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QUARTERLY JOURNAL ~ of the FLORIDA ACADEMY OF SCIENCES:
VOLUME 31
Editor PIERCE BRODKORB
Published by the
FLormpA ACADEMY OF SCIENCES Gainesville, Florida 1968
PUBLICATION DATES OF VOLUME 31
NuMBER 1: January 29, 1969 NuMBER 2: May 22, 1969 NuMBER 3: July 25, 1969 NuMBER 4: August 28, 1969
New Taxa PROPOSED IN VOLUME 31 +Eupatagus inggrZachos (Echinoidea: Spatangidae ) tAmbystoma tiheni Holman (Amphibia: Ambystomatidae ) ¢+Hyla swanstoni Holman (Amphibia: Hylidae) +Zenaidura prior Brodkorb (Aves: Columbidae )
}Pulsatrix arredondoi Brodkorb (Aves: Strigidae)
+Fossil
161
276
283
174
112
CONTENTS OF VOLUME 31
NUMBER 1
A new view of the “synthesis of life” Sidney W. Fox
Florida Academy of Sciences award for 1968
Population trends in some southern plantation counties Joseph S. Vandiver Copper limitation with penicillamine M. I. Djafar, H. R. Camberos, and G. K. Davis
Ecology of American oysters in Old Tampa Bay, Florida John H. Finucane and Ralph W. Campbell II
Capture of a tagged ridley turtle Donald E. Sweat
Returns of tagged pen-reared green turtles Ross Witham and Archie Carr
A review of Anolis angusticeps in the West Indies Albert Schwartz and Richard Thomas
A mass inshore movement of fishes on the Florida coast Carter R. Gilbert
Two birds new to the Pleistocene of Reddick, Florida Richard Brewer
Fs ™ NuMBER 2 Comets, superstitions, and history Duane Koenig Amphioxus in Old Tampa Bay, Florida Gideon E. Nelson
Reproduction and ecology of the longnose killifish Robert A. Martin and John H. Finucane
An extinct Pleistocene owl from Cuba Pierce Brodkorb The bone-eating dog, Borophagus diversidens Cope Walter W. Dalquest
Nesting status of the brown pelican in Florida in 1968 Lovett E. Williams, Jr., and Larry Martin
Hippoboscid flies from cattle egrets in central Florida John B. Funderburg, Margaret L. Gilbert, and Ernest L. Bostelman
Nitrate and ammonia in rumen of steers fed millet D. T. Buchman, R. L. Shirley, and G. B. Killinger
Oyster shell as roughage replacement in cattle diets T. A. Dunn and J. F. Hentges
ill
81 93
101 112 115
130 141 143
150
NUMBER 3
A new echinoid from the Ocala limestone Louis G. Zachos
A Pleistocene herpetofauna from Kendall County, Texas J. Alan Holman
An ancestral mourning dove from Rexroad, Kansas Pierce Brodkorb
Vertebrate fauna of Nichol’s Hammock, a natural trap Sue E. Hirschfeld
Observations on the nature of parasitism CG. E. Pice Acacia choriophylla, a tree new to Florida Taylor R. Alexander
The egg and hatchling of the Suwannee terrapin Crawford G. Jackson, Jr., and Marguerite M. Jackson
Records of the coal skink in Florida Henry M. Stevenson Vertebral anomaly in Micropogon undulatus David J. Hansen
Armadillo distribution in Alabama and northwest Florida James L. Wolfe
Tropical marine fishes from Pensacola, Florida Keitz Haburay, C. F. Crooke, and Robert Hastings
Officers and members of the Academy
NUMBER 4
Relationships of growth and age to organ weights in rats David B. Van Vleck and Virginia Gentle
Pugheadedness in the spotted seatrout Curt D. Rose and Alva H. Harris
Baby loggerhead turtles associated with sargassum weed David K. Caldwell
Lower Oligocene amphibians from Saskatchewan J. Alan Holman
Reptiles and birds of the Cay Sal Bank, Bahama Islands Donald W. Buden and Albert Schwartz
lv
161
165
173
ILE
190
197
199
205
207
209
213
220
241
268
Pl
273
290
Quarterly Journal of the
Florida Academy of Sciences
Vol. 31 March, 1968 No. 1
CONTENTS
A new view of the “synthesis of life” Sidney W. Fox Florida Academy of Sciences award for 1968
Population trends in some southern plantation counties Joseph S. Vandiver Copper limitation with penicillamine M. I. Djafar, H. R. Camberos, and G. K. Davis
Ecology of American oysters in Old Tampa Bay, Florida John H. Finucane and Ralph W. Campbell II
Capture of a tagged ridley turtle Donald E. Sweat
Returns of tagged pen-reared green turtles Ross Witham and Archie Carr
A review of Anolis angusticeps in the West Indies Albert Schwartz and Richard Thomas
A mass inshore movement of fishes on the Florida coast Carter R. Gilbert
Two birds new to the Pleistocene of Reddick, Florida Richard Brewer
Mailed January 29, 1969
QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Editor: Pierce Brodkorb
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Footnotes should be avoided. Give ACKNOWLEDGMENTS in the text and AppREss in paragraph form following Literature Cited.
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TABLES are charged to authors at $20.00 per page or fraction. Titles must be short, but explanatory matter may be given in footnotes. Type each table on a separate sheet, double-spaced, unruled, to fit normal width of page, and place after Literature Cited.
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ILLUsTRATIONS are charged to authors ($17.30 per page, $15.80 per half page. Drawincs should be in India ink, or good board or drafting paper, and lettered by lettering guide or equivalent. Plan linework and lettering for re- duction, so that final width is 4% inches, and final length does not exceed 6% inches. Do not submit illustrations needing reduction by more than one-half. PHotTocraPHs should be of good contrast, on glossy paper. Do not write heavily on the backs of photographs.
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Published by the Florida Academy of Sciences Printed by the Storter Printing Company Gainesville, Florida
QUARTERLY JOURNAL of the FLORIDA ACADEMY OF SCIENCES
Vol. 31 March, 1968 No. I
A New View of the “Synthesis of Life”
SIDNEY W. Fox
THE experimental research which is identified with our labora- tory has been carried out almost entirely within the state of Florida. The credit is properly shared with numerous devoted students and talented associates. Most of what has been done was possible because a few bioscientists in the NASA office disbursed for truly basic research a fraction of the small amount available. This came from a budget which is predominantly committed to space exploration and therefore to necessarily expensive hardware. Par- enthetically, I would like to state the opinion that this latter is itself insufficient to attain supremacy in space. Leadership in space is significant intellectually and societally if one believes, as do I, that the scientific secret of our ultimate origin is in the stars. Whoever tells us most thoroughly and accurately what and where man and his universe came from should be in position to tell us where man is going.
I wish also to express appreciation to a few Florida educators. Our experiments were begun in 1953 in another state in which one educational administrator admonished me not to use the “offensive” word evolution on a college television program. In contrast, I was able to name our activity at the University of Miami an Institute of Molecular Evolution, and I was attracted to the University in part by the fact that before I arrived, its catalog already frankly listed courses in evolution.
The scientific question of the synthesis of life can be analyzed as below.
; ( Synthesis ) Primordial gases —————————-» Small organic molecules [amino acids, (Polymerization ) N bases, ete.] ——————————- Macromolecules [prebiotic protein or nucleic
2 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
(Self Assembly ) (Reproduction and Darwinian Selection ) acid] ————__—_——___ Protocells ——@. —__—_——— s+
Contemporary cells and multicellular organisms
We see that, in the later stages, one needs to concern himself with the evolution from a primitive organism to a contemporary unicellular or multicellular organism. This aspect of evolution is one which Darwin clarified by his selection mechanism. Looking back on the experimental research in the field of abiogenesis since 1950, I believe that the Darwinian part of the answer, explaining evolution from a first organism, represented by far the most intri- cate and involved aspects. That required hundreds of millions of years. When we focus our attention on the true chemical synthesis of an organism starting from nonbiological precursors, such as ac- tivatable atmospheric gases, we see that we have narrowed our questions. We have then stripped away the most forbidding part of what was not so many years ago thought of as a hopelessly im- ponderable problem.
The first step from primitive reactive gases to amino acids, to the nitrogen bases of the nucleic acids, or to monosaccharides rep- resents the area in which the largest number of the few laboratories in the field have worked. Contributions have come from such laboratories as those of Calvin (1962), Ponnamperuma (1965), Oro (1965), Miller (1955), Orgel (Sanchez et al., 1966), Fox (1965, 1968), and others. The next step concerns the formation of the larger molecules, proteins, nucleic acids, and cellulose. Their for- mation is thought of as an appropriate type of polymerization of monomers. We can see also, by further analysis of this problem, that the following step is not one of true synthesis, but is rather one of structural organization of appropriate polymers. This kind of process has been referred to increasingly, by the biochemist, as an act of self-assembly. On this basis we should, strictly, not think and speak of the “synthesis of life”, but rather of the synthesis of precursor polymers and of their self-assembly into protocells. These two steps are the ones to which we have devoted major attention (Fox, 1965). Examples of self-assembly of organelles of cells are now numerous (Seventh International Congress of Bio- chem., 1967). (If, in fact, one may properly employ the jour- nalistic phrase, “secret of life’, that secret may well be the power of self-assembly. )
wy)
Fox: Synthesis of Life
The primitive cell, which our experiments now tell us could arise from reactant gases in less than a few hours (Fox, 1968) had then to evolve to a contemporary cell. The elegant studies that have been carried out by Goulian and Kornberg (1967) and by Spiegelman (1968) involve the dismantling of a contemporary cell and the utilization of contemporary enzymes and primer nucleic ac- ids for further synthesis of a contemporary type of RNA or of DNA, respectively. These processes do not, therefore, answer the funda- mental questions of how enzymes began in the absence of enzymes, of how cells arose in the absence of cells, or of how genes appeared in the absence of genes. Our work is aimed at these questions.
In this connection, I believe also that attempts to define life have an unscientific quality. Although the definition of life has a certain pedagogical value for beginning students, many of us who have thought about the question have come to the conclusion that life is not yet definable. The definition of life has, as Melvin Calvin (1962) stated, the quality of “subjective arbitrariness”. The
TABLE 1
Catalytic activities found in proteinoids
Year Substrate or of Reaction Authors Publication p-Nitropheny! Acetate Fox, Harada, Rohlfing 1962 p-Nitrophenyl Acetate Noguchi, Saito 1962 p-Nitrophenyl Acetate Rohlfing and Fox 1967 p-Nitrophenyl Acetate Usdin, Mitz, Killos 1967 Glucose > glucuronic Fox and Krampitz 1964 acid (GOR ATP = ADP Fox 1965 Durant and Fox 1966 p-Nitrophenyl Phosphate Oshima 1968 Pyruvic acid > acetic Krampitz and Hardebeck 1966 acia = CO, Hardebeck, Krampitz and Wulf 1968 Oxaloacetic acid SS Rohlfing 1967 pyruvic acid + CO. Amination of «- Krampitz, Diehl, and 1967 ketoglutaric acid Nakashima Krampitz, Baars-Diehl, 1968 Haas, and Nakashima Dehydrogenation of Krampitz, Haas, Baars, 1968
glutamic acid and Nakashima
ft QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
facts are not yet entirely at our disposal, at least not in a way that life scientists agree that they are. Any definition is a kind of judgment, so that one who renders a definition of life, i.e. a judg- ment, is making the judgment before having the facts. Accord- ingly, I consider such definitions to be unscientific.
Our research is centered around the production of polymers of amino acids by simple heating under conditions which are not only imputable to the primitive Earth but are wide-spread on the con- temporary Earth.
These protein-like polymers, or proteinoids have been shown in six laboratories to have many kinds of catalytic activity for natural substrates (Table 1). We are thus able to visualize how the first enzymes could have arisen in the absence of enzymes to make them. The appropriate geological environment and diverse amino acids would have been sufficient.
A somewhat unexpected characteristic of the proteinoids is that they represent quite highly ordered polymers (Fig. 1). The order
Tris-HC| 0.1N 02N 0.3N
0.6]
0.4
0.2 a O 100 200 300 400 500 Tube No. 20m.
Fig. 1. Elution pattern of 1:1:1-proteinoidamide fractionated on DEAE- cellulose.
Ot
Fox: Synthesis of Life
results internally from the selective interaction of the amino acids which are heated. We are able to understand this effect as due to the special shape and distribution of charge of each of the eighteen kinds of reactant amino acid. The evidence for this great limitation in heterogeneity has been published recently ( Fox and Nakashima, 1967). A principal significance of such results is that they suggested that prebiotic proteins might first have come into existence in the absence of nucleic acids to order the se- quences in those first proteins (Fox, 1965). This suggestion has also been made more recently by Steinman (1967) working with amino acid reactions in simpler systems. While prebiotic protein need not have had all of the properties of contemporary protein, it would have had to have sufficient to begin the line, e.g., macro- molecular order, metabolism, and cellular structure (Table 2).
