks. — Kemp has shown in his notes on the “Hippolytidae” of the Indian Museum7 that Hippolysmata moorei Rathbun8 and H. intermedia Kingsley9 are more properly placed in the genus Lysmata on the basis of the bifurcate, outer antennular flagellum. Kemp remarks (loc. cit.): “The only difference between this genus and Risso’s Lysmata is that in the latter the outer antennular flagellum is split and is composed of two unequal rami which are fused basally. In Hippolysmata the flagellum is simple. The character does not seem a very important one, but in my experience is reliable; it is, however, not improbable that further investigation will reveal such a degree of gradation that two distinct genera can no longer be recognized, and in this case all the species must take rank under Lysmata .” By the same token Hippolysmata acicula and paucidens Rathbun10 from Hawaii also must be considered as belonging to Lysmata. In rostral formula and number of carpal articles, our species seems not unlike Miss Rathbun’s Lysmata moorei from Porto Rico. The rostrum of the latter, however, is always longer than the eyes, exceeding them by about one half the length of its free portion, reaching at least to the middle of the second segment of the antennular peduncle; usually it is armed with two teeth below, and above with not more than five, the shorter branch of the bifurcate anten- nular flagellum is composed of twelve free segments and seven to nine fused ones, the free portion being the longer. Our species has the free portion the shorter or at most subequal with the fused portion. Further, in L. moorei the antennal scale is more truncate, the fifth abdominal somite is two thirds the length of the sixth, and the sixth four sevenths the length of the telson. Hippolysmata porteri Rathbun11 is a true Hippolysmata. 7 Rec. Indian Mus., vol. 10, pt. 2, no. 4, 1914, p. 112. 8 Bull. U. S. Fish. Comm., vol. 20, pt. 2, 1900 (1901), p. 115, text-fig. 23. 9Proc. Acad. Nat. Sci. Phila., vol. 30, 1878, p. (2); Bull. Essex Inst., vol. 14, 1882, pi. 28, pi. 1, fig. 4. io Bull. U. S. Fish Comm., vol. 23, pt. 3, 1903 (1906), pp. 912, 913, pi. 34, figs. 4, 6. 1] Revista Chilena Hist. Nat., vol. 11, 1907, p. 49, pi. 3, fig. 4. 168 Zoologica: N. Y. Zoological Society [V; 15 Latr eutes fucorum (Fabricius). (Gulf-weed or Sargassum Shrimp). Palaemon fucorum Fabricius, Suppl. Entom. Syst., 1798, p. 404. Latreutes ensiferus Stimpson, Proc. Acad. Nat. Sci. Phila., vol. 12, 1860, p. 27. Bate, Rept. Zool. Voy. “ Challenger ,” vol. 24, 1888, p. 583, pi. 104, figs. 1-1 g. Rathbun, Bull. U. S. Fish Comm., vol. 20, pt. 2, 1900 (1901), p. 114. Latreutes fucorum Stebbing, Trans. Roy. Soc. Edinburgh, vol. 50, pt. 2, 1914, p. 290, and synonymy. ' Verrill, Trans. Conn. Acad., vol. 26, 1922, p. 131, pi. 16, figs. 5-5 b, pi. 42, figs. 2-2 1, pi. 44, figs. 1-1 m, 2-2 n, 3, and syn- onymy. Off Florida; from Sargassum; 8 specimens (2 ovigerous). Off Cuba; from field of Sargassum; 23 specimens (17 ovigerous; 3 with parasitic isopod, Bopyrina latreuticola (Gissler)12 in branchial cavity). Locality?; 2 specimens (1 ovigerous; 1 with branchial parasite, Bopyrina latreuticola (Gissler)). Verrill remarks: “This delicate species is common in floating masses of ‘gulf-weed’ ( Sargas- sum). It is abundant as far north as Vineyard Sound, Mass. (S. I. Smith). Very common in the Gulf Stream further south. Bermuda (coll. G. Brown Goode). Near the Azores (Milne Edwards); African Coast (Krauss); Porto Rico (Rathbun); Beaufort, N. C. (Hay & Shore). It has been taken at Bermuda by nearly all collectors. It is nearly always asso- ciated with Leander [ Palaemon ] tenuicornis and the small crab, Planes minutusP Palaemon tenuicornis Say. (Common Gulf-weed or Sargassum Shrimp). Palaemon tenuicornis Say, Jour. Acad. Nat. Sci. Phila., vol. 1, 1818, p. 249. Leander tenuicornis Stebbing, Trans. Roy. Soc. Edinburgh, vol. 50, pt. 2, 1914, p. 288, and synonymy. Verrill, Trans. Conn. Acad., vol. 26, 1922, p. 143, pi. 43, figs. 4, 4a, and synonymy. Off Cuba; from field of. Sargassum; 25 specimens (1 ovigerous). This species is very common among Gulf-weed ( Sargassum ) and is very widely distributed in the tropical Atlantic Ocean (Verrill). Palaemon ritteri Holmes. Palaemon ritteri Holmes, Proc. Calif. Acad. Sci., ser. 2, vol. k, 1895, p. 579, pi. 21, figs. 29-35. Rathbun, Harriman Alaska Exped., vol. 10, 1904, p. 29. Palaemon sp. Rathbun, Proc. Washington Acad. Sci., vol. 4, 1902, p. 291. FPalaemon ritteri Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 561. South Seymour; shore pool, March 28; 1 juvenile. Tower Island; 1 specimen. Eden; rock pools, April 6; 1 juvenile. Northeast of Eden; seven fathoms, dredged, April 6; 1 9 ovigerous. The color of the juvenile specimen from a shore pool on South Seymour is given as “almost transparent in life, with numerous black lines, and with legs brilliant scarlet at the joints.” Miss Rathbun’s Palaemon from Clipperton Island surely is this species in spite of the somewhat different rostrum, longer sixth abdominal segment and blacker eyes. The rostrum looks a little abnormal as though possibly one or 12 Bopyroides latreuticola, Amer. Nat., vol. 16, p. 591, text-figs. 6-8. Bopyrina latreuti- cola Stebbing, Trans. Roy. Soc. Edinburgh, vol. 50, pt. 2, 1914, p. 301. 1924] Schmitt: Galapagos Macrura and Anomura 169 two of the most anterior teeth were wanting, either through malformation or accident, making the toothless part of the rostrum appear longer and more ascending than usual in this species; a count of the rostral teeth gives seven above of which two are on the carapace, and three below; the last dorsal tooth is a little behind the level of the median ventral one. The shorter ramus of the bifurcate antennular flagellum has eighteen free segments, and six fused, virtu- ally in agreement with the count of typical specimens from Magdalena Bay; in these latter there are sixteen to seventeen free segments and six fused ones. The longer sixth abdomen somite might.be an extreme variation, and the blacker eyes due to the preservation. The Peruvian specimen which Miss Rathbun with some hesitation iden- tified with this species, has the shorter ramus of the bifurcate antennular flagel- lum composed of twenty free and six fused segments, the carpus of the second legs is as long as the palm and half the length of the fingers together, the fingers are almost as long as the palm, and the fifth abdominal somite is contained one and two thirds times in the length of the sixth. The species ranges from San Diego, California, (type locality) to Peru (Rathbun). M acrobrachium jamaicense (Herbst). (Fresh-water Shrimp or Crayfish). Cancer ( Astacus ) jamaicensis Herbst, Naturg. d. Krabben u. Krebse, vol. 2» 1792, p. 57, pi. 27, fig. 2. Palaemon jamaicensis Ortmann, Zool. Jahrb., vol. 5, 1891, p. 729, pi. 47, fig. 7. Rev. Mus. Paulista, vol: 2, 1897, p. 208. Moreira, Archiv. Mus. Nac. Rio de Janeiro, vol. 11, 1901, p. 13, 78. Bithy ms jamaicensis Rathbun, Bull. XJ. S. Fish Comm., vol. 20, pt. 2, 1900 (1901), p. 123. Macrobrachium jamaicense Rathbun, Proc. U. S. Nat. Mus., vol 38 1910 p. 561, pi. 51, fig. 1. Colon; 1 d\ 1 $ . Gatun spillway, Colon; 1 juvenile. The juvenile specimen from the Gatun spillway is very probably this species. It is almost impossible to distinguish the young from those of Macro- brachium olfersii (Wiegmann).13 Miss Rathbun (1910) gives the distribution of this species as, “Fresh waters of Pacific slope of America from Lower California to Peru and of Atlantic slope, from Texas to Brazil, including West Indies.” Urocaris longicaudata (Stimpson). Urocaris longicaudata Stimpson, Proc. Acad. Nat. Sci. Phila., vol 12, i860 p. 39 [108], Kingsley, Proc. Acad. Nat. Sci. Phila., vol. 31, 1879 (1880), p. 424. Rathbun, Proc. Washington Acad. Sci., vol. 11, 1900, p. 155. Hay and Shore, Bull. U. S. Bur. Fisheries, vol. 35, 1918, p. 395, pi. 27, fig. 7. Off Cuba; from Sargassum; 1 $ ovigerous. This species is not infrequent, though never taken in large numbers14 in the area bounded by Beaufort, N. C., and the Bahamas on the North, Porto Rico and Culebra on the East and with Jacuma, Parahyba, Brazil as its southern limit. a , 13 Arch. f. Naturg., vol. 2, pt. 1, 1836, p. 150, = Palaemon spinimanus von Martens, Arch. f. Naturg., vol. 35, pt. 1, 1869, p. 26, pi. 2, fig. 3. H The U. S. National Museum possesses, in all, nineteen lots of this species, but two from Florida and one from the Bahamas contain fourteen or more specimens, two lots have four specimens, two, two specimens, and the rest one each. 170 Zoologica: N . Y. Zoological Society [V; 15 Petrolisthes edwardsii (Saussure). (Scarlet Tissue Crab). Porcellana edwardsii Saussure, Rev. et Mag. de Zool., ser. 2, vol. 5, 1853, p . 366, pi. 12. Petrolisthes edwardsii Nobili, Bull. Mus. Torino, vol. 16, no. 415, 1901, p. 11. Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 600. Eden; rock pools, April 6; 6 specimens. Ranges from the Gulf of California to Ecuador. Calcinus obscurus (Stimpson). Calcinus obscurus Stimpson, Ann. Lyc. Nat. Hist. N. Y., vol. 7, 1859, p. 83. Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 596. Nobili, Bull. Mus. Torino, vol. 16, no. 415, 1901, p. 26. Eden, off Indefatigable Island; 6 specimens. These specimens are much darker colored than others of the same species from farther north. The dactyls are all double banded, one near the tip and one at the base of the joint; in alcohol, the general coloration of the extremities is a dark, purplish red-brown with white or bluish maculations, the carapace, proximal third of the legs and eyestalks are quite a red brown; where the ground color is lighter, the spots or maculations appear more or less yellowish. A specimen from Chatham Island in the United States National Museum col- lections shows the coloration noted in this Eden material. Published records seem to indicate that this species ranges from Lower California to Ecuador. Coenobita clypeatus (Herbst). Cancer clypeatus Herbst, Naturg. d. Krabben u. Krebse, vol. 2, 1791, p. 22, pi. 23, figs. A, B. Coenobita diogenes Benedict, Bull. U. S. Fish Comm., vol. 20, pt. 2, 1900 (1901), p. 139. Verrill, Trans. Conn. Acad., vol. 13, 1908, p. 438, text-fig. 55, and synonymy. Coenobita clypeatus Rathbun, Rapport van de Visscherij en de Industrie van zeeproducten in de Kolonie Curacao, uitgebracht door Prof. Dr. J, Boeke, pt. 2, 1920, p. 327 [11], and synonymy. Colon; 3 9 . A common West Indian species ranging from Florida to Brazil; also at Bermuda. Coenobita compressus (Guerin). Coenobita compressa Guerin, Voy. autour du Monde sur la Coquille par Duper- rey, Zool., vol. 2, pt. 2, 1831, p. 29. Coenobita compressus Faxon, Mem. Mus. Comp. Zool.. vol. 18, 1895, p. 52. Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910; p. 596. James Island; 1 1 9. Miss Rathbun gives the distribution of this species as, “Lower California to Payta, Peru; Galapagos Islands; westward to East Africa.” Hippa deniiculatifrons (Miers). Remipes testudinarius, var. deniiculatifrons Miers, Jour. Linn. Soc. London , vol. 14, 1878, p. 318, pi. 5, fig. 2. Balss. Abh. der K. Bayer. Akad. Wissen., vol. 2, Math.-phys. Klasse Suppl., Abh. 10, 1914 p. 92, text-fig. 50. Remipes adactylus deniiculatifrons Ortmann, Zool. Jahrb. Syst., vol. 9, 1896, p. 229, and synonymy. Hippa deniiculatifrons Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 595. 1924] Schmitt: Galapagos Macrura and Anomura 171 Conway Bay, Indefatigable Island; March 28; 1 d\ Northeast of Eden; 7 fathoms, dredged, April 6; 1 cf. This species is known from the Galapagos Islands and the Indo-Pac-ific region as far north as Japan. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Soc ety. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” Zoologica Vol. V,No 16. HYMENOPTERA COLLECTED BY THE WILLIAMS GALAPAGOS EXPEDITION By S. A. Rohwer Bureau of Entomology, U. S . Department of Agriculture, Washington, D. C. The Hymenoptera listed below were forwarded to me by Dr. William Beebe for identification. They were collected in the Galapagos Islands, in April, 1923, in an expedition sent out by the New York Zoological Society. One of the species had been given a manuscript name by the late Dr. Ashmead from specimens col- lected in 1899. The series to which Ashmead had assigned the manuscript name is used as types for the new species and the material forwarded by Dr. Beebe are considered paratypes. Xylocopa colona Lepeletier Twenty-two specimens which agree very well with the description of this species and also agree perfectly with specimens in the National Collection which were collected in the Galapagos Islands in 1899 and later forwarded to the British Museum, where they were compared by the late Dr. David Sharp with specimens of this species in the collections of the British Museum. Ducke1 considers that colona Lepeletier is a variety of aurulenta Fabricius. For the purpose of this report I have preferred not to enter into the validity of this name. The above-mentioned 22 specimens were collected at the following localities: South Seymour, April 23, 1923; at Tagus Cove, Albemarle, April 6, 1923; at Conway Bay, Indefatigable, April 1, 1923. Tachysphex galapagensis sp. nov. In general appearance and habitus this species resembles many of the Nearctic forms, but the color of the abdomen is strikingly characteristic inas- much as the venter is entirely pale and the dor um pale on the apices of the tergites only. Female. — Length, 8 mm. Clypeus shining, convex, the anterior margin depressed and very gently rounded, without lateral teeth frons coriaceous, distinct impressed line from the anterior ocellus to between the bases of the antennae; interocellar area parted by a longitudinal depression; vertex with distimt, well separated punctures; superior interocular line less than half of the inferior interocular line and shorter han the comb'ned length of the pedi- 1 Deutsch. Ent. Zeit. 1910, p. 364. First form on press April 28, 1924. 173 174 Zoologica: N. Y. Zoological Society [V; 16 cellum and first joint of the flagellum; flagellum filiform, third, fourth and fifth joints subequal; scutum shining, with distinct, medium sized punctures which are separated laterally by a distance equal to the width of the puncture but medianly are much closer; scutellum with punctures like the scutum but much more widely separated; dorsal aspect of the propodeum coriaceous, with longi- tudinal wrinkles basally (the length of these wrinkles varies to some extent; in some specimens they extend beyond the middle of the propodeum); dorsal aspect of the propodeum perpendicular, transversely aciculate, the median depression elongate; sides of the propodeum finely longitudinally rugulose; inner spur of the posterior tibia two thirds as long as the basitarsus; abdomen shining, but when highly magnified feebly laminate; the apical margins of the tergites distinctly depressed; pygidium polished, with a few distinct punctures laterally, about two times as long as basal width; first abscissa of radius longer than either of the two following, second and third subequal (in certain paratypes the second abscissa is slightly longer than the third). Black; the venter of the abdomen, the apical margins of all tergites pale ferrugineous; the anterior tarsi, the fou posterior tarsi beneath and the spines pale ferrugineous; the spines on the basal part of the legs white; body densely clothed with silver pile; wings dusky hyaline, slightly iridescent; venation dark brown. Male. — Length, 5.5 mm. The anterior margin of the clypeus is not as broadly depressed as in the female, the convex portion slopes more abruptly; front more strongly coriaceous than in the female; superior interocular line half as long as the inferior interocular line, distinctly greater than the length of flagellar joints one and two but not quite as great as the length of joints one, two and three; flagellum very stout, the basal joints unusually short, the third and fourth shorter than the fifth; scutum sculptured similar to the female; dorsal aspect of the propodeum with the longitudinal wrinkles better defined than in the female; the ventral portions and the sides of the propodeum coriaceous; legs stout; the inner spur of the posterior tibiae three fourths as long as the hind basitarsus; apical tergite with a gentle, arcuate emargination. Colored as in female; wings hyaline, venation dark brown. A paratype female from South Seymour is smaller than the other females (6 mm.) and has a decidedly polished appearance; the sculpture is not as coarse, but otherwise it seems to agree. The paratype females from South Seymour have the wings almost hyaline. Type Locality. — Albemarle, Galapagos Islands. Paratype Locality. — South Seymour, Galapagos Islands. Described from six females (one type), one of which has lost its head, and fifteen males (one allotype) collected at the type locality March 21, 1899, and from three females from the paratype locality collected April 23, 1923. Type, allotype, paratypes. — Cat. No. 5513 U. S. N. M. Paratypes from the paratype locality returned to the collection of the New York Zoological Society. Aporinellus galapagensis sp. nov. This new species is very similar to the Nearctic fasciatus (Smith) but differs in minor ways, and it seems to be worthy of a name. 1924] Rohwer: Hymenoptera of the Galapagos [175 Male. — Length, 4.5 mm. Anterior margin of the clypeus truncate; frons finely granular with an indistinct median longitudinal line; vertex slightly raised between the posterior ocelli, shining, very feebly sculptured; postocellar line subequal with the ocellocular line; antenna stout, filiform; flagellum with short, dark bristles beneath; third antennal joint about one fourth shorter than the fourth; posterior margin of the pronotum broadly arcuate; pronotum, scutum and scutellum subopaque, finely granular; propodeum finely granular but when highly magnified appears transversely laminate, slightly depressed at the basal middle; abdomen finely laminate; inner spur of the posterior tibiae four fifths as long as the posterior basitarsus; legs feebly spined; first abscissa of the radius slightly shorter than the second; the second abscissa of the cubitus subequal with the second abscissa of the radius; second recurrent received well before the end of the second cubital cell; nervulus very slightly ante-f ureal. Black; densely clothed with silver pile which forms broad fasciae at the apical margins of the tergites; wings hyaline, strongly dusky beyond the end of the second cubital cell; venation black. Type Locality. — South Seymour, Galapagos Islands. Described from a single male collected April 23, 1923. Type. — Cat. No. 26600 U. S. N. M. Chalcis sp. A single male collected April 23, 1923, at South Seymour represents a small species which does not seem to agree with any of the described forms from South or North America, but inasmuch as the species of Chalcis are so difficult to dis- tinguish, and this is particularly true in the male, it seems unwise to describe this specimen until more material has been collected. This is. one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoologi- cal Society. Its title is “Galapagos; World’s End.” ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION Department of Tropical Research Contribution Number 183 . NEUROPTERA FROM THE WILLIAMS GALAPAGOS EXPEDITION ' By Nathan Banks . PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK November 8, 1924 Nrm f ark Xanlagiral 0>nrictg General Office: 101 Park Avenue , New York City (Mtrrn* Honorary President, Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer, Cornelius R. Agnew Inarh of Ufattagprn (Elaaa nf 1925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, C. Ledyard Blair, Anthony R. Kuser, Watson B. Dickerman, Mortimer L. Schiff, Frederic C. WAlcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr. (Elaaa nf 192B Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor (Elaaa nfl92r Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field #taff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant to the Director and Veterinarian; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector; Elwin R. Sanborn, Photographer and Editor. lE&itarial (HflmmittTT Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y. Zoologica, Vol. V, No. 17. NEUROPTERA1 FROM THE WILLIAMS GALAPAGOS EXPEDITION By Nathan Banks Neuroptera have not previously been reported from the Galapagos Islands; the small collection taken by William Beebe, though broken, are of much interest. None appear identical with any on the mainland, but all are related to Central American forms, the Megalomus, however, showing some affinity to certain South American species. The types are in the collection of the N. Y. Zoological Society. Family MYRMELEONIDAE Myrmeleon perpilosus sp. nov. Belonging to the group of M. crudelis, in which there are no black bristles above on the thorax. The face and vertex black, the face shining, no pale spots on the vertex. The pronotum with two pale spots each side and the anterior corners pale, and there are traces of pale in the middle of the meso- and metathorax. Pleura wholly dark. Abdomen with the tips of last two segments pale, and a pale spot near tip of some of the other segments; entire abdomen clothed with long white hair, rather longer than usual, and in the male much longer than in M. crudelis and allies. In the female the last ventral segment has black hair. Legs pale, the femora with broad brown stripe, tibia lined with brown, tarsal joints dark at tips. Wings moderately narrow, acute at tips; hyaline, veins dotted with dark, the main veins dark in streaks at all con- nections; costals simple, unforked, in cubital area three rows of cells, eight branches to radial sector, seven cross-veins before radial sector in fore wings. Length of front wings, 27 mm. Locality. — Conway Bay, Indefatigable, Galapagos Islands. Collected.— April 1, 1923. Family CHRYSOPIDAE Chrysopa nigripilosa sp. nov. Greenish, much marked with black. Palpi mostly black; a black stripe on each cheek under each eye, a black spot under base of each antenna; an- tennae dull brownish. Pronotum with many minute dark dots, from each a 1 Department of Trop. Research No. 183. First form, on press November 3, 192J/. 177 Route of the Noma , and details and location of the Archipelago. 178 1924] Banks: Galapagos Neuroptera 179 black hair, rest of thorax also with short, black hair, and a few dark spots. Basal part of the abdomen with a broad median dark stripe, beyond broken up; ventral segments five to seven dark. Legs pale, with black hair. Wings hyaline, stigma with about three dark spots; venation pale, much marked with black, the gradates and several of the basals wholly dark, the costals and radial cross veins black at ends, and nearly all connections of veins dark. The cubital vein just before the third cubital cell is broadened; the divisory vein ends much beyond the cross- vein; gradates divergent, inner row (of about 4) nearer to the radial sector than to the outer row (of about 7). Hind wings marked much as the fore wings, but less strongly; about three inner and six outer gradates. Both wings are moderately slender, and acute at tips. Length of fore wing, 11 mm. Locality. — South Seymour, Galapagos Islands. Collected. — April 20, 23, 1923. Chrysopa galapagoensis sp. nov. Pale yellowish (probably greenish alive), a black spot on each cheek under the eye, palpi marked with black. Pronotum as broad as long, narrowed in front, with rather long hair on sides. Wings hyaline, venation pale, marked with black; in the fore wing the gradates, several basal cross-veins, the radial, and some cubital cross-veins wholly dark, the costals and some median cross- veins partly dark; stigma greenish. In hind wings the gradates wholly, and costals and radials partly, dark. Both wings acute at tips, the fore wings not especially slender. The gradates in parallel series, about six or seven in each, the inner row almost as near to outer row as to the radial sector; divisory veinlet ends before the cross-vein. Length of fore wing, 10 mm. Locality. — South Seymour, Galapagos Islands. Collected. — April 23, 1923. Family HEMEROBIIDAE Megalomus darwini sp. nov. Face tawny, vertex darker, antennae pale brownish, paler on basal joint, thorax and abdomen dark; legs pale, front and mid tibiae with band before middle and another near tip, hind tibia very long. Wings rather slender for a Megalomus, nearly two and one-half times as long as broad, brownish, rather darker behind than in front, with darker clouds over some of the cross- veins; venation dark with here and there pale spots or short streaks, margin alter- nately pale and dark, gradates wholly dark; hind wings pale, the venation wholly dark, except some inner gradates which are hyaline. The fore wings have a broad costal area and most of the costal cross-veins forked. There are but four branches of the radial sector, the first however with two branches from the upper side, the gradate behind this branch is hyaline; outer gradates number about twelve. In hind wings the inner gradates are hyaline and hardly visible, and the four outer are not very distinct. Appendages short, not promi- 180 Zoologica: N. Y. Zoological Society [V ; 17 nent, lower appendages short, outer side rounded and end in a corneous point. Length of fore wing, 6 mm. Locality. — Conway Bay, Indefatigable, Galapagos Islands. Collected. — April 1, 1923. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoological Society. Its title is “Galapagos; World’s End.” ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V. NUMBER 18 Department of Tropical Research Contribution Number 184 ISOPODS FROM THE WILLIAMS GALAPAGOS EXPEDITION By Willard G. Van Name PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 11, 1924 New fork Zonlngtral gwrietg General Office: 101 Park Avenue, New York City (®titora Honorary President, Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer, Cornelius R. Agnew laarfc nf manager# (ttkaa of X 925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, C. Ledyard Blair, Anthony R. Kuser, Watson B. Dickerman, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr. flUaan of 1929 Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor dlaaa uf 192r Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field i>rirntifir i>taif William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant to the Director and Veterinarian; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research ; Lee S. Crandall, Curator of Birds ; George S. Huntington, Prosector; Elwin R. Sanborn, Photographer and Editor. lEfcttnrial Olnmmittee Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y. Zoologica, Vol. V, No. 18. ISOPODS1 FROM THE WILLIAMS GALAPAGOS EXPEDITION By Willard G. Van Name2 (Plates VI1I-X1X incl.) The isopods collected by the Harrison Williams Expedition at the Galapagos Islands comprise eleven species, and are of especial interest as no less than nine of them are terrestrial forms, while up to the present time but three terrestrial isopods (two of them widely ranging species) have been recorded from the islands. The col- lection is a small one, totaling only about fifty specimens, and of necessity was hastily made at a few stations only, yet that so many species of these small creatures, inconspicuous in appearance and secretive in habits, were obtained, speaks highly for the diligence and industry of the naturalists of the expedition, and indicates moreover that future collecting will probably add to the list of Galapagos isopods a number of terrestrial as well as marine species.3 I wish to express my thanks to Mr. William Beebe for the opportunity of studying this collection. The following are the eleven species obtained by the Harrison Williams Expedition and dealt with in the present article. Only the two which are marked with an asterisk were previously known from the Galapagos Islands or vicinity. Five of them are con- sidered to be new species. Suborder Flabellifera (Marine) Cirolana may ana Ives, 1891. ? *Meinertia gaudichaudii (Milne-Edwards), 1840. Suborder Oniscoidea (Terrestrial) Tylos latreilli Audouin and Savigny, 1826. Philoscia culebroides, sp. nov. Philoscia william si, sp. nov. Philoscia nomae, sp. nov. Porcellionides pruinosus Brandt, 1833. Rhyscotus laxus, sp. nov. * Cubans galapagoensis Miers, 1877. Cubaris beebei, sp. nov. Ligyda baudiniana (Milne-Edwards), 1840. The literature dealing with the Isopoda of the Galapagos Islands and vicinity is not extensive and refers mainly to marine species, 1 Cont. Department of Trop. Research No. 184. 2 Contribution from the Laboratory of Invertebrate Zoology of the American Museum of Natural History, New York. 3 Beebe (Galapagos, World’s End, p. 329) mentions the relative abundance of isopods among the few land invertebrates on Tower Island. 181 First form on press December 11, 19 2 it. Plate A. SKETCH MAP OF GALAPAGOS ISLANDS Route of the Noma, and details and location of the Archipelago. 