Allometric Relationships in Lions vs. Domestic Cats Author(s): D. Dwight Davis Source: Evolution, Vol. 16, No. 4 (Dec.

, 1962), pp. 505-514 Published by: Society for the Study of Evolution Stable URL: http://www.jstor.org/stable/2406182 Accessed: 25/06/2010 06:46
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ALLOMETRIC RELATIONSHIPS IN LIONS VS. DOMESTIC CATS

D. DWIGHT DAVIS Chicago Natural History Museum, Chicago, Illinois Received April 6, 1962

Gene-controlled allometric growth rates are capable of producing profound and complex evolutionary changes in the proportions of organisms. Their role in vertebrate evolution has recently been emphasized by Rensch (1948, 1960). Allometric rates may reflect the results of natural selection in two ways: (1) selection for increased (or decreased) organism size may result in exponential changes in the relative sizes of structures, and such changes may or may not be adaptive, and (2) natural selection may operate directly on the allometric rates themselves to produce changes that are by definition adaptive. Finally, (3) many organs and tissues have the capacity of responding directly to the physiological needs of the organism, and vary allometricallywithout any change in the genetic background, although the capacity for such physiological response is of course inherited. Rensch (1960) has summarized pertinent data from the literature, and Frick (1957b) has critically discussed procedural and mathematical methods. The major problem in evaluating these data is eliminating other variables. The most important of these other variables are differences in genetic background, differences in physiological requirements that are unrelated to size, and differences in the procedures used by the several investigators. Others are sex, age, and physical condition. Inadequate size differences between organisms being compared may be a limiting factor unless samples are very large. The ideal situation would be comparisons between closely related organisms with similar habits and behavior, but differing substantially in size, for then minor individual variations and differences in technique are minimized. This ideal at least approached in the Felidae, a compact
EVOLUTION

group of very similar mammals among which the largest species are more than 60 times as heavy as the smallest. Studies of allometric weight relationships between species have been limited almost entirely to a few compact organs: brain, heart, liver, kidneys, etc. No attempt has been made to evaluate more diffuse structures, such as the wet skeleton or the musculature, perhaps because preparing these for weighing presents formidable practical problems. Yet the locomotor system may account for nearly two-thirds of total body weight in a mammal, and is no less subject to allometric growth rates than are the organs of metabolism. The following data are from two captive-bred lions that were destroyed for reasons other than ill health. Both were received within a few hours after death, and all weights and measurements of soft parts (except brain and spinal musculature) were made within seven hours after the animal was received. The male was destroyed at the Chicago Zoological Park on Feb. 2, 1962. He was 3-4 years old, and was below normal size. The skeleton showed that he was somewhat rachitic, but otherwise he seemed to be in good condition although there was very little body fat. The skull of this individual is preserved in the Chicago Natural History Museum under the catalog number 0-1526. The female was received from the Lincoln Park Zoo on May 28, 1946. She was mature, but not aged, and in excellent physical condition, with considerable body fat. Her skeleton is preserved under the catalog number 54639. Both of these specimens were below the average size of wild lions. The gross weight of the male was 109.3 kg, of the female 138.8 kg. The average and ex-

16: 505-514.

December, 1962

505

506

D. DWIGHT DAVIS

treme weights given by Meinertzhagen two sample groups (skin, musculature, fe(1938) for 14 wild-killed male lions are male reproductive organs), was computed 175.9 (149.5-191.4) kg, for five females from the equation
151.4 (122.3-185.9) kg.

To make comparisons possible, the methods used by Latimer in his work on domestic cats were followed as closely as was practical. The animal was first weighed (weight of fat and gut contents was later subtracted), then the skin was removed and weighed. The viscera and all easily removable fat were next removed from the carcass, and the carcass divided into head, torso, and the four extremities. The head, torso, and extremities were weighed, then the musculature was carefully removed and the remaining moist ligamentary skeleton (with the contained central nervous system) weighed. The difference between the two weights gave the weight of the musculature. The weight of the brain was subtracted from the weight of the skeleton; the spinal cord was not removed. Data are also given for most of the body parts of a male leopard destroyed at the Chicago Zoological Park on March 8,1962. This animal was about 13 years old, and was in good physical condition, with a moderate amount of body fat. His gross weight was 47.5 kg. His skeleton is preserved under the catalog number 57415. Items up to a kilogram were weighed on a simple balance sensitive to 0.1 gram. In 1946, items between one and 15 kg were weighed on a spring balance that had been checked for accuracy by means of laboratory weights. In 1962, items heavier than one kg were weighed on a triple beam balance sensitive to one gram. All weights and measurements were made either by me or under my direct supervision.