We may turn now in more detail to the question of self-assembly and the origin of the cell. In 1954 Professor George Wald (1954) wrote, “For a time this problem of molecular arrangement seemed
TABLE 2
Properties of thermal proteinoids in common with those of contemporary proteins. Fox (1965) and bibliography. Limited heterogeneity Qualitative composition Quantitative composition Range of molecular weight Color tests Solubilities Inclusion of nonamino acid groups Optical activity Salting-in and salting-out properties Precipitability by protein reagents Hypochromicity Infrared absorption maxima Recoverability of amino acids on hydrolysis Susceptibility to proteolytic enzymes Catalytic activity Inactivatability of catalysis by heating in aqueous solution
“Nonrandom” (nonuniform) sequential distribution of residues
Nutritive quality
Morphogenicity
6 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
to present an almost insuperable obstacle in the way of imagining a spontaneous origin of life, or indeed, the Jaboratory synthesis of a living organism. It is still a large and mysterious problem, but it no longer seems insuperable. The change in view has come about because we now realize that it is not altogether necessary to bring order into this situation; a great deal of order is implicit in the molecules themselves.” This concept of the specification of mor- phology by the nature of a precursor macromolecule is an extrapo- lation of the concept, now experimentally supported, that order in primitive proteins was first determined internally by the reacting amino acids. Professor Francis Schmitt (1956) first demonstrated self-assembly of fibrils of the protein collagen (Fig. 2). Dr. A. IL. Oparin has done just one kind of experimentation in the field which he first called the origin of life, and for which I now prefer the nineteenth century phrase of spontaneous generation. Oparin used coacervate droplets (Fig. 3) made from gelatin
. 2. Electron micrograph of microfibrils assembled from collagen, Sie Do Ay lauvornn Slaiamniae (1S )56))),
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Fox: Synthesis of Life
Fig. 3. Coacervate droplets. From Oparin.
(1965) as models of self-ordering, or self-assembly, of the first cell. Among the defects, which Oparin acknowledges, this model em- ploys polymers from living things. As in other experiments, these fail to answer the question of how cells arose when there were no cells to produce them. This question is now answered in principle by the proteinoid, in a process so simple that it resembles that of making instant coffee.
When hot water is poured onto proteinoid, and the resultant clear hot solution is cooled, millions of microscopic spherules sepa- rate (Fig. 4). These are stable to centrifugation, they have a kind of osmotic property, they can be made gram-negative or gram- positive, they have catalytic powers, they can be produced so that
§ QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
©
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Fig. 4. Proteinoid microspheres. Approximately 2u in diameter.
they are motile, they bind polynucleotides as well as dyes, and they also show some selectivity in the passage of molecules through their boundaries (Fig. 5).
As Fig. 6 shows, these boundaries are structured. One may in this figure compare a section of a proteinoid microsphere with one of Bacillus cereus under the electron microscope. Experts who are uninformed on these units often guess wrong as to which is which, reportedly because the artificial particle has a thicker bound- ary. In the same figure we see that the artificial boundary is a double layer. This has permitted some broadening of our under- standing of the Danielli model (1935) of the unit membrane of the contemporary cell, especially with regard to the contribution of lipid.
In Fig. 7, we observe a cyclic phenomenon which is intrinsic to the units composed of proteinoid. In the first picture are shown microspheres which have, during a week in their liquor, developed buds which in appearance, texture, and tenacity re-
Fox: Synthesis of Life 9
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Fig. 5. Time-lapse study in ultraviolet light of diffusion of polymer outward from microsphere when pH is raised slightly. Experiment per- formed by Mr. R. J. McCauley with Dr. Philip O'B. Montgomery.
semble buds on yeast. In the second photomicrograph, the buds have been removed, a phenomenon resulting from heat, electrical, or mechanical shock. These are then stained with Crystal Violet and tranferred to a solution of proteinoid saturated at 37° and
10 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
allowed to cool to 25° over one hour. The buds grow by a kind of heterotrophic process. In the last picture, one can see one of the microspheres with a second generation bud. In this manner, we can visualize an evolution from simple physical processes acting on
Fig. 6. Electron micrographs showing section of Bacillus cereus in upper left. Section of proteinoid microsphere in upper right. Lower micrograph displays double layer in boundary of proteinoid microsphere treated with buffer to raise the pH of a suspension.
Fox: Synthesis of Life i
AOE ie.
Fig. 7. Optical micrograph of proteinoid microsphere replicating by budding and heterotrophic growth. a) Microspheres with buds, b) Buds after removal, c) Microspheres which have grown from stained buds, d) Mi- crosphere with second generation bud.
simply derived polymers to yield the minimal complexity required for reproduction.
While these experiments have not produced a fully contem- porary type of organism, they have shown how a _ proteinaceous microparticle with internally ordered macromolecules, catalytic ac- tivities, and many of the properties of a contemporary cell, includ- ing the ability to participate in a presumably primitive reproductive process can, and could be, spontaneously produced. The necessary geophysical conditions, being found in abundance on the contempo- rary Earth (Fox, 1966), should have earlier been abundant also. This rugged process to the primordial stage can be visualized as having occurred quickly, easily, and often when amino acids with a small proportion of aspartic acid, glutamic acid, or lysine were present.
Without vet defining life, we can use this physical model to help describe life. The model suggests that life is a range of associations of unique chemical materials having intrinsic and characteristic physical properties. Many or all of these properties have their
12 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
simple physical counterparts. In the growth of buds removed from microspheres, for example, we see a similarity to the growth of inorganic crystals. But these units are not inorganic crystals; they are composed of organic material which has an array of cata- lytic activities such as to what would be needed biochemically in the evolution of metabolism. We are thus dealing with complex macromolecules, and especially with supramolecular organization or systems. ‘These systems have properties of association, differ- entiation, and other behavior which are simply not to be found at the more rudimentary or molecular level.
The model answers in one way how cells could arise in the ab- sence of cells, how enzymes could come into existence in the ab- sence of enzymes to make them, and how macromolecular informa- tion could arise in the absence of nucleic acid. Experiments demonstrating one type of origin of primordial ribonucleic acid have been published from our laboratory. Based on these studies, Carl Woese (1968) has reported experiments constructed to ex- plain the origin of the code between RNA and protein, and with Dr. Waehneldt we also have other reports of selective inter- action of RNA, DNA, and proteinoid (Waehneldt and Fox, 1968). The idea that protein first arose before or with RNA and DNA is not a new suggestion; many theorists, e.g. Lederberg (1961), Thimann (1965), have advanced this conceptual possi- bility in the past. What is a new view is the detailed interpreta- tion of our physical model consistent with that suggestion.
Among studies under way are attempts at further contemporiza- tion of the proteinoid microsphere, especially through incorporation of internal mechanisms for synthesis of biopolymer, polyamino acids, and coding polynucleotide. At least two laboratories are studying the potentialities of proteinoids as food. At least two laboratories are studying the relationship of pyrolysis of amino acids to the origin of petroleum. We believe we have seen in the properties of microspheres clues to the study of models of behavior at the most primitive evolutionary level, as in chemotaxis. Some of the data obtained on the proteinoids are being interpreted in a context which is more fundamental than that of the origin of life, the relationship of entropy to general evolution. Needless to say, what has been done so far has helped to discipline plans in the search for extraterrestrial life (Fig. 8). Here we see compari-
Fox: Synthesis of Life 13
~
Fig. 8. “Organized elements” of carbonaceous chondrites (left) and proteinoid microparticles (right).
sons of “organized elements’ from a meteorite with proteinoid microparticles. These have been much republished.
Among other consequences, this study has focused attention on what the Space Research Committee at the University of Miami has titled simply The Survival of Man. Our space research group
14 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
is concerned with identifying the problems and possible solutions of the survival, terrestrial and extraterrestrial, of man as he seem- ingly hurries to his own destruction. - This is a somber note to end on, but not necessarily a pessimistic one. Anyone who has a curiosity about and a reverence for life, I believe, must be con- cerned with how it began and must also be concerned with doing what can be done to insure its continued and internally controlled evolution. In many ways, some of them indirect, study of the synthetic origins of life leads into studies of its maintenance and preservation.
LITERATURE CITED
Carvin, M. 1962. Communication: from molecules to Mars. Bull. Am. Inst. Biol. Sci., vol. 12, no. 5, pp. 29-44.
DANIELLI, J. F., AND H. J. Davson. 1935. The permeability of thin films. Jour. Cellular Comp. Physiol., vol. 5, pp. 495-508.
Fox, S. W. 1965. <A theory of macromolecular and cellular origins. Nature, London, vol. 205, pp. 328-340.
—. 1966. The development of rigorous tests for extraterrestrial life. In Biology and the exploration of Mars, National Academy of Sciences Printing and Publishing Office, Publication 1296 (C. S. Pittendrigh, W. Vishniac, and J. P. T. Pearman, eds.) pp. 223-224.
1968. How did life begin? Science and Technology, no. 74, pp. 51-61.
Fox, S. W., anp T. Nakasuma. 1967. Fractionation and characterization of an amidated thermal 1:1:1-proteinoid. Biochim. et Biophys. Acta, vol. 140, pp. 155-167.
Gou.iAn, M., AnD A. Kornserc. 1967. Enzymic synthesis of DNA. XXIII. Synthesis of circular replicative form of phage 9X174 DNA. Proc. Natl. Acad. Sci. U. S., vol. 58, pp. 1723-1730.
LEDERBERG, J. 1961. Exobiology; experimental approaches to life beyond the Earth. In Science in space. McGraw-Hill Book Co., Inc. (L. V. Berkner and H. Odishaw, eds.), pp. 407-425.
Minter, S. L. 1955. Production of some organic compounds under possible primitive Earth conditions. Jour. Amer. Chem. Soc., vol. 77, pp. 2351-2361.
Oparin, A. I. 1965. The origin of life and the origin of enzymes. Advances in Enzymol., vol. 27, pp. 347-380.
Oro, J. 1965. Stages and mechanisms of prebiological organic synthesis. In The origins of prebiological systems and of their molecular matrices. Academic Press, (S. W. Fox, ed.) pp. 137-171.
Fox: Synthesis of Life 15
PONNAMPERUMA, C. 1965. The chemical origin of life. Science Journal, no. 65, pp. 39-45.
SANCHEZ, R. A., J. P. FERRIS, AND L. E. Orcet. 1966. Cyanoacetylene in prebiotic synthesis. Science, vol. 154, pp. 784-785.
Scumitt, F. O. 1956. Macromolecular interaction patterns in biological systems. Proc. Amer. Phil. Soc., vol. 100, pp. 476-486.
SEVENTH INTERNATIONAL CONGRESS OF BIOCHEMISTRY, TOKYO, JAPAN. 1967. Symposium on the self assembly of structural subunits, and other papers in the Congress.
SPIEGELMAN, S. 1967. The synthesis of living systems. In vitro synthesis of viral RNA. Chem. Eng. News, vol. 45, no. 33, pp. 150-156.
STEINMAN, G. 1967. Sequence generation in prebiological peptide sequence. Arch. Biochem. Biophys., vol. 119, pp. 76-82.
THIMANN, K. V. 1963. The life of bacteria. 2nd ed. Macmillan, New York, p. 834.
WAEHNELDT, T. V., AND S- W. Fox. 1968. The binding of basic pro- teinoids with organismic or thermally synthesized polynucleotides. Bio- chim. Biophys. Acta, vol. 160, pp. 239-245.
Wap, G. 1954. The origin of life. Scientific Am., vol. 192, pp. 44-53.
WokeseE, C. 1968. The fundamental nature of the genetic code: prebiotic interactions between polynucleotides and polyamino acids or_ their derivatives. Proc. Natl. Acad. Sci. U. S., vol. 59, pp. 110-117.
Institute of Molecular Evolution, University of Miami, Coral Gables, Florida.
Quart. Jour. Florida Acad. Sci. 31(1) 1968( 1969 )
Florida Academy of Sciences Award for 1968
THE recipient of the Florida Academy of Sciences honors award for 1968 was Dr. Sidney W. Fox. The medal and citation were presented by Dr. Robert L. Davis at the 32nd annual meeting of the Academy at Stetson University, DeLand, March 21-23.
Dr. Fox received his Ph. D. in Biochemistry from California In- stitute of Technology in 1940. He was on the faculty of chemistry at Iowa State College from 1943-1955, and head of the chemistry section, Iowa Agricultural Experiment Station. He came to Flor- ida State University in 1955, where he was professor of chemistry and director of the Oceanographic Institute and Institute for Space Biosciences. In 1964, he became director of the Institute of Mole- cular Evolution at the University of Miami.
Dr. Fox has written many books and reviews and more than 150 research papers. He is a member of the American Society of Bi- ological Chemists, Fellow of the American Association for the Ad- vancement of Science, American Society for Cell Biology, Society for the Study of Evolution, and others.
Dr. Fox has dedicated 14 years to research on chemical origins of life, in the laboratory as well as the slopes of hot Hawaiian volcanoes. His studies have made him the major proponent of one theory of the origin of life, that of thermal proteinoids. His most pioneering contributions have been to concepts and methods of sequence determinations in proteins, initial studies of organismic synthesis of abnormal protein, and to the theory of organismic evo- lution of protein molecules.
Dr. Fox addressed the Academy on the subject “A New View of the Synthesis of Life.”
Quart. Jour. Florida Acad. Sci. 31(1) 1968(1969 )
Population Trends in Some Southern Plantation Counties
Jos—EPH S. VANDIVER
Aux the world today knows that the heritage of abuse and of neglect which the Negro American has experienced is yielding its evil harvest of bitterness and strife in the metropolitan ghettos. Their background in the plantation areas of the rural South ill- prepared the hundreds of thousands of migrants for the setting of the great city. The illiteracy or near-illiteracy of the rural Negro of the South, his poverty, his lack of skills, and his socialization in dependency combined to provide the human raw materials of the black lumpenproletariat of Megalopolis.
The rural South continues to export its surplus black workers, although, as we shall see, there are some indications that the rate of migration may very récently have slowed down. However that may be, quite unrealistic projections of future growth of the Negro population in the metropolitan centers, based upon an indefinite extension of the past volume of migration or of past growth rates, are occasionally published. The simple and rather obvious fact is that the size of the seedbed population in the rural South is no longer sufficiently large to provide a migratory stream capable of maintaining the past rates of growth of the Negro population in the metropolitan North. Unless new sources of black migrants de- velop, from the West Indies, perhaps, or from Africa, then the migration-based part of the growth of the northern Negro popula- tion must inevitably slow down.
