1590,5 FI v,22:3 cop, 2 GENERAL FUNCTION OF THE GALL BLADDER FROM THE EVOLUTIONARY STANDPOINT BY FRANK W. GORHAM RESEARCH ASSISTANT, DEPARTMENT OF PHYSIOLOGY NORTHWESTERN UNIVERSITY AND ANDREW CONWAY IVY PROFESSOR OF PHYSIOLOGY, NORTHWESTERN UNIVERSITY ZOOLOGICAL SERIES FIELD MUSEUM OF NATURAL HISTORY VOLUME XXII, NUMBER S JUNE 21, 1938 PUBLICATION 417 GENERAL FUNCTION OF THE GALL BLADDER FROM THE EVOLUTIONARY STANDPOINT BY FRANK W. GORHAM RESEARCH ASSISTANT, DEPARTMENT OF PHYSIOLOGY NORTHWESTERN UNIVERSITY AND ANDREW CONWAY IVY PROFESSOR OF PHYSIOLOGY, NORTHWESTERN UNIVERSITY A NATURAL -v^> HISTORY ^ ZOOLOGICAL SERIES FIELD MUSEUM OF NATURAL HISTORY VOLUME XXII, NUMBER 3 JUNE 21, 1938 PUBLICATION 417 PRINTED IN THE UNITED STATES OF AMERICA BY FIELD MUSEUM PRESS GENERAL FUNCTION OF THE GALL BLADDER FROM THE EVOLUTIONARY STANDPOINT BY FRANK W. GORHAM AND ANDREW CONWAY IVY It has long been known to anatomists that certain animals do not possess a gall bladder. The evidence offered has usually been from individual dissections, and the explanations of the absence have been rather unsatisfactory. Therefore, an effort has been made to accumulate from the literature and dissections1 a list of species in which the state of the organ is known. From this list certain physiological conclusions can be drawn. This work was stimulated by the physiological investigations of Schmidt and Ivy (1937) who studied domestic and wild animals, some with, others without, a gall bladder. They found that animals varied in the amount of bile produced by the liver, the size and con- centrating ability of the gall bladder, the motility of the common duct, and the resistance offered by the choledochoduodenal junction to the flow of bile into the intestine. From their data, they concluded that animals without a gall bladder produce large quantities of bile, while those with a gall bladder which concentrates well, produce small quantities of bile. Between these extremes lies a series of intermediate types, and, in general, it may be said that the amount of bile formed by the liver varies inversely with the efficiency of the concentrating apparatus associated with the biliary ducts. These observations suggested that by ascertaining the presence or absence of the gall bladder throughout the various vertebrate orders, evolutionary evidence might be obtained pertaining to the function of the organ. Also, information would become available, serving to further physiological and possibly genetic research. PHYSIOLOGICAL EVIDENCE To indicate the academic and practical significance of this problem, an outline of the physiology of the organ will be given. General Functions Assigned to the Gall Bladder. — The general functions theoretically assigned to the gall bladder are those of serv- ing (a) as a bile reservoir for digestive purposes, and (6) as a pressure 1 Field Museum of Natural History has been most generous in allowing the authors to use its collections and libraries. Without the aid of its staff members, this work could scarcely have been begun. We desire to express our thanks especially to Dr. Wilfred H. Osgood, Mr. Colin C. Sanborn, Mr. Karl P. Schmidt, and Mr. Rudyerd Boulton. 159 160 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII regulatory mechanism. These functions are suggested by the follow- ing evidence: (1) A sphincter is present at the junction of the common bile duct with the duodenum, and this sphincter is necessary for filling the gall bladder. When the sphincter is relaxed the gall bladder does not fill. (2) In some animals the secretory pressure of bile is markedly less than the pressure necessary to overcome the sphincter. To prevent injurious back pressure on the liver when the sphincter is forcefully contracted, a regulatory device is essential. (3) Cholecystectomy in animals that have a relatively powerful sphincter, leads in most instances to dilation of the common ducts. (4) Those animals which do not possess a gall bladder, physiologically have no sphincter or only a very weak one. Thus, when the gall bladder is present, a sphincter (Sphincter of Oddi; Sphincter ductus choledocus) is essential for its filling; and when an efficient sphincter is present a gall bladder is necessary for the regulation of biliary pressure. It is evident, however, that these observed facts bear more directly on pressure regulatory function than on that of bile storage for digestive purposes. Concentrating Activity of the Gall Bladder. — In some animals, as man, the dog, and the cat, the storage function of the gall bladder is augmented by its ability to concentrate hepatic bile from five to ten times. In such animals, the gall bladder is literally a reservoir itself, of small volume but of large concentrating capacity. In other animals, as cattle, the gall bladder concentrates bile only twice or less. This ability of the organ to concentrate may be interpreted as supporting either or both of the aforementioned functions. Volume Output of Bile by the Liver. — Animals vary widely in regard to the amount of bile secreted per kilo of body weight or per gram of liver. For example, the guinea pig secretes from 154 to 220 cc. of bile per kilo of body weight per day. Man, however, only secretes 20 cc. of bile per kilo per day. Physiologic Capacity of the Gall Bladder. — Since the size of the gall bladder, the bile output, and the ability of the gall bladder to concentrate varies in different animals, the physiologic capacity will vary. By this term is meant that fraction of the daily output of bile which the gall bladder is capable of storing. If it were known with certainty that this bile was stored for digestive purposes, this should be a part of the definition. A contrast in physiologic capacity is demonstrated by man, who can store the bile secreted during 12 hours, and a guinea pig, which can store only the bile secreted for 12 minutes. 1938 GALL BLADDER— GORHAM AND IVY 161 An Additional Function of the Sphincter.— It should be considered that this structure probably prevents the regurgitation of duodenal contents into the common bile duct. This prevents ascending in- fection of the liver. It suggests that animals devoid of a competent sphincter should have some compensatory mechanism. Several such mechanisms do occur. There may be a one-way valve; large quanti- ties of bile may keep the ducts flushed out; or the extra-hepatic ducts may show peristaltic activity. Investigations show that the horse, which lacks both a competent sphincter and a gall bladder, has both a valve and a liver, which forms large quantities of bile. On the other hand, the adult pigeon,1 which also lacks a gall bladder, has motile ducts, and a liver forming large quantities of bile. Practical Significance of the Problem. — It is not generally agreed that the gall bladder serves for the storage of bile for digestive purposes. This is because physiological studies have not as yet demonstrated detectable disturbances of digestion or nutrition after cholecystectomy. However, when a gall bladder which concentrates efficiently is removed, certain changes are known to occur. These are (a) incompetence of the Sphincter of Oddi, with a variable return of competence; (6) dilation of the extra-hepatic ducts; (c) small areas of local necrosis in the liver. These changes certainly indicate that the gall bladder has a function, even if its general role is not to store bile for digestive purposes. Unfortunately, it is not known why all these changes occur, but dilation of the ducts is generally attributed to the return of competence of the sphincter. If this interpretation is correct, and if no direct digestive disturbances result from cholecystectomy, then the chief function of the organ in man, the dog, and the cat, for example, is to regulate the pressure in the extra-hepatic ducts, and this regulation is rendered necessary by the Sphincter of Oddi. Hence, from an evolutionary viewpoint, a gall bladder was developed secondarily to the sphincter, and the sphincter served primarily to prevent regurgitation from the intestine. However, given an animal whose liver produces relatively small quantities of bile, it is then reasonable to assume that such an animal might possess a better digestive apparatus if a gall bladder and a sphincter were present to render possible the storage of bile for diges- tive purposes. The following facts force the consideration of such a hypothesis: (a) Bile plays an important role in the digestion and absorption of fats; (6) it is essential for the absorption of vitamin D and carotene, the precursor of vitamin A; (c) it is important in the 1 A gall bladder is seen in the embryo. 162 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII absorption of iron and calcium; (d) all domestic animals that form a small quantity of bile have a gall bladder of large physiologic capacity; (e) the absorption of bile salts from the intestine augments the secretion of bile by the liver. From this, it seems that a reserve supply of concentrated bile, which may be discharged into the in- testine in small amounts during the first 20 or 30 minutes after a meal, would have a favorable effect in initiating digestive processes in the intestine. In fact, the gall bladder bile is sometimes referred to by physiologists as "ignition bile." If it can be established that the storage of bile for digestive purposes is a function of the gall bladder, it will be obvious that cholecystectomized patients, or patients with chronic disturbances of the biliary tract, should be fed frequently with the idea of promot- ing the flow of bile by the liver, and thus keep it thin and moving as in animals where there is no gall bladder. That frequent feeding should be employed is an old clinical impression still held by some internists, but such management will not generally be recognized as essential until its physiologic importance is proved. For example, formerly it was debated whether fats and acid fruits should be avoided by the patient with acute cholecystitis. Now, since it is known that fats and acids are potent excitants of gall bladder activity, doubt no longer exists. In summary, the physiological evidence indicates that the chief function of the gall bladder is pressure regulation; but the possibility that storage of bile for digestive purposes also occurs in those forms which concentrate bile, is one which cannot be easily dismissed. While this remains unproved, it may be surmised that the gall bladder developed secondarily to the appearance of a sphincter at the junction of the common duct with the intestine. ANATOMICAL EVIDENCE Bearing the physiology of the gall bladder well in mind, it should be interesting to examine various vertebrate groups for the organ, to ascertain its presence or absence, and to correlate other facts pertaining to the economy of the animal. Thus, diet, other modifications of the digestive tract, and even taxonomic relationships, may bear on the existence of the organ in an individual species. Little effort has been made to describe variations in the form or anatomical relationships of the organ. Suffice it to say that this hardly seems necessary, since the organ is either present or absent, and no series between the extremes can be easily demonstrated. 