log Y1-log Y2 log Xl-log X2'

where Y is the organ weight in lions and cats, respectively, and X is body weight minus organ weight.' The figure in the table is the mean of the exponents independently computed for each of the two lions. Where there were more than two sample groups, the "somatic exponent" is the exponential constant in the equation Y = bXa. The equation was converted to logs, and the value of a computed by the method of least squares. The samples include published weights for various felid species, in addition to my data. The composition of each sample is described below under the organ or body part. The exponent a (and with much less confidence the constant K) expresses the rate at which the weight of the organ changes as body weight changes. When its value is less than 1, organ weight increases at a slower rate than body weight (negative allometry), when greater than 1, organ weight increases at a faster rate (positive allometry). When the exponent is 1, the relative weight of the organ is the same regardless of body size (isometry). Head.-The head was removed from the torso by cutting the neck muscles at their attachment to the back of the skull and disarticulating the occipito-atlantal joint. The tongue was cut just anterior to the hyoid and left in place in the head. Head weight therefore includes all head structures except the skin.
1 The German workers include Y in X (i.e., X = body weight rather than body weight minus organ weight). This makes no appreciable difference for structures representing only a small percentage of total body weight. For heavier structures, such as the skeleton or musculature, it considerably alters the values of the constants.

WEIGHTS The weights of the body and its parts are given in table 1. The percentages of total body weight given by Latimer for 52 domestic cats of each sex were assembled and tabulated for comparsion. The "somatic exponent," where there were only

ALLOMETRIC RELATIONSHIPS IN CATS

507

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508

D. DWIGHT DAVIS

Limbs.-The fore limbs were removed from the female lion and the leopard by severing the shoulder muscles where they attach to the torso. Thus fore limb weight for these individuals includes the scapulae and shoulder muscles. In the male lion, to make the figures comparable to Latimer's, the fore legs were severed at the scapulo-humeral articulation, and the weights therefore do not include the scapulas and shoulder muscles.2 In all three cases the hind legs were removed by cutting the hip muscles at their attachments to the pelvis. Musculature.-The musculature can be removed more completely from the skeleton of a large animal than from a small one. Therefore the relatively heavier musculature indicated for the lions than for the domestic cats may not be entirely real. Respiratory system.-As in Latimer's work, weight includes the trachea and larynx. The somatic exponent was calculated from the two lions, the leopard, and the mean of Latimer's 104 domestic cats. Heart.-The chambers were washed out with a stream of running water before the organ was weighed. The pericardium was left in place, and the great vessels cut close to their origin. The heart seems abnormally large in the male lion, and slightly small in the female. Schiller (1959) gives the weights of six preserved hearts of adult zoo lions. The mean and extremes of his figures are 733 (530-940) g. A total of 13 samples of body weight/ heart weight ratios in felids of various species and sizes can be compiled from the literature plus my data. Crile and Quiring (1940) give data for adult samples representing six species (11 individuals), and Frick (1957a) for one specimen of Felis caracal. A scatter diagram for all available body weight/heart weight ratios, with the calculated regression line, is shown in fig. 1. Digestive tube.-The stomach was emp2 The weight of the left fore leg was not recordedbefore the muscles were removed. The weight of the muscles of this leg was estimated from the corresponding weight of the right leg.