To illustrate the point, in 1910, the eleven states of the Con- federate South held approximately 80 per cent of the total Negro population of the nation, and this population was predominantly rural. Indeed, about 35 per cent of the total Negro population of the nation then lived in a discontinuous belt of 258 nonmetropolitan counties in which the plantation system had been dominant and in which Negroes made up half or more of the population. By 1960, this sweep of “Black Belt” counties was the home of 25 per cent fewer Negroes than in 1910. The entire Confederate South, by the latest estimates, now has not four-fifths, but just under one- half, of the black population of the nation, and more of these reside in the rapidly-growing cities of the South than in the countryside. Fewer than one-quarter of the nation’s Negroes now live in their
18 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
traditional homeland of the plantations and farms and hamlets and small towns of the Confederate South.
Even so, this is an important, and a most severely disadvan- taged segment of our national population. It also is a segment of our population which we tend to forget, now that so much of our attention, and so many of our anxieties, are focused upon the blacks of the ghetto. This report is a brief demographic survey of ten of the Black Belt counties of the plantation South.
The selection of the ten counties here reported upon was made as a step in a larger program of research upon the Southern Black Belt. It was deemed desirable to locate, for more intensive study than could be given to the region as a whole, ten sample counties in which the continued dominance of the cotton plantation re- mained a feature of the rural economy. The period just after World War II, before the recent revolutionary changes in planta- tion organization got underway, was used to designate the coun- ties to be followed through these decades of change.
The counties were selected to represent, not typical counties, but counties in which the cotton plantation as a going concern con- tinued to be dominant. It is important that this be clear: the counties herein studied are presumed to be, not typical in the sense of the average, but typological in the sense that regional traits are preserved in their most intense form.
However, these were not the ten counties of the entire South in which plantation dominance, as revealed by the measures used, was most complete. That basis alone for selection would have yielded ten contiguous counties, all in the alluvial valley known as the “Delta” and all in Mississippi. Rather than choose ten counties in a single block, that county in which plantation domi- nance was most marked in each of several subregions of the South was chosen. Two such subregions each were designated in the Mississippi and Arkansas Delta areas, and another in the Delta lands within Louisiana; therefore, five of the selected ten counties are found in lower Mississippi Valley. One county each was se- lected to represent the greatest intensity of plantation dominance found in upland Mississippi, in Alabama, in Georgia, and in each of the Carolinas. In this way the ten counties of this study located in seven states were chosen. The ten counties are: Marengo, Ala- bama; Crittenden and Desha, Arkansas; Terrell, Georgia; Tensas,
VANDIVER: Population Trends 19
Louisiana; Bolivar, Panola, and Tunica, Mississippi; Scotland, North Carolina; and Lee, South Carolina. <A detailed account of the means of selecting the ten counties is precluded by limitations of time and of space. This matter will be presented, however, in full in a monograph intended for publication. Meanwhile, the author can provide a detailed operational account of the means by which the chosen counties were designated.
Census materials for these counties were examined for the years 1930, 1940, 1950, and 1960; agricultural censuses for inter- vening mid-points during these decades were also consulted for certain relevant information; and vital statistics were collected through 1965, the latest year for which detailed county reports were available.
That the ten counties are indeed more Southern than the South in general can readily be ascertained by glancing at some of the census materials (Table 1). For example, in 1930 these counties contained a total population of 297,000 people, of whom 212,000 were nonwhite. Even after continued decades of heavy migra- tion from the region, the 1960 population of the counties was 267,000, of whom 163,000, or 61 per cent of the total, were non- white. These are still “Black Belt” counties in which nonwhites decidedly outnumber whites.
For all practical purposes, the terms “Negro” and “nonwhite” may be used interchangeably in reference to the total populations of the 10 counties. A few hundred persons classified as “Indian” in Scotland County, North Carolina, and an even smaller number of Chinese in the Delta counties, together account for something less than three-tenths of one per cent of the population of the ten counties, or about one-half of one percent of the nonwhite popula- tion of these counties.
The loss of population of the ten counties since the beginning of World War II would be expected on the basis of their rurality alone; these emphatically are highly rural counties. Indeed, few areas in America have approached the overwhelming rurality which the Southern plantation system produced. As recently as 1930, the total urban population of the ten selected counties was under 18,000, although these fairly populous counties then contained nearly 300,000 residents. Their population at that time was about 94 per cent rural.
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VANDIVER: Population Trends 21
In the ten counties, as in the South and the nation in general, each successive decade has shown growth of the urban population. Towns which were too small to be considered “urban” in 1930 have since passed the 2,500 mark, and the places which were then urban have grown rapidly; by 1960, two of them had become centers of more than 10,000 persons. Although the total popula- tion of the counties in 1960 was about 30,000 smaller than in 1930, the urban population, which had increased from 18,000 to 68,000 had come to represent about one-fourth of the population. By the standards of a region now predominantly urban in residence and a nation 70 per cent so, this is still very rural. From the viewpoint of residents of the ten counties, however, the growth of the towns has been remarkable, particularly when seen in the light of the declining rural base.
In this connection, however, it may be noted that the declining rural population has not meant a decline in productivity. To use only the most locally relevant index of productivity, the total cotton yield of the ten counties increased from 444,000 bales in 1929 to 476,000 bales in 1959, although cotton now required only a fraction as many workers and half as much acreage.
During the same period of time, the rural-farm population of the ten counties declined from over three-fourths of their total (in 1930) to less than 100,000 persons, or about 36 per cent of their total, in 1960. This decline in farm population was produced, here as elsewhere, from an actual decline in the number of people engaged in farming, but much of it also was caused by census classificatory procedures.
Sharecropping, so long dominant as the pattern of labor ar- rangements in the plantation South, involved the allocation of specific acreage to the cropper; the census defined such acreage as a “farm”, and residents thereon as “farm population”. In 1930, 89 per cent of all farm operators in these ten counties were classified as “tenant farmers’, and of course, most tenant farmers were crop- pers. As recently as 1950, more than two-thirds of all tenants, and the majority of all farm operators in these counties were share- croppers.
With the shift during the 1950’s and since from sharecropping to other forms of labor arrangements, the number of sharecroppers has rapidly diminished. By 1960, the combined ten counties had
22 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
fewer remaining croppers than one county among them, Bolivar, Mississippi, had recorded twenty years earlier, and since that time, sharecropping has become increasingly uncommon.
Displaced plantation workers often drifted into the towns and cities of the ten counties, providing a part of the local urban in- crease, even though they did not thereby cease to be dependent upon occasional wage labor on farms for part or most of their subsistence. Many others, however, remained in the country but were simply no longer classified as farm residents.
This can be demonstrated by the simple device of subtracting the populations of all incorporated places from the total population of the ten counties. This procedure would not work if there were unincorprated centers of any particular size. In the ten counties under consideration, however, there are hamlets, but no mill towns or other centers of any size, which are unincorporated. In most counties, the opposite tendency, very widespread in the rural South, to incorporate even very tiny villages has been at work. Thus, we find included in our “town and village” population such incorporated centers as Rohwer, Arkansas, with 86 people, and Dayton, Alabama, with 99.
When the population which lives outside incorporated towns (for convenience, called hereafter the “open-country” population ) is compared with the farm population, we note that the two figures were quite similar to each other in 1930 and 1940 (see Table 1). There always were some persons outside towns who were not farmers, so the “open-country’ population exceeded the farm population, but only by a few thousand. In 1940, for example, the rural-farm population in the 10 counties was 235,000, while the number of open-country residents was 249,000.
Between 1940 and 1960, however, the farm population declined from 235,000 to 97,000, a loss of 138,000. The open-country popula- tion also declined, from 249,000 to 164,000, or a decline of 85,000. This is a big loss, to be sure, a loss of 34 per cent, but the farm population, as recorded, lost nearly 60 per cent. It is maintained that the open-country figure, including many agricultural workers no longer classified as farm residents, is the more meaningful basis for comparison.
In only two of the counties has the growth of towns or cities been rapid enough that the total population of the county in 1960
VANDIVER: Population Trends 23
was larger than it had been 20 years earlier; the other eight coun- ties recorded actual losses. In all of them, however, the flow of migration was outward, for the gains recorded in the two counties were far smaller than the natural increase during this period. The effort to make precise estimates based upon vital statistics is largely pointless because of faulty recording of births and deaths in some counties during much of the time span under consideration and probably still. Using the data as they are, and thereby prob- ably underestimating the true amount of natural increase, one can estimate that the population of the ten counties would have in- creased, between 1940 and 1960, by 110,000 persons, had no migra- tion at all existed. They would, under such circumstances, have shown a 1960 population of 421,000 persons. The actual figure in 1960 was 267,000; on this basis, one may estimate that these ten plantation counties with a 1940 population of only 310,000 people contributed almost half that number, precisely, 154,000, to net out- migration during the two decades. Both races contributed to the net migration from these rural counties, but of the estimated migra- tion of about 154,000, only some 20,000 were white. It may well be, therefore, that the ten counties under consideration exported to ghettos north and south well more than 125,000 people, mostly poor, mostly unskilled.
The age structure of each of the ten counties clearly reflects this migration, especially among nonwhites. To cite but one example, in 1960, Bolivar County, Mississippi, had 4,588 nonwhite male children between 5 and 14 years of age, too young, presumably, to migrate on their own initiative. In the upper teens and 20's, however, migration occurs. The figure of 4,588 between ages 5 and 14 was followed by only 2,648 males between ages 15 and 24, and that ten-year span was followed by merely 1,038 nonwhite males remaining in Bolivar County in the ages of 25 through 34. A county capable of producing 4,588 nonwhite sons between 5 and 14 years of age is capable of holding fewer than one-fourth as many nonwhite males from 25 through 34!
Bolivar County does not provide the extreme combination of demographic and related data to be found among the 10 counties, however. Lee County, South Carolina, happens to combine some statistical measures in such a way as to represent perhaps the most impressive picture of rural wretchedness to be found in the entire
24 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
American South. Such data, incidentally, should not be taken too seriously in respect to a given county. The possibility of sampling errors or even of slack or haphazard enumeration procedures in a given community can distort county data when the population base is relatively small. Even so, that such statistical data can be as- sembled from American materials for the year of 1960 is its own commentary upon the affluent society.
We occasionally read descriptions of Asian and Latin American lands where the median age of the population is as low as 15, and we interpret such data to imply very high reproductive levels and very high loss, from mortality or otherwise, of adults. The nonwhite male population of Lee County, South Carolina, in 1960 recorded a median age of only 14.6 years. The median ages in each of the 10 counties were lower than those for the states in which the counties are located, but no other population segment had so low a figure as this.
The median family size among the nonwhites of Lee County was above the figure of six, and the median family income of non- whites in Lee County in the previous year (1959) had been only $912. Even if we allow for home food production not included in this dollar sum, and let us hope that it existed, it still must come as something of a shock to realize that here is a county in which the average nonwhite family of six persons subsisted on an income, in 1959 dollars, of $912, or $154 per family member per year. With such incomes, the adults sought to prepare the actual majority of the total population, which was under 15 years of age, to face the complexities of metropolitan existence in the latter part of the 20th century.
Lee County is the extreme, but the condition of poverty is usual for nonwhite families in each of the 10 counties studied. For the 10 counties combined, nearly one-third of the nonwhite families re- ported 1959 incomes of under $1,000, and 90 per cent of them had family incomes of less than $3,000.
Vital statistics for the 10 counties were assembled for the entire period of time beginning in 1939, and selected materials are pre- sented in Table 2. The census years of 1940, 1950, and 1960, and the latest available data, were the points selected for inclusion in this table. To minimize the erratic variations found in some of the counties of small base population, the examination of county data
bo Ol
VANDIVER: Population Trends
TABLE 2
Recorded births, death, and amount of natural increase in white and nonwhite population of 10 counties
1939-41 1949-51 1959-61 1963-65 Number of births Total population 21,700 28,542 23,954 22,848 White pcpulation DoD 8,170 6,758 6,316 Nonwhite population 16,125 20,372 17,196 16,532 Number of deaths Total population 9,605 9,312 8,290 8,660 White population 2,184 DAME 2.683 2,951 Nonwhite population (eA 6,700 5,607 5,709 Amount of natural increase Total population 12,095 19,230 15,664 14,188 White population 3,391 5,558 4.075 3,365 Nonwhite population 8,704 13,672 11,589 10,823
Source: U. S. Office of Vital Statistics, 1939-65.
was done through combining the census year, the year just before, and the year just after, thus somewhat smoothing meaningless variations. These consolidated data were summed for the 10 counties and are presented in Table 2, as are the figures for the three latest years currently available.
A glance indicates that the trend of reproduction among mem- bers of both races followed the so-called “baby boom” which took place nationally during the 1940's. It is almost certain that a part of this apparent increase in reproduction in the 10 counties repre- sented improved birth registration, especially in the Arkansas and South Carolina counties, where the figures for the years around 1940 were improbably low and erratic. Other sources of data, how- ever, including the age distributions reported in census materials, confirm an increased fertility in these plantation counties, as well as elsewhere in the nation during the 1940's.
Unlike the national pattern, however, the level of births in these counties began to decline after 1950. The decline was parallel to, and in about the same degree, as the concurrent de- cline in the size of the total population in the counties. The likelihood that the decline in births was largely the result of the decline in the population rather than a change in the reproductive pattern is suggested by the data for Crittenden County, Arkansas. This is the only one of the ten counties which had a slight growth
26 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
of total population during the 1950’s and the only one to show a small increase in the number of births. Jt therefore appears that the factors leading to increased reproduction during the 1940's, aided no doubt by improved registration, were sufficiently strong to causé a marked gain in the number of births to members of both races, despite the declining population base of the nonwhites. Dur- ing the 1950’s, both races declined in numbers in these counties and both races showed some decline in the number of births.
