I! jl ii m 3- m m '. un i O = CD CD lo ALASKA VOLUME XIV * SMITHSONIAN INSTITUTION HARRIMAN ALASKA SERIES VOLUME XIV Monograph of the Shallow-water Starfishes of the North Pacific Coast from the Arctic Ocean to California (WITH 110 PLATES) BY ADDISON EMERY VERRILL Professor Emeritus of Yale University PART 1. TEXT (PUBLICATION 2140) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1914 jE.crb Q^-afHmcrc (p »iLTlMORb, Ml'., V. S. J. . EDITOR'S PREFACE. The Starfishes collected by the Harriman Alaska Expedition were sent to Professor A. E. Verrill for study and report ; and through the courtesy of various museums of the United States and Canada numerous additional specimens from the West Coast were placed in his hands. With this rich material at his disposal, Professor Verrill undertook a monographic revision of the Starfishes inhabiting the shallow waters of the Pacific from Arctic Alaska sou.th to California. In the course of three or four years (by 1904) the larger part of the monograph had been written, most of the species (including Stimpson's types, in the United States National Museum) had been photographed, and more than 40 plates had been reproduced and printed. But pressure of other duties so delayed the completion of the work that a dozen years have elapsed since it was begun. The long delay, however, has not been without compensations, for much new material has come to Professor VerrilPs hands, including col- lections from Patagonia and Fuegia, enabling him to add matter of great importance on the geographic distribution of the group. In the meantime, Mr. E. H. Harriman, organizer and patron of the Expedition, having passed away, the publication of the work, with its large series of plates, has been generously provided for by Mrs. Harriman. In view of Professor Verrill's lifelong studies of the Starfishes, it is believed that the present monograph will long remain a standard authority on the subject, and that it will receive a hearty welcome from all workers on the Echinoderms. C. HART MERRIAM, Editor. WASHINGTON, D. C., May, 1913. SHALLOW- WATER STARFISHES OF THE NORTH PACIFIC COAST FROM THE ARCTIC OCEAN TO CALIFORNIA TABLE OF CONTENTS. PAGE Preface xi Introduction I Richness of the Starfish Fauna ; Causes I Food and Feeding Habits of the Starfishes 3 Activities of Starfishes ; Migrations ; Rate of Travel 4 Larval Stages; Protection of Eggs and Young; Incubation 7 Senses ; Instincts ; Memory ; Acquired Habits 9 Peculiarities of the Starfish Fauna of the Northwest Coast of America n Multiplicity of Rays ; Variability 12 Subspecies and Varieties 17 Hybridism of Associated Species 18 Faunal Relations and Distribution of the Species 18 Morphological Features of the Class Asterioidea 20 Classification of the Asterioidea 24 Order Forcipulosa or Forcipulata 24 Various Kinds of Pedicellarise 25 Families and Subfamilies of Forcipulosa 26 Family Asteriidae 27 I. Morphology and Growth of the Skeletal Ossicles (Text-fig, i). .. . 27 II. Pedicellariae of the Asteriidae 30 III. Relative Importance of Morphological Characters in Asteriidae.... 31 A. Modes of Growth and Increase in Number of Skeletal Plates 32 B. Interactinal Plates and Spines 33 C. Form and Arrangement of the Dorsal Ossicles 34 D. Dorsal Spines ; Form and Arrangement 34 E. Pedicellariae ; Special Forms (Text-fig. 2) 35 F. The Madreporite or Madreporic Plate and surrounding Spines 36 G. Jaws and Oral Spines (Text-fig. 3) 36 H. Adambulacral Spines ; Arrangement 38 I. Number of Rays ; Variability 38 J. Arrangement of the Ambulacral Feet or Sucker-tubes (Podia) 39 K. Modes of Development ; Brooding of the Young ; Position of the Genital Pores 39 Classification of Asteriidae 39 Subfamily Heliasterinae 40 Subfamily Pycnopodiinae 40 Subfamily Stichasterinae 40 Subfamily Asteriinae 42 A. Generic Subdivisions of Asteriinae 43 B. Subdivisions of the Asteriinae having Definite Longitudinal Rows of Dorsal Plates and Spines 44 C. Table of Extralimital Species of Coscinasterias and closely Allied Forms 46 D. Observations on Various Genera 46 (ix) 53 X TABLE OF CONTENTS PAGE E. Table of Principal Genera of Asteriinse with Regular Longitu- dinal Rows of Dorsal Plates 49 F. Subdivisions of Asteriinse with Reticulated Dorsal Skeletons. . 50 G. Analytical Table of the Generic and Subgeneric Groups of Northern Asteriinse having Reticulated Dorsal Skeletons 54 H. Asteriidse of the Northwestern Coast of America 55 I. Analytical Table of Genera, Subgenera, Species, and Subspecies of Asteriidae from the Northwest Coast of America 56 Asteriinse : Descriptions of Genera and Species 67 Subfamily Pycnopodiinae 19? Descriptions of Pycnopodiinse IQ7 Descriptions of Pedicellasteridae 202 Order Spinulosa 203 Morphology 203 Classification 204 Suborders Avelata and Velata 204 Families of Spinulosa 204 Descriptions of Families, Genera, and Species 205 Order Phanerozona 280 Morphology 280 Classification 282 List of Families 282 Suborder Valvulosa or Valvata 284 Descriptions of Families, Genera, and Species 285 Suborder Paxillosa 3*3 Descriptions of Families, Genera, and Species 314 Species erroneously attributed to the Fauna 337 Geographical Distribution 337 I. The Beringian Fauna 338 II. The Columbia-Alaskan Fauna 34° III. The Californian Fauna 343 IV. The South Californian Fauna 345 V. Some Comparisons with Other Faunae 347 VI. Relations of the Columbia-Alaskan and Californian Faunae 348 VII. Relations of the South Californian Fauna 349 VIII. Patagonian, Fuegian and Antarctic Shallow-water Starfishes contrasted with Those of the North Pacific Coast 351 IX. List of Patagonian and Fuegian Starfishes Found in 50 to 500 Fathoms ; mostly Related to Northern or Arctic Species 367 X. Lists of Extralimital Species Partially Described, Revised, or Figured 37° 1. North Atlantic and Arctic Species 370 2. Patagonian, Fuegian, Antarctic, and South Atlantic Species 371 3. Species from the Panamic Fauna 372 4. Indo-Pacific, Australian, and Japanese Species 373 XL List of New Genera Established 374 Bibliography 374 Index 389 PREFACE. THIS report was originally planned to include only the littoral and shallow-water species of the starfishes of Alaska and adjacent waters contained in the fine collections made on the Harriman Expedition by Dr. (now Professor) W. R. Coe, together with the collections in the Yale Museum, previously received from the same region, including a number of my original types. The subsequent reception of numer- ous other large collections from the same and adjacent regions soon led to a great extension of the work, so as to include all the shallow- water species from California to Bering Sea. The various collec- tions sent to me by the Canadian Geological Survey through the late Mr. J. F. Whiteaves, and later through Professor John Macoun have been particularly large and valuable, containing many new forms. My thanks are especially due to Professor Macoun and other mem- bers of the Canadian Survey for these favors. A small but very important collection was sent to me from the Museum of Comparative Zoology, through the kindness of the Director, Mr. Samuel Henshaw, and of Dr. H. L. Clark; I also enjoyed an opportunity to study there the entire collection of star- fishes from the Northwest coast. A large collection was also received, through Dr. C. F. Newcombe, from the Provincial Museum of British Columbia at Victoria. Professor Trevor Kincaid, University of Washington, Seattle ; Pro- fessor W. E. Ritter, University of California, Berkeley, and others have also sent useful collections ; and to all these I wish to express my obligations for assistance given. A small collection, mostly from Bering Sea, was sent to me by the U. S. National Museum, and was very useful for the more Arctic species. I am also indebted to the National Museum, through the kindness of Dr. R. Rathbun and Miss M. J. Rathbun, for photo- graphs of the type specimens of the several species early described by Dr. Wm. Stimpson, which have been of great use and are now reproduced on my plates. Several of these have not been previously figured. The report was very nearly finished, mostly in its present form, previous to 1904, and forty-three plates had been engraved and printed at that time. (xi) Xll PREFACE In the meantime, Prof. W. K. Fisher, of Stanford University, has published (1911 b) an extensive work on some of the orders of starfishes of the North Pacific, including not only very many deep- water species, but also considerable numbers from the shallow waters of the same regions, covered by my report, and including many of the species previously described and figured by me, in this work. He had the advantage of having studied my specimens, manu- scripts and plates several years ago, while visiting me at the Yale Museum. Yet in some cases our conclusions are somewhat divergent. I have not seen his types, nor have I been able to devote so much time as I would like to the correlation of his species of certain difficult genera, like Solaster and Henricia, with my own. But I trust that the figures and descriptions given in both works will enable future students to understand what the forms discussed really are, regardless of the names. For the numerous excellent drawings of miscroscopic details and for nearly all the photographs, I am indebted to the careful work of my son, Mr. A. Hyatt Verrill, who has devoted much time to the work during several years. ADDISON EMERY VERRILL. NEW HAVEN, CONN., July, 1912. Monograph of the Shallow-water Starfishes of the North Pacific Coast from the Arctic Ocean to California, with Revisions of Various Extra- limital Genera and Species. BY ADDISON EMERY VERRILL. Professor Emeritus of Yale University. INTRODUCTION. RICHNESS OF THE STARFISH FAUNA; CAUSES. THE Northwestern Coast of North America seems to be the head- quarters or metropolis of shallow-water starfishes. No other region, so far as known, has anything near so many species, nor so many generic and family types, in shallow water, nor so great an abun- dance of individuals. The species and genera confined to the deeper waters off these shores seem to be equally numerous, or more so. At present there are over one hundred shallow-water species known from that region, besides about twenty named varieties. The abundance and variety of starfishes in these shallow waters is apparently greater than at Panama, the West Indies, or the most favorable parts of the East Indies in similar depths. This profusion of starfishes does not hold good for the other classes of Echinoderms. The Sea-urchins (Echinoidea) are repre- sented by comparatively few species, of which only about five are common, and those belong to widely distributed genera. But this is about the normal number for any similar cold-water fauna. The serpent stars (Ophiuroidea) are represented by about the usual number of species, none of them of unusual types. The Crinoidea are represented in shallow water only by a single large, comatulid species, so far as known to me. The Holothurians are numerous, but not remarkably so, and mostly of common genera. This coast seems admirably adapted for the permanent occupation and evolution of starfishes, and similarly for certain groups of other marine invertebrates and fishes. 2 (i) 2 VERRILL Among the fishes the Salmonidae may be mentioned. Among the mollusca some groups, like the limpets, chitons, Buccinidae, and other families, are much more abundant than in any other regions. Nemerteans and tubicolous Annelids are also remarkably large and numerous. There is every reason for thinking that such groups, including various genera and families of starfishes, have lived and flourished on that coast for many geologic periods, and have developed in that region, by continuous evolution, most of the species found there at present, as well as great numbers that have disappeared. Indeed, I am led to conclude that this region has been, for long ages, a center of evolution, from which various generic and specific types of starfishes have been dispersed to other faunal areas, with or without subsequent changes. It is not at all improbable that some arctic and boreal species of the North Atlantic originated on the Alaskan coast and have since crossed to the Atlantic by way of the Arctic Ocean, when the arctic climate was warmer. Others may have migrated southward to the subtropical and tropical regions of Mexico and Central America, and even to the Antarctic Seas, for some of the deep-water Patagonian and antarctic species show peculiar affinities with the Alaskan species. Such migrations, across the tropics, would easily be possible for species having a considerable range in depth, for their congenial temperatures could easily be found at no great depth, even off Panama. One reason for the great variety and persistence of the starfishes and other groups on this coast is, no doubt, the fact that the tem- peratures of the waters were not seriously reduced during the Glacial Period, while those of the North Atlantic were rendered so cold that the preexisting fauna was largely exterminated, while the survivors were driven southward, except in the case of eminently arctic and deep-sea species. Another very favorable condition is the comparatively equable temperature of those waters, due to the reflex of the great Japanese tropical current of water. Its influence in modifying the climate of the whole coast, south of the Aleutian Islands, at least, is well known. For the prosperity of most marine animals, equability of temper- ature is exceedingly favorable, whether high or low, especially at the breeding season. SHALLOW-WATER STARFISHES 3 No doubt the comparative uniformity of the temperatures at all seasons, and over vast areas, in the deep sea, is the principal cause of the great abundance of starfishes and many other groups of animals in the deep sea and of their wide distribution. Constancy of temperature at the breeding season, year after year, is of the greatest importance to nearly all animals, both in the sea and out of it, as the writer pointed out many years ago in the case of birds,1 and as has since been confirmed by other investigators for birds and other groups. Another very favorable condition, for the multiplication of star- fishes on this coast, is the vast extent and broken condition and rocky character of the coast-line, with innumerable islands, bays, straits, fiords, and inlets, affording every variety of stations and any amount of shelter from severe storms, and at the same time furnishing innumerable suitable stations on the rocky shores, for the growth and increase of all sorts of plant and animal life, on which the star- fishes may feed. It has been stated that the coast-line of Alaska and the adjacent islands exceeds 26,000 miles, or more than the circum- ference of the earth. That of British Columbia is also of vast extent. FOOD AND FEEDING HABITS OF THE STARFISHES. The littoral and shallow-water starfishes are nearly all carnivorous, by preference, and feed very largely on the barnacles and mollusca that live among, or attached to, the rocks, such as mussels, oysters, limpets, chitons, small spiral gastropods, etc. Considering the great numbers and large sizes of many of the starfishes, the wonder is that they have not already entirely exter- minated those mollusca on which they largely feed. The vast amount of damage done to the cultivated oyster beds on our Atlantic coast by a single small species of starfish (Asterias forbesi) is well known. What then must be the destruction wrought to the bivalves on the Pacific coast, where there are some forty related species, with similar habits, many of them becoming over two feet across ? One of our native starfishes, six inches across, will eat over twenty small oysters in one day. Probably it would take a hundred oysters or mussels to satisfy one of the giant starfishes of Alaska or Puget Sound. 1 Proceedings Boston Soc. Nat. Hist., vol. x, p. 259, 1866. 4 VERRILL The great abundance, size, and voracity of these Pacific starfishes will always be a great hindrance to the cultivation of oysters, mus- sels, etc., on that coast, except, perhaps, where the water is dis- tinctly too brackish for starfish life. Fortunately for the mollusca, at least, most of the larger littoral starfishes are more or less cannibals. When opportunity occurs, they do not hesitate to devour each other. From the stomach of a large ten-armed Solaster, I have taken the half-digested ray of an Asterias that must have been a foot across when living. An inch or two of the ray still protruded from the mouth of the Solaster, for it was much too large to swallow entire. The various kinds of starfishes that inhabit rocky shores, clinging to the rocks, have large numbers of strong, muscular sucker-feet, each tipped with a perfect sucker for adhesion. In the ordinary five-rayed kinds, six to eight inches across, there will be four close rows of such suckers the whole length of the ray, perhaps 200 to a ray; or 1,000 on the five rays. In the case of the many larger five-rayed and six-rayed kinds, two feet across, the number of suckers may be 4,000 to 8,000 or more. How many there are on the big twenty-rayed or twenty-four- rayed star of that coast, which becomes over thirty inches across, and has the suckers mostly crowded into four rows, on each ray, nobody knows, for apparently nobody has had the patience to count them. Probably there may be more than 40,000. I refer to the great Pycno- podia or " sun-star." (See pi. xxix.) Such starfishes are able to open an oyster or mussel by a long and steady pull with these suckers, while the rays are wrapped around the victim. All such starfishes can evert the large, loose, bag-like stomach and wrap it around its prey till digestion is completed, if it be too large to swallow entire. But the mouth is also very extensible and dilatable, so that they can swallow objects sur- prisingly large. I have often taken sea-urchins an inch or more in diameter with the spines nearly all in place, from the stomachs of starfishes of no more than ordinary size. ACTIVITIES OF STARFISHES ; MIGRATIONS ; RATE OF TRAVEL. Although most starfishes, as ordinarily seen in life clinging to stones, etc., appear very sluggish and slow in their motions, they are really able to travel to considerable distances and undoubtedly, in some cases, make migrations of considerable extent, either in SHALLOW-WATER STARFISHES 5 search of food or to seek more congenial temperatures in deeper or shallower water, or for other reasons, such as breeding habits. In Long Island Sound it is a common experience of the oyster cultivators to find that many thousands of starfishes (Asterias for- besi) suddenly appear on certain oyster grounds which had been nearly or quite free of them a few days previously. If undisturbed for a few weeks they may destroy thousands of bushels of oysters and then as suddenly disappear from the devastated grounds, to visit some other locality. In the autumn, September to November, as I have myself observed, year after year, at Outer Island, of the Thimble Islands group, in Long Island Sound, they migrate upward from deeper to shallower water, and may then be found in myriads on rocky shores between tides, where none were to be seen a few days before. At such times large numbers are often caught and destroyed in the interests of the owners of oyster grounds in the vicinity. In some seasons a boy, one of my grandsons, often caught during low tide two or three bushels, morning after morning, on one small island, up to a total of fifteen or sixteen bushels, and more still came up every morning. These were mostly not more than half-grown and a bushel would usually count up to 1,000 or more. The same operation was carried on at many adjacent islands by others, with similar results. The effect of this wholesale slaughter was said to have notably decreased the loss of oysters on the oyster grounds of the district during the next year.1 At this time the starfishes were feeding mainly on the young oysters of the season, which had " set " thickly on all the rocks and ledges, below half-tide, but they were also, in many cases, feeding on the rock barnacles (Balanus balanoides), which were also abun- dant on the rocks. Later in the season they disappear, going into deeper water to avoid freezing temperatures. I have repeatedly tested their rate of travel, on the shores of Outer Island. By placing a hundred or more in a small pool of water left by the tide at about half-tide mark, it was easy to ascertain how far and in what directions they would travel during 1 Immense numbers of this same starfish are also taken in Long Island Sound and other oyster-growing waters by dragging " tangles " or " swabs " over the oyster beds, as first recommended by the writer in 1873 and 1876. A single oyster steamer by this means has often taken twenty-five barrels or more in a short time. O VERRILL about six hours, while the tide was above the outlet of the pool, which was on a nearly smooth, gently sloping, granite ledge. In brief, nearly all left the pool, and travelled downward, but in irregularly divergent lines, to various distances, varying from one to about fifteen feet. The majority followed a nearly straight course down the more sloping parts of the ledge, but were readily diverted to one side or the other by small variations in the slope. Those that travelled the greater distances probably left the pool soonest, and were perhaps in more active condition. Selecting a dozen or so of the best travellers, they might average ten feet to twelve feet in six hours, or nearly two feet an hour, providing they left the pool about as soon as it was covered by the tide, which was probably true of some of them, at least. As their motions are rather deliberate and continuous, they could, no doubt, keep up this rate for many hours, or about 336 feet in a week. Yet these were mostly rather small, from 4 to 6 inches across, and the larger ones were found to travel much faster than the young ones, for they have a much larger number of sucker-feet, which are also stronger and longer. Such observations as the above give, of course, only an approxi- mate idea of their rate of travel. These were neither impelled by hunger nor by cold, but merely by an instinctive tendency to seek a lower level. A stronger stimulus might have increased the speed. Very large starfishes, like many of the west coast species, would probably travel very much faster, and perhaps those with many rays and a vast number of suckers, like Pycnopodia and Solaster, would be still more speedy. By actual count a half-grown Pycnopodia was found by the writer to have over 22,000 ambulacral feet. At any rate, the observed speed would be quite sufficient to permit them to ascend or descend a sloping sea bottom quickly enough to avoid any ordinary changes in the temperature of the water liable to be injurious, and also sufficient to enable them to seek their ordinary food on new grounds. It is known that, in spite of their voracity, they can go for many days, or even weeks, without any food and still remain active and in good health. However, there are other families of starfishes that are far more lively, while some others are far more sluggish. The most lively starfishes known to me belong to the genus Luidia. Some of these are very large species. These live on bottoms of fine sand or mud, more commonly in rather deep water, but often close to the shore in sheltered, shallow, SHALLOW- WATER STARFISHES 7 sandy bays and harbors. Some of the species have more than six rays. In all cases the rays are long, flattened, and very flexible. Ordinarily when at rest they lie just beneath the surface of the sand, which soon forms into star-like impressions, agreeing with the star- fish in size and form. This is due to the minute currents of water produced by the ciliary motions connected with the respiration of the starfish by means of the dorsal papulae. When disturbed, the starfish glides away quickly, just under the loose superficial sand, using its " sucker-feet " or " ambulacral feet " as paddles for swimming or gliding. These ambulacral feet are much larger and longer than usual and are much flattened trans- versely, and have no terminal sucker, but end in a point. They are very muscular and can be waved back and forth, like paddles, in unison, so that the motion is more like rowing or swimming than running or creeping. While under the sand, no doubt the pressure against the sand causes them to act something like legs or feet ; but when placed in an aquarium without sand, they can swim or glide along the bottom and up the perpendicular sides with surprising rapidity. My own observations on this genus were made in 1901, at Ber- muda * on Luidia clathrata Say, a five-rayed species. Similar observations have since then been made by others on different species of Luidia, so that this mode of locomotion is prob- ably common to all the species of the genus. LARVAL STAGES; PROTECTION OF EGGS AND YOUNG; INCUBATION. The wide and rapid distribution of many, if not most, starfishes of shallow seas is, however, due to the fact that nearly all produce vast numbers of minute eggs, which develop into small, singular, bilateral, free-swimming larval forms (bipinnaria, brachiolaria) that require many days or weeks to go through their metamorphoses before set- tling down to the bottom in the starfish form. While in these larval stages, they may be drifted long distances by waves and currents and finally settle down in places far from their place of origin. Of course, in most cases they become widely dispersed and vast numbers perish, but sometimes, on the other hand, myriads of young may happen to drift along in company, by the action of a current, and so finally locate as a vast colony in a new home. 1 A. E. Verrill, 1901, p. 36, and Zoology of Bermuda, Article 10, p. 36, 1903. (See Bibliography.) 8 VERRILL Perhaps the sudden appearance of great numbers of young star- fishes on certain oyster grounds, where they were not found pre- viously, may be accounted for in this way. When well fed on young oysters, these little baby starfishes grow rapidly and soon become large enough to attract attention and do great mischief, even in one season.1 Certain genera and species of shallow-water starfishes, and per- haps a large proportion of the deep-sea species, do not produce so many minute eggs, nor do these develop into free-swimming larvae. On the contrary, the mother retains the eggs and cares for them till they pass through an abbreviated metamorphosis and develop into minute young starfishes provided with " sucker-feet " or podia to enable them to take care of themselves. Commonly, in such cases, the eggs and young are held under and around the mouth in large clusters. During the time required for their development the mother appears to be incapable of taking any food, owing to the obstruction of the oral region. The young are often attached by a larval actinal pedicel in clusters or strings. This is the mode of carrying the eggs and young observed in sev- eral species of Henricia, Anasterias, Sporasterias, and Podasterias, and in numerous small species of Asteriidse, belonging to the genus Leptasterias. I have personally observed this in L. compta, L. tenera, and L. littoralis of the New England coast, and in L. epichlora and cribraria of Alaska. The young of epichlora were collected by Dr. W. R. Coe. (See pi. LXXXV, figs. 2, a-/.) In all these .the genital pores are on the ventral side, near the mouth. In some other cases (Lepty chaster} the mother carries the eggs on the back, between the spines. Another method occurs in the genus Pteraster, and probably in all the related genera of the family Pterasteridae. In these there is a large dorsal " nidamental pouch," marsupium, or better, gono- codium, for the retention of the eggs and the development of the young, which remain in it until fully formed and sometimes up to 10 to 15 mm. or more in diameter. This gonocodium is formed by a tent-like membrane, which is supported and kept stretched by the tips of slender, elongated, radiating spinules, arising from the dorsal ossicles, and long enough * Starfishes that have such minute eggs and young usually have minute genital pores, situated in pairs, dorsally, in the interradial areas. Those that have large eggs that they carry around or over the mouth have the genital pores larger, on the ventral side near the mouth, so far as observed. SHALLOW-WATER STARFISHES 9 to leave a considerable cavity beneath it, in which the genital orifices and respiratory papulae are situated. This membrane is perforated by small pores for the ingress of water, and normally has a large, valvular, central oscule through which the water escapes. The mem- brane has muscular fibers for contraction. The eggs are laid and fertilized in the gonocodium, and when the young are matured they often, or perhaps always, escape by rup- tures of the membrane, which later heal up — a sort of natural csesa- rian operation. In this group the eggs are few in number, as usual in all cases where the young are well cared for. Several species of this family occur on the west coast. (See pi. xxxu.) SENSES; INSTINCTS; MEMORY; ACQUIRED HABITS. Comparatively little is known, from actual observation, as to the nature or acuteness of the senses in starfishes. At the tip of each ray, in the apical plate, there is a small and very simple eye, or pigmented eye-spot, often red in color during life. It is at the end of the radial nerve. Probably this kind of an eye is of use only for distinguishing different degrees of light, or possibly the shadow of a nearby moving object. Close to the eye there is a single odd ambulacral tube, generally considered a tentacle, and supposed to have, at least, the sense of touch. However, all parts of the integument, and especially the papulae and sucker-feet, are also very sensitive to touch. There- fore it may be that these tentacles have some other sense func- tion more developed than elsewhere, possibly that analogous to smell or taste, for appreciating the purity or salinity of the water. They must have a sense for heat and cold, as this determines their migrations at certain seasons. That they are somewhat sagacious and discriminating in their successful selection of the young oysters for food in spite of the labor and time required to open them, is good evidence that they have some sense analogous to taste, and have sense enough to follow it. That they can be attracted to an open oyster or other favorite food, from some distance, indicates that they have some sense simi- lar to smell, like that of many mollusca and Crustacea that are easily attracted by a dead fish or other odorous food. As they have no proper head or central brain, but only a circle of equal nervous centers, it is clear that they can have no consciousness of right or left, forward or backward. Each ray is practically equal to every IO VERRILL other, yet all the vast number of suckers, sometimes 20,000 or more, and all the rays must act in unison or there could be no progression, nor any procuring of food. Just how starfishes decide to travel in one direction rather than in another is a problem difficult to solve.1 Dr. Jennings, who has experimented extensively with reference to the behavior of starfishes, says very truly that whatever a starfish can do at all it can do in many different ways and seldom does it twice in just the same way. This is particularly applicable to its modes of righting itself when turned over, getting out from under weights, elastic bands, etc. He found that when a starfish had been forced to use the same arms to right itself very many times and for several weeks, it could thus be trained to continue to use the same arms, and thus had acquired a new habit, but if left to itself it lost the habit in about a week. Perhaps the preference of the starfish to use the same rays was mainly because, by repeated use, the muscles of those rays had become somewhat larger and stronger by the systematic " training." I know of no other successful attempts to educate a starfish. In nature they seem to show some memory and some persistency. I once placed a large active holothurian (Thy one briareus), that I wished to figure, in one end of a large aquarium, about four feet long, while there was a starfish (Asterias forbesi) at the other end. Next morning the starfish was mounted on the Thyone and had slightly eroded its skin by means of its oral spines. I disengaged the Asterias, put it at the opposite end of the tank with stones and other obstacles between, and supplied other food, such as cracked mus- sels, etc., and other Thy ones, apparently just as good. In a couple of hours it was back again, working away at the same spot on the holothurian. The same course was taken a second time, with the same result. The starfish was evidently bound to eat that particular Thyone or go without any dinner. I had to remove it from the tank to save the Thyone. Apparently the Asterias either remem- bered where his chosen dinner rested, or else it had a keen sense of smell to distinguish it from others of the same species, and from other natural food. In respect to maternal instincts, the most interesting case known to me was told to me by Prof. Louis Agassiz, while I was a student * Dr. L. J. Cole (Journ. Exper. Zool., xiv, No. i, 1913) has recently made some interesting experiments on the behavior of Asterias forbesi under special conditions, as to the relative use of the various arms as anterior or directive, etc. SHALLOW-WATER STARFISHES II and assistant with him about 1862. I am not aware that it has been published. It is recorded in a diary kept by me at that time. He said that he had a large aquarium, at Nahant, Mass., in his summer residence, I think, and placed in it a large female Solaster endeca,1 which was carrying a cluster of eggs over the oral area. As an experiment, he removed the cluster and put it at the extreme opposite end of the aquarium. In a few hours the starfish found the cluster and replaced it as at first. Mr. E. Desor (Proc. Boston Soc. Nat. Hist., vol. in, p. n, 1848) records a similar observation in the case of Henricia sanguinolenta. I have, personally, never had a chance to repeat this experiment, for though I have dredged thousands of Solasters, it has always been at the season of the year when they were not carrying eggs. Such experiments should be repeated at the various permanent seaside laboratories now in existence on both coasts. None existed during the long period of most of my field work, except, after 1881, that of the U. S. Fish Commission. PECULIARITIES OF THE STARFISH FAUNA OF THE NORTH- WEST COAST OF AMERICA. The general discussion of the geographical distribution of the genera and species and their relations to those of other regions must be left to a final chapter. There are, however, some very peculiar features of more general interest, to which it seems best to call attention in this place. The great size of many of the species, some of which are among the largest, if not the very largest species known, has already been mentioned.2 The Pycnopodia, though perhaps not the heaviest, is probably the largest starfish known. It becomes at least thirty-six inches broad ; and according to some collectors may be even four feet across. 1 As no one else has recorded the carrying of eggs by Solaster, and as its genital pores are described as minute and dorsal, I suspect that there was an error on my part in recording the name, or on his part in giving it. Perhaps Henricia was meant. 2 At least nine of the species of Asterias and allies become there over two feet in diameter. Among these large species are Orthasterias columbiana Ver., O. forcipulata Ver., Ev. troschelii (St.). var. rudis V., Ev. acanthostoma V., Pisaster ochraceus (Br.), P. giganteus (St.), P. lutkenii (St.), P. papulosus Ver., Pycnopodia helianthoides (Br.). Luidia foliolata becomes about as large. 12 VERRILL MULTIPLICITY OF RAYS ; VARIABILITY. It is a singular fact, for which no satisfactory reason has yet been given, that nearly all Echinoderms, whether living or fossil, are nor- mally five-rayed. This rule prevailed quite as constantly in the remote paleozoic ages as in the present period. The number of rays was apparently well fixed in the unknown primordial ancestors of the earliest fossil Echinoderms of most classes, for all classes of Echinoderms, except, perhaps, the Holo- thurians, appeared in the early geologic ages, with most of their more important features much as we find them now. Among Paleozoic Echinoderms, as among modern ones, certain starfishes departed from the five-rayed type by acquiring additional rays. As in some modern genera, this was apparently done in post- larval life by the interpolation, or budding in, of new rays between the older ones, in some of those species having numerous rays. At present, this is known to occur only in Pycnopodia of the Northwest coast ; in Heliaster, represented by several species on the tropical and subtropical Pacific coasts, from lower California to Chile ; and in Labidiaster, from Patagonia. It is probable, also, that it occurs in Rathbunaster Fisher, a deep-water Californian species, allied to Pycnopodia. Of Heliaster, seven species are known. When adult, the number of rays exceeds twenty and in some species is as high as forty to forty-four. In all its species new rays are gradually interpolated, rather irregularly, between the older ones, all around the circumference, but not without some order.1 The same is true of Labidiaster, which has numerous long rays when adult. In Pycnopodia, the number of rays when adult is twenty to twenty- four. It normally starts, when very small, with six equal rays. A new pair of rays then appears, one on each side of one of the primary rays ; then another pair appears just back of these, and so on in suc- cessive pairs.2 This would always produce an even number of rays. But variations from this bilateral regularity often appear, producing odd numbers. (See pis. LXXIII and LXXIV.) ' See H. L. Clark, Starfishes of the Genus Heliaster, Bull. Mus. Comp. Zool., vol. LI, pp. 25-76, pis. i-vni, 1907. In this paper all the species are fully de- scribed and their modes of adding new rays are explained. " See plates. See Professor W. E. Ritter and G. R. Crocker ( Proc. Wash- ington Acad. Sci., vol. n, pp. 247-274, 1900), who give a good account of the process. SHALLOW- WATER STARFISHES 13 All the known species of these three remarkable genera are from the Pacific coast of America and the outlying islands.1 Their resemblance to certain fossil starfishes of the Devonian is very striking, and may indicate a continuous descent from those ancient forms. If so, it would be good proof of the antiquity of the West Coast forms, as mentioned above. Aside from these remarkable and localized genera, there are other genera of the fauna that have numerous rays. The genus Solaster is represented there by five or six shallow-water species, and Crossaster by one. All these are multiple-rayed, the rays varying from eight to fifteen. In these genera, there is no evidence of the interpolation of new rays in post-larval life. They are also genera that are widely distributed in all seas. (See pis. vm and x.) They appear to be more numerous on the Northwest Coast than elsewhere, both as to individuals and species. Several additional species occur in deeper water off the coast. The group may have originated in the North Pacific, in former geologic ages. The genus Pteraster in all other seas has five rays, except P. obscurus, of the North Atlantic and Arctic, which ordinarily has six or seven. Variety octaster V., of Bering Sea, normally has eight rays, but sometimes seven. Mr. Fisher thinks that it is only a variety of P. obscurus, but even if so, it is singular that the latter should be six-rayed in the Atlantic and anywhere from six-rayed to eight- rayed in the North Pacific. So, likewise, other genera that are generally five-rayed in all other parts of the world are apt to have six-rayed species on the North Pacific coast. This is true of the genus Asterias and its subdivisions, in which we find many six-rayed species on that coast, as well as numerous five-rayed ones. Some of those that are ordinarily five- rayed are also often found with six or seven rays. This occurs in other regions, but not so frequently as there. Among the species of Asteriidae having normally six or more rays, the following may be mentioned : Pisaster grandis (St.), a very large species ; Orthasterias merriami V. ; A. aceruata St. ; A. kath- erincz Gray ; A. multiclava V. ; Stenasterias macropora V. ; Leptas- terias hexactis (St.) ; L. Vancouver! (Per.) ; L. cequalis (St.) ; L. coei V. ; L. epichlora (Br.), subsp. alaskensis V., and several varie- ties of the same species; and L. ntacouni V. Some of these have occurred with seven rays, and some with five, as abnormal variations. 1 Two species of Heliaster occur at the Galapagos Islands, and one at Juan Fernandez. These were evidently derived from Pacific coast ancestors. 14 VERRILL The following species, which are normally five-rayed, have been found with six rays : Henricia leviuscula (ST.), from Alaska. Henricia sanguinolenta (Muix.), Bering Sea. Pisaster ochraceus (Bn.), Monterey, Calif., and British Columbia. Patiria miniata (BR.). Six-rayed common; four- rayed and seven-rayed rare; California. Dermasterias imbricata (GR.), Sitka. When we consider the great geological antiquity and remarkable persistence of the five-rayed condition in echinoderms generally, it is very remarkable that so many genera and species of existing starfishes should have acquired the peculiarity of having higher numbers. It seems natural to conclude that there must be some special advantage gained by this increase in rays, which does not obtain in the other classes to the same extent. At the present time all known species of sea-urchins and holo- thurians are normally five-rayed, and the same holds good through all the geologic ages for the former class at least. Abnormal sea- urchins with four and with six rays have been observed in a number of species. Dr. Robert T. Jackson, in his recent extensive work,1 has discussed this and various other variations of sea-urchins very fully. He states that he has personally studied seventy-one cases of more or less complete variations from the perfect five-rayed condi- tion. To find these required the special examination of 50,000 speci- mens, including many species. In some species about one individual in a thousand shows varia- tions of this kind.2 The serpent-stars (Ophiuroidea) show more variation in the number of rays than the sea-urchins do. Several genera contain some species that are normally six-rayed. The common Ophiocoma puntila of Bermuda and the West Indies has about equal numbers of five-rayed and six-rayed individuals living together. Many of the species of Ophiactis have six to eight rays while young, and divide autotomously, but when mature they are usually either regularly five- rayed or six- rayed. Yet some of the species of the same genus are always five-rayed and do not divide ; so it does not seem to be a matter of great importance in this genus. 1 Phylogeny of the Echini, Mem. Boston Soc. Nat. Hist., vol. x, 1912. Variations from the pentamerous system are described on pp. 35-50. " See also A. E. Verrill, Amer. Naturalist, vol. XLIII, p. 545, for a discussion of the subject with figures. SHALLOW- WATER STARFISHES 15 Variations from the five-rayed condition occur not very rarely among living Crinoids, and were not uncommon among the ancient fossil pentremites and cystideans;1 but on the whole the crinoids have been pretty constantly five-rayed through all the geologic ages. We must conclude that all these variations originated at first as " sports," which have persisted by heredity and natural selection, because they were advantageous. It is easy to conjecture that, in the case of two starfishes, otherwise similar in size and structure, living together on a rocky shore and exposed to violent surf, the one with six rays would be able to cling more securely to the rocks than the one with five rays. Therefore, because of the increased number of ambulacral sucker-feet, it might well be the form preserved by natural selection, unless for some other important but unknown reason, the five-rayed condition has certain other more important advantages. It is certainly true that most of the shallow-water species with multiple rays live among rocks in situations exposed to the surf. This is true of the seven species of Heliaster, with very numerous rays; and of Pycnopodia with twenty to twenty-four rays, and of the various shallow-water and littoral species of Solaster and Cross- aster, which usually have nine to fifteen rays (rarely eight or less). It is also true of the numerous six-rayed species of Asterias, Pisaster and allied genera.2 However, the ability to cling tenaciously to rocks may be perfected in other ways, involving equally an increased number of sucker-feet. This is often attained by lengthening the rays, as in many species of Asterias; by crowding the suckers into more than four rows, as in some large species of Pisaster; and by increasing the size and strength of the suckers. Although it seems probable that the added protection gained against the violence of the waves by the evolution of more numerous rays and suckers is a real cause for the retention of this feature, it may not be the principal one. The attainment of four rows of sucker- 1 Some of the paleozoic Cystidea were tri-radial. Hence it has been thought by some writers that this was the more primitive condition and that the pentamerous condition was acquired by the addition of another pair of rays. See Bather, op. cit, 1901. 