tied, but the intestines were not cleaned out. The mesenteries (but not the omentum) are included in the weight of the gut. The somatic exponent was calculated from my three felids plus Latimer's data. Liver.-Somatic exponent based on 11 samples. Crile and Quiringgive six figures, and Frick one. Pancreas.-Somatic exponent based on four samples: my three felids, and Latimer's data for the domestic cat. Spleen.-Somatic exponent based on 10 samples. Crile and Quiring give data for five and Frick for one. The size of the spleen is subject to great individual variation, so the somatic exponent from the available sample may not truly represent the relations in the Felidae. Adrenals.-Somatic exponent based on 11 samples. Eight are from Crile and Quiring. Kidneys.-The difference in weight between the two lions is enormous. The figure for the male seems abnormally high, that for the female abnormally low. Individual weights of the kidneys were: male lion, right 409, left 407; female lion, right 254, left 237; leopard, right 127, left 123. Somatic exponent based on 9 samples. Four are from Crile and Quiring, and one from Frick. Bladder and urethra.-Only the proximal part of the urethra was included with the bladder. The somatic exponent is based on my three felids and Latimer's data. Reproductive organs.-For the male reproductive organs I have three samples: the male lion, the leopard, and Latimer's domestic cats. The somatic exponent was calculated by the method of least squares. For the female organs I have only two samples: the female lion and Latimer's domestic cats. Brain.-Weight of the brain includes the dura. The only weight of a lion's brain I find in the literature is 219 g for an animal weighing 119.5 kg (Weber, 1888, cited from Brummelkamp). This gives a percentage of 0.18, which is slightly lower than mine.

ALLOMETRIC RELATIONSHIPS IN CATS 3.40

509

3 3.00 3.

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Felis Felis Felis Felis Felis Felis Felis Felis Felis Felis Felis Felis Felis

oinca pardus leo (3) leo (2) concolor serval tigris tigris caracal leo df leo 9 pardus domestica

X 0

~
1.00

13 3.40 3.80 I 4.20 I 4.60. I 5.00 5.40

log(body

weight-heart

weight)

FIG. 1. Scatter diagram, with fitted regression line, for body weight/heart weight relationship in Felidae. Nos. 10-12 are my felids; no. 13 is the mean of Latimer's cats. Others are published records.

Brummelkamp(1940) compiled from the literature brain weight/body weight figures for 11 felids of various sizes. These data, together with Latimer's and mine, were plotted on a scatter diagram and a regression line fitted by the method of least squares (fig. 2). The regression exponent is 0.61, indicating a strong negative allo-

metric relationship between brain weight and body weight in the Felidae.
DISCUSSION OF WEIGHTS

The total weights of our lions are 39 (male) and 57 (female) times the mean weights of Latimer's domestic cats. According to Meinertzhagen's figures, an

510 2.60 a = -. 644 b = .611 2. 20 -

D. DWIGHT DAVIS

14

o6

1.80~~~~~~~~~~~
1. 80 1 40104. 11 / 1. Felis domestica 2. Felis domestica 3. Felis pardus Felis pardus 15. Felis pardus 6. Felis leo 7. Felis concolor 8. Felis concolor Felis conlcolor ~~~~~~~~~~~~~~9. 10. Felis catus 11. Felis canadensis Felis domestica ~~~~~~~~~~~~~~~12. Felis leoe ~~~~~~~~~~~~~~~~13. 14. Felis pardus

_ 1 _ 1. 40 c ._
=

2 12 0
r

1.00

II

3.40

3.80

4.20

4.60

5.00

5.40

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weight-heart

weight)

FIG. 2. Scatter diagram, with fitted regression line, for body weight/brain weight relationship in Felidae. Basic data from Brummelkamp (1940), to which my lion (13), leopard (14), and the means of Latimer's cats (12) have been added.

average adult male lion is about 62 times as heavy as an average adult male domestic cat. Relative weights of the locomotor system in organisms differing only in size are determined largely by physical laws. Equivalent performance in a large body demands a relatively greater quantity of skeletal muscle tissue because the power of a muscle increases as the square of its diameter whereas the mass to be moved increases as the cube of its linear dimensions. Thus a positive allometric relationship is to be expected in closely related animals. Our data indicate a slight positive allometry in the musculature of the Felidae, but at best it is so slight that the performanceof a lion is certainly distinctly inferior to that of a domestic cat. (This, of course, is likewise true of the prey of the two animals, assuming equivalent size

differences.) Apparently no previous attempt has been made to determine the relationship between organism size and the mass of the skeletal musculature in mammals. The same physical laws apply to the skeleton. Our data indicate an isometric relationship between skeleton weight and body weight in the Felidae, which would likewise make the potential performanceof a lion inferior to that of a domestic cat. As pointed out by Thompson (1952), an animal may reach a size where it is unstable dynamically, though still stable statically. Elephants probably approach this condition. From weight figures for four adult African elephants given by Robertson-Bullock (1962), the skeleton averages 20% of total body weight (less proboscis, ears, and tusks), whereas the musculature averages only 39.8%. It is customarily regarded as a general