1938 GALL BLADDER — GORHAM AND IVY 163 "PREVERTEBRATES." — Among lower forms, the liver has not consolidated and migrated to become an organ distinct from the gut. In Balanoglossus, the liver is represented by structures called "hepatic caeca," which are groups of cells surrounding numerous ductlike processes which connect with the alimentary tract. In the sea- squirts (ascidians), a group of glands communicates with the stomach by a duct. This is referred to as the digestive gland or liver. In Amphioxus the liver is similar. It is thus seen that none of the known predecessors of higher vertebrates possess an organ resembling a gall bladder. The gall bladder then becomes a typical vertebrate structure and very characteristic of the group. CYCLOSTOMATA. — These are among the simplest vertebrates, but differ from all the others in lacking true jaws. The hagfish (Myxine) and the lamprey (Petromyzon) both have a gall bladder at some time during their lives. The adult hagfish has a bilobed liver, a gall bladder, but no discrete pancreas. The lamprey has a similar liver and gall bladder in the larval form, but the organ and its ducts are absent in the adult. FISHES. — Recent fishes are divisible into six orders. Two of these, the sharks and rays (Elasmobranchii), and the chimaeras (Holocephali) have cartilaginous skeletons. The others have true bony skeletons. They are the sturgeon-like fishes (Chondrostei); forms related to the dogfish (Holostei); the true bony fishes (Teleostei); and the lungfishes (Dipnoi). The sharks and rays are representatives of a relatively primitive group which prospered during Devonian times. They all apparently possess a gall bladder.1 The lungfishes are interesting in that they represent a group similar to that from which Amphibia arose. They, too, retain the organ. The teleosts are the most successful of recent fishes and have been dominant since Cretaceous times. In them the gall bladder is occasionally absent and neither Cuvier (1835) nor Owen (1846) could offer any explanation for the variability. In those fish (Bull- heads) which have been studied, the gall bladder was found to con- tract and evacuate on the administration of fatty foods, but its concentrating ability is not known. AMPHIBIA. — These all have a gall bladder in so far as they have been examined. This general statement is only superficially attested in the present work (Table 1). From the amphibians of the Coal Measures arose the reptiles which flourished during the Mesozoic, 1 J. F. Daniel: The Elasmobranch Fishes. Univ. Calif. Press, 1922, p. 139. 164 FIELD MUSEUM OF NATURAL HISTORY—ZOOLOGY, VOL. XXII and which gave origin during this period to the first mammals. Reptiles, therefore, are of current interest. REPTILES. — All existing forms possess a gall bladder. Early workers reported its occasional absence, but their information was apparently erroneous. In Table 2 the presence of the organ is recorded in 42 families and 70 species. An attempt is made in this, as in subsequent lists, to select genera characteristic of their respec- tive families. It is perhaps unfortunate that no recent reptile approximates those from which mammals arose. But it is probable that these and, in fact, all reptiles possessed gall bladders. Therefore it is reasonable to believe that all early mammals likewise had the organ. The habit of intermittent feeding in reptiles is well known. Some of them are carnivorous and others herbivorous. These habits support the suggestion that the gall bladder is a reservoir of bile for digestive purposes. However, nothing is known concerning the physiology of the organ in this group. BIRDS. -This group appears to have been evolved from reptiles at about the same time as mammals, but it has attained a high degree of specialization and the number of the existing species is very large. With the available information, conclusions are difficult. Tendencies are perceptible, however. The carnivorous birds (hawks, owls, etc.) all retain the organ, while the herbivorous forms (parrots, pigeons) and the insectivorous forms (woodpeckers) may retain it or lose it. In no order yet examined is the organ invariably absent. It has been suggested (by R. Boulton, of Field Museum staff) that the insectivorous birds which have lost the organ were immediately derived from an old herbivorous ancestry. An example of interest here is the Kea parrot of New Zealand, whose carnivorous habits are as recent as the introduction of sheep into that region. The list of species in Table 3 is given as a matter of record, with the hope that its evident deficiencies will stimulate investigation by those who have more available material. MAMMALS. — In this paper, mammals are of prime interest. This is largely because of man, and because it is more logical to cor- relate the physiology of man with that of other mammals than with that of some lower group. The orders will be considered separately. Monotremata. — The monotremes are in many respects the most primitive of living mammals, yet in some respects they are highly 1938 GALL BLADDER — GORHAM AND IVY 165 specialized. It seems not unlikely that they represent a line which descended directly, but independently, from the mammal-like reptiles. It may be recalled that they have a bill or beak devoid of teeth, and that they lay eggs but nourish their young on milk. The spiny anteater (Echidna) and the duck-bill (Ornithorhynchus) differ in their habits. One is terrestrial, and the other is semi- aquatic; one feeds on insects, and the other on shellfish and mollusks. Both have a gall bladder, and the cystic duct and the pancreatic duct join the common duct before it enters the intestine. Nothing is known concerning the physiology of the gall bladder or bile ducts. Marsupialia. — Marsupials are considered higher than monotremes because they have abandoned the oviparous method of reproduction and now bring forth very immature young which are nursed in the marsupium, a structure characteristic of this group. The American opossum is a simple form which represents an ideal ancestor which has apparently passed almost unchanged from Cretaceous times. At the end of the Cretaceous, placentals began to appear and later became dominant. Somewhat previous to this, Australia with her mammalian fauna became isolated. With no other competition, marsupials there radiated adaptively until almost every ecologic niche was filled. Thus, we have marsupial forms which parallel super- ficially almost every other mammalian type. Food habits vary widely. The opossums (Didelphiidae) are essentially omnivorous. Forms like the Tasmanian wolf (Dasyuridae) are usually carnivorous, but other members of the family are in- sectivorous. The marsupial mole (Notoryctes) represents a mono- typic family (Notoryctidae) which is insectivorous. Species allied to the Koala (Phascolarctidae) are herbivorous and the honey- bear, itself, eats only the young shoots of a single species of Euca- lyptus. The wombats (Wombatidae) are herbivorous. The kangaroos and wallabies (Macropodidae) are herbivorous, and the large kangaroo is said to practice a kind of rumination. The interest- ing American marsupial, Caenolestes (Caenolestidae), is insectivorous with an omnivorous tendency. The anteater, Myrmecobius, is the lone representative of its family (Myrmecobiidae). As might be expected, the stomach of marsupials shows adapta- tions related to the diet. The stomach is simple in the zoophagous, entomophagous, and carpophagous forms. In those types which feed on the bulkier parts of plants, the stomach becomes complex. The gall bladder is present in all marsupials (Table 5) and is of large 166 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII size, with no tendency to reduction in volume. Nothing is known concerning its concentrating ability. It contracts weakly, but the ducts and extraduodenal ampulla of the common duct are motile. The fact that all marsupials possess a gall bladder, even those which are strictly herbivorous, is of great interest. Among placental mammals, where the herbivorous type of diet has been assumed, the organ is absent in certain species. Remembering that marsupials are rather low in the scale of mammalian evolution, it would seem that the gall bladder is a primitive structure not easily lost. Table 5 affirms the presence of the organ in 26 species, representing the 10 existing families. Most living mammals belong to a more advanced group known as the placentals, in which a true placenta is constantly found. This structure permits a longer pre-natal period, and the need for the marsupial pouch is consequently lost. Other marked differences occur. Primitive placentals are known to have existed at the end of the Age of Reptiles. These primitive placentals were insectiv- orous, and from them arose the ancestors of our existing higher mammalian fauna. Members referable to the same order (Insecti- vora) as some of the early forms, still exist. Insectivora. — Table 6 records the existence of the gall bladder in 34 species, representing all families. The name of the order implies the nature of the diet, but a number of species eat fresh flesh when it is offered. Chiroptera. — Rather closely related to the insectivores, but of ancient and unknown origin are the bats. Bats have been very successful, and many families and species exist. They fall into two suborders, the Megachiroptera being the more primitive. The majority of these are large, and subsist on fruit and pollen. The stomach is rather complex. Most of the Microchiroptera are small and more highly specialized. The construction of the stomach differs according to the diet, which includes all types. Some 17 families and about 2,000 different species and subspecies of bats are said to exist. Table 8 records the gall bladder present in 13 families and 59 species. Therefore it is very probable that it is present in all bats. Dermoptera. — This order is represented by a single living genus, Galeopithecus (or Galeopterus). This animal, known as the flying lemur, is interesting, as it represents a form possibly similar to the 1938 GALL BLADDER — GORHAM AND IVY 167 progenitors of bats. It is a fair-sized, herbivorous, tree-living form with a moderately complex stomach. The gall bladder is present (Table 7). Primates. — Paleontologically and structurally, the primates are closely related to the primitive arboreal insectivores, the chief advances having been made in the development of a large brain, and stereoscopic vision. Three suborders are distinguishable. These are represented by the lemurs (Lemuroidea), by Tarsius (Tarsoidea), and by the true monkeys and apes (Anthropoidea). The lemurs :::::« ORGAN PRESENT — ORGAN ABSENT ?