2 The family Brisingidee, however, is mostly confined to deep water. All the species are multirayed, with long rays. Some of them climb over gorgonians, as do many ophiuroids. The genus Labidiaster, of the Patagonian region, is found in shallow water among rocks and sea-weeds. It has numerous long rays. 1 6 VERRILL feet in Asteriidae, as compared with the more primitive two-rowed condition, was probably due to the same causes. I am inclined to believe that the increase of rays has been due more to the advantage gained in holding their food securely, and in opening bivalves, than for holding to the rocks, though both go together. The starfishes are the most predaceous of the echinoderms. They feed largely on bivalve shells that cling firmly to the rocks, like the oysters, and on others that must be pulled open by means of their numerous muscular suckers, part of which must be used for keeping in a suitable position and part for the opening of the oyster, at the same time, so that the more arms and suckers they have, the more rapidly and effectively they can secure their food, in case they are feeding on large bivalves. We must admit that, so far as known, the five-rayed and six-rayed individuals of a species appear to be equally well nourished and grow to equal size. Also that the normally six-rayed species of Asterias are commonly no larger, nor more robust, than the allied five-rayed species, in the same environment. Even the four-rayed individuals appear to be well nourished and of ordinary size. It is quite possible, however, that they grow more slowly. It is known that the rate of growth of starfishes is very rapid when food is abundant, but very slow when food is scarce. No observations have been made on the comparative rate of growth of six-rayed specimens. It must be remembered that the variation in the number of rays is necessarily attended by extensive changes in the number, size, and form of the skeletal plates of the body; also in the number of ambulacral feet and water tubes, nerve ganglions, nerve cords, stomach lobes, hepatic glands, and all other internal organs.1 A six- rayed specimen has twelve reproductive glands, instead of the ten in its five-rayed competitor. If the number of ovules be propor- tionately large, it would produce twenty per cent more young. So, likewise, it would have an additional stomach lobe and two more hepatic glands. This would perhaps be of advantage in the 1 This does not hold good in the cases of the multirayed species of Heliaster, according to Clark. (The Genus Heliaster, 1907.) This genus has but five stomach lobes and five pairs of gastric retractor muscles, whatever may be the number of rays. Yet the added rays contribute to digestion in that group, for each ray has a pair of hepatic glands and a pair of gonads. The term "hepatic gland" must not be understood as implying that these glands correspond precisely with the liver of vertebrates, for it is known that the digestive fluids of echinoderms are different chemically from those of the latter. Some claim that these glands of starfishes are more like the pancreas in function. SHALLOW-WATER STARFISHES I/ digestion of food and cause more rapid growth and earlier maturity, if not greater reproductive powers. It would be a very interesting experiment to try breeding six-rayed or seven-rayed varieties of some common five-rayed species. I have found five-rayed young (about 5 per cent) among those carried by a six-rayed mother, in the case of Leptasterias epichlora. Fisher states that he has found seven-rayed young in the gonocodium of an eight-rayed Pierasier obscurus, and one with nine rays. That six-rayed and seven-rayed varieties might be easily obtained by selection and isolation is very probable. I have observed that in the case of our common New England starfish (Asterias forbesi) six- rayed and seven-rayed individuals are much more common in some localities than elsewhere, indicating, perhaps, a tendency to the inheritance of these features. SPECIES, SUBSPECIES, AND VARIETIES. Within the limits of recognized species, certain groups of indi- viduals, apparently having a definite distribution and more or less permanency of characters, are regarded as bathymetrical or geo- graphic races. Such races are here called subspecies. In certain genera, for example Asterias, Henricia, Solaster, speci- mens occur which cannot be referred definitely to any species or sub- species. Some of these appear to be local variations, due to unfavor- able environments ; " sports," " freaks," or " hybrids ; " others may be abnormal individuals. They are here called varieties. If in future the characters of any of these varieties prove fairly constant, it may become necessary to elevate them to subspecific or even specific rank. To determine the status of these forms, much larger series of speci- mens than are at present available will be necessary. It is not unreasonable to think that some of these " varieties " and " subspecies " are incipient species, now in the process of evolution, and that eventually some may become fixed species, while others will disappear by the elimination of the most unfit. The Northwest Coast appears to be one vast nursery for the origi- nation and evolution of new varieties, subspecies and species of star- fishes and various other groups of marine animals. A peculiar difficulty in the determination of genera and species of starfishes consists in the fact that many, especially of the Asteriidse, do not attain their adult characters until of considerable size, so that young, or even half-grown specimens, may appear to belong to a different species, or even a different genus, for the more complex l8 VERRILL species pass through stages corresponding to the adult condition of simpler or more primitive genera, and there is often no way to tell whether a small specimen is mature or not, without comparison with a series. HYBRIDISM OF ASSOCIATED SPECIES. Owing to the fact that numerous related species of several genera are living together on the Northwest Coast, it is natural to conclude that some of them may frequently hybridize, especially if their breeding seasons are coincident. I have seen many hybrids between Astcrias vulgaris and A. forbesi of the New England coast. There is considerable evidence that this often occurs on the Pacific Coast, leading to the existence of many very puzzling specimens in some groups. I have personally seen a number of apparent hybrids between diverse species of Asterias, Pisaster, Henricia, and other genera abundant on that coast. Perhaps some of the equally puzzling speci- mens of Solaster may also be hybrids. To determine this matter satisfactorily requires large collections, and especially observations made on the living specimens in their natural environment. The littoral species of Leptasterias that carry their young, abundant on the shores, are eminently adapted for such studies, and are easily raised. This can best be done at one of the seaside biological stations, and for that reason I do not propose to discuss the matter at length in this place. Dr. Fisher, also, has found numerous varieties and connecting forms, some of which he thinks may be hybrids, among the large numbers of specimens of Henricia that he has studied. It is quite probable that some of the now well established species of starfishes of that coast originated at first as hybrids and that others are now in the process of differentiation into varieties and species. FAUNAL RELATIONS AND DISTRIBUTION OF THE SPECIES. Details of the distribution of the species must be left to the end of this report, but there are some interesting facts that may well be mentioned here. The shallow-water starfish fauna of the Northwest Coast, taken collectively, may be considered as consisting of three parts. ist. Those of more southern or tropical origin that have migrated northward to California or beyond. SHALLOW-WATER STARFISHES 19 2d. Those that are of Arctic or circumpolar distribution and extend southward into Bering Sea, and in many cases to the Aleutian Islands, or much farther south, especially in the deeper waters. Their origin may have been in the North Pacific originally, in many cases. 3d. Those that appear to have originated in the region extending from the Aleutian Islands to California. The last group is by far the larger and more important. Very likely it is possible to divide that very extensive coast line into two or more faunal districts, but for the starfishes this does not seem to be necessary for the present, for many species range throughout that whole extent of coast. The limits of distribution on that coast seem to be determined entirely by the temperature of the water, especially in the breeding season, due probably to the greater sensitiveness of the free-swim- ming larval forms. The adults can regulate their temperatures by migrating into deeper or shallower water as occasions require. Among the more southern forms are Orthasterias forreri, Pisaster capitatus, P. paucispinus, Marthasterias sertulifera, species of Astro- pecten and Luidia, Linckia Columbia, etc. Among the arctic and in part circumpolar species are Asterias acervata, A. multiclava, L. arctica, C. cribraria, Allasterias rathbuni, Henricia sanguinolenta, H. tumida, H. arctica, Solaster endeca, Crossaster papposus, Pteraster obscurus, and var. octaster, P. mili- taris, Diplopteraster multipes, Tosiaster arcticus, C. granularis, Leptychaster arcticus, Ctenodiscus crispatus, etc. The species apparently indigenous on that coast are too numerous to enumerate here. Some of them are nearly allied to those of the North Atlantic and other regions, but many are very peculiar and have no near allies in other regions, so far as known. A number of the genera and higher groups are peculiar to that coast, others have there a remarkable development in number and variety of species, showing that their evolution must have gone along continuously for vast periods of time. In many cases primi- tive and highly specialized species of a family are found associated. Among these peculiar types are Pycnopodia, Rathbunaster, several species of Pisaster, many species of Asteriinge, Dermasterias imbri- cata, Leptychaster (Glyphaster} anomalus, Pteraster tesselatus, Bunodaster ritteri, etc. Fuller lists will be given at the end of this work, with more details of their distribution. Class ASTERIOIDEA. MORPHOLOGICAL FEATURES. The Asterioidea1 have a polygonal or star-shaped body, in which the rays are direct prolongations of the body itself, and contain exten- sions of the body cavity and more or less of the viscera, especially one or more pairs of gonads and a pair of digestive glands ; generally, also, a lobe of the saccular stomach. The skeleton is made up of large numbers of ossicles or plates, mostly articulated so as to be more or less movable, giving flexibility both to the rays and to the disk, though in some species (certain Goniasteridae) the flexibility is slight, except at the tips of the rays. The actinal side of the disk and rays has deep radial ambulacral grooves, extending to the tip of the rays. The roof of the groove is supported by the two rows of ambulacral ossicles, arranged like rafters, or in close, inverted V-shaped pairs of compressed plates or bars, between which there are rows of pores for the passage of the ambulacral feet.2 In the middle line of each ray and external to the ambulacral plates are situated the radial nerve and blood-vessel. The radial water tube supplies water to the locomotive tubes through the medium of muscular ampullae, usually double, situated internally above the ambulacral plates. In Brisingidae ampullae are lacking; in Echinasteridse they are single. There is no median row of calcareous plates covering the ambu- lacral areas and radial nerve and blood-vessel, such as exists in Ophiuroidea. The tip of each ray ends in a special terminal ocular, or apical, plate, supporting a pigmented ocellus, to which the radial nerve extends. According to the studies of Fewkes, 1888, these are the first plates to appear in the young. The grooves are bordered on each side by a row of plates called adambulacral, which always bear spines. 1 This spelling of the name is preferred because it is derived from Asterias, not from Aster. 3 In many paleozoic fossil starfishes the ambulacral plates are not opposite, in pairs, but alternate. This rarely occurs in existing species, though I have noticed it in Pycnopodia as an abnormal variation in some of the rays, and also that it may occur from lateral bending. (20) SHALLOW-WATER STARFISHES 21 The mouth is central, dilatable, and surrounded by soft membrane. The so-called jaws are merely the adoral ambulacral and adambu- lacral plates, more or less modified and coalesced ; the " teeth " are only slightly modified adambulacral spines, in this work called peroral spines. (See text-fig. 3.) The stomach is very saccular and usually evertible. It usually has a lobe and a pair of digestive glands extending into the cavity of each ray; but in the case of some multiple-rayed species (Heliaster), it has lobes corresponding only to the primary five rays. In this case the five stomach-lobes do not enter the rays, but the pairs of digestive glands do. In some slender-rayed genera, also, the stomach is confined to the disk. The intestine is usually nearly or quite abortive and not functionally active. The so-called " anus " is a dorsal pore, chiefly for the discharge of secretions from the dorsal glands, or " csecal appendages," probably nephridial in function, and called nephridial glands in this work. Commonly there is a single pair of branched gonads in the proxi- mal part of each ray, with simple ducts discharging through a pair of interradial pores, which may be either ventral or dorsal. In cer- tain Brisingidse, Luidiidae, and in a few other families there are several pairs of gonads and genital pores arranged serially along the sides of each ray. The madreporite or madreporic plate is dorsal,1 excentric, and commonly single, yet in some multiradiate species there may be sev- eral. It is an organ primarily for the purpose of eliminating the excess of absorbed water from the ambulacral tubes and body-cavity. The sides of the rays and disk are generally supported by two rows of marginal plates, usually larger or thicker than the other plates, and commonly bearing special spines. They are called supra- marginals or superomarginals and inframarginals or inferomarginals. (See text-fig, i.) The upper row is sometimes much reduced, or obsolete, and rarely both rows are lacking or rudimentary. These rows of plates belong to the primary system of plates, and extend to the apical plate of the rays, like the ambulacral and adambulacral rows. Plates are constantly added to these rows by the budding in of new plates between the apical plate and the one next to it, the apical plate being pushed farther outward and the ray lengthened at the 1 In some paleozoic fossil starfishes it is said to be ventral, but it is not so in any living species. The statement by Gregory (op. cit., p. 238, 1900) that it is ventral in Asterina is erroneous. 22 VERRILL same time. The first median dorsal plate of the rays appears very early in the young, and the first marginals and adambulacrals soon after. In many genera and families, but not in all, additional rows of plates may be interpolated between the marginal rows proximally, and are called intermarginals ; or between the inferomarginals and the adambulacrals, when they are called interactinals, intermediate actinals, or simply actinal plates.1 These do not appear very early in the young, and are often without spines. These interactinal and intermarginal plates do not belong to the primary system of plates, nor do they commonly reach the apical plate. Their new plates develop mostly at the tips of the rows, as the starfish grows larger, and new rows may be interpolated till they sometimes become very numerous, in order to increase the size of the rays. The dorsal skeleton of the disk, in a five-rayed species, primarily has five basal radials (or first dorsals), five genitals, and five pairs of interradial plates, besides the plates that later become the apical or ocular plates, and the centro-dorsal plate. By the interpolation of new ossicles and plates in various ways, the structure often becomes very complex. On the rays we can usually distinguish a median or carinal row, extending from the basal radial to the apical plate. Other regular rows may develop each side of this (the dorso-laterals), or the whole surface may become covered with a tesselated arrangement, or a reticulated system of plates and transverse ossicles.2 The dorsal plates, like the marginals and interactinals, commonly bear spines or small spinules, but they may be covered with granules, or with a smooth soft integument, or even appear quite naked, being then covered only with a thin membrane. These plates and their armatures of spinules take several special names, according to their forms and structure, and are often char- acteristic of special genera and families and higher groups. When they become columnar or of hour-glass shape, and have the summit covered with a radiating cluster of small slender spinules, 1 For more details of the rows of plates and their sequence, see below, under the family Asteriidas, and text-figure, I, i-iv. For the sequence in the early stages of the development of the primary plates, see J. W. Fewkes: On the Development of the Calcareous Plates of Asterias, 1888. 2 For more details of the skeletal plates, see the discussions under the sev- eral orders and families below, especially under Forcipulata and Phaner- ozona. SHALLOW-WATER STARFISHES 23 they are called paxillae. These are most typically formed in Astro- pecten, Luidia, and allied genera, and are characteristic of the sub- order Paxillosa.1 Pedicellariae of one or more kinds are generally present, but are entirely lacking or very rare in certain families, such as the Solasteridse, Echinasteridae, Pterasteridae," and in some genera of other families. Their uses are imperfectly known. One use is to keep the body free from dirt and parasites." Another use, perhaps the most important, seems to be, according to my own observations, to retain the discharged unfertilized ova until fertilization takes place, thus preventing their dispersal and loss. This, however, needs further investigations on living starfishes. These curious organs are not found in other classes of Echino- derms, except in the Echinoidea. In the latter they are commonly mounted on long stalks and generally have three valves, sometimes two to four or even more. In the Asterioidea they are usually sessile or (Order Forcipulosa) have very short pedicels, and they are most frequently bivalved. They are of several different types, which are commonly char- acteristic of the orders and families, and often of the genera. Sometimes there may be bivalved, trivalved, four-valved, and five-valved ones on a single specimen. (See below, under Der- masterias imbricata, and pi. vi, figs. 4, 5 P, P'.}. In some cases they are not really valvular, but consist of several movable spinules, arranged in convergent groups or opposed rows (pi. xxxiv, P. P.). The sessile valvular forms are generally situated over a pore, pene- trating a plate, for the passage of nerves, etc. (Pis. XLVII, XLVIII.) The different kinds take special names. For further details of the pedicellariae and their special names, see below, under the sev- eral orders, and plates LXXV-LXXXV. In nearly all species there are numerous small, soft, tubular out- growths from the body-wall, called papulae, serving for respiratory purposes. They may be in large groups, especially between the 1 For various other special forms and their names, see below, under Phaner- ozona and Valvulosa. 3 In the last two of these three families, and perhaps in the first, the mother carries and protects the eggs and young until they are able to provide for themselves. * The large bivalve pedicellariae of Hippasteria are very muscular. I have lifted a large living starfish of this kind, weighing about a pound, entirely out of water by a toothpick, seized by the jaws of a dorsal pedicellaria. 24 VERRILL dorsal and lateral ossicles, or stand singly, or they may occur only in special areas. Rarely they are branched. Most shallow-water starfishes that have been studied in this respect have a free-swimming, bilateral larval form, known as a brachiolaria ; yet a considerable number carry their eggs and young till they develop the starfish form, as explained above. These have an abbre- viated metamorphosis. The two methods may occur in different genera of the same family (e. g., Asteriidce}. The embryology of the deep-sea species is unknown in most cases, and may be diverse, or even more abbreviated. CLASSIFICATION OF ASTERIOIDEA. In this report I have adopted the division of the class into three large orders, viz. : I. FORCIPULOSA or FORCIPULATA. II. SPINULOSA. III. PHANEROZONA. The last has two suborders, considered orders by some recent writers, viz. : I. VALVULOSA or VALVATA. II. PAXILLOSA. Order FORCIPULOSA Verrill, or FORCIPULATA Perrier. Stelleridce forcipulata PERKIER, Mem. Etoiles de Mer, pp. 166, 188, 1876. Forcipulata PERRIER, Exped. Sci. Trav. et Talisman, p. 27, 1894. Asteries Ambulacraires VIGUIER, Squellette des Stellerides, Arch. Zool. Exper., vii, p. 93, 1878. Cryptozonia (pars') SLADEN, Voy. Chall., xxx, pp. xxxiv, 397, 1889. Adetopneusia (pars') and Leptostroteria (pars) SLADEN, op. cit., p. xxxrv. The form is always stellate, often with long rays, commonly five, but often multiple. Ambulacral plates, except the orals, are usually short and closely crowded (leptostroterial) , but not in Brisingidae. The proximal pair is elongated and, except in Pedicellaster and a few others, forms the inner end of the jaw. Ambulacral feet are generally arranged in four rows, — but in two rows in Pedicellas- teridae, Brisingidae, and Zoroasteridae. In some large species of Pisaster they may form six or more rows by crowding, especially subproximally. They are always terminated by suckers. SHALLOW-WATER STARFISHES 25 Adambulacral plates are generally short and crowded, equal in number to the ambulacrals. They may each bear a single spine (monacanthid} , or two spines (diplacanthid} , or alternately one and two (subdiplacanthid} , or rarely three spines (triplacanthid} . The superior and inferior marginal plates are rather small (cryptozonial}, but are generally distinct and bear spines, often longer than those on the dorsals. In some Brisingida one row, or rarely both rows, may be lacking. They extend to the apical plate and increase in number during the whole period of growth by the addition of new plates next to the apical. The dorsal skeleton is variously constituted, but usually consists of plates and ossicles, often very unequal in size and form, so united as to move freely, leaving many papular areas between them. In some of the Brisingidse and in Pycnopodia the ossicles are mostly nearly abortive, roundish, and isolated in the integument. The plates generally bear spines, either singly or in clusters; sometimes they are reduced nearly to the form of granules ; never paxilliform. The papulae may be few, or many in large clusters. They usually occur both on the dorsal and on the ventral surfaces, — adetopneusic arrangement. The Forcipulosa, as the name indicates, are especially character- ized by the presence of very peculiar, pincer-like, two-bladed pedicel- lariae, usually of two forms, movably attached .to the dermis of the spines, or to the surface between, by means of longer or shorter flexible dermal pedicels. VARIOUS KINDS OF PEDICELLARIyE. Except possibly in a few rare cases, and in quite immature speci- mens, at least one form of these is always present. Of the two principal forms, those known as forcipate or " minor pedicellaricz" are the most abundant and occur most constantly. Such pedicellariae do not occur on starfishes of any other order. Those nearest analogous occur on certain Echinoidea. In the larger kind, called "major pedicellaria" by Stimpson;1 1 1 prefer to use ordinarily the names first given by Stimpson, in 1861, for the two principal forms of pedicellariae, viz., major pedicellarice and minor pedicellarice. These terms are sufficiently descriptive, and not so liable to be confused as those proposed later by Herapath, though the latter have some advantages. It is desirable, however, to have some additional special de- scriptive terms for varieties of each sort. Either form may be attached to the spines or to the general integument. The minor pedicellarise that sur- round spines in wreaths, may be called circumspinal; those in smaller irregu- lar clusters on the spines, epispinal; those attached to the general surface of 26 VERRILL " pedicellaires droites" by Perrier, 1869; and " forficiform pedicel- lories " by Herapath, 1866, and by Sladen, 1889, the jaws are nearly straight, articulated at the base by a simple joint to a thick basal piece, so that they open and close like forceps. These may be either dermal or attached to the spines, either singly or in clusters; they often have short pedicels. In the Brisingidce and in Pedicellaster they are usually lacking. (See pi. XLIX, figs. 3-3^; pi. xxx, etc.) The second kind, called "minor pedicellarice" by Stimpson, "pedi- cellaires croises" by Perrier, "crossed pedicellaria" by several writers, and " forcipiform pedicellarice" by Herapath, are usually much smaller and the blades are curved and crossed something like those of scissors, and so articulated that they open and close like scissors, tweezers, or pincers. These, like the others, may be attached directly to the integument, either singly or in clusters, or to the spines, to the pedicels of the larger forms, to saccular dermal growths around the spines, to the inner edge of the ambulacral grooves, or rarely even to the sucker-feet. (See plates LXXVI- LXXXIV.) They are most commonly attached by slender and some- times long pedicels, and aggregated into wreaths or clusters on the spines. They are often so abundant, in the wreaths around the dorsal spines, that they nearly or quite conceal the spines and integument in living specimens. In some cases they are attached in large num- bers to dermal sheaths or sacks loosely surrounding the spines. They often have a formidable array of minute sharp denticles, and are frequently very characteristic of species. (See text-fig. 2.) FAMILIES AND SUBFAMILIES OF FORCIPULOSA. This order now includes the following families and subfamilies : Family ASTERIIDJE. Subfamilies ASTERIINJE; STICHASTERINJE; PYCNO- PODIIN^E; HELIASTERIN^:. Family ZOROASTERIDM. Family PEDICELLASTERIDM. Family BRISINGIDCE. Subfamilies BRISINGINM; LABIDIASTERIN&. (Type, Labidiaster Liitk.) the body are dermal; those on a papular area are papular; those on the adambulacral spines or plates are adambulacral. Major pedicellariae may also frequently occur within the adambulacral grooves, attached to the inner sur- face of the adambulacral plates, where they may be called intra-adambulacral. SHALLOW-WATER STARFISHES 27 Family ASTERIIDJE Gray (emended). Asteriida: GRAY, Ann. Mag. Nat. Hist., vi, p. 178, 1840; Synopsis, p. I, 1866. Perrier, Revis. Stell., Arch. Zool. Exper. et Gen., iv, p. 302, 1875; Mem. Etoiles de Mer, pp. 167, 198, 1876. Viguier, Squelette des Stellerides, pp. 93, 99, pl- v, figs, i-io, 11-12, 1878. Asteriida! (emended) + Stichasterida; SLADEN, Voyage Chall., xxx, pp. 430, 560, 1889. Perrier, Exp. Trav. et Talisman, pp. 43, 105, 107, 128, 1894; Contrib. 1'etude des Stellerides Atlan. Nord, pp. 25, 31, 1896. Rays five to twenty or more, but most frequently five or six. Madreporic plate generally single in five-rayed and six-rayed forms, but often two or more in those with a variable number of rays. Dorsal and actinal plates various in form and arrangement. Odonto- phore usually formed of a single piece. Oral ambulacral plates elongated; those along the grooves become very short and com- pressed. Papulae occur both on the actinal and dorsal sides. Ambu- lacral feet usually crowded in four rows, sometimes more. Pedicel- lariae of two kinds are present. Dorsal ossicles generally bear spines or spinules, various in size and kind, and sometimes granules. They are never true paxillae. The mouth is large; the stomach saccular and evertible; intestine rudimentary or abortive. A subcentral, dorsal pore (nephridial pore) is present, through which is discharged the secretion of the branched or lobular " rectal " or nephridial glands, which are usually unequal and often but two or three in number. This pore is usually called the " anal pore," but rarely, if ever, functions as an anus in this family. The intestine is nearly or quite abortive, in most cases. I. MORPHOLOGY OF THE OSSICLES. Aside from the ambulacral and adambulacral plates, the skeleton of the rays consists of five fundamental or primary rows of ossicles, viz., the median dorsal or carinals; the two superomarginals; and the two inferomarginals. The latter may or may not be confined to the ventral side. (See fig. I, i.) The ossicles of adjacent rows may be articulated directly by their lobes, or either large or small, simple connecting ossicles may inter- vene. This simple or primitive type of skeleton is found in Heterasterias volsellata (Sla.), and in a few other species, especially when young. In most cases more or less numerous ossicles or rows of ossicles are interpolated during growth between the five primary rows, either above or below, or between the marginals, thus giving rise to many 28 VERRILL md. md- ffffftnirn? FIG. i. Diagrams generalized to illustrate progressive ontogenetic and phylogenetic development of the plates and spines in the rays of starfishes (Asteriide?) . Lettering as follows; the primary rows are shaded: md, Median dorsal or carinal row. m, Marginal rows, tn', Intermarginal row. _ sm, Supermarginal row. im, Inferomarginal row. ad, Adambulacral row. din, Dorsal-inter- mediate, medio-lateral, or dorso-lateral rows, p or pa, Peractinal row of plates, sa, Sub- actinal rows. These and the peractinals are the interactinal plates, collectively, o, The ocular, apical, or terminal plate. i, The simplest or most primitive condition, common in the young; uncommon in the adults, in which only the primary rows of plates are developed. The longitudinal area between ad and the next row is called the perambulacral lane or channel; the space between the marginals (m) is called the intermarginal lane or channel; the area next above the upper marginal is called the supramarginal lane or channel. These " lanes " or " channels " are always crossed by transverse connective ossicles or by over- lapping lobes of the plates, dividing them up into larger or smaller papular areas, and they usually bear more or fewer of the larger pedicellariae in the Asteriids. ii, The condition, more advanced, in which the peractinal row (p) and one medio-lateral row (din) are developed. A common condition in the Asteriinae. This is called the mono- actinoplacid condition. iii, A more advanced condition in which an additional interactinal row (sa, subactinal) and a second dorso-lateral or medio-lateral row (din) have appeared. iv, A more complex state, in which there are two subactinal rows (sa), and an inter- marginal row (m'). This is a common condition, but more medio-lateral rows are often added, as well as more subactinals, in such genera as Pisaster, Evasterias, etc. This is called the polyactinoplacid condition. All these plates usually bear spines, one or more each. They may be easily visible or they may be entirely concealed by a thick tough skin. The peractinal or sub- actinal plates, when thin or small, may be destitute of spines and stand edgewise, so as to show only the edge at the surface, or they may be so closely joined to the inferomarginals as to require cleaning with potash to be seen. SHALLOW-WATER STARFISHES 29 different styles of skeleton in the more complex genera. (Fig. I, ii-iv.) The carinal or median dorsal plates (md) usually remain distinct, but are often small and not easily recognizable. The superomar- ginals (sm) are nearly always easily recognizable, though small, and they generally bear spines larger than the dorsals. They never form a stout margin, as in the Paxillosa. The inferomarginal plates (im, fig. i) are generally very evident and carry one or more rows of spines usually longer or larger than the dorsals. They are usually separated from the superodorsals by a more or less wide, naked lateral lane or channel on which are rows of papular groups, and frequently many of the larger pedicellarise. In some cases one or more short intermediate rows of lateral plates (intermarginals, w'), bearing spines, may be interpolated in these areas, proximally. Both marginal rows normally reach the ocular or apical plate. The peractinal row (fig. I, pa) rarely actually touches the apical plate, and generally falls considerably short of it. This affords a useful character for distinguishing the inferomar- ginal from the peractinal plates.1 This name is proposed for the first or primary row of actinal plates. To designate collectively the rows of plates that may exist between the inferomarginals and the adambulacrals, I propose to use the term " inter actinals" in place of the longer one, "intermediate actinals," used by Sladen and others. I would do away with the use of the words " actinal spines " or plates used by many writers as including the inferomarginals with the true actinals. Ventrals may be less objectionable if such a collective name be needed, and is so used by Perrier and others. The upper marginal row of ossicles is usually pretty clearly defined by special spines. This superomarginal row usually diverges from the lower one, or turns upward at the base of the rays, and may run a little upon the dorsal surface to join its counterpart on the next ray at the dorsal interradial angle or axil. The plates here called subactinals (text-fig, i, so) appear later than the peractinals. They are often lacking in the young, up to 30 to 50 mm. in diameter, even when they are present in the adult. They often extend only on part of the ray, proximally. The number of rows of these in large species increases with the age. 1 It seems desirable to have special and convenient terms to designate the several series of plates and spines in this and related families. . 30 VERRILL The row of actinal plates that unites with the adambulacrals, when there are several rows, may be called synactinal, to indicate its func- tion. It may consist either of the peractinals or of subactinals. When these rows are very much alike in size and form, mistakes have often been made in descriptions, the peractinal row having often been described as the inferomarginal row or vice versa, or else no distinction has been made by the earlier writers. The most positive criterion, in doubtful cases, is to trace these rows of plates to their origin. True marginal rows originate at the ocular plate; peractinals, when present, often terminate near the ocular ; but other rows generally fail to reach the end of the ray. The inferomarginal plates may be directly joined to the adambu- lacrals by a lobe or apophysis, but more commonly there is at least one row of actinal plates between, and sometimes, besides this first or peractinal row (pa), there may be one or more shorter rows of subactinal plates (sa) interpolated proximally. All these plates usually bear spines, but in some cases all the interactinal plates are rudimentary and spineless. The secondary dorsal skeleton varies much, but is most commonly chiefly made up of unequal and more or less imbricated ossicles or lobulated plates, with the ends or lobes of adjacent plates over- lapping, so as to leave rather wide papular spaces between them. Or the plates may be broader, somewhat rhombic, lobed, and more closely imbricated, as in Stichaster. In most cases, however, they are also united transversely by smaller connective ossicles. ii. PEDICELLARIAE OF THE ASTERIID^:. Text-figure 2. Pedicellariae of both kinds are probably always present in the normal adults, though on some individuals they may be few or lacking. The major or forficulate pedicellariae are generally ovate or lanceo- late, sometimes wedge-shaped, stone-hammer-shaped, lyrate, or spatulate. The tips may be acute, plain, flat, or denticulate. They are mostly dermal, but may also occur on the ventral and oral spines, or along the inner edges of the grooves, or attached to pedi- cels in clusters within the grooves, or even on the pedicels of the ambulacral feet. They may be of several sizes and forms on one specimen. Certain species of Stichaster, Coronaster, etc., have a peculiar very large form of dermal pedicellariae, often as thick as the spines, in SHALLOW- WATER STARFISHES 3! which the broad valves terminate in a series of curved denticles, so that when closed the valves resemble a pair of clasped hands, or better still, a pair of clasped feet of a cat, with the claws protruded. These may be called felipedal or unguiculate. In many other cases similar gigantic pedicellariae have spatulate valves, with a smooth or denticulate edge. In Pisaster they are sessile, stout, erect, stone- hammer-shaped or wedge-shaped, with serrate or unguiculate edges. (See pi. XLIX, figs. 3-36.) These peculiar giant forms are usually accompanied by others of the ordinary ovate form, much smaller in size, and they are usually characteristic of special generic groups. But as such groups are sometimes widely separated, both morphologically and geographi- cally, it is not unlikely that these are survivals of a type of pedicel- lariae that, at some remote period, was common to the whole family. The minor or forcipulate pedicellariae (text-fig. 2) generally form wreaths or large or small clusters on the spines, but they may also occur, either singly or in clusters, on the integument of the dorsal or lateral plates, on the papular areas, and in other situations. III. RELATIVE IMPORTANCE OF MORPHOLOGICAL CHARACTERS IN ASTERIID;E. Perrier, Sladen, Bell, and others who have divided this family, have differed considerably in their estimation of the relative impor- tance of the various morphological characters. A. MODES OF GROWTH AND INCREASE IN NUMBER OF SKELETAL PLATES. Text-figure i, i-iv. The number of ossicles and plates of nearly all kinds, except the primary radials and orals, and the oculars, increase continually dur- ing the entire growth of most starfishes. The size and age have no definite limit, so that unusually large specimens of any species may often occur with correspondingly increased numbers of plates and spines. Therefore the exact number of plates, say of marginals, is of no great importance, except when specimens of identical sizes or ages are compared. In the early stages of growth, the first dorsal plates that appear are the central, primary interradials, the radials, and the ocular plates. A little later the median dorsal radials and the marginals appear; and the commencement of a row between the upper marginals and median row often appears at the same time or a little later. The number of the plates in all the primary rows is 32 VERRILL increased by the formation of new plates between the apical terminal or ocular plate and the plate next to it, the oculars being forced farther away as the numbers and growth of the plates continue to increase. But besides the primary rows of plates, other rows and scattered ossicles may appear, to fill up the dermal spaces produced by the increasing diameter of the rays and disk. In some forms of starfishes this is accomplished largely by the rapid growth of the primary plates, which may thus become very large, but this produces a rather inflexible skeleton. In the Asteriidse flexibility seems to be an essential feature, especially as most of the species cling to irregular stones and similar objects, and rarely live habitually on smooth mud or sand. Therefore, the skeleton has come to be com- pleted in this group by the constant addition of small ossicles movably articulated between the rows of larger ones, especially on the dorsal surface, both longitudinally and transversely. These later interpolated ossicles may form regular or irregular longitudinal rows between the median and marginal rows, or they may be so interpolated as to form a more or less regular reticu- lation ; in other cases they may be entirely irregular, with no deter- minate arrangement, seeming to be introduced wherever needed at the time. These variations in the modes of interpolation of the plates, together with their forms, which may vary from linear to broad, angular, lobate, and scale-like forms, combined with variations in the spines, give rise to the great variety of structures and forms seen in the dorsal surface of starfishes of this family. Moreover, in most species, the rapid increase of the dorsal skeleton alone apparently does not give sufficient space for the rapidly growing internal organs, especially the reproductive organs, within the bases of the rays, and therefore new rows of plates (fig. i, iii, iv) must be interpolated between the inferomarginals and the adambulacrals, and sometimes between the upper and lower mar- ginals, to increase the diameter of the rays. The most constant of these rows is that which I have designated as the peractinal (pa). This row appears very early in many species and often nearly reaches the tip of the ray, but usually not quite to the ocular plate. But the other actinal rows (subactinals or interactinals) (sa) are successively shorter and often do not reach the middle of the ray. The plates situated proximally in these rows appear earliest, and new ones are added at the distal end of each row. This is also the case with the interpolated dorsal and lateral rows. In many species, SHALLOW-WATER STARFISHES 33 as in those of Pisaster, the number of interpolated interactinal rows (sa) goes on increasing during the whole or most of the period of growth, so that in large specimens there may be four or five rows on each side. But in many other species the number of rows is fixed very early, the young, two inches in diameter, having as many rows as those six to eight inches in diameter, the subsequent increase in size being due to the enlargement of the plates and the formation of new ones dorsally and distally. But in many species with long, slender rays the peractinal plates alone are developed, and in a few even these are lacking, or else so small as not to be visible exter- nally, as in Urasterias linckii, etc. So, likewise, in other long-rayed species, the dorsal skeleton may lack more than one pair of interpo- lated rows of ossicles, as in Coscinasterias. B. INTERACTINAL PLATES AND SPINES. Text-figure i. The presence or absence of actinal plates and spines, and whether one row or several rows of these plates be developed, are characters that often seem to be of generic value. Those forms that have no such plates at maturity, or have only one imperfect row, are probably the more primitive types, or at least they have probably inherited and preserved this primitive character. The very young starfish in all the genera apparently has none of these plates, but acquires them as growth proceeds. They are lacking or rudimentary in Urasterias, Distolasterias, Stylasterias, and some other groups. They exist in only one row in Orthasterias, Coscinasterias, etc. In true Asterias and some other groups, one or two, or more, incomplete subactinal rows are usually added. In Evasterias, Pisaster, and Cosmasterias, two, three, or more rows are present. In those species that have subactinal plates, these increase in number during the whole period of growth, and new rows may also be added continually. Never- theless these plates seem to afford morphological characters of much value in classification, and they are usually coincident with other special features. It is of great importance that correct distinctions should be made between the actinal, infer omarginal, and supramar- ginal plates and spines. This has not been done in most of the earlier descriptions. Side views of the rays are necessary to show these plates properly, but such views are rarely given. In many species they are without spines and so concealed by thick skin that they cannot be determined without preparation, and therefore are not visible in alcoholic specimens. 