Death statistics show a decline in the number of deaths throughout both decades for nonwhites, a result consistent both with improved health conditions and with a declining population size. Among whites, during the 1940’s when the population was increasing, so did the number of deaths. The number of deaths among whites at the end of the decade of the 1950’s was slightly higher than the level at the beginning of the decade.
Trends in vital statistics during the 1960’s may give some clues to trends in the population base of the counties since the last census. There have been reports from the Deep South which suggest that the pace of migration northward set during the 1940's and 1950’s slowed down somewhat during the early 1960’s. For example, see Ellen S. Bryant (1963). In this set of estimates, Bryant indicated stationary or slightly increasing nonwhite popula- tions in precisely those plantation counties which had undergone heavy losses during the 1950's.
Since the technological revolution in the plantation areas has become virtually complete since 1960, with the use of herbicides reducing the need for hand labor in hoeing, the last step to yield to mechanization, there would appear to be no particular reason for Negroes to remain there. Since, however, conditions have failed to improve in the Northern ghettos during these years, it may simply be that the migration “pull” of the cities has weakened. If this is true, then a piling-up of unemployed or underemployed youths must have occurred since 1960. If, indeed, such a surplus of young persons with plenty of time and with nothing to lose has developed in the Mississippi Delta and in the Alabama Black Belt, here may be an underlying demographic factor of the activism among Negro youth in these areas in the 1960's.
If in fact such an arrest of the outward migration of upper teen- agers and young adults has taken place during the early 1960's,
VANDIVER: Population Trends Dit
one would expect, other things equal, that the number of births would increase. There is no reason to assume that youth in these highly fecund age groups, under-employed and finding no migra- tion goal in the cities, would fail to engage in some degree of reproduction. One would also expect the number of deaths, as the result of an increasing population base, to increase but, be- cause of the youthful concentration of the population, only a very little.
The actual data do not conform to this expected pattern. Among whites, births have continued to decline and deaths have increased since 1960. If this indicates anything in particular about trends, it may suggest that the aging of the white population accounts for the increased deaths and that the national drop in the birth rate is also reflected among plantation whites. Among nonwhites also, both of these trends have been evident, but in both cases the changes have been quite small, distinctly smaller than among whites. A very slow decline in the number of births, and a very slow rise in the number of deaths, has been underway among the nonwhite population of the 10 counties. It is not altogether im- possible that a piling-up of adolescents and young adults is under- way, but that the expected result of increased births failed to develop because of a stronger counterpressure toward reduced fertility, consistent with the nation wide trend underway through- out the 1960's. This is not altogether impossible, but it certainly is not demonstrated by the data.
In summary, this attempt to examine the demographic charac- teristics of ten plantation counties in seven states has indicated that these counties have lost population since 1940, that this loss was very heavy in the farm and open-country categories, and was not offset by the growth of towns and cities. Since the decline of population occurred during years of high natural increase, the total out-migration was quite large when viewed in proportion to the total size of the population base. The loss of population and the volume of migration among nonwhites has been far greater than among whites.
Moreover, the out-migration is much in evidence when the age structure of these counties is examined. All of the counties show median ages, especially for their nonwhite population, below the averages of the nation and of the states in which the counties
28 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
are located. Not only are the families large, but family incomes are very low in all of the counties. In the ten counties considered together, 90 per cent of the nonwhite families reported 1959 median incomes of under $3,000.
The national trend toward a marked increase in the number of births and in the amount of natural increase during the 1940's was strongly evidenced among both races in the ten counties. During the 1950's, and unlike the national pattern, the number of births registered distinct, although not dramatic, declines in the ten coun- ties; this was true among both racial groups. There is indirect basis for supposing that this reduction merely reflected the smaller base population of young adults remaining in these counties after years of sustained outward migration.
Although no discernible economic inducements to encourage persons to remain in these counties are evident, some reports have suggested that the outward movement, especially of nonwhites, slowed down during the 1960’s. The closing of migratory paths presumably would cause a piling-up of young persons in those ages in which migration has usually occurred, that is, in the upper teens and early 20's. Such an accumulation of youth in the popu- lation might be expected to be associated with an increased num- ber of births registered. The vital statistics offer no indication that this is taking place.
LITERATURE CITED
BRYANT, ELLEN S. 1963. Population estimates for Mississippi counties. Mississippi Agric. Exp. Sta., Bull. 693 (July, 1964).
U. S. BurEAU OF THE CENsus. 1930. Composition and characteristics for counties, cities, and townships. 15th Census of the United States, Population, vol. 3.
—. 1940. Characteristics of the population. 16th Census of the United States, Population, vol. 2.
—. 1950. Characteristics of the population. 17th Census of the United States, Population, vol. 2.
———. 1960. Characteristics of the population. 18th Census of the United States, Population, vol. 1.
1930. Type of farm: the Southern states. 15th Census of the United States, Agriculture, vol. 3, part 2.
1935. Statistics by counties with state and U. S. summaries. Census of Agriculture, vol. 1.
VANDIVER: Population Trends 29
1940. Statistics for counties. 16th Census of the United States, Agriculture, vol. 1.
1945. Statistics by counties. Census of Agriculture, vol. 1.
— —. 1950. Counties and state economic areas. 17th Census of the United States, Agriculture, vol. 1.
1954. Counties and state economic areas. Census of Agriculture, vol. 1.
1959. Counties. Census of Agriculture, vol. 1.
U. S. NATIONAL OFFICE OF VITAL Statistics. 1939-1965. Vital Statistics of the United States, [Annual Reports]. Government Printing Office.
Department of Sociology, University of Florida, Gainesville, Flor- ida 32601.
Quart. Jour. Florida Acad. Sci. 31(1) 1968( 1969 )
Copper Limitation with Penicillamine
M. I. Dyarar, H. R. CAmMBEROs, AND G. K. Davis
In work with soft tissue calcification in cattle and sheep oc- curring under field conditions it was noted that frequently, if not always, analysis of liver showed very low levels of copper (Bingley and Carrillo, 1966). It has been of interest that the elastic fibers which are the site of early calcification in the aorta are also the site of degeneration under copper deficiency condi- tions.
Efforts to develop a satisfactory laboratory animal model to study this soft tissue calcification suggested the possible need for a low copper diet, especially if very low levels of the causative agent were to be used. The guinea pig as an herbivorous animal, known to be susceptible to soft tissue calcification, was chosen for our studies.
The initial problem was to develop a satisfactory, palatable, low copper diet. The commercially available low copper diets retained sufficient copper to meet minimum needs of the animals. Our attempts at preparing such a diet faltered on the problem of palatability.
We obtained a pelleted low copper Reid-Briggs diet from Nu- tritional Biochemical Co., Cleveland, Ohio. It consisted of vitamin free casein, alphacel, sucrose, corn oil, and a copper free mineral mixture. Analysis by Bechman Atomic Absorption Spectrometer indicated 4 ppm Cu, significantly higher than our goal of 1 ppm Gut
The difficulties experienced with the production or purchasing of a palatable low copper diet suggested to us the possibility of introducing a chelating agent which might reduce utilization or produce a negative balance causing a functional if not an absolute deficiency.
Since studies with human patients with Wilson’s disease and with normal subjects showed that penicillamine increased urinary copper (Walshe, 1956, 1956a, 1964; Scheinberg and Sternlieb, 1960) and decreased the concentration of copper in the liver (Scheinberg and Sternlieb, 1960, 1963) this compound was selected for study with guinea pigs. The present report is concerned with the results
DJAFAR ET AL.: Copper Limitations With Penicillamine ol
of our efforts to ascertain the influence of orally administered penicillamine upon storage of body copper. Our eventual goal is to produce a guinea pig with low copper reserves that may be used in studies of cardiovascular calcification.
EXPERIMENTAL PROCEDURE
Young male Hartley strain guinea pigs were purchased from Simonsen Lab. Inc., White Bear, Minnesota, and were caged into stainless steel cages with stainless steel wire floors. The metabolic pans were also made from stainless steel. Purina Guinea Pig Chow was fed during the experiment and contained 11.66 ppm of copper as analyzed by the Atomic Absorption Spectrophotometer. D- penicillamine was purchased from Nutritional Biochemical Co. and used in the amount of 150 mg per day administered orally for a period of 10 days. Daily (24 hours) collection of urine was performed before penicillamine treatment for a period of 4 days and during penicillamine treatment for 10 days. Total fecal col- lection was done one time before and 3 times during penicillamine treatment.
The urine was diluted one to one with trichloroacetic acid, 10 per cent, filtered, and stored in the refrigerator for further de- termination. Feed, fecal, and tissue samples were first dried and later ashed in a furnace at 600 C for 16 hours. The ashed samples were digested for one hour on a hot plate with 50 per cent concentrated HCl, filtered, and made up to volume with triple distilled water.
The animals were killed 24 hours after the last dose of d-peni- cillamine and blood, liver, kidney, and spleen were collected.
Hemoglobin was determined by the cyanmethemoglobin method with a Bausch & Lomb Spectronic 20. Copper determina- tions in plasma, feed, urine, fecal matter, and organs were per- formed with the Perkin Elmer 303 Atomic Absorption Spectro- photometer.
RESULTS
In initial tests there was a high mortality rate with guinea pigs receiving 150 mg dl-penicillamine orally, which is apparently more toxic for the guinea pigs than the d-penicillamine, and it was decided to use the latter in this experiment.
32, QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
The effect of d-penicillamine, upon the excretion of copper in the urine during a ten day experimental period was significantly higher (P<.01) than the excretion before the penicillamine was administered to the animals (Table 1).
TABLE 1 Effect of d-penicillamine on copper retention in guinea pigs
Treatment Pretreated (4 days) Treated (10 days) Experiment No. I! 2 IL 2,
No. of Animals 8 8 8 8
Intake (mg) 264.9 320.4 243.9 285.3 Fecal (mg) 228.1 281.2 2053 245.0 Absorbed (mg ) 36.8 — 39.2 38.6 40.3 Absorbed (% of intake ) 13.9 p22 15.8 14.1 Urinary (mg) PLD) YT 46.0 44.4 Urinary (% of intake) 7.9 (ell 18.9 15.6 Retention (mg) 15.8 (G55) —7.4 —4,1 Retention (% of intake) 6.0 ‘Dell ~3.1 —1.5 Retention (% of absorption) 42.9 42.1 —19.4 —10.2
The increased excretion of copper in urine started within 24 hours and reached the highest level on the 5th and 8th day.
In order to see the after effect of penicillamine on the urinary copper excretion a second experiment was conducted. The urine copper was determined for 10 days after the last dose of peni- cillamine administration (Fig. 1).
Urinary Copper Excretion (in ppm)
1 4 1 Pre-treatment During treatment After treatment DAYS OF OBSERVATION
Fig. 1. Urinary copper excretion before, during, and after treatment with d-penicillamine.
DJAFAR ET AL.: Copper Limitations With Penicillamine 33
It appeared that urinary copper excretion reached a normal level within 48 hours after the last dose of penicillamine intake.
All the treated animals, except two (one in each experiment), showed a negative copper retention caused by penicillamine treatment. It is also possible that penicillamine had an effect on the absorption of dietary copper. Animals receiving penicillamine had a higher per cent absorption rate than the pretreated ones. However, in these experiments the difference was not statistically significant (Table 1).
The negative retention of copper due to penicillamine adminis- tration was reflected in a significant decrease in copper storage in liver and spleen, but not in kidney.
The two animals on penicillamine treatment that showed a positive retention had an amount of copper in liver and spleen similar to that of the controls and both were significantly higher than those in animals which had negative retention (Table 2).
TABLE 2
The effect of d-penicillamine on copper storage
Liver Spleen Kidney ug/gm ug ug/gm ug ug/gm ug dry weight _ total dry weight _ total dry weight _ total Treatment Normal 96.5 640*°* 39.0 8.80° tice 71.0n.s. Treated 54.5 280 31.2 5.34 61.5 65.4 fee < 05 e201
n.s. not significant
There were no significant differences in the plasma copper, hemoglobin content, and hematocrit between the treated animals 24 hours after the last dose of penicillamine and the controls.
DIscussION
In the preliminary trials we tried to produce a very low copper diet (<1 ppm) that was palatable for guinea pigs. Special feed samples from General Biochemicals Corp., Chagrin Falls, Ohio, and Nutritional Biochemicals Co. of Reid-Briggs formula for guinea
34 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
pigs were also purchased and analyzed. The diet that contained purified casein, sucrose, alphacel, corn oil, and copper free vitamin mineral mixture was given to guinea pigs after gradually changing from the Purina Chow. The pelleted purified feed was less palat- able and produced diarrhea. The copper content of this diet was determined by both the Perkin Elmer and Beckman Atomic Absorption Spectrophotometers and ranged between 2 to 4 ppm. Although we kept these animals on this low diet for more than 3 months, they continued to gain weight and did not show signs of Cu deficiency. Everson et al. (1967) succeeded in producing a laboratory guinea pig diet which contained 0.5 and 0.7 ppm copper. They used non fat dry milk solid, glucose (cerelose), EDTA treated alpha-cellulose, cotton seed oil, salts, and vitamin mineral mixture in the feed and the copper content was determined by the carbamate colorimetric method. We elected to pursue a dif- ferent technique.
Due to these difficulties in securing a diet which contained copper less than 1 ppm as determined by atomic absorption spec- trophotometry, the penicillamine treatment was used to produce animals with a low copper storage in the body.
Experiments conducted in rats have shown that dl and I- penicillamine are toxic and the LD 50 per oral administration has been found to be 365 mg/kg body weight (Aposhian and Aposhian, 1959; Wilson and du Vigneaud, 1950). This toxicity was attributed to the involvement of dl-penicillamine with the pyridoxal-5-phos- phate and the increased pyridoxine excretion in urine. D1-peni- cillamine was shown in our experiments to be toxic for guinea pigs when they received orally about 320 mg/kg of body weight. However, d-penicillamine was not toxic to guinea pigs at a dose of 400 mg per kg of body weight, although the body weight gain was lower during the 10 days treatment.