>? ORGAN VARIABLE FIG. 1. Probable history of the gall bladder in placental mammals (adapted from Romer). are the most primitive of living primates, and they possess many insectivore affinities. Between the lemurs and the true monkeys is Tarsius, possessing some of the features of each group. The anthro- poids include monkeys, apes, and man. They are separable into the New World forms (Platyrrhines) and the Old World forms (Catar- rhines). The latter include the family Hominidae, of which man is the only member. The diet of primates is quite varied. As a group they are omnivorous. The more strictly herbivorous forms subsist chiefly on the more concentrated portions of plants, such as the seeds or 168 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII fruit. The stomach of the lemurs and of Tarsius is simple. In the New World monkeys the stomach is simple except in the spider monkeys (Ateles), and the howlers (Alouatta), where there is some tendency to sacculation. In the Old World monkeys, the stomach is simple in the species which possess cheek pouches. In the langurs (Presbytis) and the guerezas (Colobu-s), which have no cheek pouches, the stomach is complex. In these monkeys, there is accumulation and retention of vegetable food in the stomach, and bezoars are not rarely found. The stomach reaches its maximum complexity in this group, which includes the sacred ape of India, one of the langurs. The gall bladder is present in representatives of all families of primates. Table 9 includes 68 species. Carnivora. Fossil evidence indicates clearly that all terrestrial fissiped carnivores arose from onecreodont family (Miacidae) probably since Eocene times. Eocene and Paleocene carnivores were known as creodonts and are directly descended from primitive insectivores of the preceding era. The ancestry of the marine forms is not known but is probably very similar. From the direct ancestry, diet, and feeding habits, these forms would be expected to possess constantly a gall bladder and this is well shown in Table 10 where 65 species representing all families were found to have the organ. Edentates. — Under the general term "edentates" convenience may allow an inclusive discussion of three unique mammalian orders, the Xenarthra, Pholidota, and Tubulidentata. Linnaeus, Cuvier, and other early investigators noted the queer dental structure and the lack of relationships with other orders, and therefore placed all these together in a distinct but somewhat heterogeneous order. Careful examination of anatomical and paleontological evidence failed to reveal any true relationships, and separate orders were indicated. The order Xenarthra includes the armadillos, anteaters, and sloths. Existing species are only a shadow of a previously large and varied fauna. Recent forms are, for the most part, highly specialized. The anteaters subsist mostly on termites; the armadillos are insectiv- orous but feed frequently on carrion; the sloths are herbivorous. As in other orders, the stomach is adapted to the diet. A gizzard- like pylorus occurs in the insectivorous forms, while the stomach tends to become complex in the sloths. The gall bladder is present in all forms except Bradypus, the three-toed sloth. It is significant that variation in this order first appears in the herbivorous types. However, the question immediately arises as to why the organ exists in Choloepus, the two-toed sloth. Has this species been long distinct 1938 GALL BLADDER— GORHAM AND IVY 169 or has it only recently diverged? The existing limited fauna with its paucity of species speaks against recent divergence and so also do marked structural differences other than the gall bladder. There- fore, Miller's (1923) classification is followed in which Choloepus stands as the only genus of a family. The order Pholidota, containing three genera, is the scaly ant- eater of the Old World. The two genera examined, Manis and Smutsia, possess a gall bladder and a gizzard-like pyloric antrum. OrycteropiLS is the single living representative of the order Tubu- lidentata. Its relationships are also unknown, but the work of Jepson (1932) suggests that the order was distinct by the end of the Eocene. The pyloric antrum is moderately thickened, and the animal possesses a gall bladder. The condition of the gall bladder among the Edentates is shown in Table 11. Artiodactyla. — Existing artiodactyls form three distinct groups: (a) Suina: pigs, peccaries, hippopotamuses. (6) Tylopoda: camels and llamas, (c) Pecora: chevrotains, deer, giraffes, prongbucks, and bovids. These suborders were all separate by the end of the Eocene, but more primitive forms are found earlier in this epoch which give evidence as to the ancestry of the whole group. Matthew (1909) and Gregory (1910) believe that the Artiodactyla have been derived from unknown creodonts allied to the Mesonychidae. These are carnivorous, so it is perhaps correct to say that a cow is more closely related to a lion than it is to a horse. Among the Suina, the true pigs (Suidae) constantly possess the gall bladder. The peccaries (Tayassuidae) have lost the organ. As to the hippopotamus, the gall bladder was absent in one of the four recorded dissections. More data would be desirable. It is interesting that the pig has a simple stomach, while that of the hippopotamus and the peccary is complex. It has even been suggested that the two latter ruminate. The Tylopoda, camels and llamas, have long been distinct from other artiodactyls. They have complex stomachs and ruminate. They lack the gall bladder. All Pecora probably arose from forms similar to existing Traguline deer, the chevrotains, which have a gall bladder. True deer (Cervidae) have probably all lost the organ, with the exception of the musk-deer (Moschus). This Asiatic form represents a species which is transitory between the true deer and their Traguline 170 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII ancestors. The retention of the gall bladder adds to the evidence afforded by the lack of horns and the excessive development of the canines, which indicates the primitive nature of this deer. The American prongbuck, not a true antelope, has a long separate history, and is the single living representative of the family Antilocapridae. It has a complex stomach and a gall bladder. The giraffes (Giraffidae) offer the curious possibility that the organ may be anomalously present in a very small percentage of cases. It has been reported in two of twenty recorded dissections. The* gall bladder is probably present in all members of the family Bovidae, with the exception of those in the subfamily Cephalophinae, a group of small African antelopes known as duikers in which it is constantly absent. The absence of the organ has been made a characteristic of this group by Pocock (1918). Crisp's (1862) report of its absence in two species of the subfamily Bubalinae should be rechecked in view of other errors made by this author. The report of the organ in two closely related groups by Garrod (1877) also needs confirmation. Tetracerus, a form which has long been juggled between the Cephalophinae and the Tragelaphinae, should by gall bladder evidence be classed with the latter, as is done by Weber (1927). In such an early distinct order, which has shown such great radiation and which has claimed so many now extinct genera, diversity of structure in existing species might be expected. The rather uniform presence or absence of the gall bladder in the various families lends significance to the gall bladder as an important struc- tural characteristic. Its occasional appearance in the giraffe and the primitive musk-deer is of interest in connection with their supposed Traguline origin. The retention of the organ in the Antilocapridae and most of the Bovidae lends support to the theory of the rather recent radiation of these families from the Tragulidae. The absence of the organ from the Cephalophinae may indicate a divergence of this subfamily from the Bovidae. At least, the presence or absence of the gall bladder follows quite well the taxonomy of this rather heterogeneous group (Table 12). "Sub-ungulates." — Three orders of mammals are generally referred to under this name. They are the Hyracoidea, Proboscidea, and the Sirenia. Although existing species show few affinities, the earlier representatives had many similarities which suggest a common origin (Romer, 1936). 1938 GALL BLADDER— GORH AM AND IVY 171 The Hyracoidea, known as conies or dassies, have one principal living genus (Procavia), confined to the Old World, and including various species. Superficially, they resemble rabbits, but dentition and foot structure unmistakably ally them to hoofed animals. Their stomach is rather complex and Lydekker reports that they ruminate. The Biblical l description is fitting: "Cheweth the cud, but divideth not the hoof." Records of dissections of these animals are confusing either because the gall bladder was variable, or the interpretation of the structure found was not uniform. The first explanation is probably correct. The order Proboscidea includes the elephants. These are highly specialized forms with a rather unique biliary anatomy. The gall bladder is absent but the common duct is wide, long, and has a reticulated mucosa. A large duodenal ampulla, called the terminal bile pouch, is present. It is divided irregularly into sacks and the pancreatic duct enters it. Owen (1866) says it is contractile. The structure is reminiscent of that found in some whales, and of the extra-duodenal ampulla of the guinea pig and opossum. The order Sirenia includes three recent families, one of which has become extinct in historic times. This last, Steller's sea-cow, apparently is the only one which lacks a gall bladder. This is difficult to explain, but the stomach is very complex. In animals which have evolved in the direction of these forms, a tendency to disappearance of the gall cyst is perceptible. In the elephants, this tendency is completely manifest; in the sirenia and hyraces it is less so. It seems that the latter are about to lose the organ. There is a pouch in the elephant which seems analogous to structures seen in some whales, but the elephant is a herbivorous type and the need for this compensatory structure is not obvious. Perissodactyla. — This order, once more abundant, is now repre- sented by horses, tapirs, and rhinoceroses. In many respects, these forms have become highly specialized. It is well to point out that the order is an ancient one, first appearing at the base of the Eocene. When curiosity is expressed concerning the reason why two animals whose habits are as similar as those of the horse and the cow differ in respect to the gall bladder, it should be pointed out that they are of diverse ancestry. Thus, the gall bladder is constantly absent in the Perissodactyla (Table 14), 1 Leviticus 11:5. 172 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII but only occasionally so among the Artiodactyla. The stomach in this order is simple, but the immense size of the caecum is almost characteristic. In the wall of the common duct are some microscopic sacculi which may have compensatory significance. These are found in the horse but not in the tapir. Cetacea.