4 34 VERRILL C. FORM AND ARRANGEMENT OF THE DORSAL OSSICLES. The form and arrangement of the dorsal ossicles vary considerably and often afford characters of much value. Aside from the stichas- terial arrangement, there are two main types, — the reticulate and the longitudinal, — in both of which the principal plates are lobed, substellate, or cruciform. The former, which is seen in true Asterias, Pisaster, etc., includes several subtypes, — as the stellate- reticulate, seen in Pisaster fissipinus; the areolate, seen in E. tro- schelii; and the irregular or indeterminate, seen in Asterias vulgaris, A. rubens, etc. These variations depend partly upon variations in the forms of the plates and partly on their arrangement and the existence of supplementary ossicles, and especially on the size and the form of the papular areas. The abactinal skeleton of the longitudinal type may consist of only three rows of stout dorsal plates, besides the marginals, with few or no supplementary ossicles, as in Coscinasterias; or the median may alone remain distinct, as in some species of Urasterias. But in other related groups there may be five or more regular rows. These last have but three rows when quite young, so that the forms with three rows are probably more primitive. Urasterias may be still more primitive. Its skeleton is very likely a degenerate type. In the stichasterial arrangement the plates become broader, more angular, and less lobed, and they are usually united directly together by their overlapping or adjacent edges in rather regular, longitu- dinal, imbricated rows ; or they may have a tesselated arrangement. The intervening papular areas are consequently small and rather regularly arranged. D. DORSAL SPINES; FORM AND ARRANGEMENT. The number, form, and arrangement of the dorsal spines vary widely. Their arrangement, when coincident with that of the plates, is a matter of much importance, but widely different appearances may occur in a single species, due to the variable number of spines that may occur on a single plate, combined with their variations in size and shape. No character is more likely to lead to mistakes in the determination of species and genera than the appearance of the dorsal spines, especially in those groups which, like restricted Asterias, have very numerous spines with an indeterminate or irregular arrangement. But in forms that have a limited number or regular arrangement of plates, the spines often conform strictly to SHALLOW-WATER STARFISHES 35 the plates, each plate bearing one or two spines, and then they may be very reliable guides, in case the specimens are adult. The number of plates or spines in a row, and often, also, the number of rows, increase with age. The supplementary or connective ossicles, when present, also increase with age, and may bear corresponding spines. Mr. Bell, in his arrangement (1881, p. 502), made a special dis- tinction between those species that bear certain spines on isolated or special dermal ossicles (autacanthid) and those that bear spines only on the regular skeletal ossicles (typacanthid). But the special der- mal ossicles, at least in most cases, seem to be merely new skeletal ossicles in process of formation, which are destined, a little later, to become articulated with the older ossicles. Hence this character seems to be of little importance and variable according to the age of the specimens, in most species. FIG. 2. Minor pedicellariae of Pycnopodia helianthoides (a, b) ; profile views. X 146. The dorsal and marginal spines may be similar or dissimilar. The spines may be large and long, or slender, short and thick, or capitate. They may stand singly on the plates, or in groups, or they may be reduced to the form of small spinules covering the plates in large numbers, or even nearly to granules, but they do not form paxillae nor pseudopaxillae. E. PEDICELLARIAE; SPECIAL FORMS. Peculiar or unusual forms of major pedicellariae occur on many species, and sometimes species of the most diverse structures and belonging to widely separate faunae will agree in having some of the felipedal type, mentioned above. Such special forms of pedicel- lariae are doubtless of great value for indicating community of origin and generic affinity, when they occur on species that are otherwise structurally similar ; but they must not be made to overshadow other structural features, for that would lead to absurd results, as bringing 36 VERRILL together such unlike forms as Coronaster of the North Atlantic and Stichaster of the South Pacific — forms that evidently belong to dis- tinct families. In the case of Pisaster, our eight species, though so different in their dorsal spines, all have similar and peculiar erect, stout, unguiculate pedicellariae of a type rarely to be found in other groups. Some of the remarkable forms of pedicellariae are probably sur- vivals of such structures present in remote ancestors, common to this and the allied families in former geologic periods, rather than structures independently developed in diverse genera, now living in different oceans. Their complexity of structure seems to be too great to have arisen independently in modern times. F. MADREPORITE OR MADREPORIC PLATE, AND SURROUNDING SPINES. Several writers have made considerable use of the character of this plate. In some of the species having more than six rays, or a variable number, there may be two or more madreporites. Such species are also, in most cases, subject to fission, as in Coscinasterias, Stephanasterias, etc. The number of madreporites is variable in each species of this kind, while closely related species may have but one. The madreporite is sometimes surrounded by a definite circle of special spines (echinoplacid Bell),1 but in closely related species no such regular circle of surrounding spines exists. In some cases different specimens of the same species may vary in this respect. Therefore it cannot be considered a character of more than specific value, and sometimes not even varietal. Bell, in his arrangement, appears to have given too much importance to it. The same remarks, as to variability, apply to a naked groove or narrow channel that sometimes surrounds the plate. The relative position of the plate and the number and arrangement of its gyri vary in different species, and also greatly with age, but are of some specific value in certain cases, as is also its color, in life. G. JAWS AND ORAL SPINES. Text-figure 3. The jaws in this family consist of an apical jaw-plate, composed of the first and second pairs of adambulacral plates closely united 1 Among the northwestern American species that are usually " echinoplacid " are L. aqualis, A. katheriruz, A. acervata, L, epichlora (variable), L. coei, E. troschelii (variable), etc. In general, species with small and numerous spines are apt to have this character. It is particularly conspicuous in Cosmas- terias lurida. SHALLOW-WATER STARFISHES 37 together and usually bearing six oral spines. The four terminal spines (peroral spines or papillae} are directed more or less hori- zontally over the mouth (a, of} ; the two outer ones (a') are diver- gent and smaller and are sometimes lacking. The outer angle of the jaw, formed by the second pair of adambulacral plates, bears an erect and usually longer pair of (epioral) spines (e, e). The characters of these oral spines and plates often afford good specific distinctions, and in some cases at least are of generic value ; but they have not yet been described or figured with care, except in a comparatively small number of species, so that at present their 11 FIG. 3- Orthasterias calif arnica V ., type, i, A jaw; a, a; a', a', apical or peroral spines with pedi- cellarix (p, p); e, e, epiorals; e', odd epioral; X 15; ii, an adambulacral spine. X 15. value is somewhat uncertain. In general characters they are remark- ably constant throughout this family, and in species that differ widely in other respects. This would indicate that their special arrangement has persisted for a very long period of time, for it must have been established before the world-wide dispersal of the family took place, and probably before the family itself was differentiated. In the calcareous peristomal ring the jaw-plates, formed by the union of two ambulacral plates, are usually more prominent than the plates alternating with them, known as the odontophores, formed by the union of two interambulacral plates. Such an arrangement is 38 VERRILL called an " adambulacral jaw" and it is characteristic of the family Asteriidse and of most other families of Forcipulosa. External to this proper jaw-plate are a series of two to ten or more pairs of adoral adambulacral plates, each pair closely pressed together (contingent), to form the adoral carina, which may be more or less compressed and thin, and is longest in those species with a large disk, like the Piasters, in which it often contains ten or eleven pairs of united plates. The spines (adorals) borne on these plates are often longer than those beyond. H. ADAMBULACRAL SPINES; ARRANGEMENT. Much importance has been given by many writers to the number of adambulacral spines on each of the plates. Many species have regu- larly a single spine on each plate, thus forming a simple regular row (monacanthid) . Others have regularly two spines to a plate, form- ing two rows (diplacanthid) . But still more frequently the spines stand irregularly, one or two to a plate, or alternately one and two, (subdiplacanthid) . In these the spines often appear to form three rows. In Allasterias Ver. the inner spine is situated higher up within the groove, on alternate plates. In some cases, as in Pisaster and Coscinasterias, the monacanthid condition is associated with other important characters, and in these it seems to be really of generic value ; but it should not be used as a character of primary importance, nor by itself, for generic divisions, for in some cases closely related species, or individuals of the same species, may differ in this respect.1 Yet it is a character ordinarily early fixed in the young starfish, and therefore important. I. NUMBER OF RAYS J VARIABILITY. The number of rays can only be used for subordinate purposes, for it is apt to vary sporadically, even in species that are the most constant, like Asterias vulgaris and A. forbesi of the Atlantic coast, in which we find, when vast numbers are examined, that a certain per cent will have six rays, and a smaller per cent seven rays and four rays, while eight-rayed and even nine-rayed examples may occur very rarely ; but those cases with seven and more rays may, in most cases, be due to abnormal repairs after injury. In certain species of Leptasterias that carry their eggs and young till they reach the period of bottom life, the young carried by the 1This is true of Coscinasterias tenuispina and C. acutispina (Stimp.)- SHALLOW-WATER STARFISHES 39 same parent may have different numbers of rays. Thus those carried by six-rayed parents of L. epichlora alaskensis, were found to be mostly six-rayed ; but about five per cent were five-rayed, and a few were four-rayed. (See below, and pi. LXXXV, figs. 2, a-/.) In other species and genera the number of rays is extremely variable, as in Coscinasterias, where they vary indifferently, from six to eleven or more. In Pycnopodia, additional rays bud in between the older ones in pairs during growth, so that the adult may have twenty to twenty-four rays. This variation is complicated in most species of the former group, and in Stephanasterias albula, by their remarkable habit of spontaneous fission (autotomy) and subsequent irregular replacement of the lost rays by each half. (See pis. LXXIII and LXXIV.) J. ARRANGEMENT OF THE AMBULACRAL FEET OR SUCKER-TUBES. These are generally arranged in four crowded rows in each groove, or sometimes in two zigzag rows, one on each side of the median line. But in many of the larger species with broad rays, as in the genus Pisaster, they may form six or even eight rows, espe- cially subproximally. Pycnopodia has four rows of sucker-feet sub- proximally, and two rows proximally and distally when adult. K. MODES OF DEVELOPMENT ; BROODING OF THE YOUNG J POSITION OF THE GENITAL PORES. Two widely diverse methods of development occur in the Asteriidae, as described above (p. 7). So far as now known, those genera that carry or brood their eggs and young (padophoric species} also have their genital pores situated on the actinal side, near the mouth ; while those that disperse their minute eggs, and whose young undergo a prolonged, free-swimming larval existence, have the geni- tal pores in the dorsal interradial areas. It is probable that this dif- ference is the most important morphological character by which the family, or at least the Asteriinse, can be divided into large generic groups. Unfortunately this feature has not been noted nor studied except in the case of a small number of the genera and species, so that at present, it can be utilized only in a limited number of genera. CLASSIFICATION OF ASTERIID^. As defined above, the Asteriidse would include, as subfamilies, Asteriinae, Stichasterinse, Pycnopodiinae, and Heliasterinae:=Helias- teridae Sla. 40 VERRILL Subfamily H ELI AST ERIN ;£. Subfamily Heliasterince VIGUIER. Family Heliasteridce Perrier; Sladen. The Heliasterinae do not seem to me to be worthy of more than subfamily rank. They agree with Asteriidae in nearly all important structural characters. They differ mainly in having very numerous short rays and a very broad disk, but in these respects they are closely approached by Pycnopodia and Coelasterias. Their dorsal, ventral, and ambulacral skeletons are distinctly like Asteriidae. The only distinctive characters of much value are the restriction of the stomach to the disk by a discobrachial septum, and the existence of double septa between the rays. The group, which includes only the genus Heliaster, is confined to the Pacific coast of America and the adjacent islands, from Chile to Lower California. (See H. L. Clark, op. cit, 1907, plates i-vm, for details of structure and descriptions of all the species.) Subfamily PYCNOPODIIN&. For the characters of this subfamily, see below, p. 197. Subfamily STICHASTERINJE. The most developed forms of Stichasterinae,1 e. g., Stichaster striatus = aurantiacus, have several regular rows of interactinal plates ; other forms have only one or two rows. Some of the As- teriinae (Ccelasterias, Pisaster, etc.) also have from three to six rows of such plates. On the other hand, some forms of Stichasterinae (Stichastrella rosea, Stephanasterias albula, and Granaster} have the dorsal ossicles more or less irregularly arranged ; and even the median row may not be distinct in S. albula. In the latter the ven- tral plates bear longer spinules. Ccelasterias V. is an intermediate genus with multiple rays. The Stichasterinae have, at present, no known representatives on the northwestern coast, unless we so reckon the Stephanasterias 1 Stichaster M. and Tr. (April, 1840) was monotypic. Its type was S. stria- tus M. and Tr. = Asterias aurantiaca Meyen (not of Linne) = Tonia atlantica Gray (Sept., 1840). Tonia is a complete synonym of Stichaster, which has about six months priority. The European species, A. rosea, usually referred to Stichaster, and taken as its type by many writers, is a distinct genus which, so far as I know, has no valid name. It may be called STICHASTRELLA, S. rosea being the type. It has two or three rows of interactinal plates ; adambulacral plates bear two or three spines irregularly. Carinal rows of plates and spinules are usually distinct, but only by the slightly larger size of the plates. SHALLOW-WATER STARFISHES 4! albula,1 which is a very widely distributed North Atlantic and Arctic species, which has been found at moderate depths off the northern Alaskan coasts. I consider it one of the Asteriinse, allied to Lept- asterias. It is autotomous while young, with a variable number of rays, but when large it is usually regularly six-rayed. The consideration of the importance of the stichasterial or imbri- cated arrangement of the dorsal ossicles early led to the separation of Stichaster as a genus, and later as the type of a family, but perhaps even that character does not always indicate close affinity nor an identical origin, for it may have been developed independently in different regions and in distinct lines of descent. As a matter of fact, we find the greater number of the species, and those that are the larger and more typical, living in the Antarctic seas and on the southwestern coast of South America, while only a few rather small and less characteristic forms exist in northern seas. It is not unlikely that the northern forms, like S. rosea and S. albula, originated entirely independently of those of South American seas, and should, therefore, be classed as distinct genera. This is indicated, moreover, by their morphological characters. S. albula, especially, differs but little from some forms of Lepastcrias, and may well have been developed from some member of that or a similar group. Its embryology is not known. However, all recent investigators admit that the stichasterial arrangement of the dorsal ossicles is at least of generic value.2 Other variations in these plates may be of equal value. The discovery of many new generic and specific types intermediate between typical Stichaster and Asterias, as already intimated, renders it difficult to define the limits of the two so-called families, typi- fied by these genera. 'This is the Asterias albula (Stimp.) = Stichaster albulus Ver. = Stephan- asterias albula Ver. = Stichaster albulus S\aden = Nanaster albulus Perrier, 1894, p. 131. It is found from the Arctic Ocean to South Carolina and Georgia, in deep water, on the east American coast, and as a littoral species it occurs as far south as the Bay of Fundy. It is also common on the northern Euro- pean coasts. 2 Sladen has even gone so far (op. cit., p. xxxvi, 1889) as to widely separate Stichasteridae and Zoroasteridse from the Asteriidae on this character, almost exclusively, placing them between Linckiidae and Solasteridae — families that do not belong to the Forcipulata. This seems to me a very unnatural arrange- ment, due to overestimating the importance of the character of the dorsal ossicles. 42 VERRILL Sladen and Perrier disagree both as to the character and limits of the two groups, because certain genera and species present characters of both groups, or have intermediate characters. Therefore it seems necessary to reunite them in one family. However, it may be convenient to retain the groups as subfamilies or sections with no very definite limits. A pseudostichasterial condition is sometimes produced by the existence of clusters of spinules in regular rows, on the transverse dorsal connective ossicles. This cannot always be determined with- out removing the investing integument. Perrier considered the characters of the dorsal spinules as of para- mount importance in this group. He would place all the genera with distinct spinules or spines in Asteriidae, and those with granule-like spinules in Stichasteridae ; 1 but in fact there are all intermediate gradations in the character of the armature, and it is often difficult to say whether these structures, in some species, should be called " granules " or " spinules." This seems to me a character only of specific value.* Subfamily ASTERIINJE. Asteriidce (restr.) SLADEN, op. cit, 430, 560, 1889. Perrier, 1894, pp. 105, 128; 1896, pp. 25-31. As here limited this subfamily corresponds nearly with the re- stricted family Asteriidse in the systems of Sladen and of Perrier. The Asteriinse3 are chiefly characterized by the more or less openly reticulated arrangement of the dorsal and lateral ossicles, especially of the dorsal ossicles, which are usually more stellate or lobulate and more slender than in the Stichaster group, and generally united either by the ends of the lobes or apophyses, or else by means of interpolated ossicles, thus allowing more flexibility in the rays, and larger papular areas. The Stichasterinae, on the contrary, are chiefly characterized by the more or less regular subtesselated or imbricated arrangement of the 1 S. gracilis and 5\ albula, with other related forms, have definite spines. "The genus Calvasterias Perrier, 1895, has about five series of lobed and imbricated, flat dorsal plates, bearing few small short spines, and entirely covered with a thick, naked, soft, canaliculated dermis. The character of the plating should cause it to be placed in the Sti- chasterinae. It is monacanthid and there is a row of small spiniferous, interactinal plates, at least in C. stolidota Sla., of Chile and the Falkland Islands. 3 This form of spelling the word seems necessary, because it is derived from Asterias, not from Aster. SHALLOW-WATER STARFISHES 43 ossicles, both on the sides and dorsal surface of the rays, the ossicles themselves being larger, broader, and more rhombic, and usually united in more definite, longitudinal rows, with smaller and more regular intervening papular areas. In either group, as at present understood, the interactinal or ventral plates may have the regular serial and subtesselated arrangement. A. GENERIC SUBDIVISIONS OF ASTERIIN^E. The genus Asterias, in the wider sense, as still used by many writers, includes a very large number and a great variety of species, found in all seas. Various attempts have been made to divide the genus into a number of genera or subgenera on structural char- acters, but hitherto there has been no general agreement among writers, as to the number or limits of most of the subdivisions. Nor do I flatter myself that my own views will be altogether acceptable. One of the best known classifications hitherto proposed is that of Sladen.1 He admitted, in the restricted family Asteriidse (our Asteriinse), five distinct genera, including Pycnopodia. Under Asterias he had six subgenera. Most of these are undoubtedly worthy of generic rank. One of them (Leptasterias) had been proposed as a genus by the present writer, many years earlier, and another (Stolasterias) is essentially identical with our Coscinasterias, 1867. His subgenus Hydrasterias is evidently a distinct genus, peculiar to the deep seas, and has already been so recognized by Perrier and myself. Perrier, in a later work,2 recognized all of Sladen's divisions as genera, but with the limits modified in some cases, and proposed four more generic subdivisions, some of which seem to have no great syste- matic value. This is particularly, true of Diplasterias, separated from Asterias mainly on account of the two regular rows of adambu- lacral spines, — a character that is variable in this group and often of no more than specific importance, taken alone. Many species of Asterias have alternately one or two adambulacral spines, and others have irregularly one and two to a plate. C. tenuispina is generally strictly monacanthid, but unusually large specimens usually bear two spines on a few of the plates proximally. Perrier, in the article referred to, recognized fifteen genera in the family. 1 Voyage Challenger, Zool., xxx, pp. 560-564, 1889. 2Exped. Scientif. du Travailleur et du Talisman, Echinodermes, pp. 108, 109, 1894. /j /| VERRILL One of his diplacanthid genera (Podasterias) had P. lutkeni Per. (not Asterids liitkenii Stimpson) as its type, and is, therefore, of special interest in this connection. In a subsequent work 1 Perrier has added two more genera: Distolasterias, for D. stichantha (Sladen), separated from Stolasterias mainly on account of its two rows of adambulacral spines ; and Sclerasterias, for S. guernei Per., a new and peculiar type. But in this later scheme he has judiciously omitted Diplasterias ( Perrier, non Kcehler, nee Ludwig) . After a careful study of a large number of species, belonging to these various divisions, I am convinced that several other genera or subgenera, based on structural characters of quite as much impor- tance, can be distinguished. As understood in this work, the genus Asterias (type A. rubens L.) will be used in a sense much more restricted than by Sladen. I propose to keep separate, as genera, most of the groups called sub- genera by Sladen — as, for example, Cosmasterias, Smilasterias, Coscinastcrias, Hydrasterias — and most of the groups more recently proposed by Perrier. Some of these generic groups need to be modified in definition or extent, and others should be added. (See below, under Geographical Distribution, for the characters of antarctic genera.) I am well aware that many students of these starfishes do not think it worth while to subdivide Asterias, even to the extent that was done by Sladen and by Perrier. But in the case of so extensive a group, with a world-wide distribution, it will be practically impos- sible to intelligently study or discuss its evolution and distribution unless we recognize the relationship between allied species by means of distinctive group-names for minor groups, recognizable by mor- phological characters. We should aim at a phyllogenetic classifica- tion, and this is more likely to be attained by the recognition of mor- phologically similar groups of species than by an indiscriminate lumping of all these diverse forms in one great genus, containing perhaps two hundred species or more. B. SUBDIVISIONS OF THE ASTERIIN;E HAVING DEFINITE LON- GITUDINAL ROWS OF DORSAL PLATES AND SPINES. Among the numerous species in which the dorsal plates and spines have a constant and characteristic arrangement, there are several divisions, apparently of generic and subgeneric value. Several of 1 Contr. a 1'etude des Stellerides de 1'Atlantique Nord, Resultats des Cam- pagnes Scientif., faits par Albert I, Prince de Monaco, fas. xi, pp. 34, 35, 1896. SHALLOW-WATER STARFISHES 45 these are not yet known from the northwestern coast, but are likely to occur hereafter, when more dredging shall have been done, and therefore it may be useful to give here a summary account of the principal groups of this kind. Genus Coscinasterias Verrill. Coscinasterias VERRILL, Trans. Conn. Acad. Sci., i, p. 248, 1867. (Type, C. muricata V. = (?) C. calamaria (Lam.).) Stolasterias (pars') SLADEN, Voyage Chall., vol. xxx, pp. 563, 583, 1889. (Type, C. tenuispina.) Non Perrier. Coscinasterias and Polyasterias PERKIER, Exped. Trav. et Talisman, p. 108, 1894. Body small, rays elongated, somewhat angular, usually carinate. Dorsal ossicles of the rays stout, four-lobed, usually arranged in three or five regular rows, besides the upper marginals, which form regular lateral rows. Actinal ossicles usually consist of one primary (peractinal) row, like the lower marginals, but they may be rudi- mentary and spineless in half-grown specimens, and only bear spines in large, mature individuals, as in C. tenuispina. Minor pedicellariae are abundant. Large forficulate or major pedi- cellariae of the usual forms are present, often in considerable num- bers, on the dorsal and lateral plates, and smaller ones occur on the marginal areas. Large unguiculate pedicellarise are lacking. Adambulacral plates monacanthid or nearly so. Often autotomous; rays variable in number. Madreporites often two or more. Stolasterias, which was proposed as a subgenus by Sladen in 1889, was nearly identical with Coscinasterias, characterized by me in 1867. He gave no reason for changing the name. It should be regarded as a synonym of the latter and be eliminated. Sladen's type was A. tenuispina, which is a Coscinasterias with the peractinal plates . rudimentary and without spines, except in the larger specimens. M. Perrier (1894) correctly retained Coscinasterias for the typical forms, like calamaria, but separated those species that are known to undergo spontaneous fission under the name of Polyasterias = typical Stolasterias Sla., though they do not appear to differ much in structure from the preceding, while he retained Stolasterias Sla., in a restricted sense, for those allied to glacialis, although C. tenui- spina was named by Sladen as the type. As C. tenuispina often divides spontaneously, it belongs to Poly- asterias in Perrier's arrangement. This would make the latter strictly synonymous with Sladen's typical Stolasterias, which should 46 VERRILL be restricted to this type as a subgenus, if retained at all. The group including glacialis had also been previously named Marthr- asterias by Jullien (Bull. Zool. Soc. France). No species of typical Coscinasterias is known to me from the northwestern coast, but a species (C. acutispina Stimp., sp.), similar to C. tenuispina, and often having some diplacanthid adambulacral plates, occurs at Ousima Island, Japan. It is autotomous. C. TABLE OF EXTRALIMITAL SPECIES OF COSCINASTERIAS AND CLOSELY ALLIED FORMS. A. More than one distinct row of dorsal spines (usually three rows). B. One row of peractinal spines and plates well developed. Rays variable, six to twelve. Mostly fissiparous. C. Rays mostly seven to twelve. Adambulacral spines strictly uniserial (monacanthid). Lower marginals bear two spines each. Usually two or more madreporic plates are present. C. muricata (VER.) 1867. (? = C. calamaria.') C. calamaria (LAM.). Australia; Indian Ocean. C. jehennesii (PER.). Madagascar. (? = calamaria, teste Sladen.) C. echinata (GRAY). Valparaiso. Rays eight. C. gemmifera (PER.). Chile; Fiji Is. ( ? = echinata.) C. (f)aster (GRAY). Rays twelve to thirteen. Five rows of dorsal spines (Gray). CC. Some of the adambulacral plates diplacanthid. Rays variable. C. acutispina (STIMP.). Ousima Island. C. tenuispina LAM. (Large specimens.) BB. No interactinal spines; peractinal plates rudimentary or lacking. AA. Only one row of large dorsal plates and spines. Adambulacral plates monacanthid. b. Rays mostly five to nine. Fissiparous. Subgenus Stolasterias (Sla. restr.). C. (S.) tenuispina (LAM.) when young. Eastern Atlantic; Brazil; West Indies. Variety atlantica (VER.). Bermuda; Cuba, bb. Rays constantly five or six. Type not fissiparous. Marthasterias (JuL.) when young. Type, M. glacialis (MULLER). D. OBSERVATIONS ON VARIOUS GENERA. Genus Heterasterias Verrill, nov. Type, Stolasterias volsellata Sla., Philippines. The rudimentary state of the dorsal skeleton and the lack of peractinal plates in the named species may entitle it to be con- sidered the type of a new generic division, as suggested by Sladen. This view is strengthened by reason of the presence of large ungui- culate dermal pedicellariae. In this character and in general appear- SHALLOW-WATER STARFISHES 47 ance it resembles Coronaster, but the latter has only two rows of ambulacral feet. I would suggest Heterasterias, as a genus; type, H. volsellata. Genus Marthasterias Jullien. Marthasterias JULLIEN, Bull. Zool. Soc. France, p. 141, 1878. Stolasterias (pars} SLADEN, Voy. Challenger, xxx, p. 563, 1889. Perrier, Voy. Trav. et Talism., pp. 108, 109; Resultats Camp. Scient., i, p. 34, 1896. Rays five, angular, normally with three dorsal radial rows, besides the superomarginal rows, of stout plates bearing large, mostly conical spines ; usually two on the superomarginals. Inferomarginal plates usually with two rows of spines. Adambulacral plates generally monacanthid, each plate with a single spine. Peractinal ossicles small and without spines, scarcely visible externally.1 When young it has only one row of dorsal spines, the median, developed; when very large there may sometimes be more than three rows. Besides the type, M. glacialis Miiller, this group includes M. africana, of South Africa, and M. rarispina (Perrier). The allied diplacanthid species were referred to another generic group (Distolasterias) by Perrier. Genus Distolasterias Perrier. Distolasterias PERRIER, Resultats des Camp. Scient., i, p. 34, 1896. This name was proposed by Perrier for a group of which Stol- asterias stichantha Sladen, of Japan, was the type. It is distin- guished from his Stolasterias= Marthasterias by the diplacanthid arrangement of the adambulacral spines ; and by having five or more regular close longitudinal rows of dorsal plates. The actinal plates are rudimentary and without spines. But the Stolasterias neglecta of Perrier (1896, p. 37, pi. i, figs. 2, 2a, 4-40) is diplacanthid and therefore does not go in that genus, as he defines it, for in the same work (op. cit, p. 35) he makes the genus monacanthid. In this respect neglecta is like his Distolasterias (op. cit., p. 34), but the type of the latter has a much more complex dorsal skeleton, with five or more rows of dorsal plates and spines. 6*. neglecta apparently belongs with a group of which A. forreri Lor. may be taken as the type. *For description and figures of skeleton, see Viguier, op. cit, pp. 100-105, pi. v, figs, i-io, 1878. 48 VERRILL Genus Orthasterias Verrill, nov. Rays long, usually five, bearing three, five, or more dorsal rows of rather long spines, besides a superomarginal row on each side. Inferomarginals bear each one or two spines, usually two. One row of interactinal or peractinal plates, with or without spines, close to the adambulacrals ; the latter are diplacanthid ; a short subactinal row may also occur rarely. Lateral and dorsal dermal pedicellarise are large, partly wedge- shaped or spatulate, often dentate, or unguiculate. Type, O. colum~ biana Ver., sp. nov. Also includes O. dawsoni Ver., nov. ; 0. tanneri Ver. ; O. biordinata Ver.; O. calif arnica Ver., etc. Subgenus Stylasterias Verrill, nov. Type, S. forreri Lor. Dorsal plates in the adult usually in three regular rows, having elongated spines. Upper and lower marginal plates large and regular, with a well marked channel between, crossed by the large descending apophyses of the upper plates, and carrying papulae and large forficiform pedicellarise. Lower marginal plates close to the adambulacrals, each bearing one or two long spines. Peractinal plates lacking or rudimentary; when present never bearing spines, and so small as to be easily overlooked, without dissection. Adam- bulacral spines diplacanthid. Among other species that belong to this group are the following : S. enopla Ver., off Nova Scotia. S. neglecta (Per.), East Atlantic. S. contorta (Per.), West Indies. Genus Cosmasterias Sladen, 1889. This name was proposed for the group having as the first species, Asterias (Cosmasterias) tomidata Sladen (op. cit, pi. cv, figs. 8-10), and including also Asterias sulcifera Perrier, 1869 = C. lurida (Phil- ippi), 1858. Perrier has adopted Cosmasterias with C. sulcifera1 (Per.), from off Patagonia, etc., as the type. The genus is diplacanthid and has several regular rows of inter- actinal spines, but it has large, unguiculate or felipedal, dermal, 1 For full synonymy and new descriptions of this species and C. tomidata, see Leipoldt, Asterioidea der Vettor Pisani Exped., 1895, PP- 552-563. See, also, below, under Geographical Distribution. SHALLOW-WATER STARFISHES 49 dorsal pedicellariae, not found in northern polyactinoplacid types. Its principal dorsal plates are in regular radial rows. This suggests an independent line of development, and hence the group may well receive a distinctive name. C. tomidata (Sla.), the type, was from Chile, in 45 fathoms. Somewhat similar, large, unguiculate or felipedal pedicellarise occur on Stichaster polygrammus Sla., off Patagonia ; S. felipes Sla., South Africa; Stolasterias volsellata Sla., Philippines; Coronaster briareus (V.), Atlantic, and some other species.1 In these they are not so sessile and rigidly erect as in Pisaster. Each of those species named is now referred to a distinct genus. E. TABLE OF PRINCIPAL GENERA OF ASTERIIN/E WITH REGU- LAR LONGITUDINAL ROWS OF DORSAL PLATES AND SPINES. I. NOT MORE THAN ONE ROW OF EVIDENT ACTINAL (PERACTINAL) PLATES, — MONACTINOPLACID OR ANACTINOPLACID. A. Monacanthid, or nearly so. B. Rays slender, variable, mostly six to eleven; often more than two madre- poric plates when adult; usually autotomous. C. One (often incomplete) row of actinal plates, closely united to adambu- lacrals, and usually bearing a simple row of spines (monactino- placid). Major pedicellariae large, compressed, ovate or pointed. Usually three or five rows of dorsal plates and spines, a. Peractinal spines usually present in the adult. Coscinasterias Ver. Type, C. muricata Ver. aa, Peractinal spines mostly absent. Section Stolasterias (Sla.), restr. Type, C. (S.) tenuispina. CC. No visible actinal plates, the inferomarginals joining the adambulacrals (anactinoplacid). Only one row of dorsal plates — the median radials. Some large, unguiculate, dermal major pedicellarias ; some of the plates may be diplacanthid. Heterasterias Ver., nov. Type, H. volsellata (Sla.) BB. Rays five or six, stout. Three or five rows of strong dorsal plates and spines. No interactinal plates with spines. Usually but one madre- poric plate; rarely two. Marthasterias Jullien. Type, M. glacialis (Mull.). AA. Adambulacral plates mostly diplacanthid. D. No evident interactinal plates that bear spines; if present, rudimentary. a. Five or more rows of stout dorsal plates and spines when adult. Major pedicellarias not felipedal. Distolasterias Per. Type, D. stichantha (Sla.). *They often resemble, more or less, a pair of miniature clasped hands with fingers interlocked or two catspaws with claws protruded and interlocking, and hence they may be called felipedal. The name of S. felipes Sla. refers to this feature. 50 VERRILL aa. Only three regular rows of dorsal plates with spines. Stylasterias Ver., nov. Type, S. forreri (Lor.), aaa. Usually only one row of dorsal plates (median) developed; they are broad, covered with a thick channeled dermis. Sclerasterias Per. Type, S. guernei Per. E. Atlantic. DD. One row of interactinal plates without or with spines (monactinoplacid). Three to five or more rows of dorsal plates and spines. Major pedicellariae of two kinds. Some compressed, lanceolate or ovate; others large, thick, spatulate or wedge-shaped. Rays five or six. Orthasterias Ver., nov. Type, O. columbiana Ver., nov. II. TWO OR MORE ROWS OF ACTINAL PLATES AND SPINES (POLYACTINOPLACID) . A. Diplacanthid. Special large dermal pedicellariae are unguiculate or feli- pedal. a. Spines vesiculated. Cosmasterias Sla. C. tomidata (Sla.). aa. Spines not vesiculated. Cosmasterias lurida (Phil.) =sulcifera Per. R SUBDIVISIONS OF ASTERIIN^E WITH RETICULATED DORSAL SKELETONS. This large group presents fewer morphological characters for gen- eric and subgeneric distinctions than the last, and some of them present gradations and intermediate conditions. However, if we separate the genus Pisaster and consider the large group to which Asterias rubens, A. vulgaris, and A. forbesi belong as typical Asterias, there will be left several more or less differentiated types of structure. Genus Leptasterias Verrill. Type, L. millleri (Sars.). This group, as separated by me, 1866, differs from true Asterias plainly in having only a few large papulae in each cluster. Its typical species are L. mulleri, L. compta, L. tenera, etc. In these the dorsal spines are slender and numerous and the interactinal spines are in one row (rarely two). The discovery of numerous additional species, of larger size, and having more papulae, now renders it diffi- cult to make any sharp distinction, in this respect, between the two groups. Hence I have sometimes considered it as of only subgeneric value, and closely connected with typical Asterias. Its typical species carry the large eggs and young attached to the oral region, and the development is direct or abbreviated. SHALLOW- WATER STARFISHES 51 The most important generic feature is the fact that the genital pores are on the actinal side,1 between the proximal inferomarginal plates, not dorsal, as in Asterias. The ovaries consist of compara- tively few large digitate or glomerate tubules which, when filled with the large ova, appear beaded. The genital ducts are short and wide. Genus Evasterias Verrill, nov. Type, E. troschelii (Stimpson). We may also distinguish this as a generic group, typified by A. troschelii. In this the dorsal plates and ossicles are small and reticulated, while there are three or more regular rows of stout inter- actinal spines and plates. The increased number of rows of inter- actinal plates and their closer union are the principal distinctions in this case, but the number of plates varies with age, so that the young specimens are essentially like Asterias in this respect. Genus Urasterias Verrill. \ Plate LXX, figures 1-4. Type, U. linckii (Miiller and Troschel.) =A. stellionura Perrier. This type differs so much in structure from all others named that it should probably be considered a distinct genus. The dorsal skeleton is very feebly developed, consisting mostly of small plates and delicate, short, rod-like or linear ossicles, united in an irregular network, with large meshes, but having a median radial row of rather larger, slightly lobed, spiniferous plates, and some similar lobed plates at the intersections of many of the meshes. The dorsal spines are slender, irregularly scattered, except the radials. Disk covered with a network of stouter, somewhat lobed ossicles, bearing solitary spines, and usually having in the type a larger flat plate, or group of such plates, at each interradial area, above, and extending to the lower side, and usually perforated by a pair of papular areas. This peculiar group of plates seems to be made up of a considerable number of plates in the adult, often united 1 This is true also of the genus Sporasterias of southern South America and the Antarctic Ocean, including 5". antarctica (Lutk.), the type = Asterias rugispina Stiinp. = A. spirabilis Bell = A. varia (Philippi) ; 5". rupicola Ver. ; and several other nominal species; and probably, also, of A. studeri Per.; Anasterias chirophora Lud. ; A. belgicce Lud. ; A. perrieri (Stud.) ; Podasterias liitkeni (Per.) ; Stichaster nutrix (Stud.), which carry their young in the same or a similar manner. They are all antarctic. 52 VERRILL together in young specimens. Upper and lower marginal plates wide and strong, each bearing, in the young, a single spine. The upper marginal plates have a large and wide descending lobe ; these join the stout upper lobe of the inferomarginals, but leave wide papular areas between them, along the wide lateral channels. The upper marginals have small side lobes, which are united together longitudinally, by smaller narrow, interpolated spineless plates. Peractinal plates are small, or rudimentary without spines. The adambulacrals generally appear to be joined directly to the infero- marginals, but minute ossicles may intervene. Adambulacral plates diplacanthid or subdiplacanthid. (PI. LXX, fig. i.) Major pedicellariae are large and numerous, especially between the inferomarginal spines and on the inner margins of the grooves, but also occur on the back. They are acute-ovate and a little com- pressed. The minor pedicellariae are unusually large and denticu- late; they form very large pedicellate clusters, especially on the inferomarginal spines. Ambulacral pores and feet very large. The lack of distinct peractinal plates and spines and the presence of the large interradial plates are very important morphological char- acters, aside from the feeble dorsal skeleton. The large size of the ambulacral pores is a notable feature. The type, U, linckii = stellionura (Per.), is found from Spitz- bergen to Nova Scotia.1 Another arctic form, U. gunneri (Dub. and Kor.), is closely allied and may be only a variety (teste Ludwig). One or both of these may very likely be found hereafter in Bering Sea, for they are probably circumpolar. Another arctic species, U. panopla (Stuxberg) seems to be con- generic with U. linckii. It has a similar, very feeble, reticulated dorsal skeleton, and lacks distinct peractinal plates and spines. Its proximal adambulacral plates in some cases bear three or four spines, but are mostly diplacanthid. The superomarginal and inferomarginal plates are cruciform and are connected by a narrow, spineless, intermediate plate. The inferomarginal ossicles usually bear two spines, some- times more in large examples. It is from Spitzbergen, Barents Sea, Kara Sea, Finmark, etc. It is remarkable for the great number and large size of the dorsal minor pedicellarise. 1 It has been found as a fossil abundant in the postglacial clays at Salem, Massachusetts. SHALLOW-WATER STARFISHES 53 Genus Parasterias Verrill, nov. Type, P. albertensis Verrill, British Columbia. Five rays, stout; dorsal plates openly reticulated, much as in Asterias; interactinal spines and plates lacking ; adambulacrals dipla- canthid. For more details see below, p. 187. Genus Ctenasterias Verrill, nov. Type, C. spitzbergensis (Dan. and Koren). Dorsal skeleton composed of an irregular network of slender, but numerous, mostly transverse ossicles, bearing large numbers of minute, uniform spines, arranged in transverse rows or combs on the sides of the rays. Marginal spines in two double rows near together. Adambulacral plates diplacanthid, or partly triplacanthid. Two forms of major pedicellariae ; the larger ones are lyrate or cylindrical and obtuse. A single row of small interactinal spines. Probably closely allied to Leptasterias. Reproduction not known. C. cribaria (Stimpson) is pretty certainly identical with the type, which is from Spitzbergen. Genus Allasterias Verrill. Type, A. rathbuni (Verrill). Bering Sea. Adambulacral spines alternately diplacanthid and triplacanthid, or monacanthid and diplacanthid; the inner spine of the larger alter- nate group is inserted on an angle of the plate within the edge of the furrow. Dorsal skeleton openly reticulated; superomarginal plates and spines form an angular margin. Genus Sporasterias Perrier. Type, 5". antarctica (Lutk.) =A. rugispina Stimp. (teste Leipoldt). Adambulacral spines regularly monacanthid. Dorsal skeleton irregularly reticulated, sometimes nearly as in some species of Lept- asterias, sometimes weakly developed.1 Genital pores ventral ; the eggs and young are carried over and around the mouth, as in Lept- asterias. For details, see below, under Geographical Distribution, p. 355- Genus Smilasterias Sladen. Type, S. scalprifera (Sladen). Off Patagonia, etc. Adambulacral plates diplacanthid or triplacanthid, the spines and those of the inferomarginal plates large, flat, and placed in transverse rows. Dorsal skeleton irregularly reticulated with small spines. 1 In 5". rupicola Ver., from Kerguelen Island, the dorsal skeleton is well developed, with short, stout ossicles and numerous small spines. 54 VERRILL Genus Meyenaster Verrill. Type, M. gelatinosus (Meyen, 1834). Chile. Meyenaster VERRILL, op. cit, 1913, p. 485. Adambulacral plates monacanthid; one row of larger interactinal spines on thick ovate plates, each in line with two similar spines on each inferomarginal plate; superomarginal plates stout, each alter- nate one with a single large spine ; a wide intermarginal lane bearing a regular row of papular areas, twice as many as superomarginal spines, and many large pedicellarias. A row of median radial stout spines, usually one on each alternate plate; in large specimens an intermediate imperfect row on each side; medio-lateral skeleton reticulated irregularly with numerous short, transverse and oblique ossicles defining five or six irregular series of large papular areas on each side. Central part of disk symmetrically stellate with a central and twelve surrounding spines, in type. Large, stout pedi- cellariae dorsal, lateral, and intra-ambulacral, compressed, ovate-tri- angular, not unguiculate ; many very small acute, major pedicellariae on papular areas ; minor pedicellariae abundant on large, broad, fleshy circumspinal sheaths. A thick, strongly canaliculated skin covers and conceals all the plates. G. ANALYTICAL TABLE OF THE GENERIC AND SUBGENERIC GROUPS OF NORTHERN ASTERIIN^ HAVING RETICULATED DORSAL SKELETONS. A. Monacanthid. Dorsal skeleton coarsely reticulate. Interactinal spines and ossicles well developed, in several rows. Large unguiculate erect pedicellarise are present; rays five or six. Pisaster (Ag.). Type, P. ochraceus (Brandt). AA. Usually diplacanthid, or alternately diplacanthid and monacanthid, rarely triplacanthid. Interactinal plates and spines present or absent, or rudimentary. B. Adambulacral spines confined to the outer or actinal surface of the plate. C. Dorsal plates mostly angular or lobed. Interactinal plates and spines forming at least one row (peractinal), usually with other shorter proximal rows. Both rows of marginals well developed. Asterias, sens ext. D. Dorsolateral plates and transverse ossicles openly reticulated. a. Interactinal plates strong, closely united, or imbricated in three or more rows in the adult, bearing several regular close rows of nearly uniform spines, like the inferomarginals ; intervening papular areas small. Two kinds of major pedicellariae usually present Dorsal spines unequal, several on a plate, reticulate or areolate. Genus Evasterias V. Type, E. troschelii (St.). SHALLOW-WATER STARFISHES 55 aa. Interactinal plates not so closely united; usually one or two rows, with larger papular areas between. Major pedicellariae usually of but one kind. Dorsal spines rather irregularly arranged, usually scattered, sometimes areolate or acervate. b. Papular areas large, bearing numerous small papulae. Larvae are usually free-swimming brachiolariae ; genital pores dorsal. Asterias, restr. Type, A. rubens L. bb. Papular areas smaller and bearing fewer large papulae. Eggs and larvae are usually carried by the mother around the mouth. No free larval stages ; genital pores ventral. Genus, Leptasterias Ver. Type, L. mulleri (Sars). aaa. Interactinal plates lacking or rudimentary; papular areas large, numer- ous. Parasterias, nov. Type, P. albertensis Ver., nov. DD. Dorsolateral plates and ossicles wide, irregular, rather closely joined, leaving only small papular areas in longitudinal lines, but the ossicles do not form regular rows. All the dorsal and marginal plates are covered with close clusters of very small spines, longer on the inferomarginal and interactinal plates. Autotomous; rays mostly two or three to nine ; finally six in adult. Stephanaslerias Ver. Type, S. albula (Stimp.). Circumpolar; North Atlantic; Bering Sea. CC. Dorsal plates mostly rod-like. Interactinal plates small or rudimentary, mostly spineless. Usually two kinds of major pedicellariae. c. Dorsal skeleton feeble, composed mostly of small, linear or roundish, iso- lated and reticulated ossicles. Dorsal spines few and mostly scat- tered, but usually forming a median radial row. Both rows of mar- ginal plates well developed and rather large, bearing single or double rows of stout spines, separated by a wide lateral channel. Dermal major pedicellariae large and abundant, wedge-shaped. Minor pedicellariae abundant on the spines and skin, attached to a large fleshy sheath on the spines. Alternate superomarginals with- out spines. Urasterias Ver. Type, U. linckii (Mull, and Trosch.). cc. Dorsal skeleton composed of nearly uniform short linear rods, irregularly reticulated, and covered with numerous small, uniform spinules, often in transverse rows or combs on the sides. Two rows of marginal plates near together, in the type bearing double rows of spines. Larger major pedicellariae obtuse, large. Ctenasterias Ver. nov. Type, C. spitzbergensis (D. and K.)