182 1924] Van Name: Galapagos Isopods 183 many of them from deep water. It comprises two short articles by Richardson (1901, 1913; see bibliography at the end of this article) and Hansen’s (1897) account of those collected by the U. S. Albatross in 1891. Other references and descriptions are scattered, and are noted in the following pages under the species to which they apply. I have found the following isopods, only sixteen in number even when deep sea forms are included, recorded from the islands and neighboring waters. Suborder Chelifera Family TANAIDAE Tanais stanfordi Richardson. Tanais stanfordi Richardson, 1901. Proc. Washington Acad. Sci., Ill, p. 565, figs. 58-60. Clipperton Island lagoon. Family APSEUDIDAE Apseudes galapagensis Richardson. Apseudes galapagensis Richardson, 1913. Proc. U. S. Nat. Mus., XLII1, p. 159, figs. 1, 2. Albatross Station 2807, off Chatham Island, 812 fathoms. Suborder Flabellifera Family AEGIDAE Aega acuminata Hansen. Aega acuminata Hansen, 1897. Bull. Mus. Comp. Zool. Harvard, XXXI, p. 104, pi. II, figs. 3-3 b. Albatross Station 3403, Lat. 0° 58' 30" S., Long. 89° 17' W., 384 fathoms. Aega longicornis Hansen. Aega longicornis Hansen, 1897. Bull. Mus. Comp. Zool. Harvard, XXXI, p. 106, pi. II, figs. 5-5 b; pi. Ill, figs. 1-1 a. Albatross Station 3402, Lat. 0° 57' 30" S., Long. 89° 3' 30" W., 421 fathoms. Aega plebeia Hansen. Aega plebeia Hansen, 1897. Bull. Mus. Comp. Zool. Harvard, XXXI, p. 105, pi. II, figs. 4-4 d. Albatross Station 3402, Lat. 0° 57' 30" S., Long. 89° 3' 30" W., 421 fathoms; also other stations to depths of 978 fathoms. Family CYMOTHOIDAE Meinertia gaudichaudii (Milne-Edwards). Ceratothoa gaudichaudii Milne-Edwards, 1840. See Schioedte and Meinert, 1881-1883, Naturhist. Tidsskr. (3) XIII, p. 335, pi. XIII, figs. 11-15; pi. XIV, figs. 1-5. 184 Zoologica : N. Y . Zoological Society [V; 18 Widely distributed in the Pacific; parasitic in mouth of fish of the genus Thunnus and allies. Black Bight, Albemarle Island and 200 miles north of Wenman Island (Richardson, 1901, p. 568). Young specimen, probably this species, collected by the Harrison Williams Expedition. Cymothoa exigua Schioedte and Meinert. Cymothoa exigua Schioedte and Meinert, 1884. Naturhist. Tidsskr. (3) XIV, p. 232, pi. VI, figs. 7, 8, and Richardson, 1905, p. 250, fig. 261. Charles Island, from mouth of fish Citharichthys sordida. Aegathoa excisa Richardson. Aegathoa excisa Richardson, 1901. Proc. Washington Acad. Sci., Ill, p. 567, fig. 61. Nierstrasz, 1915, Zool. Med. Rijks Mus. Nat. Hist. Leyden , p. 103. Monod, 1922, Assoc. Franc. Avanc. Sci., Congres de Montpellier, 1922, pp. 405, 409. From the fin of a dolphin ( Coryphaena hippurus) in Lat. 5° N., Long. 90° W. Suborder Valvifera Family ARCTURIDAE Ardurus abyssicola Beddard. Arciurus abyssicola Beddard, 1886. (See Rept. Voy. Challenger XVII, part 1, p. 98, pi. XXI, figs. 5-8.) (Description by Beddard from a specimen dredged near the Low Archipelago in 2885 fathoms and doubtful speci- mens from between Australia and New Guinea, 1400 fathoms.) Albatross Station 2807, off Chatham Island, 812 fathoms (Richardson, 1913, p. 159). Suborder Asellota Family MUNNOPSIDAE Eurycope pulchra Hansen. Eurycope pulchra Hansen, 1897. Bull. Mus. Comp. Zool. Harvard, XXXI, p. 97, pi. I, figs. 1-1 i. Albatross Station 3413, Lat. 2° 34' N., Long. 92° 6' W., 1360 fathoms, also one other station in 1471 fathoms. Eurycope scabra Hansen. Eurycope scabra Hansen, 1897. Bull. Mus. Comp. Zool. Harvard, XXXI, p. 100, pi. I, figs. 2-2 d; pi. II, fig. 1. Albatross Station 3413, Lat. 2° 34' N., Long. 92° 6' W., 1360 fathoms. Munnopsis longiremis Richardson. Munnopsis longiremis Richardson, 1913. Proc. U. S. Nat. Mus., XLIII, p. 161, figs. 3, 4. Albatross Station 2807, off Chatham Island, 812 fathoms. 1924] 185 Van Name: Galapagos Isopods Suborder Epicaridea Family BOPYRIDAE Cryptione elongata Hansen. Cryptione elongata Hansen, 1897. Bull. Mus. Comp. Zool. Harvard, XXXI, p. 112, pi. Ill, figs. 5-5 a; pi. IV, figs. 1-1 g; Richardson, 1899, Proc. U. S. Nat. Mus., XXI, 1899, p. 869; Ann. Mag. Nat. Hist. (7) IV, p. 338; Bouvier, 1900, Trav. Stat. Zool. Wimereux, VIII, p. 285; Richardson, 1904, Proc. U. S. Nat. Mus., XXVII, p. 87; 1905, Bull. No. 54, U. S. Nat. Mus., p. 520, fig. 567 (after Hansen). Albatross Station 3407, Lat. 0° 4' S., Long. 90° 24' 30" W., 885 fathoms, parasitic on the shrimp N ematocarcinus agassizii Faxon. Suborder Oniscoidea Family ONISCIDAE Porcellio laevis Latreille. Porcellio laevis Latreille, 1804. (Very widely distributed, especially in the warmer parts of the world, but also in temperate regions. See Richardson, 1905, p. 614 and Budde-Lund, 1885, p. 138, for synonyms, distribution, etc.) Chatham Island (Hansen, Bull. Mus. Comp. Zool. Harvard, XXXI, p. 124; Richardson, 1905, p. 615, fig. 666). Cubans galapagoensis Miers. Cubans galapagoensis Miers, 1877. Proc. Zool. Soc. London, 1877, p. 74, pi. XII, figs. 2-2 c. Charles Island (Miers). Collected also by the Harrison Williams Ex- pedition. Family LIGYDIDAE Ligyda exotica (Roux). Ligia exotica Roux, 1828. Widely distributed on the shores of tropical and subtropical regions. See Richardson, 1905, p. 676, figs. 716-718, and Van Name, 1918, Bull. American Mus. Nat. Hist., XLIII, p. 72, for syno- nyms). “Guadaloupe and Clipperton Island” (Richardson, 1901, p. 568, under name Ligia exotica). DESCRIPTIONS OF SPECIES COLLECTED Suborder Flabellifera Family CIROLANIDAE Genus Cirolana Leach, 1818 Cirolana mayana Ives. Cirolana mayana Ives, 1891, Proc. Acad. Nat. Set. Philadelphia, ann. 1891, p. 186, pi. VI, figs. 3-10; Richardson, 1901, Proc. U. S. Nat. Mus., XXIII, p. 512; Moore, 1902, Bull. U. S. Fish Comm., XX, part 2, p. 166, pi. VIII, figs. 1-5; Richardson, 1905, Bull. 54, U. S. Nat. Mus., p. 87, figs. 66-70; Boone, 1921, Univ. of Iowa Studies, IX, No. 5, p. 92. 186 Zoological N. Y. Zoological Society [V; 18 antlL ont I Plate VIII. 1-3, Cirolana mayana Ives, 1891. Dorsal and lateral views, X 11, and front view of head, X 15; ant. antennae; cl. clypeus; fl. frontal lamina. (Plate VIII, figs. 1-3). This is a species widely distributed on the West Indian region and the Caribbean Sea, and reported also from San Francisco Bay, Lower California, by Richardson (1905, p. 87), who examined a large number of specimens from the last named locality without being able to detect any specific differences separating them from West Indian localities. It is represented in the Galapagos collection by a single specimen 8.3 mm. long, which I have compared with specimens from Porto Rico, Dominica, and Andros Island, Bahamas, likewise without finding any basis for separating the Pacific and West Indian forms. The head has its anterior border produced into a small triangular point between the bases of the first antennae. This point meets the frontal lamina extending up from below; the latter, seen in an anterior view, forms a small 19241 Van Name : Galapagos Isopods 187 convex keystone-shaped plate separating the bases of the second antennae; in a strictly dorsal view this plate is only narrowly visible and projects but little beyond the curve formed by the basal segments of the first antennae which form the front outline of the body when seen from that aspect. The first antennae have the two basal segments of the peduncle broad and flattened, though their width shows chiefly in an anterior view of the head; the third is much smaller and narrower; the flagellum, which is slender and has 13 articles in the present specimen, reaches near the posterior end of the first thoracic segment when drawn back. The second antennae have five segments in the peduncle, but the basal one is reduced and entirely concealed by the second in a dorsal view, and almost entirely so even in a front view of the head; the flagellum is stout and reaches well back along the fourth thoracic segment; it has 23 articles in this specimen. The anterior end of the clypeus is free and forms a rounded- triangular, downwardly and forwardly projecting process on the underside of the head in front of the mouth. The legs are stout, laterally compressed and provided with numerous stout spines. The abdomen is wide; its segments are all distinct; only the fourth segment has the lateral angles produced into pro- minent posteriorly directed points. The telson bears, as do the branches of the uropoda, a fringe of rather short hairs on the posterior part of the border. The telson has a pair of shallow depressions on the basal part of the upper surface and is rounded behind, but rather more narrowly so than is represented in the previously published figures (this is true of the West Indian specimens examined also). The internal branches of the uropoda scarcely exceed the telson; they are wide and have the external border emarginate. The external branch appears to present a good character for distinguishing this from other American species of the genus in its unusual length, as it exceeds the internal branch in length by about one-fifth. Color yellowish, with blackish pigment in irregular stellate dots on the back. For further details the reader is referred to Richardson, 1905, p. 87. The Galapagos specimen, the first record from those islands, was taken in damp sand on the shore of Indefatigable Island (Collector’s number 2014). It is not fully grown; the species reaches over 12 mm. in length. Family CYMOTHOIDAE Genus Meinertia Stebbing, 1893 ? Meinertia gaudichaudii (Milne-Edwards). Cymothoa gaudichaudii Milne-Edwards, 1840, Hist. Nat. Crust., Ill, p. 271. Ceratothoa rapax Heller, 1865, Reise Novara, Crust., p. 146, pi. XII, fig. 17. Ceratothoa gaudichaudii Schioedte and Meinert, 1883, Naturh. Tidsskr. (3) XIII, pp. 335-3M), pi. XIII, figs. 11-15; pi. XIV, fig. 5. Meinertia gaudichaudii Stebbing, 1893, Hist. Crust., p. 345; Richardson, 1899, Proc. U. S. Nat. Mus., XXI, p. 829; 1899, Ann. Mag. Nat. Hist., (7) IY, p. 171; 1901, Proc. Washington Acad. Sci., Ill, p. 568; Stebbing, 1902, in Willey, Zool. Results, p. 643; Richardson, 1905, Bull. 54, U. S. Nat. Mus., p. 237, figs. 241-245; 1910, Proc. U. S. Nat. Mus., XXXVIIT. p. 79, figs. 1, 2. 188 Zoologica: N. Y. Zoological Society [V; 18 Plate IX. 4-5, young individual, probably Meinertia gaudichaudii (Milne-Edwards), 1840. Dorsal and lateral views, X about 18. (Plate IX, figs. 4-5). The larval isopod, 4.1 mm. long, shown in Figs. 4 and 5 is in too early a stage for sure identification in the absence of other specimens in stages to connect it with the adult, but I refer it here provisionally, rather than to any allied species, largely on the ground that M. gaudichaudii is a widely distributed and apparently common Pacific species that has been twice recorded from the Galapagos Islands and their vicinity. As far as I am aware, the only figure and description of the young of this species is that given by Schioedte and Meinert (1883, Naturhist Tidsskr. (3) XIII, p. 339, PI. XIV, fig. 5; figure reproduced in Richardson 1905, Fig. 243) which represents a younger individual only 3.7 mm. long. The differences between this and the present specimen are, I think, explainable by the greater age and more advanced development of the latter. In alcohol the specimen is yellowish with the usual irregularly stellate black pigment spots on the upper parts. The first antennae reach, when drawn back, to about the rear corner of the head and are somewhat thick and inflated, consisting of 7 segments. These arise quite close together. The second antennae are longer and much slenderer, consisting of 9 segments and reaching half way along the first thoracic segment. The eyes are large and well pigmented and contain 7 rows of ocelli, with 8 in the longest row. 1924] Van Name: Galapagos I so pods 189 The six pairs of thoracic limbs are rather long and, excepting the first pair, rather stout. They are laterally compressed, with well developed hooked dactyli, which on the three anterior pairs have a few rather poorly developed dentations on the basal third of their concave aspect, those of the others appear practically smooth. Spines were not demonstrated on the propodus of any limb; the carpus of the posterior pairs, however, is provided with a distally situated spine. The propodus of the second pair is noticeably expanded. The posterior three legs have the thighs keeled, and are noticeably larger and longer than the anterior ones, increasing successively in length and stoutness. The telson is moderately wide and smoothly rounded behind. It is ex- ceeded a little by the rather narrowly oval branches of the uropoda, which, as well as the rear border of the telson itself, bear a fringe of hairs. Meinertia gaudichaudii is reported to be found parasitic in the mouth of species of Thunnus (tunnies). Its previously reported range includes the American Pacific coast from Mazatlan to Chile, and the Louislade Archipelago southeast of New Guinea (Richardson, 1905, 1910), as well as the following Galapagos localities (Richardson, 1905): Black Bight, Albemarle Island, and 200 miles north of Wenman Island. The present immature specimen was found attached just behind the gills of a fish ( Trachinotus paloma) in Conway Bay, Indefatigable Island, March 29, 1923. Suborder Oniscoidea Family TYLIDAE Genus Tylos Latreille, 1829 The isopods of this genus, which constitutes a family by itself, are dis- tinguished from the other terrestrial forms by having the uropoda modified to form, in conjunction with inwardly directed plate-like processes of the fourth and fifth abdominal segments, a cover or operculum for the other abdominal appendages which they entirely conceal. The uropoda are therefore visible only in a ventral view of the abdomen. Tylos latreilli Audouin and Savigny. Tylos latreilli Audouin and Savigny, 1826, Descript. Egypte, p. 285, pi. XIII, fig. 1 ( fide Richardson) ; Budde-Lund, 1885, Crust. Isop. Terrest., p. 273; Richardson, 1902, Trans. Connecticut Acad. Sci., XI, p. 300, pi. XL, fig. 56; 1905, Bull. No. 54. U. S. Nat. Mus., p. 586, figs. 646, 647. (Plates X-XI, figs. 6-10). See Budde-Lund, 1885, p. 273, for other synonyms and references. This is a species widely distributed in the Mediterranean region. It has been recorded also from Odessa, Russia and in the New World from the Bermudas, Santa Marta, Colombia, and Florida. There are specimens in the Galapagos collection which I do not feel justified in separating specifically from this form. For comparison I have had available specimens from Adria, Italy, received from Dr. Karl W. Verhoeff, and others from Porto Rico and Key Largo, Florida, in the American Museum collections. From the Italian and Porto Rican specimens, the Galapagos examples differ in a few minor points, notably in having the narrow upper end of the 190 Zoologica: N. Y. Zoological Society [V ; 18 epistome less prominently raised above the surface of the forehead, into which it passes with a less conspicuously impressed suture. Moreover in the Gala- pagos specimens the lobes of the head below the eyes are always wide and broadly truncated, though more variable in form in the others, and the rear end of the body (and of the opercular valves) is somewhat more broadly rounded. The telson moreover has its lower or terminal border always straight without any trace of the slight tendency to emargination usually noticeable in the specimens from the other localities. Nevertheless these differences are slight Plate X. 6-7, Tylos latreilli Adouin and Savigny, 1826. Dorsal and side views, X 8.75. and apparently more or less inconstant, and I am unwilling with the material available to attribute much weight to them, since the Florida specimens are more or less intermediate, approaching more those from the Galapagos. It is not unlikely that a species so widely distributed will prove divisible into geo- graphical races, but I do not feel that such a division should be attempted with- out much more material, collected at more numerous localities, than I have at hand. As in the case of many other widely distributed and supposedly well known species, the published figures and descriptions leave a good deal to be desired, so that the following notes on, and figures of, the Galapagos specimens seem worth including here. 1924] Van Name: Galapagos Isopods 191 Body of oblong outline, rounded before and behind when seen from above. It can be rolled up into a ball. Back highly arched; body surface minutely granular and slightly uneven, but no actual tubercles are developed except a few minute ones on the front of the epistome and clypeus. The surface of the body, limbs and antennae bears rather thinly scattered short, stiff hairs or minute setae, rising from minute depressions, which latter are more conspicuous and form a definite row along the rear border of the segments. The largest specimen, probably a male, is 10 mm. long. The others are all considerably Plate XI. 8-10, Tylos latreilli Audouin and Savigny, 1826. 8, ventral view of ab- domen and opercular plates formed by the uropoda {up) and segments 3, 4, and 5 of the abdomen, X 18. 9, front view of head X 14.5. 10, rear end of body, X 14.5. smaller and evidently not adult. Color white with scattered minute blackish pigment spots. Head well set back into the thorax; narrowed toward the front and provided with a truncated obliquely projecting lobe under each eye. In front of this lobe (between it and a wide notch occupied by the base of the second antennae) there is a small pointed lobe. The extreme front of the head is formed by the somewhat triangular epistome, below which the clypeus forms a transversely oblong plate. Eyes well developed, with nearly 30 ocelli. Second antennae rather short, with a five-jointed peduncle, unless a small movable plate on the an- terior surface of the head aboye each lateral wing of the epistome may be the rudiment of an additional joint. The terminal joint of the peduncle is the longest, and bears a flagellum which about equals that joint in length and which is divided into four articles. The first article is somewhat curved and 192 Zoologica: N. Y. Zoological Society [V; 18 bears at its proximal end a small tubercle on its upper outer aspect. The third article is the longest; the fourth is reduced to a mere conical point. The thoracic segments each have a considerable degree of individual cur- vature in an antero-posterior direction, and have the exposed part considerably but not abruptly raised above the part which slips under the segment in front* The thoracic segments except the first have the epimera distinctly articulated. The first segment has the posterior lateral corner well extended back and narrowly rounded at the apex of the somewhat acute angle which it forms. The inferior margin of the segment is not reflected outward to form a border, but it is double, owing to the existence of a narrow sulcus which extends along the anterior four-fifths of its length. The sulcus is slightly oblique in position so that it is visible in a lateral as well as in a ventral view of the segment, and it gradually closes posteriorly without forming a notch. None of the succeeding segments bear coxopodite processes. The thoracic epimera are rounded, except those of the sixth and seventh which are more squarely cut off. The epimera of the third, and especially of the fourth segments, are so small as to suggest that they have been injured or deformed, but this is normal. The legs are rather short and fairly stout, and of somewhat compressed cross section. They are unusually spinous. The telson is transversely oblong, with the upper margin arched. A rough transverse ridge or elevation crosses its surface near the rear or inferior border which projects a little farther ventrally than the border of the other abdominal segments. The form of the opercular plates is shown in fig. 8. Those of the fifth abdominal segment do not come together on the median line. Five specimens were collected under dead wood and slabs of lava, on Tower Island, April 29th, 1923. Collector’s number 2471. I may add that in Tylos spinulosus Dana, 1853, from Tierra del Fuego, (Dana, 1853, p. 717, Atlas, 1855, PI. XVIII, figs, la-le) the antennae have a three-jointed flagellum and Dana’s figure of the antennae shows an appendage entirely different in proportions and segmentation from those of the present species, so that it appears to be distinct generically. Family ONISCIDAE Genus Philoscia Latreille, 1804 This genus, which comprises a large number of species, many of them of minute size and distinguishable from each other only with considerable dif- ficulty, is represented in tropical and South America by many forms, the larger part of them still undescribed, while only more or less inadequate de- scriptions and illustrations are available in the case of many of those that have come to the notice of zoologists. In the Galapagos collection the genus is represented by six specimens, all of them more or less unsatisfactory on account of loss of parts or immaturity, but no less than three apparently perfectly distinct species are represented. I cannot identify them with any previously described forms, at least not with the information about the latter that is now available. 1924] Van Name: Galapagos Isopods 193 While under ordinary circumstances the description of new species of this perplexing group on the basis of such inadequate material would be entirely inexcusable, the exceptional interest which pertains to the land fauna of the Galapagos Islands seems to justify an exception in this case, especially since the remote and narrowly limited habitat greatly diminishes the probability of introducing more confusion into the already unsatisfactory understanding that we have of this genus, and will probably make the recognition of the forms by future collectors on the islands easy, even though the limitations in the amount and condition of the material prevent the descriptions being as full as they should be. Plate XII. 11-12, Philoscia culebroides, new species. Dorsal view and lateral outline of body of male, X 24. Philoscia culebroides, sp. nov. (Plate XII, figs. 11-12). The single specimen in the collection is a male a little under 2.5 mm. long. The color is the usual purplish brown above with irregular oblong light markings on the dorso-lateral regions, and whitish below, that is commonly found in this group. The body surface is quite smooth and shows little trace of pubescence, though the antennae bear scattered, short, stiff hairs. The integument is ex- tremely soft, perhaps because of recent moulting. The body is narrow, the abdomen large and but little narrower than the thorax at its anterior end. It forms nearly one-fourth of the body length. The head is wide and has the front outline sinuously curved when seen from above so that it is slightly prominent in the middle and under each eye. The sides of the head reach down into a large, somewhat square lobe-like ex- 194 Zoologica: N. Y. Zoological Society [V ; 18 tension below each eye, but this is appressed to the side of the head and does not project either laterally or forward. The eyes are well developed and pig- mented. The second antennae are fairly long, reaching beyond the third thoracic segment when well drawn back. Their flagellum is elongate and has the two first articles about equal; the third or terminal article is considerably the longest and bears a strong terminal bristle. The first three thoracic segments have the posterior lateral angles more or less rounded and not extended backward. The others are not rounded off, and extend backward in a progressively increasing degree. The legs are moderately long, rather stout, and have well developed spina- tion. I could not distinguish any special modification of the anterior pair or pairs of legs, except that, as usual, they are shorter and proportionately stouter than those farther back. The abdomen does not taper so much as in many allied species. The posterior lateral angles of the third, fourth and fifth abdominal segments are only produced backward to an insignificant extent. The telson is obtusely triangular with the tip considerably rounded off and the sides a little concave. The inner branches of the uropoda are small, but the outer ones, which are of more or less terete cross section, are large and long, about equaling in length the telson and the three preceding segments taken together. This specimen is very close to Ph. culebrae Moore, 1902 (See Bull. U. S. Fish. Comm., XX, Part 2, p. 176, PI. XI, figs. 13-17; also Richardson, 1905, p. 604, Fig. 660), from Culebra Island near Porto Rico, but I cannot refer it to that species on account of differences in the form and proportion of certain parts of the body. In the Galapagos specimen the head is proportionately wider, the downwardly projecting lobes under the eyes larger and more extended at the posterior ventral angle; the antennae longer, the first two joints of their flagellum more nearly equal; the abdomen somewhat larger and telson longer in proportion to its width. The label bears the collector’s number 2416. The specimen was taken on the underside of a slab of lava on Tower Island, April 28th, 1923. Philoscia williamsi, sp. nov. (Plate XIII, figs. 13-15). Named for Mr. Harrison Williams, patron of the expedition. This species appears to be a member of the same division of the genus as the one last described. As in that species, the front outline of the head is slightly (though only very slightly) sinuous, less even than in Ph. culebroides, and the sides of the head are prolonged downward below the eyes as small, somewhat square appressed lobes, but smaller than in that species. In colora- tion it is similar also, but the body surface is less smooth and is noticeably pubescent with short stiff hairs. The largest specimen is a female bearing a number of large young in the marsupium, which is distended so that it is dif- ficult to see how the animal could have walked. Body length a little over 3.5 mm. There are also two considerably smaller specimens, one a male, one a female, which are evidently immature. As is well shown by a comparison of the figures of the two species, the body 1924] Van Name: Galapagos Isopods 195 is broader in the present one, the head narrower and smaller, the abdomen much smaller (unusually small in fact), and more tapering, the telson shorter and more broadly rounded at the tip, and the branches of the uropoda proportionately very short and small. The antennae in the only adult individual are unfortunately broken off. The smaller specimens have the antennae, which are pubescent and quite long, but as the relative length of the antennae to that of the body varies with age, their length in the adult can only be guessed at and their restoration (in broken lines) in the figure must be regarded as somewhat conjectural in that respect. The first article of the flagellum is quite short and is equaled or a little exceeded by the second, and much exceeded by the third, which bears a terminal bristle. The eyes are well pigmented and have few though large ocelli. Plate XIII. 13-15, Philoscia williamsi, sp. nov. 13 and 14, dorsal view and lateral outline of body of female, X 18. 15, seventh leg X 21. As compared with Ph. culebroides this species has the lateral ends of the last four thoracic segments more squarely cut off and with the angles a trifle sharper. The corresponding angles of the third, fourth and fifth abdominal segments are produced back into more prominent triangular points than is the case in that species. I was unable to distinguish any special modification of the anterior pair or any pair of legs in the male. Though the immaturity of the specimen may make this observation somewhat inconclusive, it appears, when taken with the resemblances in the head, to support the view expressed above that this form is also a member of the same section of Philoscia as Ph. culebrae Moore, 1902. 