Walshe (1964) in his clearance studies in Wilson's disease pa- tients supports his hypothesis that penicillamine depletes the body stores of copper. Direct evidence for a fall in the concentration of copper in liver was shown by Scheinberg and Sternlieb (1960). Walshe (1964) suggests the probable immediate action of penicilla- mine is to render the plasma copper available for filtration at the glomerulus by breaking the copper-albumin linkages.
In this experiment the increased urinary copper during the
DJAFAR ET AL.: Copper Limitations With Penicillamine 35
ten days caused a significant decrease of copper storage in the liver and spleen, but not in the kidney. The copper present in the glomeruli and tubules of the kidney, perhaps in process of excretion may have been the cause of the nonsignificant difference in the copper content between the treated animals and the con- trols.
It is our belief that penicillamine given to guinea pigs for a longer period may cause a depletion of the body copper stores and that these animals can then be utilized in experiments re- quiring low copper reserves.
SUMMARY
Guinea pigs treated with d-penicillamine orally for 10 days excreted in their urine 2 to 4 times the copper found in the urine of the controls. The d-penicillamine was not toxic as compared to the dl-penicillamine at a level of approximately 400 mg per kg body weight. The copper storage in liver and spleen was signifi- cantly decreased.
This treatment may be used as the alternative of feeding low copper diet in conditioning guinea pigs for further experiments to study the relationship of copper deficiency to soft tissue calcifica- tion.
LITERATURE CITED
AposHIAN, H. V., anb M. M. Aposnian. 1959. N-acetyl-dl-penicillamine, a new oral protective agent against the lethal effects of mercuric chloride. Jour. Pharmacol. Exptl. Therap., vol. 126, pp. 131-135.
BINGLEY, J. B., AND B. J. Carrmxo. 1966. Hypocuprosis of cattle in the Argentine. Nature, vol. 209, pp. 834-835.
EVERSON, G. J., H. C. Tsar, anp T. I. Wanc. 1967. Copper deficiency in the guinea pig. Jour. Nutr., vol. 93, pp. 533-540.
SCHEINBERG, I. H., anp I. STERNLIEB. 1960. Metal binding in Medicine.
Eds: M. J. Seven and L. A. Johnson. J. B. Lippincott Co., Philadel- phia.
1963. The dual role of the liver in Wilson’s disease. Med. Clin. N. Amer., vol. 47, p. 815.
WatsHE, J. M. 1956. Penicillamine, new oral therapy for Wilson’s disease. Amer. Jour. Med., vol. 21, p. 487.
1956a. Wilson’s disease new oral therapy. Lancet, vol. 1, p. 25.
36 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
1964. Endogenous copper clearance in Wilson’s disease: a study of the mode of action of penicillamine. Clinical Sci., vol. 26, p. 461.
Witson, J. E., anp V. pu VicNEAUD. 1950. Inhibition of the growth of the rat by l-penicillamine and its prevention by aminoethanol and related compounds. Jour. Biol. Chem., vol. 184, pp. 63-70.
Division of Biological Sciences, University of Florida, Gaines- ville, Florida 32601. Supported in part by NIH grant AM 09089.
Quart. Jour. Florida Acad. Sci. 31(1) 1968(1969 )
Ecology of American Oysters in Old Tampa Bay, Florida
Joun H. FINUCANE AND Rap W. CAMPBELL If
THE purpose of this study was to obtain information on spawn- ing, settlement, growth, and survival of the American oyster, Cras- sostrea virginica Gmelin, for use in the commercial production of oysters in Tampa Bay, Fla. Since Old Tampa Bay produces most of the commercial oysters from the Tampa Bay estuarine system, knowledge of oyster ecology will be important in future evaluation of man-made changes to this area. Our investigation of the oyster fishery is part of a Laboratory program to evaluate estuarine re- sources in the eastern Gulf of Mexico and to develop methods of increasing the productive capacity of these areas, particularly those which have been subjected to drastic alteration by contamination and hydraulic engineering.
Fossil shell deposits and early fishery records attest to the fact that oysters have flourished in Tampa Bay over a long period (Cooke, 1945; Dawson, 1953). Commercial production of oyster meats approached 500,000 pounds annually by the late 1800's; but, from 1902 to 1962 the fishery gradually declined to a yield of 5,000 pounds or less per year. In more recent years, oyster land- ings have increased; in 1964, 147,487 pounds valued at $41,252 were marketed from Tampa Bay (Welch, 1965). This rise in production can be attributed mainly to the successful use of cultch on leased oyster grounds that now cover more than 1,000 acres in Old Tampa Bay.
Old Tampa Bay forms the northwestern segment of the Tampa Bay estuarine system. It is 13 miles long, 2 to 6 miles wide, and has a surface area of 78 square miles (Olson and Morrill, 1955). Sixty per cent of the area is 6 to 18 feet deep, and the bottom is generally firm and sandy (Goodell and Gorsline, 1961). Mean annual salinity is approximately 25 0/00 where most oyster leases are now held. Water circulation in Old Tampa Bay is principally tidal and brackish conditions are created by runoff from streams, springs, and surface water. The embayment is re- markably free of industrial pollution, although about 10 million gallons of treated domestic sewage enter the bay each day. Counts of coliform bacteria are within acceptable limits in all seasons and in most areas (U.S. Public Health Service, 1965, unpublished manu-
38 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
script entitled “Data review and literature search for Old Tampa Bay sanitary survey—phase I”). A recent survey indicates that about one-third of the bottom of Old Tampa Bay, or about 15,000 acres, is favorable for the cultivation of oysters (Engle, James B., 1965, unpublished, “Preliminary report on examination of upper Tampa Bay, Florida, for shellfish cultivation potential,” on file at Bureau of Commercial Fisheries, Shellfish Advisory Service, Ox- ford, Md.).
METHODS
Six sampling stations were selected in oyster-producing areas of Old Tampa Bay, three (stations I-III) above and three (stations IV-VI) below Courtney Campbell Parkway (Fig. 1). These were visited weekly, from April through December 1965, for determina- tion of salinity, water temperature, and the placement of spat col- lectors. Surface water samples were taken with a plastic bucket,
and water temperature was measured with a mercury thermometer |
to the nearest 0.1 C. Samples for salinity were transferred into 4 oz. prescription bottles, sealed and later analyzed for salinity by the Mohr-Knudsen titration method.
Spat collectors and wooden holding frames were constructed
according to the design of Butler (1954). Collectors were of cement-board, measuring 4-3/4 by 7 inches. They were “aged” in fresh water before use, to leach out possible repellent residues. One surface of the collector was smooth and the other rough. Two collecting plates were positioned horizontally and placed 1 inch apart in slotted holding frames so that the smooth surfaces were facing each other. Holders were weighted with a brick and suspended 1 ft. from the bottom. Collecting plates were replaced with new ones every 7 days. To obtain long-term information on survival and growth rates, additional “seasonal” collectors were left at each station for 1 to 3 months. Two plates were removed monthly from these collectors.
Spat counts were made with the aid of a dissecting stereomicro- scope. An acetate overlay etched in l-cm squares was placed over the collecting plate to facilitate counting. The average oyster set per 100 cm? was recorded for each station on the basis of total spat appearing on all four plate surfaces. An estimation of oyster
ee
FINUCANE AND CAMPBELL: Ecology of Oysters 39
OYSTER SPAT STATIONS® LEASED OYSTER BEDS
OLDS MAR
I
SAFETY HARBOR
ST. PETERSBURG
82°35’
Fig. 1. Sampling stations and major commercial oyster-producing areas in Old Tampa Bay, Fla. mortality was determined by dividing the number of survivors on each seasonal plate by the average weekly spatfall at the corres- ponding station for the same month. Fouling of the seasonal plates by barnacles, algae, and encrusting silt reduced the efficiency of this cultch as compared to the weekly plates and may have affected the survival and growth of the oysters. Because of the limited foul-
40 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
ing period, we believe that survival of young oysters was compa- rable to oysters growing on natural cultch or in shell bags. Only the seasonal plates at Safety Harbor (station II) and Rocky Point (station IV) remained throughout the investigation and provided some information on comparative oyster mortality north and south of Courtney Campbell Parkway.
OystTER SPATFALL
Over the 9-month sampling period, 618,776 spat were counted on the cement-board panels. Sixty-two per cent came from sta- tions I, H, and III (Fig. 1). The total accumulated set per square centimeter for each station varied from 84 to 450 (Table 1), and most of the spat set within a period of 21 to 30 weeks (Fig. 2). At Oldsmar (station I) over 50 per cent of the set was recorded between May 13 and June 17. The peak setting period was later (June 17 to July 8) at Safety Harbor (station II) and in the lower bay (July 15 or August 1 to August 26).
The spawning season of oysters in Tampa Bay corresponded with that in Santa Rosa Sound in northwest Florida (Butler, 1965).
STATION-1 STATION-II
STATIONAI
STATIONIV STATION YV
STATION-VI
$ 10152025 5 1018 2025 5 1015 20 25 5 1015 2025 51015 20 25 } 5 10152025
APRIL MAY JUNE JULY AUG SEPT OcT NOV DEC
Fig. 2. Duration of oyster set, and (shaded portion of bar) period of 50 per cent spatfall at six stations in Old Tampa Bay.
The greatest number of spat recorded in Old Tampa Bay, however, was 4 to 5 times higher than reported by Butler (1965) and 10 to 15 times higher than reported for Chesapeake Bay (Bahr, 1964).
Ecology of Oysters
FINUCANE AND CAMPBELL:
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QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
42
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FINUCANE AND CAMPBELL: Ecology of Oysters A3
HyproLocy oF OLp TAMPA Bay IN RELATION TO OYSTER CULTURE
The range of salinity in Old Tampa Bay was between 16.3 and 30.3 o/oo but averaged 23.35 o/oo. This average is within the the optimum salinity range of 10.0 to 28.0 0/oo for Crassostrea virginica given by Loosanoff (1965) in Long Island Sound. Salinity was lowest at Oldsmar (Station I) and highest in lower Old Tampa Bay (Fig. 3). Maximum weekly changes were usually less than 2.00
AREA-| AREA-2
STATION | STATION I s STATION IY STATION ¥ STATION Wt
ELE es Pay a7.
30
25 20 S15 10
: °
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1 i
u%
| Dee een Racetrack a pena ul sn |
ete SP at Peele ee Pa bk ee i aw 5 Corb G8 ABCERMDR eGR AED My 9k Ss 88
Fig. 3. Mean monthly salinity (top) and temperature (bottom) in Old Tampa Bay, Fla. April-December 1965.
0/oo. Further evidence of stability in salinity comes from 24 hour studies completed near Safety Harbor (Station II) in 1962 and 1963 where daily changes of only 0.09 to 1.59 0/00 were recorded ( Kelly and Finucane, unpublished manuscript, 1966, entitled “Diel hydro- graphic observations from Tampa Bay, Florida, November 1962 to December 1963,” on file at Bureau of Commercial Fisheries Biologi- cal Laboratory, St. Petersburg Beach, Fla.). This degree of change can be readily tolerated by oysters in Tampa Bay. Ingle and Daw- son (1950) reported values ranging from fresh water to 42.5 0/00 and daily changes as great as 10 0/00 in Apalachicola Bay, Florida, but noted that such marked changes in salinity may impair survival of oysters and quality of the meat.
During our study, water temperature ranged from 9.6 C in December to 34.0 C in July, and averaged 25.9 C. In general, the first spattall was at about 25.0 C, but the first mass spatfall at Oldsmar (Station I) and later in the lower bay began usually at a water temperature near 28.0 C. This finding agrees well with spawning temperatures recorded for oysters in other parts of Flor-
44 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
ida (Ingle and Dawson, 1953). Along the east coast in Long Island Sound and Chesapeake Bay, spawning has occurred at much lower temperatures ( Loosanoff and Engle, 1950; Nelson, 1928).
SURVIVAL AND GROWTH
Comparison of counts from seasonal collectors with those from regular weekly collectors at the same stations yielded estimates of mortality. From June 17 to July 15, survival was only 4.6 per cent at Safety Harbor (Station II), compared with 89.9 per cent at Rocky Point (Station IV). Over a 2 month period, June 17 to August 19, survival at these stations was 3.6 and 39.2 per cent, respectively; after 3 months, mortality was over 96 per cent at both stations. In sections of the country where spatfall is light, such high mortality may cause the fishery to fail, but in Old Tampa Bay high mortality among young oysters appears to act advantageously by reducing survivors to a number that may mature rapidly. Mortality of oyster spat is apparently greatest during the first 2 months after settlement and may reach 86 to 100 per cent (Loosa- noff and Engle, 1950). Chances of survival improve with increase in size, although larger oysters are still subject to disease and preda- tion.
All of the causes of oyster mortality in Old Tampa Bay are not known but a fungus parasite, Dermocystidium marinum, is preva- lent, and a number of oyster predators are present. Sammy M. Ray, Galveston, Texas (personal communication), found that all oysters (mean length 75 mm) collected in Old Tampa Bay in De- cember 1962 were infected with D. marinum. This pathogen is probably the most dangerous parasite of adult oysters in the waters of the southern states (Galtsoff, 1964). We also observed natural predation on both spat and adult oysters by the crown conch (Melongena corona), the drill (Thais haemastoma floridana), the left-handed whelk (Busycon perversum), the blue crab (Callinec- tes sapidus ), and the stone crab (Menippe mercenaria).