- BB. The inner adambulacral spine, on alternate plates, is attached to the inner edge of the plate, or within the groove. Peractinal plates feebly developed or lacking. Dorsal spines small and numerous. Allasterias Ver. Type, A. rathbuni V. H. ASTERIID;E OF THE NORTHWESTERN COAST OF AMERICA. This faunal region contains a remarkable assemblage of the repre- sentatives of this family. A few species from north of the Aleutian Islands are of arctic or circumpolar origin, but most are endemic. The most remarkable genus is Pycnopodia, strictly confined to this 56 VERRILL region. The genus Pisaster is here represented by seven or eight large species, some of them being among the largest and most mas- sive starfishes known. It is doubtful if the restricted genus occurs in any other fauna. The genus Orthasterias, with seven or eight species, is uncommon elsewhere. Its nearest allies are Atlantic species and Distolasterias of Japan. A large proportion (fifteen) of the species of this region are normally six-rayed, which is a remarkable peculiarity. Over twenty nominal species of Asteriinae had been already de- scribed from the Northwest Coast, between southern California and the Arctic Ocean. In the collections studied by me there are about thirty additional species, besides twenty well-marked new varietal forms, or a total of about seventy. Probably no other fauna is so rich in species of this group.1 It is not improbable, however, that some of the supposed species may prove to be only local varieties, when larger series can be carefully studied. This may possibly prove true of some of the larger forms described by Stimpson, allied to Pisaster ochraceus and P. capitatus, but on the other hand there are probably many additional species to be found by dredging in moderate depths. Nearly all the known species of this region are littoral or very shallow-water forms. Deep-water species are not described in this report. I. ANALYTICAL TABLE OF THE GENERA, SUBGENERA, SPECIES, SUBSPECIES AND VARIETIES OF ASTERIIDJE FROM THE NORTHWEST COAST OF AMERICA. 1. Dorsal skeleton continuous, formed mostly of lobed plates and smaller ossicles overlapping at the ends; dorsal plates not isolated. Rays five or six, rarely more, unless autotomous. 2. Adambulacral spines all attached to the actinal face of the plates; none attached higher within the grooves. 3. Adambulacral spines in a single regular row, one to a plate, in the adult (monacanthid). A. Interactinal plates present, at least in adults, and usually bearing spines. C. Disk thick, rather large ; rays five or six, stout, thick at base. Dorsal os- sicles strong, numerous, not confined to three or five rows. Dorsal skeleton reticulate or irregular. Dorsal and lateral dermal major pedicellariae sessile, large, stout, erect, ovoid or wedge-shaped, often nearly as thick as the spines, and usually strongly denticulated at the tips of the valves (unguiculate). Genus Pisaster M. and Tr., p. 67. Type, P. ochraceus (Br.). *The only other region that can be compared with this in the number and variety of Asteriinae is the southern coast of South America, especially around Tierra del Fuego and Patagonia, whence numerous genera and species have been described, mostly very unlike the northern forms. (See below, p. 351.) SHALLOW-WATER STARFISHES 57 D. Interactinal ossicles numerous, stout, closely united in three to six longi- tudinal rows, usually bearing three to six or more close rows of strong actinal spines. E. Dorsal spines arranged in a reticulate or areolate manner, or else grouped on the nodes ; but often forming a median row on the rays. Rays normally five, stout. a. Dorsal spines stout, coarse, very unequal, capitate, acervate, forming con- spicuous angular reticulations with or without notable nodular groups and often with median radial rows. P. ochraceus (Brandt), p. 68. Calif, to Middle Alaska, a'. Dorsal spines mostly in very prominent nodes or clusters ; reticulations less conspicuous. Var. nodiferus V., nov., p. 71. Calif, to Alaska. aa. Dorsal spines small, or rather small, acute or obtuse, not strongly capitate, usually forming evident reticulations. b. Dorsal spines exceedingly numerous, very small, obtuse or acute, forming conspicuous reticulations, with or without radial rows and a cen- tral pentagon. Actinal spines stout, obtuse, but not strongly grooved externally. P. confertus (Stimp.), p. 73. Calif, to Vancouver I. bb. Dorsal spines small, slender, not very numerous, forming an openly reticu- lated pattern with a central pentagon and distinct median radial rows. Actinal spines deeply gouge-shaped and often bifid. Disk large. P. fissipinus (Stimp.), p. 76. Oregon. EE. Dorsal spines scattered, mostly without regular order, not reticulated, but sometimes forming radial rows. Rays usually stout, five or six. d. Rays normally five. Dorsal spines large, with capitate or ovoid tips, not very numerous. e. Dorsal spines few, large, nearly equal, obtuse or strongly capitate and radially striate. Rays short. P. capitatus (Stimp.), p. 81. S. California. ee. Dorsal spines more numerous, unequal, the tips ovoid or subconic and striated, usually forming imperfect radial rows. Rays long, rounded. Size large. P. liitkenii (Stimp.), p. 83. S. Calif, to Vancouver I. Var. australis Ver., p. 88. dd. Rays normally six; dorsal spines very numerous, short, stout, conical, acute or obtuse, striated, nearly uniformly scattered ; no evident ra- dial rows. Interactinal spines in four to six rows. Disk large. Size large. P. giganteus (Stimp.), p. 89. DD. Interactinal ossicles fewer, usually forming only two rows, and mostly bearing simple rows of spines. Dorsal ossicles openly reticulated. Dorsal spines few. f. Rays normally five. g. Dorsal spines rather sparse, short, thick, subconic, striated, scattered or clustered, with distinct median radial rows. Actinal spines short, obtuse, often flat, or slightly gouge-shaped at tip. P. brevispinus (Stimp.), p. 77. S. Calif, to Vancouver I. 58 VERRILL gg. Rays thick and high at base, rapidly tapered. Dorsal spines few, stout, blunt, or conical, mostly isolated, but forming imperfect median radial rows and a regular pentagon on the disk. Superomarginal spines one to a plate. Inferomarginal row double. Large dermal pedicellariae numerous, stout, stone-hammer-shaped, with serrate jaws. P. papulosus Ver., p. 91. Puget Sd., Brit. Columbia. ff. Rays normally six, rarely five; more slender. Dorsal spines short, rather numerous, obtuse or capitate, irregularly arranged, often in groups of two or three. Interactinal spines in two or three rows. Major pedicellarise acute-triangular. P. grayi Ver. = katherinse Per. (non Gray), p. 97. Oregon. CC. Rays five; angular, more slender. Dorsal spines in three or five regular rows. Dorsal ossicles stout in longitudinal and transverse rows. h. One or two rows of interactinal plates and spines. Major pedicellarise large, ovate; minor pedicellariae not very abundant. Superomar- ginal spines one to a plate. P. paucispinus (Stimp.), p. 98. Calif, to Puget Sd. AA. Dorsal spines in three regular rows, bearing dense subapical clusters of minor pedicellariae. Superomarginal spines only on alternate plates. Ventral spines in two or three rows. Interactinal plates feebly developed. Marthasterias sertulifera (Xantus), p. 100. Gulf of Calif, to S. Diego. 3'. Adambulacral spines two to a plate, arranged in two rows (diplacanthid) ; or else in irregular rows, when some of the plates have one and some two spines (subdiplacanthid). B. At least one row of visible interactinal plates generally present in adults, often spineless; sometimes several close rows. F. Dorsolateral plates rather small, narrow, strongly lobed, usually reticulated or irregularly arranged, and united by their own lobes and by smaller transverse ossicles. G. Interactinal plates usually form one long row (peractinal), much like the lower marginals, and one or two short proximal rows (subactinals), but the latter may be mostly lacking or rudimentary, especially in the young, so that there may be only two simple rows of ventral spines ; or the lower marginals and peractinal rows may be doubled proximally, two or more spines standing on each plate; thus the number of actual rows of these spines may sometimes be four or more. Dorsal ossicles variously arranged, most often openly reticu- late. Major pedicellarise usually small, ovate or lanceolate, some- times unguiculate. Rays normally five or six. H. Dorsal papular areas usually large; papulae small and numerous, in groups of twelve to fifty or more, in the adult. Size generally rather large. Larva of many, if not all, species is a free-swimming brachiolaria. Genital pores dorsal. Genus Asterias (L.), closely restricted, p. 101. i. Rays normally five, ossicles and spines reticulate in arrangement. SHALLOW-WATER STARFISHES 59 j. Rays five, rather long, rounded. Primary dorsal spines few, short, stout (2 to 3 mm. thick), capitate, forming a median row, with few irregularly scattered ones of similar size and form. Two marginal and two interactinal, regular, mostly simple, rows of short, stout, blunt spines; adambulacrals subdiplacanthid. A. victoriana Ver., p. 102. Vancouver I. jj. Rays long, slender, acute; dorsal spines of moderate length, slender, fluted, mostly one to a plate, arranged in quincunx, or in about seven obscure alternating rows; median row with longer spines, distinct; both marginal rows single; peractinal row double, with spines longer than marginals; major pedicellariae adambulacral, rather large, lanceolate. A. namimensis Ver., p. 105. Brit. Columbia. ii. Rays normally six. k. Dorsal spines very diverse in form and size; primary ones very large, ir- regularly placed, but not acervate, very stout (2 to 4 mm.), with nipple-shaped, sulcated tips; smaller spines numerous, unequal, capitate or round-topped, with abundant minor pedicellariae. Mar- ginal and interactinal spines smaller, in about four close rows proximally; adambulacrals mostly two to a plate; the inner ones and furrow margins bear abundant clusters of small major pedicel- lariae. A. polythela Ver., p. 104. Arctic O. kk. Primary dorsal spines not unusually large; not nipple-shaped. I. Primary dorsal spines more or less unequal, capitate; the larger usually acervate or irregularly grouped. Pedicellariae small, numerous; major ones ovate, forming clusters on the adambulacral spines, a few also on the interactinal areas. One complete row of peractinal spines, separated from the adambulacrals by a naked zone contain- ing papular areas. Subactinal spines form a short series, or may be lacking. A. acervata Stimp., p. 107. Bering Sea; Arctic O. II. Principal dorsal spines not very unequal nor distinctly acervate. m. Dorsal spines rather small, irregularly reticulated; adambulacrals sub- diplacanthid. A. katherina Gray (non Per.), p. 112. Oregon; Gulf of Georgia. mm. Dorsal spines very numerous, in distinst radial rows, not very diverse in size, capitate, several on a plate ; the median row crowded and more or less clustered ; both marginal rows double ; two interactinal rows. A. multiclava Ver., p. 114. Siberia. HH. Papular areas small; papulae few, large, mostly standing singly or in small groups of one to five, rarely ten or more. Dorsal plates usually lobed and united by overlapping, sometimes in radial rows. Adambulacral spines alternately one or two to a plate, or else biserial. Minor pedicellariae as in Asterias; large, serrate, dermal ones may occur. Eggs and young usually carried in clusters around the mouth of the mother; genital pores ventral. Size small. Rays normally five or six. Genus Leptasterias Ver., in part, p. 116. (See below, p. 60.) 6O VERRILL I. Rays normally five, sometimes varying individually to six. n. Disk somewhat enlarged, thickened or depressed; rays not very slender; spines unequal and uneven; superomarginal row mostly single; inferomarginal row double. o. Disk and rays depressed ; dorsal spines very unequal ; larger ones capitate, forming a prominent median radial row and irregularly linear groups ; ventral spines all rather slender. L. inequalis Ver., sp. nov., p. 117. Orca, Alaska. oo. Disk thickened or plump ; rays strongly convex ; dorsal spines numerous, capitate or clavate, usually areolated, or arranged partly in clusters, often forming a median row ; marginal and peractinal spines thick- ened. L. epichlora (Br.), p. 132 (five-rayed form). Aleutian Is. to Puget Sd. nn. Disk smaller, rays more slender or terete. Dorsal spines subequal, clus- tered, scattered, or somewhat in longitudinal rows. p. Dorsal spines not remarkably small ; not in transverse lateral combs. At least one row of interactinal spines proximally. q. Rays very slender, terete; dorsal spines small, capitate, or clavate, numer- ous, mostly one or two to a plate, not in evident rows. Ventral spines slender, both marginal and peractinal rows single; major pedicellariae large, with long spatulate blades. L. leptalea Ver., sp. nov., p. 119. Virgin Bay, Alaska, qq. Rays less slender. Dorsal spines small, clavate, in about three longitudi- nal rows, not crowded; both marginal rows regular and distinct, the spines longer and standing singly; interactinal row imperfect. L. arctica (Murdoch), p. 120. Arctic O. ; Bering Sea. pp. Dorsal spines very slender, numerous irregularly arranged ; superomarginal and inferomarginal spines longer, mostly one to a plate, in regu- lar rows. L. epichlora miliaris Ver., nov., p. 138 (five-rayed form). British Columbia. ppp. Dorsal and marginal spines very numerous, small, subequal, arranged in close clusters, and largely concealed by the large dense wreaths of minor pedicellariae ; marginal plates of both series have two or three small spines, nearly concealed by pedicellariae. Leptasterias obtecta Ver., sp. nov., p. 144. Bering Sea. nnn. Dorsal and dorso-marginal spines very small, numerous, nearly equal, the latter arranged in transverse lines on the narrow ossicles defin- ing large transverse papular areas. Minor pedicellarise small, around bases of spines, not unusually abundant. Ctenasterias cribraria (Stimp.), p. 148. Circumpolar; Bering Sea; Arctic O. ; North Atlantic. II. Rays normally six, sometimes five or seven ; not autotomous. r. Disk small ; rays slender, not enlarged at base ; papulae few, single or in small groups. Ambulacral pores of the usual size and form, more or less crowded, in four rows. Genus Leptasterias Ver., in part, p. 116. (See above, p. 59.) t. Dorsal spines small, slender, clavate, not crowded nor clustered, pretty regularly arranged in quincunx, surrounded by thick wreaths of SHALLOW-WATER STARFISHES 6l pedicellariae; marginal spines longer; rows mostly simple; no peractinal spines. L. coei Ver., sp. nov., p. 123. Off Berg Bay, Alaska. tt. Dorsal spines longer, scattered; one or two to a plate, except in evident median radial row. Inferomarginal row double, two to a plate, longer than dorsals, subacute ; superomarginal row simple ; three interactinal rows proximally, one spine to a plate. L. macouni Ver., sp. nov., p. 124. Vancouver I. ttt. Dorsal spines mostly clustered, several on each plate, u. Rays six or seven, elongated. Dorsal spines small, obtuse, arranged in five longitudinal ranges of groups, each group with three or four spines ; others are on the transverse ossicles, so that they form " a vague reticulation." Ventral spines in two rows, pointed, longer than adambulacrals. L. vancouveri (Per.), p. 125. Vancouver I. uu. Dorsal spines small and numerous, all of about the same height, usually clavate, obtuse, standing in crowded rows. Marginal and inter- actinal spines notably longer, forming three to five or more close rows. v. Dorsal spines small, slender, subequal, mostly clavate, not much crowded, standing singly or in small groups on the ossicles, forming median radial rows, and sometimes subradial series. Major pedicellariae few, rather large, long, ovate. L. hcxactis (St.), p. 126. Calif, to Brit. Columbia. vv. Dorsal spines mostly capitate, very small and very numerous, standing in large groups on each plate, about equal in height, and thus pre- senting a nearly even surface. Papular pores in nearly regular radial rows. Marginal and interactinal spines much longer, often bent, numerous, arranged in regular rows. Peroral spines strong. L. aqualis (St.), p. 128. Calif, to Puget Sd. w. Dorsal spines very short, equal, clavate or capitate, crowded, many on each plate. Marginal and interactinal spines form about four rows. Rays short, obtuse. Var. nana Ver., nov., p. 132. California; Oregon; British Columbia, ww. Dorsal spines very numerous and crowded on the plates, distinctly capitate, with rounded tips. Marginal and interactinal spines more numerous, forming five to eight rows, several on each plate. Var. compacta Ver., nov., p. 130. California. www. Dorsal spines very numerous, crowded, the clusters arranged in three or five radial bands separated by rows of very distinct papular areas. Var. concinna Ver., nov., p. 132. California. rr. Rays normally six, sometimes five, of moderate length, thick or wide at base, not slender, rather obtuse; disk somewhat wide or swollen, skeleton firm; principal dorsal ossicles rather large. x. Dorsal ossicles not very closely united, irregularly reticulated, covered with numerous short, mostly capitate or clavate spines, usually reticulate in arrangement, often showing a more or less evident median row or band ; three rows of interactinal spines often present proximally in adult, only one in small specimens. Adambulacral spines irregu- 62 VERRILL larly diplacanthid, crowded; sucker-feet numerous, crowded in four rows. Large, denticulate, dermal major pedicellarise often present in adult. Eggs and young carried by mother. Papulae often in dorsal clusters of five to twelve or more. L. epichlora (Br.), p. 132. Calif, to Aleutian Is. a. Rays commonly five; dorsal spines very numerous, very unequal, strongly capitate, acervate. Median row not very evident Var. subnodulosa Ver., nov., p. 139. Wrangel, Alaska. aa. Rays normally six. b. Superomarginal spines stand two or three, or sometimes more, to a plate. c. Dorsal spines quite unequal in size, mostly capitate; arrangement mostly areolated or reticulated. d. Dorsal spines do not form a very evident median row, and are not notably acervate; reticulate arrangement conspicuous; larger ones capitate. Large denticulate dermal pedicellariae often present, as well as the usual form ; minor sort abundant. Subspecies alaskensis Ver., nov., p. 136. Puget Sd. to Aleutian Is. dd. Dorsal spines form an evident median row or band of larger spines. e. Dorsal spines numerous, mostly clavate, not crowded, reticulate, with the median radial row very evident. Var. carinella Ver., nov., p. 137. Alaska. ee. Median radial dorsal spines decidedly larger, capitate, others small, crowded, reticulate and areolate. Var. siderea Ver., nov., p. 137. Puget Sd. to Aleutian Is. eee. Sometimes five-rayed. Dorsal spines crowded, very small, slender, cla- vate or slightly capitate; several small ones around each larger; median ones slightly larger; superomarginals unequal, one larger and several smaller on a plate. Subspecies miliaris Ver., nov., p. 138. Alaska. cc. Dorsal spines, mostly equal or subequal, short, capitate, numerous, crowded, areolate, often forming circles around the papular areas. f. Dorsal spines are in five or more evident radial bands separated by rows of large papular areas, several spines standing on each of the larger ossicles, median band evident, but not prominent; three to four or more spines on each superomarginal plate ; two on inferomarginals, stouter; one on peractinal, stout, clavate. Papular areas large. No large pedicellarise on type. Subspecies plena Ver., nov., p. 140. British Columbia. 8. Dorsal spines very even, stout, capitate, crowdedly areolate, but not forming evident radial rows ; median row irregular or obscure ; papular areas conspicuous, but not in evident radial rows; mostly three spines on superomarginal plates; two on inferomarginals, all similar and near together, clavate, larger and longer than dorsals; peractinals similar. Large, erect, serrate major pedicellarise common between marginal rows of spines. Subspecies pugetana Ver., nov., p. 142. Puget Sd. bb. Superomarginal spines mostly stand singly on the plates. A distinct dorsal median row. SHALLOW-WATER STARFISHES 63 g. Dorsal spines small, subequal, slender, clavate. Van regularis Ver., nov., p. 139. Alaska. gg. Dorsal spines short, mammilliform, arranged somewhat in rows. Var. subregularis Ver., nov., p. 139. Alaska. xx. Dorsal and marginal ossicles stout, closely united, overlapping; median dorsal radial row prominent, bearing the larger capitate spines in a conspicuous irregular row; other spines of the larger sort form an irregular radial row on each side; inferomarginal spines longer and larger, two or three to a plate; single row of peractinals simi- lar. Disk is rather wide, depressed. L. dispar Ver., nov., p. 142. Aleutian Is. GG. Interactinal and submarginal plates rather stout, lobate, closely joined in three to five regular longitudinal rows; each plate may bear one, two or more spines, forming, in the adult, four to eight or more rows of uniform or similar ventral spines. Rays five, rounded, usually long and tapered. Dorsal ossicles mostly reticulate. Papu- lar areas large. Unguiculate dorsal pedicellarix lacking. Genus Evasterias Ver., nov., p. 151. i. Rays five, gradually tapered. Dorsal spines numerous, unequal, areolate or reticulate, often clustered; larger ones short, capitate or trun- cate; smaller ones acute, capitate or clavate. Marginal spines in regular rows ; longer than the dorsals ; interactinal spines still longer, forming four to six or more regular close rows, curving strongly upward proximally; adoral spines elongated. E. troschelii (St.), p. 151. Calif, to Alaska. j. Dorsal spines acervate, or forming very evident nodular groups with larger central capitate spines, and often, also, a median row. Var. subnodosa Ver., nov., p. 163. Puget Sd. to Alaska, jj. Dorsal spines not acervate. k. Dorsal spines smaller, very numerous, mostly clavate or subacute, few capitate, rough, openly reticulate; papular areas large; numerous crowded interactinal spines. Size large. Var. rudis Ver., nov., p. 158. Vancouver I., etc. kk. Dorsal spines nearly all capitate, not very unequal. I. Dorsal spines nearly uniform, capitate, forming close reticulations, the spines in single lines on the ossicles. Var. densa Ver., nov., p. 161. Vancouver I. II. Rays of moderate length, dorsal spines all capitate, reticulations coarse; both marginal rows of spines simple or nearly so, spines not crowded ; two or three actinal rows, not crowded. Var. alveolata Ver., nov., p. 162. Vancouver I. to Alaska, kkk. Dorsal spines not very unequal, nearly all slender, clavate or subclavate, areolate, not acervate; marginals larger, subcapitate. Pedicellarise numerous. Var. parvispina Ver., nov., p. 163. Alaska. ii. Rays five, long and large, acute at tips. Dorsal spines very numerous, small, unequal, the larger obtuse. The smaller ones are mostly acute and arranged in clusters and short transverse lines or combs, producing an imperfectly reticulate pattern, without distinct radial 64 VERRILL rows. Interactinal and submarginal spines form five or six, mostly double, crowded rows, with about eight to ten in each transverse range, in the adult Proximal adambulacral spines much elongated. Size large. E. acanthostoma Ver., nov., p. 165. Alaska. FF. The dorso-lateral spines, or groups of spinules, and more or less of the plates, have an evident arrangement in longitudinal rows. Inter- actinal plates present or rarely absent; seldom more than one row. J. Dorsal plates strong, lobate, reticulated so as to form rather obvious longitudinal rows ; papular areas usually large, often nearly rec- tangular. Each of the principal plates usually bears a single rather large spine, rarely two, forming three to five or more dorsal rows of spines; smaller spines may occur on the intermediate transverse ossicles. Marginal plates stout, bearing regular rows of spines, much like the dorsals. A row of similar peractinal plates, usually bearing a simple row of spines, often spineless, rarely lacking ; some- times a short row of subactinals proximally. Adambulacrals usually diplacanthid, sometimes subdiplacanthid. Major pedicellariae large ; usually of two or more kinds ; the larger dermal ones denticulate, or unguiculate. Rays five or six. Genus Orthasterias Ver., nov., p. 168. Type, O. columbiana Ver., sp. nov. K. A row of interactinal plates is present, with or without spines ; they may be small and not visible unless dermis be removed. L. One row of peractinal spines, at least proximally; inferomarginals with two rows, making three ventral rows altogether. a. Rays normally five, rarely six, elongated. b. Dorsal spines in at least five rows, besides some intermediate scattered ones on the connective ossicles ; papular areas rather small ; dorsal radial plates rather large and thick, firmly united. c. Five dorsal radial rows of spines obvious. d. Apical oral spines not notably enlarged. Dorsal spines obtuse and fluted. Wreaths of minor pedicellariae on thick dermal-sheaths, often above the middle of the spine. e. Adambulacral spines flattened; adoral spines longer, not very slender; dorsal spines obtuse or truncate, rough or fluted. O. columbiana Ver., sp. nov., p. 168. Vancouver I. to Yakutat, Alaska. ee. Adambulacral spines all very slender and subequal ; regularly diplacan- thid, strongly and evenly divergent, thus forming two remarkably regular pectinate rows, those of the inner row usually horizontal. O. biordinata Ver., nov., p. 173. British Columbia. dd. Apical oral spines notably stronger than others. Dorsal spines tapered, not fluted. Minor pedicellariae are notably minute, forming dense basal circumspinal wreaths. Major pedicellariae are very large, stout, often as thick as spines, wedge-shaped, erect, often dentic- ulate. O. californica Ver., nov., p. 174. California. SHALLOW-WATER STARFISHES 65 cc. Longitudinal rows of spines more than five, not very obvious, due to crowding ; spines appear as if in quincunx. f. Dorsal spines long, slender, slightly grooved. Adambulacral spines slender, orals longer. Major pedicellariae large, lanceolate, acute, dentate at tip. O. kcehleri Lor., p. 175. Vancouver I. bb. Dorsal spines in three rows, long, not fluted; ossicles openly reticulated, leaving large papular areas ; spines of two inferomarginal rows and peractinal row slender, elongated, tapered; adambulacral spines rather slender, tapered or slightly flattened. Major pedicellariae large. O. dawsoni Ver., nov., p. 175. Brit. Columbia. aa. Rays six, rounded and elongated. Dorsal spines large, in three or five rows, with smaller ones interpolated, all with a thick fleshy sheath (in alcohol) bearing a large wreath of minor pedicellariae at about mid-height or above. Both marginal rows of spines are regular and mostly simple. A regular peractinal row of spines and in mature specimens a short subactinal row proximally. O. merriami Ver., nov., p. 177. Alaska. LL. Peractinal spines generally lacking; the ossicles are rudimentary or small; inferomarginals with two rows of spines. Subgenus Stylasterias Ver., nov., p. 179. g. Dorsal spines mostly in three longitudinal rows, often with some scattered intermediate ones. h. Minor pedicellariae remarkably large and unguiculate, about half as long as major ones, forming large, loose, basal circumspinal wreaths. Larger major pedicellarise often nearly as stout as the spines. i. Spines long, inferomarginals in two rows and, like the outer adambulacral spines, often notably flattened and spatulate or gouge-shaped. Outer adambulacral spines are the larger; papular areas large; peractinal plates very small. O. forreri (Lor.), p. 179. California. ii. Adambulacral plates and spines crowded ; spines slender. Minor pedicel- lariae remarkably large and abundant; dorsal ones in large groups on the dermis and around bases of spines; dorso-lateral ossicles rather slender, openly reticulated; papular areas large; major pedi- cellariae very stout. Subspecies forcipfilata Ver., p. 180. British Columbia. hh. Minor pedicellariae not notably large; they form large, dense, basal circum- spinal wreaths; inferomarginal and adambulacral spines flattened, not notably spatulate nor strongly gouge-shaped. Dorsal spines well spaced, elongated, tapered, not fluted; alternate superomarginal plates mostly spineless. O. gonolena Ver., nov., p. 184. S. California to Gulf of Calif. gg. Dorsal spines in more than three rows ; rays slender, terete. Dorsal spines in about five rows, long, nearly uniform, tapered; papular areas large ; rays slender ; inferomarginals tapered, obtuse ; adambulacrals small, slender, outer ones a little larger and longer, slightly clavate. Major pedicellariae large, denticulate. ^ • 66 VERRILL O. leptolena Ver., nov., p. 182. British Columbia. K. No visible interactinal plates or spines. (See below, p. 66.) Genus Distolasterias Per., in part, p. 185. Dorsal spines numerous, in about seven to nine irregular alter- nating rows; rather short, fluted, blunt or clavate. Larger major pedicellarias partly stout, wedge-shaped ; partly elongated, spatulate, unguiculate, with few large curved teeth. Adambulacral spines slender, crowded, regularly diplacanthid. D. chelifera Ver., nov., p. 185. Vancouver I. JJ. Dorso-lateral plates mostly wide, lobed, overlapping or imbricated in few close rows (stichasterial arrangement), covered with clusters of minute spinules. A distinct peractinal row of plates. Disk small, rays slender, six in type. Ambulacral pores unusually large, sub- triangular or cordate, not crowded. Dorsal spines very small, short, scattered, obtuse or clavate, a distinct median radial series. Stenasterias macropora Ver., p. 145. Alaska. BB. Interactinal plates lacking or rudimentary; no interactinal spines. M. Dorsal skeleton weak, composed mostly of slender, openly reticulated ossicles, some of which do not unite in a continuous skeleton. Dor- sal spines few, rather large, in few open rows or scattered ; upper marginal spines regular and well separated from lower marginals; the latter with large and long spines ; alternate plates without spines. Minor pedicellariae large and very numerous, in sacculated wreaths or clusters. Size large. Genus Urasterias Ver., p. 187. Type, U. linckii (M. and T.). Arctic; North Atlantic. b. Several open rows of long dorsal spines ; adambulacral plates not crowded ; lower marginal spines large and well spaced; minor pedicellariae not remarkably large; bunched in large wreaths on the large, loose, saccular sheaths of the spines. Size large. U. linckii (M. and T.), p. 187 (extralimital?). N. Atlantic and Arctic. MM. Dorsal skeleton well developed. c. Dorso-lateral plates and ossicles openly reticulated. Rays regularly five, rather short, stout, depressed; dorsal spines numerous, small, slender, obtuse, arranged on small, slender, reticulated ossicles; no distinct median row; marginal spines usually two to a plate in both rows, in form like the dorsals, but rather longer; adambulacrals regularly diplacanthid. No interactinals visible. Genus Parasterias nov., p. 187. Type, P. aibertensis Ver. Brit Columbia. cc. Dorso-lateral spines and plates in more or less obvious longitudinal or transverse rows ; spines either solitary or clustered. d. Rays regularly five, large and long; dorsal plates bear large and mostly solitary spines and large pedicellariae. Genus Distolasterias Per., in part, p. 185. (See above, p. 65.) dd. Rays slender, variable in number, due to autotomy; disk small; autoto- mous ; finally six or five. Median and marginal plates little wider, imbricated in longitudinal rows; dorso-lateral ossicles and plates short, irregularly reticulated, but leaving regular rows of papular SHALLOW-WATER STARFISHES 67 areas. All the plates are covered with close clusters of small, nearly equal spinules, longer beneath. Pedicellariae of two sorts; dermal, not abundant ; few on spines. Stephanasterias albula (St.), p. 147. Circumpolar; North Atlantic; Bering Sea. ? The inner adambulacral spine, on alternate plates, is attached to a lobe of the inner edge of the plate, higher up within the groove. Dorsal spines small and numerous, scattered. Interactinal plates feebly developed or lacking. Allasterias Ver., p. 188. Type, A. rathbuni Ver. Bering Sea. f. Dorsal ossicles small, forming reticulations ; adambulacral spines alternately two and three to a plate. Superomarginal spines clustered. Dermal major pedicellarise large and numerous, pointed. Rays five, wide, depressed ; margin prominent. A. rathbuni Ver., p. 189. Bering Sea. See also, A. forficulosa V., nov., p. 194. Japan. g. Inferomarginals not very stout, flattened, blunt, not deeply channeled. Subspecies nortonensis Ver., p. 191. Bering Sea. gg. Inferomarginal and actinal spines stout, channeled or gouge-shaped. A. anomala Ver., p. 193. Siberia. 1'. Dorsal skeleton incomplete or rudimentary; ossicles partly isolated; dorsal surface covered by soft skin. Disk large; rays multiple, variable in number, increasing with age by interpolation in pairs up to twenty to twenty- four in the adult. Adambulacral spines monacanthid. Marginal plates distinct. Dorsal minor pedicellariae in large clus- ters, dermal and epispinal. Genus Pycnopodia Stimp., p. 197. Type, P. helianthoides (Br.), p. 198. Calif, to Aleutian Islands. ASTERIIN.E: DESCRIPTIONS OF GENERA AND SPECIES. Genus Pisaster A. Agassiz, ex Miiller and Troschel. Pisaster MULLER and TROSCHEL, Syst. Ast, p. 20, note, 1842. A. Agassiz, North American Starfishes, p. 96, 1877 (type, ochraceus) . Cosmasterias (pars') SLADEN, Voy. Challenger, xxx, p. 562, 1876. Large starfishes with a thick and usually broad disk and either five or six stout tapering rays. Adambulacral spines form a single regular row, one to each plate (monacanthid). Three to six rows of closely united or tesselated, thick actinal and lower marginal plates, bearing several close rows of stout ventral spines. The syn- actinal plates are closely united to the adambulacrals. Oral region usually deeply sunken. Interbrachial septa well developed. Special, very large, stout, erect, sessile, forficulate pedicellariae, with interlocking serrations at the ends of the valves (unguiculate), stand on the lateral channels, and often between the actinal and dorsal spines ; larger ones often stand in the interradial areas below. Dorsal skeleton variable, but well developed, the ossicles generally 68 VERRILL stout and openly reticulated. Dorsal spines few or many, and variously arranged, often in reticulate or acervate patterns, but sometimes scattered or in irregular rows. One of the important structural features, characteristic of most of the species, is the increased number of rows of actinal plates and spines, and their close imbricated or tesselated arrangement; but in certain species (P. papulosus') , otherwise related to the type, there are usually only two rows of actinal plates that bear spines below the inferomarginal plates, as in typical Asterias; very large specimens may have more rows. But there is another character, perhaps, of more value, common to all the species from the Northwestern American Coast related to P. liitkenii and P. ochraceus, which seems to warrant the separation of this group, as a distinct genus. This is the existence of the peculiar form of large, stout, erect, sessile, dermal pedicellarise, ovoid or wedge-shaped in form, with broad, unguiculate or denticu- lated, interlocking jaws. These occur especially along the naked spaces between the ventral and lateral spines, but they are also frequently found scattered between the dorsal spines, or between the bases of the actinal spines. They are often equal to the adjacent spines in diameter, but are much shorter. More slender, lanceolate or ovate, major pedicellariae, of the ordinary type, always coexist with these large forms, while the latter are not present on starfishes of the Asterias rubens type, nor on those belonging to most of the other divisions.1 It is probable, therefore, that this group had its origin upon the Pacific coast of North America, where it has subsequently had time enough to develop into many diverse species, some of which may have migrated to other regions and left descendants. This, then, is one of the most characteristic groups of this fauna, but it is not so strikingly peculiar as Pycnopodia and Dermasterias. Mr. A. Agassiz (1877) referred to the name Pisaster as having been proposed by Professor L. Agassiz, for the group of species allied to A. ochracea, but he did not consider the group as more than a subdivision of Asterias, though he fully described the characteristic structures in the skeleton of P. ochraceus, as contrasted with a typical Asterias. Miiller and Troschel had, however, proposed this 1 A few similar large pedicellarise often occur on Leptastcrias cpichlora (Br.), but not on the other allied species. They are often wanting, and may be due to hybridizing with P. ochraceus, associated with it. Similar large ones are found on Orthasterias columbiana and allied species. SHALLOW-WATER STARFISHES 69 name, for the same type, in 1842, but did not subsequently use it. This name should, therefore, take precedence of all others, whether it be considered a genus or a subgenus. The mode of reproduction is unknown. PISASTER OCHRACEUS (Brandt) A. Agassiz. Plate xxi, figures I, 2; plate XLIX, figures 3-3^ (pedicellarias) ; plate LVI, figures 3, 30 (variety). Aster tas ochracea BRANDT, Prod. Descr. Anim. Mertens, p. 69, 1835. Asterias janthina BRANDT, op. cit., p. 69, 1835 (descr. insufficient; color var. ?). Asterias ochracea STIMPSON, Journ. Boston Soc. Nat. Hist., vi, p. 87, pi. xxm, fig. 2, 1867. Verrill, Trans. Conn. Acad. Sci., i, pp. 325, 326, 1867. Whit- eaves, Trans. Royal Soc. Canada, iv, p. 116, 1887. A. Agassiz, North Amer. Starfishes, Mem. Mus. Comp. Zool., v, p. 96, pi. xi, figs. 1-7 (structure), 1877 (Pisaster). H. L. Clark, op. cit., 1907, p. 67, pi. vi, fig. 3 (structure). Loeb, Publ. Univ. Calif., Physiol., u, pp. 5-30, 1904 (hybridization). Asteracanthion margaritifer MULLER and TROSCHEL, Syst., p. 20, 1842 (t. Bell). Disk rather large, strongly reticulated by the prominent ossicles and spines ; rays normally five, sometimes six, stout, rapidly tapered. Two Alaskan specimens, differing considerably in form, measure as follows: (a) lesser radius, 36 mm.; greater radius, 120 mm.; ratio, 1:3.33; (b) lesser radius, 32 mm.; greater radius, 118 mm.; ratio, i : 3.68. The dorsal spines are numerous, unequal in size, but nearly all are capitate; the larger ones are short and stout, strongly capitate with flattened striated tops. They are arranged in short rows form- ing a conspicuously reticulated pattern, with large, deep, angular papular areas between them. They usually stand so closely in the rows that they nearly touch; on the disk they form a conspicuous pentagon, enclosing subordinate angular groups or partial reticula- tions and a central cluster. A zigzag, irregular median row is often distinct on the basal part of the rays. The smaller spines are similar in form but not so stout ; they are partly grouped around the larger ones, and partly serve to subdivide the primary reticulations. Each primary papular area usually contains several large clusters of papulae. Large, short, very stout, obtuse, wedge-shaped, unguiculate, dermal major pedicellariae are more or less numerous on the dorsal surface, and still more so on the sides of the rays; they are rather less in diameter than the adjacent spines. Those in the lateral chan- nels and interradial spaces are larger and stouter than the dorsal ones. Minor pedicellariae form wreaths around the bases of the 7O VERRILL dorsal and lateral spines and also occur in clusters on the papular areas. The interactinal spines form four or five crowded rows ; they are large, stout, mostly clavate, with obtuse, smaller, often flattened, striated or sulcated tips; those of the synactinal (inner) row are less stout, but not shorter; those of the interradial areas are longer and more fusiform. Toward the mouth the adambulacral rows of ossicles are prolonged as a narrow ridge to the sunken mouth, unac- companied by actinal spines. The actinal spines bear clusters of minute minor pedicellariae. The interactinal ossicles, as exposed in a denuded specimen, are small and closely arranged in about five rows at the base of the arm, with some additional ones irregularly inter- polated. Large, pedicelled, lanceolate major pedicellariae are attached within the margins of the furrows. The marginal spines are capi- tate and striated like the dorsal spines. VARIATIONS. In the Museum of Comparative Zoology I have seen two six- rayed specimens (No. 1423) from Crescent City, Calif. These have the dorsal spines unusually numerous, and strongly reticulated or areolated. Two specimens sent to me by Mr. J. F. Whiteaves, of the Canadian Geological Survey, from Vancouver Island, are remarkable for their great size. They are 20 to 22 inches in diameter. Radii of one are 75 mm. and 250 to 275 mm. ; ratios about 1 : 3.75. Radii of the other are 70 mm. and 240 to 260 mm. ; ratios about i : 3.6. The disk is abnormally flattened in both, by imperfect preservation. They agree with ordinary specimens in all essential features, except such as are naturally due to greater size. The dorsal spines are unequal and strongly clustered, but have lost more or less of their reticulated arrangement, except distally, owing to the breaking up of many of the lines of spines and the interpolation of new clusters. The spines and pedicellariae are of the ordinary forms. The large, serrate pedicellariae are very abundant on the dorsal, lateral, and actinal surfaces. The largest are on the actinal inter- radial areas, where some equal the spines in diameter. Mr. Whiteaves also sent a short-rayed specimen, from the Queen Charlotte Islands, which has the dorsal spines larger, more capitate, and more numerous than usual, so that they form very prominent nodular clusters and imperfect reticulations. Its large unguiculate pedicellariae are unusually abundant. (Var. nodiferus.) SHALLOW- WATER STARFISHES 71 In the same collection was a large six-rayed specimen from Dis- covery Sound, 1885, which differs in no other respect from the ordinary five-rayed specimen. A similar one, in our collection, was collected near Monterey, Calif., by Dr. W. R. Coe. The smallest specimen that I have seen has the radii 14 mm. and 41 mm. ; ratio about i : 2.8. The disk is elevated. The dorsal sur- face is thickly covered with characteristic nodular groups and imper- fect reticulations of capitate spines, like those of larger specimens, but smaller. On the disk they form a distinct stellate pentagon and a central cluster. The upper marginal row is simple. The inferomarginal row is double. There are also two simple rows of actinal spines ; all these forming four very regular, close-set rows of nearly equal obtuse ventral spines. On the lateral channels, proximally, there are a few of the large, erect, wedge-shaped pedicellarise, characteristic of the large specimens. Its color is variable, and often very showy; most frequently it is dark or light orange, varying to yellow, to chocolate-brown, and to violet; spines are paler. This is the most common of the species of Pisaster, from Mon- terey and San Francisco to Sitka. In most places it is by far the most abundant large starfish of the shores. It occurs at low tide among rocks and in shallow water. It was collected by the Harriman Expedition at Sitka, Virgin Bay, Wrangel, and other localities in southern Alaska. Mr. J. F. Whiteaves, of the Canadian Geological Survey, formerly sent me specimens from the following localities : Sooke, Vancouver Sound, one very large, 21 inches across (Macoun, 1887) ; Barclay Sound, one very large, 20 inches across (1887) ; Discovery Sound, one large six-rayed example, otherwise normal (G. M. Dawson, 1885) ; Queen Charlotte Islands (Dawson, 1885). I have also examined many specimens from Victoria, B. C. ; Puget Sound; Tomales Bay; Monterey; San Luis Obispo Bay; San Diego, Calif., and various other places on the coast of California. Its southern range extends certainly to Santa Barbara, Calif., and probably beyond San Diego. This can scarcely be confounded with any other species, for its conspicuously acervate and reticulate, capitate dorsal spines are characteristic. The reticulation of E. troschelii and L. epichlora is much less conspicuous, and the former has more slender rays; but both, even when very young, differ in having biserial adambu- lacral spines. Larger specimens could hardly be mistaken. 