ALLOMETRIC RELATIONSHIPS IN CATS

511

rule that in homoiothermous vertebrates the relative weights of the internal organs vary inversely with total body weight, i.e., that there is a negative allometric relationship between organ weight and body weight (Rensch, 1949, 1960; Frick, 1957a). This generalization has been called "Welcker's rule." It is based on comparisons, by several different investigators (summarized by Rensch and by Frick, 1957a), between more or less closely related species of mammals and birds. A similar relationship among individuals of the same species is distinguished as the "series rule." Obviously some structures in the Felidae do not follow Welcker's rule. Rensch (1948), Frick (1957a), and others have assumed that for each organ there is a somatic exponent that is valid for all homoiothermousvertebrates, a sort of optimal physiological relation. These authors have tried to determinesuch values for various organs. This mean or "universal" value is then substituted for a in the allometric equation, and the corresponding values for the constant b calculated for groups of related species. The resulting value of b is called the "organ coefficient." It is believed to represent the constant of proportion for that organ in a genus, family, or order. The value of b, when based on a universal a, varies from group to group; in a family or genus characterized by a "large" heart it would be relatively large, in a group characterized by a "small" heart it would be relatively small. This method was taken over from the earlier work of Dubois and others on relative brain size, where it seems to have some validity. To me it seems unrealistic to assume a universal growth constant (a) for organs, such as the heart, whose size is directly related to performance. It is possible, of course, to determine a mean value as Frick and others have done, but it does not follow that the mean value has any biological significance. The potential performance of a very large active animal would be seriously impaired if the relative

size of the heart were reduced at an exponential rate. Relative growth is subject to natural selection, and we would expect the somatic exponent for many organs to be higher among active predators that periodically mobilize their reserves for bursts of energy than among relatively sedentary non-predatorsor forms that may use energy as sustained high levels. If the universal somatic exponent has no biological validity, then the "organ coefficient" derived from it has none either. The data for the organs of the Felidae may be divided into two groups: those in which the relation to body weight is constant or increases with increasing body weight, and those in which relative weight decreases as body weight increases. In the first category are the skin, skeleton, musculature, respiratory organs, heart, digestive tube, spleen, adrenals, and reproductive organs. These include the locomotor apparatus and those organs (heart, respiratory organs, adrenals) directly associated with performanceof the locomotor apparatus. In the second category are the liver, pancreas, kidneys, bladder and urethra, brain, and eyeballs. These are all organs associated with metabolism or with the central nervous system. Where comparable data are available (liver, kidneys), the somatic exponents of the metabolic organs in the Felidae agree closely with the "universal" exponents of Rensch and Frick. The relative size of brain and eyeballs decreases at a faster rate than any other organ measured. My exponent for the brain is somewhat higher than the widely accepted "universal" mammalian exponent of 0.56. The thyroid was obviously pathologically enlarged in my female lion and in several of the felids weighed by Crile and Quiring. I therefore did not try to calculate a somatic exponent for this organ. Thus in the Felidae those structures associated directly or indirectly with motor performance do not follow Welcker's rule, whereas structures associated with metab-

512
TABLE 2.

D. DWIGHT DAVIS Linear measurements, in mm, of two captive-bred lions

Male Length Per cent body length Length Female Per cent body length Mean

Body length (nose-anus) 1,448 Tail length 787 Spine length 1,005 265 Skull, condylobasal length Bizygomatic diameter 179 Tranv. diam. post. to forelimbs 214 Dorsoventral diam. post. to 350 forelimbs Circumf. post. to forelimbs 1,016 Circumf. at xiphisternal 1,054 junction Interacetabular diameter 73
R3 L3 R