— Whales are highly specialized marine mammals of very wide distribution. Unlike most other aquatic mammals, they are incapable of any locomotion on land. Structural vestiges, how- ever, indicate that they were not always so confined and paleontolo- gists agree that they were probably derived from terrestrial carnivores some time in the Paleocene. By the end of the Eocene, they consti- tuted a distinct mammalian order. Thus, as were the horses so were the whales, exposed to evolutionary tendencies over a long period. Table 15 shows that the gall bladder was absent in all except one of the species recorded, and examination of this exception (Williams, 1838) leaves the impression that a mistake might well have been made. The absence of the gall bladder in the Cetacea is indeed odd, for these forms are highly carnivorous. They have very complex stomachs like ruminants, but the proventriculus is aglandular. The whale-bone whales (Mysticeti) feed on small marine invertebrates like cuttle-fish and squid; the toothed whales (Odontoceti) feed on fish and at least one genus, Orca, the killer, feeds on seals. The nature of the whale's diet makes continuous feeding highly improb- able. But the complex stomach and the observation that whole food is regurgitated when the animals are harpooned, led Beddard (1900) to suggest that the animals feed hurriedly and store their food in an accessory stomach. The digestion of this food would then be protracted over several hours. In Tursiops (Hein, 1915), and probably a good many other Odontoceti (Weissberg, 1933), a dilated bile reservoir is found in the course of the common duct. Into this drains the pancreatic duct, and between the reservoir and the intestinal outlet the common duct contains a valve similar to the valvula spiralis of the human cystic duct. This structure is very interesting and may represent a true compensatory mechanism. Were these animals of herbivorous ancestry, such biliary anatomy might be explained by suggesting that the whale was an animal which lost the gall cyst, changed its habits, and, when the need for the organ reappeared, developed the structure which is described above. With a proven carnivorous ancestry, 1938 GALL BLADDER— GORH AM AND IVY 173 however, explanation is difficult, but it may be associated with the complex stomach, or a depth-pressure modification. Rodentia. — Rodents are known from a long paleontological record, and they were early divisible into two suborders. One, Simplici- dentata, has two upper incisors; the other, Duplicidentata, has four. The rabbits and their allies comprise the latter group. More superficial divisions are quite natural. The squirrel-like forms, Sciuromorpha, are the simplest and the most primitive genera. Radiation at an early era resulted in our present diverse sciuromorph fauna, including six rather distinct families. Mouselike rodents, Myomorpha, are a more recent offshoot from early rodent lineage. Structural modifications indicate a degree of specialization not seen in the squirrel-like forms. Three families are included among the existing genera. The dormice, Myoxidae, retain a simplicity of structure indicating that forms similar to them may have been ancestral to the more highly specialized Dipodidae and Muridae. The jerboas, Dipodidae, are saltatorial forms of un- known ancestry. However, the greatest number of living rodents are found in the true rats and mice, the Muridae. Excepting man, they are the most successful of recent mammals, and, like man, their terrestrial range is practically unlimited. Radiation was apparently from the Old World. The other division of simplicidentate rodents, the Hystrico- morpha, includes a variety of forms of which the porcupine and the guinea pig are representative. The hystricid rodents are largely confined to South America, on which continent a great multiplicity of species has developed. The rabbits and hares have been placed in a distinct order (Lagomorpha) by many authorities. The possession of a pair of extra incisors was a differential point in Oligocene times, but morphological similarities probably best find recognition if a single order is used. Rodents are essentially herbivorous, but a number of them will accept a more omnivorous diet. The families may be considered separately: Aplodontiidae: These live near mountain streams and in dense vegetation of the Pacific northwest. They feed on various green plants. Sciuridae: These feed chiefly on nuts, seeds, and grass. They probably occasionally eat birds' eggs and insects. 174 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII Castoridae: Beavers are aquatically adapted forms which fell trees. The bark of the trees is utilized for food. Heteromyidae: Kangaroo rats and mice of the Americas which live abundantly in arid, subdesert regions where plants bloom freely for only a few days during the year. The result is a large number of seeds. These are gathered and stored by the animals for future consumption. Apparently they hibernate. Geomyidae: Pocket gophers are fossorial and eat roots, bulbs, grass, and seeds. They may feed more or less constantly. They do not hibernate, but tunnel extensively under the snow when necessary. Pedetidae: These are fossorial and saltatorial forms, the Cape Jumping Hares. Their food seems to be entirely of a vegetable nature. Myoxidae: Arboreal and nocturnal creatures of small size. They feed on nuts and seeds, and probably hibernate. Dipodidae: Feed upon buds, leaves, twigs, and many kinds of plants; on seeds, grain, wild berries, chestnuts, acorns, grass, and bark. Spalacidae: The Cape Mole-rats live in subterranean burrows, which they dig in search for bulbs and roots. Muridae: This family includes a variety of rats and mice, whose habits were probably originally herbivorous. They easily modify their needs to the available supply. Their diet has in many cases become that of man, whom they parasitize. Bathyergidae: Members of this group are fossorial and subsist on a vegetable diet. Hystricidae: Old World porcupines whose food is entirely vegetable and consists mainly of roots. Erethizontidae: Arboreal forms, which eat the bark of trees. They do not hibernate. They are restricted to the New World. Dasyproctidae: The agoutis are tropical rodents, whose food consists of foliage, roots of ferns, fallen fruit, and possibly nuts. Caviidae: The best-known member of this family is the common guinea pig. In their natural state, the cavies feed on roots, corn, and other vegetable substances. Chinchillidae: Grass and roots form the chief substance of their diet. Long arid seasons may deprive these animals of water for considerable periods, but they seem to survive on the dried grass. 1938 GALL BLADDER— GORHAM AND IVY 175 Capromyidae: Arboreal forms which live in the dense forest. Feed on fruits, leaves, and bark, but may also eat the flesh of small animals, particularly that of a kind of lizard. Octodontidae: The octodonts are represented by both African and South American variations. Both of these are herbivorous. Thryonomyidae: The African cane-rat which digs for roots and ground nuts. Extensive runway systems are formed under the grass and reeds. Ctenodactylidae: An African diurnal form. It lives among rocks and is herbivorous. Leporidae: The rabbits are diurnal and strictly herbivorous. Ochotonidae: These interesting mammals live in alpine rock slides. They gather grass which they stack and allow to dry. This supply keeps them during the winter months. It has been impossible to investigate thoroughly all the available rodent material, but a few noteworthy points may be mentioned. The stomachs of rodents vary markedly in their form and even in their histologic structure. This variation has been the subject of at least one paper.1 Arrangement of rodent families into two groups, one with simple stomachs, and the other with complex stomachs, failed to reveal any correlation with the presence or absence of a gall bladder. The gall bladder is present in some families, absent in others, and variable in still others (Table 16). Further investigations are desirable and will probably alter these lists: Present Castoridae Spalacidae Capromyidae Anomaluridae Bathyergidae Thryonomyidae Myoxidae Dasyproctidae Leporidae Dipodidae Cayiidae Ochotonidae Chinchillidae Absent Aplodontiidae Heterpmyidae Petromyidae Geomyidae Pedetidae Erethizontidae Inconstant Sciuridae Hystricidae Muridae Octodontidae Conclusions are not accurate with such meager evidence. The whole order seems to be in a rather plastic state. Evolutionary tendencies are perceptible. The gall bladder is absent in the Aplodontiidae. In the Sciuri- dae it is usually present. The Pedetidae are intermediate between 1 K. Toepfer. Die Morphologic des Magens der Rodentia. Morph. Jahrb., 17. 176 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII the Sciuromorpha and Myomorpha and, as they are in other respects highly specialized, the gall bladder absence is not surprising. How- ever, the apparently close relationship which these forms have to the Anomaluridae, which are cholecystous, and the variation of the organ in the existing Myoidea, suggest an origin of this latter group from separate forms, some in which the organ was present and others in which it was absent. The geomyids and heteromyids are closely related and have been distinct since Oligocene times. Among the Myomorpha, the Myoxidae, Dipodidae, and the Spalacidae show simpler dentition than the Muridae. These three possess a gall bladder. The Muridae claims more genera than any other mammalian family and probably more individuals. In this family, the gall bladder is constantly present in some genera and absent in others. Its presence or absence may even be found in different species of the same genus. Some tendencies among the subfamilies are distinguishable. The Cricetinae, Lophiomyinae, and Hydromyinae usually do not possess the organ, while the Gerbillinae apparently tend to retain it. Among the Murinae proper, available records (nine genera) indicate vari- ability in this subfamily. The present distribution and the large number of species and subspecies show that these rodents are not only very successful, but that they are at a morphologically plastic stage in their development. The habits of the group and the variable presence of the gall cyst hint at its eventual disappearance. Hystricomorph rodents generally possess a gall bladder, but in a few instances the organ is absent. The New World porcupines, the Erethizontidae, all lack the organ, but it was present in six out of eight dissections of the Old World forms. The Erethizontidae have been distinct since the Oligocene. DISCUSSION From the above evidence, several facts are forthcoming. In the first place, it will be seen that the gall bladder is present in all carnivorous forms (except whales), and that it may be lacking among omnivorous or herbivorous forms. A carnivorous animal of neces- sity consumes only occasional meals, and these have a high fat content. From a physiological point of view, storage of concentrated bile should be a great aid in the digestion of such meals, and it is significant that the organ is constantly present in animals having these habits. 