196 Zoologica : N. Y. Zoological Society [V ; 18 Collector’s number of the three specimens 2232. They were collected under lava blocks on Tower Island, April 18th, 1923. Philoscia nomae, sp. nov. (Plate XIV, figs. 16-18). With Philoscia culehroides, described above, was a mutilated example of a much broader species, having the body surface very smooth and shining and very long legs with well developed spines. The head and antennae, part of the first thoracic segment, and the uropoda are missing. The specimen is a female. Though it resembles Ph. williamsi, just described, in color and in the small tapering abdomen, it differs in the following characters: Larger size; the specimen if entire would measure about .5 mm. long; much longer and slender legs (compare figs. 15 and 18 representing the seventh legs of each) ; only the last three instead of the last four thoracic segments have the posterior angles noticeably extended back (their apices are also less sharp than in Ph. williamsi ); the telson is obtusely triangular, the apex being little rounded off and the sides practically straight. Being a female the specimen gives no information as to possible secondary sexual characters in the anterior legs, and its incomplete condition makes it difficult to determine its nearest relationship, or identify it with any described species, but its position in this genus seems to be evident. Collector’s number 2416. Taken under lava on Tower Island, April 28th, 1923. Genus (or subgenus of Genus Porcellio Latreille, 1804) Porcellionides Miers, 1877 Stebbing (Rec. Indian Mus. Calcutta, VI, p. 188, 1911) has shown that the commonly accepted name Metoponorthus Budde-Lund, 1879, is antedated 1924] Van Name: Galapagos Isopods 197 by and a synonym of Porcellionides Miers, 1877, which must therefore be used for this group whether it be treated as a genus or only a subgenus. Porcellionides pruinosos (Brandt). Porcellio pruinosus Brandt, 1833, Bull. Soc. Imp. Nat. Moscou, VI, p. 19. Porcellio maculicornis Koch, 1835-1844, Deutschl. Crust., p. 34. Metoponorthus pruinosus Budde-Lund, 1885, Crust. Isop. Terrest., pp. 169, 171; Sars, Crust. Norway, II, p. 184, pi. LXXX, fig. 2; Richardson, 1905, Bull. 54, U. S. Nat. Mus., p. 627, fig. 674; Racovitza, 1908, Arch. Zool. Exper. et Gen. (4) IX, p. 386. Porcellionides pruinosus Stebbing, 1911, Rec. Indian Mus. Calcutta, VI, p. 189. Plate XV. 19, Porcellionides pruinosus Brandt, 1833. Dorsal view of female, X 7.5. (Plate XV, fig. 19). Many other of the numerous synonyms and references to this species are iisted in the above works of Budde-Lund and Richardson. This species of almost world wide distribution, common in nearly every country in the vicinity of human habitations, hardly needs description here. It is of rather flattened form, the back with slightly developed small granular tubercles. The legs of the posterior part of the body are long and slender, but those of the anterior part are much shorter. The long antennae with a two- jointed flagellum in which the proximal joint is long and slender, and the ter- minal one considerably shorter, are one of the most conspicuous characters by which the species may be recognized. Six specimens (the largest a female only about 8 mm. long and probably not fully grown) were obtained by the Galapagos Expedition. Five specimens (Collector’s numbers 2403, 2416, 2471) were taken in April, 1923, on Tower Island; one specimen (number 2306) was collected on Indefatigable, April 22nd, 1923. All were found resting motionless on the under- side of lava slabs. Genus Rhyscotus Budde-Lund, 1885 A small but widely distributed group of very small isopods distinguished by the great bulbous enlargement of the epistome which is well marked off from 198 Zoologicci: N. Y. Zoological Society [V; 18 the rest of the head, and the very wide, short maxillipeds which have the palp and molar portions also very wide and proportionately short. For a diagnosis and synopsis of the genus see Budde-Lund, 1906, in Voeltzkow, Wiss. Ergeb. Reise in Ostafrika, II, p. 298 ff. That author has made it the type and only genus of a subfamily (Rhyscotinae) of the Oniscidae, and in the above work divides it into two sections, to which however he does not assign names. Hyyergnathus Richardson, 1905, is regarded by him as insufficiently distin- guished from Rhyscotus. It is based on a species from Texas. Rhyscotus laxus, sp. nov. (Plate XVI, figs. 20-22). Body long and narrow, and moreover so loosely articulated that consider- able motion of the segments in a longitudinal direction is possible, while the soft integument permits of a varying degree of lateral spread of the free lateral ends of the segments. The illustration here given shows the segments quite closely approximated, so that, seen from above, the outline is that of a long narrow ellipse. Many of the preserved specimens are more relaxed and longi- tudinally extended, so that they exhibit a more parallel-sided outline. Body surface evenly, but not very thickly, covered with short hairs or setae, visible only on considerable magnification. The antennae, uropoda and legs are also more or less setose. Along the free borders of the segments the setae are a little longer and stouter and form a regular row, closer together than on the general surface of the body. In the alcoholic specimens, the back, excepting a border along the free margins of the segments, an irregular area of variable but considerable extent near each lateral end of each segment of the thorax, also numerous smaller oval spots on the median parts of the thoracic segments and on the head, is slaty gray; below considerable gray pigment is also found on the maxillipeds, first thoracic legs, and on the abdominal segments and pleopoda, except the first pair; the antennae are also quite deeply pigmented above and below, except the second and third joints. Most of the under parts and limbs and the entire uropoda, both above and below, as well as the above mentioned spots and areas on the back, are unpigmented, appearing whitish or translucent with little or no tinge of yellow. There is much individual variation in the relative extent of the pigmented and unpigmented parts. The length of the largest specimens ranges from 4.25 to about 5 mm., depending on the degree of relaxation of the muscles and consequent extension of the intersegmental membranes. This also causes the length-to-width ratio to vary greatly. Head considerably narrower than the first thoracic segment into which it is rather deeply set back. A nearly straight transverse furrow marks off the epistome from the main or posterior part of the head. The latter is produced downward and a little forward into an obtuse lobe on each side below the eyes, which have few, apparently usually about 10, well developed ocelli. The epi- stome forms a large rounded projecting bulbous expansion of the anterior median part of the head between the sockets of the second antennae. It is covered, like the rest of the body and head, with scattered short setae and shows 1924] Van Name : Galapagos Isopods 199 on careful examination a number of very faint transverse furrows on its an- terior aspect, but these are so shallow and poorly marked as to easily escape notice altogether. The second antennae are long and stout, the flagellum is long and composed of two articles, the first being considerably the shorter. The mouth appendages form a prominently projecting mass. Plate XVI 20-22, Rhyscolus laxus, sp. nov. 20, 21, dorsal and lateral -views of female, X 16 and 18 respectively. 22, ventral view of abdomen of female, X 21. The thoracic segments vary comparatively little in length. Their lateral ends are cut off in the arc of a large circle; only the last three have their posterior lateral angles noticeably extended backward, this occurring to a rapidly in- creasing degree from the fifth to the seventh. The first three have the posterior lateral angles broadly, the others quite narrowly rounded off. The portion of each segment that fits under the segment in front of it is slightly but quite abruptly lower than the rest, the change of level being marked by a noticeable 200 Zoologica : N. Y. Zoological Society [V; 18 line. The legs are, in proportion to the slenderness and small actual bulk of the body, rather long and large. They are more or less setose, but their spina- tion is rather weakly developed. The first pair are stouter than the two next pairs but exhibit no special modification. The dactylus of all the legs ends in a very minute claw on the lower side of which is a minute vesicle-like pad that is usually in a more or less collapsed condition in these specimens. The abdominal segments except the first two have the posterior angle produced back into a small acute point. The telson is triangular with straight converging sides and a rounded tip. Excepting one very minute individual whose peculiarities are evidently due to immaturity, the specimens all have pleopoda with outlines like those shown in the figure, the first pair being long, narrow and sharply pointed, while the others all have a more or less acutely pointed median posterior angle, especially those of the fifth segment. Long, narrowly produced pleopoda are found in the males only in most terrestrial isopods, but many of these specimens are certainly females, having marsupial plates under which (usually one under each plate) eggs or embryos are carried. These plates are of squarish outline and overlap very little at the median line, hence they are not very conspicuous. I am not certain whether or not any of the specimens are males, for although some lack the marsupial plates, and differ in their somewhat smaller size and in having the first joint of the peduncle of the antennae much shorter than the second, possibly these differences are to be attributed to immaturity or sterility instead of to sexual difference. The above peculiarity of the first pleopoda and the comparatively inconspicuous marsupial plates may explain Budde-Lund’s failure to recognize females (except one mutilated individual lacking the entire posterior part of the body and hence also the pleopoda) among the specimens of this genus that he examined. Collectors’ numbers of the specimens obtained: 2297 — 15 specimens, sifted from dead leaves, South Seymour, April 22nd, 1923. 2403 — 1 specimen (female, type), under lava, Tower Island, April 27th, 1923. 2416 — 1 specimen, under lava, Tower Island, April 28th, 1923. 2471— 3 specimens | under j Tower Island Apri] 2m, 1923. 2422— 3 specimens j This form belongs to the first of the two sections into which Budde-Lund (1906, in Voeltzkow, Wiss. Ergeb. Reise in Ostafrika, II, pp. 299, 301) divides the genus Rhyscotus. The species comprising it are distinguished by having the telson triangular with straight (not concave) sides, the thoracic legs ter- minated with a very small claw beneath which is a vesicle-like pad, and the basal segment of the uropoda equal in length to the inner branch it bears. To these characters this species conforms. Budde-Lund in his synopsis enumerates four species in this section. The present one differs from R. parallelus from Venezuela (see Dollfus, Ann. Soc. Entom. France, LXII, p. 342, PL IX, figs, 6-6d) in the front of the epistome showing only very faintly indicated transverse furrowing and in the lobes of the head in front of the eyes being less acute and prominent, and from R. ortonedae from Equador (Budde-Lund, 1906, op. cit., p. 299, PI. XVII, figs. 11-31) in the much less globular epistome and more 1924] Van Name: Galapagos Isopods 201 angular outlines of the body segments as seen in a lateral view. From R . linearis from the Comoro Islands (Budde-Lund, 1906, op. cit.., p. 300, PI. XVII, figs. 32, 33) a very incompletely described form, the present one would seem to be distinguished by the less broadly rounded lateral angles of the segments, a slightly broader telson, and by not having the setae on the borders of the segments conspicuously large, though as a matter of fact they are little longer than those scattered over the body surface. The widely separated localities are against their identity also. The fourth species of the section, R. cubensis (Budde-Lund, 1906, op. cit., p. 300) from Cuba, very briefly described from a single mutilated specimen comprising only the head and four body segments, is said to have the rear border of the third and fourth thoracic segments “in medio leviter incurvo,” while in this species the border is straight or even very slightly convex. It therefore seems necessary to consider this form as new. Genus Cubans Brandt, 1833 Cubans galapagoensis Miers. Cubans galapagoensis Miers, 1877, Proc. Zool. Soc. London , 187V, p. 74, pi. XII, figs. 2-2 c. Armadillo galapagoensis Budde-Lund, 1885, Crust. Isop. Terrest., p. 40; 1904, Rev. Crust. Isop. Terrest., pi. Ill, p. 108. (Plate XVII, figs. 23-27). This species was described by Miers from a single specimen 11 mm. long, collected in 1875 at Charles Island by the expedition under Commander W. E. Cookson. The present collection contains also only a single specimen, a male, considerably smaller than the original one, though in its rolled up condition it cannot be accurately measured. Miers’ description being brief, and in some respects vague, the following details are given here. Body seen from above oblong, narrowly rounded behind; in front the outline of the head forms a curve only gently arched. Body hard and compactly articulated; its surface minutely granular under magnification and raised also into rounded and elongate tubercles arranged with some regularity. These form a well defined transverse row along, though a little removed from, the posterior margin of each thoracic segment. In each of these transverse rows about six of these tubercles occupying the median region are small and rounded, those in the dorso-lateral parts (about four on each side) are larger and more elongate; at the junction of the main and epimeral portion of the segment there is a single somewhat elongate one, and on each epimeron a single obliquely placed one. A few smaller, less well defined tubercles occur in front of this row on the median portion of each segment, and on the first segment, which is much longer than the others, there are additional elongate ones and a pair of exceedingly large ones close together, one each side of the median line on the anterior part of the segment. The forehead bears a number of small tubercles. The surface of the abdomen is practically smooth except for a tubercle on each epimeron of the third, fourth and fifth segment, a pair on the anterior part of the telson, with a poorly defined one somewhat asymmetrically placed behind or below this pair. Whether this represents a somewhat misplaced median 202 Zoological N. Y. Zoological Society [V; 18 tubercle or one of a pair the other of which is not developed, I do not know. The specimen cannot be straightened out without danger of breaking it, but would probably not much exceed 6 mm. in length. The color in alcohol is rather dark gray above with the margins of the segments and the tubercles Plate XVII. 23-27, Cubaris galapagoensis Miers, 1877. 23, 24, dorsal and lateral views of male, X 14. 25, rear end of body, X 19. 26, ventral side of anterior segments, X 15.5. 27, front view of head, X 17.5. light grayish or brownish white (unpigmented), as are also the legs and most of the under parts. Head only moderately wide, eyes small with about 15 ocelli. Upper edge of epistome gently arched when seen in an anterior view, and form- ing a narrow projecting border clear across the head. Antennae of moderate length and quite slender; the flagellum is slender, shorter than the last joint of the peduncle, and consists of two articles, the basal one about one-third the length of the second or terminal one. 1924] Van Name: Galapagos Isopods 203 Thoracic segments having their exposed part noticeably (but not very abruptly) raised above the part fitting under the next segment in front, and the rear border forming a somewhat prominent ridge. Their rear lateral angles are considerably produced backward, and except the last three, conspicuously rounded in outline. The first segment has the lateral border curved as seen in a side view, and its anterior two-thirds are rolled outward to form a prominent but not very thick projecting margin. No distinctly defined groove marks this off from the lateral face of the segment. The rear lateral angles have a small nearly equal-sided cleft to receive the second segment when the body rolls up; this is not continued forward into a perceptible sulcus on the underside of the rolled out margin except for a short distance. The second segment bears a small, short, rather bluntly pointed coxopodite process between which and the inner face of the epimeron the inner division of the cleft rear angle of the first segment fits when the body rolls up. The third segment merely has the an- terior border of the epimeron thickened, but bears no process. The legs are only moderately long, rather weak and not very spiny. The abdomen has the lateral ends of the segments squarely truncated and slightly flared outward. The telson is somewhat broader than long in its upper portion; its terminal border, which is somewhat convex, projects a trifle beyond the outline formed by the other abdominal segments and is nearly two-thirds the width of the upper part. The middle portion is a little constricted. The exposed parts of the basal joints of the uropoda are longer than wide. Their inner branches (seen only in a ventral view) are exceedingly short; the outer branches are reduced to minute rudiments, short and thick, each borne on a small tubercle close to the inner margin of the outer face of the basal joint, some distance frPm the rear margin. The specimen was found under a stone on Eden Island. Cubans beebei, sp. nov. (Plate XVIII, figs. 28-30). Body oblong when seen from above, the front outline of the head and first segment, and that of the abdominal segments and uropoda, forming broadly rounded curves in front and behind. Back highly arched, its surface without any coarse tuberculation (though very slightly uneven in the dorso-lateral regions), but under considerable magnification it exhibits evenly, though not very thickly distributed scabrous punctations. Color gray-brown above with the usual light markings; legs and under parts not pigmented. Length of the largest specimens, which are perhaps not fully grown, about 5 to 6 mm. The exposed part of each thoracic segment is somewhat elevated (though not abruptly so) above the part overlapped by the segment next in front. Their posterior lateral angles are but little extended backward, those of the first three or four are rounded; the other segments become successively more squarely truncated proceeding toward the rear. Upper margin of the epistome only very gently arched and turned up to form a very narrow but distinct projecting border clear across the front of the head. This border is separated from the forehead by a very narrow impressed groove or furrow. Eyes rather small, with about 12 ocelli. Second antennae rather short and small, their flagellum is two-jointed; the basal joint is only about one-third that of the terminal one in length. 204 Zoological N. Y. Zoological Society [V; 18 Lateral border of first thoracic segment turned obliquely up (rather widely in front, the reflected part diminishing to nothing as the rounded rear angle of the segment is approached) so as to form between itself and the surface of the main part of the segment a narrow shallow groove. Posterior lateral corner of the first segment with a small cleft, but this is not extended forward as an appreciable sulcus on the inferior aspect of the margin except for an insignificant Plate XVIII. 28-30, Cubans beebei, sp. nov. 28, side view of female. X 15.5. 29, rear end of body. X 20. 30, ventral side of anterior segments, X 17.5. distance. The epimeron of the second segment bears on its inner aspect a small coxopodite process ending in a flattened bluntly rounded tip. No processes on the third segment. Legs rather small and weak, their spines rather few and small. Lateral ends of the third to fifth abdominal segments truncated and forming a smooth continuous outline with the basal segments of the uropoda and telson. They are not noticeably bent or flared outward. Telson broader than long by nearly one-fourth and considerably con- stricted in the middle; the truncated rear end is about two-thirds the width of the upper part. On the middle line near the upper end there is an elongate depression or shallow pit with a slight elevation on either side. Basal segments of uropoda a little longer than wide. Their inner branches, Visible only from below, are very short and rather wide; the outer branches are reduced to very small rudiments borne close to the inner margin (that next to the telson) of the basal joint, some distance from the rear end. Eight specimens, representing individuals of both sexes, are in the col- lection. Three of them have the collector’s number 2297 and were taken in siftings of dead leaves on South Seymour, April 22nd, 1923. The others (in- cluding the type) are labeled “ Under stones, Eden.” One of the largest females has been selected as the type. 30 1924] Van Name: Galapagos Isopods 205 This species is very closely related to several of those already described from the warmer parts of America, but most closely of all to C. grenadensis (Budde-Lund), 1893, Entomol. Meddel., ann. 1893, p. 115, from Grenada, W. I., also described briefly and somewhat crudely figured by Dollfus, 1896 (Proc. Zool. Soc. London, ann. 1896, p. 392, figs. 5a-5d), who records it from two additional islands (Becquia and Balthazar) of the West Indies. I cannot however regard the two forms as identical. In C. grenadensis the sulcus on the inferior margin of the first segment is considerably better developed, reaching half the length of the border before fading out, the body surface is “minutissime et densissime reticulate punctata” according to Budde-Lund, which would hardly be correctly applied to the rather scattered scabrous punctations of the Galapagos form; the upper margin of the epistome is described as “in medio levissime, vix memorabiliter reflexo,” while in the Galapagos specimens it is nearly as much reflected at the center as at the sides and has a decidedly more convex outline as seen from above or below than the figures of Dollfus indicate is the case in the West Indian species. His figure also shows the inner branch of the uropoda much longer and narrower than in the present specimens, and the size is much larger (14 mm. long) accord- ing to Dollfus. From another closely related species, C. dugesi (Dollfus), 1896, from Mexico, the form of the telson serves as a distinguishing character, as in that form it is as long as wide. C. pisum (Budde-Lund), 1885, from Florida, is distinguished by its much longer antennae and moreover has the minute outer branch situated near the end (“apice propius”) of the basal joint of the uropod. C. dumorum (Dollfus), 1896, from Mustique Island, W. I., has the coxopodite process of the second segment large and square. C. vincentis (Budde-Lund) syn. C. cincta , C. perlata and C. silvarum, all described by Dollfus (1896) from the West Indies, as well as C. galapagoensis Miers, 1877, redescribed above in this paper, are all near allies of the present form, but distinguished either by the tuberculation of the body surface or by the much greater length of the sulcus on the margin of the first segment of the thorax (in the case of C. perlata by both these characters). Family LIGYDIDAE Genus Ligyda Rafinesque, 1814 (= Ligia auct. plur.) Ligyda baudiniana (Milne-Edwards). Ligia baudiana Ives, 1891, Proc. Acad. Nat. Sci. Philadelphia, p. 185, pi. VI, fig. 2. Ligia baudiniana Milne-Edwards, 1840, Hist. Nat. Crust., Ill, p. 155; Saussure, 1858, Mem. Soc. Phys. Hist. Nat. Geneve, XIV, p. 476; Stuxberg, 1875, Ofvers. K. Svensks. Vetensk.-Acad. Forh., XXXII, No. 2, pp. 43, 46, 48; Ives, 1891, Proc. Acad. Nat. Sci. Philadelphia, pp. 199, 200; Richardson, 1901, Proc. U. S. Nat. Mus., XXIII, p. 574; 1902, Trans. Connecticut Acad. Sci., XI, p. 306, pi. XL, fig. 61; 1904, Proc. U. S. Nat. Mus., XXVII, pp. 24, 30; Chilton, 1916, Mem. Indian Mus. Calcutta, V, pp. 464, 466, 472, 473; Jackson, 1922, Proc. Zool. Soc. London, pp. 689, 698, pi. II, figs. 17, 18. Ligia exotica (not Roux, 1828) + L. exotica var. hirtitarsis Dollfus, 1890, Bull. Soc, Etudes Sci. Paris, XII, p. 7, figs. 5, 6. Ligia hirtitarsis Dahl, 1892, Ergeb. Plankton-Exped., I, pi. 1, p. Ill, pi. Ill, figs. 1, 6, 7, 11, 12. 206 Zoologica : N. Y. Zoological Society [V ; 18 Ligia gracilis Moore, 1902, Rept. U. S. Comm. Fisheries, XX, part 2, p. 175, pi. XI, figs. 7-12. Ligyda baudiniana Richardson, 1905. Bull. 54, U. S. Nat. Mus., p. 678, figs. 719-723. The following references appear to apply to this species more or less doubtfully: Ligia baudiana Miers, 1877, Proc. Zool. Soc. London , pp. 670, 671. Ligia baudiniana Bate, 1868, Ann. Mag. Nat. Hist. (4) I, pp. 443, 446. (Plate XIX, figs. 31-36). The Galapagos collection contains specimens which I am unable to separate specifically from this species inhabiting the West Indian region and Atlantic coast of America from Florida to Rio Janeiro, although not reported on the Pacific side. The largest specimen (fig. 31) from the Galapagos has more elongate antennae, which reach to or beyond the end of the body, and longer uropoda than are credited to this species in the descriptions except in the somewhat doubtful one of Miers, 1877, (which may refer to another species, L. exotica Roux, 1828), but few museum specimens have these parts (especially the uropoda) attached and complete, so that we actually have very little infor- mation as to how long these parts normally become in L. baudiniana, although the specimens available show that there is considerable variation individually and with age. Under these circumstances I cannot base a specific distinction on this character. In its general form and character this species resembles the well known and widely distributed L. exotica (Roux), 1828, but is distinguished by its smaller size, somewhat wider head and more elongate eyes, its obtusely ending telson, the more complete fusion of the epimera with the main portion of the thoracic segments, and especially by the first legs of the male, which lack the small lateral process at the distal end of the propodus that is present in L. exotica but have the merus and carpus flattened and provided on the thin inferior borders with a single row of short, close-set spiny hairs. These joints are also provided with a file-like area of fine, parallel, obliquely transverse ridges and furrows on their anterior aspect. The second and third legs have the corpus swollen and provided with a narrow file-like area, but the fringe of spiny hairs is present only on the first pair. In the female the three anterior legs are similar to the fourth and the more posterior pairs which are alike in both sexes. Some other particulars not shown in the accompanying figures are as follows: The body surface is covered with scattered, very small, low tubercles; slightly larger ones form a row along the rear edge of the segments. The color is greenish gray, due to minute, thickly but unevenly scattered, irregularly stellate pigment spots. The largest male and female (13.5 and nearly 12 mm. long respectively) have antennae reaching beyond the end of the body when drawn back and having 43 or 44 articles in the flagellum. The smaller specimens have the antennae proportionately much shorter (not reaching much beyond the end of the thorax) and with fewer articles (sometimes less than 30). The uropoda were lost except in two specimens. In the large male they are long, the basal Plate XIX. 31-36, Ligyda baudiniana (Milne-Edwards), 1840. 31, dorsal view Of female, X 6.4. 32, first leg of female X 8.4. 33, first leg of male X 7.2. 34, second leg of male X 7.2. 35, second leg of female X 8.4. 36, styloid process of right second pleopod of male, ventral view, X 12. 207 208 Zoologica : N. Y. Zoological Society [V; 18 joint almost equaling the exposed part of the abdomen measured on the median dorsal line, and the inner branch (the outer is missing in both cases) consider- ably exceeding the basal joint in length. In one of the smaller specimens the uropoda are, however, proportionately much shorter. I have compared the stylets of the second pleopoda in males of the Galapagos and West Indian specimens and find them closely similar, but they are very different from those of L. exotica, which have the ends enlarged and rounded. In L. baudiniana the organ tapers to a not very acute tip which is twisted slightly inward toward the median line. The ventral surface bears a shallow groove along the entire length of the stylet. This groove has its outer margin more prominent than the inner. From a small cleft on the dorsal surface near the distal end, a taper- ing flexible fleshy process of varying length protrudes. Six specimens are in the Galapagos collection. Collector’s numbers: 2471 — 3 small specimens (not adult). 2232 — 1 male. 2416 — 1 female. These were all taken under lava blocks on Tower Island, in April, 1923. Conway Bay, Indefatigable Island, April 1, 1923. One male (largest specimen). This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, .are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoological Society. Its title is “Galapagos; World’s End.” BIBLIOGRAPHY Beebe, W. 1924 Galapagos, World’s End. New York, 443 pp., 107 illusts. Bouvier, J. 1900 Contribution a l’6tude des Epicarides — les Bopyridae. Trav. Stat. Zool. Wimereux, VIII, pp. 1-396, pis. I-XLI, Paris. Budde-Lund, G. 1885 Crustacea Isopoda Terrestria per familias et genera et species descripta. Hauniae, pp. 1-319. 1904 A Revision of Crustacea Isopoda Terrestria with additions and illustrations. Part II. — Spherilloninae; Part III. — Armadillo. Copenhagen, pp. 33-144, pis. VI-X. Hansen, H. J. 1897 Reports on the Dredging Operations off the West Coast of Central America to the Galapagos, to the West Coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U. S. Fish Commission Steamer ‘Albatross’ during 1891. Part XXII. — The Isopoda. Bull. Mus. Comp. Zool., XXXI, pp. 95-129, pis. I- VI, map. Jackson, H. G. 1922 A revision of the isopod genus Ligia (Fabricius). Proc. Zool. Soc. London, 1922, pp. 683-707, pis. I, II. Miers, E. J. 1877 Crustacea, in Gunther, A., Account of the Zoological Collection made during the visit of H. M. S. ‘ PetereV to the Galapagos Islands. Proc. Zool. Soc. London, 1877, pp. 73-75, pi. XII, figs. 2-2c. Monod, Th. 1922 Remarques sur le genre “Aegathoa” Dana suivies de la de- scription d’Ae. indicatrix, nov. sp. Assoc. Fran?, p. Avanc. des Sci., Congres de Montpellier, 1922, pp. 405-413, figs. 1-12. Nierstrasz, H.