Another possible cause of oyster mortality in Tampa Bay is the “oyster leech,” Stylochus inimicus. This parasitic flatworm was abundant on the commercial oyster beds in central and lower Tampa Bay during the summer and fall of 1965. In laboratory ex- periments, as few as three specimens of S. inimicus were found to
FINUCANE AND CAMPBELL: Ecology of Oysters A5
be capable of killing and eating an adult oyster within 1 week. The presence of this animal has been reported periodically in Tampa Bay and other Florida estuaries (Ingle and Dawson, 1953).
The weekly growth increment at all stations was 2.0 to 2.6 mm during the first month. Shell length averaged 17 mm at Rocky Point after 2 months, and 10 mm at Safety Harbor after 3 months. During these periods spat plates were heavily silted, and the outer plate surfaces showed some evidence of predator browsing which affected the growth and survival of these oysters. After 7 months, the average size of the oysters at Safety Harbor was only about 23 mm due to mortality of the older oysters. This rate of growth would produce a marketable oyster (4 inches) in about 2 years; oysters on cultch apparently grow faster, however, for local oyster- men have recovered shells more than 4 inches long in a single year. Robert M. Ingle, Tallahassee, Florida (personal communica- tion), stated that cultch planted at Rocky Point during a favorable growth period from May until February produced some 5 inch oysters. In general, the average growth of oysters in Old Tampa Bay appeared to be less than that reported by Ingle and Dawson (1952) for oysters grown on cultch in Apalachicola Bay. They re- corded a harvest of marketable oysters in about 18 months. In contrast, about 48 months are required to produce a 4 inch oyster in Long Island Sound (Churchill, 1921).
LITERATURE CITED
Baur, L. M. 1964. Final report of Maryland oyster observations for 1964. Chesapeake Biol. Lab. Bull., no. 2, 6 pp.
Butter, P. A. 1954. Selective setting of oyster larvae on artificial cultch. Proc. Nat. Shellfish Assn., vol. 45, pp. 95-102.
1965. Reaction of some estuarine mollusks to environmental factors. Health Educ. Welfare Dep., U.S. Public Health Serv. Publ. 999-WP-25. In Biological problems in water pollution, 3rd Seminar, 1962, pp. 92- 104.
CHURCHILL, E. P., JR. 1921. The oyster and the oyster industry of the Atlantic and Gulf Coast. Rep. U.S. Comm. Fish., 1919, pp. 1-51.
Cooxe, C. W. 1945. Geology of Florida. Florida Geol. Serv., Geol. Bull. no. 29, 339 pp.
Dawson, C. E., Jr. 1953. A survey of the Tampa Bay area. Florida Bd. Conserv., tech. ser., no. 8, 39 pp.
46 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Ga.tsorF, P. S. 1964. The American oyster Crassostrea virginica Gmelin. US. Fish. Wildl. Serv., Fish. Bull. vol. 64, 480 pp.
GoovELL, H. G., anp D. S. Gorstinr. 1961. A sedimentologic study of Tampa Bay, Florida. Internat. Geol. Congr., XXI Sess., Norden, Den- mark, 1960, pp. 75-88.
INGLE, R. M., anp C. E. Dawson, Jr. 1950. Variation in salinity and its relation to the Florida oyster. Salinity variations in Apalachicola Bay. Proc. Gulf & Carib. Fish. Inst., 3rd Ann. Sess., pp. 35-42.
. 1952. Growth of the American oyster, Crassostrea virginica (Gmelin), in Florida waters. Bull. Mar. Sci. Gulf & Carib., vol. 2, no. 2, pp. 393-404.
. 1953. A survey of Apalachicola Bay. Florida Bd. Consery., tech. ser., now LO Sopp:
LoosaNorr, V. L. 1965. The American or eastern oyster. U.S. Fish Wildl. Serv., Circ. 205, 36 pp.
LoosaNnorrF, V. L., AND J. B. ENGLE. 1950. Spawning and setting of oysters in Long Island Sound in 1937, and discussion of the method for predicting the intensity and time of oyster setting. U.S. Fish Wildl. Serv., Fish. Bull., vol. 49, pp. 217-255.
Nextson, T. C. 1928. Relation of spawning of oysters to temperature. Ecology, no. 9, pp. 145-154.
Ouson, F. C. W., AnD J. B. Morritt, Jr. 1955. Literature survey of the Tampa Bay area. Oceanogr. Inst. Florida State Univ., 66 pp.
Wetcu, E. 1965. Summary of Florida commercial marine landings, 1964. Mar. Fish. Res. Rep. to Florida Bd. Conserv., 77 pp.
Bureau of Commercial Fisheries Biological Laboratory, St. Pet- ersburg Beach, Florida 33706. Contribution No. 38.
Quart. Jour. Florida Acad. Sci. 31(1) 1968(1969 )
Capture of a Tagged Ridley Turtle
DonALD E.. SWEAT
On December 10, 1966, the shrimp boat Miss Marathon netted a ridley turtle (Lepidochelys olivacea kempii) between the Mar- quesas Keys and the Dry Tortugas. The turtle bore a small metal tag on the posterior edge of the left front flipper which read, “Premio por Devolucion, Remitir: Dir. Gral. de Pesca, México, D. F.” on one side and “A 1071” on the other side.
The turtle was brought to a Marathon fish house on December 12, 1966, and the Florida Board of Conservation was notified. All pertinent data were sent to the proper Mexican authorities and the turtle was put in the Key West Municipal Aquarium for further observation.
Dr. Archie Carr of the University of Florida was also notified, and he gave the following pertinent information. The turtle, a female, was released by Mexican fisheries officers on May 12, 1966, between Barra de las Calabazas and Cachimba, Tamaulipas (a spot about 90 miles north of Tampico), Mexico. Its carapace measured 65 cm when released.
The turtle was captured on December 10, 1966, after 212 days at liberty... During this time its carapace length had increased by 4 cm (69 cm at capture) and it had traversed approximately 955 miles, if travel was in a straight line.
The tagging and release of this turtle is but a small part of the current Mexican research on sea turtles. As part of the Pro- grama Nacional de Marcado de Tortugas Marinas ( Montoya, 1966) in Mexico, 285 ridley turtles were tagged and released during the months of April through July 1966 (Chavez, 1966). Recovery data on 10 of these 285 turtles were published in January of this year (Chavez, 1967). Four of these first 10 recoveries were made off the Mexican coast, two others were taken off the Texas coast, and four from Louisiana waters. To my knowledge, this capture represents the first of this group of tagged turtles taken in Florida waters.
LITERATURE CITED
Cuavez, H. 1966. Propositos y finalidades. Bol. Progr. Nac. Marc. Tortugas Mar., Mexico, vol. 1, no. 1, pp. 1-16.
48 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
——. 1967. Nota preliminar sobre la recaptura de ejemplares marcados de tortuga lora, Lepidochelys olivacea kempii. Bol. Progr. Nac. Marc. Tortugas Mar., México, vol. 1, no. 6, pp. 1-5.
Montoya, A. E. 1966. Programa nacional de marcado de tortugas marinas. Inst. Nac. Invest. Biol-Pesq., México, pp. 1-39.
Florida Board of Conservation Marine Laboratory, St. Peters- burg, Florida. Contribution No. 1138.
Quart. Jour. Florida Acad. Sci. 31(1) 1968( 1969 )
Returns of Tagged Pen-Reared Green Turtles
Ross WITHAM AND ARCHIE CARR
OnE of the unsolved puzzles in the natural history of sea turtles is the disappearance of the young during the year after hatching. This gap not only hinders studies of the life cycle of the species but also prevents evaluation of the success of restoration projects involving the transplantation of young turtles and the reduction of predation upon the early stages. Hatchling predators are of two kinds: those that attack the eggs and young on shore and those that intercept the hatchlings after they enter the sea. The former can be circumvented by fencing or careful policing of sites of heavy nesting, or by moving eggs to protected hatcheries. For some time it has seemed possible that survival might be further augmented by rearing the young turtles to sizes at which such smaller predators as gulls, robalo, and jackfish would be unable to plague them in the water. At the same time, however, it has appeared possible that this move might actually decrease survival by blocking normal behavioral and ecologic development and making the young turtle unfit to go through its regular life cycle. Within recent months a few data suggesting that this is not true have accumulated. Two relevant cases are reported below.
Ninety-eight green turtles sent to Florida from the hatchery of the Caribbean Conservation Corporation at Tortuguero, Costa Rica were kept for one year in concrete tanks at the House of Refuge Museum at Stuart. These were released into the Indian River on November 10, 1964. Each was tagged with a Monel poultry-wing tag, fastened to the back edge of the right front flipper near the body. Two of these turtles have now been re- covered. The first was caught by Mr. Jack A. Scammell on January 15, 1965, after 64 days of freedom, in the Indian River about seven miles north of the release point. Size and weight were not determined.
The second turtle was recovered by Mr. Vincent Russell off Sandy Cay, Grand Bahama Island, on May 13, 1967, after having been at large for 30 months. It had traveled at least 65 nautical miles, and had crossed the Gulf Stream. It weighed 14 pounds (6350 grams) when retaken; the length was not reported. At the time this turtle was released the carapace length was 187 mm,
50 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
and although it was not weighed, two other turtles with carapace lengths of 184 mm, released at the same time, weighed 810 and 850 grams (1.78 and 1.87 pounds). Thus, weight-gain in the re- captured turtle during the 30 months of free life was about 12 pounds (5500 grams).
As improved tagging techniques are developed more yearlings will be tagged and released, in the hope of substantiating this bit of evidence that pen-reared turtles may be able to adapt to the normal ecologic regimen of the species.
Research and restoration work was supported by the National Science Foundation, the Office of Naval Research, and the Carib- bean Conservation Corporation.
Florida Board of Conservation Marine Laboratory, St. Peters- burg, Florida; Department of Zoology, University of Florida, Gainesville, Florida 32601.
Quart. Jour. Florida Acad. Sci. 31(1) 1968( 1969 )
A Review of Anolis angusticeps in the West Indies
ALBERT SCHWARTZ AND RICHARD THOMAS
In 1856 Hallowell described Anolis angusticeps from Cienfue- gos, Las Villas Province, Cuba. In 1894 a closely related species, Anolis oligaspis Cope, was named from New Providence Island, Bahamas. These two species were ultimately combined (Barbour, 1937, p. 128) as two subspecies of A. angusticeps, the nominate form occurring on Cuba and the Isla de Pinos, and A. a. oligaspis in the Bahamas (New Providence, Andros, Long islands). The Bahaman subspecies has generally been regarded as rare and has been much less well represented in collections than A. a. angustt- ceps. Two additional subspecies have more recently been named, A. a. chickcharneyi Oliver (1948) from South Bimini island in the northwestern Bahamas, and A. a. paternus Hardy (1967) from the Isla de Pinos. Hardy (1967) summarized the pertinent data on all specimens of A. angusticeps available to him but made little comment on the validity of A. a. chickcharneyi. Since we have had considerably more experience in the Bahamas with A. angusti- ceps than previous workers, since the senior author collected the species in Cuba and the Isla de Pinos (under National Science Foundation grants G-3865 and G-6252), and especially since we have made several pertinent observations on the habits of this supposedly rare species, we have attempted to review the accumu- lated data on the variation and habits of Anolis angusticeps.
We have studied 276 specimens of A. angusticeps. Many of the Bahaman specimens are in the Albert Schwartz Field Series (ASFS); the balance of the lizards have been borrowed from the following institutions: Academy of Natural Sciences of Philadel- phia (ANSP), American Museum of Natural History (AMNH), Carnegie Museum (CM), Museo y Biblioteca de Zoologia de la Habana (MBZH), Museum of Comparative Zoology (MCZ), Mu- seum of Zoology, University of Michigan (UMMZ), University of Florida collections (UF/FSM), United States National Museum (USNM), and the Instituto de Biologia, Academia de Ciencias de Cuba (IB). We wish to thank the following persons in charge of these collections: James Boehlke, Edmond V. Malnate, Charles M. Bogert, George W. Foley, Neil D. Richmond, Miguel Jaume, Clarence J. McCoy, Jr., Ernest E. Williams, Walter Auffenberg,
52 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Lewis D. Ober, Charles F. Walker, Doris M. Cochran, James A. Peters, and Orlando H. Garrido. In the field we have had the assistance of Edwin B. Erickson, John R. Feick, William H. Gehr- mann, Jr., Ronald F. Klinikowski, David C. Leber, James D. Small- wood, Barton L. Smith, Willard M. Stitzell, and George R. Zug, whose help in collecting these anoles in Cuba, the Isla de Pinos, and the Bahamas is gratefully acknowledged. We are especially fortunate in having Mr. Malnate check various data for us on the holotypes of A. angusticeps and A. oligaspis, both of which are in the Academy of Natural Sciences of Philadelphia. We have ex- amined the holotypes of A. a. chickcharneyi and A. a. paternus ourselves. The large series of A. angusticeps taken by Thomas W. Schoener on South Bimini and now in the Museum of Compara- tive Zoology has added materially to the quantity of Bahaman specimens.
Hardy (1967) used five characters in his discussions of varia- tion in A. angusticeps. These are 1) presence or absence of ven- tral keeling; 2) number of scales in the first and tenth caudal ver- ticils; 3) number of scales between the seventh canthals (seventh canthals as counted by Oliver, 1948, but first canthal as counted by us, beginning at the anterior margin of the orbit); 4) number of postmental scales; and 5) color. We found other counts useful for differentiation of subspecies in other anoles, and accordingly we have taken data on the number of loreals on one side, the minimal number of scales separating the supraorbital semicircles, the number of scales between the supraorbital semicircles and the interparietal on each side (written as a fraction, i.e., 1/1), number of fourth toe lamellae on phalanges II and III, presence or absence of sculpture on head scales, and presence or absence of keeling on the scales on the anterior face of the thigh.