72 VERRILL PISASTER OCHRACEUS, Var. NODIFERUS Verrill, nov. Plate LVI, figures 3, 30. This name is proposed for that variety of this species which has the dorsal spines collected into large detached groups. The form is the same as in the typical reticulated variety, and specimens occur of all sizes from two inches up to a foot or more in diameter. A strongly characterized small specimen from Monterey has the radii 16 mm. and 52 mm. ; ratio, i : 3.25. The dorsal ossicles are reticulate, but so many are without spines that the latter show no regular reticulate pattern. The Geological Survey of Canada sent me several specimens with the nodular character even more prominent. The dorsal spines are all capitate and somewhat unequal; they form a conspicuous pentagon with a central cluster on the disk and five prominent median radial rows proximally. Between the median and superomarginal rows there are ten to twelve large irregularly placed and very prominent clusters of capitate spines, with about five to eight spines in each group. The superomarginal spines form a simple regular row in the type ; they are about equal in size to the dorsal spines, but more conical. The ventral and adambulacral spines and the pedicellariae are like those of the typical form. A very young specimen, also from Mon- terey, has the radii 8 mm. and 24 mm. ; ratio, 1 : 3. This small one agrees with the preceding in all essential characters, but the spines are correspondingly smaller and there are but two distinct rows of interactinal spines. The pentagon on the disk is well marked. This variety occurs commonly from Monterey, Calif., north to Sitka, Alaska. It is the predominating variety on some parts of the coast of British Columbia, on rocks at low tide. Six-rayed specimens are not very rare. HYBRIDS ? There are indications that this species may hybridize with L. epichlora and with others, for where they live associated, as at Sitka, specimens occur that seem to belong to Leptasterias epichlora, but have some of the characters of ochraceus, in a greater or less degree. One young L, epichlora from Wrangel, Alaska (var. subnodu- losa), has only five rays, and the dorsal spines are more strongly acervate and much larger than usual, giving it a close superficial SHALLOW- WATER STARFISHES 73 resemblance to the young of ochraceus, var. nodiferus, of similar size (diameter about 4 inches), but it has the biserial adambulacral spines and other characters of epichlora, and lacks the giant serrate pedicellariae. (See under L. epichlora, var. subnodulosa.) Other specimens have these variations in less degree, and many have more or less of the giant lateral pedicellarise, which may indi- cate an infusion of ochraceus blood, so to speak, for they are not constantly present in epichlora and do not exist in the allied species. Where so many closely related species are associated together, as on the Alaskan coasts, occasional hybridization is to be expected. It is well known to occur on certain middle portions of the New Eng- land coast, between A. vulgaris and A. forbesi, where the ranges of these two species overlap, as well as their breeding seasons, as at Wood's Hole and Vineyard Sound, Mass., just south of Cape Cod. The Asterias janthina Brandt (Prod., p. 269, 1835), probably described from a colored drawing only, was so poorly characterized that no one has been able to identify it with certainty. Dr. Stimpson thought that it might be only a color-variety of ochraceus, and that is not unlikely, » from Brandt's statement. His description was as follows : "A species quite similar to the preceding (ochraceus} but dif- ferent. Diameter of the disk less, rays a little longer, with more numerous and more crowded spines, much more numerous in the center of the disk, and not forming a pentagonal star. Back all light violaceus ; spines white." The relative length of the rays in P. ochraceus is quite variable, and so is the number and arrangement of the dorsal spines. Some of our Sitka and Vancouver Island specimens have the spines closely crowded. The color in most species of this group may vary between yellow, orange, red, and purple. This species is often violet or purple, in life. PISASTER CONFERTUS (Stimpson). Plate xxxvm, figures i, 2 (type) ; plate LIII, figure 2. Asterias conferta STIMPSON, Proc. Boston Soc. Nat. History, vin, p. 263, 1862. Verrill, Trans. Conn. Acad. Sci., I, p. 326, 1867. Perrier, Revision, p. 335 [7i], 1875. Sladen, op. cit., p. 820. Bell. Proc. Zool. Soc. London, p. 494, 1881. Whiteaves, Trans. Royal Soc. Canada, iv, p. 116, 1887. De Loriol, Mem. Soc. Phys. et Hist. Nat., Geneve, xxxn, part 2, p. 17, pi. in, figs, i-ig, 1897. Dr. Stimpson's original description was as follows: " Rays five, stout, rounded, and dilated at base ; disc large. Pro- 74 VERRILL portion of smaller to greater diameter, i : 3.5. Ambulacral furrows broad at the base of the ray, where the pores are crowded into six rows. Ambulacral spines in one regular row, slender, compressed; subcylindric, as long as the ventral spines, somewhat tapering, with bluntly-rounded tips. Ventral spines crowded, very numerous, in six or seven rows near the base of the ray ; short, subcylindric, a little bent outward, and slightly flattened externally, with more or less longitudinal striation near the tip. Beyond the ventral spines there are no regular channels, but the surface is covered with pedicellarise and minute spines. The dorsal spines are numerous, equal in size, but very small, — not more than one-twentieth of an inch in length, scarcely capitate, but more or less pointed. They are uniformly dis- tributed over the disc and rays in a reticulating manner, the inter- spaces being from one-tenth to one-half inch in diameter, and thickly crowded with groups of papulae and minor pedicellariae ; the latter generally occurring about the bases of the spines, but not forming wreaths or crowns. The major pedicellariae are very short and stout, regularly conical or somewhat wedge-shaped, with very broad valves. They are very numerous, particularly on the disc, where they often stand in groups of ten to twenty together. The largest are found isolated in the angles of the rays below, where one occurred having a diameter of eight-hundredths of an inch, and a regularly conical form, with a square base. Diameter of our only specimen, ten inches. " It is allied to A. ochracea, but differs in its more numerous spines, pedicellariae, and ambulacral pores. " Habitat, Puget Sound — North West Boundary Commission. Dr. C. B. Kennedy." I have reproduced large photographs of the type of this species, now in the National Museum. These were sent by Dr. R. Rathbun. (See pi. xxxvni, figs, i, 2, reduced.) In this specimen the radii are about 45 mm. and 162 mm. ; ratio, about as i : 3.6. The rays are thick at base and taper rapidly, though they differ considerably in this respect, indicating that the skeleton in life is rather flexible and weak. The crowded six-rowed or eight- rowed arrangement of the proximal ambulacral pores is very evident. The slender adambulacral spines near the mouth are much longer than those farther out, as in some other species; they stand singly on the plates, in one regular row. The ventral spines are pretty uni- form in size and form, and are nearly smooth and blunt ; they stand mostly in six rows proximally, but form only four or five rows at the SHALLOW-WATER STARFISHES 75 middle of the ray. Probably there are at least four rows of inter- actinal ossicles. The dorsal spines, as stated, are small, mostly subacute, and arranged in a distinctly areolated or reticulated pattern, many of them forming regular transverse or oblique rows or combs on the reticulating ossicles; but they differ rather more in size than indi- cated in the original description, some of them being much smaller, more slender and very acute. But they are far more uni- form than in P. ochraceus. They do not form a definite median row, The dermal minor pedicellariae are numerous. The dorsal, sessile, dermal major pedicellariae are often larger than the spines on the dorsal surface, especially at the interbrachial angles above, and also below, and along the lateral channels. The actinal spines bear thick clusters of minor pedicellariae on their outer sides. VARIATIONS. A specimen of this species from Malaspina Inlet, coll. G. M. Dawson, 1885, received from the Geological Survey of Canada through J. F. Whiteaves, agrees well with the type in most respects, but it has distinct radial bands of small spines. Radii, 28 mm. and no mm.; ratio, 1:4; but the dried specimen is considerably flat- tened, due to imperfect preservation. The dorsal surface is very irregularly reticulated; the numerous small spines have a tendency in many parts to form rather long transverse or oblique rows. Most of the spines are short and blunt, or subacute, but some of the smaller ones are slender and very acute. Minor pedicellariae of very small sizes are exceedingly abundant, both in wreaths around the bases of the spines and on all the inter- mediate surface. The superomarginal spines form a somewhat irregular band of small spines, scarcely larger than the dorsals, sev- eral standing on each ossicle. The inferomarginal spines are similar, a trifle longer, blunt, and mostly striated distally, two often standing on one plate. The interactinal spines, which form three or four rows proximally, are similar in form, but increase in size toward the grooves. The adambulacral spines are long, slender, and regularly tapered. The giant pedicellariae occur mostly on the interradial areas and lateral channels. Some of them are stouter than the inter- actinal spines, but shorter. They are usually stout and blunt, very obtusely conical or blunt wedge-shaped, with slightly denticulate tips. The ambulacral pores are crowded into eight rows proximally. 76 VERRILL M. de Loriol (op. cit., 1896, p. 17, pi. in, figs, i-ig) has described and figured a specimen, evidently of this species, from Vancouver Island. But his general figure (i) makes the dorsal reticulation much more regular than in the type, and the madreporic plate is put in the center of the disk, which is probably the fault of the artist. I have received several other very similar specimens from near Victoria (C. F. Newcombe, coll.) and from Departure Bay (Geo- logical Survey of Canada) . None of these agree perfectly with the type. This species is rare in collections and seems to be known only from Puget Sound, Vancouver Island, and British Columbia, where it is said to be common. It is closely related to P. ochraceus and P. fissipinus by the char- acter of the ventral and adambulacral spines, and by the reticulated dorsal surface. But the dorsal spinulation is even more like that of E. acanthostoma V. The latter has much longer rays, a much feebler skeleton, two rows of adambulacral spines, and lacks the very large, dentate, dermal pedicellarise. PISASTER FISSISPINUS (Stimpson). Plate xxxix, figures i, 2 (type). Asterias fissispina STIMPSON, Proc. Boston Soc. Nat. Hist., vin, p. 264, 1862. Verrill, Trans. Conn. Acad. Sci., i, p. 326, 1867. Pisaster fissispina VERRILL, Amer. Journ. Sci., xxvni, p. 63, 1909. Dr. Stimpson's original description is as follows: " Five rays, short, and dilated at base ; disc large. Proportion of the diameters, i : 3. Ambulacral pores near base of ray crowded, alternating, indistinctly six or eight rowed. Ambulacral spines in one regular row, as long as the ventrals, and flattened on the outer side. Ventral spines sub-equal, stout, sub-cylindrical, truncated, with fissured tips, and a deep, longitudinal sulcus on the outer side ; they form five regular rows. The marginal dorsal spines are as large as the ventrals, capitate, with striated sides and pinched tips, and form an irregular row, of much fewer spines than occur in a ventral row. The spines of the back are few in number, and of only half the size of the ventral spines. They are shaped like the marginal spines, from which they are not ordinarily distinct, and are arranged on reticulating ridges, forming a rather open net-work. On the disc they form a pentagon, from each angle of which extends a median row reaching to one-third the length of the ray. Within the penta- gon, close to its periphery, the madreporic plate is situated. The SHALLOW-WATER STARFISHES 77 dorsal spines increase in size toward the tips of the rays. Papulae ing groups. Minor pedicellariae nearly as in A. ochracea and A. con- ferta; major pedicellariae smaller and far less numerous than in those species, but of similar short and stout form. Diameter, thir- teen inches. " Habitat Shoalwater Bay, Oregon Coast. Northern Pacific Rail- road Expedition. Dr. J. G. Cooper." Large photographs of the type of this species, which is still in the U. S. National Museum, were furnished by Dr. R. Rathbun. They indicate that the average radii are 60 mm. and 170 mm.; ratio, about as 1 : 2.83. The rays taper rapidly from a broad disk. The slender adambulacral spines are compressed and much crowded, one to a plate. The peractinal and synactinal spines are mostly gouge-shaped, though often bifid at tip ; but on the distal half of the ray, many are only sulcate and flattened. The dorsal surface was very accurately described by Dr. Stimpson, as may be seen by comparison with our figures of his type. This species is more nearly allied to P. ochraceus than to any other. It differs in the peculiar gouge-shaped spines, in the more open reticulation of the dorsal skeleton, and in the much smaller and more slender dorsal spines. The locality given by Stimpson is the only one positively known to me for this species. PISASTER BREVISPINUS (Stimpson). Plate XLI, figures i, 2 (type); plate XLIV, figures I, 2; plate XLV, figure i; plate LXIX, figure 3; plate LXXVI, figures i-ib (details). Asterias brevispina STIMPSON, Journ. Boston Soc. Nat. Hist., vi, p. 88, pi. xxm, fig. 3 (ventral spines), 1857. Pisaster brevispina VERRILL, Amer. Journ. Sci., xxvin, p. 63, 1909. The original description by Dr. Stimpson is as follows : " Rays five, each equalling in length twice the diameter of the disk. Upper surface covered with very short, blunt, nearly uniform spines, moderately numerous, sometimes forming an irregular row along the middle of the ray, and showing a tendency to reticulation on the sides. Beneath, there is a single row of slender ambulacral spines, which are blunt and somewhat irregular in length; between these and the marginal channel there are four rows of short compressed spines, gouge-shaped, or notched by an oblique concavity at their truncated extremities. Madreporic body large. Color yellowish. Diameter, six inches. Taken from a sandy bottom in ten fathoms near the mouth of San Francisco Bay." 78 VERRILL Dr. Rathbun has sent nearly natural-size photographs of the original type of this species, which seems to be almost unknown in many later collections. The disk is of rather large size and elevated ; the five rays taper rapidly from rather wide and swollen bases. The radii are about 30 mm. and 116 mm., if the photographs are correctly marked as to reduction ; ratio, about as 1 : 3.87. The adambulacral spines are small and slender, mostly compressed, crowded in one row, one to a plate ; those near the mouth are longer. The inferomarginal and actinal spines are very much larger, rather short and very stout, mostly clavate or subspatulate, and with a deeply grooved or gouge-shaped tip ; but many are simply sulcate and little flattened. They form four or five crowded rows near the bases of the rays, the inferomarginals bearing two spines, but at about the middle they are reduced to three or four regular rows. Most of them bear close clusters of minor pedicellariae on their outer side. The superomarginal spines are smaller, fusiform or subconical, sub- acute; they form a pretty regular row, separated from the infero- marginals by a rather wide channel. The dorsal spines are short, thick, nearly equal, with conical, mostly subacute tips. They are rather irregularly distributed, in short broken rows or singly ; they form a somewhat evident median radial series, of two or three close irregularly alternating rows on the basal part of the rays ; on the sides many of them are in obliquely transverse short rows, and sometimes short longitudinal rows can be traced. They bear small basal groups of minor pedicellariae. Dermal minor pedicellarise everywhere scattered between the spines. The sessile, dermal, major pedicellariae appear to be few and mostly along the submarginal channel; they are rather small, short, sub- conic, obtuse. A few lanceolate, acute, major pedicellariae occur along the edges of the ambulacral grooves. VARIATIONS. This species varies considerably in appearance, according to age, locality, and mode of preservation. When dried carefully, from alcohol, the disk is high and convex, while the mouth and jaws are deeply sunken; the rays are stout and high at base but rapidly tapered. There is usually a distinct median dorsal row of spines, simple in the younger specimens, but often double- or triple-ranked in the older ones. SHALLOW-WATER STARFISHES 79 On the medium-sized specimens the pentagon of the disk is very evident, but in large specimens it is obscured by the numerous inter- polated scattered spines. In those about six to eight inches in diameter, the central plate, five radial, and five interradial primary plates are evident. Each bears a cluster, usually of four to six short, thick, clavate spines, with ovate or acorn-shaped striated tips. They are similar, in size and form, to those of the median radial rows. Other somewhat smaller, but similar, spines are scattered within and around the primary pentagon, and over the dorsal surface gen- erally. But the dorsal spines are not numerous in the young. They do not form distinct rows, but are irregularly scattered, and are mostly pretty nearly equal in size and form. In the larger specimen, ten to twelve inches in diameter, the dorsal spines become decidedly more numerous, and many are grouped two to four or more on one plate. The superomarginal spines are similar to the dorsals, usually with rather more conical tips, which are strongly striated. They form a regular, close row, simple in the smaller specimens, but often double proximally in the larger ones. The inferomarginal row is very regular and nearly always double, two spines standing obliquely on each plate, sometimes three in the larger specimens. They are rather stouter and more obtuse than the upper marginals, usually with the tips striated in small specimens; but in the larger ones these spines mostly lose their striations, and many become more or less flattened distally, while some are apt to have a distal furrow on the upper side ("gouge-shaped"), though this is by no means constant. In nearly all the specimens there are but two simple, regular actinal rows of spines, making, with the inferomarginals, four close, unusually regular rows. The actinals are short and stout, similar to the inferomarginals. In the larger specimens some of them are somewhat flattened, and some are often grooved or slightly bilobed distally. The adambulacral spines are slender, four or sometimes five corre- sponding to each synactinal plate. In young examples they are mostly regularly tapered and form a very regular even row ; in the larger ones, many of them are often stouter, obtuse, cylindric, a little flattened, or even slightly clavate. They become longer proxi- mally, near the mouth. Major pedicellarise of rather large size occur on the back, sides, and interradial axils, on the adambulacral spines, and on the inner 80 VERRILL margins of the grooves. Those of the actinal side and ambulacral furrows are mostly acute-lanceolate or acuminate, compressed, with the sharp curved tips often overlapping. They often exceed the adambulacral spines in diameter. Those of the dorsal surface are shorter, stouter, ovate, with narrow obtuse tips, which are slightly denticulate. Those on the intermarginal furrow are largest and stoutest, with stronger denticulations. Minor pedicellarise are very small. They form small wreaths around most of the dorsal spines, and clusters on the outside of the ventral ones; small clusters also occur on the inside margin of the adambulacral plates and on some the larger major pedicellarise. On the dorsal surface they form mostly rather small clusters, pretty evenly scattered over the whole surface; in large specimens they often become very numerous and pretty evenly distributed. The papulae are very small and apparently not very numerous, except in large examples. The integument is thick and firm, so that it conceals the outlines of the plates. The mouth is deeply sunken. The jaws are elongated, narrow or much compressed, nearly perpendicular within ; the adoral spines form two close rows of about eight each ; the apical oral spines are rather stouter than the rest; there is often but one to a jaw. The color, in the only case noted, was pale purple or pink. A good specimen from off Monterey, Calif. (No. 18290, M. C. Z.), differs somewhat from all the others. Radii, 33 mm. and 130 mm. ; height in middle, 40 mm. The central and ten primary radial and interradial plates are covered with clusters of short capitate spines with obtuse tips, about eight to ten in a cluster. About ten smaller plates within the pen- tagon are indicated by small clusters of simple spines. The median radial rows are conspicuous. On the proximal third of the rays they mostly contain three regular rows of spines; further out the three rows are less regular. On the distal fourth there is usually but one row. The other dorsal spines are mostly grouped in clusters of three to six, the clusters being irregularly scattered, largest on the sides of the rays. The spines are all similar, short, thick, with more or less conical, obtuse or ovate striated tips. The marginal and actinal rows of spines are very regular and even. They stand in close rows. There is a wide and deep intermarginal channel. The superomar- ginal row of spines is simple. The inferomarginal row is double. The two interactinal rows are simple and nearly equal. SHALLOW- WATER STARFISHES 8 1 MEASUREMENTS. Radii Serial number. Lesser. Greater. Locality. 13040 50 mm. 168 mm. Off San Francisco. 13046 26 mm. 73 mm. Off San Francisco. 1304*: 18 mm. 60 mm. Off San Francisco. 1304^ 42 mm. no mm. Off San Francisco. 18216 22 mm. 63 mm. Off Santa Cruz, Calif. 18200 33 mm. 130 mm. Off Monterey, Calif. This species ranges from south of San Francisco to Puget Sound and northward to Departure Bay, B. C. In the Museum of Comp. Zoology at Cambridge, Mass., I have studied a good series from off San Francisco (No. 1304) ; Crescent City, Calif. (No. 1303) ; Gulf of Georgia, A. Agassiz (No. 1301) ; Monterey Bay (No. 1820, see figs.); off Santa Cruz I., Calif. (No. 1821). It appears to be common in shallow water on the Californian coast. Very few authentic localities have been recorded hitherto, except that given by Dr. Stimpson (Bay of San Francisco). A single large specimen has been sent to me from Departure Bay, B. C. (coll. H. C. Young, Geological Survey of Canada). PISASTER CAPITATUS (Stimpson). Plate xxxvi, figures 3, 4 (type) ; plate LVI, figure 4. Asterias capitata STIMPSON, Proc. Boston Soc. Nat. Hist., vm, p. 264, 1862. Verrill, Trans. Conn. Acad. Sci., i, p. 327, 1867. ? Perrier, Arch. Zool. Exper., rv, p. 335, 1875 (six-rayed specimen, no descr.). Dr. Stimpson described this species as follows: " Rays five, not contracted at base ; disc large. Proportion of diameters, i : 4.5. Ambulacral pores rather narrow, in four regular rows. Ambulacral [adambulacral] spines in one regular row, linear, compressed, and blunt. Ventral spines as long as the ambulacrals, capitate, with bluntly-rounded heads, elegantly striated on the con- vex inner face and tip, and with a median sulcus on the outer side. They are arranged in four rows, those of the outer row being largest ; and there are some minor pedicellarise on the outer sides of the spines in all of the rows. The dorsal spines are not very numerous, but are for the most part large, their regularly globular and beauti- fully striated or radiated heads being about eight-hundredths of an inch in diameter, and larger than those of the ventral spines. They are arranged without order, standing about one-seventh of an inch apart; but five or six longitudinal rows may be obscurely traced, the marginal row being most distinct, containing eighteen or twenty 82 VERRILL spines. On the disc, there is a central tubercle, but scarce any indi- cations of a pentagon. Around the bases of the dorsal spines there are regular wreaths of minor pedicellariae. The major pedicellariae, which are of the short, conical or sub-globular form, with broad valves, are scattered between the spines as in A. ochracea. Papulae in groups. " Color in life purple, according to Dr. Newberry. The spines are probably white, or at least of a lighter color. Diameter, five and a half inches. "This very pretty species differs from A. ochracea in its larger dorsal spines, which are not arranged in a reticulating pattern ; and from A. Liitkenii in its shorter and more numerous ventral spines, as well as in the presence of major pedicellariae on the back. " Habitat, San Diego, Cal. Colorado Expedition. Dr. J. S. New- berry." The photographs of the type, sent by Dr. R. Rathbun, are six inches in longest diameter; radii, 18 mm. and 80 mm.; ratio, nearly i : 4.5. Little need be added to Stimpson's description of the type. The adambulacral spines are often flattened. The ventral spines are unequal ; those of the inner or synactinal row are distinctly smaller than the others, and the outer or peractinal are largest. Most of these spines are stout and clavate, rather than capitate, the tips being mostly obtusely conical or ovoid and strongly striated; only a part are grooved ; distally they become much more slender and distinctly clavate. The superomarginal spines are considerably stouter than the ventral ones, but not much longer. They form a regular simple row of rather widely spaced spines, each one on the raised node of a marginal ossicle. The dorsal spines are mostly quite similar to the laterals in size and form. The larger ones have large, rounded, capitate, and strongly striate tops ; but those toward the ends of the rays become longer and more slender, cylindrical or clavate, rather than capitate, but striated in the same way. The spines are distantly spaced, generally one on each raised node of the reticulating ossicles. Usually, in our specimens, five or six ridges, indicating the lobes of the ossicles, radiate out from the base of each spine, so that the skeleton has a pretty regularly reticu- lated structure. The spines usually form more or less evident median, radial rows, and sometimes two less distinct rows can be traced, between the median and marginal rows. SHALLOW-WATER STARFISHES 83 In the Museum of Yale University there is a specimen of this species collected at Monterey, Calif., which agrees very closely with the type in size and structure. The radii are 20 mm. and 70 mm. ; ratio, 1 : 3.5. The rays are shorter than in the type. The spinulation agrees in all essential points with the type, though the distal dorsal spines of the rays appear longer and less stout, and more of them have the clavate form. The wreaths of minor pedicellariae about the spines are not con- spicuous, but they contain very large numbers of very minute pedi- cellariae. Others of similar small size are scattered between the spines or grouped in dermal clusters on or around the large papular areas. The papulae are small and form large groups, above and below. The dorsal and lateral dermal major pedicellariae are moderately large and stout, ovoid or subconic, with obtuse and usually denticu- lated jaws. They are not nearly so numerous nor so large as in the type. Most of the actinal spines are clavate and many lack the sulcus on the outside; the synactinal spines are smaller and more cylin- drical; between them are large papular areas. The dorsal skeleton is firm and distinctly reticulated, with a tendency to a six-rayed arrangement around the principal spines, which are not so numerous as in the type. Monterey, Calif. (R. E. C. Stearns, Mus. Yale Univ.) ; San Diego (Stimpson, Ives, and others) ; San Diego (Mus. Yale Univ. coll.). This species is closely allied to P. liitkenii. It differs especially in having much fewer dorsal spines, which are also larger and more distinctly capitate. The ventral (actinal) spines are more unequal, stouter, shorter, and more capitate. PISASTER LUTKENII (Stimpson). Plate XL, figures i, 2 (type). Asterias liitkenii STIMPSON, Proc. Boston Soc. Nat. Hist, vnr, p. 265, 1862. Verrill, op. cit., 1867, p. 32. Perrier, Revis., p. 70, 1875. Bell, Genus Asterias, Proc. Zool. Soc. London, 1881, p. 495. Sladen, op. cit., pp. 566, 824, 1889. De Loriol, Mem. Soc. Phys. et Hist. Nat., Geneve, xxxu, p. 15, pi. ii [xvn], figs, i-i/t, 1897 (description). Pisaster liitkenii VERRILL, op. cit., p. 63, 1909. Asterias exquiseta DE LORIOL, Rev. Zool. Suisse, 1888, p. 463, pi. xvin. fig. 2 (young). When full-grown this is one of the largest starfishes on the coast. Dr. Stimpson's type was not half-grown. The following is his description : 84 VERRILL " Rays five, rather broad, with blunt tips ; disc moderately large. Proportion of the diameters, i : 4.25. Ambulacral pores in four regular rows. Ambulacral [adambulacral] spines in one regular row, slender, long (but not longer than the ventrals), not com- pressed, but tapering to a blunt point. Ventral spines in four approximated rows, elongated, scarcely capitate; heads elongated, subtruncate, striated within, and often sulcated along the middle on the outer side, where there are always semicircular clusters of minor pedicellariae. Beyond the ventral spines there are distinct lateral channels. The dorsal spines are uniform in size and distance, being about one-fourth of an inch apart in one specimen, and arranged without order, except in the regular marginal row, which consists of about thirty spines; and they form no pentagon on the disc. They are one-eighth inch high, and half that in breadth, capitate, with the heads striated and conical, with pinched tips. Around the base of each there is a rather narrow ring of minor pedicellariae. There are no major pedicellarise scattered among the dorsal spines, but a few occur in the lateral channels, similar to those of A. ochracea and capitata, but of smaller size and with narrower valves. Papulae in groups. Diameter, one foot. It differs from A. gigantea in having only five rays, and in other particulars. " Habitat, Coast of Oregon. Mus. Smithsonian." Photographs of the types of this species were also kindly sent by Dr. R. Rathbun from the U. S. National Museum. As marked, the photographs give the radii as 24 mm. and 96 mm. ; ratio, 1 : 4. The description by Dr. Stimpson is so good, in this case, that little need be added to it. The upper marginal spines form a pretty regu- lar row ; they are about as large as the ventrals, but more conical, and like the dorsal spines in form, though rather larger. The median dorsal row of spines is not often very evident ; in some places indi- cations of four or five other longitudinal rows can be seen on either side, so that there may be ten to twelve spines in a very irregular transverse series, but there are no distinct transverse rows. On the distal part of the rays the wreaths of minor pedicellariae, around the spines, become larger and crowded ; there are also large clusters of dermal minor pedicellariae scattered over the whole dorsal surface, many of them on the papular areas, besides many single pedicellariae. Two very large and perfect specimens of this species are in the Museum of Yale University, collected at Pacific Grove, California, by Professor W. R. Coe, 1901. SHALLOW-WATER STARFISHES 85 One of these (a) agrees very closely with the type, except in features due to its much greater size. Its radii are 60 mm. and 270 mm. ; ratio, 1 : 4.5. The dorsal surface is nearly uniformly covered with conical spines, not forming definite rows. About twenty-one to twenty- five may be counted in an irregular transverse series at the base of the rays. The upper marginal plates mostly bear two spines at the base of the rays, but only one distally. The lower marginals bear two com- plete rows, to the tips of the rays. There are three regular rows of actinal spines proximally, mostly one to a plate ; the first is complete to the tips ; the second ceases near the tip ; the third at about the distal third. The inferomarginal row and first two actinals are all close together and much alike; they are rather short, very stout, clavate, with rounded, striate tips, channeled on the outer side. The syn- actinal spines are longer and not quite so stout, tapered and chan- neled on the outer side. On the interradial areas some of the actinal spines are also longer, conical, and subacute. The adambulacral spines are slender and tapered, becoming decidedly longer toward the mouth and on the adoral plates. The adoral carinae are about 14 mm. long, strongly compressed, and con- sist of about eleven or twelve closely united pairs of adambulacral plates, sloping upward, and normally bearing each one long spine; but in many cases one is lacking in a pair, and on some of the carinse two to four pairs are destitute of spines. Those in the last erect pair (epiorals) on the jaw are longer and stouter than the others. The terminal or peroral pairs of spines are horizontal, short, stout, flattened, enlarged or subspatulate at the blunt tips. A short, thick, strongly lateral divergent spine, not half as long, occupies the side angles. The adambulacral pores are large, and on the proximal part of the rays, within the radius of the disk, they form eight distinct rows; they are reduced to four regular rows a little beyond the border of the disk. On about three adoral pairs of plates, they also decrease, the innermost being a single pair. The minor pedicellariae are abundant on nearly all the spines, but the wreaths around the bases of the dorsal and marginal spines are not so large as in var. australis. Numerous, large, erect, sessile, stout-ovoid and stone-hammer-shaped pedicellariae, with strongly serrate, blunt jaws, are scattered everywhere over the back, on the 86 VERRILL lateral channels and among the ventral spines. Along the inner edges of the adambulacral grooves, attached by pedicels, there are many compressed and longer ones, some of which are larger than the dorsals, others much smaller. These are mostly long-ovate, often with incurved margins and blunt at the tips, which are den- tate ; others are acute-lanceolate, and many are quite small. In the Museum of Yale University there are also several smaller dry specimens of this species, agreeing very closely with the type. The best medium-sized specimens, with the radii 40 mm. and 180 mm., have the ratio as i : 4.5. The adambulacral spines are slender, terete, subacute, arranged in one simple row. Those near the mouth are not much lengthened; more distally many of them are somewhat flattened, with obtuse or slightly spatulate tips. They are mostly without minor pedicellariae, though small groups occur on some of them. A few bear single, rather large, ovate-lanceolate major pedicellarise, like the adambulacral pedicellarise. The latter are rather numerous, attached to the inner edge of the adambulacral plates, at or within the margin, by a long pedicel, which often bears a cluster of minute acute pedicellariae at and below the base of the large one ; the latter is unusually large, compressed-ovoid, subacute or obtuse, with the tips of the jaws often denticulate. Their pedicels are often as long as the adambulacral spines. The actinal spines are large, subequal, and form four or five regular rows, proximally, but usually only four beyond the middle of the ray; they become much more slender distally ; those of the interbrachial region become longer, fusiform, and often acute. Most of them are stout, clavate or subcapitate, strongly sulcated on the outside and with the tip finely striated all around. Most of them bear a small cluster of minor pedicellarise on the outside. There are, apparently, only four rows of peractinal and synactinal plates. The outer (peractinal) ones often bear two spines. The marginal spines are larger and rather longer, similar to the dorsal spines, and with subconic striated tips; they form a very regular row, standing singly and well spaced on a row of somewhat raised ossicles, and leave below them a wide and well defined mar- ginal channel, which becomes dilated at the interbrachial region. The dorsal spines are similar in form but unequal in size, numer- ous, arranged without any very evident order, but pretty evenly spaced. They mostly stand singly on the nodes of the reticulating ossicles, which form five to seven radial ranges and nine to ten in transverse series. The dorsal spines bear a thin basal wreath of SHALLOW-WATER STARFISHES 87 minor pedicellariae. There are also many small clusters of dermal minor pedicellarise. Papulae small, very numerous, in clusters. The dorsal major pedicellariae are numerous, scattered among the spines and especially in the lateral channels. They are short, thick, ovoid, obtuse, with the jaws denticulated at the apex; they often equal the smaller spines in diameter, but are shorter. This species can only be confounded with P. capitatus, to which it is evidently closely allied. The latter, however, not only has much larger and more capitate dorsal spines, but they are much fewer in number, there being usually only about five or six spines in a trans- verse series, besides the laterals. The supposed absence of large major pedicellariae on the back of P. liitkenii, referred to by Stimpson, does not hold good, as shown by our specimens, above described. This species appears to range, in its typical form, from Van- couver Island to Monterey, California, and as a variety farther south to San Diego. Oregon (Stimpson) ; Saanich Inlet, Vancouver Island (de Loriol) ; San Diego and Monterey, California (Yale Museum) ; Vancouver Island (Canadian Geological Survey) ; off Pacific Grove, near Monterey, taken on fish lines (Dr. W. R. Coe. Two very large specimens, a, b). When full-grown this is one of the largest starfishes on the North- west Coast. It is equalled, in this genus, only by P. achraceus, P. giganteus, and P. papulosus. VARIATIONS. One of the very large specimens (b) obtained from off Monterey by Dr. Coe, 1901, differs considerably from the more typical ones in the character of its spinulation, due perhaps, to a superabundance of food. Its radii are 55 mm. and 284 mm. ; ratio, about 1 : 5.2. It is noticeable on account of the great development of its actinal and inferomarginal spines, which are decidedly more numerous and longer than in the other specimen of equal size and from the same place, described above. The proximal inferomarginal and most of the actinal plates bear two, three, or even more, stout, often divergent spines, so that one can count eight to ten of these spines in each irregular transverse series. But one ray (doubtless a reproduced one) is only about two- thirds as long as the rest, though nearly as stout ; on this the ventral 88 VERRILL spines form only four or five rows, though similar in size. On the distal half of all the rays the ventral rows decrease to about four to six, the plates bearing only two spines, or but one. The ventral spines are unusually crowded proximally. They are also longer and less stout, though mostly blunt, except the syn- actinals and those on and near the interradial areas, where many of them become decidedly longer and acute. Many in the synactinal row are deeply grooved on the outside, and the distal ones become more slender and subacute. The adoral spines are long, slender and acute. Large serrate major pedicellarise are scattered over the dorsal and lateral surfaces, but are not half so numerous as on the other large specimen. The dorsal spines are less numerous, larger, more acute, and more regularly spaced, their average intervals being 8 mm. to 10 mm. wide. Minor pedicellariae form dense wreaths around all the dorsal and upper marginal spines. Off Pacific Grove, California (Dr. W. R. Coe, 1901). PISASTER LUTKENII Var. AUSTRALIS Verrill, nov. Two very large specimens (c, d) of this variety are in the Museum of Yale University. One has the radii 56 mm. and 252 mm. ; ratio, 1:4.5. The oral adambulacral spines are one-fourth longer than those of the mid-ray ; they are long, slender, regularly tapered. The ventral spines, near the base of the rays, are in four and five crowded rows ; but on the mid-ray there are about four regular rows ; those of the inner and middle rows are very stout and blunt, often nearly as thick as long, irregular in form, often pinched, lobed, or sulcated, and sometimes slightly bifid; the tips are often clavate, swollen, or truncate. The inferomarginals, or two outer rows, and those near the mouth, are longer, not so stout, and more regular, mostly with obtuse or ovoid tips. Distally they become longer and more fusi- form, and many are deeply channeled on the outer side. The inferomarginal spines stand two on a plate, nearly or quite to the tips of the rays, and thus form the two outer rows of ventral spines. There are proximally three rows of stout actinal plates, each usually bearing a single spine, thus forming three rows; the first row (peractinal) extends to the tip of the ray; the third row (syn- actinal) only extends to about the distal third of the ray. SHALLOW-WATER STARFISHES 89 The dorsal spines are much less numerous than in the typical variety; they are evenly scattered, and scarcely form any distinct radial rows. About thirteen to fifteen spines occur in irregular transverse series, besides the upper marginals, but they do not form any definite transverse rows. The upper marginals are much like the dorsals, but larger. They mostly stand singly on the large plates, so that they are well apart. Between the upper and lower marginals there is a rather wide, naked channel, bearing large pedicellariae; at the base of the rays it expands into a rather large, triangular, interradial area, destitute of spines. Minor pedicellariae, in very large dense wreaths, surround the upper marginal and dorsal spines, the clusters becoming so large distally that they are in contact. In dry specimens they seem to be attached outside the swollen base of the spines, which is covered by a thick skin, but in life they probably rise up at least to mid-height of the spine. On the ventral spines they form large clusters on the outer surface. Very large, ovoid, or blunt stone-hammer-shaped, erect, sessile, major pedicellariae, with strongly serrate jaws, occur in considerable numbers on the lateral channels, between the dorsal spines, and among the ventral spines. Other major pedicellariae, some of them nearly as large, but others much smaller, somewhat compressed, mostly ovate and obtuse, occur on the adambulacral spines and on the inner edge of the ambulacral grooves. Many of these have large numbers of small, acute, forficulate pedicellariae on their pedicels. San Diego, California (Dr. Edw. Palmer, Yale Museum). PISASTER GIGANTEUS (Stimpson) Verrill. Plate xxxvn, figures i, 2 (type). Asterias gigantea STIMPSON, Jour. Boston Soc. Nat. Hist, vi, p. 88, pi. xxni, figs. 4-6, 1857. Verrill, Trans. Conn. Acad., i, p. 327, 1867. ? Bell (pars), Proc. Zool. Soc. London, 1881, p. 564; = in part A. katherina Per., non Gray. Dr. Stimpson's incomplete original description is as follows: " Body very large, swollen ; rays six in number, in length some- what less than twice the diameter of the disk. Upper surface cov- ered with numerous short, blunt, equidistant spines, uniform in size and regularly distributed; these spines are somewhat conical in shape, but truncated at the tip and constricted at the base, with the sides longitudinally furrowed. The spines of the lower surface QO VERRILL are short and thick, but slightly compressed and notched at the extremity. .Diameter, two feet. Taken in Tomales Bay, by Mr. Samuels." Large photographs of the type, which is in the U. S. National Museum, have been furnished by Dr. R. Rathbun, for reproduction in our plates. Dr. Stimpson's description was very incomplete, for he scarcely described the ventral side at all. The disk is rather large and the six rays taper rapidly to rather acute tips. They are variously bent, and wrinkled, indicating a rather weakly reticulated dorsal skeleton. The radii are about 73 mm. and 304 mm. ; ratio, about 1 : 4.75. The adambulacral spines are rather small, slender, compressed, crowded, and stand in a single row, one to a plate; they are much smaller and shorter than the crowded ventral spines. The latter form about eight close rows proximally, but these decrease to four or five at the distal third of the rays. They differ much in form. Those nearer the adambulacrals are partly fusiform and sometimes subacute, but more often flattened and subacute at the tip ; those of the middle rows are stouter, either blunt, clavate, or flattened ; those of the outer rows are still shorter and stouter, with blunt tips, but those near the base of the rays are apt to be fusiform and subacute. The lateral or superomarginal row of spines is not very distinct; its spines are subacute and much smaller than the actinals. They form in some places two irregularly alternating rows, and differ but little, in size or form, from the dorsal spines, though they may be rather longer and more acute. The dorsal spines are small, either obtuse or acute, roughly striated, very numerous, nearly uniformly scattered over the whole surface, mostly standing singly, and do not form reticulations nor any distinct median row. On the sides of the rays, in some places, they form small transverse groups or combs on the connective ossicles. About twenty to twenty-four can be counted in an irregular series across the rays at the base, but they form no evident trans- verse rows. The spines become rather larger, longer, and more crowded near the ends of the rays, where they are surrounded by close wreaths of minor pedicellariae. On the basal part of the rays and on the disk the minor pedicellarias form small clusters around the bases of the spines, and many other dermal ones are scattered between the spines. A few rather small, stout, subconical, dermal major pedicellariae are scattered on the dorsal and lateral surfaces, and especially on the SHALLOW- WATER STARFISHES 91 interbrachial areas and intermarginal channels. The actinal spines bear small clusters of minor pedicellarise on their outer sides. This species appears to be very distinct from all others. Perrier placed it as a synonym of his A. katherince = P. grayi Ver., to which it bears some little resemblance in form. He was probably misled by Stimpson's very poor description. Nothing is known concerning its range. Stimpson's original record is the only authentic locality known to me. Some of the localities recorded are due to errors. PISASTER PAPULOSUS Verrill. Plate XLH, figure i; plate XLIII, figure i; plate LX, figure i; plate LXXVI, figures 2-2d (details) ; plate LXXX, figure 4. Pisaster papulosus VERRILL, Amer. Journ. Sci., vol. xxvm, p. 63, 1909. The type of this species is a large dry five-rayed specimen, in excellent condition, taken in Puget Sound and sent to me by Pro- fessor T. Kincaid. A specimen received from the Provincial Museum of British Columbia, through Dr. C. F. Newcombe, but not in so good preservation, is much larger (diameter, 28 inches or 660 mm.). Thus, when full-grown it is one of the largest starfishes yet discovered. The disk is large and remarkably high and plump, with a raised central pentagon, bearing ten groups of spines on the angles and sides, besides a central group ; from the radial angles carinal radial rows of spines run to the tips of the arms, which are long, rather stout at the base, but regularly tapered to acute tips. The radii of the type are 42 mm. and 215 mm. to 225 mm.; ratio, about i : 5.25 ; breadth of rays at base, 45 mm. ; elevation of disk, 46 mm. ; length of dorsal spines, 4 mm. ; diameter, i mm. ; length of larger ventral spines, 3 mm. to 4 mm. ; diameter, 2 mm. to 2.5 mm. The dorsal surface is covered with an integument that conceals the ossicles. The papular areas are large and numerous ; a conspicuous row runs along each side of the median radial row of spines, and there are several less regular rows on each side. The papulae are very numerous and small, darker than the surrounding integument, which is light yellowish orange in the dried specimen; the papular areas brown. The dorsal spines are large, few, and mostly isolated, but they form a distinct, simple, radial or carinal row, one to a plate, proxi- mally, but on the distal third of the ray these spines become more distant and the row less distinct. 