-F. 1915 Die Isopoden-Sammlung im naturhistorischen Reichsmuseum in Leiden. Part I. — Cymothoidae. Zool. Mededeel. Rijks Mus. Nat. Hist. Leyden, ann. 1915, pp. 71-108, pis. Ill, IV. Richardson, H. 1899 Key to the Isopods of the Pacific Coast of North America with descriptions of twenty-two new species. Proc. U. S. Nat. Mus., XXI, pp. 815-869. (Reprinted in Ann. Mag. Nat. Hist. (7) IV, pp. 157-187, 260-277, 321-338). 1901 Papers from the Hopkins-Stanford Galapagos Expedition. VI. — The Isopods. Proc. Washington Acad. Sci., Ill, pp. 565-568, figs. 58-61. 1904 Contributions to the Natural History of the Isopoda. Proc. U. S. Nat. Mus., XXVII, pp. 1-89, 657-681, 131 text-figs. 1905 A Monograph of the Isopods of North America. Bull. 54, U. S. Nat. Mus., pp. i-liii, 1-727, text figures 1-740. 209 210 Zoologica: N. Y. Zoological Society [V; 18 Richardson, H. ( Continued ) 1910 Report on Isopods from Peru collected by Dr. R. E. Coker. Proc. U. S. Nat. Mus., XXXVIII, pp. 79-85, figs. 1-6. 1913 Descriptions of two new Isopods, an Apseudes and a Munnopsis, both from the Galapagos Islands. Proc. U. S. Nat. Mus., XLIII, pp. 159-162, figs. 1-4. Schioedte, J. C. and Meinert, F. 1884 Symbolae ad Monographiam Cymothoarum Crustaceorum Isopodum Familiae. Part IV. — Cymothoidae. Naturhist. Tidsskr. (3) XIV, pp. 221-454, pis. VI-XVIII. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION PARASITIC COPEPODS FROM THE WILLIAMS GALAPAGOS EXPEDITION By Charles B. Wilson PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK December 15, 1924 fork ^nnlngtral ^orirtg General Office: 101 Park Avenue , New York City WffxttVB Honorary President, Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer , Cornelius R. Agnew Boartu of IManagera aHaan of 1925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, C. Ledyard Blair, Anthony R. Kuser, Watson B. Dickerman, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr. (Elasa nf 102fi Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor mo&B of 102 7 Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field #ripnttlfc William T. Hornaday, Director of the Zoological Park ; W. Reid Blair, Assistant to the Director and Veterinarian; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector; Elwin R. Sanborn, Photographer and Editor. EMtnral ©Dmmtttpp Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y . Zoologica, Vol. V, No. 19. PARASITIC COPEPODS1 FROM THE WILLIAMS GALAPAGOS EXPEDITION By Charles B. Wilson (Plate XX). The seventh expedition of the Department of Tropical Research of the New York Zoological Society was known as the Williams Galapagos Expedition. They visited the little group of islands named, which have become biologically famous, and Mr. William Beebe, who acted as Director of Scientific Work, collected quite a number of parasitic copepods. These were forwarded to the present author with the request that they be identified and such notes added as might be of interest. So little is known of the copepod fauna of that region that even a bare list of the species obtained would be well worth publication. In the present instance also the species are themselves of considerable interest and their presence here on the equator in the Pacific Ocean greatly modifies our pre- vious knowledge of their distribution. Caligus parvus Bassett-Smith, A single female was obtained from the outside surface of a large brown grouper at Conway Bay, Indefatigable Island, April 1, 1923. The only previous record of this species was made by Bassett-Smith who obtained both sexes from a puffer in Bombay Harbour, India. He said of them . . . “They were often seen actively moving about, and were of a pinkish color, both sexes being equally common.” The present specimen still retains in the preservative a decided pinkish hue. Lepeophtheirus disBimulatus Wilson. Fifteen specimens, including both sexes, were taken in company with the preceding species upon the same grouper. This species has been found twice before, the original type specimens being taken from a white-spotted serranus, Epinephelus labriformis, at Charles Island, another of the Galapagos group. A second lot was obtained from the red grouper, Epinephelus morio, at the Bermuda Islands in 1903. These two lots, together with the present one, indicate that the species is fairly common upon the groupers of both the Atlantic and Pacific Oceans in the tropics. Caligus irritans Heller. A single female was taken from the mouth of a large-eyed blue fish in Conway Bay, Indefatigable Island. This species was originally described by *Cont. Department, Trop. Research No. 185. 211 First form on press December 15, 1925 Plate A. SKETCH MAP OF GALAPAGOS ISLANDS Route of the NoYna, and details and location of the Archipelago. 212 1924] Wilson: Galapagos Parasitic Copepods 213 Heller from specimens taken from the gills of a grouper off the coast of Brazil. Both sexes were also found by the present author upon the gills of the red- mouthed grunt and the crevalle at Jamaica in the West Indies. This specimen from the Galapagos Islands indicates that the species may be looked for in the tropics of both oceans. Panclarus satyrus Dana. Three females and a male were taken from the outside skin of a nine-foot shark captured at night in Seymour Bay, Indefatigable Island. Another female was found upon the skin of a six-foot shark at Tower Island. This parasite had worn a rather deep depression into the skin of its host. A single female was also taken from the outer skin of an eight-foot shark at Seymour Bay, Indefatigable Island. Dana’s original type specimens came from a large shark captured south of Tongatabu, one of the Friendly Islands. The U. S. National Museum contains a single lot of 15 females taken from the sides and pectoral fins of a blue shark, Prionace glauca, at the Hawaiian Islands. The male was first described from two specimens taken from a large shark in the tropical Atlantic. It seems to be a very common parasite at the Galapagos Islands. Paralebion elongatus Wilson. Three females and two males were taken from a nine-foot shark at Inde- fatigable Island. The types of this species were obtained from a shark caught in Chesapeake Bay in the summer of 1910. Nessipus costatus sp. nov. (Plate XX, figs. 9-16). Host and record of specimens. — Two females without egg strings and one male were taken from a nine-foot shark at Indefatigable Island, April 21, 1923. There is no record of the part of the body of the host, upon which they were found, but if we may judge from the records of other species of this genus the male was probably found on the outside surface or fins, while the females came from the throat or gill cavity. Costatus, ribbed, alluding to the frontal plates. Specific characters of female. — Carapace nearly orbicular, a trifle wider than long; lateral areas of medium width; posterior lobes short, wide and bluntly rounded; cephalic area large; frontal margin evenly rounded, with a small incision at the center; frontal plates wide and distinctly ribbed transversely, a very distinctive character. Second and third thoracic segments fused and furnished with a single pair of lateral plates, well rounded and reaching behind the posterior lobes of the carapace; no dorsal groove indicating the line of separation between these segments; posterior margin of third segment three-lobed. Fourth segment contracted anteriorly but not forming a neck as in the following species; its two dorsal plates fused with no indication of their dual origin, the combined plate slightly wider than the genital segment and without a trace of a posterior invagination. Genital segment one-half longer than wide, with straight parallel sides and broad evenly rounded lobes at the posterior corners; posterior invagination wide 214 Zoologica: N. Y. Zoological Society [V; 19 and shallow, showing the abdomen; the latter is one-jointed and its posterior margin just reaches the tips of the posterior lobes on the genital segment; anal laminae fairly large, each armed with four plumose setae. First antennae of the usual pattern; second pair long and slender, with a pad at the base but none on the second joint, the terminal claw without accessory spines. First maxillae similar to those of other species but more evenly rounded, with two small spines at the tip and a minute palp armed with three tiny spines. Second maxillae without accessory spines on the second joint and at the tip. Maxillipeds without a claw but with wart-like processes shutting together as in some species of Pandarus. Swimming legs as in other species, the first pair with five spines on the outer margin of the terminal joint of the exopod, the fourth pair with three spines on the outer margin of the exopod; these spines all taper to a sharp point. Specific characters of male. — Carapace elliptical, longer than wide, with narrow lateral areas and a large cephalic area; posterior lobes curved inward at their tips; frontal margin more convex than in the female, with a slit-like central invagination; frontal plates distinctly ribbed as in the female; two small ellip- tical lenses, transparent and placed diagonally just behind and on either side of the tripartite eye. Second, third, and fourth segments about the same length but diminishing regularly in width, the last one wider than the genital segment and nearly twice as wide as long. Genital segment one-fourth wider than long, with prominent conical lobes at the posterior corners. Abdomen one-jointed; annal laminae small and angular, each armed with four plumose setae a little larger than those on the female. Antennae, mouth parts, and legs similar to those of the female. Color (preserved material). — Both sexes a uniform yellowish-white. Total length of female 5 mm., of male 4 mm. Carapace of female 2.80 mm. wide, 2.60 mm. long. Carapace of male 2.30 mm. wide, 2.50 mm. long. Remarks . — The conspicuous transverse ribs of the frontal plates furnish the distinctive character which is the easiest to recognize, but the general makeup of the body and the details of the appendages are considerably different from those of other species. Nesippus occultus sp nov. (Plate XX, figs. 1-8). Host and record of specimens. — One female and nine males were taken from a nine-foot shark at Indefatigable Island, April 21, 1923. As with the preceding species no statement was made in regard to the location of the specimens upon the shark’s body. Usually in this genus the males frequent the outside surface while the females are found in the mouth or throat. Ordinarily also only one or two females are found upon a single shark and this would explain somewhat the disparity in the specimens of the two sexes. Occultus, hidden, alluding to the abdomen. Specific characters of female. — Carapace transversely elliptical, one-eighth wider than long; frontal plates distinct and relatively very narrow; frontal 1924] Wilson: Galapagos Parasitic Copepods 215 margin evenly rounded without any incision at the center; posterior lobes short, wide, and rather pointed; lateral areas narrow for this genus, cephalic area large. Plate XX. 1, Nesippus occultus , dorsal surface of female; 2, dorsal sur- face of male; 3, second antenna; 4, mouth tube and first maxillae, the latter not in their normal position (see text); 5, second maxilla; 6, maxilliped; 7, first leg; 8, fourth leg; 9, Nesippus costatus, dorsal surface of female; 10, dorsal sur- face of male; 11, second antenna; 12, first maxilla; 13, second maxilla; 14, maxil- liped; 15, first leg; 16, fourth leg. 216 Zoologica : N . Y. Zoological Society [V; 19 Second and third thoracic segments fused as usual, with only a single lateral plate on either side, but with a fairly distinct dorsal groove between the seg- ments. Fourth segment contracted into a narrow neck anteriorly and ex- panded posteriorly into a pair of dorsal plates, which are about half the width of those on the second and third segments. Here again the two are so thor- oughly fused as to give no evidence of their dual origin either by a longitudinal dorsal groove or by a posterior sinus on the midline. Genital segment elongate elliptical, twice as long as wide, with the lateral margins somewhat evenly rounded and with a deep posterior sinus, one-fourth of the entire length of the segment; posterior lobes broadly and evenly rounded, overlapping slightly on the midline. Abdomen and anal laminae entirely con- cealed, the former wider than long with parallel sides, the latter rather large and somewhat triangular, each armed with four plumose setae. Egg strings narrow and twice the length of the body. First antennae like those in other species; second pair relatively large with a small adhesion pad on the ventral surface at the base of the terminal claw. The latter is large and curved into a half circle, with the concave surface flat- tened and armed with a row of minute teeth along either margin and with two setae near its proximal end. First maxillae short and wide laminae, more or less angular, with a single short spine at the tip and near the center of the inner margin a minute palp tipped with three tiny spines. These laminate maxillae stand on edge, projecting from the ventral surface parallel with each other. In the figure (4) here given they were cut loose from the ventral surface of the head, the posterior portion of. each was turned outward away from the mouth tube, and they were turned onto their sides in order to show their shape. Second maxillae long, the basal joint quite stout, the terminal portion slender and apparently jointed at the center. There are two small spines at the distal end of the basal portion on the inner surface and a tiny claw at the tip of the terminal portion, attached like a fingernail. This female was evidently securely attached to her host, since one maxilliped was pulled off and the tip of the other was broken. Enough was left, however, to indicate that the maxillipeds of the female are like those of the male. The swimming legs are like those of other species of the genus, but each of the spines on the terminal joint of the exopod of the first legs is swollen at the end, and is tipped with a minute secondary spine. Specific characters of male. — Carapace relatively wider than in the female and with a slight median incision on the frontal margin; lateral areas also wider, cephalic area the same. Thoracic segments like those of the female with the second and third segments distinctly separated; fourth segment contracted anteriorly into a neck shorter and wider than in the female. The posterior expansion of this segment is no wider than the anterior portion of the genital segment, while in the female it projects considerably on either side. The genital segment is quadrangular with nearly parallel sides, the posterior corners produced into minute spines, the posterior margin with a convex double curve. Abdomen one-jointed, two-fifths of the width of the genital segment; anal laminae very large, each of them being twice the area of the abdomen and armed with four large plumose setae. 1924] Wilson: Galapagos Parasitic Copepods 217 The antennae and mouth parts are like those of the female. The terminal joint of each maxilliped is much swollen until it is as wide as, or wider than, long. It is tipped with a stout claw, strongly curved and armed with a second- ary spine near the center of the concave margin. Opposite the base of the claw is a spherical knob over which the tip of the claw shuts down tightly. The surface of this knob is corrugated and anything like a fold of skin, caught between the claw and the knob, is held as if in a vise. Color (preserved material). — Both sexes a uniform yellowish-white, tinged with brown over the reproductive organs; egg strings light brown. Total length of female, 6 mm., of male, 5 mm. Carapace of female, 2.60 mm. long, 3 mm. wide. Carapace of male, 2.50 mm. long, 3 mm. wide. Genital segment of female, 2.80 mm. long, 1.50 mm. wide. Genital segment of male, 1 mm. long, 1 mm. wide. Remarks . — This species resembles Heller’s Nesippus crypturus more nearly than any other of the described forms, but shows the following specific differ- ences. In crypturus the fused second and third thoracic segments are con- tracted posteriorly into a neck of the same width as that of the fourth segment; here there is no neck at all on the third and fourth segments. In crypturus the fourth segment is about the same width and length; here it is twice as wide as long. In crypturus the terminal claw of the second antenna is but slightly curved and its lateral margins are smooth; here it is bent into a half circle and each lateral margin is armed with a row of fine teeth. In the second maxilla Heller’s species showed a single spine on the outer margin of the second joint at the distal end and the terminal claw tapered to a sharp point. Here there are two spines on the inner margin of the second joint near the distal end, and the terminal claw is bluntly tipped with an accessory spine attached like a toe- nail. The types of both these new species of Nesippus are deposited in the Department of Tropical Research of the New York Zoological Society. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoological Society. Its title is “Galapagos; World’s End.’’ ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V. NUMBER 20 Department of Tropical Research Contribution Number 186 COLEOPTERA FROM THE WILLIAMS GALAPAGOS EXPEDITION By Andrew J. Mutchler PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK January 20, 1925 Nrw fork Xnologtral 8>orirtg General Office: 101 Park Avenue, New York City Honorary President, Henry Fairfield Osborn; Vice-Presidents, Madison Grant and Frank K. Sturgis; Secretary, Chairman, Exec. Committee, Madison Grant; Treasurer, Cornelius^R. Agnew loarb of fftattagprn dlasa of 1925 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, C. Ledyard Blair, Anthony R. Kuser, Watson B. Dickerman, Mortimer L. Schiff, Frederic C. Walcott, Beekman Winthrop, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr. flllaaa of 192B Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor flllaiia nf 192 7 Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman Du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field ^taff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant to the Director and Veterinarian; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds ; George S. Huntington, Prosector; Elwin R. Sanborn, Photographer and Editor . lE&ttnnal Committpp Madison Grant, Chairman; William T. Hornaday Charles H. Townsend William Beebe Elwin R. Sanborn, Sec’y. Zoologica, Vol. V, No. ZO. COLEOPTERA* 1 FROM THE WILLIAMS GALAPAGOS EXPEDITION By Andrew J. Mutchler (Figs. 42-46 incl.) The first records of Coleoptera from the Galapagos Islands were based on specimens collected by Charles Darwin while on the famous 'Beagle’ Expedition. There were twenty-nine species in the series, one of which was described as new by the Reverend F. W. Hope (1837). The remainder were reported on by Mr. George R. Waterhouse (1845). He erected three new genera for forms which could not be placed in the then known genera and described twenty-two species as new. Of the other six species he made the following remarks: "But four species amongst the Gala- pagos Coleoptera occur, so far as I have been able to ascertain, in any other quarter, and of these, two ( Dermestes vulpinus and Corynetes rufipes 2) are insects which, feeding upon dried meat and such substances, have been carried to all parts frequented by ships; the third is a wood feeding insect (genus Apaie), and might be trans- ported for a considerable distance by floating timber; and the fourth is a water-beetle which appears to me clearly identical with Hydro- philus lateralis (genus Tropisternus of Solier), an insect found in the United States, Mexico, and some of the West Indian islands. 1 should observe, moreover, there is in the collection a second, minute, species of Hydrophilus closely resembling the Philhydrus affinis of our English collections, but which is rather smaller, less distinctly punctured, and of a darker hue.” The above refers to five of the species; of the sixth, which is a staphylinid, he says: "Three speci- mens found under a dead bird in Chatham Island. These specimens approach very nearly in size and form Cr[eophilus] maxillosus of Europe, and the C. villosus of North America.” These islands were visited again in 1852 by members of the Swedish frigate ' Eugenie ’ and from the Coleoptera obtained during their stay, Boheman (1858-1859) described six new species; but, if any of the species collected by Darwin were among this material, records of them were not included in the account. The visit of H.M.S. 'Petrel,’ in 1875, brought forth additional records of the Coleopterous fauna of these islands. The beetles collected on this 1 Gont. Department, Trop. Research No. 186. 1 Necrobia rufipes. 219 Plate A. SKETCH MAP OF GALAPAGOS ISLANDS Route of the Noma , and details and location of the Archipelago, 220 1924] Mutchler: Galapagos Coleoptera 221 expedition were treated by Mr. Charles Waterhouse (1877)3 who described three new genera and six new species. He also lists the species known to him to have occurred in these islands and cites, when known, the particular islands on which they were collected. The list contains thirty-nine described species and one questionable form of Philhydrus. The next records were by Dr. L. 0. Howard (1890) who, in an “Annotated Catalogue of the Insects Collected in 1887-1888, by naturalists of the U. S. Fish Commission, Steamer 'Albatross/ ” lists nine species with the specific names. One of these, Calosoma galapagoum Hope, was found to be incorrectly determined as the form described by Hope, and was later treated as a new species by Lined. Dr. Howard also included a question- able species of Mallodon and mentions three unidentified specimens of Curculionidae. The species listed by Dr. Howard were deter- mined by Martin Lined. The undescribed species contained in the above material were treated by Lined (1898) who also included the results of another ‘Albatross’ Expedition in 1891 and the collection made by Dr. G. Baur of Clark University during the same year. In the paper by Lined, which was printed posthumously, there are fourteen species described as new and six previously described species not included in former papers. There are also references with notes to species which had been recorded from the islands by Hope, Geo. R. Waterhouse, Boheman, and Howard. There is also a record by F. X. Williams (1907) of the Expe- dition of the California Academy of Science, in which a list of the families of beetles, collected by that expedition, are given. The account is as follows: “ Coleoptera, 150 species; one nocturnal species of Cicindela; Calosoma plentiful; Dytiscidae, 5 species; Gyrinidae, 1 species; Hydrophilidae, 1 species; few Staphylinids were found on rotting cactus and carrion; Coccinellidae, 3 species; Dermestidaewere common; Histeridae not common; Elateridae, 5 species; Buprestidae, one small species; Cleridae, 1 species; several Ptinids; Scarabaeidae, 2 species — one a Trox; Cerambycidae, 10 species; Chrysomelidae, 2 small species; Tenebrionidae were common. Rhyncophora numer- ous.” He also states that Coleoptera were abundant in cereals, especially on the vessel. The specimen of Cicindela mentioned in the above article was listed by Dr. Walther Horn in ‘ Genera lnsecto- rum’ (1915), fasc. 82c, as Cicindela galapagoensis who gives as the authorities "Van Dyke, in litt.; Williams in litt.” 1 believe the s This paper was not included in the records by Martin Linell. 222 Zoologica: N. Y. Zoological Society [V; 20 brief description of the species on pp. 238, 241, 251, 397 and 399, given by Dr. Horn in the above publication, are sufficient to estab- lish the species and credit should be given him as the author. There is also a more complete description of the species in Archiv for Zoologi, 1920, XIII, No. 11, p. 17. 1 have inquired of Dr. E. C. Van Dyke concerning the material collected by the California Academy of Sciences and have been informed that the above species is the only beetle, of the lot col- lected by F. X. Williams on the Galapagos Islands, which has been described. The material before me was collected by the Harrison Williams Galapagos Expedition of the New York Zoological Society under the directorship of William Beebe. It includes twelve forms previously recorded from the islands, one species which is more or less cos- mopolitan but not heretofore recorded and eight forms which 1 am herewith describing as new. It seems remarkable that six of the new species are contained in genera which have not heretofore been recorded from these islands, one of which, apparently, cannot be placed in any previously described genus. 1 have also included in the following pages the description of a species of Calosoma which was collected by Dr. G. Baur and wrongly determined by Linell. 1 wish to thank Mr. William Beebe for his kindness in allowing me the privilege of identifying this material, also to express my gratification to Prof. William Morton Wheeler for notes on the environment of many of the species. The determination of the species has been greatly facilitated through the courtesy of the U. S. National Museum in allowing me the privilege of comparing specimens with the Linell material and the kindness. of Mr. K. G. Blair in comparing specimens with the Waterhouse types in the British Museum. Calosoma linelli sp. nov. Calosoma galapageium (Hope) Linell, 1898, Proc. U. S. Nat. Mus., XXI. p. 250. This species was determined as C. galapageium Hope by Linell. The one specimen on which this determination was made was later sent to Mr. K. G. Blair of the British Museum, who returned it with the following remarks: “It . . . seems possible that Linell was mistaken as regards galapageium as the type is 16 mm. in length, is more shining than the other and does not in the least suggest a Cychrus ” 1924] Mutchler: Galapagos Coleoptera 223 The specimen was so fully described by Linell that I believe it better to repeat his diagnosis than to attempt to make a new description: “Form and size of Cychrus stenostomus, apterous, smooth, and very shining. Head black, impunctate, mandibles piceous; labrum and palpi ferruginous. Antennae ferruginous, slightly darker outward, finely rufo-pubescent from the fifth joint, reaching the elytra to one-fourth the length from the base. Thorax black, aeneous at the base, entirely impunctate, slightly wider than long, sub- cordate, somewhat wider at apex than at base; disk feebly convex, not depressed at the sides; median line distinctly impressed; basal fovae rounded, deep, ap- proximate to the sides; base truncate; posterior angles prolonged and deflexed. Elytra at base slightly wider than the thorax at middle, ovate, one-half longer than broad, dark cupreous green; humeri rounded; disk slightly convex, feebly (at sides and apex obsoletely) punctato-striate; intervals nearly flat, smooth; the third, seventh, and eleventh with feebly convex, elongate elevations, separ- ated by rounded very shallow foveae, each fovea with a couple of punctures. Epipleura and ventral surface reddish brown, smooth. Legs ferruginous; tibiae sparsely and finely spinose, the intermediate ones strongly arcuate (male), expanded at apex, pubescent beneath and prolonged into a spine as long as the spurs; anterior tarsi (male) with the first three joints strongly dilated and densely spongy beneath, the first joint campanulate, the second widest, quad- rate, the third strongly transverse, the fourth short, emarginate, two-thirds as broad as the third, with a few small spines and a trace of sponginess beneath, fifth joint narrow, cylindrical. Posterior coxae oval obtuse. Length, 12.5 mm.; width, 5 mm.” The above description was made from one male specimen, collected by Dr. G. Baur on Chatham Island, which I have made the type of the new species linelli. The type and only representative, as far as known, of this new form, is in the collection of the U. S. National Museum. Calosoma howardi Linell. One specimen, James Island, April 5, and one, South Seymour, April 23. Running about singly on soil. Both of these specimens are probably females. This species was described by Linell from ninety-two specimens: two from Duncan Island, twelve from Chatham Island, and seventy-eight from Charles Island. Collected by the Albatross Expedition in 1888 and 1891 and by Dr. G. Baur during 1891. The type of the species is in the U. S. National Museum. Tachys beebei sp. nov. (Fig. 42) Brownish black, antennae slightly longer than the head and thorax, three basal joints testaceous, darker at the extreme apex, fourth joint darker at the apical half, fifth to eleventh dark throughout. Palpi testaceous. Mandibles testaceous, margined with darker. Head narrower than the thorax, black, closely and somewhat finely punctate, labrum darker and more coarsely punctate than other parts of head; frontal grooves extending back beyond the middle of the eye. Pronotum black, subquadrate, wider at apex than base, widest at about the apical third, sides distinctly margined, front angles acute, slightly produced, hind angles obtuse; very finely and obsoletely punctate, basal im- pression moderately deep and outlined by a single row of punctures which begin at the basal margin about half-way between the angles and the middle and curve slightly towards the center, median impression indistinct, slightly 224 Zoological N. Y. Zoological Society [V ; 20 Fig. 42 .