1938 GALL BLADDER— GORHAM AND IVY 177 The absence of the organ in species of other dietary habits, is indicated by the evidence of Schmidt and Ivy. They found species in which the organ had no apparent or demonstrable function. It may be said that the organ was "physiologically absent." Species possessing this type of gall bladder are usually members of an order in which there are other species in which the organ is anatomically absent. Further, the gall bladder is not absent from those species which have descended most directly, or with the least specialization in regard to body form, from the ancestor of mammals. Another factor which may influence the existence of the gall bladder in a given species is the proximity of this species to the ancestral form. It will be noted that those species, families, or orders which most closely resemble the ancestor of a given group tend constantly to retain the gall cyst, while those which vary most, tend to lose the organ. The fact that other parts of the digestive tract adapt themselves quickly to dietary habits, suggests that the gall cyst may be the least plastic of the digestive organs. It is true, however, that it has completely disappeared in some mammalian families, whereas the stomach, caecum, and colon have not, although they frequently have undergone marked variations in form. The stable nature of the gall bladder suggests that it may have taxonomic value. In only nine families was the organ found to be present in some members and absent in others. In at least six of these the variation occurred between species or individuals. In the remaining three, it was possible to arrange subfamilies, and here, for the most part, definite tendencies following the lines of arrange- ment were demonstrable. Individual variations in the organ are probably comparable to the anomalies occasionally seen in the human cadaver. The frequency with which these anomalies occur may be a measure of the declining "need" for the organ. Thus in the dissections of Procavia, three observers found a gall bladder; five did not. On the other hand, with the giraffe the records show only two gall bladders out of about twenty dissections. From these records, it might be concluded that these animals will soon lose the organ completely. It is interesting that closely related species, particularly among the Muridae, may differ in the possession of the organ. Besides this, the occurrence of anomalous absences in individuals of the same species suggests that it might be possible to study the mechan- ism of the inheritance of the organ. 178 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII CONCLUSIONS CD The gall bladder is a typical vertebrate structure, found first in larval agnathostomes and generally throughout the rest of the group. (2) All reptiles and amphibians have a gall bladder. (3) In birds the presence of the gall cyst is extremely variable, but carnivorous birds generally retain it. (4) The gall bladder is a primitive mammalian structure. It is not easily lost. It is not as readily lost as the form of the stomach, caecum, and colon is changed; i.e. a complex stomach may appear in a line of descent without the gall bladder's being lost. It is usually retained in forms which most closely resemble the ancestral type; forms which vary most widely may lose it. (5) Carnivorous mammals (except whales) possess a gall bladder, while mammals with other dietary habits may lose the organ. This apparently is related to the intermittent feeding habits of carnivorous types. Herbivorous forms with continuous feeding habits are most likely to lack the organ. (6) From the evidence of the dietary habits of cholecystous and acholecystous forms, one may conclude that if the gall bladder is removed from a given individual of any species, digestive efficiency will be diminished only if (a) the organ has a large physiologic capacity, and (6) intermittent meals of a high fat content are given. (7) There is no anatomical evidence to indicate that the Sphincter ductus choledochus developed before a gall cyst. Yet the presence of such a sphincter renders a gall bladder essential as a pressure regulatory apparatus, unless the common bile duct manifests definite peristaltic activity, as, for example, in the guinea pig. 1938 GALL BLADDER— GORHAM AND IVY 179 TABLES Unless otherwise indicated all numbers are catalogue numbers of specimens in Field Museum. The presence of the gall bladder is indicated by + its absence by — . The name in parentheses is the one used by the original author. Refer- ences to Gorham, 1936, are to Field Museum specimens dissected but not preserved. TABLE 1. — AMPHIBIA Apoda Caecilidae + Typhlonectes compressicauda ] + Typhlonectes natans [ Fuhrmann 1914. + Typhlonectes dorsalis J Salientia Pipidae +Pipa pipa (P. americana) Beddard 1895a. Pelobatidae -+Megophrys monlana (Xenophrys monticola) } +Megophrys hasseltii (Leptobatrachium) > Beddard 1907a. +Megophrys feae (Megalophrys) J Rhinophrynidae -\-Rhinophrynus dorsalis Giinther 1858. Leptodactylidae +Pseudis paradoxa (Rana) Hunter 1861. Microhylidae +Breviceps verrucosus Beddard 1908. Ranidae +Rana pipiens Gorham 1936. Caudata Necturidae -\-Megalobatrachus japonicus Beddard 1903. +Cryptobranchus alleghaniensis (Menopoma) Sirenidae + Siren lacertina Amphiumidae +Amphiuma means (A. didactylum) . . . . Salamandridae -\-Salamandra salamandra (S. maculata) Hunter 1861. TABLE 2. — REPTILIA Sauria Gekkonidae +Gekko gecko 8914. Eublepharidae +Coleonyx mitratus 5051. Pygopodidae +Lialis jicari 13869. Agamidae +Agama agama 19815. +Calotes mystaceus 14492. -i-Chlamydosaurus kingi Beddard 1905. Iguanidae -\-Crotaphytus collaris 637. +Phrynosoma blainvilli 8056. Cordylidae +Cordylus giganteus 19258. +Platysaurus guttatus 17322. -\-Chamaesaura aenea. . . . 17465. 180 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 2. — REPTILIA (Continued) Sauria (continued) Anguidae +Gerrhonotus infernalis 11200. +Anguis fragilis 22895. +Ophisaur us apus 1 5680. Anniellidae +Anniella pulchra Coe & Kunkel 1906- Helodermatidae +Heloderma suspectum Shufeldt 1890. Varanidae + Varanus niloticus Beddard 1907. Xantusiidae -\-Lepidophyma flavomaculata 21788. Teiidae +Ameiva ameiva 16585. Amphisbaenidae -\-Amphisbaena alba 17802. Lacertidae +Lacerta viridis 15749. Gerrhosauridae +Gerrhosaurus nigrolineatus 13506. Scincidae +Eumeces schneideri 19636. Feyliniidae -\-Typhlosaurus vermis 16031. Dibamidae -\-Dibamus novae-guineae 14251. Chamaeleontidae -i-Chamaeleo pumilis (Chamaeleon) 1 +Chamaeleo par Mob™ (Chamaeleon) .. : Beddard 190Tb> +Chamaeleo dilepis (Chamaeleon) i * — Chamaeko verrucosus (Chamaeleon) i +Chamaeleo verrucosus (Chamaeleon) 18277. -\-Rhampholeon spectrum 19855. Serpentia Typhlopidae + Typhlops punctatus 21081. Leptotyphlopidae -\-Leptotyphlops albifrons 87. Boidae +Charina bottae Cope 1898. +Constrictor constrictor 1 1404. +Sanzinia madagascarensis (Corallus) Beddard 1906a. Pythonidae +Python bivittatus 8925. +Python sebae Beddard 1904. Aniliidae +Anilius scytale 16943. +Anilius scytale (Ilysia) Beddard 1906. + Anilius scytale (Boa) Owen 1833a. •Beddard could not find the organ. 1938 GALL BLADDER— GORHAM AND IVY 181 TABLE 2. — REPTILIA (Continued) Serpentia (continued) Uropeltidae +Rhinophis blythii • \ peters 1861. +Rhinophis oxyrhynchus ) Xenopeltidae / ^523 +Xenopeliis unicolor | Thompson 1913. Achrochordidae +Chersydrus granulatus } Colubridae I Cope 1898. +Elaphe quadrivitlata +Heterodon contortrix • J +Coluber constrictor Gorham Dasypeltidae +Dasypellis macrops Homalopsidae +Enhydris enhydris Boigidae -\-Boiga dendrophila Disteiridae +Pelamis platurus Elapidae +Naja nigricottis • • • • • • • • +Naja hannah Beddard +M icrurus fulmus Amblycephalidae + Amblycephcdus moellendorffii Viperidae + Vipera berus -17bb. Crotalidae +Crotalus riridis (C. confluentus) ^OP« ^ y8- +Crotalus horridus Owen 1833a. Testudinata Dermochelidae +Dermochelys coriacea Burne lyuo. Chelhydridae -rChelydra serpentina Martin 1830b. Testudinidae +Emys orbicularis (E. lutraria) Hunter 1 bl. + Testudo elephaniina (T. indica) Martin 1 30. + Testudo elephantina (T. elephantopus) Hunter 1 61. + Testudo graeca Martin 1830a. +Geoemyda trijuga (Emys) • j Anderson 1879. -f Kachuga ahongoka J +Clemmys leprosa • j Kollman 1912. -\-Testudo mauritanica J Cheloniidae +Chelonia mydas (Chelone) Hunter 1861. Crocodilia Crocodilidae +Crocodylus cataphractus (C. leptorhyncus) Martin 183ob. +Crocodylus niloticus ("Crocodile du Nile") Geoffrey 1803. +Crocodylus acutus Owen 1831d. Rhynchocephalia Sphenodontidae +Sphenodon punctatus (Hatteria) Giinther 1867. 182 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 3. — AVES Struthioniformes Struthionidae f Garrod & Darwin 1872, -Slruihio camelus \ Hunter 1861. Rheiformes I Rothschild 1900. - Rheidae Rothschild 1900 Casuariiformes Casuariidae +Casuarius ] Dromiceiidae | Rothschild 1900 + Dromiceius J -\-Dromiceius novae-hollandiae Boulart 1900 Apterygiformes Apterygidae +Apteryx aiistralis Owen 1838 Sphenisciformes Spheniscidae -\-Aplenodyles patagonica Reid 1835 Procellariiformes +Procellariidae . . Forbes 1882a Pelecaniformes Phaethontidae +Phaethon lepturus (P. flavirostris) Beddard 1897a Pelecanidae +Pelecanus rufescens Martin 1835a -\-Pelecanus occidentalis (P. fuscus) -\-Pelecanus onocrotalus Phalacrocoracidae +Phalacrocorax carbo J Anhingidae +Anhinga anhinga (Plotus) Garrod 1876b. Ciconiiformes Ardeidae +Ardea cinerea. +Ardea purpurea (Nycticorax). Hunter 1861. +Botaurus stellaris. +Nyclicorax nycticorax (N. europaeus) . Cochleariidae +Cochlearius cochlearius (Cancroma) Murie 1867a. Balaenicipitidae f Beddard 1888. +Balaeniceps rex . . -j Fox 1929. Scopidae 1 Mitche11 1913- +Scopus umbretta Beddard 1884. Ciconiidae +Leptoptilos crumeniferus Mentzer 1929. Phoenicopteridae -\-Phoenicoplerus ruber Hunter 1861. Anseriformes Anhimidae +Anhima cornida (Palamedea corniculaia) Beddard 1894. 1938 GALL BLADDER— GORHAM AND IVY 183 TABLE 3. — AVES (Continued) Anseriformes (continued) Anatidae -\-Cygnus olor \ u..nt.or i ofi1 +Branla canadensis (Anser) / Hunter 1 +Branta bernicla Fox 1923. +Branta leucopsis \ u, . , aft1 +Cairina moschata (Anser) / Hunter 1 +Eulabeia indica Fox 1927. + Mergellus albellus (Mergus) Kuhl 1820. Falconiformes Accipitridae +Gyps fulvus (Vultur) , +Aquila chrysaetos. +Haliaeetus albicilla +Accipiter nisus (Astur) +Accipiter gentilis (Astur palumbarius) 4-Aviceda leuphotes Hunter 1861. .97002. Falconidae — Falco peregrinus Kuhl 1820. Galliformes Cracidae +Penelope purpurascens (P. cristata) Hunter 1861. Phasianidae +Arborophila brunneopeclus 97003. Numididae +Numida meleagris Hunter 1861. Opisthocomidae — Opisthocomus hoazin (O. cristatus) Young 1888. Gruiformes Turnicidae -Turnix tanki : 97007. Gruidae +Anlhropoldes virgo (Grus) Hunter 1861. Aramidae +Aramus scolopaceus Garrod 1876a. Rallidae -\-Porphyrio albus Hunter 1861. +Notornis mantelli Benham 1899. Heliornithidae — Podica senegalensis Beddard 1890a. Rhynochetidae +Rhynochetos jubatus Murie 1867a. Cariamidae +Cariama cristatus (Dicholophus) Martin 1836. Otidae +Otis tarda Fox 1929. Charadriiformes Jacanidae + Jacana spinosa (Parra) Forbes 1881a. Charadriidae + Vanellus vanellus (V. vulgaris) . Scolopacidae I K ,, -Erolia alpina (Tringa) f Kutl -\-Calidris canutus (Arenaria calidria) I 4- Numenius arqualu Hunter 1861. -\-Limnodromus griseus } Phalaropodidae [ Gorham 1936. +Lobipes lobatus J 184 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 3. — AVES (Continued) Charadriiformes (continued) Laridae + Larus marinus Hunter 1861. •\-Larus argentatus Hill 1926. + Lanis ridibundus Kuhl 1820. Alcidae + ('ria aalge (iroile) Hunter 1861. + Uria aalge Kuhl 1820. +SynthUborhamphus antiques 1 g^g^t 1887 + Brachyramphus marmoratus / Columbiformes Pteroclididae + Pterocles orientalis (P. arenarius) | Garrod 18741 + Syrrhaple8 Garrod 1874. Columbidae +Ptilinopus insolitus (Aedirhinus) \ u -.....-]] -i ooo +Coryphoenas crassirostris (Turacoena) / ^ — Columba .... -Turtur — Macropygia . — Ectopistes . . . — Chamaepelia. — Metriopelia . . — Zenaida — Caloenas ... — Didnnculus. . — Chalcopelia . — Tympanistria Garrod 1874. — Ocyphaps. — Leucosarda . . — Phaps — Phlogoenas . . — Starnocnas . . — Geopelia .... — Goiira +Carpophaga . +Lopholaemus. -\-Ptilonopits . . — Treron — Columba vitiensis halmaheira (lanthoenas leucolaema) . . . . \ r . 