92 VERRILL On either dorso-lateral side there are similar, but rather sparsely scattered, spines, which mostly show no definite arrangement ; but in some cases, distally, they have a tendency to form about three imperfect rows on each side. These spines are all rather large, up to 3 mm. to 4 mm. high, and about i mm. thick, tapered or conical, subacute or acuminate, and often pinched up near the tips and strongly sulcated. They bear large and dense wreaths of minor pedicellariae placed around their bases as dried, and there are numer- ous large, dense clusters of similar pedicellariae thickly scattered over the surface between the spines, and around the papular areas, as if taking the places of the spines that are present in other species. Many of these clusters are as large as those around the spines, or up to 3 mm. to 4 mm. in breadth. They are abundant on both the disk and rays, but become more numerous on the distal half of the rays, where they are often so crowded that in life they must form a nearly continuous coating. Large, stout, erect, dentate, dermal, major pedicellariae are numerous, scattered over the whole dorsal and lateral surfaces of the rays. They are rather quadrangular in an end view; in a side view scarcely compressed, blunt wedge-shaped or stone-hammer-shaped. The tip of the jaws are serrate or unguiculate and subtruncate. The larger ones are about 1.5 mm. high and nearly I mm. broad. The pentagon of the disk is formed by five radial clusters of stout conical spines, like the dorsal radials, standing about three together, and five similar interradial groups, standing a little nearer the center. The large, brown madreporic plate takes the place of most of those in one interradial space. A cluster of five spines occupies the center. The large dorsal interradial areas of the disk are mostly without spines, but have an abundance of pedicellariae and papulae. The upper marginal rows of spines are like the median, but they become more regular and the spines closer together distally. The channel below them is wide proximally, and bears many of the large, serrate pedicellariae and numerous minor ones. The row of infero- marginal spines is strong, double, and regular, each plate usually bearing two spines, especially distally, to the tips of the rays. The distal spines are subacute and sulcate, much like the dorsals, but those on the proximal half are mostly stouter (2 mm. or more), with obtusely rounded and sulcated tips. Close to these, but with an intervening papulose channel, is a regular, mostly single row, of peractinal spines, continuous to the tips of the rays. The spines are like the inferomarginals. A SHALLOW-WATER STARFISHES 93 shorter row of similar synactinal spines, extending to the distal third, intervenes between the peractinals and the adambulacrals, but is separated from the latter by a definite channel, having papulae proximally. Close to the base of the rays some of the infero- marginal and actinal plates often bear two or three small extra spines, and all bear dense clusters of minor pedicellarise on the outer side. The adambulacral spines form a single very regular row; they are slender, tapered, and subacute; about five correspond to a single actinal spine. The adoral ones become somewhat longer. Many of them bear thick clusters of forficulate pedicellariae of various sizes in a single cluster. Between their bases and on the inner edges of the ambulacral grooves, and on the oral spines, and interradial areas are many large major pedicellariae, part of which are unguiculate, like those of the back, but others are longer, com- pressed, lanceolate, with the tips subacute, or terminating in one or two hook-like teeth. The pedicels of these, and also of those attached to the spines, are often surrounded by large clusters of small forficulate pedicellariae of nearly the same form, but of various sizes, many being as minute as the ordinary minor pedicellariae. Friday Harbor, Puget Sound (Professor Kincaid). The paucity of spines on the back of the specimen above described might be thought to be due in part to some injury and lack of subsequent restoration of the spines. But on the distal part of the rays, at least, the spines seem to be certainly normally arranged. There are no naked ossicles or tubercles where spines are likely to have been attached. I have examined four additional specimens (No. 1904, Mus. Comp. Zool.) from the same district and also sent by Professor Kincaid. They are all larger. One of them agrees closely with the type specimen described above. Its radii are 1 10 mm. and 330 mm. Others, equally large, have the disk and rays flattened in drying and appear to have more numerous and larger spines and fewer pedicellariae. One has the radii 85 mm. and 330 mm. This species is rather closely related to P. brevispinus (Stimpson). But the characters of the dorsal spines and pedicellariae and of the actinal spines, and the larger spines and papular areas, seem to require their separation. Of course a much larger series of specimens than I have studied might show intermediate forms, as in other cases. The difference in the number of rows of actinal spines is greater than we should expect in specimens of similar size. The dorsal spines are much less numerous in our species and the rays much longer 94 VERRILL and more tapering. The characters of the adoral carina and oral spines are also peculiar. VARIATIONS. A very large dry specimen of this species has been sent to me by Dr. C. F. Newcombe from the Provincial Museum of British Columbia. It was dredged in ten fathoms, off Victoria, 1894. Its radii are 64 mm. and 330 mm. ; ratio, 1 : 5.14. The disk is distorted in drying and is probably larger than the normal size. The rays are thick at base, but taper to long and relatively slender tips. The dorsal surface is covered with a thick integument, dark reddish brown in the dry specimen, and has but few distant spines. But numerous large, stout, thick, unguiculate pedi- cellariae are scattered over the surface, which is also covered with small minor pedicellarise, thickly scattered over nearly the whole surface and also forming wreaths around the bases of the spines. Probably a large part of these may have been rubbed off by repeated dustings, for the specimen has evidently been exposed to dust and mold; and owing to its great size, it probably was not very well preserved originally. Hence it is probable that the dermal minor pedicellariae originally existed in still larger clusters, as they do in depressed spots, where the coating of dust and mold has not been removed, especially distally. But as numerous large major pedi- cellariae are still scattered over the surface, it is evident that no severe cleansing has taken place, sufficient to remove all those on the spines. The dorsal skeleton is coarsely and irregularly reticulated, but the ossicles are concealed by the thick integument. The spines are mostly conical and acute or subacute, about 1.5 mm. to 2.5 mm. long, with but few smaller ones. They form distinct but incomplete simple radial rows, in which they usually stand 2 mm. to 5 mm. apart, where continuous. A very few similar spines are irregularly scattered over the disk and rays. The madreporic plate is large, with fine gyri, its diameter is 13 mm. The superodorsal plates are well down on the sides of the rays. They are relatively small and not very distinct, and bear a simple row of spines, rather larger than the dorsals, many of them blunt with grooved tips. They are close to the inferomarginals, except in the interradial areas, where they diverge a little, leaving a spine- less area. The ventral spines proximally consist of four or five rows close together and nearly equal. They are short (3 mm. to 4 mm. long), SHALLOW-WATER STARFISHES 95 stout, mostly blunt or truncate and striated at the tip ; but close to the bases of the rays they become longer. Some are acuminate and subacute. On the distal third of the rays there are only three rows, besides the superomarginals, there being but one row of actinals distally. The inferomarginal ossicles are rather large, thick and convex, roundish on the outer end, and usually bear two or three divergent spines, which are mounted on low mammillae with a central pit. The peractinals are similar in form externally, and mostly bear but one spine, but sometimes two proximally. The principal row of sub- actinal plates, proximally, are large and transversely broad in their internal part, but show only a rounded exterior surface, smaller than the other plates and bearing a single spine. Between these, proximally, there are a few smaller wedge-shaped or irregular sub- actinal ossicles, interpolated irregularly and mostly without spines. The inferomarginal and actinal plates are closely crowded together. There is a rather wide naked channel between the synactinal and adambulacral plates, due to the great transverse length of the latter externally to the single spine that stands on the inner edge. The adambulacral spines are slender and tapered, mostly 5 mm. to 7 mm. long, becoming still longer near the mouth. Large major pedicellariae, like those of the type, occur among the actinal adambulacral spines and along the inner edge of the grooves. These are long-ovate or lanceolate, with acute or subacute jaws, often toothed. With these there were others of various smaller sizes, of nearly the same forms. THE YOUNG. A small specimen, from Sitka, has the radii 10 mm. and 45 mm. ; ratio, i : 4.5. The dorsal spines are few and sparse, except along the median radial rows, where they form pretty close and nearly regular simple rows. A very imperfect, irregular row also occurs on each side, about midway between the median and superomarginal rows. Ten small groups of about two or three spines each form a central pentagon, with three central spines. Elsewhere dorsal spines are mostly lacking. All these spines are short, cylindrical or slightly clavate, with obtuse, sulcated tips. The dorsal ossicles are rather large and broad; even those that are without spines are often lozenge-shaped; those that bear spines are convex centrally, with a mammilla and pit. The superomarginal spines form a regular simple 96 VERRILL row ; they are smaller than the dorsals, but of the same form. The inferomarginal spines stand mostly two on a plate and with the per- actinals form three regular nearly equal rows. They are larger than the superomarginals and either cylindrical or slightly tapered, obtuse, finely sulcate. The peractinals are rather smaller than the rest and less obtuse. Adambulacrals form a single row; they are about as long as the actinals, but much more slender and evenly tapered. Three or four of the inner adoral plates, forming the compressed carinae, are with- out spines. A few large, ovate, unguiculate pedicellariae occur on the lateral channels and adambulcral plates. Some smaller acute-lanceolate ones occur on the inner edges of the grooves. Minor pedicellariae are rather thickly scattered everywhere over the dorsal and lateral integument. Another young specimen of larger size, with the radii 16 mm. and 82 mm., was sent by Dr. C. F. Newcombe. It was taken at Kuper Island, B. C., February, 1894. This agrees closely with the smaller one, described above, in the number and character of its dorsal spines, but the ventral spines are relatively stouter and form four rows proximally. The dorsal minor pedicellariae are much more numerous, both on the integument and around the spines, especially distally, where they form large dermal clusters. The papular areas are also very large, with numerous papulae. Thus it has in most respects the characters of the adult. Large major pedicellariae are rather numerous on the interradial areas and sides of the rays ; and many large, very acute, lanceolate ones occur on the inner edges of the grooves, sometimes in regular rows, one to a plate, for some distance. Most of these are sur- rounded at base by clusters of smaller major pedicellariae of about the same shape, but of various sizes. This specimen is remarkable for having two spines on some of the adambulacral plates, here and there, the second spine being smaller and attached back of the regular one. This condition has not been noticed in the other specimens. The adoral carinae are rather long and strongly compressed, de- scending abruptly to the mouth in all our specimens, and destitute of spines on three or four adoral plates. Apical peroral spines long, tapered, about equal to adorals, side-spines apparently lacking, or replaced by a cluster of pedicellariae; epioral spines lacking on our specimens. SHALLOW-WATER STARFISHES 97 This species ranges from Puget Sound to Sitka. Puget Sound (Professor Kincaid) ; Kuper Island and Victoria (C. F. New- combe) ; Sitka (Harriman Expedition, W. R. Coe). PISASTER GRAYI Verrill, nom. nov. Asterias katherina PERKIER, Steller. du Mus., Archiv Zool. Exper., iv, p. 331, 1875 (non Gray, 1840). Asterias dubia VERRILL, Amer. Naturalist, XLIII, p. 545, 1909 (non Clark). Perrier evidently erred in his determination of Gray's type of A. k&tkerin$*. robusta (Ludwig, as Stolasterias, p. 228), is from off the Galapagos, in 704 meters. The three species referred to Sporasterias do not seem to belong to that genus as herein restricted. S. mariana is from off Las Tres Marias Islands, in 1244 meters; 5. cocosana is from 245 meters, off Cocos Island ; S. galapagensis is from off Chatham Island, in 704 fathoms. The dorsal plates and connective ossicles are broad, imbricated, and have a regularly stichasterial arrangement in the last two species, which are otherwise very distinct. , S. mariana has five rows of dorsal plates and no visible interactinals. It may belong to Distolasterias. Hydrasterias diomedea is from 155 meters and 121 meters, off Panama and Cocos Island. None of these species are nearly allied to any of the shallow water species of California. The subfamily Asteriinae is again abundantly represented by shal- low-water species on both coasts of southern South America from Chile and northern Patagonia to Cape Horn and the Antarctic Islands. Yet nearly all the species of these regions belong to general not found in the North Pacific and North Atlantic. SHALLOW- WATER STARFISHES 351 VIII. PATAGONIAN, FUEGIAN, AND ANTARCTIC SHALLOW- WATER STARFISHES CONTRASTED WITH THOSE OF THE NORTH PACIFIC COAST.1 As already remarked, the Antarctic starfishes are abundant and diversified, especially those of the subfamily Asteriinae. Most of the latter, found in shallow water, belong to genera peculiar to the southern hemisphere. One notable peculiarity is found in the large proportion of species that carry and incubate their eggs and young, being brooders, or padophoric species. This peculiarity extends, also, to many of the Antarctic Echinoidea, Ophiuroidea, and Holothurioidea. The genera of starfishes of the family Asteriidse, having this habit, differ, also, in anatomical details and in the position of the genital pores from those that produce minute eggs that develop into free- swimming larvae, as is commonly the case with northern genera. Most of the pcedophoric Asteriidse carry their young in clusters under and around the mouth, as in Leptasterias of the north, but Stichaster nutrix Studer is described as carrying them in pouch-like diverticula of the stomach, at least in part. This p&dophoric habit is not confined to shallow-water and littoral species, for it has been observed in some species from considerable depths, such as Anasterias belgicce Ludwig, and A. chirophora Lud- wig, taken in 450 to 560 meters, in the Antarctic Ocean (see below). In these species the young are attached together in large groups, like clusters of grapes, by means of their adoral peduncles and a central stalk. Although many of the Antarctic species agree with Leptasterias of the north, in this habit of carrying their young, and some of them, also, in general appearance, they do not appear to be nearly allied.3 Most of them are regularly monacanthid, while Leptasterias is never truly monacanthid. Some are diplacanthid, as in the case of Podas- terias steineni and P. liitkenii (see below, p. 361). In the case of Leptychaster kerguelenensis Smith, the eggs and young are carried on the back, under the protection of the projecting spinules of the paxillse, which is more analogous to the method in Pterasteridse, in which the supraspinal membrane gives additional protection, and for a longer period of development. 1 In the following review I have omitted some imperfectly described forms and all the abyssal species. * It may be safely assumed, I believe, that all species that carry their eggs and young around the mouth, also have their genital pores on the ventral side, and therefore cannot be congeneric with those that have them dorsal, as in typical Asterias. 352 VERRILL Another striking peculiarity of the Asteriinae of this region, and of the Antarctic Ocean generally, is the large percentage of species that have the skeletal plates, and sometimes the spines, covered with a thick and soft, but rather tough, dermis, which is usually canalicu- lated, and often entirely conceals the plates in alcoholic preparations, or even in dry ones.1 Even the spines, if small, may be submerged in the dermis, or appear as mere papillae. In some cases such starfishes have the skeletal plates, more or less aborted, especially the dorsal ones, but sometimes even the upper marginals, as in Anasterias, Padasterias, and Adelasterias. FORCIPULOSA. No doubt the most peculiar and remarkable species, pertaining to these southern waters, is the Labidiaster radiosus Liitken, belonging to the Brisingidae. It has a small disk with from twenty-six to forty-two rays, the number increasing with age, and grows to a large size. Unlike the other forms of Brisingidae, it lives in shallow water, as well as at a considerable depths. It is found on both coasts of Patagonia and off Cape Horn, etc. The only other described species is L. annulatus Sladen. It oc- curred off Kerguelen Island and Heard Island, in 75 to 150 fathoms, and in the Arafura Sea ( ?) in 800 fathoms. It has forty to forty-five long, slender rays. It differs considerably in structure from L. radio- sus, especially in having the dorsal and superomarginal plates nearly abortive distally, on the rays, beyond the genital regions. This last character, with others, should require a generic separation. Therefore I propose for it the new generic name, Labidastrella, with L. annulata as its type. Aside from Labidiaster, the most characteristic species belong to the Asteriidae and Ganeriidae. There are several genera of Asteriidae that do not occur in northern seas, while most of the northern genera are entirely lacking. Thus there are no species of true Asterias, Pisaster, Orthasterias, Evasterias, Urasterias, Allasterias. It is doubtful if there is any true Leptasterias, though some antarctic 1 The apparent thickness of the dermis depends much on the mode of pres- ervation, which should be stated in descriptions. It is hardened and con- tracted by strong alcohol, and still more when dried, especially after being in alcohol. In alcohol of about 70 per cent it is more naturally preserved. In very weak alcohol and in formalin it may swell up and become soft. Some of the difficulties in the identification of such starfishes are thus due to the mode of preservation. In more cases it is due to the negligence of the describer in not cleaning and describing the skeletal plates and pedicellariae. SHALLOW-WATER STARFISHES 353 species have been described that closely resemble that genus, and may possibly belong to it. On the other hand, there are found in the Patagonian fauna, and farther south, numerous species and varieties of monacanthid star- fishes, most of which are known to carry their eggs and young (pado- phoric} and to have, in those species dissected, ventral genital pores. These have been referred to the genera Anasterias Perrier, 1875 ; Calvasterias Perrier, 1875; Sporasterias Perrier, 1894; Padasterias Verrill, gen. nov. Another group of pcedopkoric species includes several diplacanthid species, belonging to the genera Podasterias Perrier and Granaster Perrier. None of these are known in the northern hemisphere. The genera Meyenaster Verrill, May, 1913 (monacanthid), and Cosmasterias Sladen, 1889 (diplacanthid), also characteristic of this region, have not yet been observed to carry their young. The most common and abundant littoral and shallow-water star- fishes of both coasts of Terra del Fuego and Magellan Strait belong to a monacanthid generic group with a reticulated dorsal skeleton, for which Sporasterias Perrier, 1894, seems to be the earliest tenable name, with S. rugispina (Stimpson, 1862, as Asterias) for its type. This group, which we may also designate as the rugispina — or ant- arctica — group, has commonly been called Anasterias Perrier. The latter was originally based solely on A. minuta Perrier, a small, doubt- less very young starfish, with a thick dermis, rudimentary dorsal skeleton, and few dorsal spines, and thus quite unlike the group that we call Sporasterias. (See Anasterias, below, p. 354.) F. Leipoldt (op. cit., 1895, p. 563) proposed to unite nearly all the then known monacanthid starfishes of the Antarctic seas into one comprehensive and exceedingly variable species. This would be a very convenient arrangement, perhaps, if it would only prove true. No doubt, however, many of the nominal species of this group are synonymous ; and the species are certainly variable, as are all others that carry and incubate their eggs and young. Among the forms united by Leipoldt, under the name Asterias rugispina, the following belong apparently to Sporasterias, having a reticulated dorsal skeleton and many spines, not submerged in a thick dermis: Sporasterias antarctica (Lutken, 1856) ;S. rugispina (Stimp- son, 1862, as Asterias) ; S\ varia (Phil., 1870) ; S. rupicola (Verrill, 1876, as Asterias} ; S. perrieri (Smith, 1876, six-rayed, as Asterias, non A. perrieri Studer, 1884) ; S. cunninghami (Perrier, 1875, as Asterias) ; 5\ spirabilis (Bell, 1881, as Asterias) ; S. hyadesi (Per- 24 354 VERRILL rier, 1886, as Asterias). Of these, 5". rugispina, appears to be dis- tinct by reason of its large, wide, obtuse minor pedicellariae, and few, capitate and sulcate dorsal spines. S. perrieri Smith, from Kerguelen Island, is regularly six-rayed ; 5\ rupicola Verrill, from the same place, a smaller, five-rayed form, with many spines and few and minute pedicellarise, is apparently distinct. (See below, p. 356.) All the others may well belong to a single species, for which S. antarctica (Liitken) seems to be the correct name. ANASTERIAS (Perrier) restricted. In addition to all the above forms, Leipoldt includes in the same composite species the following, which have a rudimentary or partly abortive dorsal skeleton ; few small imbedded spinules ; and a thick dermis that conceals the plates, and largely the spinules. These constitute, with others, the restricted genus Anasterias (Lysasterias Fisher) : Anasterias ntinuta Perrier, 1875 (type) ; A. perrieri (Stu- der), 1884 (now Smith) ; A. verrillii (Bell, 1881, as Asterias). Also Calvasterias stolidota Sladen. Fisher (1908, p. 88) proposed the name Lysasterias to replace Anasterias Ludwig, with Asterias perrieri Studer, of South Georgia, as the type, assuming that A. minuta belongs to a distinct genus and is identical with S. rugispina, etc. However, the original description by Perrier shows that it has only isolated dorsal plates, covered with a thick dermis ; few imbedded dorsal spinules ; numerous pedicellarise, and most of the other characters as in A. perrieri Studer and A. studeri Perrier. Therefore, it is most likely the young of one of these or of some other closely allied species of the same group, and not the young of rugispina, nor of spirabilis, as has been thought, for the young of that group, when of similar size, have a reticulated dorsal skeleton, abundance of spines, and the dermis not thick enough to conceal the plates. Hence, I consider Lysasterias Fisher a synonym of Anasterias. Since the specific name, Asterias perrieri Studer (1884), was pre- occupied by A. perrieri Smith, 1876, it requires a new name. There- fore, I propose to call it ANASTERIAS LYSASTERIA. To the genus Anasterias (restr.) Ludwig (1905, p. 42) added, besides the two species above named, three new species from deeper water, southwest of Cape Horn. The described species are as follows : SHALLOW-WATER STARFISHES 355 A. rninuta Per., 1875. Patagonian and Fuegian; shallow-water. A. lysasteria Ver., nom. nov. = A. perrieri (Std. HOW Smith). South Georgia, cast ashore; also Fuegian (t. Perrier). A. studeri (Per.), 1891. Falkland Is., 320 meters ; carried young. A. verrillii (Bell, 1881, p. 513, pi. XLVII, fig. 3, as Astcrias). Fuegian ; Gregory Bay, etc., littoral. A. lactea Lud. (1903, p. 50). About S. lat. 71°, in 450 meters. A. belgicce Lud. (1903, p. 51). About S. lat. 70°, in 560 meters ; carried young. A. tenera (Koehler, 1905). Wandel I. ; carried young. The attached young of A. belgica were described by Ludwig, 1903, p. 53, pi. vi, figs. 61-65 ; pi. vii, figs. 66, 67. For the following species, placed in Anasterias by Ludwig, I pro- pose to establish a new genus : Genus Paedasterias Verrill, nov. Type A. chirophora (Ludwig, op. cit., 1903, p. 43, plates v, vi, vii). This is monacanthid and incubates its young, and therefore prob- ably has ventral genital pores. Its skeleton is more reduced than in Anasterias, the upper marginal plates being absent, except as rudi- ments distally. It is covered by a thick dermis and bears large, wide, felipedal pedicellarise. It is found at from 450 to 560 meters, in the Antarctic Ocean, south- west of Cape Horn, about S. lat. 70° to 71°. Ludwig has carefully described and figured the attached young. Genus Sporasterias Perrier. Type, Asterias rugispina Stimpson. Sporasterias PERRIER, 1896, p. 55. Anasterias (pars) PERRIER, 1891 (non 1875). Sporasterias LUDWIG, 1905. This is a monacanthid genus with a reticulated dorsal skeleton, not replaced by a thick dermis, bearing numerous small, scattered spines, and usually numerous pedicellarige of both kinds. Interactinal spines are usually absent; sometimes there is an imperfect row. Inferomarginal spines are usually two to a plate; superomarginal ones usually one to a plate ; interactinal papulae usually solitary, enlarged proximally; jaws and adoral ridge narrow, elongated. 356 VERRILL SPORASTERIAS PERRIERI (Smith). Asterias perrieri SMITH, Ann. and Mag. Nat. Hist., ser. 4, xvn, p. 106, 1876 (now Studer, 1884; non Anasterias perrieri Ludwig, 1905)- This is a true Sporasterias, with well developed skeleton and numer- ous spines. It is reguarly six-rayed. Smith states that his largest specimen carried hundreds of attached young, all of which were six- rayed. It appears to be clearly distinct from the rugispina-antarctica group. The type was from Kerguelen Island, in 5 to 10 fathoms. SPORASTERIAS RUPICOLA Verrill. Asterias rupicola VERRILL, op. cit., 1876, p. 74. This is a small, short-rayed form. The types, which were from Kerguelen Island shore, have the following characters when dry : The dorsal skeleton is pretty well developed, openly reticulated, with numerous moderately large papular areas, evenly distributed, larger on the basal part of the rays, mostly with two to four or five papulae; single ones occur distally. The dorsal plates are not con- cealed by the dermis ; the dorsal spines are evenly distributed, short, clavate, subequal, with no evident larger median row, one to three on the small nodal plates. The superomarginal spines form a distinct simple row, one to a plate. The plates being stout and imbricated. The inferomarginal plates mostly bear two larger and longer clavate spines. A wide lane between the upper and lower marginal rows, broadening proximally, has a row of large papular areas, with three to five papulae separated by stout descending apophyses of the superomarginal plates, some of which carry a single small spinule. A short interactinal row of small plates exists proximally, some of them bearing a single spine. Between these is a row of single papulae ; the proximal ones are often larger than the spines and inflated, as in other allied species. The adambulacral spines form a single regular row; they are smaller and more slender than the inferomarginals, and are distinctly clavate. The interradial axils are covered with a thicker canaliculated dermis without spines. The jaws are narrow. Major pedicellariae are few and small; some along the furrow margins and on the oral spines are thick, ovate or blunt-lanceolate ; a few smaller, more acute ones are on dorsal spines. Minor pedicellariae appear to be lacking in the several types. They are abundant in S. rugispina (Stimpson) and most of the other allied forms, and there are other notable differences in the spines and skeleton. SHALLOW-WATER STARFISHES 357 Genus Calvasterias Perrier. Type, C. asterinoidcs Perrier; Sladen, 1889, p. 589. Calvasterias PERRIER, op. cit., 1875, p. 348. This is another genus in which the skeleton, and also, largely, the spines, are covered with a thick, soft dermis, entirely concealing the plates. It is monacanthid. An imperfect spiniferous interactinal row of plates is present; usually there are two rows of inferomarginal spines and a single row of superomarginals. Position of genital pores unknown ; nor is it known whether the young are incubated. The dorsal skeleton in the type consists of wide, lobed or substellate, imbricated plates, overlapping in both directions by .their lobes, as in Stichaster. Descriptions and figures of other species do not show the character of the plates, and some may not be congeneric, if the plates prove to be abortive or rudimentary. In the type the dorsal spines are few, short, capitate, nearly con- cealed by the dermis and an abundance of papulae of similar size. The type was supposed to have been from Torres Strait, but Sladen records the species from the Falkland Islands. CALVASTERIAS (?) STOLIDOTA Sladen. C. stolidota SLADEN, 1889, p. 590, pi. ci, figs. 3, 4; pi. cm, figs, n, 12. This has longer rays than the type, with more evident, scattered dorsal spines, which are short, rough, capitate. The skeleton was not described, and so it may not be a true Calvasterias. Leipoldt united it to Sporasterias rugispina, with doubt. This is from both coasts of Patagonia and the Falkland Islands, in 5 to i o fathoms. MEYENASTER GELATINOSUS (Meyen, 1834, as Asterias). Asterias rustica GRAY, 1840, p. 179; Synopsis, p. i, 1866. Asterias gelatinosa CLARK, op. cit, 1910, p. 337, pi. vi, fig. 2. Meyenaster gelatinosus VERRILL, Amer. Journ. Science, xxxv, May, 1913, p. 348. See also Introduction, above, p. 54, for generic characters. This is a large, and a very characteristic species on the west coast of Chile, from Iquique southward. It grows to the diameter of 20 inches or more. It is monacanthid and usually six-rayed, yet five- rayed and seven-rayed specimens are frequently found. The only other species of the genus, known to me, is supposed to be from the Society Islands (No. 1427, Coll. Mus. Comp. Zool.). That 358 VERRILL is a large, five-rayed species, similar as to spinulation and thick dermis, but with very unlike pedicellariae. Its major pedicellariae are remarkably slender, numerous, part of them large, narrow-lanceolate and acute. The minor pedicellariae are minute and very abundant, both on the dermis and on thick sheaths around the sulcated dorsal spines. This genus somewhat resembles Marthasterias. Genus Cosmasterias Sladen, 1889. Type, Asterias sulcifera Perrier, 1869, = C. lurida (Phil.). Diplasterias (pars) PERKIER, 1891. This is also a genus apparently peculiar to the Patagonian and Fuegian region, where one species is abundant in shallow water and often at low tide. It is diplacanthid ; jaw-spines and adorals are elongated ; the dorsal plates form several longitudinal bands or rows, and in the adult each plate bears a group of short, unequal spines. There is at least one row of interactinal spines; often three or more rows, when adult. Large unguiculate or felipedal pedicellariae are found, above and below. Neither of the species has been observed to carry the young. COSMASTERIAS LURIDA (Phil., 1858) Ludwig, 1905. Cosmasterias sulcifera (PERRIER, 1869, as Asterias) Sladen, 1889. To this species Leipoldt, 1895, unites the following: Asteracanthion clavatutH Phil. ; A. fuhutn Phil. ; A. spectabile Phil. ; A. mite Phil. ; and Stichaster polygrammus Sladen, 1887, 245 fathoms. The species last named seems to me clearly distinct. (See below, p. 360.) To the synonyms of this species it will probably be necessary to add Asterias obtusispinosa Bell, 1881, p. 92, pi. ix, fig. 3 ; and A. neglecta Bell, op. cit., p. 94, pi. ix, fig. 4. Both are Patagonian ; the former from Sandy Point, 9 to 10 fathoms. C. alba (Bell), loc. cit., p. 92, pi. ix, fig. 2, also from Sandy Point, may not be distinct, though it has somewhat longer spines. C. sulcifera has been considered the type of Cosmasterias Sladen and Diplasterias Perrier. (See, also, Introduction, p. 48.) It is diplacanthid and has several (two or more) rows of inter- actinal plates ; its dorsal plates form unequal longitudinal rows, each plate, in the adults, bearing a transverse group of short, unequal spines, becoming numerous in large specimens. The minor pedicel- lariae are abundant around the spines and on the dermis, and there are many large, scattered major pedicellariae, some of them unguicu- late or dentate. SHALLOW-WATER STARFISHES 359 It varies much in appearance, according to age, and has many local or casual varieties. Whether all the forms referred to it by Leipoldt and others really belong together seems to me somewhat uncertain. I have studied a series of young and old from Eden Harbor, Pata- gonia, sent by the Museum of Comparative Zoology. These show wide variations, especially in the character of the dorsal rows of spines and the number of spines on each plate. The number increases rapidly with age, so that the larger specimens (radius, 175 mm.) have very numerous, crowded, transverse groups of spines and pedi- cellariae, the median and some of the others forming thick, wide, longi- tudinal rows. The madreporic plate is surrounded by a circle of small special spines, increasing in number with the age, from six or seven up to twenty or more. The major pedicellarise are large and stout. Some are obtusely lanceolate or ovate, others are blunt and unguiculate, with few teeth. They are abundant on the edge of the grooves, where many have long, slender pedicels, and between the ventral and lateral spines, and on the dorsal papular areas, where they are often stouter than the adja- cent spines. Minor pedicellariae are abundant on the dorsal dermis, crowded, especially over the transverse ossicles and between the spines. The larger specimens may have six to eight interactinal spines in a transverse row, arising, apparently, from three, to five, rows of plates. Young specimens, with the greater radius 50 mm., have only two rows of interactinal plates, each with one or two spines, and the spines on both marginal rows stand partly singly, partly two to a plate ; there are but three dorsal rows, and the spines in these stand partly in pairs, partly singly. This species occurs on both coasts of Patagonia and Fuegia, from low tide apparently to 348 fathoms. COSMASTERIAS TOMIDATA (Sladen). Cosmasterias tomidata SLADEN, 1889, p. 576. This has more regular rows of small, clustered dorsal spines, and very numerous dermal minor pedicellariae, larger than usual. The major pedicellariae are numerous, large and stout, often much larger than the spines ; mostly strongly felipedal. Gulf of Penas, 45 fathoms, Sladen ; Porto Lagunas, etc., 50 to 80 meters, Leipoldt ; off Argentina, S. lat. 44° 52', 55 fathoms, coll. Mus. Comp. Zoology. 360 VERRILL COSMASTERIAS POLYGRAMMUS (Sladen, 1889) Verrill. Stichaster polygrammus SLADEN, op. cit., 1889, p. 434, pis. c, cm. A diplacanthid species, easily distinguished by the conspicuous transverse or banded arrangement of the dorsal and lateral spines and plates. The dorsals and superomarginals form about seven longi- tudinal rows. The spines are small and numerous, arranged in nar- row transverse rows of three to seven on the ossicles. The major pedicellariae are large; some of them are felipedal. Papular areas are in regular rows. Ventral plates bear four or five spines in the transverse rows. Position of genital pores unknown. The type was from 245 fathoms, west of Magellan Strait. Leipoldt considered this a form of Cosmasterias sulcifera. The differences are notable. COSMASTERIAS BRANDTI (Bell) Verrill. Cosmasterias brandti (BELL, 1881, p. 91, pi. ix, fig. i, as Asterias). The dorsal and lateral plates are said to be about eleven in each transverse row, and to bear small spines, in transverse series. It may be the same as C. polygrammus. Trinidad Channel, at 30 fathoms. COSMASTERIAS FERNANDENSIS Meissner. Cosmasterias fernandensis MEISSNER, 1896, p. 104, pi. vi, fig. i, as Asterias. [Polyasterias] fernandensis de Loriol, op. cit., 1904, p. 41, pi. in, figs. 4-8 (rays unequal, five or six). This has been recorded by Loriol from Gulf San Mathias, E. Pata- gonia. The type was from Juan Fernandez. It appears to be auto- tomous, though often five-rayed. GASTRASTER STUDERI de Loriol. Gastraster studeri DE LORIOL, 1904, p. 34, pi. iv, figs. 3-3/. This has four irregular rows of podia proximally, but only two rows on the distal half of the rays. Its relations are obscure. It may be allied to Granaster biseriatus Koehler. From Gulf San Mathias. Genus Adelasterias Verrill, nov. Type, A. papillosa (Koehler, 1905, as Diplasterias). This is diplacanthid and is covered everywhere with a thick der- mis, which rises into numerous papilliform processes, both on the SHALLOW-WATER STARFISHES 361 back and on the sides, which enclose rudimentary or nearly abortive spinules. The skeleton is almost entirely abortive ; marginal plates are not present, unless as rudiments. The type is from the Antarctic Ocean. Genus Podasterias Perrier, 1896. Type, P. lutkeni Perrier. Diplasterias (pars) PERRIER, 1891, p. K, 77. Ludwig, Voy. Belgica, 1903. This genus is here extended to include several paedophoric dipla- canthid starfishes having a reticulated dorsal skeleton, not concealed by a thick dermis, and usually two or more rows of interactinal plates and spines. Thus they have a general resemblance to typical Asterias, like A. rubens. As some of the species are known to carry their young, the genital pores are presumed to be ventral, but I do not know that any species has been dissected with reference to this. Among the species that appear to belong here are the following : P. liitkeni (Perrier, 1891, p. K, 81, as Diplasterias). Fuegian and Falkland Is., 95 to 220 meters. Podasterias steineni (Studer, 1885, pi. i, figs. 40, b, as Asterias). Diplasterias steineni Perrier, 1891, p. K, 84. South of Cape Horn, 99 meters (Perrier). P. loveni (Perrier, 1891, p. K, 80, as Diplasterias). S. lat. 52° 44' 31", 320 meters. P. spinosa (Perrier, 1891, p. K, 82, as Diplasterias). S. lat. 47° 29', depth not recorded. P. meridionalis (Perrier, 1875, p. 340, as Asterias). Antarctic Expedition. Some of the several species, from the same region, described by Bell in 1881, probably also belong here, but the skeletal structure was not described. The first two species have been found carrying their young. The name Diplasterias Perrier is not adopted for this genus because, as originally defined, its only character was the existence of at least two rows of adambulacral spines. This made it synonymous with typical Asterias, which was the evident intention of Perrier, for he restricted Asterias to the monacanthid species. In the second place, his first species was A. sulcifera, which he evidently regarded as the type. But Sladen's report was published while his was being put in type. So that in an Appendix, p. K, 160, he admits the priority of Cosmasterias for sulcifera and abandons both Diplasterias and Podasterias (MS. name) and refers lutkeni (non Stimpson) and its congeners to Asterias. This will not hold 362 VERRILL good, for his liitkenii is pasdophoric. In a later work (1896, p. 35) he revived Podasterias, with P. lutkeni Perrier as the type. Fisher, 1908, p. 89, considered lutkeni Perrier a Pisaster, and sug- gested a change of name, owing to the priority of Pisaster liitkenii (Stimpson). They are not congeneric, the latter being monacanthid, and therefore no change is needed in the specific name. Genus Cryptasterias Verrill, nov. Type, Diplasterias turqueti Koehler, 1905, p. 465. This is diplacanthid and has a reticulated dorsal skeleton entirely concealed by a thick dermis. The dorsal surface bears soft dermal papillae, some of which bear a few pedicellariae ; others, scattered, con- tain rudimentary spinules. There is a simple row of superomarginal spines and a double row of inferomarginals ; apparently there are no interactinal plates. It is supposed to incubate its young. Antarctic. STICHASTER STRIATUS Miiller and Troschel, 1840. Asterias aurantiaca MEYEN, Reise urn die Erde, I, p. 292, 1834 (non Linne). Stichaster striatus MULLER and TROSCHEL, op. cit, 1840, p. 321 (non Asterias striatus Lam.). Stichaster aurantiacus VERRILL, 1867, p. 293. Clark, 1910, p. 337, pi. vm, fig. i. Tonia atlantica GRAY, op. cit., 1840, p. 180; Synopsis, 1866, p. 2. This is, perhaps, the most abundant littoral starfish found on the entire coast of Chile, south to Talcahuano, extending northward to Callao, Peru (Yale Museum, F. H. Bradley). Common at Valpa- raiso, on rocks. It grows to large size, up to 12 inches in diameter or more. It is the true type of the genus Stichaster, being the only species named when that genus was proposed in 1840. Perrier ( 1894, p. 131 ; 1896, p. 27) was in error in making Asterias rosea the type. (See also Introduction, p. 40.) It is not generically related to any North Pacific species. It is per- haps the only known species that can be properly referred to the genus Stichaster. Nearly all those referred to it by Sladen and others have already been separated generically,1 except, perhaps, S. polyplax of the Australian region. This should clearly be generically distinct. I would propose for it the name Allostichaster. 