—TACHYS BEEBEI sp. nov. more distinct basally. Scutellum black. Elytra wider than the thorax, sub- lateral margins broadly pale, sutural portion black, the black covering about one-half the width of the elytra at the base, then becoming abruptly narrower, covering about one-third the width for the same distance longitudinally, from this point the black color gradually curves outwardly, the widest portion being slightly behind the middle where it again gradually curves inwardly to the suture at about the apical fifth, lateral margin dark to about the apical fifth; finely striate, the two inner striae more or less distinct, first dorsal puncture at the middle, second about one-fifth from the apex. Under surface black. Mouth-parts testaceous. Legs pale testaceous, coxae slightly darker. Length 2.25 mm. Described from eight examples from South Seymour, April, 1923. Col- lected under stones in damp mud around pools. Type No. 28055 and seven paratypes No. 28056. Coll. Amer. Mus. Nat. Hist. Eretes sticticus (Linnaeus). One specimen collected on Chatham Island, April 7. In small pools among lava blocks. This species was first listed, under the name Eunectes occidentalis Erichson, as occurring in the Galapagos Islands by Charles Waterhouse (1877, p. 77). 1924] Mutchler: Galapagos Coleoptera 225 The record is as follows: “six examples, which agree very well with the brief description of this species, Hab. Charles Island, Cookson.” The species was later listed by Sharp (1882, p. 697) as Dytiscus sticticus Linnaeus. It may be well to repeat the following remarks by Dr. Sharp: “Widely distributed in the warmer parts of the Old World, apparently rare in America. It is worthy of note that this species is found in a greater number of islands than any other of the Dytiscidae.” Linell, in his paper, did not refer to the species. Thermonectes basilaris (Harris). One male and one female specimen. South Seymour, April. In small pools among lava blocks. These specimens seem to be the same form as that listed by Charles Waterhouse (1877, p. 77) from Charles Island as Acilius incisus Aube var. (a synonym of Thermonectes basilaris Harris). The following differences noted by Waterhouse apply also to the specimens which I have before me: “The male agrees perfectly with that of A. incisus [ T . basilaris ]. The female differs in having the thorax more punctured, and in having the elongate punctures on the elytra much stronger and more close than in any examples of incisus [ basilaris ], and the punctures although diminishing in strength and density, extend nearly to the apex.” I was at first inclined to consider the specimens as representing a new form but, after reading Waterhouse’s remarks, I have come to the conclusion that it may be more advisable to consider them as belonging to the above species, as only the female varies from the typical form. I have compared the specimens with examples of basilaris in the American Museum collection and find that, although there is considerable variation in the density and depth of the punctures on the thorax and elytra of the females, none are as strongly punctured as in the specimens from the Galapagos. Tropisternus lateralis Fabricius. Three specimens, South Seymour, April 23. In small pools among lava blocks. This species has been recorded from North and South America, including the Antilles, and is abundant in the United States from New York southwards. It is also abundant in Mexico and Central America. Geo. R. Waterhouse recorded the species as collected by Darwin but did not state on what island it was found. Charles Waterhouse records it as collected on Charles Island by Darwin and Cookson and says that it appears to be common in the Galapagos. Bledius aequatorialis sp. nov. (Fig. 43) Male. — Elongate, testaceous. Antennae with basal joint elongate, second and third joints subequal in length, fourth joint shorter, fifth to tenth short, as wide or slightly wider than long, joint eleven longer, basal joints paler than the apical. Mandibles large, with a large tooth on the inner margin slightly before the middle. Head brownish black, front including labrum closely punctate, base more sparsely punctate, basal protuberances of antennae large and extending forward beyond the front margins of the eyes. Pronotum longer than wide, yellowish brown, slightly darker apically, somewhat coarsely 226 Zoologica: N. Y . Zoological Society [V; 20 punctate; middle of apex prolongated in a sharp, slightly downwardly curved horn, which reaches over the head slightly beyond the clypeal margin; apical angles narrowly rounded, basal angles very broadly rounded, disk with a narrow median f ovae extending from base to apex. Elytra short, a little longer than the thorax, yellowish, covered with a very short, fine pubescence. Exposed dorsal segments with somewhat long hairs on the lateral regions and a few scattered hairs on the disk. Under surface and legs pale yellow. Length 4 mm. Female. — Slightly darker than the male. Antennae not paler basally. Pronotum with apex squarely truncate. Otherwise as in the male. Length 4.5 mm. Described from two specimens collected under stones in moist mud around pools on South Seymour, April, 1923. Holotype male No. 28057 and allotype female No. 28058. Coll. Amer. Mus. Nat. Hist. Alloxacis seymourensis sp. nov. Elongate, chestnut-brown, sparsely covered with a pale pubescence. Right mandible bifid at the tip; apical part especially on the outer margins black, base testaceous. Palpi testaceous, apical joint of the maxillary dilated, tri- angular. Antennae inserted close to the eyes, testaceous, about one-half the length of the body; first joint clavate, slightly curved outwardly, second joint short, less than one-half the length of the first, third and following joints sub- equal in length, eleventh joint constricted at the apical half. Head finely and sparsely punctate, basal margin more or less faintly rugose, eyes narrowly emarginate at the front, moderately coarsely granulate; labrum transversely impressed just behind the apical margin. Pronotum widest about the middle, curved at the apical part, more or less oblique behind, being narrowest at the basal margin, basal margins prominent; surface moderately sparsely punctate, pubescence more dense at the sides. Elytra subparallel, indistinctly costate, punctures fine and not very closely placed, pubescence short and fine. Under surface generally slightly darker than the upper, finely and not thickly punctate, pubescence short and fine. Legs paler than the under surface, tibiae on all of the legs with two spurs at the apex, tarsi with fourth joint dilated and spongy beneath, claws with a small tooth at the base. Length 6 mm. Described from two specimens collected at South Seymour, April 23, on flowers of Cordia lutea. Type No. 28059 and paratype No. 28060. Coll. Amer. Mus. Nat. Hist. This species bears a superficial resemblance to the North American species of Oxacis, but on examination I find that the right mandible is bifid. It also has a well-defined, sharp tooth at the base of the claws similar to species of 1924] Mutchler: Galapagos Coleoptera 227 the genus Asclera, but I have followed Dr. Geo. R. Horn and considered that the species belongs in the genus Alloxacis from the fact that the right mandible is bifid and the left entire, which is the only character used in separating the genera Oxacis and Alloxacis . Coptostethus williamsi sp. nov. (Fig. 44) Female. — Wingless, dark brown, pubescent, with an intermixture of longer erect hairs. Antennae about as long as the head and thorax, serrate from the third joint onward, apex of last joint rounded, first joint thickened and about one-half longer than the second, third joint longer than the second, fourth and following joints shorter than the third and subequal in length. Head moder- ately finely punctate, inserted in the thorax to beyond the base of the eye. Thorax somewhat sparsely, coarsely punctate, sericeous pilose with large porrectly directed hairs, apical margin produced at the lateral angles, basal margin slightly bisinuate with lateral angles ending in a somewhat acute point, basal incisure strong. Scutellum cordiform. Elytra punctate-striate, each elytron with nine striae, interstices flat, slightly rugose, punctures large, elongate, deep; apical angles conjointly rounded. Underside of thorax coarsely and sparsely punctate. Abdominal segments more finely and more closely punctate. Legs slightly paler than the under surface. Tarsi with joints one to three of 228 Zoologica: N. Y. Zoological Society [V ; 20 approximately the same length, joints four and five each about two-thirds as long as joint three. Length 3 mm. Width 1.2 mm. Described from one specimen from South Seymour, collected April, 1923. Holotype No. 28061. Coll. Amer. Mus. Nat. Hist. Necrohia rufipes (De Geer). One specimen, Tower Island, April 23. This species is cosmopolitan. It was recorded by Geo. R. Waterhouse, under the generic name Corynetes, as collected by Darwin on James Island. Dermestes carnivorus Fabricius. Two specimens, South Seymour, April 23. This species is generally distributed over North and Central America. Linell records one example collected on Chatham Island by the Albatross Ex- pedition in 1891. Stomion laevigatum Geo. R. Waterhouse. Twenty-five specimens collected on Daphne Major, April 22, and sixty- nine collected on Tower Island during April. This species was very abundant under stones in bottom of crater and were associated with Ammophorusobscurus. Representatives of this lot were sent to Mr. K. G. Blair, of the British Museum, for comparison with the Waterhouse types. The following reply concerning the species was received from Mr. Blair: “ Your Stomion laevigatum do not quite agree with the type, in which the thorax is more constricted towards the base, and more convex towards the base, i.e. viewed sideways the thorax and elytra make more of an angle; the whole insect is decidedly narrower. The sculpture, however, seems to be identical and it seems that such differences as these are only sexual, at any rate yours seem to be both $ and mine both cf.” After receiving the above reply I examined the specimens which I had determined as this species and find that some of them have the thorax slightly less constricted toward the base or, if viewed sideways, the thorax and elytra make less of an angle. I also find that some specimens are broader than others, but in the whole series I cannot find any difference in sculpture on which to separate them into different forms and therefore believe that they all belong to the above species. Waterhouse, in the original description, did not mention on which island the species was collected. Stomion galapagoensis Geo. R. Waterhouse. There is one specimen collected on South Seymour in soil about roots of the large cactus, April 23, which agrees with the description of the above species in so far as it covers the parts described. The original diagnosis did not make any mention of the legs, which seem to be so different from other species ex- amined that I think it well to give the following short description of them: legs coarsely punctured and with bristles arising from the punctures. The bristles on the femora moderately coarse, those on the tibiae and tarsi intermixed with coarser, and the tarsal ones being relatively coarser than those on the femora or tibiae. Front tibiae with a somewhat decided inward curve on the inner side; middle and hind tibiae straight. 1924] Mutchler: Galapagos Coleoptera 229 This species was originally described by Waterhouse from two specimens collected by Darwin, but no mention was made to the island on which they were obtained. Linell records seven examples on Chatham Island, six by the Albatross Expedition, 1888, and one by Dr. G. Baur. I have compared the one specimen collected by the Williams Expedition with those determined by Linell, but our specimen does not agree with them in all respects. Further study and com- parison with the type material may show that either the Linell or our material represents a new form. Ammophorus obscurus Geo. R. Waterhouse. One hundred and seventy-eight specimens of this species were collected on Daphne Major during the month of April, under stones at the bottom of crater* associated with Stomion laevigatum. I was somewhat doubtful of my determination of this species, from the description, and therefore compared examples with the specimen recorded by Linell as collected on southern Albemarle Island and found that it was an entirely different form. This led me to send a representative of our lot to Mr. K. G. Blair, of the British Museum, with the request that he compare them with the Waterhouse types. The following reply was received from Mr, Blair: “ Am- mophorus obscurus is correctly determined. If you can let me see one of the Linell specimens I shall be glad to compare it with the type of other species.” The Linell specimen was later submitted to Mr. Blair who determined it as A. bifoveatus Geo. R. Waterhouse. Pedonoeces pubescens Geo. R. Waterhouse. One specimen collected on Tower Island, April 28. This specimen agrees with the description of the type from Chatham Island, in most of the essential points, but the color is more pitchy brown than piceous black and the antennae and legs are ferruginous instead of piceous black. Trox suberosus Fabricius. One specimen, Conway Bay, Indefatigable Island, April 1. Neither this genus nor species has heretofore been listed from these islands. The species has been recorded from North, Central, and South America, also from the West Indies and the Cape Verde Islands. Trox seymourensis sp. nov. Oblong-ovate, brownish black. Clypeus subangulate at the middle. Pronotum with surface roughly outlined, about one-third broader than long, sides curved and with an indentation near the basal angles, apex narrower than the base, apical margin produced at the sides, disk with an impression at each side which begins at the apical margin and extends obliquely from the apical angle to near the basal margin; central portion impressed. Scutellum longer than wide, sides subparallel, apex rounded. Elytra each with four rows of elongate tubercles slightly separated, thus forming interrupted carinae, intervals with three more or less interrupted rows of rounded tubercles, setae very short 230 Zoologica: N. Y. Zoological Society [V; 20 and sparse, being visible only under a high-power lens. Front femora broad basally, inner portion somewhat coarsely punctate and with a somewhat dense mat of brown hairs on the basal two-thirds, outer portion coarsely granulate- punctate, lateral margins fringed with stiff, bristle-like hairs, those on the outer margin arising from somewhat deep-set punctures. Front tibiae with a some- what large median tooth and with two smaller tooth-like projections on the basal half, the projection nearest the median tooth being the larger. Middle and hind legs more or less coarsely punctured and with a row of bristles on the margins. Length 10.5 mm. Described from one specimen collected on South Seymour, April 23. Type No. 28062. Coll. Amer. Mus. Nat. Hist. There is also in the collection one elytron taken on Tower Island, April 28, which is no doubt a fragment from a specimen of the same species. Stenodontes moliarius (Bates). One specimen. This species was listed by Howard as Mallodon sp.? The same specimens were identified by Linell as Mallodon ( Stenodontes ) moliarium, whose records are as follows: “The Albatross expedition in 1888 collected on Charles, Chatham, and Duncan islands seventeen examples of this large Prionid, which is distributed through Mexico and Central America to Panama.” Docema darwini sp. nov. Oblong, subparallel. Upper surface black, glabrous, shining. Antennae yellow with apical joints slightly darker; sparsely covered with a pale yellow pubescence which becomes more dense on the apical joints. Palpi dark. Eyes somewhat coarsely faceted. Head not noticeably punctate. A median carina extends from the frontal margin of the head to slightly beyond the bases of the antennae. At the back of the head is a V-shaped excavation, the apical portion of which connects with the basal end of the carina, thus forming a Y-shaped outline with the carina as the stem of the Y. Pronotum broader than long, slightly narrowed apically, finely but not very closely punctate, lateral margins somewhat broad, antebasal impression deep and extending to near the lateral margins. Scutellum transverse. Apex broadly rounded. Elytra moderately coarsely punctate on the basal half becoming finer apically, the punctures ar- ranged in more or less distinct rows; humeral angles somewhat prominent with a raised line extending from the umbone to the apical third; lateral margins well developed, epipleurae broad at the base, gradually narrowing and becoming obsolete at the apex. Under surface black, sparsely covered with pale pubes- cence. Legs brown, sparsely covered with pale pubescence; femora generally dark brown, almost black in some specimens, tibiae and tarsi paler than femora, apical tarsal joints darker in some examples. Length 2.2-2. 5 mm. Described from fifteen specimens collected on Tower Island during April and one specimen from Eden, April 4. All taken on low bushes along the beach. Type No. 28063 and paratypes 28064. Coll. Amer. Mus. Nat. Hist. Pantomorus galapagoensis Linell. One male and two females, Conway Bay, Indefatigable Island, April 1. 1924] Mutchler: Galapagos Coleoptera 231 Fig. 45 . — NEOPEN TAR THR UM TO WERENSIS sp. nov. Described by Linell from one male and three females, collected on Chatham Island. The type of this species is in the U. S. National Museum. Neopentarthrum gen. nov. Rostrum moderately long and somewhat broad, slightly arched, apical part at sides in front of antennae parallel; basal part slightly constricted; scrobes oblique, extending backwards to the under margins of the eyes. Antennae short, inserted in front of the middle, scape moderately short, club-shaped; funicle five-jointed, first joint thick, about as long as the second and third taken to- gether, joints two, three, and four gradually shorter and not as thick as the first joint, the five joints with bristle-bearing punctures; club moderately large and as long as joints two, three, and four combined, pubescent, apex with two or more annulations. Eyes moderately large, situated at the base of the rostrum, convex, facets moderately coarse. Prothorax narrowed and constricted in front, more gradually narrowed behind. Scutellum small, distinct, rounded at apex. Elytra oblong, wider at base than the thorax, subparallel to apical third where they become slightly but gradually narrower, apices rounded. Legs short and stout, femora inflated, tibiae slightly triangular with prominent hooks at the apex. Tarsi with first joint somewhat elongate, second joint short, third joint deeply emarginate and a little longer than the second, claw joint about as long as the second and third combined. Body beneath slightly convex, glabrous. I have erected this genus to include a new species, toiverensis, of Cossoninae found in the Galapagos Islands by the Williams Expedition. There is, seem- ingly, no genus in the pentarthrid group which agrees with the generic characters found in this species. Neopentarthrum towerensis sp. nov. (Fig. 45) Subcylindrical, piceous, shining. Antennae short and thick, scape short, clavate, funicle five jointed, first joint large; other joints closely united and gradually smaller, club slightly paler than the other part, appearing annulate under a high power. Beak moderately densely and somewhat finely punctate. 232 Zoologica : N. Y. Zoological Society [V; 20 Fig. 4,6.— PLATYPUS SANTACRUZENSIS sp. nov. Eyes situated at base of beak, small, slightly convex. Pronotum slightly longer than wide, apex constricted behind the apical margin; from this constriction the sides gradually curve outwardly to about the middle where they again grad- ually curve inwardly to the base; disk moderately coarsely punctate, those on the central portion being the coarser. Scutellum small, rounded at apex. Elytra slightly wider than the base of the thorax, cylindrical, surface feebly striate, striae with coarse punctures, intervals flat, each with a row of small punctures. Body beneath somewhat coarsely and sparsely punctured. Legs short, femora swollen apically; tibiae broadening apically ,apex with the usual claws; tarsi with the basal joint longer than the second, second and third about equal in length, claw joint nearly as long as the other three taken together. Front coxae narrowly separated. Length 2-2.75 mm. Described from seven specimens taken on Tower Island, April 28. Bur- rowing in numbers under bark of bushes. Type No. 28065 and paratypes No. 28066. Coll. Amer. Mus. Nat. Hist. Platypus santacruzensis sp. nov. (Fig. 46) Male. — Elongate, cylindrical, flavous. Mandibles with apex and outer margin darker. Head slightly wider than the thorax, sparsely covered with paler hairs; front flat, sloping forward, moderately coarsely and closely granu- late-punctate; basal portion smooth, median black line extending from the basal 1924] Mutchler: Galapagos Coleoptera 233 margin to the sloping portion. Pronotum longer than wide, very finely but not closely punctate, and with a few scattered hairs along the apical and front part of the lateral margins; disk with a short basal median line. Scutellum strongly depressed and acuminate. Elytra somewhat deeply striate-punctate and with a few scattered hairs which become more closely placed at the lateral margins and at the apex; apical part dark, narrowed, sutural angle squarely truncate, lateral angle prolonged, apex tridentate. Under surface, except abdominal segments, finely and somewhat sparsely punctate, abdominal segments more closely and coarsely punctate, sparsely covered with somewhat stiff hairs. Front tarsal joint prolonged as usual in Platypodidae. Length 4.75 mm. Described from one specimen which was collected while it was flying along the shore of Seymour Bay, Indefatigable Island. Holotype No. 28067. Coll. Amer. Mus. Nat. Hist. LIST OF COLEOPTERA KNOWN TO OCCUR IN THE GALAPAGOS ISLANDS The following list contains, as far as I have been able to ascertain, all of the species of Coleoptera4 which have been reported as being found in the Galapagos Islands. There are seventy-three species included therein, or more than twice as many as Wallace5 6 * recorded in his reference to the insects and land shells found in this archipelago. His remarks on the paucity of the insect fauna are as follows: “The insects are very scanty; the most plentiful group, the beetles, only furnishing thirty-five species, belonging to twenty-nine genera and eighteen families. The species are almost all peculiar, as are some genera. They are mostly small and obscure insects, allied either to American or world wide groups. The Carabidae and Heteromera are the most abundant group, the former furnishing six and the latter eight species.”8 The numerical proportions of groups are not very different in the present list but the Carabidae are fewer as, in the addition of forty forms this family has furnished only five species, or slightly more than twelve per cent. The Tenebrionidae (Heteromera of Wallace) has furnished eight additional forms, or twenty per cent of the total addition. The species which have been added to the list give us an addition of only three families (Wallace lists only eighteen families but he included, as the families are now known, three in the Malacodermes and three in the Necrophaga) but they have raised the number of genera to fifty-five, or an addition of twenty- two (some of the species listed by Wallace as belonging in one genus were found to belong in two or more genera). There has been much discussion as to the introduction of animals on these islands, whether by land bridges or on floating masses. 4 I have not considered the California Academy of Sciences material as this material has not been definitely determined. 6 ‘Island Life,’ 1880. In this paper Wallace lists thirty-seven species, taken from the records of Waterhouse, 1877, Proc. Zool. Soc. London. 6 Wallace did not include Calosoma galapageium Hope in his list of Carabidae. He also omitted Stomion galapagoensis Geo. R. Waterhouse, Ammophorus galapagoensis Geo. R. Waterhouse, Pedonoeces costatus Geo. R. Waterhouse and Pedonoeces morio Geo. R. Water- house in his list of Heteromera. 234 Zoologica : N. Y. Zoological Society [V; 20 It is noteworthy (1) that even the present enlarged list includes but a small part of the fauna to be expected if there had been a land bridge and (2) that the habits of the species now on the islands are such as would be ex- pected of insects that had been introduced either by floating masses or by man. In this connection the following theory expounded by Wallace may prove interesting: “The observation of Captain Collnet, quoted by Mr. Darwin in his Journal, that drift-wood, bamboos, canes, and the nuts of a palm, are often washed on the southeastern shores of the islands, furnishes an excellent clue to the manner in which many of the insects and land-shells may have reached the Galapagos. Whirlwinds also have been known to carry quantities of leaves and other vegetable debris to great heights in the air, and these might be then carried away by strong upper currents and dropped at great distances, and with them small insects, and mollusca, or their eggs.” The probable introductions, if any, through the agency of man are the scavenger and wood-boring forms. The reason for including the latter here is that six conspicuous species of Cerambycidae appear to be of recent introduction, as they were not reported by the earlier writers and these beetles would almost certainly have been found, had they been there. They were probably taken over in wood which formed a part of boat’s cargo or of the boats themselves. The references to literature merely refer to the original description of the species and such papers which treat on their distribution in the Galapagos Islands. Species marked by the asterisk are those which are not indigenous to the Galapagos Islands. Family CICINDELIDAE Cicindela galapagoensis W. Horn, 1915, pp. 238, 241, 251, 397, 399 and 402. 1920, p. 17. Listed as Cicindela sp.? by F. X. Williams, 1907, p. 260. Family CARABIDAE Calosoma galapageium Hope, 1837, p. 130. Calosoma howardi Linell, 1898, p. 251. Listed by Howard, 1889, p. 191, as Calosoma galapagoum? Hope. Calosoma linelli sp. nov. Listed as Calosoma galapageium Hope, by Linell, 1898, p. 250. Scarites galapagoensis Linell, 1898, p. 253. Tachys beebei sp. nov. Pterostichus calathoides (Geo. R. Waterhouse), 1845, p. 21. Described under the generic name of Feronia. Listed by Charles Waterhouse, 1877, p. 82, as Feronia. Howard, 1889, p. 191, listed under the generic name Poecilus. Lined, 1898, p. 252, listed under the generic name Pterostichus. Feronia insulars Boheman, 1858, p. 14. Charles Waterhouse, 1877, p. 82. Lined, 1898, p. 255. Notaphus galapagoensis (Geo. R. Waterhouse), 1845, p. 23. Charles Water- house, 1877, p. 81. Lined, 1898, p. 255. Amblygnathus obscuricornis Geo. R. Waterhouse, 1845, p. 22. Charles Water- house, 1877, p. 82. Lined, 1898, p. 255. Platynus galapagoensis (Geo. R. Waterhouse), 1845, p. 21. Described under 1924] Mutchler: Galapagos Coleoptera 235 the generic name of Feronia. Listed by Charles Waterhouse, 1877, p. 82, as Feronia. Linell, 1898, p. 252, lists it under the generic name of Platynus. Selenophorus galapagoensis Geo. R. Waterhouse, 1845, p. 22. Charles Water- house, 1877, pp. 77 and 82. Howard, 1889, p. 191. Linell, 1898, p. 254. Family DYTISCIDAE Copelatus galapagoensis Geo. R. Waterhouse, 1845, p. 23. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 255. * Thermonectes basilaris (Harris), 1829, p. 8. Listed by Charles Waterhouse, 1877, pp. 77 and 82, as Acilius incisus Aube var. * Eretes sticticus (Linnaeus), 1766, p. 666. Listed by Charles Waterhouse, 1877, pp. 77 and 82, as Eunectes occidentalis Erichson. Sharp, 1882, p. 699, lists it as Dytiscus sticticus. Family HYDROPHILIDAE * Tropisternus lateralis (Fabricius), 1775, p. 228. Geo. R. Waterhouse, 1845, p. 26. Charles Waterhouse, 1877, pp. 78 and 82. Linell, 1898, p. 255. Philhydrus species? Geo. R. Waterhouse, 1845, p. 26. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 255. Family SILPHIDAE Acribis serrativentris Charles Waterhouse, 1877, pp. 78 and 82. Family STAPHYLINIDAE Bledius aequatorialis sp. nov. * Creophilus villosus (Gravenhorst), 1802, p. 160. Geo. R. Waterhouse, 1845, p. 24, as Creophilus sp. Charles Waterhouse in 1877, p. 82, records the species as villosus. Linell, 1898, p. 255, not knowing of the Waterhouse article, says: “This is probably Creophilus villosus Gravenhorst, introduced from North America.” Family MELYRIDAE Ablechrus flavipes Charles Waterhouse, 1877, p. 79. [Listed by Charles Water- house, 1877, p. 81, as Ablechrus darwinii.] Family CORYNETIDAE * Necrobia rufipes (De Geer), 1775, p. 165. Geo. R. Waterhouse, 1845, p. 26 and Charles Waterhouse, 1877, p. 81, refer to the genus as Corynetes. Linell, 1898, p. 257. Family OEDEMERIDAE Oxacis galapagoensis Linell, 1898, p. 266. Alloxacis seymourensis sp. nov. Family ELATERIDAE Anchastus galapagoensis Geo. R. Waterhouse, 1845, p. 25. Charles Waterhouse, 236 Zoologica : N. Y. Zoological Society [V; 20 1877, p. 82. Linell, 1898, p. 256. [Described by Geo. R. Waterhouse and listed by Charles Waterhouse and Linell under the generic name Physo - rhinus.] Heterocrepidius puberulus Boheman, 1858, p. 66. Linell, 1898, p. 256. Coptostethus williamsi sp. nov. Family DERMESTIDAE * Dermestes carnivorus Fabricius, 1775, p. 55. Linell, 1898, p. 256. * Dermestes vulpinus Fabricius, 1781, p. 64. Geo. R. Waterhouse, 1845, p. 26. Charles Waterhouse, 1877, p. 81. Linell, 1898, p. 256. Family PHALACRIDAE Phalacris darwinii Charles Waterhouse, 1877, pp. 78 and 82. Family COCCINELLIDAE Scymnus galapagoensis Geo. R. Waterhouse, 1845, p. 41. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 256. Family ALLECULIDAE Lobopoda galapagoensis Linell, 1898, p. 266. This species was listed by Howard, 1889, p. 192, with the following remark: “Two specimens of a species Allecula probably new from Charles Island.” Family TENEBRIONIDAE Stomion galapagoensis Geo. R. Waterhouse, 1845, p. 29. Charles Waterhouse, 1877, pp. 79 and 82. Howard, 1889, p. 192. Linell, 1898, p. 262. [Ac- cording to determinations made by Mr. K. G. Blair Stomion piceum Linell, 1898, p. 262, and Stomion carinipenne Linell, 1898, p. 262, also belong to this species.] Stomion bauri Linell, 1898, p. 263. Stomion helipioides Geo. R. Waterhouse, 1845, p. 30. Charles Waterhouse, 1877, p. 82. Howard, 1889, p. 192. Linell, 1898, p. 263. Stomion laevigatum Geo. R. Waterhouse, 1845, p. 30. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 263. Ammophorus galapagoensis Geo. R. Waterhouse, 1845, p. 30. Charles Water- house, 1877, p. 82. Linell, 1898, p. 263. Ammophorus bifoveatus Geo. R. Waterhouse, 1845, p. 31. Charles Waterhouse, 1877, p. 81. Howard, 1889, p. 192. Linell, 1898, p. 263 [and in error as A. obscurus on p. 264]. Ammophorus cooksoni Charles Waterhouse, 1877, pp. 80 and 82. Ammophorus caroli Linell, 1898, p. 264. [This species is without much doubt synonymous with cooksoni .] Ammophorus obscurus Geo. R. Waterhouse, 1845, p. 32. Charles Waterhouse, 1877, p. 82. [The Linell record of this species, based on the determination made by Mr. K. G. Blair, has been credited to A. bifoveatus .] Pedonoeces galapagoensis Geo. R. Waterhouse, 1845, p. 35. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 265. 1924] Mutchler : Galapagos Coleoptera 237 Pedonoeces costatus Geo. R. Waterhouse, 1845, p. 35. Linell, 1898, p. 265. Pedonoeces mono (Boheman), 1858, p. 92. [Described under the generic name of Tessaromma .] Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 265. Pedonoeces pubescens Geo. R. Waterhouse, 1845, p. 36. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 265. Pedonoeces bauri Linell, 1898, p. 265. * Gnathocerus cornutus (Fabricius), 1798, p. 51. Linell, 1898, p. 266. Phaleria manicata Boheman, 1858, p. 92. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 266. Family BOSTRICHIDAE * Tetrapriocera longicornis (Olivier), 1790, p. 15. Howard, 1889, p. 191, Lists this species with the following note: “One specimen of Tetrapriocera was collected on Indefatigable Island. Although the same habitus it is probably different from our Florida species, T. longicornis Oliv., which is known to have a wide distribution in Central and South America.” Linell, 1898, p. 256. * Amphicerus cornutus (Pallas), 1772, p. 18. Linell, 1898, p. 256, as A. puncti - pennis. Amphicerus cornutus subspecies galapaganus Lesne, 1910, pp. 183-186. The first record of the representatives of this species was based on three speci- mens found by Charles Darwin on the dead branches of a Mimosa tree in Chatham Island. They were listed by Geo. R. Waterhouse, 1845, p. 36, as belonging in the genus Apate but no specific designation was given to them. This same series of specimens was later recorded by Charles Water- house, 1877, p. 82, as Bostrichus unicantus with a footnote, referring to the above, which reads as follows: “The Apate mentioned by Waterhouse, Ann. and Mag. Nat. Hist., 1845, XVI, p. 36.” Linell, 1896, p. 256, not knowing of the determination by Charles Waterhouse, combined the records of Darwin’s specimens with a specimen collected by Dr. G. Baur on Albe- marle Island, which Linell determined as Amphicerus punctipennis = cornutus. The Darwin specimens were later studied by Lesne, who de- scribed them as Schistoceros = Amphicerus cornutus subspecies galapaganus. In his description of this new form Lesne says that he has not seen the specimen collected by Dr. G. Baur but has studied the four (2 male and 2 female) specimens [Waterhouse lists only three specimens] collected by Darwin. In his article Lesne makes no mention of Charles Waterhouse’s determination of the species. The specimen collected by Dr. G. Baur may also belong in this sub- species but, as it has not been determined as such, I have followed Linell and listed it as cornutus. Family SCARABAEIDAE Copris lugubris Boheman, 1858, p. 42. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 258. Parapseudoryctes galapagoensis (Geo. R. Waterhouse), 1845, p. 26. Charles Waterhouse, 1877, p. 82. Howard, 1889, p. 191. Linell, 1898, p. 258, 238 Zoologica: N. Y. Zoological Society [V; 20 described the genus Pseudorydes for this species, the former authors having listed it under the genus Orydes. I have herewith changed the generic name to Parapseudorydes as the name Pseudorydes is a homonym, it having been used by Sharp (1873, Rev. et Mag. Zool., (3) I, p. 267) for species found in Australia, which are not congeneric with the Galapagos forms. * Trox suberosus Fabricius, 1775, p. 31. [The record which I have embodied in this paper is, to my knowledge, the first from these islands.] Trox seymourensis sp. nov. Family PASSALIDAE * Neleus tlascala Percheron, 1835, p. 45. Linell, 1898, p. 257. Family CERAMB Y CIDAE * Stenodontes moliarius (Bates), 1879, p. 9. Howard, 1889, p. 191, as Mai - lodon sp.? Linell, 1898, p. 259, under the generic name Mallodon. Achryson galapagoensis Linell, 1898, p. 259. Eburia lanigera Linell, 1898, p. 259. Eburia bauri Linell, 1898, p. 260. Eburia amabilis Boheman, 1859, p. 150. Charles Waterhouse, 1877, p. 82. Howard, 1889, p. 192. Linell, 1898, p. 261. Acanthoderes galapagoensis Linell, 1898, p. 261. Family CHRYSOMELIDAE Docema galapagoensis Geo. R. Waterhouse, 1845, p. 39. Described as Haltica galapagoensis. Charles Waterhouse, 1877, pp. 81 and 82. [In this paper Charles Waterhouse erects the genus Docema for this species.] Linell, 1898, p. 262, not knowing of the Charles Waterhouse paper, lists the species under the genus Haltica. Docema darwini sp. nov. Longitarsus lunatus Charles Waterhouse, 1877, p. 81. Diabrotica limbata Charles Waterhouse, 1877, pp. 81 and 82. Family PLATYSTOMIDAE Ormiscus variegatus Geo. R. Waterhouse, 1845, p. 37. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 268. Family CURCULIONIDAE Otiorhynchus cuneiformis Geo. R. Waterhouse, 1845, p. 38. Charles Water- house, 1877, p. 82. Linell, 1898, p. 267. Pantomorus galapagoensis Linell, 1898, p. 268. [This species was listed but without a name by Howard 1889, p. 192, with the following remark: “Three specimens belonging to this family [Curculionidae] were collected on Chatham Island.”] Anchonus galapagoensis Geo. R. Waterhouse, 1845, p. 39. Charles Waterhouse, 1877, p. 82. Linell, 1898, p. 268. Neopentarthrum towerensis sp. nov. 1924] Mutchler: Galapagos Coleoptera 239 Family PLATYPODIDAE Platypus santacruzensis sp. nov. Family SCOLYTIDAE Linell, 1898, p. 268, records a single specimen without elytra, belonging in the group Hylurgi. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoological Society. Its title is “Galapagos; World’s End.” BIBLIOGRAPHY Bates, H. W. 1879 ‘Biol. Centr. Amer. Coleoptera,’ V, [1879-1886]. Boheman, C. H. 1858 and 1859. ‘Kongliga Svenska Fregatten Eugenies,’ Zoologi I, Insecta, [1858-1868]. De Geer, Carl. 1775 ‘ Memoirs pour servir a l’historie des Insectes,’ V. Fabricius, J. Christ. 1775 ‘Systema Entomologie.’ 1781 ‘Spec. Ins.,’ I. 1798 ‘Suppl. Ent. System.’ Gravenhorst, J. L. C. 1802 ‘Coleoptera Microptera Brunsvicensia.’ Harris, T. W. 1829 ‘New England Farmer.’ Hope, F. W. 1837 Trans. Ent. Soc. London, II. Horn, Walther. 1915 ‘Genera Insectorum,’ fasc. 82c. 1920 Arkiv for Zoologi, XIII, No. 11. Howard, L. O. 1889 Proc. U. S. National Museum, XII, [1890]. Lesne, P. 1910 Bulletin du Museum d’Histoire Naturelle, XVI. Linell, Martin. 1898 Proc. U. S. National Museum, XXI, [1899]. Linnaeus, C. 1766 ‘Systema Natura,’ 12th Ed., I, part 2. 240 Zoologica: N. Y. Zoological Society [V ; 20 Olivier, G. A. 1790 ‘Entomologie,’ IV, No. 77. Pallas, Peter S. 1772 ‘Specilegia Zoologica,’ fasc. 9, [1767-1774]. Percheron, A. 1835 * Monographie des Passales.’ Sharp, David. 1882 Trans. Royal Dublin Soc., (2) II, [1880-1882]. Waterhouse, Charles. 1877 Proc. Zool. Soc. London. Waterhouse, Geo. R. 1845 Ann. Mag. Nat. Hist., XVI. Williams, F. X. 1907 Entomological News, XVIII. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY DEPARTMENT OF TROPICAL RESEARCH WILLIAMS GALAPAGOS EXPEDITION VOLUME V. NUMBER 21 Department of Tropical Research Contribution Number 187 HEMIPTERA-HETEROPTERA FROM THE WILLIAMS GALAPAGOS EXPEDITION By H. G. Barber PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK March 10, 1925 Nrw fork Ifimlogtral Swirly General Office: 101 Park Avenue , New York City GDfTtmB President, Madison Grant; Vice-Presidents, Frank K. Sturgis and Henry D. Whiton; Chairman, Executive Committee, Madison Grant; Treasurer, Cornelius R. Agnew. laard of iHattag^ro CElaBB nf 192H Henry Fairfield Osborn, Lispenard Stewart, Charles F. Dieterich, George F. Baker, Wm. Pierson Hamilton, Robert S. Brewster, Edward S. Harkness, William B. Osgood Field, Edwin Thorne, Percy A. Rockefeller, John E. Berwind, Irving K. Taylor. aka* of 192r Madison Grant, Wm. White Niles, Frank K. Sturgis, Ogden Mills, Lewis R. Morris, Archer M. Huntington, George D. Pratt, Coleman du Pont, Henry D. Whiton, Cornelius R. Agnew, Harrison Williams, Marshall Field. Cla00 of 1938 Percy R. Pyne, George Bird Grinnell, Cleveland H. Dodge, Anthony R. Kuser, Mortimer L. Schiff, Frederic C. Walcott, George C. Clark, Jr., W. Redmond Cross, Henry Fairfield Osborn, Jr., Arthur A. Fowler, George Gordon Battle, Bayard Dominick. S>rinttiftr iStaff William T. Hornaday, Director of the Zoological Park; W. Reid Blair, Assistant to the Director, and Veterinarian; Charles H. Townsend, Director of the Aquarium; Raymond L. Ditmars, Curator of Reptiles; William Beebe, Honorary Curator of Birds and Director of Department of Tropical Research; Lee S. Crandall, Curator of Birds; George S. Huntington, Prosector; H. C. Raven, Associate Prosector ; Elwin R. Sanborn, Photographer and Editor . SE&itaml (fammittr? Madison Grant, Chairman; William T. Hornaday, Charles H. Townsend, William Beebe, Elwin R. Sanborn, Sec’y. Zoologica, Vol. V, No. 21 HEMIPTERA-HETEROPTERA1 FROM THE WILLIAMS GALAPAGOS EXPEDITION By H. G. Barber (Figs. 47-50 incl.) The Hemiptera-Heteroptero us fauna from the Galapagos Islands so far as known is rather limited. I have been able to find only twenty-one species so far listed from the islands and two species of Halobates from neighboring waters. Twelve of these are restricted to the islands and ten are also neotropical species. From the col- lections made by William Beebe on this expedition of the New York Zoological Society, I have been able to add nine more species to the island fauna, seven of which are new, and one new species of Halo- bates. The drawings have been made by myself. Family PENTATOMIDAE Acrosternum ( Nezara ) viridans Stal. Acrosternum ( Nezara ) viridans Stal. Stal, Freg. Eugen. Resa-Insects, 228, 1859; Enum., II, 41, 1872. Howard, Proc. U. S. Nat. Mus., XII, 194, 1889. Heidemann, Proc. Washington Acad. Sci., Ill, 365, 1901. Two from James Island, April 5, and two from Chatham Island, April 7. This has been recorded by Stal also from Panama and Peru. Podisus ( Armd ) sordidus Stal. Podisus ( Arma ) sordidus Stal. Stal, Freg. Eugen. Resa-Insects, 221, 1859; Enum. I, 51, 1870. Two from Indefatigable Island, April 22. Also known from Peru. Family COREIDAE Stenocephalus insularis Dallas. Stenocephalus insularis Dallas. Dallas, List Hem. II, 482, 1852. Stal, Enum. I, 218, 1870. Heidemann, Proc. Washington Acad. Sci., Ill, 365, 1901. Ten from James Island, April 4 and 5, and Albemarle Island, April 6. Dallas described this species from material obtained by Charles Darwin on his voyage of the Beagle. Harmostes disjunctus sp. nov. ( = H. serratus Heidemann nec Fabr.) Color yellow-brown, with paler costal margin of corium and the femora spotted with brown; membrane obscurely spotted. Head finely granulate, as 1 Cont. Department Trop. Research No. 187. 241 Plate A. SKETCH MAP OF GALAPAGOS ISLANDS Route of the Noma, and details and location of the Archipelago. 242 1924] Barber: Hemiptera-Heteroptera 243 long as wide; tylus elevated, acute, about attaining middle point of the first antennal segment; antenniferous tubercles acute, nearly porrect, slightly turned outwards towards apex, extended forward on a line with the apices of the jugae; vertex with a narrow median sulcus extending from line of the ocelli to near base of the tylus. Antennae 3.75 mm. long, with granulated basal seg- ment stout, 1.5 mm. long; second and third segments much more slender, cylindrical, neither especially incrassate at their apices, second segment 1 mm., third 1.5 mm. long; piceous terminal segment spindle-shaped, one-half as long as third segment. Head paler beneath with a piceous streak behind the buc- culae, the latter disappearing on a line with the front margin of the eyes. Rostrum extending only to the anterior part of the posterior coxae, the basal segment reaching to a line drawn across posterior margins of the eyes. Pro- notum only a trifle shorter than head and just twice as wide across humeral angles as it is long; lateral margins slightly and evenly arcuate, the edge reflexed and provided with a few rather weak, pale teeth, the anterior ones forming an acute anteriorly directed process; surface, except for the cicatrices and along the narrow margin before these, closely, almost confluently, coarsely punctate; provided with a median longitudinal pale carina; humeral angles rounded or obtusely angled, not projecting much beyond costal margin of corium; humeri somewhat roundly elevated within and these connected by a somewhat evident transverse ridge, placed just before the posterior margin of the pronotum. Scutellum more shallowly punctate; apical part narrow, evenly rounded, with the edge pale, elevated, the surface within scooped out. Corium and clavus concolorous or the former slightly variegated with pale brown ; these parts evenly, coarsely, almost confluently punctate; the pale impunctate costal margin reflexed and provided with 11 to 12 almost equi-distant dark brown spots; the outline of the costal margin, viewed from above, scarcely at all concavely sinuate, the greatest width across the hemielytra being two-thirds the distance from the base; apex of the corium, at least in the male, reaches to the apex of the abdomen. Membrane hyaline, with scattered indistinct brownish spots. Beneath paler than above. Central disk of sternites with a piceous spot, widest on the mesosternum; meso- and meta-pleura closely, shallowly punctate with faint indications of darker maculations. Venter pale yellow-gray, with numerous rosy red spots and with a short transverse piceous fascia on the sides of the 4th, 5th and 6th segments, placed closer to the lateral margins than to the median line; anteriorly very obscurely sulcate in the middle. Legs pale ochraceous, the femora, except at base, much spotted with ferrugineous-brown, the posterior ones about twice the diameter of the others and armed beneath with three strong white spines with smaller, close set, dark spines between the distal two; apex reaches back only as far as the apex of the corium. Length male 6 mm. Type and paratype males: Indefatigable Island. This species is very close to H. affinis described by Dallas from an unknown locality and fixed by Van Duzee in 1909 for the Florida species. I have Florida as well as numerous West Indian specimens for comparison. There is no doubt that this Galapagos species is what Heidemann, 1901, misidentified as serratus Fab. I have seen Heidemann’s specimen in the collection of U. S. N. M . 244 1924] Barber: H emitter a-Heteropter a 245 H. disjunctus differs from serratus especially by its more quadrate head, shorter rostrum, less pronounced spinous process of tylus and antenniferous tubercles. From a finis it may readily be separated by the following characters: the different relative lengths of the second and third antennal segments, straighter costal margins, corium reaching to apex of the abdomen, shorter posterior femora and almost non-sulcate venter. By its denticulate pronotal margin it differs from the following South American forms: raphimerus Spin., minor Spin., marmoratus BI., apicatus Stal and prolixins Stal. Corizus hyalinus Fabricius. Corizus hyalinus Fagricius. Fabricius, Ent. Syst. 168, 1794. Four from Daphne Major April 22, and three from Tower Island April 28. These are all dark specimens of this widely distributed species, not hitherto reported from these islands. C. lugens Stal is unknown to me. Nysius ( Cymus ) marginalis Dallas. Nysius {Cymus) marginalis Dallas. Dallas, List Hem. II, 556, 1852. Stal, Freg. Eugen. Ins., 252, 1858; Enum. IY, 122, 1874. Butler, Proc. Zool. Soc. London, 88, 1877. Heidemann, Proc. Washington Acad. Sci., Ill, 366, 1901. Eight from Daphne Major, April 22. So far as known this is an endemic species. Heraeus pacificus sp. nov. (Fig. 47) Color dull fusco-ferrugineous, with anterior lobe of pronotum, scutellum and sternum black; posterior lobe of pronotum with four streaks and humeral angles brown; corium with narrow costal margin to just beyond middle, a sub- apical spot, two others near posterior margin and the inner field fasciate with testaceous; membrane brown with pale veins. Head and anterior lobe of pronotum with a few long setae. Head equally long as pronotum and slightly narrower across eyes than width of posterior lobe of pronotum, transversely finely granulose with sparse covering of very fine appressed, grayish pubescence; apex of tylus reaching midway on basal antennal segment, anteocular distance to apex of antenniferous tubercles two-thirds the length of an eye, postocular region gradually contracted. Antenna with extreme base of second and third segments pale, slightly enlarged basal segment but slightly longer than the anteocular part of head, second segment about twice as long as first and one- third longer than third, fourth segment subequal to second; all segments except basal with fine short hairs. Rostrum reaching to middle of hind coxae, the enlarged basal segment dark castaneous, remainder, except at apex, pale; first segment reaching mid-way on postocular region of head, second segment one- third longer, third a little shorter than second. Pronotum as long as head, with the two lobes of nearly equal length, the anterior one dull black provided with a few erect hairs, posterior lobe sparsely, finely punctate, one-third wider than anterior lobe; rounded humeri and four longitudinal streaks paler. Scutellum dull black, disk somewhat paler either side of the middle, extreme apex tes- taceous. Hemielytra non-pilose; clavus finely, closely punctate with outward pale, smooth ruga reaching nearly to apex; corium sparsely punctate between the veins. Membrane dark brown with distinct pale veins. Sternum dark 246 Zoological N. Y. Zoological Society [V; 21 with rim of all acetabuli and the posterior flange of the meta-pleura ferrugineous, enlarged fore femora shining dark castaneous with only the apex pale, armed beneath with several strong teeth beyond the middle; fore tibia slightly curved, pale stramineous, infuscated at apex; middle and hind femora pale stramineous with a pre-apical castaneous ring; tibiae pale, infuscated at tip; all tarsi pale, infuscated apically. Venter ferrugineous with coating of fine appressed pale hairs. Length 6 mm. Type. — Male, James Island, April 4; allotype, — James Island, April 4; two paratypes, male and female, James Island, April 4. This species is rather closely related to H. plebejus Stal, but is darker in color and less pilose, the an- tennae longer and the diameter of corium relatively wider. Orthaea insularis sp. nov. (Fig. 48) Sparsely setose on head, pronotum and scutellum. Dull black. Hemi- elytra with somewhat more than anterior half, a preapical subcrescentic fascia and wide posterior margin of meta-pleura sordid whitish; remainder of the corium fusco-ferrugineous with the area near inner apical angle frequently ferrugineous. Membrane heavily infuscated, with a broad, pale patch at apex. Legs cas- taneous with the intermediate and posterior femora pale at base and the tarsi testaceous. Head faintly rugulose with fine appressed gray hairs, equally long as width across eyes, one-fifth shorter than anterior lobe of pronotum; tylus rufous, reaching midway on basal segment of antennae. Antennae testaceous, with terminal segment somewhat darker, second segment twice as long as first, third, one-third to one-fourth shorter, fourth segment very nearly as long as second. Rostrum castaneous, basal segment darker, apex of fourth segment reaching just past intermediate coxae. Pronotum with anterior lobe dull black, sparsely setose, twice as long and one-third narrower than the posterior lobe, the latter dark velvety-brown, anteriorly next to the sinus some- what pruinose and there with a few scattered punctures; humeri concolorous; posterior margin very concave before the base of the scutellum. Sternum dull black with the posterior expanded margin of the metasternum white. Legs with the anterior femora much incrassate, more so in the male, armed except for a short distance at base with 8-10 short, stout teeth arranged somewhat in a double series; tibia and tarsi paler. Intermediate and hind legs with nearly basal half of femora and the trochanters whitish, remainder of femora and the tibia castaneous, with the tarsi sordid stramineous. Scutellum dull black, sparsely punctate, apical half depressed. Hemielytra with clavus fuscous at base and apex and provided with several rows of fuscous punctures; the inner longitudinal area sometimes smudged with fuscous; the anterior whitish part of the corium sparsely punctate with fuscous and with an abbreviated infuscated patch midway along the claval suture; the preapical crescentic whitish fascia reaching to the costal margin and provided with a few scattered fuscous punc- tures; the broad fusco-ferrugineous fascia sharply defined anteriorly and about* the preapical fascia. Membrane dark smoky-brown, with the veins towards base and a broad patch occupying nearly the whole apex beyond the middle, Fig. 48. ORTHAEA INSULARIS sp. nov. 247 248 Zoologica: N. Y. Zoological Society [V; 21 pale. Venter shining castaneous with sparse covering of fine appressed hairs. Length 6.5 mm. Type: Male and allotype, James Island, April 4, 1923. Paratypes: three males and nine females from James Island, male and female from South Seymour Island, female from Albemarle Island and two nymphs from Indefatigable Island. This species is related to the dark form of O. bilobata Say, but is quite differently colored and marked. The anterior lobe of the pronotum in the male is relatively longer than in the female and often more swollen. Family PYRRHOCORIDAE Dysdercus concinnus Stal. Dysdercus concinnus Stal. Stal, Oefv. Vet Ak. Fork p. 198, 1861; Enum. Hem. I, p. 121, 1870. Distant, Biol. Cent. Amer. Rhynch. I, p. 231, PL 21, figs. 11, 12, 15, 1883. Dysdercus mundus Walk., Cat. Hemipt. Br. Mus., V, p. 188, 1872. A single specimen was taken on Indefatigable Island. It is a fairly common neotropical species not hitherto reported from the Galapagos Islands, in fact it is the only member of the family yet found there. Family MIRIDAE Creontiades fuscosus sp. nov. (Plate 49) Head, pronotum, scutellum, hemielytra anteriorly and posteriorly, antennae with basal segment, narrow base and apical third of second, first segment of rostrum, most of sternum and venter, femora and posterior tibia, dark brown or fuscous; posterior half of clavus and broad transverse area posteriorly ex- tended along the costal margin, pale yellow-white. Head shading to brown on the vertex, .75 mm. long, .8 mm. wide across the eyes; diameter of vertex very nearly equal to diameter of an eye; median longitudinal sulcus distinct; clypeus separated from the vertex by a distinct furrow. Antenna with basal segment .75 mm. long, second very nearly 2 mm., third 1.5 mm. and the fourth about .5 mm. long; the last two smoky-brown in color with the base of the third paler. Rostrum sordid stramineous, reaching to apices of posterior coxae. Pronotum somewhat shining, rugose, shading into paler brown on the disk, sparsely coated with fine appressed hairs, 1.5 mm. wide and 1 mm. long, lightly depressed behind and between the slightly evident callosities. Scutellum very distinctly transversely depressed before base, posterior to which the disk is quite convex. Hemielytra impunctate, with sparse fine coating of appressed pale hairs; apex of corium not reaching tip of abdomen. Membrane dark smoky-brown. Sternum and venter fuscous-brown. Coxae, trochanters, odoriferous orifices and outer margin of the meta-pleura stramineous. Anterior and intermediate tibia pale stramineous with a faint prebasal, premedian and apical band, brown; tarsal segments tipped with brown. Length, male 4 mm.; female, 5.5 mm. Type: female, Indefatigable Island; allotype topotypic. Paratypes: male, Indefatigable Island and female, James Island. 