SYSTEMATIC ACCOUNT
Anolis angusticeps angusticeps Hallowell, 1856
Anolis augusticeps (sic) Hallowell, 1856, Proc. Acad. Nat. Sci. Philadelphia, p. 228 (Cienfuegos, Las Villas Province, Cuba; holotype ANSP 7789).
Definition. A subspecies of A. angusticeps characterized by smooth ventral scales, modally 7 scales between first canthals, modally one row of scales between supraorbital semicircles, median
SCHWARTZ AND THoMaAs: Review of Anolis 53
dorsal scales in first caudal verticil moderate in number, postmen- tal scales modally 4, femoral scales variably keeled or smooth, head scales usually smooth in males, sinuously rugose in females, and ventral color white to whitish, not yellow.
Distribution. Cuba; intergrades between A. a. angusticeps and A. a. paternus known from five localities in Pinar del Rio Province in western Cuba (Fig. 1).
Size. Largest male (UMMZ 70046, vicinity of Soledad, Las Villas Province) 52 mm snout-vent length; largest female (MCZ 11146, Soledad, Las Villas Province ) 43 mm.
Variation. The sample of 107 A. a. angusticeps may be divided into three separate groups for further discussion: 1) Pinar del Rio Province (west), 2) Habana-Matanzas-Las Villas provinces (cen- tral), 3) Camagiiey-Oriente provinces (east). When the entire lot of Cuban material ‘is so divided, certain trends in scalation, especially in the arrangement of the head scales, are shown.
The number of scales between the first canthals varies from 3 to 10. The three samples have the following ranges, means and modes: 1) 3-9; mean 6.4, mode 6; 2) 5-10; mean 6.8, mode 7; 3) 5-9; mean 7.2, mode 7. There is an increase in mean number of snout scales between the first canthals from west to east, with the highest value in the Camagiiey-Oriente sample. The low mean in Pinar del Rio reflects the relationships of the western sample with A. a. paternus of the Isla de Pinos.
The number of loreals varies between 14 and 41, with no ob- vious tendencies toward higher or lower numbers in any region. Means are 25.2 (Pinar del Rio), 25.3 (Habana-Matanzas-Las Villas), and 25.0 (Camagiiey-Oriente ).
All three samples of A. a. angusticeps have the supraorbital semicircles modally separated by one row of scales; this condition occurs in 17 of 33 Pinar del Rio lizards, 30 of 45 lizards from Habana-Matanzas-Las Villas, and 23 of 27 lizards from Camagiiey- Oriente. On the other hand, the semicircles are in contact in some specimens, the number in the three samples being 14 in the west, 13 in the central sample and 4 in the eastern sample. Two lizards in both the Pinar del Rio and the Habana-Matanzas- Las Villas samples have the semicircles separated by two scales; no Camagiiey-Oriente lizard has this condition. The highest inci- dence of the non-modal condition of semicircles in contact occurs
QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
54
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SCHWARTZ AND THoMas: Review of Anolis
in the western Pinar del Rio sample; as in the case of the number of scales across the snout between the first canthals, the high incidence of semicircles in contact in Pinar del Rio reflects the relationship of A. a. paternus to the Pinar del Rio sample. A. a. paternus modally has the semicircles in contact.
The number of scales between the interparietal and the supra- orbital semicircles varies between 0/0 and 2/2, with asymmetrical conditions (0/1, 1/2) also occurring. The mode is strongly 1/1 in all samples. The number of supraorbital scales in contact with the interparietal is modally 0/0 in all samples, with counts of 0/1, 1/1, 1/2, and 2/2 also encountered. None of these latter cate- gories closely approaches the frequency of 0/0 in any sample.
Number of fourth toe lamellae on phalanges IJ and III varies between 14 and 22, with means of 17.6 in the west and east, and 17.4 in the central sample. No geographical trend is present.
Scales in the median dorsal row in the first caudal verticil range from 4 to 7, with means of 5.3 in the west and central lizards, and 5.4 in the Camagiiey-Oriente sample. Tenth verticil scales vary between 3 and 6, with means of 4.4 in the western and eastern samples and 4.1 in the central sample.
Number of postmental scales varies geographically. The total variation in this character is 3 to 8; the means, from west to east are 5.7, 5.1 and 4.7, showing a distinct reduction of number of postmentals from west to east. The modal number is 6 in the Pinar del Rio lizards and 4 in the Camagitiey-Oriente region; the central sample is bimodal, with equal numbers of lizards having 4 and 6 postmentals, and an almost equal number of lizards having 5 postmentals.
From the above data the west-east cline in some scale charac- ters (snout scales between first canthals, contact of supraorbital semicircles, number of postmental scales) is clearly demonstrated. The influence of A. a. paternus on the Pinar del Rio lizards (or, preferably, the intermediate nature of the Pinar del Rio lizards between the subspecies angusticeps and paternus) is reflected in the counts and arrangement of the head scales in the western sample. The ventral scales in some Pinar del Rio lizards will be noted below.
The dewlap color in A. a. angusticeps is variable but apparently not correlated with geography. The basic color is pale orange or
56 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
yellowish orange (peach, apricot; Maerz and Paul, 1950, Pl. 10 D 7, is a good reference, recorded for a specimen from Las Villas). A male from Pinar del Rio had the dewlap yellow, one from the Sierra de Trinidad in Las Villas Province had the dewlap orange, and another male from Trinidad was recorded as having the dew- lap pale orange (Pl. 2 A 10). Ruibal (1964, p. 488) reported _ that the dewlap color of Cuban examples was peach (yellow-pink ), and Collette (1961, p. 139) likewise considered the dewlap peach- colored.
Special mention is necessary of two specimens (IB 864-65) from Cayo Cantiles in the Archipielago de los Canarreos, the string of islands and cays which extends eastward from Punta del Este on the eastern coast of the Isla de Pinos. These two male lizards, which might most properly be expected to represent the Isla de Pinos subspecies paternus, are assignable to the Cuban A. a. angusticeps. Both have the hindlimb and ventral scales smooth. Such a peculiarity of distribution suggests that A. an- gusticeps may have reached the Archipiélago de los Canarreos from mainland Cuba rather than serially from the Isla de Pinos to the west. Additional specimens from this island chain are greatly to be desired; it is even possible that Archipiélago A. angusticeps will ultimately be shown to be a distinctive subspecies restricted to that region. Several other reptiles (Leiocephalus cubensis, Dromicus andreae) have distinctive subspecies in the Canarreos, and A. angusticeps may follow suit.
Specimens examined. Cuba, Pinar del Rio Prov., Cabo de San Antonio, 1 (USNM 51847); north shore, Ensenada de Corrientes, 45 km W Cayuco, 1 (AMNH 81343); Ensenada de Corrientes, 1 (MCZ 55551); Las Martinas, 1 (MCZ 50154); San Waldo, 4 km N Cortés, 1 (IB 1302); Sierra de Guane, 2 (MCZ 11149-50); 3.5 km NE Guane, 1 (AMNH 81344); Luis Lazo, 1 (MCZ 12220): near Vinales, 2 (MCZ 55568-69); pinelands near Vinales, 1 (IB 1060); San Vicente, 6 (AMNH 76510-11, AMNH 78233, AMNH 81342-43, AMNH 81345); 5.6 mi. NW San Vicente, 1 (AMNH 78231); Cueva del Cable, San Vicente, 2 (AMNH 78232, AMNH 78437); San Diego de los Banos, 2 (AMNH_ 58913-14); 1 km N Herradura, 8 (MCZ 59235-42); Rangel, 1 (AMNH 83089); Rio Santa Cruz (not mapped), 1 (USNM 54416); Habana Prov., Marianao, 22 (USNM 160915-23, USNM 160925-37): Bosque de la Habana, 1 (USNM 160924); Cueva de Cotilla, 9 km SW San José de las Lajas, 1 (AMNH 76512); Giines, 1 (AMNH 46520); Cueva de Rincén de Guanabo, 2 mi. E Playa de Guanabo, 1 (AMNH 96498); Sitio Perdido (not mapped), 1 (USNM 75816); Cayo Cantiles, Archipiélago de los Canarreos, 2 (IB 864-65); Matanzas Prov., Matanzas,
SCHWARTZ AND THQMAs: Review of Anolis 57
1 (UMMZ 73924); Las Villas Prov., 5 km SE Paso Caballo, 1 (AMNH 78234): Limones, Cienfuegos, 2 (MCZ 42317-18); Soledad, 2 (MCZ 11146, MCZ 92102): vicinity of Soledad, 1 (UMMZ 70064); Buenos Aires, Sierra de Trinidad, 1 (MCZ 42578); Guajimico, 16 mi. SE Soledad, 1 (AMNH 78238); Trinidad, 3 (AMNH 78235-36, AMNH 81347); 6 km W Trinidad, 1 (MCZ 59254); 1.8 mi. S Topes de Collantes, 1 (AMNH 96499); Cayo de Lanzanillo, 1 (IB 882); cliffs at San José del Lago, 1 (AMNH 78237); Sierra de Jatibonico, 1 (MCZ 7956); Camagiiey Prov., Los Paredones, Sierra de Cubitas, 1 (MCZ 73953); Rio Jigiiey, between Esmeralda and Jaronu, 1 (MCZ 59256); 20 km W Camagiiey, 2 (AMNH 81322, AMNH 83608): Finca San Pablo, ca. 15 km SW Camagiiey, 2 (MCZ 59257-58); Finca Santa Teresa, 9 km W Camagiiey, 4 (MCZ 59244, MCZ 59246-47, MCZ 59262); Granja San Lucas, 9 km W Camagitey, 1 (IB 1208); Playa Santa Lucia, 1 (AMNH 83609); 15 km S Playa Santa Lucia, 3 (MCZ 59259-61 ); Marti, 1 (UMMZ 70992); Oriente Prov., Birama, 32 km SW Victoria de las Tunas, 2 (MCZ 59251-52): near San Ramon, west of Campechuela, 3 (MCZ 59248-50 ); coast south of Pico Turquino, 1 (MCZ 42469); Playa Juragua, 3.7 mi. E Siboney, 1 (AMNH 96500); upper Rio Ovando, 1 (MCZ 52526); La Florida, Baracoa, 1 (IB 883); ca. 9 km SE Moa, + 1000 feet, 1 (MCZ 59253); Cuchillas de Guajimero (not mapped), 1 (MCZ 42558): specimens with no locality data other than Cuba, 6 (ANSP 7997, AMNH 46512, AMNH 46563-65, USNM 83933); data for holotype of A. a. angusticeps (ANSP 7789 ) incorporated into analysis.
Anolis angusticeps paternus Hardy, 1967
Anolis angusticeps paternus Hardy, 1967, Carib. Jour. Sci., 6 (1-2), p. 25 (vicinity of Nueva Gerona, Habana Province, Isla de Pinos: holotype
USNM 142156).
Definition. A subspecies of A. angusticeps characterized by keeled ventral scales, modally 6 scales between first canthals, modally supraorbital semicircles in contact, median dorsal scales in first caudal verticil low in number, postmentals modally 6, fe- moral scales keeled, head scales smooth to weakly sinuously rugose in males, sinuously rugose in females, and ventral color yellow.
Distribution. Isla de Pinos, where known only from the north- ern two-thirds of the island, north of the Ciénaga de Lanier; specimens intermediate between A. a. paternus and A. a. angus- ticeps in Pinar del Rio Province, Cuba ( Fig. 1).
Size. Largest male (AMNH 81326) 49 mm snout-vent length: largest females (AMNH 81334, USNM 142171, MCZ 11143) 38 mm.
Variation. The sample of 49 A. a. paternus is constant in
58 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
several distinctive features of the subspecies, namely the keeled ventral scales and keeled scales on the anterior face of the thigh. The thigh keeling may be weak in some specimens, but it is al- ways present. The number of scales between the first canthals varies between 3 and 8 (mean 6.1, mode 6), and the loreals range from 15-37 (mean 27.9). The supraorbital semicircles are more often in contact than not (55.1 per cent). There are most often 1/1 scales between the interparietal and the semicircles (22 individuals) but an almost equal number (17 lizards) have 0/0 scales (= interparietal in contact with semicircles on both sides ) in this position. Counts of 0/1, 1/1, 1/2, 2/2, 2/0, and 3/1 also are found, with 1/2 having the highest frequency of these. Fourth toe subdigital scales vary from 15 to 22 (mean 18.8). Median dorsal scales in the first caudal verticil range from 4 to 7 (mean 4.6) and in the tenth caudal verticil are either 4 or 5 (mean 4.1). Postmentals vary between 5 and 8 (mean 6.2, mode 6).
Hardy has described the distinguishing features of A. a. pater- nus and our examination of the material agrees completely with his diagnosis. Field notes on series collected by the senior author likewise confirm the presence of the yellow ventral color, which may be very bright in intensity. The dewlap is variable, from pale pink (PI. 1 B 10) to pale orange.
Discussion. A. a. paternus is obviously a derivative of the western populations of A. a. angusticeps, as Hardy pointed out. Ventral keeling occurs in some A. a. angusticeps from Pinar del Rio Province (specimens from San Waldo, Vinales, San Vicente, Herradura, and the vicinity of Guane), with the highest incidence of keeling at the latter two localities. The relationships of the Isla de Pinos herpetofauna to that of Pinar del Rio have been pointed out on several previous occasions, and the situation with A. a. paternus reinforces the closeness of the fauna of these two geographical regions. In the discussion of variation in A. da. angus- ticeps we have pointed out that several features of scutellation are clinal in nature, with the Pinar del Rio populations showing affinities with A. a. paternus. Aside from the differences in ventral keeling and ventral color, the differences between the Cuban and Isla de Pinos subspecies are modal as are the scale distinctions in many subspecies of anoles.
The large number of specimens of A. a. paternus in contrast
SCHWARTZ AND THOMAS: Review of Anolis 59
to the relatively smaller number from all of Cuba intimates that the lizard is more common in the Isla de Pinos. This was indeed the experience of the senior author, who encountered far more A. a. paternus on the Isla than on Cuba itself.