1 Perrier, 1894, proposed the genus Granaster for Stichaster nutrix Std. ; but G. biscriatus Koehler (antarctic) seems to be a distinct genus, as it has only two rows of podia. It may be called HEMIASTERIAS with biseriatus as the type. SHALLOW-WATER STARFISHES 363 Genus Allostichaster Verrill, nov. Type, A. polyplax Miiller'and Troschel. It is diplacanthid and multiple-rayed, with one to five madreporic plates, and is probably autotomous. The two rows of marginal plates are stout and imbricated ; dorsal plates and spines form five somewhat irregular longitudinal rows, several short, obtuse spines on each plate. Minor pedicellariae are dermal, usually not circumspinal. SPINULOSA. The Echinasteridse are well represented by Henricia, several species ; Poraniopsis, one species ; on the coast of Chile, by Echinas- ter; at the Falkland Islands, by Cribraster sladeni Perrier. Henricia seems to have several species and subspecies, and these vary much as do the northern species. H. obesa (Sladen, 1889) was described as from a depth of from 12 to 245 fathoms. Magellan Strait ; Falkland Islands. H. pagenstecheri (Studer) was originally from South Georgia. Shallow water ( ?). H. hyadesi (Perrier 1891, as Cribrella), Clark, 1910, p. 337, pi. n, fig. 5. This was recorded by Perrier from Fuegia, etc., and by Clark from Chile (ex. Meissner) ; 35 to 200 meters (Perrier). H. studeri (Perrier, 1891), from Magellan Straits, in 9 to 183 meters. Ludwig, 1905, seems to consider all these as variations of H. pagen- stocheri. To me most of them seem as distinct, if correctly described and figured, as some of them are from the corresponding Arctic forms. These species of Henricia are all pretty closely related to the northern and Arctic species, contrary to those of the Asteriidse. Cribraster is known only from the Falkland Islands; depth not known, probably littoral. C. sladeni Perrier, 1901 (p. K, 104, pi. xi, figs. 2a, 2b, and Appendix, p. K, 161). Echinaster antonioensis Lor., 1904, and E. lepidus Lor., 1904, have been described from the Gulf of San Mathias. E. lepidus is said to lack interradial papular pores, and in that case it belongs to Rhopia Gray, near R. sepositus, of the Mediterranean. The former species is related to E. brasiliensis. Poraniopsis echinaster Perrier, 1891 (p. K, 106, pi. x, figs. 2, 20, 95 meters). This is nearly related to P. inflata Fisher, from the North Pacific. Poraniopsis mira (=Lahillia mira de Loriol, op. cit., 1904, p. 33, pi. HI, figs, i-i/i). 364 VERRILL This is a large species with short, stout rays, from the Gulf San Mathias, E. Patagonia. Perknaster densus Sladen. Off Kerguelen Island; 127 fathoms. P. fuscus Sladen. Off Heard Island and Kerguelen Island ; 25 to 75 fathoms. The family Acanthasteridae, pertaining almost entirely to tropical seas, and always rather rare, is said to be represented by a species in the Magellan Straits, but the only record seems to be an ancient one and may not be reliable. It has not been found there by any recent expedition. Acanthaster Solaris (Schr.) Duj. et Hupe (pars) was described by Gray, 1866 (as Echinaster solans), from this region, but apparently without any modern record. According to him, it has twenty-one rays and ten madreporic plates. Duj. and Hupe united all the known species under this name. A. ellisii (Gray) occurs in the Panamic fauna. It usually has from eleven to thirteen rays. The oldest known species, A. planci (Linne, Syst. Nat., ed. X, Appendix, p. 823, as Asterias planci}, occurs apparently throughout most of the Indo-Pacific Oceans, in the tropical zone. The type was described by Linne as having fifteen rays. It came from Goa, Por- tuguese India, and the figure of Columna ( Phytobasanos, pi. xxxm, fig. A) was referred to as its basis. The figure is characteristic and the name should be adopted. It is the same as A. echinus (Ellis and Sol.) =A. echinites of Lam. and of many recent writers. The number of rays and madreporic plates is variable. Usually there are from fifteen to twenty very spinose rays. The family Asterinidse is unusually well represented. On the coast of Peru and southward Patiria chilensis (Liitken) and P. gayi (Perrier) are recorded. They are closely related and probably not dinstinct species; also, Patiriella calcarata (Perrier) and P. pusilla (Perrier). Desmopatiria fiexilis Verrill probably occurs on the same coast, but its origin is not positively known. Farther south, on the coasts of Chile and Magellan Strait, and around Tierra del Fuego to Cape Horn, the very common, small, lit- toral and shallow- water species is Patiriella, fimbriata (Perrier) or P. bispinosa (Perrier), if the latter, a supposed variety, proves to be distinct. P. fimbriata, as originally described, was labelled as from Bourbon Island, but Perrier himself expressed doubt as to whether the locality labels were correct, owing to the close agreement with specimens from Chile. SHALLOW-WATER STARFISHES 365 The type had mostly a single spine on the ventral plates, while the Patagonian form usually has two spines on many of the plates. There are, also, some differences in the dorsal spinulation, so that they may prove to be distinct; yet I am not aware that it has been found in tropical seas by later collectors, and the ancient Bourbon Island label may have been wrong. I have examined numerous specimens from Magellan Strait, and found but little variation, but have seen no authentic tropical specimens. It has a thicker dermis than usual, and is not a typical Patirietla. P. calcarata and P. pusilla are, however, more nearly allied to sev- eral African and Australian species than to any in the northern hemi- sphere. Asterinides perrieri=Asterina perrieri de Loriol, 1904, a small species from E. Patagonia, Gulf of San Mathias, appears to belong to the genus Asterinides, but may be closely related to P. fimbriata or bispinosa. Enoplopatiria marginata, which has been recorded from near the eastern end of Magellan Strait, is a common Brazilian species, extend- ing northward to the West Indies, and recorded also from the west coast of Africa. A closely related new species, E. siderea, in our col- lection, is labeled as from Panama (donor, Captain Dow, of the Panama and California steamship line). E. siderea Verrill is a rather large species, radii, 24 mm. and 48 mm., with stout, short, broad rays and well defined papular areas, with numerous forked dorsal pedicellarias ; ventral plates with a comb of three elongated spines ; four webbed furrow spines. It differs from E. marginata in having the spinules of the inter- radial dorsal plates longer, widely divergent, stellate and paxillif orm, and in other characteristics ; but it is evidently closely related, so much so that I formerly suspected the correctness of the label, thinking that it might have come from Colon, on the Atlantic side, and might be only a large variety of E. marginata. See plate cix, figure 4. The family Ganeriidae is very characteristic of the Patagonian- Fuegian fauna. Indeed, nearly all the known genera and species are from that region and adjacent districts. Of the genus Ganeria Gray, 1847, Perrier, 1891, recognized four species; viz.: G. falklandica Gray, 1847, Falkland Islands; east of Magellan Strait, 55 fathoms, Sladen; G. robusta Perrier, 1891, 28 meters; G. hahni Perrier, 1891, 138 meters ; G. papillosa Perrier, 1891, depth not given. 366 VERRILL The genus Lebrunaster Perrier, 1891, p. K, 116, type L. paxillosus Perrier, from Patagonia, depth unknown, is known only from this fauna. Numerous nominal species of Cycethra have been described from Magellan Strait and the coasts of Patagonia and Terra del Fuego, where the genus is common and variable. Ludwig, 1905, reduces them mostly to varieties of C. verrucosa (Phil.). This species includes C. nitida Sladen, Meissner -f- C. elec- tilis Sladen, Leipoldt -f- C- simplex Bell, Meissner -f- C. pinguis Sladen -f- C. ganeriodes Perrier -j- C. elongata Perrier -}- C. media Perrier + C. asterina Perrier+C. subelectillis Perrier +C. calva Per- rier -f- C. regularis Perrier -j- C. astcriscus Perrier. The last eight forms were regarded by Perrier himself as mere varieties of C. sim- plex, as well as some of the others. They are common in Magellan Strait and southward to Cape Horn, from low tide to 40 fathoms. Cycethra lahillei de Loriol, 1904, p. 25, pi. II, fig. 2, from Gulf of San Mathias, appears to be a distinct species, with short rays, imperfectly developed marginal plates, and slender paxilliform spinules. One of the generic types, common to this region and the North Pacific, is Peribolaster, of the family Korethrasteridae, of which only two species are known. P. foliculata Sladen occurred in 45 fathoms, west of Patagonia. P. biserialis Fisher, 1911, ranges from Cali- fornia to Bering Sea, in from 57 to 313 fathoms. The family Solasteridae is represented in moderate depths by Solaster australls (Perrier) and Lophaster stellans Sladen, both of which have occurred in from 30 to 40 fathoms. In greater depths several additional species of those genera occur. (See list of deeper- water species below, p. 368.) These are genera widely distributed in all seas, especially in rather deep water, coming into shallow water only in the colder regions. Some of the Antarctic species are closely allied to those of the North Pacific and Arctic Ocean, like those of other families from deep water. The Pterasteridse are also well represented by several species of Pteraster and Diplopteraster in waters of somewhat greater depths. These are also allied to the corresponding species of the North Atlantic and North Pacific. (See list of deeper-water species below, p. 368.) The genus Retaster, which is represented by two species, does not occur in northern waters, but is found in the Indo-Pacific region, from Cape Good Hope to Australia. The Patagonian species are as follows : SHALLOW-WATER STARFISHES 367 R. gibber Sladen, 1889; Ludwig, 1905, pi. v, figs. 6, 7. This occurred in from 14 to 245 fathoms, in Magellan Strait. R. verrucosus Sladen. This occurred in 55 fathoms, in Magellan Strait. PHANEROZONA. The family Odontasteridae is better represented in the Fuegian fauna than in most regions, some species occurring commonly even at low tide and in shallow water. Several nominal species have been described, some of which Ludwig (1905) and others reduce to syno- nyms. Ludwig recognizes three shallow-water species ; viz. : 1. O. grayi (Bell, non Perrier) = Gnathaster grayi Sladen — O. pedicellaris (Perrier). 2. O. penicillatus (Phil.) + O. pilulatus (Sladen, as Gnathaster) -j- O. rneridionalis (Smith). Leipoldt, 1895. 3. Asterodon singularis (Miiller and Troschel) Leipoldt, 1895, pi. xxxi, figs. ?a-c, as Odontaster = A. granulosus Perrier, 1891, pi. xi, figs. 40, b. A. belli (Studer) Perrier is probably not distinct from A. singularis. The first species is from a depth of 10 to 97 meters. The second species ranges from 5 to 150 fathoms. Several other antarctic species have been described from deeper water. (See Sladen ; also Koehler, 19050 Luidia magellancia Leipoldt, 1895, p. 610, pi. xxxn, figs. na-c. This is remarkable as inhabiting cold waters, for all the other numer- ous species inhabit tropical or temperate seas. The type had the radii 208 m. and 35 m; ratio, 1 : 6. Its nearest related species are L. bellonce Liitken, which ranges from Chile to Mexico, and L. phragma Clark (1910, p. 329, pi. n, fig. i), which was supposed to be from Peru. This may, therefore, be considered as one of the few that were probably of Panamic origin. IX. LIST OF PATAGONIAN AND FUEGIAN1 STARFISHES FOUND IN FROM 50 TO 500 FATHOMS, MOSTLY RELATED TO NORTHERN OR ARCTIC SPECIES. In the somewhat deep, cold waters on the " continental slopes " (50 to 500 fathoms), a considerable number of genera and species have been found that are closely related to those of the North Pacific and North Atlantic and the Arctic Ocean. Many of these have been recorded by Sladen, 1889; Perrier, 1891; Leipoldt, 1895; Ludwig, 1905; Kcehler, 1905, and others. Among them are the following: 1 Some species from Kerguelen Island, etc., are here included, but some arc common to the two regions. 368 VERRILL Coscinasterias candicans (Lud., 1905, as Stolasterias). S. lat. 70° to 71° ; 450 to 560 meters. Pedicellaster antarcticus Lud., 1905. 430 to 450 meters. P. scaber (Smith) Perrier. Kerguelen I., etc. Henricia simplex (Sladen, 1889, as CribreUa). Antarctic ; 50 to 310 fathoms. H. prastans (Sladen, as Cribrella). 210 fathoms. H: hyadesi (Perrier, 1891). 35 to 200 fathoms. Echinaster smithii Ludwig. 450 meters. Solaster octoradiatus Ludwig. 450 to 500 meters; S. lat. 70° to 71°. Solaster subarcuatus Sladen. 150 fathoms. Solaster regularis Sladen. 175 fathoms. S. australis (Perrier). 65 to 198 meters. Lophaster stellans Sladen. 40 to 1325 fathoms. Includes L. pentactis Per., 200 meters. Pteraster stellifer Sladen. 245 fathoms. P. rugatus Sladen. 150 fathoms. P. ingouffi Per. Fuegian; 270 meters. P. lebruni Per. Fuegian; 80 meters; 450 meters, Ludwig; S. lat. 71° 24'. Diplopteraster verrucosus (Sladen, as Retaster). East of Magellan Strait ; 55 fathoms. D. peregrinator (Sladen, as Retaster). Kerguelen I. ; 127 fathoms. Hymenaster perspicuus Lud. 400 to 450 meters. Porania antarctica Smith. Kerguelen I. ; South Georgia ; Magellan Strait ; Antarctic Ocean, S. lat. 71° 18', 450 meters; Fuegian, 10 to 1,600 fathoms. Includes P. mo- gelhcenica Studer and P. patagonica Perrier. P. glaber Sladen. 30 to 127 fathoms. Odontaster cremens Ludwig. 450 meters. Mimaster cognatus Sladen. 245 to 1325 fathoms ; 450 meters, S. lat. 70° 23', Ludwig. Pseudarchaster patagonicus (Per., 1891, p. K, 125, pi. xm, figs. 20, zb, as Astro gonium). 283 meters. SHALLOW-WATER STARFISHES 369 Hippasteria hyadesi Per. 326 meters. Ceramaster patagonicus (Sladen). 55 to 245 fathoms. C. austro-granularis (Per., 1891, as Pentagonaster). 340 meters. Leptychaster kerguelenensis Smith. Kerguelen and Marion Islands; 10 to 100 fathoms. L. antarcticus Sladen. 210 fathoms. Psilaster fteuriaisi (Per., 1891, as Goniopecten). 198 to 283 meters. Cheiraster gerlachei Lud. 450 to 560 meters. Pontaster planeta Sladen. 245 fathoms. Ctenodiscus australis Liitk. 55 to 600 fathoms. C. procurator Sladen. 40 to 1325 fathoms. All the species (thirty-four) included in this list are generically allied to northern or arctic species. In many cases the resemblance is very close specifically. This is so especially in the cases of some species of Henricia, Solaster, Pteraster, Diplopteraster, Porania, Hippasteria, Ceramaster, and Ctenodiscus. These facts clearly indicate that extensive migrations of the ances- tors of these species have occurred, in the colder deep waters, in one or both directions, at no very remote geologic periods. It appears, with our present knowledge of oceanic currents and temperatures, that the greater part of the migration would have been from the Arctic Ocean southward to the Antarctic. But no barriers exist, at the depths that these species inhabit, to prevent a northward migration, for abyssal species are not included in the list. The shal- low-water and littoral species, on the other hand, show no evidence of such comparatively recent migrations, for much the larger part are peculiar to this region and the adjacent Antarctic seas and islands, with a few additions from the Indo-Pacific region. Scarcely any of the strictly shallow-water species seem to have been able to cross the tropical regions, on either side of America, unless in remote geologic time. Those that can descend to the depths where very cold water exists, even in tropical seas, have been able to migrate from polar seas across the tropics, probably in both directions, no doubt with extreme slow- ness, and yet with but little change in appearance or structure. Such 25 37O VERRILL migrations have probably been going on during vast geologic periods and, no doubt, are still in progress. Many of the existing genera of starfishes are doubtless of great antiquity, geologically speaking, and show wonderful persistence in many of the details of structure and habit. X. LISTS OF EXTRALIMITAL SPECIES PARTIALLY DESCRIBED, REVISED, OR FIGURED. Many other extralimital species, merely included in the faunal lists, or only mentioned by name under their genera, are not included in these lists. I. NORTH ATLANTIC AND ARCTIC SPECIES. Asterias vulgaris Ver. New England. Asterias forbesi Desor. New England, etc. A. acervata borealis = A. polaris. Arctic and North Atlantic. Orthasterias tanneri Ver. Off East coast United States. PI. XLVIII, fig. 6. O. subangulosa Ver. (nom. nov.) —A. angulosa Per., preoccupied. West Indies. Stylasterias neglecta (Per. non Bell). Coscinasterias tenuispina (Lam.) Ver. Europe; Bermuda. Marthasterias glacialis (Mull.) Jul. Europe, etc. Sclerasterias guernei Per. East Atlantic. Leptasterias littoralis (Stimp.) Ver. New England. L. compta (Stimp.) Ver. New England. L. tenera (Stimp.) Ver. New England. Ctenasterias spitzbergensis (D. and Kor.)- Arctic. Urasterias linckii (M. and Tr.) Ver. Nova Scotia ; Arctic Ocean ; post-glacial fossil at Salem, Mass. PL LXX, figs. 1-4. U. panopla (Stuxb.) Ver. Arctic. Stichastrella rosea (Mull.) Ver. Europe. Coronastcr briareus Ver. West Atlantic. SHALLOW-WATER STARFISHES 371 Henricia sanguinolenta miliaris Ver. New England. H. antillarum (Per.). West Indies. H. sexradiata (Per.) Ver. West Indies. Crossaster korenii Ver. (nom. nov.). Arctic and North Atlantic. Asterina gibbosa (Penn.) Gray. Europe. A. pygmaa Ver. = Porania. (Young). New England. Asterinides -folium (Liitk.) Ver. West Indies; Bermuda. Enoplopatiria marginata (Hupe) Ver. West Indies ; Brazil ; West Africa, etc. Tremaster mirabilis Ver. Newfoundland Banks, etc. Hippasteria phrygiana (Parel.) Ag. New England; Europe; Arctic Ocean. PI. xLvn, fig. i; pi. XLVIII, figs. i-S ; pi. XLIX, fig. 6. Mediaster bairdii Ver. West Atlantic. PI. n, fig. 2; pi. in, fig. 2 (type). Blakiaster conicus Per. West Indies. Luidia clathrata (Say). Southern coast United States; Bermuda; West Indies, etc. PL cm, fig. i. 2. PATAGONIAN, FUEGIAN, ANTARCTIC, AND SOUTH ATLANTIC SPECIES. Labidiaster radiosus Liitk. Pedicellaster antarcticus Lud. Sporasterias rugispina (Stimp.) Per. S. antarctica (Lutk.) Per. 5". perrieri (Smith). 5". rupicola (Ver.). Anasterias minuta Per. A. lysasteria (Ver.), nom. nov. — A. perrieri Std. (non Per.). A. belgica Ludwig. A. studeri (Perrier). A. tenera Koehler. A. verrillii (Bell). Padasterias (gen. nov.) chirophora (Lud.) Ver. Calvasterias asterinoides Per. C. (?) stolidota Sladen. M eyenaster gelatinosus (Meyen) Ver. Smilasterias scalprifera Sladen. Cosmasterias lurida (Phil.) Lud. Includes C. sulcifera, etc. C. tomidata Sladen. C. polygrammus (Sladen, as Stichaster). 372 VERRILL C. brandti (Bell, as Asterias) Ver. C. fernandensis (Meiss., as Polyasterias) Ver. Gastraster studeri Kcehler. Adelasterias (gen. nov.) papillosa (Koehler, 1905, as Asterias). Podasterias lutkeni Per. P. meridionalis (Perrier, as Asterias). P. spinosa ( Perrier, as Diplasterias) . P. steineni Per. Cryptasterias (gen. nov.) turqueti (Koehler, as Asterias} Ver. Antarctic. Granaster nutrix (Studer) Per. S. Georgia. Hemiasterias biseriatus (Koehler, as Granaster) Antarctic. Stichaster striatus (M. and Tr.) = Asterias aurantiaca Meyen (non Linne). Peru and Chile. Heliaster helianthus (Lam.) D. and H. Ecuador to Chile. Henricia obesa (Sladen). H. pagenstecheri (Studer). H. hyadesi (Per.) Clark. Cribraster sladeni Per. Echinaster antonioensis Kcehler. E. (Rhopia) lepidus (Kcehler). Poraniopsis echinaster Per. P. mira (Koehler, as Lahillia). Acanthaster Solaris (Schr.) D. and Hupe. Panamic, etc. Patina chilensis (Liitk.) Ver. P. gayi (Per.) Ver. Patiriella calcarata (Per.) Ver. P. pusilla (Per.) Ver. P. fimbriata (Per.) Ver. P. bispinosa (Per.) Ver. Asterinides perrieri (Kcehler, as Asterina) Ver. Desmopatiria fiexilis Ver. Porania antarctica Smith. Ganeria falklandica Gray. G. papillosa Per. Lebrunaster paxillosus Per. Cycethra verrucosa (Phil.). Includes C. simplex Bell; C. electilis Sladen, etc. C. lahillei Koehler. Peribolaster foliculata Sladen. Solaster australis (Per.) Lophaster stellans Sladen. Retaster gibber Sladen. R. verrucosus Sladen. Odontaster grayi (Bell). O. penicillatus (Phil.). Includes O. pilulatus Sladen. Aster odon singularis (M. and Tr.). Includes O. granulosus Per. Leptychaster kerguelenensis Smith. Luidia magellanica Leipoldt. Astropectenides (gen. nov.) mesacutus (Sladen). South Atlantic. SHALLOW-WATER STARFISHES 373 3. SPECIES FROM THE PANAMIC FAUNA. Coscinasterias alexandri (Ludwig, as Stolasterias}. Stylasterias robusta (Ludwig, as Stolasterias). Acanthaster ellisii (Gray). Mithrodia bradleyi Ver. PI. cvm, fig. i (type). Enoplopatiria siderea Ver. PI. cix, fig. 4. Echinaster tenuispinus Ver. PI. evil, fig. 2, type. Asterinides modesta Ver. Callopatiria obtusa (Gray) Ver. Amphiaster insignis Ver. PI. cvm, fig. I (type). Astropecten cerstedii Gray. A. ormolus Gray. 4. INDO-PACIFIC, AUSTRALIAN, AND JAPANESE SPECIES. Labidastrella (gen. nov.) annulata (Sladen, as Labidiaster) . Australasian; off Heard Island and Kerguelen Island, in 75-150 fathoms. Heterasterias (gen. nov.) volsellata (Sladen, as Asterias}. Japan. Coscinasterias acutispina (Stimp.). Japan. C. muricata Ver. New Zealand. Allostichaster (gen. nov.) polyplax (M. and Tr.) Ver. Australasian. Distolasterias stichantha (Sladen) Ver. Japan. Allasterias forficulosa Ver. Japan. PI. LXXXIII, figs. 3-3^ ; pi. LXXXIV, fig. I. A. versicolor (Sladen) Ver. Japan. A. amurensis (Liitk.) Ver. Siberia. A. migrata (Sladen, 1878, as Asteracanthion rubensvar.). Straits of Korea. Ccelasterias australis Ver. New Zealand. Acanthaster planci (Linne)=/4. echinites of authors. Indo-Pacific. Henricides (gen. nov.) heteractis (Clark, as Henricid). Lord Howe I. ; Australasian. Henricia densispina (Sladen, 1878). Straits of Corea ; 40 fathoms. H. japonica Ver. Japan. Patiria coccinea Gray. Indo-Pacific. Asterinopsis penicillaris (Lam.) Ver. Indo-Pacific ; Australasian. Patiriella regularis Ver. New Zealand. 374 VERRILL XI. LIST OF NEW GENERA ESTABLISHED AND DESCRIBED. Brisingidae : Labidastrella. Type Labidiaster annulata Sladen. Asteriidae : Evasterias. Type, E. troschelii (Stimpson). Orthasterias. Type, O. columbiana Verrill. Stylasterias. Type, O. forreri (de Loriol). Heterasterias. Type, H. volsellata (Sladen). Stenasterias. Type, 5". macropora Ver. Ctenasterias. Type, C. spitsbergensis (Danielssen and Koren). Tosiaster. Type, T. arcticus Ver. Allostichaster. Type, A. polyplax (Miiller and Troschel). Pcedasterias. Type, P. chirophora (Ludwig). Adelasterias. Type, A. papillosa (Kcehler). Cryptasterias. Type, C. turqueti (Koehler). Parasterias. Type, P. albertensis Verrill. Stichastrella. Type, 5". rosea (Miiller). Hemiasterias. Type, H. biseriatus (Koehler, as Granaster}. Echinasteridae : Henricides. Type, H. heteractis (Clark). Astropectinidae: Astropectinides. Type, A. mesacutus (Sladen). BIBLIOGRAPHY OF STARFISHES OF NORTHWEST COAST, AND OTHER WORKS REFERRED TO. As a rule, general works and text-books are not included in this list, unless specially referred to in the text. AGASSIZ, ALEXANDER and Mrs. E. C. Sea Side Studies in Natural History, Boston, 1865. AGASSIZ, ALEXANDER. North American Starfishes. Memoirs Museum Com- parative Zoology, Harvard College, vol. v, no. I, 4°, 136 pp., 20 pis., Cam- bridge, Mass., 1877. AGASSIZ, L. Prodrome d'une Monographic des Radiares ou Echinodermes. Mem. Soc. Sci. Nat. Neuchatel, vol. i, pp. 168-199, 1835. BAKER, C. F. Some Echinoderms Collected at Laguna. First Annual Report of Laguna .Marine Laboratory, California. No. 5. BATHER, F. A. What is an Echinoderm? J. Lond. Coll. Soc., vol. vin, pp 22, 23, text cuts, igoi. A discussion of the theories of the primitive evolution of the different classes. BELL, F. JEFFREY. Note on Abnormal (Quadriradiate) Specimen of Ambly- pneustes formosus. Linn. Soc. Journ. (Zool.), vol. xv, pp. 126-129, pi- VF 1880. . Contributions to the Systematic Arrangement of the Asteroidea; part I, The Species of the Genus Asterias. Proc. Zool. Soc. London, pp. 492-515, pis. XLVII, XLVIII, 18810. SHALLOW- WATER STARFISHES 375 — . Account of the Zoological Collections Made During the Survey of H. M. S. "Alert" in the Straits of Magellan and on the Coast of Patagonia. Echinodermata. Proc. Zool. Soc. London, 1881, pp. 87-101, pis. vni, ix, i88i&. — . Descriptions of new or rare Species of Asteroidea in Coll. Brit. Museum. Proc. Zool. Soc. London, pp. 121-124, pi- vi, 1882. — . On the Generic Name of Asterias sanguinolenta, O. F. Miiller. Ann. and Mag. Nat. Hist., ser. 6, vol. vi, pp. 472-473, 1890. — . Catalogue of the British Echinoderms in the British Museum. Lon- don, pp. 202, 16 pis., 1892. — . On Odontaster and the Allied or Synonymous Genera of Asteroid Echinoderms. Proc. Zool. Soc., pp. 259-262, London, 1893. Echinoderma. Report on the Collections of Natural History made in the Antarctic Regions during the Voyage of the Southern Cross. Lon- don, 1902, pp. 214-220, pis. xxvi-xxvin. BRANDT, J. F. Prodromus descriptions animalium ab H. Mertensio in orbis terrarum circumnavigatione observatorum, Fasc. I, 4°, 72 pp., St. Peters- burg, 1835. In Recueil des Actes Acad. Imp. St. Petersburg, p. 199, 1834. . Bemerkungen iiber die Asteriden und Echiniden des Ochotskischen, Kamtschatkischen und Behringschen Meeres. In A. Th. v. Middendorff, Reise in den aussersten Norden und Osten Sibiriens, Bd. II, Teil I : Wir- bellose Thiere, pp. 27-34, St. Petersburg, 1851. BUSH, KATHERINE J. Catalogue of Mollusca and Echinodermata Dredged on the Coast of Labrador by the Expedition under the Direction of Mr. W. A. Stearns, in 1882. In Proc. U. S. Nat. Mus. vol. vi (1883), pp. 236- 247 (Echinodermata, pp. 245-247), 9 pis., Washington, 1884. CLARK, HUBERT LYMAN. Echinoderms from Puget Sound. Proc. Boston Soc. Nat. Hist., vol. xxix, no. 15, pp. 323-377, 4 pis., 1901. — . Papers from the Hopkins Stanford Galapagos Expedition, 1898-99, vol. xn, Echinodermata. In Proc. Washington Acad. Sci., vol. rv, pp. 521-531, 1902. . The Echinoderms of the Woods Holl Region. Bull. U. S. Fish Comm., vol. xxn, pp. 547-576. pis. i-xiv, 1904. — . Starfishes of the Genus Heliaster. Mus. Comp. Zool. Bull., vol. LI, pp. 27-76, pis. i-vin, 1907. On pi. vi, fig. 3, Asterias (Pisaster) ochracea is figured with inter- radial partitions exposed. . Scientific Results of the Trawling Expedition of the Thetis. Mem. Austral. Mus., vol. iv, part 2, 1909. The Echinoderms of Peru. Bull. Mus. Comp. Zool., vol. LJI, no. 17, pp. 321-358, pis. i-xiv, 1910. CU£NOT, L. Contribution a 1'Etude Anatomique des Asterides. In Arch. Zool. Exper. et Gener. (2), vol. v, bis, 2, Mem., 144 pp., 9 pis., 1888. CROCKER and MONKS. See Ritter, W. E., 1902. DANIELSSEN, D. C, and KOREN J. Fra den norske Nordhavsexpedition, i. In Nyt Magazin for Naturvidenskaberne, Bd. xxm, 39 pp., 5 pis., Chris- tiania, 1876. — . The same, vol. xxvi, pp. 177-194, 2 pis., 1880. (Asterias spits- bergensis. etc.) VERRILL . The same, vol. xxvn, pp. 267-299, 4 pis., 1882. . The same, vol. xxvm, pp. i-io, 2 pis., 1883. . Asteroidea. Norwegian North Atlantic Expedition, 1876-78. Zoology, 4°, 118 pp., 15 pis., map, 1884. DESOR, EDOUARD. Zoological Investigations among the Shoals of Nantucket. In Proc. Boston Soc. Nat. Hist., vol. in, for 1848-51, pp. n, 17, 65-68, Cam- bridge, 1851. Records egg-carrying habits of Henricia. DODERLEIN, LUDWIG. Einige Beobachtungen an arktischen Seesternen. In Zool. Anzeiger, Bd. xxn, pp. 337-339- 1899. . Die Echinodermen der Olga-Expedition. Wiss. Meeresuntersuch. (N. F.) iv. Abteilung, Helgoland, Heft 2, pp. 197-224, pis. iv-x, 1900. Excellent plates of arctic Asteriidae, from photographs. DUBEN, M. W., and KOREN, J. Oefversigt af Skandinaviens Echinodermer. In Kongl. Vetensk. Akad. Handlingar for 1844, pp. 229-328, pis. vi-xi, Stockholm, 1846. DUJARDIN ET HUPIE. Historic nat. des Zoophytes, Echinodermes, in Suites a Buffon. (Largely a translation of Miiller and Troschel, Syst. Aster.) 1862. DUNCAN, P. MARTIN, and SLADEN, W. PERCY. Report on the Echinodermata Collected during the Arctic Expedition, 1875-76. In Ann. Mag. Nat. Hist. (4), vol. xx, pp. 449-470, 1877. . Echinodermata. In G. S. Nares, Narrative of a Voyage to the Polar Sea during 1875-76, vol. n, pp. 260-282, London, 1878. . A Memoir on the Echinodermata of the Arctic Sea to the West of Greenland, 82 pp., 6 pis., London, 1881. D'URBAN, W. S. M. The Zoology of Barents Sea. In Ann. Mag. Nat. Hist. (5), vol. vi, pp. 253-277, London, 1880. FABRICIUS, OTHO. Fauna Groenlandica, 8°, 452 pp., Hafnias et Lipsise, 1780. FEWKES, J. WALTER. On the Development of the Calcareous Plates of Asterias. Bull. Mus. Comp. Zool., vol. xvii, pp. 1-56, pis. i-v, Jan., 18880. . On the Serial Relationship of the Ambulacral and Adambulacral Calcareous Plates of the Starfishes. Proc. Boston Soc. Nat. Hist., vol. xxrv, pp. 96-115, text cuts, i888&. Contains an account of the Development of Asterina gibbosa. . An Aid to a Collector of the Coelenterata and Echinodermata of New England. In Bull. Essex Inst., vol. xxm, 92 pp., with figures, Salem, Mass., 1891. FISCHER, F. Echinodermen von Jan Mayen. In Die Internationale Polarfor- schung, 1882-83; Die osterreichische Polarstation, Jan Mayen, Bd. in, 4°, 10 pp., Wien, 1886. FISHER, WALTER K. New Starfishes from Deep Water off California and Alaska. Bull. Bureau Fisheries for 1904, vol. iv, June 10, pp. 291-320, 1905. . Two New Starfishes from Monterey Bay, California. Zoologischer Anzeiger, vol. xxx, no. 10, June 19, pp. 229-302, 19060. . Note on Eremicaster, a Genus of Starfishes. Zoologischer Anzeiger, vol. xxxn, no. I, July 23, pp. 12-14, 1907. . Necessary Changes in the Nomenclature of Starfishes. Smithsonian Miscell. Collections, vol. LII, pp. 87-93, 19080. . Some Necessary Changes in the Generic Names of Starfishes. Zoologischer Anzeiger, vol. xxxni, no. n, August 18, pp. 356-359, ioo8&. SHALLOW-WATER STARFISHES 377 — . New Pterasteridae from the North Pacific. Ann. and Mag. Nat. Hist, ser. 8, voL v, February, pp. 167-170, 19100. — . New Genera of Starfishes. Ann. and Mag. Nat. Hist., ser. 8, vol. v, February, pp. 171-173, — . New Starfishes from the North Pacific; I, Phanerozonia. Zoolo- gischer Anzeiger, vol. xxxv, March 29, pp. 546-553, I9ioc. — . New Starfishes from the North Pacific; II, Spinulosa. Zoologischer Anzeiger, vol. xxxv, March 29, pp. 568-574, 19100*. — . The Genus Blakiaster. Bull. Mus. Comp. Zool., vol. LIV, no. 4, pp. 161-165, pis. i, n, 19110. Discusses also Leptychaster and Bunodaster. — . Asteroidea of the North Pacific and adjacent waters. Bull. U. S. Nat. Mus. No. 76, Part I. Phanerozonia and Spinulosa. 4°, 419 pp., 122 pis., Washington, 191 ib. FORBES, EDWARD. On the Asteriadae of the Irish Sea. Memoirs of the Wer- nerian Society, Edinburgh, vol. vm, part i, pp. 114-129, 2 pis., 1839. . A History of British Starfishes and other Animals of the Class Echinodermata, 8°, 269 pp., London, 1841. Published in six monthly numbers, the first of each month, from October, 1840, to March, 1841 (teste Bell, op. cit., 1890, p. 493). 1840-41. Notes on Animals of the Class Echinodermata. In Peter C. Suth- erland's Journal of a Voyage in Baffin Bay and Barrow Straits in the Years 1850-51, vol. n, 8°, Appendix, pp. ccxiv-ccxvi, London, 1852. GANONG, W. F. The Echinodermata of New Brunswick, Bull. no. 7, Nat. Hist Soc. New Brunswick, pp. 1-57, i pi., St. Johns, 1880. GRAY, JOHN EDWARD. A Synopsis of the Genera and Species of the Class Hypostoma (Asterias Linnaeus). Ann. Mag. Nat. Hist., vol. vi, London, 1841 (pp. 175-184, November, 18400; pp. 275-290, December, i84O&). . Ann. Mag. Nat. Hist., vol. xx, 18470. . Descriptions of Some New Genera and Species of Asteriadae, Proc. Zool. Soc. London for 1847, pp. 72-83, i847&. . Description of Platasterias, a New Genus of Astropectinidee, from Mexico. Proc. Zool. Soc. London, February, 1871, p. 136, pi. ix, 1871. GREGORY, J. W. The Stelleroidea. A treatise on Zoology, edited by E. Ray Lankester, part 3, pp. 237-259, 1900. GRIEG, JAMES A. Undersogelser over dyrelivet i de vestlandske Fjorde; II, Echinodermer, Annelider, etc., fra Moster. Bergens Museums Aars- beretning, 1888, n pp. i pi., Bergen, 1889. . Om Echinodermfaunaen i de vestlandske Fjorde. Bergens Museums Aarbog, 1894-95, no. 12, 13 pp., Bergen, 1896. . Skrabninger i Vaagsfjorden og Ulvesund ytre Nordfjord. Bergens Museums Aarbog, 1897, no. 16, 27 pis., Bergen, 1898. . Oversigt over det nordl. Norges Echinodermer. In Bergens Mus. Aarbog, for 1902, pp. 1-38, pi. i, 1902. Echinodermen von dem norwegischen Fishereidampfer Michael Sars in den Jahren 1900-1903 gesammelt; III, Asteroidea. Bergens Museums Aarbog, 1905, no. 13, 87 pp., 2 pis., 1906. . Echinodermata. Report of the Second Norwegian Arctic Expedition in the Fram, 1808-1902, no. 13, 26 pp., 3 pis., 1907. GRUBE, AD. EDOUARD. Beschreibungen neuer oder weniger bekannter Seesterne und Seeigel. Nova Acta Ac. Caes. Leop. Carol. Nat. Cur., vol. xxvn, 50 pp., 3 pis., 1857. i I 378 VERRILL . Diagnoseu einiger neuen Echinodermen. Archiv f. Naturgesch., pp. 340-344, i857&. . Ueber einige Seesterne des Breslauer Museums. 42. Jahresber. d. schlesischen Ges. f. vaterlandische Cultur, p. 52, 1865. . Einige neue Seesterne des hiesigen Zoologischen Museums. 43. Jahresber. d. schlesischen Ges. f. vaterlandische Cultur, p. 59, 1866. HAMANN, OTTO. Echinodermen. Bronn's Klassen und Ordnungen des Tier- reichs, Heft 2, Abth. 3, pp. 657-719, 1899. HARTOG, MARCUS M. The True Nature of the " Madreporic System " of Echinodermata, with Remarks on Nephridia. Ann. and Mag. Nat. Hist., ser. 5, vol. xx, pp. 321-326, November, 1887. HERAPATH, W. B. On the Pedicellarise of the Echinodermata. Quarterly Journ. Micros. Sci., vol. v, p. 175, 1866. HEUGLIN, M. TH. VON. Reisen nach dem Nordpolarmeer in den Jaren 1870 und 1871, 3. Teil, 8°, Braunschweig, 1874. HOFFMANN, C. K. Die Echinodermen, gesammelt wahrend der Fahrten des Willem Barents in den Jahren 1878 und 1879. Niederland. Archiv f. Zool., Suppl., Bd. i, 20 pp., I pi., 1882. IVES, J. E. On Two New Species of Starfishes. Proc. Acad. Nat. Sci. Philadelphia, p. 421, 1888. The species are Pteraster tesselatus and Coronaster bispinosus. . Variations in Ophiura panamensis and Ophiura teres. Op. cit, p. 76, 18890. O. panamensis, dark variety, from San Diego. . On a New Genus and Two New Species of Ophiurans; op. cit, p. 143, 18896. Ophioncus granulosus and Ophioglypha lockingtoni; localities uncertain. . Catalogue of the Asteroidea and Ophiuroidea in the Collection of the Academy of Natural Sciences of Philadelphia; Proc. Acad. Nat. Sci. Philadelphia, for 1889, p. 169 (distribution), 18890. . Echinoderms and Arthropods from Japan. Proc. Acad. Nat. Sci. Philadelphia, for 1891, pp. 210-223, pis. vn-xii, 18910. . Echinoderms and Crustaceans Collected by the West Greenland Expedition of 1891. Proc. Acad. Nat. Sci. Philadelphia, for 1891, pp. 479- 481, 18916. . List of the Echinoderms and Crustacea in the Cabinet of Frederick Stearns, Detroit, Mich. Private publication, not dated; 1891 (?). On p. ii is a List of West Coast Asterioidea; on p. iii, a List of Echinoidea; on p. v, a List of Ophiuroidea. JARZYNSKY, TH. See Wagner, Nicolas. JENNINGS, H. S. Formation of Habits in the Starfish. Johns Hopkins Univ. Circulars. Notes from Biol. Laboratory, 1907, no. 3, p. 16 [188], 19070. . Behavior of the Starfish Asterias forreri de Loriol. Univ. Calif. Pub., Zoology, vol. iv, no. 2, pp. 53-185, 19 text figs., 19076. . Features in the Behavior of the Starfish Illustrating the Grounds for the Attribution, by older Authors, of Intelligence to Lower Animals. Proc. Seventh Internat. Zool. Cong., Boston, August, 1907, pp. 145-146, Cambridge, 1912. Relates to experiments on Asterias forreri (= Orthasterias forreri). SHALLOW-WATER STARFISHES 379 JULLIEN, JULES. Description d'un Nouveau Genre des Stellerides, de la Familie des Asteriadees ; in Bull. Soc. Zool. de France, p. 141 (Marth- asterias}, 1878. KALISCHEWSKIJ, M. Zur Kenntnis der Echinodermenfauna des Sibirischen Eismeeres. Resultats Sci. Exped. polaire Russe en 1900-03, sous la Direction du Baron E. Toll, vol. i, sect. E, Zool. i, pt. 4, 3 pis., Bull. 1907 ; Mem. de 1' Academic Imperiale des Sciences de St. Petersbourg, ser. 8, Gasse Physico-Mathem., vol. xvm, no. 2, 1907. KNIPOWITSCH, N. Eine zooligische Exkursion im norwestlichen Theile des Weisen Meeres im Sommer 1896. Annuaire du Musee Zoologique de 1'Acad. Imp. des Sciences de St. Petersbourg, pp. 278-326, 1806. KGEHLER, RENE. Note preliminaire sur les Echinodermes recueillis par 1'Ex- pedition Antarctique Franchise du Dr. Charcot. Bull. Mus. d'Hist. Naturelle, for 1905, no. 6, pp. 464-470, 1905. . Echinoderms. Exped. Antarct. Frangaise (1903-05), commandee par le Dr. Jean Charcot, pp. 1-27, pis. i-rv, December, 1906. . Note preliminaire sur quelques Asteries et Ophiures Provenant des Campagnes de la " Princesse Alice." Bull. 1'Institut Oceanographique, no. 99, April, pp. 1-40, 19070. — . Asteries, Ophiures et Echinides recueillis dans les Mers Australes par la " Scotia " (1902-04). Zoologischer Anzeiger, vol. xxxn, no. 6, Sep- tember 17, pp. 143-145, IQ07&- — . Asteries, Ophiures et Echinides de 1'Expedition Antarctique National Ecossaise. Trans. Roy. Soc. Edinburgh, vol. XLVI, part 3, no. 22, De- cember 6, pp. 529-581, 628-632, pis. i-vin, xn, 1908. An Account of the Deep-sea Asteroidea Collected by the Royal Indian Marine Survey Ship Investigator, 1909. Echinoderma of the Indian Museum, Part V, Asteroidea, i, 143 pp., 13 pis., Calcutta, 19090. — ' . Echinodermes provenant des Campagnes du Yacht Princesse-Alice (Asteries, Ophiures, Echinides, et Crinoides). Result. Campag. Sci. du Albert I, Prince de Monaco, fasc. xxxiv, April 30, pp. 1-136, pis. i-xxin, I9O9&. KCEHLER, R., and VANEY, C. Mission des Pecheries Cote Occid. d'Afrique, II, Echinodermes. Act. Soc. Linn., Bordeaux, ix, 9 pp., pis. rv-vi, 1905. — . An account of the Shallow-water Asteroidea. Echinoderma of the Indian Museum, Part VI, Asteroidea, 11, June, 191 pp., 20 pis., Cal- cutta, 1910. KOREN, J., and DANIELSSEN, D. C. Fauna littoralis Norvegise, 3. Heft, folio, 163 pp., 16 pis.. Bergen, 1877. LAMARCK, J. B. P. A. DE. Histoire Naturelle des Animaux sans Vertebras. First ed., vol. n, Paris, 1816. LEIPOLDT, F. Asteroidea der " Vettor-Pisani "-Expedition (1882-85). Zeitschr. f. wiss. Zool., vol. LIX, pp. 545-654, pis. xxxi, xxxu, 1895. — . Ueber die Geschlechtsorgane eines brutpflegenden Seesternes, der Asterias rugispina Stimpson. Sitzungsberichte der Niederrheinischen Gesellschaft fur Natur- und Heilkunde zu Bonn, 1896, pp. 100-104. LEVINSEN, G. W. R. Kara-Havets Echinodermata. In Liitken, Dijmphna- Togtets zoologisk-botaniske Udbytte, 38 pp., pis. xxxiv, xxxv, Kjoben- havn, 1886. LJNCK, JOH. HENR. De Stellis marinis, folio, 107 pp., 42 pis., Lipsiae, 1733. 380 VERRILL LORIOL, P. DE. Notes pour Servir a 1'fitude des Echinodermes, no. 2. In Recueil Zoologique Suisse, vol. iv, pp. 363-407, pi. xviii, Geneve, 1888. Includes descriptions of Asterias forreri, p. 401, pi. xvm, fig. i., and Asterias exquiseta, p. 403, pi. xvm, fig. 2, both from California. . Notes pour Servir a 1'fitude des Echinodermes. No. 3. Mem. Soc. Phys. et Hist. Nat. Geneve, vol. centenaire, suppl., 1890, p. 22, pi. in, 1891. The same, no. 5. In Mem. Soc. Phys. et Hist. Nat. Geneve, vol. xxxn, part 2, no. 9, pp. 1-26, pis. xvi, xvn, [i-iii], 1897. Includes descriptions and figures of several starfishes from Vancouver I. ; viz., Asterias kcehleri, A. saiinichensis, Crossaster vancouverensis, Astropecten rubidus (Mexico), etc. . Notes pour Servir a 1'Etude des Echinodermes, no. 7. Mem. Soc. Phys. et Hist. Nat. Geneve, vol. xxxin, part 2, no. I, pp. 12-29, pis. n, in, 1899. . The same. Seconde Serie, fasc. n, pp. 1-68; pis. i-iv, 1904. Includes various starfishes, some new, from Gulf San Mathias, East Patagonia, with good figures. LUDWIG, HUBERT. Echinodermen des Beringsmeeres. Zoolog. Jahrb., Bd. I, for 1886, pp. 275-296, pi. vi, 1886. . Die Seesterne des Mittelmeeres, 4°, 491 pp., 12 pis., Berlin, 1897. . Arktische Seesterne. In Fauna Arctica (Romer and Schaudinn), vol. i, article xiv, pp. 445-502, Jena, 1900. . Seesterne. Resultats du Voyage du S. Y. Belgica. Zoologie, pp. 1-72, 7 pis., 1903. Brutpflege bei Echinodermen. In Zoolog. Jahrb. for 1904, pp. 683- 699, 1904. . Asteroidea. In Reports on an Exploration off the West Coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer Albatross, during 1891, etc. Mem. Mus. Comp. Zool., vol. xxxn, July 17, pp. i-xii, 1-292, pis. i-xxxvi, 19050. . Asterien und Ophiuren der schwedischen Expedition nach den Magalhaenslandern, 1895-1897. Zeitschr. f. wiss. Zool., vol. LXXXII, pp. 39-79, pis. v, vi, IQ05&. . Notomyota, eine neue Ordung der Seesterne. Sitzungsber. k. preuss. Akad. Wiss., vol. xxm, April, pp. 435-466, 1910. . Ueber die J. E. Grayschen Gattungen Pentagonaster und Tosia. Abdr. Zoolog. Jahrb., Suppl., xv, i Band, 44 pp., 1912. LUTKEN, CHR. FR. De ved Danmarks kyster levende Pighude; Videns- kabelige Meddelelser fra den naturhist. Forening, Kjobenhavn, for 1856, pp. 88-110, 18570. . Oversigt over Gronlands Echinodermata, 8°, 109 pp., i pi., Kjoben- havn, 1857^. . Bidrag til Kundskab om de ved Kysterne af Mellem og Syd-Amerika levende Arter af Sostjerner. Vidensk. Meddel. naturhist. Foren., Kjoben- havn, 1859. . Kritiske Bemserkninger om forskjellige Sostjerner (Asterider), med Beskrivelse af nogle nye Arter. Vidensk. Meddel. naturhist. Foren., Kjobenhavn, for 1864, pp. 123-169, 1865. SHALLOW- WATER STARFISHES 381 — . Fortsatte kritiske og beskrivende Bidrag til kundskab om Sostjer- nerne (Asteriderne), Vidensk. Meddel. naturhist. Foren., Kjobenhavn, for 1871, pp. 227-304, pis. iv, v, 1871. Nogle temmelig uventede Forogelser af den norske Havfauna. Vidensk. Meddel. naturhist. Foren., Kjobenhavn, 1889. MCANDREW, R., and BARRETT, L. List of the Echinodermata dredged between Drontheim and the North Cape. Ann. Mag. Nat. Hist. (2), vol. xx, pp. 43-46, London, 1857. MARENZELLER, E. v. Die Colenteraten, Echinodermen und Wiirmer der k. k. osterreichisch-ungarischen Nordpol-Expedition. Denkschriften d. math.- naturw. Klasse d. Kais. Akad. d. Wiss., Bd. xxxv, 4 pis., Wien, 1878. MASTERMAN, A. T. The Early Development of Cribrella oculata, with Re- marks on Echinoderm Development. Trans. Roy. Soc. Edinburgh, vol. XL, pp. 373-418, pi. i-v, 1902. Abstracts in Nature, LXV, p. 551, April, 1902; and American Naturalist, vol. xxxvii, p. 121, February, 1903. MEISSNER, MAXIMILIAN. Die von Herrn Dr. L. 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Acanthasteridae 204. acanthostoma, Evasterias, new species ii, 64, 76, 161, 165, 341, 348; pis. 20, 24. acervata, Asterias 13, 36, 59, 107, 108, 338; pis. 27, 106. acervata borealis, Asterias 370. acutispina, Coscinasterias 38, 46, 373. Adelasterias, new genus 352, 360, 371, 374- Adelasterias papillosa 360, 372. Adetopneusia 24. aqualis, Asterias 128. aequalis, Leptasterias 36, 128; pis. 25, 56. aequalis, Mediaster, 291, 296, 343, 344, 346, 348 ; pis. 2, 3, 5. sequalis, var. compacta, Leptasterias, new variety 61, 130, 341, 344; pi. 16. aequalis, var. concinna, Leptasterias, new variety 61, 132, 344. aequalis, var. nana, Leptasterias, new variety 61, 132, 341, 344, 345. afhnis, Asterias 259, 261. affinis, Crossaster 261. affinis, Pteraster 278. alaskensis, Leptasterias epichlora, new subspecies 13, 39, 61, in, 136, 338, 341 J pis. 16, 28, 84. alba, Cosmasterias 358. albertensis, Parasterias, new species 53, 55, 187, 342, 348; pis. 57, 70. alboverrucosa, Asterias 259, 261, 262. albula, Asierias 41. albula, Stephanasterias 39, 40, 41, 55, 67, 147, 338. albulus, Asteracanthion 147. albulus, Nanaster 41. albulus, Stichaster 41, 147. aleutica, Henricia 213. Alexandraster 205. Alexandraster inflatus 206. alexandri, Coscinasterias 350, 373. alexandri, Stolasterias 372. Allasterias 38, 53, 55, 66, 188, 352. Allasterias amurensis 373. Allasterias anomala 67, 193, 338, 347 ; pis. 59, 60, 69, 78. Allasterias forficulosa, new species 67, 194, 373 ; pis. 77, 83, 84. Allasterias migrata 373. Allasterias rathbuni 53, 55, 66, 67, 189, 337, 338, 347- Allasterias rathbuni nortonensis 67, 191, 338, 347; pl. 78. Allasterias versicolor 196, 373. Allopatiria ocellifera 263. Allostichaster, new genus 362, 363, 373, 374- Allostichaster polyplax 373. alveolata, Evasterias troschelii, new variety 63, 157, 162, 341 ; pl. 62. Amphiaster 294. Amphiaster insignis 294, 373 ; pl. 98. amurensis, Allasterias 373. amurensis, Asterias 194. Anasterias 8, 352, 353, 354. Anasterias belgicae 51, 355, 371. Anasterias chirophora 51, 355. Anasterias lactea 355. Anasterias lysasteria, new name 354, 355, 37L Anasterias minuta 354, 355, 371. Anasterias perrieri 51, 354, 355. Anasterias studeri 355, 371. Anasterias tenera 355, 371. Anasterias verrillii 354, 355, 371. (389) 390 INDEX angulosa, Asterias 370. annulata, Labidastrella 373. annulata, Labidiaster 352, 373, 374. anomala, Allasterias 67, 193, 338, 347 ; pis. 59, 60, 69, 78. anomalus, Glyphaster 316, 328, 340, 347; pis. i, 6. Antarctic species 2, 51, 350, 351. antarctica, Porania 368, 372. antarctica, Sporasterias 51, 53, 353, 354, 371. antarcticus, Leptychaster 369. antarcticus, Pedicellaster 367, 371. Anthenea 286, 287. Anthenea granulifera 288. Antheneidce, 282, 299. Antheneinae 283. antillarum, Henricia 210, 370. antonioensis, Echinaster 363, 372. aporus, Pteraster 277, 278. aporus, Pterasterides 278, 339, 347. Archaster arcticus 326. Ar chaster bairdii 298. Archaster dawsoni 312. Archasteridae 283. arctica, Asterias 120. arctica, Henricia, new species 239, 339, 347; PL 87. arctica. Leptasterias 120, 338, 347 ; pis. 56. 71-72, 83. arctica, Tosia 292. arcticus, Archaster 326. arcticus, Astropecten 326. arcticus, Ceramaster 292. arcticus, Leptoptychaster 326. arcticus, Leptychaster 326, 339. arcticus, Solaster dawsoni 347. arcticus, Tosiaster 292, 339; pis. 50, 99- arcuatus, Pteraster tesselatus 269. armatus, Astropecten 373. armatus, Astropecten 317, 319. aspera. Henricia 241, 339, 342, 346, 348. Asteracanthion 101, 114. Aster acanthion albulus 147. Asteracanthion clavatum 358. Asteracanthion fulvum 358. Asteracanthion katherina? 112. Asteracanthion margaritifer 69. Asteracanthion mite 358. Asteracanthion problema 147. Asteracanthion rubens 373. Asteracanthion spectabile 358. Asteracanthium camtschaticum i 14. Asterias 4, 11, 16, 17, 18, 33, 34, 43. 44, 55, 58, 68, 101, 352. Asterias acervata 13, 36, 59, 107, 108, 338; pis. 27, 106. Asterias acervata borealis, new name 107, in, 370. Asterias aqualis 128. Asterias affinis 259, 261. Asterias alboverrucosa 259, 261, 262. Asterias albula 41. Asterias amurensis 194. Asterias angulosa 370. Asterias arctica 120. Asterias aurantiaca, 362, 372. Asterias borealis 107, 113. Asterias brachiata 152, 156. Asterias brandti 371. Asterias brevispina 77. Asterias camtschatica no. 114. if 6, 127, 132, 135. Asterias capitata 81. Asterias conferta 73, 77. Asterias (Cosmasterias} tomidata 48. Asterias cribraria 148. Asterias crisp atus 330. Asterias cunninghami 353. Asterias decemradiatus 254. Asterias (Diplasterias) epichlora 152. Asterias douglasi 107, 113, 114. Asterias dubia 97. Asterias echinophora 207. Asterias endeca 244. Asterias epichlora 132, 158. Asterias exquiseta 83. Asterias nssispina 76. Asterias forbesi 3, 5, 10, 18, 38. 188, 370. Asterias forreri 47. 