54.4 69.4 18.3

1,610 730 1,021 273 224 206 400

45.3 63.4 17.0

49.9 66.4 17.7

1,095 108
L R L R L

Forelimb length Humerus length Radius length Mc III length Hind limb length Femur length Tibia length Mt III length
2

567
258

557
255

56.41
45.52

55.41
45.82

613
269

611
269

59.81
43.92

59.81
44.02

59.81
44.82

208 101 669 297 257 115

200 102 668 296 258 114

36.7 17.8 66.61

44.42

35.9 18.3 66.61

44.32

38.4 17.2

38.6 17.1

241 103 693 310 266 117

241 101 694 309 268 117

39.3 16.8 68.01 44.72 38.4 16.9

39.4 16.5 68.01

44.52

37.8 17.4 68.0'

44.52

38.6 16.9

38.5 17.0

'Per cent of spine length. Per cent of limb length. I R = right; L = left.

olism do (except the digestive tube, where surface area may be the critical factor). The somatic exponents of the metabolic structures appear to agree closely with those of other mammals. Other factors must be responsible for the high somatic exponents for the skin, spleen, and reproductive organs.
LINEAR MEASUREMENTS

Linear measurements are given in table 2. Body length, tail length, and diameters and circumferences of the thorax were measured on the body before it was skinned. Spine length is the length of the fresh ligamentary spine, measured along the curves, from the anterior margin of the atlas at the dorsal midline to the posterior margin of the centrum of the last lumbar vertebra. (The spine is short in the female lion because she had only six lumbars instead of the usual seven; the seventh was incorporated into the sacrum.) To determine the difference in length between a

fresh ligamentary spine and the same spine after it is cleaned and disarticulated, the spine of the female lion was re-articulated and measured 16 years after it was cleaned. Length was 1030 mm, a difference of 9 mm (less than 1%) from the original fresh ligamentary length. All measurementsof individual bones of the male lion were made on the fresh disarticulated skeleton, those of the female on the dried disarticulated skeleton. Lengths of limb bones are the more meaningful "functional length" (Howell, 1944), rather than the usual greatest length. Limb length is the sum of the lengths of the three segments. Linear measurements of the digestive tube are given in table 3, together with the mean relative lengths for Latimer's domestic cats. In measuring the small intestine of the male lion and the leopard, the mesentery was cut, the intestine extended in a straight line (but not stretched), and the length measured with

ALLOMETRIC RELATIONSHIPS IN CATS

TABLE 3.

513

Linear measurements in mm and relative lengths of digestive tube in felids

Felis leo ci Length Per cent body length Felis leo 9 Length Per cent body length Felis pardus a' Per cent Length body length Felis domestica Per cent body length d' Y 521 390 276

Body length Digestive tube

esophagus small intestine

1,448 10,059
610 8,077 v

695
558

1,610 8,500
-

1,257 528
426
}

509
378 262

6,850

3,864 480 3,000

307 239

colon
c

1,100 1,372 80 95 300 75 87 384 31 51 49

rectum caecum

graph are the means of these five individuals. Accurate measurements of the length of fresh intestines of large mammals are almost impossible, and the recorded difference between the lions is probably not DISCUSSION OF LINEAR MEASUREMENTS real. Since even the smaller ratio, for gut I tried measuring limb segments on the length/body length in the female, is much carcass, as Latimer did, but abandoned greater than in the domestic cat, a positive this in favor of the much more accurate allometric relationship is indicated. The measurements of the bones. This, of remarkably short digestive tube of the course, has the disadvantage of making my leopard, which is relatively shorter than in the domestic cat, suggests on the condata incomparablewith Latimer's. Body proportions may be conveniently trary that there may be no consistent ratio expressed in a pictorial graph, in which in the Felidae. spine length is taken as 100 and lengths SUMMARY of head, limbs, and tail are represented as 1. The cats (family Felidae) provide percentages of spine length (fig. 3). excellent material for studying allometric Measurements of three dry skeletons were relationships,because great range in organadded to those of the two individuals reism size is associated with minor differported in this paper. The figures on the ences in body form, habits, and behavior. A full-grown lion is about 60 times as 100 heavy as an average adult domestic cat. 2. In the Felidae, body weight/organ weight ratios fall into two categories: (a) relative organ weight is constant or increases with increasing body size (isometry or positive allometry) (b) relative organ weight decreases with increasing body size (negative allometry). N=5 3. Structures associated directly or inFIG. 3. Pictorial graph of body proportions in Felis leo. Limb lengths are indicated as percent- directly with locomotor performance are ages of spine length (= 100). Proportions of the isometric or show positive allometry. three limb segments are indicated as percentages 4. Structures associated with metaboof total limb length. The dotted lines connect lism or with the central nervous system elbow and knee, wrist and ankle, and distal ends show negative allometry. of third metapodials.
---0 \