187r; -Alectroenas pulcherrima (Erythroenas) / u Psittaciformes f Cuvier 1835. - Psittacidae -I Beddard 1898. { Fox 1923. -\-Kakaioe goffini (Cacatua) 1 +Kakaloe moluccensis (Cacatua) I /-arrofi isyyv, -\-Kakatoe haematnropygia (Cacatua philippinarum) [ +Nymphicus hollandicus (Calopsitta novae-hollandiae) . . . . j Cuculi formes Musophagidae +Gallirex porphyreolophus (Corythaix) Owen 1834. + Turacuf persa (Corythaix buffoni) Martin 1836a. Cuculidae +Carpococcyx radiatus Beddard 1901a. + Scythrops notae-hollandiae Beddard 1898a. +Cucnlus canorus Hunter 1861. +Ccnlropn* sinensis 97014. 1938 GALL BLADDER— GORHAM AND IVY 185 TABLE 3. — AVES (Confirmed) Strigiformes Strigidae + Scolopelia peli Murie 1871. + Phodilus badius Beddard 1890. +Asio otus (Otus aurifa) Hunter 1861. Caprimulgiformes Steatornithidae +Steatornis Beddard 1886a. Podargidae +Batracho8tomus Blyth 1866. + Batrachostomus . +Podargus Aegothelidae + Aegotheles . Beddard 1886a. Caprimulgidae -\-Caprimulgus — Chordeiles sp +Chordeiles minor minor Gorham 1936. +Nyctidromus Beddard 1886a. Micropodiformes Trochilidae — Archilochus colubris Crisp 1862a. — Campylopterus ensipennis 97208. Trogoniformes Trogonidae +Harpactes erythrocephalus 97017. Coracii formes Alcedinidae — Halcyon smyrnensis 97019. Meropidae +Melittophagus erythrocephalus 97187. Leptosomatidae +LeplosomuS discolor. . . { ^andld^and Milne-Edwards 1875. Bucerotidae +Bucorvns abyssinicus Garrod 1876. +Dichoceros bicornis (Buceros cavatus) Owen 1833. Piciformes Capitonidae + Trachylaemus goffinii 97232. -\-Megalaema trirens +Cyanops franklinii (Megalaema) -\-Xaniholaema rosea Ramphastidae +Aulacorhynchus prasinus (Aulacorhamphus) . +Ramphasto8 discolorus +Ramphastos ritellinus -\-RamphasiGs piscivorous (R. carinatus) . -\-Pteroglossus aracari (P. wiedi) +Selenidera maculirostris Picidae — Picumnus squamulatus 97216. Forbes 1882. 186 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 3.— AVES (Continued} Passeriformes Formicariidae -\-Myrmotherula schisticolor ..................................... 97214. Tyraniidae +M ecocerculus leucophrys ...................................... 97220. Pittidae -Pitta oatesi Philepittidae -PhilepHta ............................................ Forbes 1880b. Hirundinidae -\-Psalidoprocne fuliginosa .................................. .... 97260. Campephagidae +Pericrocotu8 flammeus ....................................... 97029 . Dicruridae ^Chaptea aenea .............................................. 97032. Oriolidae +Oriolus chinensis ............................................ 97033. Corvidae +Chripsirhina temia ........................................... 97042. +Corvus brachyrhynchos ................................. Gorham 1936. +Pyrrhocorax pyrrhocorax ............................... Hunter 1861. Paridae — Aegithaliscus annamensis ..................................... 97142. Timeliidae +Mesia argentauris ........................................... 97109. — Alcippe nipalensis ......................................... < — Mixornis rubricapilla ...................................... < Q^QOI' +Garrulax leucolophus ......................................... 97048. -\-Garndax vassali ............................................. 97045. +Stachyris nigriceps. . . . < Q-AKO' y < uoo. rycnonotidae +Criniger gularis ............................................. 97129. -\-Oiocompsa flanventris ...................................... / 97118. m j-j 97144. Turdidae ; 97149 +Luscinia calliope .......................................... i Q"! 59 Sylviidae { 97153. +Orlhotomus sutorius .......................................... 97186. Muscicapidae — Siphia parva ................................................ 97166. +M uscicapula rubeculoides ..................................... 97172. +Culicicapa ceylonensis ....................................... 97163. Motacillidae +Anthus hodgsoni ............................................ 97186. Bombycillidae - Bombycilla garrula ...................................... Kuhl 1820. Laniidae +Laniarius atroflarus ......................................... 97257. +Lanius excubitor ...................................... Hunter 1861. Nectariniidae — Cinnyris reichenom ......................... . < Q»-OOO' ^ y i -o-. — Cinnyris jugularis ........................................... 97189. — Aethopyga saturata ........... . .97194. 1938 GALL BLADDER— GORHAM AND IVY 187 TABLE 3. — AVES (Continued) Passeriformes (continued) Dicaeidae +Dicaeum concolor ............................................ 97201. Zosteropidae +Zosterop8 palpebrosa .......................... -\-Zosterops virens ............................................. 97284. Ploceidae +Estrilda melpoda ............................................ 97254. +Munia striata ............................................... 97205. Icteridae +Zarhynchus wagleri ......................................... 97229. Thraupidae +Calospiza chrysophrys guttala ................................. 97217. +Thraupis cyanocephala subcinerea ............................ 97218. Fringillidae +Allapele$ semirufus .......................................... 9721 1. TABLES 4-16.— MAMMALIA TABLE 4 Monotremata Ornithorhynchidae f Owen 1838. +Ornithorhynchus anatinus ............................. \ Crisp 1862. [ Flower 1872. +Ornithorhynchus analinus ("Platypus") ............... Mackenzie 1918. Tachyglossidae f + Tachyglossus aculeatus (Echidna hystrix) ............. i Flower 1879 [ Chapman 1887. + Tachyglossus aculeatus (Echidna sp.) .................. Mackenzie 1918. TABLE 5 Marsupialia Didelphiidae [ Mackenzie 1918. -4-Didelphis marsupialis .......................... \ Hunter 1861. 1 Flower 1872. +Didelphis paraguayensis (D. azarae) ................... { taws™ 834^*' +Didelphis virginianus .................................. Gorham 1936. +Monodelphis brevicaudatus (Didelphis hunteri) ............. Hunter 1861. Dasyuridae +Dasyurus quoll (D. tiverrinus) ....................... Mackenzie 1918. ,. , . /Alston 1880. +Antechinomys lamger .............................. | Beddard 1908b. + Thylacinu9 sp ........................................... Crisp 1862. +Phascogale tapoatafa (P. penciUata) ...................... Hunter 1861. -\-Sarcophilus harrisi ................................. Mackenzie 1918. Notoryctidae +Notorycies lyphlops ................................... Carlsson 1904. Paramelidae +Chaeropu» castanotis ................................... Parsons 1903. Phalangeridae +Phalanger maculatus (Cuscus) ........................... Forbes 1881. + Tricho8urus mlpecula (Phalangista mtlpina) ...... | j^ckenzieVgiS. +Pseudochirus sp .................................... Mackenzie 1918. -\-Schoinobale8 volans (Petaurus taguanoides) ................ Hunter 1861 188 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 5— Continued Marsupialia (continued) Phascolarctidae [ Forbes 1881. -f Phascolarctos cinereus \ Sonntag 1921. Wombatidae I Mackenzie 1918. + Wombatus ursinus (Phascolomys tvombat) Forbes 1881. + Wombatus ursinus ("Wombat") Cleland 1869. + Wombatus ursinus (Phascolomys sp.) Mackenzie 1918. Macropodidae / jjunter ^ggj -\-Macropus cangaru (M. major) < /-i • _ , uco ^ v_ nsp ioO— . +Macropus cangaru (M. giganteus) Flower 1872. +Macropus parryi \ Mackenzie 1918 +M acropus walabatus J •i-Megaleia rufa (Macropus ruber) Crisp 1862. +Dendrolagus inustus | Owvn 1852. -\-Dorcopsis luctuosa (Halmaturus luctuosus) Grarrpd 1875. +Dorcopsis luctuosa (Halmaturus xanthopis) Crisp 1862. +Polorous tridactylus (Hypsiprymnus murinus) Hunter 1861. Caenolestidae +Caenolestes obscurus Osgood 1921. Myrmecobiidae +Myrmecobius fasciata . . Mackenzie 1936. TABLE 6 Insectivora Tenrecidae +Oryzorictes hova 1 nnh<,on 1 009 +Centetes caudatus / Dobson 1882- Potomogalidae *+Polomogale velox Seabra 1901. Solenodontidae +Solenodon paradoxus Allen 1910. +Atopogale cubanus (Solenodon) 1 Chrysochloridae +Chrysochloris asiatica (C. aurea) [• Dobson 1882. +Chrysospalax dobsoni (Chrysochloris villosa) j +Amblysomus hottentotus (A. rutilans) j +Amblysomus obtusirostris (Chrysochloris) Peters 1852. Erinaceidae i Dobson 1881. +Erinaceus europaeus { piQ^L^J i^o ' { Hill 1926.'" +Echinosorex albus (Gymnura rafflesi) Allen 1910. -+Paraechinus deserti (Erinaceus algirus) ) +Paraechinus micropus (Erinaceus pictus) -i-Paraechinus amir (Erinaceus macracanthus) +Paraechinus blanfordi (Erinaceus jcrdoni) Dobson 1882. -+Paraechinus niger . + Hemiechinus grayi (Erinaceus grayi) -rAtelerix albiventris (Erinaceus) •±Aethechiiius angolae (Erinaceus diadematus) j + Ericulu$ Mfairi (Echinops) Martin 1838. Soricidae •i Sore x pulgaris Arnback 1907. +Sorex araneus Hill 1926. +Crocidura hirla Peters 1852. + Blarina brevicauda 16545. •Called "Vesicula vermiculair." 1938 GALL BLADDER— GORHAM AND IVY 189 TABLE 6 — Continued Insectivora (continued) Talpidae • -\-Desmana moschata (Myogale) ......................... \ nnhann -f Scapanus latimanus (Scapanus townsendi) .............. JL Tupaiidae + Tupaia belangeri ...................................... Garrod 1879. +Ptilocercus lowii ........................................ Clark 1926. +Dendrogale frenata ........................................... 46628. Macroscelididae +Rhynchocyon cirnei ................................... ] -\-\asilio brachyrhynchus (Macroscelides fuscus) ............ [ Peters 1852. +Petrodromus tetradactylus .............................. J + Macroscelides rozeti ................... Duvernoy and Lereboullet 1840. TABLE 7 Dermoptera Galeopithecidae +Galeopterus temmincki (Galeopithecus volans) ............ Chapman 1902. TABLE 8 Chiroptera Pteropidae +Pteropus rufus rufus (P. edivardsi) ....................... Flower 1872. +Pteropus giganteus giganteus (P. medius) ................ \ „ _K. , cc, +Pleropus subniger (P. rubicoUis) ........................ / KoDm l( 81* +Pteropus edwardsi (P. edwardsii) ......................... Peters 1852. Cynopteridae *+Cynopterus brachyotis (C. scherzeri) *-\-Penthetor jagori (Cynopterus) . *-\-Epomops franqueti (Epomorphus comptus) Robin 1881. *+Hypsignathus monstrosus. *+Nyctymene cephalotes (Eonycteris) *+Roussettu$ amplexicaudaius (Cynonycteris) Robin 1881. +Rou$settus collaris (Cynonycteris) \ PQ, _, , flr-9 +Epomophorus crypturus / * Rhinopomidae +Rhinopoma microphyllum. Emballonuridae * -\-Emballonur a nigrescens. Robin 1881. *-(-/?/! ynchiscus naso (Rhynchonycteris) ................... * -\-Balantiopteryx plicata (Saccopleryx) ................... *+ Taphozus melanopogon .............................. +Coleura afra (Emballonura) .............................. Peters 1852. Xoctilionidae +Noctilio leporinus ...................................... Robin 1881. +Noclilio albiventer ...................................... Cuvier 1835. Nycteridae x Robin 1881. •+ A yctens reiout ....................................... / +Xycteris hispida (N. villosa) ............................. Peters 1852. * The abeence of the gall bladder was not noted. 