249 250 Zoologica: N. Y . Zoological Society [V; 21 I am indebted to Dr. H. H. Knight who has examined this and the following species and placed them generically for me. Psallus insularis sp. nov. Color sordid stramineous with sparse coating of depressed hairs, fringed with a few long hairs along the lateral margin of the pronotum. Head some- what shining, nearly .25 mm. long and .5 mm. wide. Antennae a little dusky, short, with the two basal segments relatively stout; first segment reaching just beyond apex of the head, second segment .65 mm. long, third one-half the length of the second, the fourth a trifle over one-half the length of third. Rostrum, except at apex, pale, reaching to apices of posterior coxae. Pronotum .75 mm. wide and .3 mm. long. Scutellum with width equal to length, some- what shining, frequently a little dusky. Hemielytra somewhat shining, with the costal edge nearly straight. Membrane lightly infuscated. Sternum and venter concolorous, pale. Legs pale, femora with a few fine black hairs beneath and tibia provided with a few short black setae. Tarsi, except at apices, pale. Length, 2 mm. Type: male, James Island; allotype, topotypic. Paratypes: three and eight females, James Island. The specimens are all in rather poor condition. Family REDUVIIDAE Repipta annulipes sp. nov. Male moderately elongate, sparsely pilose, somewhat shining; stramineous in color. A broad fascia on the sides of the head before and behind the eyes, apices of the jugae and tylus, narrow rim about the ocelli, anterior pronotal tubercles, a broad fascia on the propleura, two subparallel fascia on the meso- pleura meeting anteriorly, a single fascia on the meta-pleura, a broad longitu- dinal fascia along the sides of the venter continuous with the fascia of the pleura, the outer margins of ventral segments 2-5 posteriorly and the costal margins fusco-piceous. The posterior lobe of pronotum, bases of the four spines and two discal spots posteriorly on the anterior lobe and the hemielytra dark brown or fuscous. The antennae and legs are pale stramineous, the femora twice banded beyond middle with brown, the tibiae with two sub-basal bands and the apices brown. Head equally long as the pronotum, shining, provided with two mediocre pale spines at base of the antenniferous tubercles, these not as long as one-third the length of an eye; postocular region a little longer than anteocular, much narrowed posteriorly to form a short collum, somewhat swollen behind the eyes, the sides lightly rounded and slightly converging posteriorly towards collum, the latter rather sharply set off. Antennae with basal segment stra- mineous, sparsely setose, three and one-half mm. long, slightly embrowned at apex; second segment slightly embrowned at apex, one mm. long, third segment slightly incrassate, narrowly pale at base, 5 mm. long, fourth segment brown, about 1.5 mm. long. Rostrum pale with little difference between the length of the first and second segments. Pronotum sparsely covered with incumbent grayish hairs, especially on the posterior lobe; anterior lobe shining, the deep 1924] Barber: Hemiptera-Heteroptera 251 central longitudinal groove evanescent just before the anterior margin; the posterior lobe finely rugulose, the four spines mediocre but slightly longer than the preocular ones, pale at apices. Scutellum shining stramineous, produced into a short, somewhat elevated, acute process. Hemielytra brown with the costal margins infuscated, as on the posterior lobe of the pronotum, the surface has a sparse covering of incumbent grayish hairs, the apex sordid stramineous; the apical quadrangular cell over twice as long as wide. Membrane hyaline, extending well beyond apex of abdomen, the median vein embrowned. Con- nexival margins of abdomen unarmed. Venter sparsely setose, with a broad black longitudinal vitta placed nearer to lateral margin than to median line, this provided anteriorly on segments 2 to 5 with an oblique pale depression. Genital segment provided with a rather stout upwardly curved spine. Legs densely pilose. Length, 11 mm. Type: Male, Indefatigable Island. Family NABIDAE Nabis punctipennis Blanch. One female, James Island, April 5. I follow Heidemann in the identi- fication of this species. Without specimens for comparison it is impossible to be sure of the identity of this species. It certainly is very close to N. ferus. Family TINGIDAE Corythaica renormata sp. nov. Pale cinereous, with antennae (only the two basal segments present) and legs pale stramineous. The following parts infuscated: hood dorsally, especially the areoles of it, a few of the veins of the paranota very faintly, the tumid elevation of the corium inwardly, costal areole and veins bounding it in the expanded part, a cross-vein at the rather abrupt termination of the arcuated part, the fourth cross-vein beyond this, all of the veins bounding the areoles along the posterior margin of the corium, broadly and those bounding the areoles of the membrane so broadly as to make these parts clouded; a pale translucent spot in the middle of the membrane occupying two large areoles. Hood somewhat elongate, nearly twice as long as its greatest diameter behind the middle, acute anteriorly, rounded behind, provided with about seven sunken areoles, carinate in the middle; seen from the side it is more arched dorsally and does not extend so far forward beyond the head as in C. carinata. The lateral margins of the pronotum strongly elevated, com- pressed and strongly arcuated before the acutely produced, almost erect paranotal angles; these provided with only about four large areoles confined to the margins. The median carina a little longer than the hood, more arched before the middle, not more markedly elevated than the lateral carina and not distinctly areolate except for one or two large areoles in the widest part; the disk anteriorly embrowned, with large obscure shallow punctures, posteriorly to which the disk is provided with small, clean-cut punctures, the posterior prolongation obscurely areolate. Hemieleytra strongly, arcuately expanded 252 50. HALOBATES ROBUST US sp. 1924] Barber: Hemiptera-Heteroptera 253 one-third way from base, the costal area wide, provided with a single row of about sixteen areoles, a double areole just posterior to the widest part; the costal areoles are subquadrate anteriorly and posteriorly with the four cells just behind the arcuated part wider than long; the subcostal area with three irregular rows of shallow areolate-like punctures; the discal area very nearly equally long as the remainder of the hemielytra, provided with small shallow areoles, not as evident as in C. planaris; the tumid elevation is very pronounced opposite to the widest part of the hemielytra, almost evenly declivous anteriorly and posteriorly and provided with about three rows of areoles at the highest part. The membrane is provided with large areoles. Body beneath dark castaneous, with margins of bucculae and pleurae in part paler. Length: 3 mm.; width of pronotum, .95 mm.; width of hemielytra, 1.15 mm. Type: female, Daphne Major, April 24. This differs from carinata in being relatively much wider with different shape of the hood and in the acute paranotal angles, etc. It is more closely related to planaris from which it can readily be distinguished by its narrower form, less expanded paranotal angles, less elevated pronotal carinae and dif- ferences in the arrangement of the areoles of the coastal region. Dr. C. J. Drake who has examined this specimen pronounces it as new. Family SALDIDAE Pentacora rubromaculata Heid. Pentacora rubromaculata Heid. Heidemann, Proc. Wash. Acad. Sci. Ill, 368, 1901. Only a single specimen was obtained. Family GERRIDAE Halobates robustus sp. nov. (Fig. 50) Form rather robust, widest across the apex of the mesonotum. Antennae and fore legs shining brownish black, with the usual coating of fine hairs; the intermediate and hind legs somewhat shining, brownish black, covered with fine hairs. Body black, slightly shining, densely coated with fine pale pubes- cence; on the sides and ventrally, with a much denser coating of fine incumbent hairs, giving a silvery white appearance to those parts. Base of the head between the eyes furnished with two oblique yellowish-brown fascia which do not quite meet in the central line. Dorsally with the posterior margins of the exposed abdominal segments narrowly and more or less distinctly yellowish-brown; ventrally with the posterior margins of these segments more distinctly bordered with yellow-brown and the first genital segment in the female is posteriorly and on the sides and the two genital plates of the second for the most part, yellowish; in the male the last genital segment is black. The head is almost twice as wide as it is long; the apices of the antenniferous tubercles are rather acutely prominent but not extended quite as far forward as the median lobe of the head. The antennae are rather slender, almost two- thirds as long as the body, the first segment only a trifle shorter than the other 254 Zoologica : N. Y. Zoological Society [V ; 21 three taken together; length of the respective segments as follows: male, I 1.35 mm., II .5 mm., Ill .45 mm., IV .5 mm.; female, I 1.6 mm., II .6 mm., Ill .5 mm., IV .56 mm. The pronotum is four times as wide as long, just anterior to the middle with a shallow transverse depression on either side. The mesonotum abruptly widened in front just behind the pronotum from whence the sides are gradually rounded to the widest part just before the base of the middle legs; the greatest width in the male is 1.80 mm., in the female 2.30 mm.; in the male the dorso-ventral dimension is 1.22 «im., in the female 1.65 mm. The fore legs are rather stout with the apical tibial tooth well-developed; the length of the respective segments as follows: male, femur 1.6 mm., tibia 1.35 mm., tarsus .66 (first segment .165 mm., second .495); female, femur 2 mm., tibia 1.76 mm., tarsus 1.1 mm. (first segment .38 mm., second segment .72 mm.). The middle legs are 11.49 mm. long in the male and 12.31 mm. long in the female, with the greatest diameter .25 mm. one-quarter way from the base; the length of the respective segments is as follows: male, femur 4.67 mm., tibia 4.4 mm., tarsus 2.42 mm. (first segment 1.65 and the second .77 mm.); female femur 5.06 mm., tibia 4.5 mm., tarsus 2.75 mm. (first segment 1.98 and the second .77 mm.); the apical two-thirds of the tibia provided with a fringe of fine curved hairs over twice as long as the diameter of the tibia and agglutinated at their apices. The hind legs are much shorter and more slender, being about 6.71 mm. long in the male and 7.72 mm. in the female, with the greatest diameter .165 mm.; the relative length of the respective segments is as follows: male, femur 3.85 mm., tibia 2.2 mm., tarsus .66 mm. (first segment .44 mm., the second .22 mm.). The abdomen consists of the usual nine segments which may best be^shown in the illustration. Length: male, 3.85 mm., width 1.80 mm.; female, 4\5 mm., width 2.3 mm. Tyjpe: male, Conway Bay, Indefatigable Island. Allotype and 24 paratypes taken with the type. This species seems most closely related to sericeus but besides its difference in size the relative lengths of the segments of the antennae and legs will serve to distinguish it. This is one of the series of scientific papers of the Harrison Williams Gala- pagos Expedition, under the directorship of William Beebe, sent out by the Department of Tropical Research of the New York Zoological Society. The general account and narrative of the expedition, together with the natural history and photographs of the fauna, are embodied in a volume by William Beebe, published by G. P. Putnam’s Sons, under the auspices of the Zoological Society. Its title is “Galapagos; World’s End.” ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY MUSg- INDEX TO VOLUME V DECEMBER 1923-1925 MARCH Numbers 1-21 Inclusive PUBLISHED BY THE SOCIETY THE ZOOLOGICAL PARK, NEW YORK 1927 Zoologica, Vol. V, Nos. 1-21. Snbex Ablechrus, 235 darwinii Charles Waterhouse, 235 flavipes Charles Waterhouse, 235 Acanthoderes galapagoensis Linell, 238 A carina, Galapagos Is., 93-95 Achelous, 159 ruber Milne Edwards, 159 tumidulus Stimpson, 159 Achryson galapagoensis Linell, 238 Acilius incisus Aube, 225, 235 Acribis serrativentris Charles Waterhouse, 235 Acronyctinae, 24-25 Williams Galapagos Expd., 24-25 Acrosternum (Nezara), 241 vividans Stal., 241 Acrotelsa galapagoensis Banks, 67, 71-73 description, 71-72 (Plate IY, figs. 13-17; Plate Y, figs. 18- 20), 70, 72 Aedes, 87 niger Gibs, 87 portoricensis Ludlow, 87 taeniorhynchus, 87 portoricensis (Ludlow), 87 Aega, 183 acuminata Hansen, 183 longicornis Hansen, 183 plebeia Hansen, 183 Aegathoa excisa Richardson, 184 Aegidae, 183 Galapagos Is., 183 Agraulis, 58 vanillae galapagensis Holland, 55, 57, 59 Agromyzidae, 90 Galapagos Is., 90 Agrotinae, 23-24, 32-33 Williams Galapagos Expd., 23-24 description of new species, 32-33 Agrotis ypsilon, 51 Albatross, U.S.S., 85, 101, 183, 221, 223, 228, 230 visit to Galapagos, 85, 101, 109, 118, 183, 221, 223, 228, 230 Catalogue of insects collected by, 221 Alcidamea, 131 Aldrich, Dr. J. M., 92 Allecula, 236 Alleculidae, 236 Galapagos Is., 236 Alloxacis seymourensis sp. nov., 226-227, 235 description, 226-227 Alpheus bouvieri var. chilensi Coutiere, 162 Amblygnathus obscuricornis (Geo. R. Water- house), 234 Amblyomma, 94-95 darwini, Hirst, 95 hirtum , 95 williamsi, sp. nov. 94-95 description, 94-95 (Fig. 16), 94 Amblyrhynchus, 11, 95 Galapagos Islands, 11, 95 (Fig. 6), 10 American Museum collections from Williams Galapagos Expedition, 3 Ammophorus bifoveatus Geo. R. Waterhouse, 229, 236 caroli Linell, 236 cooksoni Charles Waterhouse, 236 galapagoensis Geo. R. Waterhouse, 236 obscurus Geo. R. Waterhouse, 228, 229, 236 Ammotrecha solitaria Banks, 95 Amobiinae, 91 Galapagos Is., 91 Amphicerus, 237 cornutus (Pallas), 237 subspecies galapaganus Lesne, 237 punctipennis, 237 Amyna, 25, 37, 54 insularum sp. nov., 25, 37, 54 description of species, 37 (Plate I), 34 octo Guenee, 37 Anchastus galapagoensis Geo. R. Water- house, 235 Anchonus galapagoensis Geo. R. Waterhouse, 238 Andrena, 128 Annual Garden Party of the Zoolo gica Society, 3 Anomis 27, 42 doctorum Dyar, 42 professorum sp. nov., 27, 42 description, 42 (Plate II), 40 Anomura The Macrura and Anomura Collected by the Williams Galapagos Expedition, by Waldo R. Schmitt, 161- 171 (Figs. 39-41 incl.), 162, 164, 166 Anthoecia, 23 cystiphora Wallgrn., 23 inflata Wallgrn., 23 onca Wallgrn., 23 Anthrax , 87 lateralis, Thomson, 87 nudinscula Cog., 87 primitiva Walker, 87 tincta Thomson, 87 Ants, 53, 101-122 Galapagos Ils., 53, 101-122 Collected by Albatross, 101 Collected by Expd. of Calif. Acad., 101 Collected by Williams Expd., 101- 122 Apate, 219, 237 Aphiochaeta, 88 scalaris (Loew), 88 Aporinellus, 174-175 fasciatus (Smith), 174 galapagensis sp. nov. 174-175 description, 174-175 Apseudes galapagensis Richardson, 183 Apseudidae, 183 Galapagos Is., 183 Apterygota of the Hopkins Stanford Expd. to Galapagos, 67 Apterygota of the Williams Galapagos Ex- pedition, by J. W. Folsom, 67-76 (Plates III, IY, Y), 68, 70, 72 Arachnida of the Williams Galapagos Ex- pedition, by Nathan Banks, 93-99 (Figs. 15-18 inch), 93, 94, 96, 98 Araneida, 95-99 Galapagos Is., 95-99 [255] 256 Zoologica [Vol. V Arctiidae, 23 Williams Galapagos Exp., 23 Arcturidae, 184 Galapagos Is., 184 Arcturus abyssicola Beddard, 184 Argiope argentata Fabr., 97 Argyrodes jucundus Cambr., 97 Ariadne tarsalis Banks, 95 Aristotelia, 30 howardi Walsingham, 30 Asclera, 227 Asellota, 184 Galapagos Is., 184 Ashmead, Dr., 173 Asilidae, 87 Galapagos Is., 87 Asio galapagensis, 139 Asyndetus, 88 versicolor sp. nov., 88 description, 88 Atomus, 94 Atteva, 31, 53, 54 hysginiella Wallengren, 31, 53, 54 sylpharis Butler, 31 Attidae, 98-99 Galapagos Is., 98-99 Audouin, J. V., 134 Baccha, 88 clavata Fabr., 88 fascialis Thomson, 88 Banks, Mr. Nathan, 67, 71, 93 Banks, Nathan Arachnida of the Williams Galapagos Expedition, 93-99 (Figs. 15-18 inch), 93, 94, 96, 98 Banks, Nathan Neuroptera from the Williams Galapa- gos Expedition, 177-180 Barber, H. G. Hemiptera-Heteroptera from the Wil- liams Galapagos Expedition, 241-254 (Figs. 47-50 inch), 244, 247, 249, 252 Bassett-Smith, 211 Bates, H. W., 239 Baur, Dr. G., 110, 221, 222, 223, 229, 237 Beagle, H.M.S., 85, 219, 241 visit to Galapagos Is., 85, 219, 241 Beauregard, H., 129, 134 Beck, 51 Beck, Rollo, 81, 84 Beebe, William, 3, 46, 63, 67, 73, 76, 77, 85, 101, 125, 137, 143, 173, 181, 211, 222, 241 Galapagos, World’s End, 7, 20, 63, 209 Beebe, William Notes on Galapagos Lepidoptera, 50-59 Beebe, William, 3, 46 Williams Galapagos Expedition, 3-20 Brief narrative, 7-20 (Figs. 1 to 10 inch), 4, 5, 6, 8, 9, 10, 12. 16, 18, 20 Plate A, map of Galapagos Islands, 2 Resume, 3-7 Beebea, gen. nov., 30, 46, 54 description, 46 guglielmi sp. nov., 30, 46, 54 description, 46 (Plate II, 40) Bee-flies, 87 Galapagos Is., 87 Birds Williams Galapagos Expedition, 3, 5, 11, 54, 55-57 Bird-infesting Mallophaga Collected by the Williams Galapagos Expedition, by H. E. Ewing, 81-84 (Fig. 12), 83 Bithynis jamaicensis, Rathbun, 169 Blair, Mr. K. G., 222, 228, 229, 236 Bledius aequatorialis sp. nov. 225-226, 235 description, 225-226 (Fig. 43), 226 Boheman, C. H„ 219, 221, 239 Bombyliidae, 87 Galapagos Is., 87 Boobies Galapagos Islands, 15, 19 Bopyridae, 185 Galapagos Is., 185 Bopyrina latreuticola (Gissler), 168 Bostrichidae, 237 Galapagos Is., 237 Bostrichus unicantus, 237 Bouvier, J., 209 Brachyuran Crabs collected by the Williams Galapagos Expedition, by Mary J. Rathbun, 153-159 (Plate VII, Fig. 38), 154, 157 Broking, Mr. drawings on Williams Galapagos Ex- pedition, 5 Brues, Charles T., Triungulin Larvae from the Williams Galapagos Expedition, 125-136 bibliographv, 134-136 (Figs. 28-32 inch), 126, 127, 131, 132 Brues, Prof. C. T., 103, 132, 134, 135 Bruesia Perk., 133 Budde-Lund, G., 189, 198, 209 Bugnion, E., 130, 134 Buprestidae, 221 Galapagos Is., 221 Bursera, 69, 144, 145 graveolens Trian. and Planch., 108, 110, 112, 118 Busck, A., 30 Buteo galapagensis, 91, 139 Tick-flies from, 91 Butorides sundevalli, 91 Galapagos Is., 91 Tick-flies from, 91 Butterflies, Galapagos Islands, 55-59 migration, 57-59 Cactus Galapagos Islands, 16, 17, 108 (Fig. 8), 16 Calappa, 153, 159 conveza Saussure, 153, 159 flammea (Herbst.) Bose., 159 Calcinus obscurus (Stimpson), 170 California Academy of Sciences, Expedition to Galapagos Islands, 51, 221 Coleoptera collected by, 222, 233 C aligns, 211-212 irritans Heller, 211-212 parvus Bassett-Smith, 211 Callidryas eubule, 54, 55, 56, 57 Callinectes danae Smith, 158-159 Callipharixenos, 133 Callitroga macellaria Johns, 89 Calosoma Galapagos Islands, 53, 221, 223 galapageium Hope, 221, 222, 234 howardi Linne, 53, 223, 234 linelli sp. nov., 222-223, 234 description, 223 Calotermes pacificus Banks, 103, 112, 115, 118 Camponotus formciformis Forel, 111 Camponotus Mayr., 53, 111-121 Galapagos Is., 53, 111-121 macilentus F. Smith, 112, 113, 120 var. Albemarlensis, 120 var. barringtonensis, 121 var. bindloensis, 121 var. castellanus var. nov., 116-118, Nos. 1-21] Index 257 121 description, 116-118 (Fig. 25), 117 var. duncanensis, 120 var. hoodensis, 121 var. jacobensis, Wheeler, 116, 118, 121 var. narboroensis, 120 var. pervicus var. nov., 115-116, 120 description, 115-116 var. sapphirinus var. nov., 114- 115, 120 description, 114-115 (Fig. 24), 114 var. vulcanalis, 120 planus F. Smith, 53, 112, 113, 120 var. fernandinensis, 120 var. fidelis, 121 var. indefessus, 119, 120 var. isabelensis Wheeler, 53, 119, 120 var. peregrinus Emery, 53, 113, 118, 119, 120 var. pinzonensis, 120 var. sansalvadorensis var. nov. 119- 120 description, 119-120 (Fig. 27), 120 var. santacruzensis Wheeler, 119, 120 (Fig. 26), 118 senex F. Smith, 111, 112 Cancer clypeatus Herbst, 170 flammeus, Herbst, 150 grapsus Linn., 156 jamaicensis Herbst, 169 minutus Linn., 157 Capsidae, 53 Galapagos Is., 53 Carabidae, 233, 234-245 Galapagos Is., 233, 234-235 Cardiosoma guanhumi Lat., 156 Cassia , 54 Catabena, 25 Caterpillars, food for birds, 57 Catocalinae, 26, 41 Williams Galapagos Expd., 26, 41 description of new species, 41 Cattle, wild Galapagos Islands, 13 Centipedes, 137-141 Galapagos Is., 137-141 Cerambycidae, 221, 234, 238 Galapagos Is., 221. 234, 238 Ceratopogon galapagensis Coquillett, 86 Ceratothoa, gaudichaudii Milne-Edwards, 183 gaudichaudii Schiodte and Meinert, 187 rapax Heller, 187 Ceroplastes, 152 Chalcis sp., 175 Chamberlain, Ralph Y. Chilopods of the Williams Galapagos Expedition, 137-141 (Plate VI, figs. 1-4), 138 Champion, G. C., 131, 134 Characoma, 26 nilotica Rogenh., 26 Chelifera, 183 Galapagos Is., 183 Chilopods of the Williams Galapagos Ex- pedition, by Ralph Y. Chamberlain, 137- 141 (Plate VI, figs. 1-4), 138 Chironomidae, 86 Galapagos Is., 86 Chironomous, 86 Chloridea, 23-24 cystiphora Wallgrn., 23 virescens Fabr., 24 Chloropidae, 90 Galapagos Is., 90 Chobaut, A., 132, 134 Chrysomelidae, 221, 238 Galapagos Is., 221, 238 Chrysomyia quadrisignata Coq., 89 Chrysopa, 112 Chrysopidae, 177-179 galapagoensis sp. nov., 179 description, 179 nigripilosa sp. nov., 178-179 description, 178-179 Cicindela, 221, 234 galapagoensis Van Dyke, Williams, 221 234 Cicindelidae, 234 Galapagos Is., 234 Cirolana may ana Ives, 181, 185-187 description, 186-187 (Plate VIII, figs. 1-3), 186 Cirolanidae, 185-187 Galapagos Is., 185-187 Cirphis, 24 cooperi, sp. nov., 24 description, 33 (Plate I), 34 pyrrhias Meyr., 33 Cissites testaceus Fabr., 130, 131 triungulin of, 130, 131 (Fig. 31), 131 Citharichthys sordida , 184 Cleridae, 221 Galapagos Is., 221 Clipperton Island, 137 Clubiona, 96 Clubionidae, 96 Galapagos Is., 96 Coccidae, The, of the Williams Galapagos Expedition, by Harold Morrison, 143- 152 (Figs. 33 to 37 inch), 144, 146, 147, 149, 151 Coccinae, 152 Galapagos Is., 152 Coccinellidae, 221, 236 Galapagos Is., 221, 236 Cochlionmyia, 89 macellaria (Fabricius), 89 macellaria Towns., 89 Coenobita, 170 clypeatus (Herbst), 170 compressa Guerin, 170 compressus (Guerin), 170 diogenes Benedict, 170 Coleoptera from the Williams Galapagos Ex- pedition, by Andrew J. Mutchler, 219- 240 Bibliography, 239-240 (Figs. 42-46 inch), 224, 226, 227, 231, 232 Coleoptera Galapagos Is., 125-136, 219-240 List of Coleoptera known to occur in Galapagos Islands, 233-239 Collembola, Williams Galapagos Expd., 67 Collnet, Captain, 234 Colpocephalum Nitzsch unciferum Kellogg, 82 Compsomyia macellaria Arrib., 89 Conolophys, 11, 19 Galapagos Islands, of, 11, 19 (Fig. 4), 8 Conomyrma, 110 Conway Bay, Indefatigable Is., 9, 11 Cookson, W. E„ 111, 201, 225 258 Zoologica [Vol. V Cooper, Miss Isabel, 5, 33, 41, 65 water colors on Williams Galapagos Expedition, 5 Copelatus galapagoensis Geo. R. Waterhouse, 235 Copepods, Parasitic, from the Williams Galapagos Expedition, by Charles B. Wilson, 211-217 (Plate XX), 215 Copris lugubris Boheman, 237 Coptostelhus williamsi sp. nov., 227-228, 236 description, 227-228 (Fig. 44), 227 Coquillett, D. W„ 85, 87, 89, 91 Cordia, 54, 58, 226 lutea Lam., 108, 112, 226 Coreidae, 241 Galapagos Is., 241 Corizus hyalinus Fab., 245 lugens Stal, 245 Cormorant, Farallone, 84 lice from, 84 Cormorant, flightless, 3, 12, 14 Galapagos Islands, 3, 12, 14, 15, 84 (Fig. 7), 12 lice from, 81, 83 Corynetes rufipes, 219, 228, 235 Corynetidae, 235 Galapagos Is., 235 Coryphaena hippurus, 184 Corythaica, 251, 253 carinata, 251, 253 planaris, 253 renormata sp. nov., 251, 253 description, 251, 253 Coutiere, Prof., 162 Crab, Brachyuran Crabs of Williams Gala- pagos Expedition, 153-159 Box Crabs, 159 Fiddler Crabs, 155, 156 Ghost Crabs, 155 Land Crabs, 156 Mangrove Crab, 157 Oblong Crab, 158 Red-Bristle Crab, 158 Rock Crab, 156 Sand Crab, 155 Sargassum Crab, 158 Stone-crab, 158 Sculptured Crab, 157 Swimming Crabs, 158, 159 Crambinae, 29 Williams Galapagos Expd., 29 Crane-flies, 86 Galapagos Is., 86 Crangon bouvieri var. chilensis (Coutiere), 162 Crayfish, 161, 169 Creagrus Gulls, 19 Creontiades, 248-250 fuscosus sp. nov., 248-250 description, 248 (Fig. 49), 249 Creophilus, 219, 235 maxillosus, 219 villosus (Gravenhorst), 219, 235 Crocidosema, 31 plebiana Zeller, 31 Cronius, 159 bispinosus Miers, 159 ruber (Lamarck) Stimpson, 159 tumidulus (Stimpson), 159 Cros, Aug., 128, 129, 130, 135 Crustacea • Galapagos Is., 153-159, 161-171, 181- 210, 211-217 Cryptione elongata Hansen, 185 Cryptopidae, 137-139 Galapagos Is., 137-139 Cryptops oeebei sp. nov., 137-139 description, 137-139 navigans Chamberlain, 137, 139 Ctenidae , 98 Galapagos Is., 98 Ctenolepisma Galapagos Is., 75 Cubans , 181, 185, 201-205 beebei sp, nov., 181, 203-205 description, 203-205 (Plate XVIII, figs. 28-30), 204 cincta Dollfus, 205 dugesi (Dollfus), 205 dumorum (Dollfus), 205 galapagoensis Miers, 181, 185, 201-203 description, 201-203 (Plate XVII, figs. 23-27), 202 grenadensis (Budde-Lund), 205 perlata Dollfus, 205 pisum (Budde-Lund), 205 silvarum Dollfus, 205 vincentis (Budde-Lund), 205 Cuckoos, 57 Culicidae, 87 Galapagos Is., 87 Cupido parrhasionides Wallen., 55 Curculionidae, 221, 238 Galapagos Is., 221, 238 Cychrus stenostomus, 222, 223 Cyclosa conica Clerck, 97 Cylindromyrmex, 101-107 key to 7 known species, 106 boliviae sp. nov. description, 104-105 (Fig. 20), 105 brasiliensis Emery, 104, 106 godmani Forel, 104, 105, 106 longiceps Ern. Andre., 106 meinerti Forel, 106 striatus Mayr., 101, 102, 104, 106 williamsi sp. nov. description, 101-104 (Figs 19, b & c), 102 Cylindromyrmicini, 104 Cymothoa, 184, 187 exigua Schioedte and Meinert, 184 gaudichaudii Milne-Edwards, 187 Cymothoidae, 183-184, 187-189 Galapagos Is., 183-184, 187-189 Cymus marginalis Dallas, 245 Cypselurus furcatus (Mitchill), 63 nigricans (Bennett), 63 Dactylopiinae, 145-152 Galapagos Is., 145-152 Danais plexippus (Linne), 55 Daphne Major Is. Galapagos Archipelago, 19 Darwin Bay, Tower Island, 6, 19 (Fig. 3), 6 Darwin, Charles, 13, 85, 219, 225, 228, 229, 234, 237, 241 visit to Galapagos Islands, 13, 85, 219, 225, 228, 229, 241 Davidson, A. C., 131, 135 De Geer, Carl, 239 Deilephila lineata Fab., 54, 55, 57, 59 Dermestes 219, 228, 236 carnivorus Fabricius, 228, 236 vulpinus Fabricius, 219, 236 Dermestidae, 221, 236 Galapagos Is., 221, 236 Diabrotica limbata Chas. Waterhouse, 238 Dicranomyia, 86 floridana O. S., 86 Dictyna remota sp. nov., 97 description, 97 (Fig. 17), 96 Nos. 1-21] Index 259 Dictynidae, 97 Galapagos Is., 97 Dilophonota PI obscura conformis Roth & Jord., 54 Diptera, collected by Charles Darwin on Gala- pagos Islands, 85 described from Galapagos Is. by F. Smith in 1877, 85 by C. G. Thompson in 1868, 85 by Francis Walker in 1849, 85 total species recorded from Galapagos Is., 85 Diptera of the Williams Galapagos Expedi- tion, by Charles W. Johnson, 85-92 (Figs. 13-14), 86 Docema, 230, 238 darwini sp. nov., 230, 238 description, 230 galapagoensis Geo. R. Waterhouse, 238 Dogs, wild Galapagos Islands, 13 Dolichoderinae, 110 Galapagos Is., 110 Dolichopodidae, 88 Galapagos Is., 88 Donkeys, wild Galapagos Islands, 13 Dorymyrmex pyr amicus Roger, 110 subsp. albemarlensis Wheeler, 110 (Fig. 23), 110 Dove, 3 Galapagos Islands, 3, 19 Drake, Dr. C. J., 253 Drassidac, 96 Galapagos Is., 96 Dromicus dorsalis, 54 Galapagos Is., 54 Drosophila, 90 Galapagos Is., 90 pallida Williston, 90 willistoni Sturt., 90 Drosophilidae, 90 Galapagos Is., 90 Duck, wild Galapagos Islands, 11 Dufour, L., 129, 135 Dysderidae, 95 Galapagos Is., 95 Dystiscidae, 221, 235 Galapagos Is., 221, 235 Dytiscus sticticus Linnaeus, 225, 235 Eburia, 238 amabilis Boheman, 238 bauri Linell, 238 lanigera Linell, 238 Eden Island, Galapagos, 2, 9, 11 lava slopes, 10 (Fig. 6), 10 Elasmopalpus ?, 30, 46 galdinella sp. nov., 30, 46-47 description, 46-47 (Plate II), 40 Elateridae, 221, 235 Galapagos Is., 221, 235 Elenchus, 133 Emenadia, 133 Ennominae, 28 Williams Galapagos Expd., 28 Epeira labyrinthea Htz., 97 prompta Htz., 97 Epeiridae, 97 Galapagos Is., 97 Epicaridea, 185 Galapagos Is., 185 Epidromia, 27, 43 zephyritis sp. nov., 27, 43 description, 43 (Plate II), 40 Epinephelus, 211 labrifrons, 211 morio, 211 Epinotia, 31 3 species from Galapagos Islands, 31 Erebus odor a, 51 Erestes sticticus (Linnaeus), 224-225, 235 Eriococcus Targ., 145-147 papillosus sp. nov., 145-147 description, 145-157 (Fig. 34), 146 palmeri Ckll., 147 Eriphia granulosa Milne-Ed wards, 158 hispida Stimpson, 158 Eriphides hispida (Stimpson), 158 Eromene, 29, 53, 54 ocellea Haworth, 29, 53, 54 Erythraeidae, 93-94 Galapagos Is., 93-94 Erythraeus remotus sp. nov., 93 description 93-94 (Fig. 15), 93 Esthiopterum Harrison acutifrons (Rudow), 84 faralloni (Kellogg), 84 helleri (Kellogg and Kuwana 82 nannopteri sp. nov., description, 82-84 (Fig. 12), 83 pelagicum (Denny), 84 potens var. minor var. nov., 84 description, 84 Etiella, 30 Zinckenella Treit., 30 Eucosomia, 28 albosignata Packard, 28 impauperata Walk., 28 stellata Guen., 28 Eudamus galapagensis, Williams, 55, 58 Eugenics , Swedish frigate, 85, 219 visit to Galapagos, 85, 219 Etl7Yl€Tl6S 133 Eunectcs occidentals Erichson, 224-225, 235 Eupomacentrus beebei sp. nov., 63-64 description, 63, 64 Eurycope, 184 pulchra Hansen, 184 scabra Hansen, 184 Euteliinae, 26, 38 Williams Galapagos Expd., 26, 38 description of new species, 38-41 Euxoa, 24 williamsi sp. nov., 24 description, 32 (Plate I), 34 Ewing, H. E. Bird-infesting Mallophaga Collected by the Williams Galapagos Expedition, 81-84 (Fig. 12), 83 Fabre, M., 135 Fabricius, J. Christ, 239 Feronia. 