a tape. The method of measuring the intestine of the female lion was not recorded. In all cases the stomach was omitted in computing length of the digestive tube, as was also done by Latimer.

514

D. DWIGHT DAVIS ments of the digestive system of the adult cat. Anat. Rec., 68: 469-480. . 1938. The weights of the brain and of its parts, of the spinal cord and of the eyeballs in the adult cat. J. Comp. Neur., 68: 395-404. . 1939. The weights of the hypophysis, thyroid, and suprarenals in the adult cat. Growth, 3: 435-445. . 1940. The prenatal growth of the cat. X. The weight of the spleen in the fetal period and in the adult. Growth, 4: 259-265. . 1941. The prenatal growth of the cat. XI. The weight of the integument in the fetus and in the adult cat. Growth, 5: 285292. * 1942. The prenatal growth of the cat. XII. The weight of the heart in the fetal and in the adult cat. Growth, 6: 341-349. . 1943. The prenatal growth of the cat. XIII. The weights of the lungs, trachea, and larynx in the fetal and in the adult cat. Growth, 7: 239-250. . 1944a. The prenatal growth of the cat. XIV. The weight of the skeleton in the fetal and in the adult cat. Growth, 8: 149-158. . 1944b. The prenatal growth of the cat. XV. The weight of the musculature in the fetal and in the adult cat. Growth, 8: 205219. R. MEINERTZHAGEN, 1938. Some weights and measurements of large mammals. Proc. Zool. Soc. London, 108 (A): 433-439. RENSCH, BERNHARD.1948. Organproportionen und Korpergrosse bei V6geln und Saugetieren. Zool. Jhb. (allg.), 61: 337-412. . 1960. Evolution above the species level. Columbia Univ. Press, New York. 419 p. W. ROBERTSON-BULLOCK, 1962. The weight of the African elephant Loxodonta africana. Proc. Zool. Soc. London, 138: 133-135. SCHILLER, H.-J. 1959. Das Herz des Lowen (Felis leo L.). Morph. Jhb., 100: 163-184. THOMPSON, D. W. 1952. On growth and form. Ed. 2. Cambridge Univ. Press. v. 1, 464 p.

5. Weight of the digestive tube is isometric, but length shows no consistent relation to body length. 6. "Universal" or mean somatic exponents for body weight/organ weight ratios in mammals are of dubious value, since in some cases they are incompatible with physiological requirements.
AKNOWLEDGMENTS

My thanks are due Mrs. Dorothy Foss, former osteologist of the Museum, for assistance in processing the first lion. Mrs. Edward Levin assisted in processing the second lion and the leopard, and did much of the calculating. Dr. George Rabb assisted with the leopard.
LITERATURE BRUMMELKAMP, CITED

R. 1940. Brainweight and bodysize (a study of the cephalization problem). Verh. Ned. Akad. Wet., (2) 39, no. 5: 1-57. CRILE, GEORGE, AND D. P. QUIRING. 1940. A record of the body weight and certain organ and gland weights of 3,690 animals. Ohio J. Sci., 40: 219-259. FRICIK, H. 1957a. Quantitative Untersuchungen an athiopischen Saugetieren. Anat. Anz., 104: 305-333. 1957b. Betrachtungen ilber die Beziehungen zwischen Korpergewicht und Organgewicht. Zeitschr. Saug., 22: 193-207. HOWELL, A. B. 1944. Speed in animals. Univ. Chicago Press, 270 p. LATIMER, H. B. 1936. Weights and linear measurements of the adult cat. Amer. J. Anat., 58: 329-347. 1937. The weights and linear measure-