190 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 8 — Continued Chiroptera (continued) Megadermidae *+Megaderma spasma 1 Rhinolophidae ' '+Rhinolophus ferrum-equinum. Robin 1881. +Rhinolophus hipposideros . *+Rhinolophus euryale +Rhinolophus lobatus Peters 1852. Hipposideridae + Hipposideros diadema (Phyllorhina) \ P..L; , aai *+Hipp08ideros commersoni (Phyllorhina) / * -\-Hipposideros commersoni (Phyllorrhina vittata) Peters 1852. * -\-Hipposider os armiger (Phyllorhina) , Phyllostomidae *+Phyllostomus hastatus *+Macrotus waterhousii. Robin 1881. *+Glossophaga soricina. *+Carollia perspicillata (C. brevicauda). . . *+Artibeus jamaicensis (A. perspicillatus) . + Vampyrus spectrum Cuvier 1835. Desmodontidae + Desmodus rotundus (D. rufus) Flower 1872. Vestertiliolidae -\-Vespertilio murinus. +Myolis myotis (Vespertilio) Kuhl 1820. -\-Myotis mystocinus (Vespertilio) } *+Scotophilus temminckii > Robin 1881. *+Pipistrellus kuhlii (Vesperugo) J +Pipistrellus pipistrellus (Vespertilio) Cuvier 1835. ( Flower 1872. +Nyctalus noctula (Vesperugo) \ Hill 1926. ( Cuvier 1835. *-}-Lasiurus borealis (Atalapha noveboracensis) j *+Barbastella barbastella (Synotus) [ Robin 1881. *+Plecotus auritus J +Plecotus auritus 1 P • iaq- +Eptesicus serotinus ("Vespertillion") J +Eptesicus serotinus (Vespertilio) Kuhl 1820. +Scotophilus nigrita (Nycticejus planirostris) Peters 1852. *-\-M iniopterus schreibersii ) *+Kerivoula hardunckii Molossidae *+Af olossus obscurus *+Chaerephon plicatus (Nyctinomus) +Cheiromeles torquatus *+Tadarida brasiliensis (Nyctinomus) *+M ormopterus acetabulosus (Nyctinomus) +Nyctinomous limbatus (Dysopes) Peters 1852. +Nyctinomus taeniotus (Dinops cestoni) Cuvier 1835. TABLE 9 Primates Lemuridae -\-Lemur mayottensis . . Beddard 1884a. > Robin 1881. +Lemur macaco Martin 183h * The aboence of the gall bladder was not noted. 1938 GALL BLADDER— GORHAM AND IVY 191 TABLE 9 — Continued Primates (continued) Lemuridae (continued) +Lemurcatta .......................................... Hunter 1861. +Lemur fulvus .......................................... Fl°^er }g?- +Lemurfuhus (L. albifrons) ............................. Hunter 1861. . , / Flower 1872. + Lemur vanegatus (L. vanus) ..... j Grandidier and Milne-Edwards 1875. +Microcebus murinus .................................... Flower 1872. +Microcebus murinus (M. smithi) ........ ... ...... . . . . . ... .Huge iww. +Propithecus diadema .............. Grandidier and Milne-Edwards 1875. +Hapalemur griseus .................................. Bedda rd 1884a. +Hapalemur simus .................................... Beddard 1901. Indrisidae +Indris indris (Avahi laniger) ---- Grandidier and Milne-Edwards 1875. Daubentoniidae -\-Daubentonia madagascarensis (Chiromys) ..... Mjvart and Murie 1865. +Daubentonia madagascarensis ("aye-aye") ................. Owen 1866. Lorisidae +Loris tardigradus (Nyclicebus) ......................... • . - Ruge 1902. / Hunter 1861. +Loris tardigradus ................................... | Flower 1872. +Loris tardigradus (Nycticebus) ................. Mivart and Murie 1865. +Loris tardigradus (Stenops gracilis) ........................ Kuhl 1820. -\-Nycticebus javanicus ................................. j Flower 1872. +Arctocebus calabarensis ............................... J +Perodicticus potto .................................... | Flower 1872. -\-Galago madagascarensis .................................. Kuhl 1820. + Galago crassicaudatus .................................. Flower 1852. +Galago mohali .......................................... Smith 1849. v / Beddard 1901. +Myoxicebus griseus (Hapalemur) . j Grandidier and Milne-Edwards 1875. Tarsiidae ( Ruge 1902 + Tarsias fuscus (T. spectrum) ........................ \ Sonntag 1924. [ Woollard Callitrichidae +0edipomidas oedipus (Midas) .......................... ('^°W^T 186l' +Callithrix jacchus (Hapale) ........................... Beattie 1927. ( r lower loTZ. r .., f Sonntag 1924. Cebldae Crisp 1862. +Cebus capucinus .................................... \ Ruge I902a. I Flower 1872. +Cebus apella .......................................... Hunter 1861. +Ateks geoffroyi ........................................ Flower 1872. +Ateles cucullatus ....................................... Murie li 65. +Ateles ater ............................................. Ru?e 1902a- +Ateles belzebuth ......................................... Kuhl l%®' +Pithecia pithecia ...................................... Sonntag 1924. +Pitkecia monacha ...................................... 1?°^!! JOQA +Cacajao rubicundus (Brachyurus) ........................ Forbes 1880. +Alouatta senicula ...................................... Flower 1872. +Saimiri sciurea (Callithrix sciureus) ...................... Martin 1833. 192 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 9 — Continued Primates (continued) Cercopithecidae , ... 109Q -\-Cercopithecus albogularis . . . . { Owenl839a •i-Cercopithecus erythrogaster Murie 1866. i-Cercopithecus cynosurus i-Cercopithecus talapoin i-Cercopithecus cephna (C. cephus) i-Cercopithecus nictitans J •i-Cercopithecus callithricus (C. sabaena) Hunter 1861. •i-Cercopithecus petaurista Ruge 1906. -f Cercopithecus sinicus } T/V.UI i oon •i-Cercopithecus aethiops / -\-Cercocebus fuliginosus Bradley 1903. •i-Erythrocebus patas (Cercopithecus) Ruge 1906. -\-Colobus vellerosus Flower 1872. •i-Theropithecus rueppelli (Gelada) Garrod 1879a. -f Papio lestes Mentzer 1929. +Papio sphinx Ruge 1906. + Papio maimon. -\-Papio anubis . . +Papio porcarius. Ruge 1906. Hunter 1861. Ruge 1906. Hunter 1861. Flower 1872. Hunter 1861. -\-Papio leucophaeus (Mandrillus) ........................ Sonntag 1922. -\-Papio hamadryas .................................... Schrieber 1932. -f- Macaco maura ......................................... Murie 1872. -\-Macaca brunnea ..................................... Anderson 1872. -f Macaco albibarbala (M. silenus) . . . \ „„ „ -,QC1 +Macaca mulatta (M. rhesus) .......................... / * +M acaca mulatta (M. rhesus) ........................... Linback 1933. +M acaca cynomolga ..................................... } + M acaca sinica ........................................ }• Ruge 1906. +Macaca nemestrina .................................... J Pongidae f Ruge 1906a. +Hylobates lar ....................................... \ Hunter 1861. [ Flower 1872. +Hylobates lar (H. leuciscus) ........................... Chapman 1900. Owen 1830. +Pongo pygmaeus ("orang") ......................... \ Sonntag 1924a. Ruge 1906a. +Pan satyrus ("chimpanzee") .......................... +Pan satyrus (Anthropopithecus troglodytes) ............... Sonntag 1923. f Flower 1872. +Gorilla gorilla ....................................... \ Fox 1930. ( Ruge 1906a. Hominidae +Homo sapiens ............................................ A.U-C. TABLE 10 Carnivora Canidae +Cants familiaris ...................................... Gorham 1936. -(-Cants aureus ........................................ Mentzer 1929. +Canis lupus ....................................... Macalister 1867. +Speothos venaticus (Icticyon) ............................ Flower 1880. +Lycaon pictus .............. . / Crisp 1855 1 Garrod 18<8a. 1938 GALL BLADDER— GORHAM AND IVY 193 TABLE 10 — Continued Carnivora (continued) Canidae (continued) +Otocyon virgatus (Cam's otocyon virgatus) ................. Mentzer 1929. + Urocyon cinereoargenleus (Canis) ........................ Hunter 1861. -j-Cuon dukhunensis ..................................... Murie 1872c. +Fennecus zerda (Canis) ................................. Hunter 1861. +Nyctereutes procyonoides ............................... Garrod 1878a. Procyonidae / Crisp 1862. +Procyon lotor . ( Macalister 1867. +Polos flatus (P. caudivolvulus) .. . { {£2^}^ +Nasua rufa or narica .................................. Hunter 1861. +Bassaricyon gabbi (B. alleni) .......................... Beddard 1900a. -f Ailurus fulgens ....................................... Flower 1870. +Aeluropoda melanoleuca ..................................... 47432. Ursid*' / Hunter 1861. + Unusardos.. \Macalisterl867. -1- Ursus maritimus ........................................ Crisp 1862. + Ursus malayanus ........................................ Rex 1888. +Melursus ursinus (Ursus labiatus) ....................... Flower 1872. Mustelidae +Mustfla putorius (Putorius furo) ................... \ Mapai:,t_P 18fr +Mustela erminea (Putorius) ........................ / Macallster l »b , . +Mustela vison ........................................ Gorham 1936. +M ellivora sagulata ................................... 1 + Taxidea taxus (Meles labradoria) ...................... ( w „ n OC1 +Meles meles (M. taxus) .............................. +Marles martes (Mustela) ............................. J + Maries pennant i (Mustela canadensis) ................... Martin 1833b. +Mydaus marchei (M. meliceps) ........................ \ TT 1 fiP1 +Grison vittata (Mustela grison vittaius) .................. fa + Tayra barbara (Galera) .................................. Crisp 1862. +Helictis subaurantiaca ................................. Garrod 1879c. -\-Heliclis personata ................................... Beddard 1905a. +Lulra lulra (L. vulgaris) .............................. { cri^fse!61" +Gulo sp. (Ursus gulo) .................................... Crisp 1862. Viverridae + Viverra zibetha ........................................ Hunter 1861 . -f Viverra civetta ........................................ Mivart 1882. +Suricata suricata (S. telradactyla) ..................... / ' +Paguma larvata (Paradoxurus) .......................... Mivart 1882. -f Nandinia binotaia .................................... Carlsson 1900. — Nandinia binotata ..................................... Mivart 1882. +Crossarchus obscurus ................................... Martin 1834. +Arctictis binturong .................................... Garrod 1873a. +Hemigalus derbyanus .................................. Mivart 1882. +Galidea elegans ..................................... Beddard 1909a. "1™"1882- +Cryptoprocta ferox .................................... Beddard 1895. Hyaenidae +Hyaena brunnea ........................................ Murie 1867. +Hyaena vulgaris ....................................... Hunter 1861. +Crocuta crocuta (Hyaena) ................... Watson and Young 1879a. +Proteles cristata ........................................ Flower 1869. 194 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 10 — Continued Carnivora (continued) Felidae +Felis domestica ........................................ Mivart 1881. + Felis pardcdis ......................................... I /-_•__ i ofi.? + Fdis nebulosa (F. macrocelus) ........................... f L +Panthtra leo (Felis lea) . . { ggjj/f &. +Panthera pardus (Felis leopardus) ....................... Hunter 1861. -\-Panthera pardus (Felis pardus) ......................... Mentzer 1929. +Panthera onca (Felis) .................................. Martin 1832. -\-Lynx caracal (Felis) ................................... Hunter 1861. -i-Acinonyx jubala ..................................... Mentzer 1929. 4-Felis concolor ........................................ Martin 1833a. Otariidae -\-Eumetopias jubata (Otaria) .............................. Murie 1868. -\-Arclocephalus hookeri (Otaria) .......................... Murie 1867a. Odobaenidae +Odobaenus rosmarus (Trichechus) ........................ Murie 1870a. Phocid^t Hunter 1861. +Phoca ntuhna. . f Macalister 1867. +Phoca groenlandica ............................... { Crisp 1862. 1 Murie 1870b. TABLE 11 Xenarthra Bradypodidae , Hunter 186L — Bradypus tndactylus. . .......................... s Buriet 191 1. — Bradypus cuculliger (B. cuculli) .......................... Burlet 1911. / Wislocki 1928. -Bradypus gnseus .................................. ( Sonntag 1921a. Choloepodidae f Hunter 1861. +Choloepus didactylus (Bradypus) .................... j £aPP 18o2- 1 Burlet 1911. [ Sonntag 1921a. +Choloepus hoffmani ................................... Wislocki 1928. Myrmecophagidae -\-Myrmecophaga jubata ................................... Owen 1854. f Burlet 1911. -{-Cyclopes didactylus (Cyclothurus) ..................... I Flower 1872. ( Wislocki 1928. + Tamandua sp ........................................ Wislocki 1928. + Tamandua tetradactyla ................................ Beddard 1909. Dasypodidae ( Hunter 1861. +Dasypus novemcinctus ................................ ] Crisp 1862. ( Rapp 1852. -\-Dasypus novemcinctus (D. peba) ......................... Owen 1831a. -\-Euphractu8 sexcinctus (Dasypus) ....................... | 9-^n 186^° + Tolypeutes tricinctus ................................... Garrod 1878. + Tolypentes sp ......................................... Murie 1872a. +Chlamydophorus truncatus ........................... Macalister 1873. Pholidota Manidae +Manispentadaclyla.. . { Hunter 1861. -\-Smutsia temminckii (Manis) ............................ Peters 1852. -\-Manis javanica ........................................ Adams 1859. 1938 GALL BLADDER — GORHAM AND IVY 195 TABLE 11 — Continued Tubulident«va Orycteropidae i jaeger 1837. +0rycteropus afer (O. capensis) . . { gonntag 1925. +Orycteropus afer (O. afro) F°* 1930. TABLE 12 Artiodactyla Tayassuidae , Hunter 1861. -Pecan torquatus (Dicotyles) . . . | Macalister 1867. Suidae +Babirussa alfurus (B. babirus) Flower 1872. +Phacochoerus africanus Mentzer 19-29. -\-Phacochoerus aethiopicus (P. pallasii) Owen 1851. Hippopotamiidae [ Garrod 1879b. -{-Hippopotamus amphibius \ Weissberg 1932. [ Chapman 1881. -Hippopotamus amphibius • Chapman 1881. Camelidae -Corned bactrianus Flower 1872. -Lama guanicoe (L. pacos) ... -I crjsp igg2. — Lama vicugna (Auchenia sp.) J Tragulidae + Tragulus kanchil Hunter 1861. + Tragulus javanicus (T. napu) Macalister 867. -\-Dorcatherium aquaticum (Hyomoschus aquaticus) Flower 1867. Cervidae +Cervus axis Crisp 1862. / Hunter 1861. -Cervus axis \ Raven 1936. — Cervus unicolor unicolor (C. hippelaphus) . \ crjsp 1862. — Cervus unicolor moluccensis (C. moluccensis) / — Cervus unicolor sunnhoii (C. swinhpii) — Cervus unicolor marianus (C. mariannus) — Cervus elaphus — Cervus kukulii . Garrod 1877. — Cervus timoriensis (C. molucensis) — Cervus duvauceli — Cervus alfredi - Cervus porcinus -\-Mazama superciliaris (Cervus) -Mazama rufus (Cervus) .. , J Crisp . -Odocoileus hemionus (Cervus auntus) | Gorham 1936. — Odocoileus mexicanus (Cervus) • \ crisp 1862. - Alces alces (Cervus) • / -Alces alces (A. machlis) Watson and Young 18,9. — Muntiacus muntjac (Cervulus) • \ Garrod 1877. — Af untiacus reevesi (Cervulus reevesi) / - Hydropotes inermes Garrod 187 < a. -Pudu pudu (Cervus) Garrod 18«. -Pudu pudu Flower 1875. — Elaphodus cephalophus J — Elaphodus cephalophus (Lophotragus michianus) }• Garrod 187 /. — Blastoceros bezoarticus (Cervus campestris) J ' Garrod 1877. +Moschus moschiferus ] Crisp 1862- 196 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 12 — Continued Artiodactyla (continued) Giraffidae j Owen 1838b 1+Giraffa camelopardalis < Garden 1787. Owen 1838b. Crisp 1862. Neuville 1914. — Giraffa camelopardalis . Murie 1872c. Chapman 1875a. Joly and Lavocet 1845. — Giraffa camelopardalis (Camelopardalis giraffa) . . < pox ^929. -Giraffa reticulata Lonnberg 1912. Antilocapridae , Murie 1870b +Antilocapra amencana . . ' Lonnberg 1909. Bovidae Bovinae +Bo8taurus. -\Hunterl861. +Bison bison (B. amencanus) J +Syncerus coffer (Buffdus c. radcliffei) Mentzer 1929. Caprinae +Ovis musimon Crisp 1 862. -f Copra hircus (C. picta) Garrod 1877. *— Copra angoriensis (3 specimens) . . Crisp 1862. +Hemitragus jemlahicus (Copra jemlaica) Garrod 1877. Rupicaprinae + Rupicapra rupicapra Rex 1888. -i-Budorcas taxicolor Lander 1919. Ovibovinae +Oribos moschatus Lonnberg 1900. Bubalinae — Bubalis buselaphus (Antelope) \ cr}sp jggo — Damaliscus pygargus (Antelope) j -4-Damaliscus pygargus (Damalis) Garrod 1877. +Connochaetes albojubatus Mentzer 1929. +Connochaetes gnu (Catoblcpas) Garrod 1877. Cephalophinae , ^ • 1QC., /- T i it- / A i i \ i Crisp 18bJ. — Lephalophus maxima (Antelope) . . . . < QaT^O(i 1377 — Cephalophus monticola (C. pygmaeus) Garrod 1877. — Cephalophus harveyi keniae Lonnberg 1900. — Cephalophus melanorrhaeus Lonnberg 1909. -Cephalophus natalensis. •••.-,•,• \ L5nnberg 1900. — Cephalophus abyssinicus hinder j — Cephalophus grimmia (Antelope mergens) Crisp 1862. ,, , , , .,, . ' Lonnberg 1900. -( ephalophus ogilbyi | Lonnberg 1909. Oreotraginae +Ourebia nigricaudatus (Nanotragus) Garrod 1877. +Raphicerus campestris (R. neumanni) < jJp^^r^i 909 +Neotragus pygmaeus Pocock 1918. +Rhyncotragus cavendishi (R. kirkii) Mentzer 1929. t Six males, three females, others unsexed. •Communication with breeders of Angora goats, in this country and abroad, failed to reveal any breed in which the gall bladder was absent. The number of specimens indicates that Dr. Crisp prob- ably was guilty of mis-statement, rather than dissection of anomalous specimens. 1938 GALL BLADDER— GORHAM AND IVY 197 TABLE 12 — Continued Artiodactyla (continued) Bovidae (continued) Reduncinae /-,;«,« i««9 +Redunca arundinum (Antelope tsabelhana) ............... ... crisp l»b£. +Kobus defassa ....................................... Mentzer 1929. •4-Kobus ellypsiprymnus ................................... Smith 1! STsaT0a tartarica ........................................ Murie 1870c. Antelopinae ,, . , Q<,9 +Antelope cervicapra (A. bezoartica) ...................... • • • v"8^ *$?*• J Crisp lobJ. -\-Gazella dorcas (Antelope) ............................. ^ Garrod 1877. +Ga»Ua bennetti (Antelope) .. ... -CrP 1862. +Gazella granh ...................................... \ Garrod 1877. +Gazella subgutturosa .................................. 1 +Gazella muscatensis ..... . . . . 1 Garro(j 1877. -f Gazella rufifrons ..................................... +Gazella arabica ...................................... J 0 +Antidorca marsupialis (Antelope euchore) ............... ... .crisp I8t>-. +Litfiocranius walleri ................................. Lonnberg 1900. Oryginae r-««, +Oryx leucoryx (Antelope) .............................. • • • V™ J Crisp +Addax naso-maculatus ............................... \ Garrod 1877. Tragelaphinae r crisp 1862. + Tragelaphus scriptus (Antelope) ....................... | Garrod 1877. + Strepsicerus strepsicerus (S. Kudu) ...................... Garrod 1877. + Taurotragus oryx ..................................... Mentzer 1929. + Taurotragus oryx (Antelope oreas) ....................... • • cusp i»w. + Taurotragus oryx (Oreas caana) .......................... Crarroa 18 / /. +Boselaphus tragocamelus (Antelope picta) ................. ^untej j^i' +Boselaphus tragocamelus (Portax picta) .................. XT "n 1007' +Limnotragus gratus (Tragelaphus gratus) ................. Neuville l»y/. + Tetracerus quadricornis (T. subquadricornatus) ............. Garrod 18 / /. TABLE 13 Sirenia + Trichecus manatus (Manatee americanus) ................ - Murie 1870. + Trichecus manatus (Manatus inunguis) ................. Beddard 189 1. -+Trichecus manatus ("Manatee") ...................... Chapman 1875. Dugongidae +Dugong australis (Halicore sp.) .......................... ^>wen 1] Stellerl749. Hyracoidea Procariidae +Procaria capensis Mentzer* 1929 Raven 1936. Crisp 1862. -Procavia capensis i f}°^er* .. Q.0 I Huxley 1872. t Martin 1835. •Mentzer examined two specimens, one with a gall bladder fined with liver flukes. He notes that the gall bladder, with a central and two lateral sacs, as reported by Owen and Macalister, ma_y be found only as a dilation or a lateral pouch of the common duct, at the point where the four hepatic ducts join it. ** Flower says that this animal has no gall bladder, but in some specimens the common duct dilated to great size. 198 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. XXII TABLE 13— Continued Proboscidea Elephantidae f Hunter 1861. — Elephas maximus (E. indicus) \ Crisp 1862. 1 Forbes 1879. — Loxodonta africana (Elephas africanus) Forbes 1879. / Eales 1929. - Elephant <[ Chapman 1875b. TABLE 14 Perissodactyla Tapiridae — Tapinis terrestris (T. americanus) Crisp 1862. - Tapirus indicus Parker 1882. - Tapiru* indicus Murie 1871b. Rhinocerotidae D, . . . f Beddard 1887. - Rhinoceros umcorms < Qwen 1350 — Rhinoceros sumatrensis (Ceratorhinus) Garrod 1873. Equidae — Equus caballus 1 — Equus asinus [ Hunter 1861. — Equus zebra j -Equus burchelli 44391. — Equus quagga Mentzer 1929. TABLE 15 Cetacea Mysticeti Balaenidae — Balaena mysticetus Hunter 1840. Balaenopteridae , „ *. -\oa-i - Balaenoptera acutirosfrata (B. rostrata) { Eschricht 1849 -Megaptera longimana (M. boops) { gajg.^ ^4a9Calister 1868! Odontoceti Platanistidae — Platanisla gangetica Anderson 1879. Physeteridae — Physeter catodon (P. macrocephalus) Hunter 1840. Ziphiidae — Ziphiorrhynchus cryptodon(l] Burmeister 1866. — Hyperoodon ampullatus (Delphinus bidens) Hunter 1861. Delphinapteridae — Delphinaplerus leucas (Belugia catodon). . . .Hepburn and Waterson 1901. - Monodon monoceros . . { Hjjjjj }^0. Delphinidae — Neophocaena phocaenoides (Neomeris) Chi Ping 1926. — Phocaena phocaena (Delphinus) Hunter 1840. -Phocaena phocaena (P. communis) { ggJS^Ji WaterSon 1901. — Orca gladiator (Delphinus orca) Turner 1899. — Orcella brevirostris (Orcella) Anderson 1879. ( Gulliver 1853. — Globiocephalus melaena (G. melon) <^ M™91"! R^ [ Weissberg 1932a. 1938 GALL BLADDER — GORHAM AND IVY 199 TABLE 15-C«•• — Tursiops truncatus (Delphinus tursio) .................... Hunter 1861. +Gramphidelphis risii (Globiocephalus risii) ............... Williams 1838. — Gramphidelphis risii (Grampus rissoanus) ................ Murie 1871b. Rodentia TABLE 16 Aplodontiidae - Aplodonlia rufa .................................. Hall, MVZ 22623. Sciuridae ( Crisp 1862. +Sciurus vulgaris ..................................... { Rex 1888. I Cuvier 1835. — Sciurus carolinensis ..................................... Crisp 1862. +Sciurus carolinensis (S. cinereus) ........................ Hunter 1861. — Sciurus indicus (S. maximus) r + Tamias striatus ........................................... < 1544° -\-Marmota marmota (Arctomys) .......................... .Hunter 1861. +Marmota marmota (Arctomys alpinus) ............... | MTcalister'lSe?. + Marmota monax ("Marmottes de Canada") ............... Cuvier 1835. +Citellus tridecemlineatus ................................ Higgins 1928. +Citellus suslicka ("Spermophile souslick") ................ Cuvier 1835. •\-Glaucomys volans (Pteromys volucella) ................... Yarrell 1831a. +Glaucomys volans (Sciuropterus volucella) ................. Hunter 1861. +Glaucomys volans (Sciurux volucella) ....................... Crisp 1862. +Glaucomys volans ("Hassapan") ....................... } r