179. Asterias gelatinosa 357. Asterias gigantea 89, 97. Asterias granularis 290. Asterias helianthoides 198. Asterias he.ractis 126. Asterias hyadesi 354. Asterias janthina 69, 73. INDEX 391 Asterias katherinae 13, 36, 59, 112, 340, 344, 348; pis. 51, 83. Asterias lurida 337, 346. Asterias liitkenii, 44, 82, 83. Asterias mcridionalis 361, 372. Asterias militaris 272. Asterias miniata 264. Asterias minuta 107. Asterias multiclava 59, no, 114, 338, 347 ; pis. 58, 59, 69, 84. Asterias nanimensis, new species 59, 105,340; pi. 61. Asterias neglecta 358. Asterias obtusispinosa 358. Asterias ochracea 69, 77, 82. Asterias oculata 226. Asterias papillosa 372. Asterias papposa 259. Asterias paucispina 98. Asterias perrieri 354, 356. Asterias pertusa 226. Asterias planci 364. Asterias polaris 107, 108, 113, 370. Asterias polythela 59, 104, 338, 347; pis. 55, 70, 72, 79, 84. Asterias rosea 40, 362. Asterias rubens 44, 55, 68, 188, 337, 346. Asterias rubens 107, 189. Asterias rugispina 51, 53. 353, 355- Asterias rupicola 356. Asterias rustica 357. Asterias saanichensis 132, 135. Asterias sanguinolenta 226. Asterias sertulifera TOO. Asterias spirabilis 51, 354. Asterias spitsbergensis 148, 150. Asterias steineni 361. Asterias stellionura 51. Asterias (Stichaster) polyplax 125. Asterias studeri 51. Asterias sulcifera 48, 358, 361. Asterias troschelii 114, 151, 156. Asterias turqueti 372. Asterias (Urasterias) forcipulata 180. Asterias vancouveri 13, 125. Asterias varia 51. Asterias victoriana 59, 102, 340, 348; Pis. 53- 54, 69, 82. Asterias violacea 107. Asterias vulgaris 38, 370. Asteriidae 27, 30, 39, 42, 43, 56. Asteriidte, Sladen's 43. Asteriinae 26, 39, 42, 43, 49, 5«. Asterina 262. asterina, Cycethra 366. Asterina gibbosa 262, 263, 371. Asterina miniata 264. Asterina minuta 262. Asterina perrieri 365, 372. Asterina pygmcea 371. Asterinidae 204, 262. Asterinides folium 263, 371. Asterinides modesta 373. Asterinides perrieri 365, 372. asterinoides, Calvasterias 357, 371. Asterinopsis penicillaris 263, 373. Asterioidea, classification 24. Asterioidea, derivation of name 20. Asteriscus 263. asterisctts, Cycethra 366. Asteriscus miniatus 264. Asterodon belli 367. Asterodon granulosus 367. Asterodon singularis 367, 372. Asteropidas 282, 284, 304. Asteropinae, new subfamily 305. Asteropsis 305. Asteropsis imbricata 306. asthenactis, Henricia 213. asthenosoma, Luidia 336, 345, 346. Astrogonium 287, 289, 300. Astrogonium granulare 290. Astrogonium granularis 287. Astrogonium patagonicus 368. Astropecten 281, 316, 317, 348. Astropecten arcticus 326. Astropecten armatus 373. Astropecten armatus 317, 319. Astropecten calif ornicus 319. 320, 345, 346; pis. 50, 100, 101, 102. Astropecten ctenophorus 321. Astropecten erinaceus 347. Astropecten ierstedii 318, 319, 346, 373, Astropecten ornatissimus 320, 346, 349- Astropecten siderealis, new species 317. 345. 346, 349; pl- 50- Astropectinidae 283, 285, 314. 392 INDEX Astropectinides, new genus 316, 317, 321, 372, 374. Astropectinides mesacutus 316, 321, 372. atlantica, Tonia 40, 362. attenuata, Henricia leviuscula 217, 218, 342, 344. aurantiaca, Asterias 362, 372. aurantiacus, Stichaster 40, 362. australis, Coelasterias 373. australis, Ctenodiscus 333, 369. australis, Pisaster lutkeni, new variety 85, 88, 345. australis, Solaster 366, 368, 372. austro-granularis, Ceramaster 369. austro-granularis, Pentagonaster 369. Avelata, suborder 204. bairdii, Ar chaster 298. bairdii, Isaster 298. bairdii, Mediaster 298, 371 ; pis. 2, 3. belgicae, Anasterias 51, 355, 371. belli, Asterodon 367. bellonse, Luidia 367. bellula, Callopatiria 263. Benthopectinidse 283, 310. Benthopectinidce 311. Benthopectininae 283, 310, 311. Beringian fauna 338, 347. Bibliography 374-388. bifascialis, P hat aria (Linckia) uni- fascialis var. 309. biordinata, Orthasterias, new species 48,64, 172,341; pis. 63, 82. biserialis, Peribolaster 366. biserialis, Phataria 346. biseriatus, Granaster 360, 362, 372. biseriatus, Hemiasterias, new genus 362, 372. bispinosa, Patiriella 364, 372. Blakiaster 322, 324. Blakiaster conicus 322, 371. borealis, Asterias 107, 113. borealis, Asterias acervata 107, in. 370. borealis, Henricia 235, 236, 239. borealis, Henricia tumida, new sub- species 236, 339, 342, 347; pis. 12, 86, 88. brachiata, Asterias 152, 156. Brachiolaria, larval form 24. bradleyi, Mithrodia 373 ; pi. 107. brandti, Asterias 371. brandti, Cosmasterias 360, 372. brasiliensis, Echinaster 363. brasiliensis, Enoplopatiria 263. brevispina, Asterias 77. brevispina, Luidia 347. brevispina, Pisaster 77. brevispinus, Pisaster 57, 77, 93, 34°, 344, 345; pis. 41, 44, 45, 69, 76. briareus, Coronaster 49, 370. briareus, Thyone 10. Brisingidse 15, 24, 26, 374. Brisinginas 26. Brooding of the young 39. Bunodaster 316, 321. Bunodaster ritteri 316, 322, 345, 348; pis. 86, 104, 105. calcarata, Patiriella 364, 365, 372. Californian fauna 337, 343. californica, Orthasterias, new species 48, 64, 174, 185, 344, 350; pis. 68, 70, 80, 81. californicus, Astropecten 319* 320, 345, 346; pis. 50, ioo, 101, 102. californicus, Rathbunaster 197. Callopatiria bellula 263. Callopatiria obtusa 373. calva, Cycethra 366. Calvasterias 42, 353, 357. Calvasterias asterinoides 357, 371. Calvasterias stolidota 42, 354, 35 7» 371- camtschatica, Asterias no, 114, 116, 127, 132 135. camtschaticum, Asteracanthium 114. candicans, Coscinasterias 367. candicans, Stolasterias 367. Cannibalism among starfishes 4. capensis, Pteraster 278. capitata, Asterias 81. capitatus, Pisaster 19, 57, 81, 344, 345; pis. 36, 56. carinella, Leptasterias epichlora, new variety 61, I37» 34*- carribaeus, Pteraster 278. Ceramaster 289. INDEX 393 Ceramaster arcticus 292. Ceramaster austro-granularis 369. Ceramaster granularis 290, 293, 339, 343 ; pis. 4, 50. Ceramaster japonicus 292. Ceramaster leptoceramus 292. Ceramaster patagonicus 291, 339, 343, 347, 349, 368. Chaetasteridae 283. Cheiraster gerlachei 369. Cheirasteridce 311. chelifera, Distolasterias, new species 66, 185, 342; pis. 81, no. chilensis, Patiria 348, 364, 372. chirophora, Anasterias 51, 355. chirophora, Pasdasterias, new genus 355, 371- Chitonasterinae 283. Cladaster 300. clarki, Henricia 213. clathrata, Luidia 7, 371 ; pi. 103. clavatum, Asteracanthion 358. coccinea, Patiria 263, 373. cocosana, Sporasterias 350. coei, Leptasterias, new species 36, 61, 123, 178, 340, 348 ; pis. 9, 17. Coelasterias 40. Coelasterias australis 373. cognatus, Mimaster 368. columbise, Linckia 309, 337, 349. columbiana, Orthasterias, new spe- cies n, 48, 50, 64, 168, 178, 183, 341, 344, 346, 348; pis. 24, 35, 65, 78, 79, 109. compacta, Leptasterias aequalis, var. 61, 130, 341, 344. compta, Leptasterias 8, 121, 370. concinna, Leptasterias aequalis, new variety 61, 132, 344. conferta, Asterias 73. confertus, Pisaster 57, 73, 167, 340; pl. 38. conicus, Blakiaster 322, 371. constellatus, Solaster 257, 343, 348; pis. 46, 90, 91, 93, 94. corniculatus, Ctenodiscus 331. Coronaster 30. Coronaster briareus 49, 370. Coscinasterias 33, 34, 36, 38, 39, 43, 44, 45, 46. Coscinasterias acutispina 38, 46, 373. Coscinasterias alexandri 350, 373. Coscinasterias candicans 367. Coscinasterias echinata 46. Coscinasterias gemmifera 46. Coscinasterias jehennesii 46. Coscinasterias muricata 46, 49, 373. Coscinasterias (Stolasterias) ten- uispina 46, 49. Coscinasterias tenuispina 38. 43, 45, 370. Cosmasterias 33, 44, 48, 353, 358. Cosmasterias 67. Cosmasterias alba 358. Cosmasterias brandti 360 372. Cosmasterias fernandensis 360, 372. Cosmasterias lurida 36, 48, 50, 358, 3/1. Cosmasterias polygrammus 360, 371. Cosmasterias sulcifera 48, 358, 360, 361, 371. Cosmasterias tomidata 48, 50, 359, 371- crassa, Henricia leviuscula 217. crassa, Parapatiria 263. crassus, Odontaster 304, 345, 346, 349- cremens, Odontaster 368. Cribella 209. Cribella hyadesi 363. Cribella oculata 226. Cribella rosea 226. cribraria, Asterias 148. cribraria, Ctenasterias 53, 60, 148, 338; pl. 25. cribraria, Leptasterias 8. Cribraster sladeni 363, 372. Cribrella 211. Cribrella leviuscula 215. Cribrella cculata 226. Cribrella pectinata 230. Cribrella prcestans 368. Cribrella sanguinolenta 223, 226. Cribrella simplex 368. Cribrella spiculifera 220. Cribrellse 129. cribrosus, Pteraster 278. crispatus, Asterias 330. crispatus, Ctenodiscus 330, 340, 343, 345, 347, 349 ; pis. 5, 49- 394 INDEX Crossaster 13, 15, 258. Crossaster 243. Crossaster affinis 261. Crossaster korenii 261, 371. Crossaster papposus 249, 259, 339, 343 ; Pis. 5, 8, 9, 49. Cryptasterias, new genus 362, 372, 374- Cryptasterias turqueti 362, 372. Cryptozonia 24. Ctenaster 263. Ctenasterias, new genus 53, 55, 148, 374- Ctenasterias cribraria 53, 60, 148, 338 ; pi. 25. Ctenasterias spitzbergensis, 53, 55, 148, 270. Ctenodiscime 283, 329. Ctenodiscus, 315, 329, 330. Ctenodiscus australis 333, 369. Ctenodiscus corniculatus 331. Ctenodiscus crispatus 330, 340, 343, 345, 347, 349 ; pis. 5- 49- Ctenodiscus krausei 331. Ctenodiscus polaris 330. Ctenodiscus procurator 333. Ctenodiscus pygmcsus 330. ctenophorus, Astropecten 321. Culcita 288. cunninghami, Asterias 353. cunninghami, Sporasterias 353. Cycethra asterina 366. Cycethra asteriscus 366. Cycethra calva 366. Cycethra electilis 366. Cycethra elongata 366. Cycethra ganeriodes 366. Cycethra lahillei 366, 372. Cycethra media 366. Cycethra nitida 366. Cycethra pinguis 366. Cycethra regularis 366. Cycethra simplex 366, 372. Cycethra subelectilis 366. Cycethra verrucosa 366, 372. Cystidea, number of rays 15. danse, Pteraster 278. dawsoni, Acantharchaster 312. dawsoni, Archaster 312. dawsoni arctica, Solaster, new sub- species 252, 253, 347; pi. 87. dawsoni, Luidiaster 312, 339, 343, 349 ; pis. 33, 34, 35- dawsoni, Orthasterias, new species 48, 65, 172, 175, 342; pis. 23, 75, 80, 8x. dawsoni, Solaster 249, 339, 342, 348; pis. 46, 90, 91, 92. decemradiatus, Asterias 254. decemradiatus, Solaster 254. densa, Evasterias troschelii, new variety 63, 161, 341. densispina, Henricia 223, 373. densus, Perknaster 364. Dermasterias 68. Dermasterias imbricata 14, 306, 343, 345, 348; pis. 50,86,97. Dermasterias imbricata, var. valvuli f era 305. 3o8, 343 ; pi. 29. Dermasterias inermis 306. Desmopatiria 263. Desmopatiria flexilis 263, 364, 372. Development, 39. diomedeae, Hydrasterias 350. Diplasterias 43, 44, 101, 358. Diplasterias loveni 361. Diplasterias liitkeni 361. Diplasterias papillosa 360. Diplasterias spinosa 361, 372. Diplasterias steineni 361. Diplasterias turqueti 362. diplax, Linckia 309. Diplopteraster 203, 267, 273, 279. Diplopteraster multipes 279, 339. Diplopteraster peregrinator 368. Diplopteraster verrucosus 368. dispar, Leptasterias, new species 63, 115, 142, 341; pi. 16. Distolasterias 33, 47, 66, 185, 350. Distolasterias chelifera, new species, 66, 185, 342; pis. 81, no. Distolasterias stichantha 44, 49, 185, 373- Distribution, geographical 18, 337. Dorsal ossicles, form 34. Dorsal spines, form 34. douglasi, Asterias 107, 113 114. dubia, Asterias 97. INDEX 395 dyscrita, Henricia leviuscula 323, 342, 344- Dytaster 285. Echinaster 101, 206. Echinaster 209. Echinaster antonioensis 363, 372. Echinaster brasiliensis 363. Echinaster eschrichtii 226. Echinaster lepidus 363. Echinaster oculatus 226. Echinaster (Othilia) robustus, new species 207. Echinaster (Othilia) tenuispinus 346. echinaster, Poraniopsis 349, 363, 372. Echinaster (Rhopia) lepidus 372. Echinaster robustus 342. Echinaster sanguinolenta 226. Echinaster sarsii 226. Echinaster scrobiculata 227. Echinaster sentus 207. Echinaster smithii 368. Echinaster Solaris 364. Echinaster spinosus 207. Echinaster tenuispinus 208, 346, 349, 373 ; pi. 107. Echinasteridae 204, 205. Echinasterina 205, 242. echinata, Coscinasterias 46. e rhinites, Acanthaster 364, 373. echinophora, Asterias 207. echinus, Acanthaster 364. Eggs and young 7, 9, n. electilis, Cycethra 366. ellisii, Acanthaster 364, 373. elongata, Cycethra 366. endeca, Asterias 244. endeca, Solaster n, 243, 244, 248, 249, 339; pis. 9, 87, 89. Enoplopatiria brasiliensis 263. Enoplopatiria marginata 263, 365, 371. Enoplopatiria siderea 365, 373; pi. 109. epichlora, Asterias 132, 156. epichlora, Asterias (Diplasterias) 152. epichlora alaskensis, Leptasterias, new subspecies 13, 39, 61, in, 136, 338, 341 ; pis. 16, 28, 84. epichlora alaskensis, var. carinella, Leptasterias, new variety 61, 137, 341 ; pi. 16. epichlora alaskensis, var. siderea, Leptasterias, new variety 61, 137, 341 ; pi. 16. epichlora, Leptasterias 8, 17, 36, 60, 62, no, 132, 34i. 348. epichlora miliaris, Leptasterias, new subspecies 60, 61, 138, 341. epichlora plena, Leptasterias, new subspecies 61, 140, 341. epichlora pugetana, Leptasterias, new subspecies 61, 142, 341. epichlora, var. regularis, Leptasterias, new variety 63, 139, 341. epichlora, var. subnodulosa, Leptas- terias, new variety 62, 73, 139, 341- epichlora, var. subregularis, Leptas- terias, new variety 63, 139, 341. equestris, Goniaster 286. equestris, Hippasteria 286. erinaceus, Astropecten 347. eschrichtii, Echinaster 226. eschrichtii, Henricia 212, 236, 329. eustyla, Orthasterias 168. Evasterias, new genus 28, 33, 51, 54, 63, 151, 352, 374- Evasterias acanthostoma, new species ii, 64, 76, 161, 165, 341, 348; pis. 20, 24. Evasterias troschelii n, 36, 51, 54, 63, 151, 341, 344, 348; pis. 22, 25, 26, 106. Evasterias troschelii, var. alveolata, new variety 63, 157, 162, 341 ; pi. 62. Evasterias troschelii, var. densa, new variety 63, 161, 341. Evasterias troschelii, var. parvispina, new variety 63, 163, 341. Evasterias troschelii, var. rudis, new variety 63, 158, 341. Evasterias troschelii, var. subnodosa, new variety 63, 163, 341. exquiseta, Pisaster liitkenii 345. exquiseta, Asterias 83. Eye or eye-spot of starfishes 9. 396 INDEX falklandica, Ganeria 365, 372. Faunal relations and distribution 18. felipes, Stichaster 49. fernandensis, Cosmasterias 360, 372. fernandensis, Polyasterias 360, 371. fimbriata, Patiriella 364, 365, 372. nssispina, Asterias 76. iissispina, Pisaster 76. fissispinus, Pisaster 34, 57, 76, 344 ; pi- 39- flavescens, Hosea 287. Aeuriaisi, Goniopecten 369. fleuriaisi, Psilaster 369. flexilis, Desmopatiria 263, 364, 372. foliata, Luidia n, 334, 343, 345, 346; pis. 100, 103, 105. foliculata, Peribolaster 366, 372. folium, Asterinides 263, 371. Food and feeding habits 3, 16. forbesi, Asterias 3, 5, i°. I7, 3$, 188, 370. Forcipulata 24, 41- forcipulata, Asterias (Urasterias) 180. forcipulata, Orthasterias n. forcipulata, Orthasterias forreri 65, 180, 342; pis. 62, 70,88. Forcipulosa, order 24, 26. forficulosa, Allasterias, new species 67, 194, 373; pis. 77, 83, 84. forreri, Asterias 47, 179- forreri forcipulata, Orthasterias 65, 1 80, 342; pi. 62, 70, 88. forreri, Orthasterias 19, 65, i?9» 344, 345 ; pis. 65, 66, 70, 77, 80. forreri, Orthasterias (Stylasterias) 342. forreri, Stylasterias 48, 50. Fossil starfishes 2, 12, 13, 14, 20, 197- franciscus, Linckia 211. fulvum, Asteracanthion 358. fuscus, Perknaster 364. galapagensis, Sporasterias 350. galaxides, Solaster 248, 342, 348; pis. 46, 87, 89. Ganeria falklandica 365, 372. Ganeria hahni 365. Ganeria papillosa 365, 372. Ganeria robusta 365. Ganeriidse 204. ganeriodes, Cycethra 366. Gastraster studeri 360, 372. gayi, Patiria 364, 372. gelatinosa, Asterias 357. gelatinosus, Meyenaster 54, 357, 371. gemmifera, Coscinasterias 46. Genital pores 39. Geographical distribution 18, 337-352. gerlachei, Cheiraster 360. gibber, Retaster 366, 372. gibbosa, Asterina 262, 263, 371. gigantea, Asterias 89, 97. giganteus, Pisaster n, 57, 84, 87, 89, 98, 344; pi. 37- glaber, Porania 368. glacialis, Marthasterias 46, 47, 49, 370. Glyphaster 316, 325, 327. Glyphaster anomalus 316, 328, 340, 347; pis. i, 6. Gnathaster 303. Gnathaster grayi 367. Gnathaster pilulatus 367. Gnathaster ince 283, 302. Goniaster 300. Goniaster equestris 286. Goniaster granularis 290. Goniaster nodosus 286. Goniaster obtusangulus 287. Goniaster reticulatus 286. Goniaster tesselatus 286. Goniasteridae 281, 282, 285, 286. Goniasterinae 283, 289. Goniodiscides seba 288. Goniodiscus 288. Goniodiscus pentagonulus 287. Goniopecten neuriaisi 369. Goniopectinidae 283. gonolena, Orthasterias, new species 65, 184, 344, 345, 3491 pis. 67, 68, 69, 82. gracilis, Pteraster 271, 343, 348. gracilis, Stephanasterias 146. Granaster 40, 353, 362. Granaster biseriatus 360, 362, 372. Granaster nutrix 372. grandis, Pisaster 13. INDEX 397 granulare, Astrogonium 290. granularis, Asterias 290. granularis, Astrogonium 287. granularis, Ceramaster 290, 293, 339, 343 ; pis. 4, 50. granularis, Goniaster 290. granularis, Pentagonaster 290. granularis, Tosia (Ceramaster) 290. granulifera, Anthenea 288. granulosus, Asterodon 367. granulosus, Odontaster 367, 372. grayi, Gnathaster 367. grayi, Odontaster 367, 372. grayi, Pisaster 58, 91, 97, 344. guernei, Sclerasterias 44, 50, 370. guildingii, Linckia 309, 310, 337, 346. gunneri, Urasterias 52. Gymnasteriidce 304. hahni, Ganeria 365. hebes, Pteraster 270, 343 ; pi. 96. helianthoides, Asterias 198. helianthoides, Pycnopodia II, 35, 198, 342, 344, 348; pis. 29, 30, 31, 73, 74,88. helianthus, Heliaster 372. Heliaster 12, 13, 15, 40. Heliaster helianthus 372. Heliasterinae 26, 39, 40. Hemiasterias, new genus 362, 374. Hemiasterias biseriatus 362, 372. Henricia 8, 17, 18, 144, 209. Henricia aleutica 213. Henricia antillarum 210, 370. Henricia arctica, new species 239, 339, 347; pi. 87. Henricia aspera 241, 339, 342, 346, 348. Henricia asthenactis 213. Henricia borealis 235, 236, 239. Henricia clarki 213. Henricia densispina 223, 373. Henricia eschrichtii 212, 236, 239. Henricia heteractis 210, 373. Henricia hyadesi 363, 368, 372. Henricia japonica 223, 373. Henricia leviuscula 14, 213, 215, 338, 342, 344, 346 ; pis. 12, 13. Henricia leviuscula, var. annectens 224, 344, 346. Henricia leviuscula, var. attenuata 217, 218, 342, 344. Henricia leviuscula, var. crassa 217. Henricia leviuscula, var. dyscrita 223, 342, 344. Henricia leviuscula, var. inequalis, new variety 219, 342; pi. 88. Henricia leviuscula, var. leviuscula 217, 342. Henricia leviuscula, var. lunula, new variety 213, 218, 342, 344; pi. 88. Henricia leviuscula, var. multispina 220, 222, 338, 342. Henricia leviuscula, var. pectinata 229, 230. Henricia leviuscula, var. spatulifera, 224, 342, 344, 346; pi. 5, 14. Henricia leviuscula spiculifera 220, 338, 342 ; pis. 87, 107. Henricia longispina 342, 348. Henricia obesa 363, 372. Henricia oculata 226. Henricia pagenstecheri 363, 372. Henricia pectinata 212. Henricia polyacantha 213. Henricia praestans 368. Henricia sanguinolenta n, 14, 212, 226, 338, 342 ; pis. 49, 88. Henricia sanguinolenta miliaris, new subspecies 234, 370; pi. 88. Henricia sanguinolenta, var. pectinata 229, 230, 338 ; pi. 49. Henricia sanguinolenta, var. rudis, new variety 233, 338, 347 ', pi- 86. Henricia sexradiata 371. Henricia simplex 368. Henricia spatulifera 224. Henricia spiculifera 220. Henricia studeri 363. Henricia tumida 214, 234, 239, 240, 338, 347 ; pis. 12, 87. Henricia tumida borealis, new sub- species 236, 339, 342, 347 ; pis. 12, 86,88. Henricides, new genus 210, 373, 374. Henricides heteractis 210, 373. heteractis, Henricia 210, 373. Heterasterias, new genus 46, 373, 374. Heterasterias volsellata 27, 47, 49, 373. INDEX hexactis, Asterias 126. hexactis, Leptasterias 13, 61, 125, 126, 340, 344, 348; pi. 25. hexactis, Pteraster 274. hexactis, Pteraster (Temnaster) 274. hexactis, Temnaster 267, 274. Hexaster 267. Hexaster obscurtts 274, 275. Hippasteria 300. Hippasteria equestris 286. Hippasteria hyadesi 368. Hippasteria phrygiana 286, 301, 371 ; pis. 47, 48, 49. Hippasteria spinosa 301, 339, 343, 345, 346, 348; pis. 50, 98. Hippasterias Bell 300. Hippasteriinas 283, 299. hispidus, Odontaster 349. Hosea 287. Hosea flavescens 287. hyadesi, Asterias 354. hyadesi, Cribella 363. hyadesi, Henricia 363, 368, 372. hyadesi, Hippasteria 368. hyadesi, Sporasterias 354. Hybrids or freaks 17, 18. Hydrasterias 43, 44, 350. Hydrasterias diomedeas 350. Hymenaster 267. Hymenaster obscurus 267. Hymenaster perspicuus 368. Hymenasterinse, new subfamily 266, 267. Ilyaster 315. imbricata, Aster op sis 306. imbricata, Dermasterias 14, 306, 343, 345, 348 ; pis. 29, 50, 86, 97. imbricata, var. valvulifera, Dermas- terias 305, 308, 343. Incubation 7. inequalis, Henricia leviuscula, new variety 219, 342; pi. 88. inequalis, Leptasterias, new species 6b, 117, 143, 340; pi. 73- inermis, Dermasterias 306. inermis, Parastropecten 325. inflata flexibilis, Poraniopsis 206. inflata, Poraniopsis 206, 344, 346, 349, 363. inflates, Alexandraster 206. ingouffi, Pteraster 368. insignis, Amphiaster 294, 373; pi. 98. Isaster 295. Isaster bairdii 298. janthina, Asterias 69, 73. Japanese current 2. japonica, Henricia 223, 373. japonicus, Ceramaster 292. Jaws and oral spines 36. jehennesii, Coscinasterias 46. katherina, Asteracanthion 112. katherinae, Asterias 13, 36, 59, 113, 340, 344, 348; pis. 51, 52, 83. katherince, Asterias Per. 58, 89, 91, 97. kerguelenensis, Leptychaster 316, 325, 3Si, 369, 372. koehleri, Orthasterias 64, 175, 183, 342 ; pi. 75. korenii, Crossaster 261, 371. Korethrasterinas 242. krausei, Ctenodiscus 331. Labidastrella, new genus 352, 373, 374. Labidastrella annulata 373. Labidiaster 12, 15, 26. Labidiaster annulata 352, 373, 374. Labidiaster radiosus 352, 371. Labidiasterinas 26. lactea, Anasterias 355. lahillei, Cycethra 366, 372. Lahillia 205. Lahillia mira 363. Larval stages 7. Lebrunaster paxillosus 365, 372. lebruni, Pteraster 368. Leiaster 308. lepidus, Echinaster 363. lepidus, Echinaster (Rhopia) 372. leptalea, Leptasterias, new species 60, 119, 340, 348. Leptasterias 8, 38, 41, 43, 50, 53, 55, 56, 60, 72, 116, 351. Leptasterias aaqualis 36, 128; pis. 25, 56. Leptasterias aequalis, var. compacta, new variety 61, 130, 341, 344; pi. 16. INDEX 399 Leptasterias aequalis, var. concinna, new variety 61, 132, 344. Leptasterias aequalis, var. nana, new variety 61, 132, 341, 344, 345- Leptasterias arctica 120, 338, 347 ; pis. 56, 71, 72, 83. Leptasterias coei, new species 36, 61, 123, 178, 340, 348; pis. 9, 17. Leptasterias compta 8, 121, 370. Leptasterias cribraria 8. Leptasterias dispar, new species 63, 115, 142,341; pl. 16. Leptasterias epichlora 8, 17, 36, 60, 62, no, 132, 341, 348. Leptasterias epichlora alaskensis, new subspecies 13, 39, 61, in, 136, 338, 341 ; pis. 28, 85. Leptasterias epichlora alaskensis, var. carinella, new variety 61, I37t 341 ; pl. 16. Leptasterias epichlora alaskensis, var. siderea, new variety 61, 137, 341 ; pl. 16. Leptasterias epichlora miliaris, new subspecies 60, 61, 138, 34*- Leptasterias epichlora plena, new vari- ety, 61, 140, 341 ; pl. 58. Leptasterias epichlora pugetana, new subspecies 61, 142, 341. Leptasterias epichlora, var. regularis, new variety 63, 139. 341- Leptasterias epichlora, var subnodu- losa, new variety 62, 73, 139, 341. Leptasterias epichlora, var. subregu- laris, new variety 63, 139, 341. Leptasterias hexactis 13, 61, 125, 126, 340, 344, 348; pl. 25. Leptasterias inequalis, new species, 36, 60, 117, 143, 340; pl. 73- Leptasterias leptalea, new species 60, 119, 340, 348; pl. 18. Leptasterias littoralis 8, 370. Leptasterias macouni, new species 13, 61, 124. Leptasterias macropora 145. Leptasterias mulleri 55, 124. Leptasterias obtecta, new species 60, 144. 338. Leptasterias tenera 8, 370. Leptasterias vancouveri 61, 125, 340. leptoceramus, Ceramaster 292. Leptogonasterinae 283. leptolena, Orthasterias, new species 66, 182, 342; pis. 64, 77. Leptoptychaster 325, 326. Leptoptychaster arcticus 326. Leptostroteria 24. Leptychaster 8, 316, 322, 325. Leptychaster antarcticus 369. Leptychaster arcticus 326, 339. Leptychaster kerguelenensis 316, 325, 351, 369, 372. Leptychaster millespina 326. Leptychaster pacificus 326, 340, 347 ; pl. 74- leviuscula, Cribrella 215. leviuscula, Henricia 14, 213, 215, 338, 342, 344, 346; pis. 12, 13. leviuscula, Linckia 215. leviuscula, var. annectens, Henricia 224, 344, 346. leviuscula, var. attenuata, Henricia 217, 218, 342, 344. leviuscula, var. crassa, Henricia 217. leviuscula, var. dyscrita, Henricia 223, 342, 344; pl. 12. leviuscula, var. inequalis, Henricia, new variety 219, 342; pl. 88. leviuscula, var. leviuscula, Henricia 217, 342. leviuscula, var. lunula, Henricia, new variety 213, 218, 342, 344; pl. 88. leviuscula, var. pectinata, Henricia 229, 230. leviuscula, var. spatulifera, Henricia 224, 342, 344, 346; pis. 5, 14. leviuscula spiculifera, Henricia 220, 338, 342 ; pis. 87, 107. Linckia 211, 309. Linckia 209. Linckia columbiae 309* 337, 349- Linckia dispar 309. Linckia franciscus 211. Linckia guildingii 309, 310, 337, 346 Linckia leviuscula 215. Linckia oculata 226. Linckia ornithopus 310. Linckia pertusa 226. Linckia typus 211. 400 INDEX (Linckia) unifascialis, var. bifascialis, Phataria 309. Linckia variolata 211. Linckiadce 282, 308. linckii, Urasterias 33, 51, 55, 66, 181, 187, 370 ; pi. 70. Linckiidae 41, 308. littoralis, Leptasterias 8, 370. longispina, Henricia 342, 348. Lophastcr pentactis 368. Lophaster stellans 366, 368, 372. loveni, Diplasterias 361. loveni, Podasterias 361. ludwigi, Luidia 336, 345, 346. liltkeni, Diplasterias 361. liitkeni, Podasterias 44, 51, 351, 361, 372. lutkenii, Asterias, 44, 82, 83. lutkenii, Pisaster n, 57, 68, 83, 344, 345, 362 ; pi. 40. lutkenii, var. australis, Pisaster, new variety 85, 88, 345. lutkenii, var. exquiseta, Pisaster 345. Luidia 6, 281, 315, 334. 34& Luidia asthenosoma 336, 345, 346. Luidia bellonas 367. Luidia brevispina 347. Luidia clathrata 7, 371 ; pi. 103. Luidia foliolata n, 334. 343, 345, 346; pis. 100, 103, 105. Luidia ludwigi 336, 345, 346. Luidia magellanica 367, 372. Luidia phragma 367. Luidiaster 311. Luidiaster dawsoni 312, 339, 343, 349; pis- 33, 34, 35- Luidiidae 283, 333. Luidiina 333. lunula, Henricia leviuscula, new vari- ety 213, 218, 342, 344; pi. 88. lurida, Asterias 337, 346. lurida, Cosmasterias 36, 48, 50, 358, 37i. lysasteria, Anasterias, new name 354, 355, 371. Lysasterias 354. macouni, Leptasterias, new species 13, 61, 124. macropora, Leptasterias 145. macropora, Stenasterias 13, 66, 145, 341, 348; pis. 50, 74, 84. Madreporite plate 36. magelhanica, Porania 368. magellanica, Luidia 367, 372. margaritifera, Asteracanthion 69. marginata, Enoplopatiria 263, 365, 371. mariana, Sporasterias 350. marsippus, Pteraster 273, 339, 347. Marthasterias 46, 47, 100, 358. Marthasterias glacialis 46, 47, 49, 370. Marthasterias sertulifera 19, 58, 100, 345, 349- Maternal instincts 10. media, Cycethra 366. Mediaster 285, 295. Mediaster sequalis 291, 296, 343, 344, 346, 348 r pis. 2, 3, 5. Mediaster bairdii 298, 371 ; pis. 2, 3. Mediaster stellatus 298. Mediasterinse 283, 294. meridionalis, Asterias 361, 372. meridionalis, Podasterias 361, 372. merriami, Orthasterias, new species 13, 65, 177, 342; pis. 18, 19, 75. mesacutus, Astropectinides 316, 321, 372. Meyenaster 54, 353. Meyenaster gelatinosus 54, 357, 371. migrata, Allasterias 373. Migratory habits 2, 4. miliaris, Henricia sanguinolenta 234, 37o; pl. 88. miliaris, Leptasterias epichlora, new subspecies 60, 61, 138, 341. militaris, Asterias 272. militaris, Pteraster 272, 278, 339, 343. millespina, Leptychaster 326. Mimaster cognatus 368. Mimasteridse 282. miniata, Asterias 264. miniala, Asterina 264. miniata, Patiria 14, 264, 343, 344, 346, 347, 348, 349 ; pl- 7, 108, 109. miniatus, Asteriscus 264. minuta, Anasterias 354, 355, 371. minuta, Asterias 107. minuta, Asterina 262. mira, Lahillia 363. mira, Poraniopsis 363, 372. INDEX 401 mirabilis, Tremaster 371. mite, Asteracanthion 358. Mithrodia bradleyi 373 ; pi. 107. Mithrodiidae 204. modesta, Asterinides 373. Mollusca, food for starfishes 3. Morphological features 20, 31. Mouth of starfishes 21. mulleri, Leptasterias 55, 124. multiclava, Asterias 59, no, 114, 338, 347 ; pis. 58, 59, 69, 84. multipes, Diplopteraster 279, 339. multipes, Pter aster 278, 279. multipes, Retaster 279. multispina, Henricia leviuscula 220, 338. multispinus, Pteraster 271, 343, 348. muricata, Coscinasterias 46, 49, 373- Myonota, suborder 283, 310. Myxasteridae 204. nana, Leptasterias aequalis, new vari- ety 61, 132, 341. 344, 345- Nanaster 146. Nanaster albulus 41. nanimensis, Asterias, new species 59, 105, 340; pl.6i. Nectria 287. Nectriinae 283. neglecta, Asterias 358. neglecta, Stolasterias 47. neglecta, Stylasterias 48, 370. nitida, Cycethra 366. nitida, Stephanasterias albula 147- nodiferus, Pisaster ochraceus 57, 72, 340, 344 ; pi. 56. nodosus, Goniaster 286. nodosus, Oreaster 286. nortonensis, Allasterias rathbuni 67, 191, 338, 347; pl- 78. Notomyota, order 310. nutrix, Granaster 372. nutrix, Stichaster 51, 362. obesa, Henricia 363, 372. obscurus, Hexaster 274, 275. obscurus, Hymenaster 267. obscurus, Pteraster 13, 17, 274, 339. obscurus, var. octaster, Pteraster 13, 276, 339, 347- 27 obtecta, Leptasterias, new species 60, 144, 338. obtusa, Callopatiria 373. obtusangulus, Goniaster 287. obtusangulus, Pseudoreaster 287. obtusispinosa, Asterias 358. ocellifera, Allopatiria 263. ochracea, Asterias 69, 77, 82. ochraceus, Pisaster n, 14, 57, 69, 72, 340, 344, 345; pl. 21. ochraceus, var. nodiferus, Pisaster 57, 72, 340, 344- octaster, Pteraster 274. octoradiatus, Solaster 368. oculata, Asterias 226. oculata, Cribella 226. oculaia, Cribrella 226. oculata, Henricia 226. oculata, Linckia 226. oculatus, Echinaster 226. Odontaster 303. Odontaster crassus 304, 345, 346, 349. Odontaster cremens 368. Odontaster granulosus 367, 372. Odontaster grayi 367, 372. Odontaster hispidus 349. Odontaster pedicellaris 367. Odontaster penicillatus 367, 372. Odontaster pilulatus 367, 372. Odontaster robustus 349. Odontasteridae 283, 302. Odontophores 37. cerstedii, Astropecten 318, 3i9, 346, 373- Ophiactis 14. Ophidiaster 309. Ophidiasteridae 282, 285, 308. Ophiocoma pumila, number of rays 14. ornatissimus, Astropecten 320, 346, 349- ornithopus, Linckia 310. Oreaster nodosus 286. Oreaster reticulatus 286. Oreasteridae 281, 282, 286. Orthasterias, new genus 48, 64, 168, 352, 374- Orthasterias biordinata, new species 48, 64, 172, 34i ; pis. 63, 82. 402 INDEX Orthasterias californica, new species 48, 64, 174, 185, 344, 350; pis. 68, 70, 80, 81. Orthasterias columbiana, new species ir, 48, 50, 64, 168, 178, 183, 341, 344, 346, 348 ; pis. 24, 35, 65, 78, 79, 109. Orthasterias dawsoni, new species 48, 65, 172, 175, 342; pis. 23, 75, 80, 81. Orthasterias eustyla 168. Orthasterias forcipulata II. Orthasterias forreri 19, 65, 179, 344, 345 ; pis. 65, 66, 70, 77, 80. Orthasterias forreri forcipulata 65, 1 80, 342; pis. 62, 70, 88. Orthasterias gonolena, new species 65, 184, 344, 345, 349; pis. 67, 68, 69,82. Orthasterias koehleri 65, 175, 183, 342. Orthasterias leptolena, new species 66, 182, 342; pis. 64, 77. Orthasterias merriami, new species 13, 65, 177, 342; pis. 18, 19, 75. Orthasterias (Stylasterias), new sub- genus 65. Orthasterias (Stylasterias) forreri 342. Orthasterias subangulosa, new name 168, 370- Orthasterias tanneri 48, 168, 348, 370; pis. 48, 109. Ortmannia 205. Ossicles, morphology of 27. Othilia 206. (Othilia) robustus, Echinaster, new species 207. Oyster beds damaged by starfishes 3. Oysters, food of starfishes 3, 5. pacificus, Leptychaster 326, 340, 347; pi. 74- Psedasterias, new genus 352, 353, 355, 374- Paedasterias chirophora, new genus 355, 371- pagenstecheri, Henricia 363, 372. Paleozoic Echinoderms 12. panopla, Urasterias 52, 182, 370. papillosa, Adelasterias 360, 372. papilla sa, Asterias 372. papillosa, Diplasterias 360. papillosa, Ganeria 365, 372. Papposa, Asterias 259. papposus, Crossaster 249, 259, 339, 343 ; pis. 5, 8, 9, 49. papposus, Solaster 259. papposus, Solaster (Polyaster) 259. papulosus, Pisaster n, 58, 68, 91, 340; pis. 42, 43, 60, 76, 80. Parachasteridce 310. Parapatiria crassa 263. Parasterias, new genus 53, 55, 66, 187, 374- Parasterias albertensis, new species 53, 55, i87, 342, 348; pis. 57, 70. Parastropecten 325. Parastropecten inermis 325. parvispina, Evasterias troschelii, new variety 63, 163, 34*- Patagonian species 2, 12, 15, 56, 353. patagonica, Porania 368. Patagonicus, Astrogonium 368. patagonicus, Ceramaster 291, 339, 343, 347, 349, 368. patagonicus, Pentagonaster 291. patagonicus, Pseudar chaster 368. Patiria 264. Patiria chilensis 348, 364, 372. Patiria coccinea 263, 373. Patiria gayi 364, 372. Patiria miniata 14, 264, 343, 344, 346, 347, 348, 349 ; pis. 7, 108, 109. Patiria pectinifera 349. Patiriella 372. Patiriella bispinosa 364, 372. Patiriella calcarata 364, 365, 372. Patiriella fimbriata 364, 365, 372. Patiriella pusilla 364, 365, 372. Patiriella regularis 263, 373. paucispina, Asterias 98. paucispinus, Pisaster 19, 98, 340, 344, 345 ; pl. 36. Paulia 287. paxillatus, Solaster 258, 339, 343, 347. Paxillosa, suborder 24, 283, 313. Paxillosa 280, 281. Paxillosce 280. paxillosus, Lebrunaster 365, 372. pectinata, Cribrella 230. INDEX 403 pectinata, Henricia 212. pectinata, Henricia sanguinolenta 229, 230; pi. 49. pectinifera, Patiria 349. Pectinodiscus 329. Pedicellariae of Asteriidse 23, 30, 35. pedicellaris, Odontaster 367. Pedicellaster 120, 127, 202. Pedicellaster antarcticus 367, 371. Pedicellaster scaber 368. Pedicellaster typicus 202. Pedicellasteridse 24, 26, 202. penicillaris, Asterinopsis 263, 373. penicillatus, Odontaster 367, 372. penicillatus, Solaster 262. Pentaceros 286. Pentacerotida 282, 285. pentactis, Lophaster 368. Pentagonaster 286, 288. Pentagonaster 289. Pentagonaster austro-granularis 369. Pentagonaster granularis 290. Pentagonaster patagonicus 291. Pentagonaster pulchellus 286. Pentagonasterince 283, 285, 289. pentagonulus, Goniodiscus 287. peregrinator, Diplopteraster 368. peregrinator, Retaster 368. Peribolaster biserialis 366. Peribolaster foliculata 366, 372. Perknaster densus 364. Perknaster fuscus 364. perrieri, Anasterias 51, 354. perrieri, Anasterias 355. perrieri, Asterias 354, 356. perrieri, Asterina 365, 372. perrieri, Asterinides 365, 372. perrieri, Sporasterias 353, 354, 356, 371- Persephonaster 327. perspicuus, Hymenaster 368. pertusa, Asterias 226. pertusa, Linckia 226. Phanerozona, order 24, 280, 282. Phataria 309. Phatatria biserialis 346. Phataria (Linckia) unifascialis, var. bifascialis 309. Philomaster 289. phragma, Luidia 367. phrygiana, Hippasteria 286, 301, 371; pis. 47, 48, 49- pilulatus, Gnathaster 367. pilulatus, Odontaster 367, 372. pinguis, Cycethra 366. Pisaster 18, 24, 28, 31, 33, 38, 39, 54, 56, 67, 72, 352. Pisaster brevispina 77. Pisaster brevispinus 57, 77, 93, 340, 344, 345; pis. 41, 44, 45, 69, 76. Pisaster capitatus 19, 57, 81, 344, 345 ; pis. 36, 56. Pisaster confertus 57, 73. 167, 340; pi. 38. Pisaster fissispina 76. Pisaster fissispinus 34, 57, 76, 344; pi. 39- Pisaster giganteus n, 57, 84, 87, 89, 98, 344J pl. 37- Pisaster grandis 13. Pisaster grayi 58, 91, 97, 344. Pisaster katherince Per. 58, 89, 91, 97. Pisaster liitkenii n, 57, 68, 83, 344, 345, 362. Pisaster liitkenii, var. australis, new variety 85, 88, 345- Pisaster liitkenii, var. exquiseta 345- Pisaster ochraceus n, 14, 57, 69» 72> 340, 344, 345 J pis. 21, 49, 56. Pisaster ochraceus, var. nodiferus 57, 72, 340, 344 ; pl. 56. Pisaster papulosus n, 58, 68, 91, 340 ; pis. 42, 43, 60, 76, 80. Pisaster paucispinus 19, 98, 340, 344, 345 ; pl. 36. planci, Acanthaster, 364, 373. planci, Asterias 364. planeta, Pontaster 369. Plates, skeletal 23, 28, 31. plena, Leptasterias epichlora, new sub- species 61, 140, 341. Plutonaster 281. Podasterias 8, 353, 361. Podasterias loveni 361. Podasterias liitkeni 44, 51, 351, 361, 372. Podasterias meridionalis 361, 372. Podasterias spinosa 361, 372. Podasterias steineni 351, 361, 372. polaris, Asterias 107, 108, 113, 370. 404 INDEX polaris, Ctenodiscus 330. polyacantha, Henricia 213. (Polyaster) papposus, Solaster 259. Polyasterias 45. Polyasterias fernandensis 360, 371. polygrammus, Cosmasterias 360, 371. polygrammus, Stichaster 49, 358, 360, polyplax, Allostichaster 373. polyplax, Asterias (Stichaster) 125. polyplax, Stichaster 362. polythela, Asterias 59, 104, 338, 347; pis. 55, 70, 72, 79, 84. Pontaster planeta 369. Pontasterinas 283, 31 1. Porania 281. Porania antarctica 368, 372. Porania glaber 368. Porania magelhcenica 368. Porania patagonica 368. Porania pygm&a 371. Poraniidce 304. Poraniinae 305. Poraniopsis 205. Poraniopsis echinaster 349, 363, 372. Poraniopsis inflata 206, 344, 346, 349, 363. Poraniopsis inflata flexibilis 206. Poraniopsis mira 363, 372. Porcellanaster 315. Porcellanasteridae 283, 328. Porcellanasterinae 283, 329. prastans, Cribrella 368. praestans, Henricia 368. problema, Asteracanthion 147. procurator, Ctenodiscus 333. Pseudarchaster 285. Pseudarchaster patagonicus 368. Pseudarchasterinae 283. Pseudoreaster obtusangulus 287. Psilaster 315. Psilaster fleuriaisi 369. Pteraster 8, 267. Pteraster afifinis 278. Pteraster aporus 277, 278. Pteraster capensis 278. Pteraster carribaeus 278. Pteraster cribrosus 278. Pteraster danae 278. Pteraster gracilis 271, 343, 348. Pteraster hebes 270, 343 ; pi. 96. Pteraster hexactis 274. Pteraster ingouffi 368. Pteraster lebruni 368. Pteraster marsippus 273, 339, 347. Pteraster militaris 272, 278, 339, 343. Pteraster multipes 278, 279. Pteraster multispinus 271, 343, 348. Pteraster obscurus 13, 17, 274, 339. Pteraster obscurus, var. octaster 13, 276, 339, 347- Pteraster octaster 274. Pteraster pulvillus 273, 278, 339. Pteraster reticulatus 268. Pteraster rugatus 278, 368. Pteraster semireticulatus 278. Pteraster stellifer 278, 368. Pteraster (Temnaster) hexactis 274. Pteraster tesselatus 268, 271, 339, 343, 348 ; pis. 32, 86, 97. Pteraster tesselatus arcuatus 269. Pteraster tesselatus hebes 270. Pterasteridae 204, 266. Pterasterides 277. Pterasterides aporus 278, 339, 347. Pterasterinae 266, 267. pugetana, Leptasterias epichlora, new subspecies 61, 142, 341. pulchellus, Pentagonaster 286. pulvillus, Pteraster 273, 278, 339. pusilla, Patiriella 364, 365, 372. Pycnopodia 4, 6, 12, 15, 39, 40, 43, 67, 68, 197. Pycnopodia helianthoides n, 35, 198, 342, 344, 348; pis. 29, 30, 31, 73, 74,88. Pycnopodidce 197. Pycnopodiinae 26, 197. pygmaea, Asterina 371. pygmcea, Porania 371. pygmaus, Ctenodiscus 371. Pythonaster 267. Pythonasteridae 204. radiosus, Labidiaster 352, 371. Randasia 287. Rathbun, Richard, species dedicated to 190. Rathbunaster 12. Rathbunaster californicus 197. INDEX 405 rathbuni, Allasterias 53, 55, 66, 67, J89, 337, 338, 347- rathbuni nortonensis, Allasterias 67, 191. 338, 347; pl- 78. Rays, development 28. Rays, number and variability 12-15, 38. Rays, spontaneous fission 39. regularis, Cycethra 366. regularis, Leptasterias epichlora, new variety 61, 139. 341- regularis, Patiriella 263, 373. regularis, Solaster 368. Reproductive organs 32. Retaster gibber 366, 372. Retaster multipes 279. Retaster peregrinator 368. Retaster verrucosus 366, 368, 372. reticulatus, Gonaster 286. reticulatus, Oreaster 286. reticulatus, Pteraster 268. Rhopia 209. Rhopia sepositus 363. ritteri, Bunodaster 316, 322, 345, 348; pis. 86, 104, 105. robusta, Ganeria 365. robusta, Stolasterias 373. robusta, Stylasterias 350, 373. robustus, Echinaster 342. robustus, Echinaster (Othilia), new species 207. robustus, Odontaster 349. rosea, Asterias 40, 362. rosea, Cribella 226. rosea, Stichastrella 40, 370. rosea, Stichaster 40. rubens, Asteracanthion 373. rubens, Asterias 44, 45, 68, 188, 337, 346. rubens, Asterias 107, 189. rudis, Evasterias troschelii, new vari- ety 63, 158, 341. rudis, Henricia sanguinolenta, new variety 233, 338, 347 ; pl. 86. rugatus, Pteraster 278, 368. rugispina, Asterias 51, 53, 353, 355- rugispina, Sporasterias 353, 356, 357, 371- rupicola, Asterias 356. rupicola, Sporasterias 51, 53, 353, 35 6, 371. rustica, Asterias 357. saanichensis , Asterias 132, 135. sanguinolenta, Asterias 226. sanguinolenta, Cribrella 223, 226. sanguinolenta, Echinaster 226. sanguinolenta, Henricia n, 14, 212, 226, 338, 342 ; pl. 49, 88. sanguinolenta miliaris, Henricia, new subspecies 234, 370; pl. 88. sanguinolenta, var. pectinata, Henricia 338; pl. 49- sanguinolenta, var. rudis, Henricia, new variety 233, 338, 347 ; pl. 86. sarsii, Echinaster 226. scaber, Pedicellaster 368. scalprifera, Smilasterias 53, 371. Sclerasterias guernei 44, 50, 370. scrobiculata, Echinaster 227. sebce, Goniodiscides 288. semireticulatus, Pteraster 278. Senses, instincts, memory 9. sentus, Echinaster 207. sepositus, Rhopia 363. sertulifera, Asterias 100. sertulifera, Marthasterias 19, 58, 100, 345, 349- sexradiata, Henricia 371. siderea, Enoplopatiria 365, 373 ; pl. 109. siderea, Leptasterias epichlora alas- kensis, new variety 61, 137, 341. siderealis, Astropecten, new species 317. 345, 346, 3495 pl- 50. simplex, Cribrella 368. simplex, Cycethra 366, 372. simplex, Henricia 368. singularis, Asterodon 367, 372. Skeletal plates, growth of 31. Skeleton of starfishes 20. sladeni, Cribraster 363, 372. Smell, sense of 9. Smilasterias 44, 53. Smilasterias scalprifera 53, 371. smithii, Echinaster 368. Solaris, Acanthaster 364, 372. Solaris, Echinaster 364. Solaster 4, 13, 15, 17, 18, 242. Solaster australis 366, 368, 372. 406 INDEX Solaster constellatus 257, 343, 348; pis. 46, 90, 93, 94. Solaster dawsoni 249, 339, 342, 348; pis. 46, 90, 91, 92. Solaster dawsoni arctica, new sub- species 252, 253, 347 ; pi. 87. Solaster decemradiatus 254. Solaster endeca n, 243, 244, 248, 249, 339 ; pis. 9, 87, 89. Solaster galaxides 248, 342, 348; pis. 46, 87, 89. Solaster octoradiatus 368. Solaster papposus 259. Solaster paxillatus 258, 339, 343, 347 Solaster penicillatus 262. Solaster (Polyaster) papposus 259. Solaster regularis 368. Solaster stimpsoni 250, 254, 339, 348; pis. 10, 11, 15, 46, 86, 94, 95. Solaster subarcuatus 368. Solaster vancouverensis 254. Solasteridae 41, 204, 242. Solasterince 242. spatulifera, Henricia 224. spatulifera, Henricia leviuscula, var. 224, 342, 344, 346 ; pis. 5, 14- Species, determination of 17. spectabile, Asteracanthion 358. spiculifera, Cribrella 220. spiculifera, Henricia 220. spiculifera, Henricia leviuscula 220, 338, 342; pis. 87, 107. Spines, 34, 36, 38. spinosa, Diplasterias 361, 372. spinosa, Hippasteria 301, 339, 343. 345, 346, 348; pis. 50,98. spinosa, Podasterias 361, 372. spinosus, Echinaster 207. Spinulosa 24, 203. spirabilis, Asterias 51, 354. spirabilis, Sporasterias 354. spitsbergensis, Asterias 148, 150. spitzbergensis, Ctenasterias 53, 55, 148, 370. Sporasterias 8, 53, 350, 353, 355- Sporasterias antarctica 51, 53, 353, 354, 371- Sporasterias cocosana 350. Sporasterias cunninghami 353. Sporasterias galapagensis 350. Sporasterias hyadesi 354. Sporasterias mariana 350. Sporasterias perrieri 353, 354, 356, 371- Sporasterias rugispina 353, 356, 357, Sporasterias rupicola 51, 53, 353, 356, 37i. Sporasterias spirabilis 354. Sporasterias varia 353. Sports or hybrids 17. steineni, Asterias 361. steineni, Diplasterias 361. steineni, Podasterias 351, 361, 372. stellans, Lophaster 366, 368, 372. stellatus, Mediaster 298. Stelleridse f orcipulatae 24. Stelleridae Spinulosse 203. stellifer, Pteraster 278, 368. stellionura, Asterias 51. Stellonia 101, 206. Stenasterias, new genus 145, 374. Stenasterias macropora 13, 66, 145, 341, 348; pis. 50, 74, 84. Stephanaster 286. Stephanasterias 36, 55, 130, 146. Stephanasterias albula 39, 40, 41, 55, 67, I47» 338. Stephanasterias albula, var. nitida 147. Stephanasterias gracilis 147. stichantha, Distolasterias 44, 49, 185, 373- stichantha, Stolasterias 47. Stichaster 30. Stichaster 146. Stichaster albulus 41, 147. Stichaster aurantiacus 40, 362. Stichaster f elipes 49. Stichaster nutrix 51, 351. Stichaster polygrammus 49, 358, 360, 37i. Stichaster polyplax 362. Stichaster rosea 40. Stichaster striatus 40, 362, 372. Stichasteridae 27, 42. Stichasterinae 26, 39, 40, 42, 146. Stichastrella, new genus 40, 374. Stichastrella rosea 40, 370. stimpsoni, Solaster 250, 254, 339, 348 ; pis. 10, 11, 15, 46, 86, 94, 95. INDEX 407 Stolasterias 43, 44, 46, 350. Stolasterias 45, 47. Stolasterias alexandri 372. Stolasterias candicans 367. Stolasterias neglecta 47. Stolasterias robusta 372. Stolasterias stichantha 47. (Stolasterias) tenuispina, Coscinas- terias 46, 49. Stolasterias volsellata 46. stolidota, Calvasterias 42, 354, 357, 371- Stomach of starfishes 21. striatus, Stichaster 40, 362, 372. studeri, Anasterias 355, 371. studeri, Asterias 51. studeri, Gastraster 360, 372. studeri, Henricia 363. Stylasterias, new subgenus 33, 48, 65, 179, 374- Stylasterias forreri 48, 50. Stylasterias neglecta 48, 370. Stylasterias robusta 350, 373. subangulosa, Orthasterias, new name 168, 370. subarcuatus, Solaster 368. subelectilis, Cycethra 366. subnodosa, Evasterias troschelii, new variety 63, 163, 34*. subnodulosa, Leptasterias epichlora, new variety 62, 139, 341. subregularis, Leptasterias epichlora, new variety 62, 73, 139, 341. Subspecies, definition of 17. Sucker-feet 4, 6, 15, 39. sulcifera, Asterias 48, 358, 361. sulcifera, Cosmasterias 48, 358, 360, 361, 37i. tanneri, Orthasterias 48, 168, 348, 370 ; pis. 48, 109. Temnaster 267. Temnaster hexactis 267, 274. Temperature during Glacial Period 2. Temperature of coast waters 2, 3. Temperature of water limits distri- bution 19. tenera, Anasterias 355, 371. tenera, Leptasterias 8, 370. tenuispina, Coscinasterias 38, 43, 45, 370. tenuispina, Coscinasterias (Stolaster- ias) 46, 49. tenuispinus, Echinaster 208, 346, 349, 373 ; pl. 107. tesselatus arcuatus, Pteraster 269. tesselatus, Goniaster 286. tesselatus hebes, Pteraster 270. tesselatus, Pteraster 268, 271, 339, 343, 348 ; pis. 32, 86, 97. Thyone briareus 10. tomidata, Asterias (Cosmasterias) 48. tomidata, Cosmasterias 48, 50, 359, 371. Tonia atlantica 40, 362. Tosia 287, 288, 289. Tosia arctica 292. Tosia (Ceramaster) granularis 290. Tosiaster, new genus 292. Tosiaster arcticus 292, 339; pis. 50, 99. Touch, sense 9. Tremaster mirabilis 371. troschelii, Asterias 151. troschelii, Evasterias n, 36, 51, 54, 63, 114, 151, 156, 341, 344, 348; pis. 22, 25, 26, 106. troschelii, var. alveolata, Evasterias, new variety 63, 157, 162, 341 ; pl. 62. troschelii, var. densa, Evasterias, new variety 63, 161, 341. troschelii, var. parvispina, Evasterias, new variety 63, 163, 341. troschelii, var. rudis, Evasterias, new variety 63, 158, 341. troschelii, var. subnodosa, Evasterias, new variety 63, 163, 341. tumida, Henricia 214, 234, 239, 240, 338, 347; pis. 12,87. tumida borealis, Henricia, new sub- species 236, 339, 342, 347 ; pis. 12, 88. turqueti, Asterias 372. turqueti, Cryptasterias 362, 372. turqueti, Diplasterias 362. Tylaster 305. typicus, Pedicellaster 202. typus, Linckia 211. 408 INDEX unifascialis, var. bifascialis, Phataria (Linckia) 309. Uraster 101. Urasterias 33, 34, 51, 55, 66, 187, 352, 374- ( Urasterias) forcipulata, Asterias 180. Urasterias gunner! 52. Urasterias linckii 33, 51, 55, 66, 181, 187, 370; pi. 70. Urasterias panopla 52, 182, 370. Valvaster 283. Valvasteridas 282. Valvasterin