91, 234, 235 americana Leach, 91 insularis Boheman, 234 spinifera Leach, 91 Filistata fasciata Banks, 95 Filistatidae, 95 Galapagos Is., 95 Finch, black, Galapagos Islands, 15 Fish collected by Williams Galapagos Ex- pedition, 5, 20, 62-65 flying, 63 surgeon, 64 Flabellifera, 181, 183-184, 185-189 Galapagos Is., 181, 183-184, 185-189 260 Zoologica [Vol. V Flamingo Galapagos Islands, 14 Flies Galapagos Is. Bee-flies, 87 Crane-flies, 86 Flesh-flies, 88 Flower-flies, 88 Horse-flies, 187 Hump-backed, 88 Long-legged flies, 88 Pomace-flies, 90 Robber-flies, 87 Tick-flies, 91 Flycatcher, 11, 54, 57 Galapagos Islands, 11, 54, 57 Folsom, J. W. Apterygota of the Williams Galapagos Expedition, 67-76 (Plates III, IV, V), 68, 70, 72 Formicidae, The, of the Harrison Williams Galapagos Expedition, by William Morton Wheeler, 101-122 (Figs. 19-27 incl.), 102, 105, 107, 109, 110, 114, 117, 118, 120 list of Galapagos Formicidae with their distribution, 122 Formicinae, 111-121 Galapagos Is., 111-121 Fregata aquila, 82, 91, 110 lice from, 82 tick-flies from, 91 Fresh Water Bay, Chatham Is., 15 Frigate birds Galapagos Islands, 14, 19, 82, 91, 110 lice from, 82 tick-flies from, 91 Fundella, 30, 47 agapella sp. nov., 30, 47 description, 47 (Plate II), 40 Gahan, C. J., 130, 135 Galapagomyia gen. nov., 86 description, 86 longipes sp. nov., 86 description, 86 (Figs. 13-14), 86 Galapagos Heterocera, by W. Schaus, 22-48 new species, description of, 32-48 (Plate I), 34 (Plate II), 40 Galapagos Islands Beagle, visit of, 85, 219, 241 California Academy of Sciences Expedi- tion to, 51, 221, 222, 223 Darwin, Charles, visit to, 13, 85, 219, 225, 228, 229, 241 fauna of, see Williams Galapagos Ex- pedition fresh water, 8, 11, 12, 14, 15 “Galapagos; Worlds End,” 20, 48, 59 Hopkins Stanford Expedition, 81, 137, 140 lighthouses, 15 map of, (Plate A), 2, 22, 50, 62, 178, 182, 212 Williams Expedition, 2-20 Gasteracantha insulana Thorell, 97 Gecarcinus lateralis (Freminville), 156 lateralis Guerin, 156 Gelechia, 31 bosquella Chambers, 31 Gelechiidae, 30-31 Williams Galapagos Expd., 30-31 Genera Insectorum, 221 Geograpsus lividus (Milne Edwards) Stimpson, 156 Geometridae, 28 Williams Galapagos Expd., 28 Gerridae, 253-254 Galapagos Is., 253-254 Gifford, E. W„ 56 Gigantotheca galapagensis, 92 Gnathocerus cornutus (Fabricius), 237 Goats, wild Galapagos Islands, 13, 17 Goniopsis cruentata (Latreille), 157 cruentatus, Rathbun, 157 Gonodonta, 21 biarmata Guen., 27 elegans Druce, 27 Gossypium, 54 Grapsus, 156 cruentatus Latreille, 157 grapsus (Linn.) Ives, 156 lividus Milne Edwards, 156 Grasshoppers, Galapagos Islands, 57 Gravenhorst, J. L. C., 239 Grouper, red, 211 Grunt (fish), 63 Guilding, Rev. Landsdown, 130, 131, 135 Gull, 3, 11 Galapagos Islands, 3, 11, 19 Guy Fawkes Rocks, Galapagos, 11 Gyrinidae, 221 Galapagos Is., 221 Hadeninae, 24, 33 Williams Galapagos Expd., 24, 33 description of new species, 33-37 Hadruroides lunatus Koch, 95 Haemulon scudderi (Gill), 63 Halobates, 241, 253-254 robustus sp. nov., 253-254 description, 253-254 (Fig. 50), 252 sericeus, 254 Haltica galapagoensis, 238 Hansen, H. J., 183, 209 Harmostes disjunctus sp. nov., 241-245 affinis Dallas, 243 apicatus Stal, 245 disjunctus sp. nov., 241-245 description, 241-245 marmoratus B., 245 minor Spin., 245 prolixius Stal, 245 raphimerus Spin., 245 serratus Fab., 243, 245 Harris, T. W„ 239 Harrison Cove, Indefatigable Is., 10 Harrison Williams Galapagos Expedition, see Williams Galapagos Expedition Harrisonia, gen. nov., 25, 36 williamsi sp. nov., 25 description of genus, 36 description of species, 36-37 (Plate I), 34 Hawk, 3, 91 Galapagos Is., 3, 91 Tick-flies from, 91 Heinrich, C., 30 Heliocontia, 25 margana Fabr., 25 Heliotropium parviflorum, 145, 146 Heller, 211, 213 Hemerobiidae, 179-180 Galapagos Is., 178-180 Hemiptera, 53, 241-254 Galapagos Islands, 53, 241-254 Hemiptera-Heteroptera from the Williams Galapagos Expedition by H. G. Barber, 241—254 (Figs. 47-50 incl.), 244, 247, 249, 252 Hemirobiid, 54 Heraeus, 245-246 pacificus sp. nov., 245-246 description, 245-246 Nos. 1-21] Index 261 (Fig. 47), 244 plebejus Stal, 246 Heron, Galapagos Ts., 91 Tick-flies from, 91 Herse cingulata Mer., 55 Heterocera collected by Williams Galapagos Ex- pedition, 22-48, 51-59 Heterocera, Galapagos, by W. Schaus, 22-48 new species, description of, 32-48 (Plates I & II), 34, 40 Heterocrepidius puberulus Boheman, 236 Heterolepisma, 67-69 intermedia sp. nov., 67-69 description, 67-69 (Plate II, figs. 1-8; Plate IY, figs. 9-12), 68, 70 pampeana Silv., 69 Heteromera, 233 Heteropoda venatoria Linn., 98 Heymons, 74 Hippa 170, 171 denticulate rons (Miers), 170 denticulatif rons, Rathbun, 170 “Hippelates plague,” 90 Hippelates pusio Loew., 90 Hippoboscidae, 91 Galapagos Is., 91 Hippolysmata, 167 acicula Rathbun, 167 intermedia Kingsley, 167 moorei Rathbun, 167 paucidens Rathbun, 167 porteri Rathbun, 167 Hippolyte, 161, 163-165 caiiforniensis Holmes, 165 gracilis Heller, 165 obliqui-manus Dana, 165 pleuracantha (Stimpson), 164, 165 williamsi sp. nov., 161, 163-165 description, 163-165 (Fig. 40), 164 Histeridae, 221 Galapagos Is., 221 Hoffman, Harry oil paintings on Williams Galapagos Ex- pedition, 5 Holcoponera whymperi, 103 Homoptera of the Williams Galapagos Ex- pedition, by Herbert Osborn, 77-79 Hope, F. W., 219, 221, 239 Hopkins Stanford Galapagos Expedition, 81, 85, 91, 137, 140, 161 Horia auriculata, 130, 131 Horia maculata Swed., 126 (Figs. 28-30 inch), 126, 127 from Xylocopa transitora Perez, Gala- pagos Is., 125 mandibles with teeth on upper surface (Figs. 29, 30), 126, 127 triungulins of, 126-129 (Figs. 28, 29, 30), 126, 127 Horiini, 130 Horn, Dr. Geo. R., 227 Horn, Dr. Walther, 221, 239 Hornia, 129 Horseflies, 87 Galapagos Is., 87 Howard, Dr. L. O., 221, 237, 238, 239 Hungerford, H. B., 136 Hydrocampinae, 29 Williams Galapagos Expd., 29 Hydrophilidae, 221, 235 Galapagos Is., 221, 235 Hydrophilus lateralis , 219 Hylurgi, 239 Hymenoptera collected by the Williams Galapagos Expedition, by S. A. Roh- wer, 173-175 Hypergnathus Richardson, 198 Hypocylindromyrmex subgen. nov., 106-107 Hyponomeutidae, 31 Williams Galapagos Expd., 31 Hypsypops rubicundus, 64 Ichthyology of the Galapagos, A contribution to, by J. T. Nichols, 62- 65 (Fig. 11), 65 Indefatigable Island, Galapagos Archipelago (Fig. 10), 20 Insects collected by Williams Galapagos Ex- pedition, 5, 11, 23-48, 51-59, 67-92, 101-122, 125-136, 143-152, 173-175, 177-180 Isolepisma Esch., 69 Isopods from the Williams Galapagos Ex- pedition, by Willard G. VanName, 181- 210 Bibliography, 209-210 (Plates VIII-XIX inch), 186, 188, 190, 191, 193, 195, 196, 197, 199, 202, 204, 207 Ixodidae, 94-95 Galapagos Is., 94-95 Jackson, H. G., 209 James Bay, James Island, 11, 13 Jassus auratus, 78 galapagoensis sp. nov., 77 description, 77-78 Jay, 3 Johnson, Charles W. Diptera of the Williams Galapagos Ex- pedition, 85-92 (Figs. 13-14), 86 Kellogg, Y. L., 81, 84 Knight, Dr. H. H., 250 Kuwana, S. I., 81 Laphygma, 25 frugiperda Sm.-Abb., 25 Larentiinae, 28 Williams Galapagos Expd., 28 Lathrodectes apicalis Butler, 97 Latreutes, 168 ensiferus Stimpson, 168 fucorum (Fabr.) Stebbing, 168 Leander tenuicornis Stebbing, 168 Lepeophtheirus dissimulatus Wilson, 211 Lepidocyrtus, 67, 75-76 intermixtus sp. nov. description, 75-76 (Plate V, figs. 27-29), 72 Lepidoptera, 23-48, 51-59 Williams Galapagos Expd., 23-48, 51-59 melanism, 59 migration, 57-59 variation, 59 Lepisma Galapagos Is., 67, 71 galapagoensis Banks, 71 insularis Banks, 67 Lepismid Eggs and Nymphs, 73-75 (Plate V, figs. 21-26), 72 Williams Galapagos Expd., 73-75 Leptodius lobatus Milne Edwards, 158 Lesne, P., 237, 239 Lice, Bird, 81 Ligia baudiana, Bate Miers Richardson, Ives, Milne-Edwards, 205, 206 exotica, Roux, 185, 205, 206 gracilis Moore, 206 hirtitarsis Dahl, 205 262 Zoologica [Vol. V Ligyda baudiniana (Milne-Edwards), 181, 205 description, 206-208 (Plate XIX, figs. 31-36), 207 exotica (Roux), 185, 206 Ligydidae , 185, 205-208 Galapagos Is., 185, 205-208 Linnaeus, C., 239 Linell, Martin, 221, 222, 223, 225, 228, 229, 230, 237, 238, 239 Lizard, 3, 56 Galapagos Islands, 3, 8, 10, 19, 54, 56 (Fig. 4, 6), 8, 10 Lobopoda galapagoensis Linell, 236 Locusts, Galapagos Islands, 53, 57 Longitarsuslunatus Charles Waterhouse, 238 Long-legged flies, 88 Galapagos Is., 88 Lucilia quadrisignata Thom., 89 Lupa sayi Gibbes, 158 Lycophotia, 24, 32 lubricans Guenee, 33 oceanica sp. nov., 24, 32 description, 32-33 (Plate I), 34 Lygaeidae, 53 Galapagos Islands, 53 Lysmata, 161, 165-167 acicula Rathbun galapagensis sp. nov., 165-167 description, 165-167 (Fig. 41), 166 intermedia Kingsley, 167 moorei Rathbun, 167 paucidens Rathbun, 167 Lyttini, 130 Machilis mutica Banks Galapagos Is., 67 Macrobrachium, 169 jamaicense (Herbst.) Rathbun, 169 olfersii (Wieg.), 169 Macrosiagon, 133 Macrura and Anomura, The, collected by the Williams Galapagos Expedition, by Waldo L. Schmitt, 161- 171 (Figs. 39-41 inch), 162, 164, 166 Magusa, 24 orbifera Walk., 24 Malacodermes, 233 Mallodon 53, 221, 230, 238 Galapagos Islands, 53, 221, 230 molarium Bates, 53, 230 Mallophaga, Bird-infesting, Collected by the Williams Galapagos Expedition, by H. E. Ewing, 81-84 (Fig. 12), 83 Mallophaga, Revillagigedo Islands, 81 Mammals collected by Williams Galapagos Ex- pedition, 3 Margarodes Guilding, 143-145 rileyi Giard, 145 similis sp. nov., 143 description, 143-145 (Fig. 33), 144 Margarodinae, 143-145 Galapagos Is., 143-145 Marptusa , 99 Martin, purple, 54, 55, 56, 58 Mayet, V., 135 Maytenus obovata Hook, 103, 112, 115 Mecistocephalus parvus Chamb., 137 Megalomus darwini sp. nov., 179-180 description, 179-180 Megalops of Ocypode gaudichaudii, 153-155 (Plate VII), 154 Meinert, F., 210 Meinertia, 181, 183, 187-189 gaudichaudii (Milne-Edwards), 181, 183, 187-189 description, 188-189 (Plate IX, figs. 4-5), 188 Melander, A. L., 132, 135 Melipotis, 27, 42, 51, 54 famelica Guen., 43 harrisoni sp. nov., 27, 42-43, 57 description, 42-43, (Plate II), 40 indomita Walk., 27 nigrescens Grote & Rob., 27, 51, 54 sinualis, 51 Meloe, 129 cavensis, 130 Meloidae, 128, 129, 131, 132 Melyridae, 235 Galapagos Is., 235 Menippe nodifrons Stimpson, 158 Menopon auri-fasciatum Kellogg, 82 navigans Kellogg, 82 Menoponidae, 82 Galapagos Is., 82 Metacylindromyrmex, 104 Metacyrba insularis Banks, 98 Metoponorthus pruinosus Budde-Lund, 197 Microlepidoptera, 30-31 Williams Galapagos Expd., 30-31 Midges, 86 Galapagos Is., 86 Miers, E. J„ 201, 209 Migrations, of moths and butterflies, 57-58 Miridae, 53, 248-250 Galapagos Is., 53, 248-250 Mitchell, Dr. James, 7 Mites, Galapagos Is , 93-99 Mithrax, 153 bellii Gerstaecker, 153 ursus Bell, 153 Mods, 26, 41 incurvalis sp. nov., 26, 41 description, 41, (Plate I), 34 repanda Fabr., 26 Mockingbird, 3, 11, 16, 54 Galapagos Island, 3, 11, 16, 54, 57 Monkeys, 3 Monod, Th., 209 Monomorium floricola, 101, 108 Morrison, Harold, The Coccidae of the Williams Galapagos Expedition, 143-152 (Figs. 33 to 37 inch), 144, 146, 147, 149, 151 Mosquitoes, 53, 87 Galapagos Is., 53, 87 Moths, collected by Williams Galapagos Ex- pedition, 22-48, 51-59 descriptions of new species, 32-48 (Plates I & II), 34, 40 eaten by birds, 56-57 migration, 57-59 Munnopsidae, 184 Galapagos Is., 184 Munnopsis longiremis Richardson, 184 Murray, M., 133, 135 Musca, 88, 89 macellaria Fabr., 89 occidua Fabr., 88 ochricornis Wied., 89 phanda Walker, 89 Muscidae, 89 Galapagos Is., 89 Mutchler, Andrew J. Coleoptera from the Williams Galapagos Expedition, 219-240 Nos. 1-21] Index 263 Bibliography, 239-240 (Figs. 42-46 inch), 224, 226, 227, 231, 232 Mycterodus productus Stal., 78 Myrmamblys, 113 Myrmeleon, 177 crudelis, 177 perpilosus sp. nov., 177 description, 177 Myrmeleonidae, 177 Galapagos Is., 177 Myrmicinae, 108-110 Galapagos Is., 108-110 Myrmocladoecus Wheeler, 113, 114 bidens Mayr, 113 bispinosus Mayr, 113 latangulus Rog., 113 mucronatus Emery, 113 planus F. Smith, 118-120 quadrilaterus Mayr, 113 Myrmorhachis Forel, 113 Nabidae, 251 Galapagos Is., 251 Nabis, 251 ferus, 251 punctipennis Blanch, 251 Nannophilus, 140 Nannopodellus, gen. nov. description, 140 purpurascens sp. nov. description, 14-141 (Plate VI, figs. 1-4), 138 Nannopterum harrisi 3, 12, 14, 15, 81, 83, 84 Necrobia ruftpes (DeGeer), 219, 228, 235 Necrophaga, 233 Neleus tlascala Percheron, 238 Nematocarcinus agassizii Faxon, 185 Nemognatha scutellaris Lee., 131 Neopentarthrum gen. nov., 231, 238 description, 231 towerensis sp. nov., 231-232, 238 description, 231-232 (Fig. 45), 231 Nephila clavipes Linn, 97 Neptunus sayi Milne Edwards, 158 Nessipus, 213-217 costatus sp. nov., 213-214 description, 213-214 (Plate XX, figs. 9-16), 215 crypturus Heller, 217 occultus sp. nov. description, 214-217 (Plate XX, figs. 1-8), 215 Neuroptera from the Williams Galapagos Expedition, by Nathan Banks, 177-180 Nicetiodes gen. nov., 30, 48 description, 48 apianella sp. nov., 30, 48 description, 48 (Plate II), 40 Nichols, J. T. A contribution to the Ichthyology of the Galapagos, 62-65 (Fig. 11), 65 Nierstrasz, H. F., 209 Noctuidae, 23-27, 32-45 Williams Galapagos Expd., 23-27 descriptions of new species, 32-45 Noctuinae, 27, 42-45 Williams Galapagos Expd., 27, 42-45 description of new species, 42-45 Noma, ship, route of, 2 Williams Galapagos Expedition, 2-20 Notaphus galapoensis (Geo. R. Waterhouse). 234 Nylanderia, 111 Nysius marginalis Dallas, 245 Ocypoda lateralis Freminville, 156 Ocypode, 154-155 albicans Bose., 155 albicans Rathbun, 155 arenarius Say, 155 guadichaudii Milne-Edwards and Lucas, 154-155 (Plate VII), 154 Odinia williamsi sp. nov., 90 description, 90-91 Odo insularis Banks, 98 Odynerus, 133 Oecophylla, 111 Oedemeridae, 235 Galapagos Is., 235 Olethreutidae, 31 Williams Galapagos Expd., 31 Olfersia, 91 fossulata Macquart, 91 intertropica Waker, 91 spinifera (Leach), 91 Olios galapagoensis Banks, 98 Olivier, G. A., 240 Ommatius, 87 marginellus Fabr., 87 saccas Walker, 87 Oniscidae, 185, 192-205 Galapagos Is., 185, 192-205 Oniscoidea, 181-183, 185, 189 Galapagos Is., 181-183, 189 Opossum, 3 Op sophy top sis insularis, 91 Opuntia helleri, 108, 112 Ormiscus variegatus G. R. Waterhouse, 238 Ornithodoros talaje Guerin, 95 Ornithomyia intertropica Walker, 91 Ornithoponus, 91 albipennis Say, 91 americana Leach, 91 americanus (Leach), 91 intertropicus (Walker), 91 Orphnaeus brevilabiatus (Newport), 137 Ortalidae, 89 Galapagos Is., 89-90 Orthaea, 246-248 bilobata Say, 248 insularis sp. nov., 246-248 description, 246-248 (Fig. 48), 247 Orthezia galapagoensis Kuw., 145, 146 Ortheziinae, 145 Galapagos Is., 145 Orthonotomyrmex Ashmead, 113 Oryctes, 238 Osborn, Herbert, Homoptera of the Williams Galapagos Expedition, 77-79 Otiorhynchus cuneiformis Geo. R. Water- house, 238 Owl Galapagos Islands, 11 Tick-flies from, 91 Oxacis, 54, 226, 227, 235 galapagoensis Linell, 235 Ozius verreauxii Saussure, 158 Paectes, 26, 38-41 indefatigabilis sp. nov., 26, 38-39 description, 38-39, (Plate I), 34 isabel sp. nov., 26 description, 39-40 (Plate I), 34 Palaemon, 168 fucorum Fabr., 168 jamaicensis Rathbun, 169 ritteri Holmes, 168 tenuicornis Say, 168 Pallas, Peter S., 240 Pandarus satyrus Dana, 213, 214 Pantomorus galapagoensis Linell, 230-231, 238 Paralebion elongatus Wilson, 213 Parapenaeus kishinouyei Rathbun, 161 Parapseudoryctes galapagoensis (Geo. R. Waterhouse), 237 264 Zoologica [Vol. V Parasitic Copepods from the Williams Galapagos Expedition, by Charles B. Wilson, 211-217 (Plate XX), 215 Pareuxesta, 89-90 hyalinata Coq., 90 intermedia Coq., 89 latifasciata Coq., 89, 90 obscura Coq., 90 Parrakeet, 3 Passalidae, 238 Galapagos Is., 238 Pectiniunguis albemarlensis Chamberlain, 139, 140 americanus Chamberlain, 139 Pedonoeces, bauri Linell, 237 costatus Geo. R. Waterhouse, 237 galapagoensis Geo. R. Waterhouse, 236 morio (Boheman), 237 pubescens Geo. R. Waterhouse, 229, 237 Pelecanus sp. lice from, 82 fuscus calif ornicus, 91 tick-flies from, 91 Pelican, Galapagos Islands, 11, 15 Pelican, brown Galapagos Is., 91 tick-flies from, 91 Penaeopsis kishinouyei (Rathbun), 161 “ de Man, 161 Penguin, 3 Pentacora rubromaculata Heid., 253 Pentatomidae, 53, 241 Galapagos Islands, 53, 241 Pentozocera, 133 Percheron, A., 240 Perigea, 24, 35, 36 apameoides Guen., 24 ebba sp. nov., 25, 36 description, 36 (Plate I), 34 ruthae, sp. nov., 24, 35 description, 35 (Plate I), 34 vacillans Walker, 36 Perixera ?, 28, 51 impudens Warr., 28, 51 Perkins, R. C. L., 133, 135 Petrel, British S.S., 85, 111, 219 visit to Galapagos Is., 85, 111, 219 Petrolisthes, 170 edwardsii Nobili, 170 edwardsii (Saussure), 170 Phalacridae, 236 Galapagos Is.. 236 Phalacris darwinii Chas. Waterhouse, 236 Phalacrocorax dilophus albociliatus, 84 lice from, 84 sulcirostris, 84 lice from, 84 Phalaena, rhexia Smith & Abb., 24 Phaleria manicata Boheman, 237 Phanias distans sp. nov., 98-99 description, 98-99 (Fig. 18), 98 Pheasants, kaleege, 57 Pheidole williamsi Wheeler, 108 var. seymourensis var. nov., 108 megacephala Fabr., 108 Phenacoccus Ckll., 147-148 parvus sp. nov., 147-148 Philaeus pacificus Banks, 98 Philatis Stal. cinerea sp. nov., 78 description, 78-79 major sp. nov., 79 description, 79 productus, 78 Philhydrus, 219, 221, 235 affinis, 219, 221 Philopteridae Burmeister, 82-84 Galapagos Is., 82-84 Philopterus Nitzsch breviformis Kellogg and Kuwana, 82 Philoscia, 181, 192-198 culebrae Moore, 194 culebroides sp. nov., 181, 193-194 description, 193-194 (Plate XII, figs. 11-12), 193 nomae sp. nov. 181, 196 description, 196 (Plate XIV, figs. 16-18), 196 williamsi sp. nov., 181, 194-196 description, 194-196 (Plate XIII, figs. 13-15), 195 Phlegathontius galapagensis nigrita Roth & Jord., 55 Phora, 88 scalaris Loew, 88 Phoridae, 88 Galapagos Is., 88 Phycitinae, 30. 46-48 Williams Galapagos Expd., 30, 46-48 description of new species, 46-48 Phytometra, 26 oo Cram, 26 Phytometrinae, 26, 41 Williams Galapagos Expd., 26, 41 description of new form, 41 Pierce, W. D., 132, 133, 135 Piesmopoda, 30, 47 carpasella sp. nov., 30, 47 description, 47 (Plate II), 40 Pigs, wild Galapagos Islands, 13 Piletocera, 29 bufalis Guen., 29 Pilocrocis, 28, 45 chathamalis sp. nov., 28, 45 description, 45 (Plate II), 40 cyclostigma, 45 ramentalis Lederer, 28 Pink eye (disease), 90 Planes minitus (Linn.) White, 157, 168 Plankton collected by Williams Galapagos Ex- pedition, 5 Plants collected by Williams Galapagos Ex- pedition, 5 Platynus galapagoensis (Geo. R. Water- house), 234 Platypodidae, 239 Galapagos Is., 239 Platypus santacruzensis sp. nov., 232-233, 239 description, 232-233 (Fig. 46), 232 Platystomidae, 238 Galapagos Is., 238 Plexippus paykulli Aud. et Sav., 98 Podisus (Arma), 241 sordidus Stal, 241 Poecilus, 234 Polyrhachis, 111 Pomace-flies, 90 Galapagos Ts., 90 Pomocentrids, 64 Ponera beebei sp. nov., 107 description, 107-108 (Fig. 21), 107 Ponerinae, 101 Galapagos Is., 101 Nos. 1-21] Index 265 Porcellana edwardsii Saussure, 170 Porcellio, 185, 197 laevis Latreille, 185 maculicornis Koch, 197 pruinosus Brandt, 197 Porcellionides pruinosus Brandt, 181, 196- 197 description, 196-197 (Plate XV, fig. 19), 197 Portunus, 158 ruber Lamarck, 159 sayi (Gibbes), 158 Poulton, Professor, 56 Prenolepis, 111 fulva Mayr. subsp. nesiotis Wheeler, 111 vividula Nyl., Ill subsp. guatelmalensis Forel var. edenensis . var. nov , 111 Prionace glauca 213 Prionids, 230 Prodenia, 51 Progne modesta, 56, 82 lice from, 82 Prosthetocirca cana, 92 Psallus insular is sp. nov., 250 description, 250 Pseuderiphia hispida, Milne Edwards, 158 Pseudococcus, 148-152 galapagoensis sp. nov., 148-150 description, 148-150 (Fig. 36), 149 insularis sp. nov., 150-152 description, 150-152 (Fig. 37), 151 Pseudocolobopsis Emery, 113 macilentus F. Smith, 114-119 Pseudoryctes Linell, 237-238 Psorya gen. nov., 27, 43 description, 43-44 hadesia sp. nov., 27 description, 44 (Plate II), 40 Pterophoridae, 30 Williams Galapagos Expd., 30 Pteroporid, 30 Pterostichus calathoides (Geo. R. Water- house), 234 Ptinids, 221 Pyralidae, 28-30, 45-48 Williams Galapagos Expd., 28-30, 45- 48 description of new species, 45-48 Pyralinae, 30, 46 Williams Galapagos Expd., 30, 46 description of new species, 46 Pyrausta, 29, 45 eneanalis sp. nov., 29, 45 description, 45 (Plate II), 40 Pyraustinae, 28-29, 45-46 Williams Galapagos Expd., 28-29, 45-46 description of new species, 45-46 Rathbun, Mary J., 153, 161, 162, 169, 170 Brachyuran Crabs collected by the Williams Galapagos Expedition, 153- 159 (Plate VII, Fig. 38), 154, 157 Red grouper, 211 Reduviidae, 250-251 Galapagos Is., 250-251 Remipes, 170, 171 adactylus denticulatifrons Ort., 170 testudinarius var. denticulatifrons Miers, 170 Repipta annulipes sp. nov., 250-251 description, 250-251 Reptiles collected by Williams Galapagos Ex- pedition, 3, 5 Rhipidius, 132, 133 denisi Chobaut, 132 Rhipiphoridae, 128, 129, 131 Rhipiphorus, 131 solidaginus Pierce, 132 (Fig. 32), 132 Rhopalocera, Galapagos Is., 55 Rhyncophora, 221 Galapagos Is., 221 Rhyscotinae, 198 Rhyscotus, 181, 197-201 laxus sp. nov., 181, 198-201 description, 198-201 (Plate XVI, figs. 20-22), 199 ortonedae Budde-Lund, 200 parallelus Dollfus, 200 Rhyzostylops inquirendus Silv., 132, 133 (Fig. 32), 132 Richardson, H„ 183, 184, 209, 210 Riley, C. V., 129, 136 Rivula ?, 27, 44-45 dubiosa sp. nov., 27, 44-45 description, 44-45 (Plate II), 40 Robber-flies, 87 Galapagos Is., 87 Rohwer, S. A. Hymenoptera collected by the Williams Galapagos Expedition, 173-175 Rose, Miss Ruth, 5, 35 records, notes and catalogues of Wil- liams Galapagos Expedition, 5 Runula albolinea sp. nov., 64-65 description, 64-65 (Fig. 11), 65 azalea, 64, 65 Saldidae, 253 Galapagos Is., 253 Sarcophaga, 88, 89 inoa Walker, 88, 92 obtusifrons Thomson, 89 reversa Aldrich, 89 violenta Walker, 88 Sarcophagidae, 88 Galapagos Is., 88 Sarcophagini, 92 Galapagos Is., 92 Sarcophagula occidua (Fabr.), 88 Sargassum crab, 157, 158 Sargassum shrimp, 161, 168 Sarothromyiops cinctus, 92 Sarrothripinae, 26 Williams Galapagos Expd., 26 Scarabaeidae, 221, 237-238 Galapagos Is., 221, 237-238 Scarites galapagoensis Linell, 234 Schaus, W., 52 Galapagos Heterocera, 22-48 new species, description of, 32-48 (Plates I & II), 34, 40 Scharz, 90 Schendylidae, 137, 139-141 Galapagos Is., 137, 139-141 Schioedte, J. C., 210 Schistocera melanocera, 53 Galapagos Islands, 53 Schistoceros cornutus, 237 subspecies galapaganus, 237 Schmitt, Waldo L. The Macrura and Anomura collected by the Williams Galapagos Expedition, 161-171 (Figs. 39-41 inch), 162, 164, 166 Scolopendra galapagoensis Bollman, 139 galapagoensis Chamberlain, 139 Scolopendridae, 139 Galapagos Is., 139 266 Zoologica [Vol. V Scohjtidae, 239 Galapagos Is., 239 Scorpionida, 95 Galapagos Is., 95 Screw- worm fly, 89 Scymnus galapagoensis Geo. R. Waterhouse, 236 Scytodes, 95 fusca Walck, 95 hebraica Simon, 95 Scytodidae, 95 Galapagos Is., 95 Sea-lion Galapagos Islands, 9, 11, 14, 20 (Fig. 5), 9 Selenophorus galapagoensis (Geo. R. Water- house), 235 Sericosema, 28, 51 lignata Warr., 28, 51 Serranus, 211 Saymour Islands Galapagos Archipelago, 17, 18 Shark, 213, 214 blue, 213 Sharp, Dr. David, 173, 225, 237, 240 Shoemaker, Miss Amalia, 143 Shrimp fresh- water, 161, 169 sargassum, 161, 168 Sicaroides ultriformis Butler, 95 Silphidae, 235 Galapagos Is., 235 Silvestri, F., 132, 133, 136 Simopone marleyi Arnold, 103 Sira, 76 Sitarini, 128, 129 Sitaris, 129 Smith, F., 85, 111 Snakes, Galapagos Is., 54 Snodgrass, R. E., 81, 83, 85, 140 Solenopsis globularia F. Smith., 108 subsp. pacifica Wheeler, 108-109 description, 108-109, (Fig. 22), 109 Solpugida, 95 Galapagos Is., 95 Sore eye (disease), 90 Sparassidae, 98 Galapagos Is., 98 Sphingidae, Galapagos Islands, 51, 54-55 Spiders, Galapagos Is., 93-99 Spragueia, 25, 38 creton sp. nov., 25, 38 description, 38 (Plate I), 34 plumbeata sp. nov., 25, 38 description, 38 (Plate I), 34 Squash bugs Galapagos Islands, 53 Stal, 241 Stanford Galapagos Expd., 137, 140 Staphylinidae, 235 Staphylinids, 219, 221 Stenocephalus insularis Dallas, 241 Stenodontes molarius (Bates), 230, 238 Sterrhinae, 28 Williams Galapagos Expd., 28 Stichotrema, 133 Stomion, 228, 229, 236 bauri Linell, 236 galapagoensis Geo. R. Waterhouse, 228- 229, 236 helipioides Geo. R. Waterhouse, 236 laevigatum Geo. R. Waterhouse, 228, 236 Sir epsicr ales, 31 smithiana Walsingham, 31 Strepsiptera, 125-136 Stylops, 133 californica Pierce, 133 Sula nebouzii, 82, 84 lice from, 82, 84 piscator, 82, 84 lice from, 82, 84 Surgeon fish, 64 Sylepta, 29 elevata Fabr., 29 gordialis Guen., 29 silicalis Guen., 29 Synalphens, 162-163 fritzmulleri, 163 neptunus Rathbun, 162 nobilii Coutiere, 162 (Fig. 39), 162 sanlucasi Coutiere, 162, 163 Syngrapha, 26 egena galapagensis form nov., 26 description, 41 (Plate II), 40 Syrphus clavata Fabr., 88 Syrphidae, 88 Galapagos Is., 88 Tabanidae, 87 Galapagos Is., 87 Tabanus lineola Fabr., 87 vittiger Thomson, 87 Tacky s beebei sp. nov., 223-224, 234 description, 223-224 (Fig. 42), 224 Tachysphex galapagensis sp. nov. 173-174 description, 173-174 Taeniorhynchus, 87 niger Gibbs, 87 Tagus Cove, Albemarle Is., 14, 15 Tanaidae, 183 Galapagos Is., 183 Tanais stanfordi Richardson, 183 Tanypus, 86 Tapinoma melanocephalum Fabr., 110 Tee-Van, Mr. photographs and moving pictures from Williams Galapagos Exp., 5 Tenebrionidae, 221, 233, 236-237 Galapagos Is., 221, 233, 236-237 Terns Galapagos Islands, 15 Tessaromma, 237 Tetramorium guineense Fabr., 109-110 Tetrapriocera longicornis (Oliver), 237 Teuthis crestonis (Jordan & Starks), 64 Theridiidae, 97 Galapagos Is., 97 Thermesia, 27 gemmatalis Hubn., 27 Thermonectes basilaris (Harris), 225, 235 Thomson, C. G., 85, 89 Thunnus, 184, 189 Thysanura, Williams Galapagos Expd., 67 Tingidae, 251-253 Galapagos Is., 251-253 Tipulidae, 86 Galapagos Is., 86 Tortoise Galapagos, 19 Townsend, Dr. C. H. T„ 91, 92 new genera and species of Diptera from Galapagos Is., 91-92 Trachea, 24, 33 roseae sp. nov., 24, 33 description, 33-35 (Plate I), 34 Trachinotus paloma, 189 Triungulin Larvae from the Williams Gala- pagos Expedition, by Charles T. Brues, 125-136 Nos. 1-21] Index 267 bibliography, 134-136 (Figs. 28-32 inch), 126, 127, 131, 132 Tropidurus, Galapagos Is., 54, 95 Tropisternus Solier, 219, 225, 235 lateralis Fabricius, 225, 235 Trox, 221, 229, 230, 238 Seymour ensis sp. nov., 229-230 description, 229-230, 238 suberosus Fabricus, 221, 229, 238 Tunnies, 184, 189 Tylidae, 189 Galapagos Is., 189 Tylos, latreilli Audouin and Savigny, 181 description, 189-192 (Plates X-XI, figs. 6-10), 190, 191 spinulosus Dana, 192 Uca heterochelos, 153 galapagensis Rathbun heterochelos Kingsley, Lamarck, Rath- bun, 153, 156 Urocaris longicaudata (Stimpson), 169 Utetheisa, 23, 51 galapagensis Wallgrn, 23 ornatrix Linn., 23, 51 Valvifera, 184 Galapagos Is., 184 Van Dyke, Dr. E. C., 222 Van Name, Willard G. Isopods from the Williams Galapagos Expedition, 181-210 bibliography, 209-210 (Plates VIII-XIX inch), 186, 188, 190, 191, 193, 195, 196, 197, 199, 202, 204, 207 Verhoeff, Dr. Karl W., 189 Villa, 87 primitiva (Walker), 87 tincta (Thomson), 87 Volcanoes, Galapagos Islands, 14, 19, 20 Walker, Francis, 85, 87, 89 Wallace, 233, 234 Warren, W., 51 Waterhouse, Charles, 221, 224, 225, 237, 238, 240 Waterhouse, George R., 219, 221, 225, 228, 229, 240 Waterhouse types (of Coleoptera) in British Museum, 222 Westermann, B. W., 136 Westwood, J. O., 136 Wheeler, William Morton The- Formicidae of the Harrison Wil- liams Galapagos Expedition, 101-122 (Figs. 19-27 incl.), 102, 105, 107, 109, 110, 114, 117, 118, 120 list of the Galapagos Formicidae with their distribution, 122 Wheeler, Prof. W. M., 17, 67, 69, 73, 128, 222 Williams Galapagos Exp., 17, 67, 73, 101-122 Williams, Francix X., 51, 54, 103, 109, 110, H2, 116, 221, 222, 240 ^ Coleoptera collected by, on Galapagos w Is., 222 Williams Galapagos Expedition, 2-20, 48, 59, 65, 76, 79, 84, 92, 99, 121, 134, 141, 152, 171, 175, 180, 208, 217, 239, 254 collections of, 23-254 Anomura, 161-171 ants, 101-122 Apterygota, 67-76 Arachnida, 93-99 beetles, 125-136, 219-240 birds, 3, 14, 15, 19, 54, 55-57 Brachyuran Crabs, 153-159 bugs, 241-254 centipedes, 137-141 Chilopods, 137-141 Coccidae, 143-152 Coleoptera, 125-136, 219-240 copepods, 211-217 crabs, 153-159, 161-171 Crustaces, 153-159, 161-171, 181- 210, 211-217 Diptera, 85-92 fish, 62-65 Formicidae, 101-122 Hemiptera, 53, 241-254 Heterocera, 23-48, 51-59 Homoptera, 77-79 Hymenoptera, 173-175 ichthyology, 62-65 insects, 23-48, 51-59, 67-92, 101- 122, 125-136, 143-152, 173-175, 177-180, 219-240, 241-254 Isopods, 181-210 Lepidoptera, 23-48, 51-59 Macrura, 161-171 Mallophaga, 81-84 mammals, 3, 13, 14, 20 mosquitoes, 53, 87 moths, 23-48, 51-59 Neuroptera, 177-180 parasitic copepods, 211-217 reptiles, 3, 8, 10, 15, 19, 54 snakes, 54 spiders, 93-99 triungulin larvae, 125-136 Williams, Harrison, Esq., 3, 32, 43, 63, 101, 103, 137 Wilson, Charles B. Parasitic Copepods from the Williams Galapagos Expedition, 211-217 (Plate XX), 215 Wohlfahrtia inoa Walker, 88 Wreck Bay, Chatham Is., 15 Xanthias insculpta Stimpson, 153 insculpta (Stimpson), 153, 157 insculptus Rathbun, 157 (Fig. 38), 157 Xanthodius lobatus (Milne Edwards), 158 Xenos nigrescens Brues, 133 Xylocopa, 129, 130, 131, 173 aurulenta Fabricius, 173 colona Lepeletier, 173 or pi f ex, 131 tenuiscapa Westw. 130 transitoria Perez, 125 (Fig. 28), 126 violacea, 105, 112 Zanon, V., 136 Zelotes galapagoensis Banks, 96 Zinckenia, 28 fascialis Cramer, 28 Zonitini, 128, 129 Zonitis bilineata Say, 128 Zoological Park, collections from Williams Galapagos Expedition, 3 Nrtu fork Zaalagiral ^orirty OBJECTS OF THE SOCIETY A Public Zoological Park. A Public Aquarium. The Preser- vation of our Native Animals. The Promotion of Zoology. ZOOLOGICA VOL. 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II 1— Two Series of Amphibians (Deckert) 25 2 — Tetrapteryx Stage, Ancestry Birds (Beebe) 25 3— 4 — Notes, Birds, Para, Brazil; Fauna of Jungle Debris (Beebe) 25 5 — The Gaff-Topsail Catfish (Gudger) 25 6 — Mammals of Australia in the Zoological Park (W. H. D. Le Souef) .75 7— 8-9 — Vertebrates, British Guiana (Beebe).. 25 10 — Habits of Sage Grouse (Horsfall) 25 11 — Eclipse Plumage, Domestic Fowl (Crandall) .25 12 — Life History of the Puffer (Welsh-Breder) 25 13 — Hermaphroditism of the Croaker (Breder) 25 14 — Food of Certain Minnows (Breder-C raw ford) 25 15 — Fishes of Sandy Hook Bay (Breder) 25 16 — Weaving of Weaver Birds (Friedman) 25 17 — Scales of Whitefishes (Mellen-V an Oosten) 30 18 — The Anderson Tree Frog (Noble) 25 Postage, Zoologica, 5 cents ZOOLOGICA VOL. Ill 1-2 — Objects, Tropical Research Station ( Osborn ) Contributions Tropical Research Station (. Beebe ) 3-11 — Social Beetles, British Guiana ( Wheeler ) Tachigalia Ants ( Wheeler ) Habits of Cucujidae ( Wheeler ) New Coleoptera ( Schwarz-Barber ) Larvae, Pupae, Social Beetles ( Boving ) New Diadiplosis (Felt) New Blepyrus (Brues) Tachigalia Membracids ( Herbert Osborn) New Entomobrya (Folsom) 12 — Fetuses, Guiana Howling Monkey (Schultz) 13 — Mammals Collected Tropical Research Sta. (Anthony) . . 14— 15 — New Batrachians (Noble). New Lizards (Noble) . . . 16 — Descriptiones Termitum in Anglorum Guiana Report- orum (Silvestri) 17 — New Genera and Species, Termitophilous Coleoptera (Mann) 18 — Glandular Structure, Abdominal Appendages, Termite Guest (Mclndoo) 19 — Termitophilous Apterygota (Folsom) 20 — Three Apparently New Species, Termitaphis (Morrison) 21 — Termitophilous Millipeds (Chamberlin) 22— 23 — A Termitophilous Braconid (Brues). Two Myrme- cophilous Phoridae (Brues) Postage, Zoologica, 5 cents Paper .25 1.00 .25 .25 .25 .75 .25 ZOOLOGICA VOL. IV 1— Fishes, Amphibians, Reptiles (Breder) 15 2 — Frozen Mammoths (Lang) 25 ZOOLOGICA VOL. V 1 — Williams Galapagos Expedition (Beebe) 2 — Galapagos Heterocera (Schaus) 3 — Galapagos Lepidoptera (Beebe) 4— 16 — Ichthyology (Nichols) Apterygota (Folsom) Homoptera (Osborn) Mallophaga (Ewing) Diptera (Johnson) Arachnida (Banks) Formicidae (Wheeler) Triungulin Larvae (Brues) Chilopods (Chamberlin) Coccidae (Morrison) Brachyuran Crabs (Rathbun) Macrura and Anomura (Schmitt) Hymenoptera (Rohwer) 17 — Neuroptera (Banks) 18 — Isopods (Van Name) 19— Copepods (Wilson) 20 — Coleoptera (Mutchler) .15 .20 .15 .50 .15 .25 .15 .20 Cloth ZOOLOGICA VOL. VI 1 — Ecology of Kartabo (Beebe) 2 — Birds of Kartabo (Beebe) 3 — Membracidae of Kartabo (Haviland) 4 — Termites of Kartabo (Emerson) Publications of the Society for sale at the Zoological Park, 185th Street and Southern Boulevard, New York City.