Specimens examined. Isla de Pinos: Nueva Gerona, 24 (USNM 27921-23, USNM 142156-73, UMMZ 60238, MCZ 11147-48); pinelands at Santa Barbara, 2 (MBZH 33); just W Nueva Gerona, east base Sierra de Casas, 17 (AMNH 81323-26, AMNH 81328-40); 8.8 mi. SSW Nueva Gerona, 1 (AMNH 81327); Santa Fé, 1 (USNM 160914); Los Indios, 1 (MCZ 11143).
Anolis angusticeps oligaspis Cope, 1894
Anolis oligaspis Cope, 1894, Proc. Acad. Nat. Sci. Philadelphia, p. 430 (New Providence Island, Bahama Islands; holotype ANSP 26119).
Anolis angusticeps chickcharneyi Oliver, 1948, Amer. Mus. Novitates, 1383:2 (western end of South Bimini Island, Bahama _ Islands; holotype AMNH 68620).
Definition. A subspecies of A. angusticeps characterized by usually smooth ventral scales, modally 9 scales between first can- thals, modally one row of scales between supraorbital semicircles, median dorsal scales in first caudal verticil high in number, post- mental scales modally 6, femoral scales usually smooth but some- times keeled, head scales usually smooth in both sexes, and ven- tral color white, not yellow.
Distribution. Islands of the Great Bahama Bank; specimens examined from North and South Bimini, Andros (including Man- grove Cay), Berry Islands (Frazers Hog Cay), New Providence, Eleuthera, Great Exuma, Long Island and Cat Island; the species has not previously been reported from the Berry Islands nor Great Exuma (Fig. 2).
Size. Largest males (AMNH 115617-VV-2515, Eleuthera; MCZ 93340, South Bimini) 53 mm snout-vent length; largest female (MCZ 93352, South Bimini) 47 mm.
Variation. Considering the entire sample of A. a. oligaspis first, the snout scales between the first canthals vary between 6 and 12, the number of loreals between 21 and 44, the supraorbital semicircles are almost always (94 of 113 lizards) separated by one row of scales, modally there are 1/1 scales between the inter- parietal and the semicircles, and 0/0 supraorbitals in contact with the interparietal, fourth toe subdigital lamellae vary between 15 and 22, first caudal verticil median dorsal scales range from 5 to
60 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Fig. 2. Map of the Bahama Islands. Solid circles indicate stations whence A. a. oligaspis has been examined. Individual localities on New Providence and the Bimini Islands have not been plotted.
9, tenth caudal verticil median dorsals vary from 4 to 6, and post- mentals are 4 to 8.
All Bahaman lizards have been separated into five samples on the basis of geography and for convenience of discussion; these samples are 1) North and South Bimini, the sample is composed of South Bimini lizards with one exception; 2) Andros and the Berry Islands; 3) New Providence; 4) Great Exuma, Long, Cat; 5) Eleuthera. The above survey of the characteristics of A. a. oligaspis can be further broken down on the basis of these samples; such a procedure is necessary to determine the status of A. d. chickcharneyi.
Means of snout scales between the first canthal scales vary from 8.1 (New Providence) to 9.3 (Andros, Eleuthera), with Bimini lizards having a mean of 8.4. Modes of snout scales are 7 (New Providence), 8 (bimode on Andros) or 9 ( Bimini, Andros bimode, Great Exuma, Eleuthera). The Andros sample (range
SCHWARTZ AND THOMAS: Review of Anolis 61
7-12) includes the total range for this scale character in the entire Bahaman lot, and other samples lack only one or the other ex- treme.
Mean number of loreals varies from 28.9 (Bimini) to 33.9 (Eleuthera). The supraorbital semicircles are almost always sepa- rated by one row of scales; exceptions are supraorbitals in contact (13 Bimini, two Andros, two New Providence including the holo- type, two Great Exuma); the only specimen from Cat Island and one from South Bimini have the semicircles separated by 2 scales.
The scales between the interparietal and the supraorbital semi- circles usually are 1/1 (modal condition in Bimini, Andros, Great Exuma), but on New Providence, the nine specimens are evenly divided between the 1/1, 1/2, 2/2 categories and there is thus no mode; on Eleuthera the mode is 2/2 (six specimens) with two lizards having 1/1 and one lizard 1/2.
The number of supraorbitals contacting the interparietal is usually 0/0 (only condition observed on New Providence and Eleuthera ); variants are 0/1 (three lizards), 1/1 (four), 2/2 (two) on Bimini, 0/1 (one) on Andros, 0/1 (one) on Great Exuma.
Fourth toe subdigital lamellae means vary from 18.2 ( Bimini) to 19.6 (Great Exuma). Scales in the first caudal verticil have means from 5.5 (New Providence) to 6.3 (Andros) with Bimini lizards having a mean of 5.8. Means of scales in the tenth caudal verticil range from 4.7 (Great Exuma) to 5.1 (Andros).
Postmentals vary in mean from 4.7 on Bimini to 6.2 on Eleu- thera, with New Providence intermediate (5.4). The modal num- ber of postmentals is six in all populations except that on Bimini, with a mode of 4.
The recognition of A. c. chickcharneyi depends (Oliver, 1948 ) on four characters: 1) six to eight scales between the first (= seventh sensu Oliver) canthals, 2) 24 to 32 loreals, a number intermediate between 17 to 23 in oligaspis and 35 to 38 in angus- ticeps, 3) 34 to 36 lamellae on the fourth toe (a number inter- mediate between 33 or 34 in angusticeps and 36 to 40 in oligaspis ), and 4) 4 postmentals (in contrast to 4 to 6 in angusticeps and 6 in oligaspis). We have not taken total lamellar counts on the fourth toe. There is no doubt that A. a. oligaspis differs (at least modally) from A. a. angusticeps; the status of A. a. chickcharneyi
62 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
in relation to A. a. oligaspis is in question. Therefore, there is no purpose in comparing A. a. chickcharneyi with A. a. angusticeps, and we confine our comparisons of the Bimini populations with those from elsewhere in the Bahamas. Oliver was hampered in his comparisons by having very little material for comparison with his five Bimini lizards; he examined three specimens of - a. angus- ticeps and two specimens of A. a. oligaspis.
Although the number of snout scales between the first canthals is not as Oliver stated (7 to 11 rather than 6 to 8), the Bimini sample does average low (8.5) in this scale feature. However, Bimini lizards are intermediate in mean number of snout scales between 8.1 (New Providence topotypes of oligaspis) and _ all other Bahaman samples (means of 9.0 to 9.3). The modal number (9) of snout scales on Bimini occurs as a bimode on Andros (8 or 9) and is also intermediate between the low mode of 7 (New Providence ) and 9 (Eleuthera, Great Exuma).
Bimini lizards average less (28.9) loreals than any other popu- lation of oligaspis (32.1 to 33.9); the range of the Bimini loreal counts (22 to 34) is completely embraced by those of oligaspis from Andros (22 to 40), Great Exuma (21 to 44) and practically included by those from New Providence ( 24 to 37).
Fourth toe subdigital lamellae on phalanges II and III average less for the Bimini sample (18.2; range 16-21), with means for other samples varying between 18.3 (Eleuthera) and 19.6 (Great Exuma). The combined ranges of fourth toe lamellae of popula- tions other than Bimini are 15-22, so that the counts on Bimini are included within the balance of the counts for oligaspis.
The Bimini sample is the only one which has 4 postmentals as the modal condition; all others have 6 postmentals modally. The Bimini mean of this character (4.7) is coordinately low com- pared with those of other samples (5.4 on New Providence to 6.2 on Eleuthera ).
In summary, we feel that the only claim to recognition for A. a. chickcharneyi is the low number of postmentals. Although both the mean and mode are low in the Bimini lizards, the range of variation of “chickcharneyi’ is enclosed by that of A. a. angus- ticeps and A. a. oligaspis, and virtually so by A. a. paternus. As far as we can determine, there are no chromatic differences be- tween A. angusticeps from Bimini and elsewhere in the Bahamas.
SCHWARTZ AND THOMAS: Review of Anolis 63
Acceptance of A. a. chickcharneyi might necessitate the naming of at least one other Bahaman population, as will be discussed below, and this is a course which we are not prepared to take at this time. Considering the variation in the various samples (some admittedly small) of A. a. oligaspis, we feel that A. a. chickcharneyi does not merit recognition.
Ventral keeling in A. a. oligaspis is usually absent, but Hardy (1967, p. 27) noted the occurrence of keeling in a specimen from Bimini. In addition to Bimini, specimens with keeled ventrals were encountered on lizards from Andros (one with weak keeling ) and Eleuthera (five of nine specimens with keeling). Keeling of the scales on the anterior face of the thigh is even more prevalent in the Bahamas; 13 specimens from Bimini, six from Andros, two from the Berry Islands, two from New Providence, one from Eleu- thera, five from Great Exuma, and three from Long have some degree of keeling of the scales on the anterior face of the thigh.
Considering all of the above information, the lizards from Eleu- thera are unique among A. a. oligaspis in that they modally have 2/2 scales between the semicircles and the interparietal (2/2 occurs only as a minor variant in all other samples) and that they in- clude a high number of individuals with keeled ventral scales. Eleuthera oligaspis also have a high mean number of scales be- tween the first canthals (9.3, which is also the mean on Andros), the highest mean (33.9) number of loreals, always have the semi- circles separated by one row of scales (a feature which is not constant in any other sample of A. angusticeps throughout its range), and have the highest mean (6.2) of postmental scales. Increasing familiarity with the Bahaman herpetofauna makes it clear that reptiles on Eleuthera have a strong tendency to differ from their relatives elsewhere in the Bahamas. In two instances (Sphaerodactylus decoratus, Thomas and Schwartz, 1966; Anolis distichus Schwartz, 1968a) the Eleuthera populations have reached a level of subspecific difference from the balance of the Bahaman populations.
If we accept A. a. chickcharneyi as a valid subspecies, we would be reluctant to leave the Eleuthera A. angusticeps unnamed. Eleuthera lizards differ more from A. a. oligaspis than do Bimini lizards. However, the small series of A. angusticeps from Eleu-
64 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
thera causes us to be circumspect; we regard all Bahaman popula- tions as A. a. oligaspis.
Comparisons: A. a. oligaspis differs from A. a. angusticeps and A. a. paternus in several scale characters. The higher number of snout scales between the first canthals (3-10 in angusticeps, 3-8 in paternus, 7-12 in oligaspis), and the greater number of scales in the first caudal verticil (4-7 in angusticeps and paternus, with means by area from 4.6-5.4; 5-9 in oligaspis with means by island between 5.5 and 6.3) are distinctive. The usually smooth head scales in both sexes of oligaspis serve also to distinguish the subspecies from both angusticeps and paternus.
Dorsal color and pattern of A. a. oligaspis is variable. Notes taken on two living specimens will serve to demonstrate this varia- bility; 1) ASFS V7241, female, dorsal ground color gray with a series of transverse black markings in the style of crossbars but only irregularly so; a pair of pale gray dorsolateral lines; posterior dorsum suffused with rich wood brown; tail banded gray and black; 2) ASFS V6803, female, dorsal ground color brown with yellow longitudinal striae on flanks superimposed on brown bars on a gray ground; color when caught gray with dark mottlings. Males tend to lack the dorsolateral lines ascribed to the females noted above, but in some color phases males have these lines. In some phases, there is also a black, butterfly-shaped lumbar spot. Dewlaps of A. a. oligaspis have been recorded in life as Pl. 10 C 6 (pale peach) on Great Exuma, PI. 2 A 10 (peach) iiagamsouth Bimini, and Pl. 9 B 7 and 9 C 7 (dark peach) on South Bimini. Although there is no evidence that A. a. oligaspis shows variation in dewlap color comparable to that of A. a. angusticeps, our data for the Bahaman subspecies are too limited to be conclusive.
Specimens examined. Bahama Islands, North Bimini, Alicetown, 1 (MCZ 46066): South Bimini, 1 mi. S Alicetown (North Bimini), 6 (UF/FSM 16602-07); west end, 9 (ASFS V2448, CM 32553, CM 32600, AMNH 68616-17, AMNH 68619-20, MCZ 49736, MCZ 93334); northeastern part, 2° (ASFS V10752-53); near Nixon’s Harbour, 1 (CM 32616) oad. to Nixon’s Harbour, west end, 3 (MCZ 93335-37); road to airport, west end, 2 (MCZ 93338-39); 1 to 1.5 mi. SSE northwest point, 16 (MCZ 93340-43, MCZ 93345-46, MCZ 93354-61, MCZ 93365-66); 1.5 mi. S northwest point, 1 (MCZ 93344); 0.5 to 1.5 mi. SSE northwest point, 5 (MCZ 93347-51); 0.75 to 1 mi. WNW airstrip, 2 (MCZ 93352-53); 1 mi. SSE northwest point, 3 (MCZ 93362-64); no data other than South Bimini, 7 (ASFS V7116-
SCHWARTZ AND THoMAs: Review of Anolis 65
19, MCZ 80129-31); no data other than “Bimini Island” or “Bimini”, 2 (UMMZ 118302, UF/FSM 7711); Andros, Fresh Creek, Coakley Town, 1 (MCZ 41990); Little Creek, 2 (UMMZ 118010); Lisbon Creek shore, 1 (AMNH 76312); Bastian Point, 2 (AMNH 76310-11); Mangrove Cay, 11 (AMNH 74487-96, UMMZ 109221); no data other than Andros Island, 1 (USNM 49533); Berry Islands, Frazer's Hog Cay, near center of north- eastern arm, 2 (ASFS V10669-70); New Providence (localities not mapped), 7 mi. W Nassau, 1 (AMNH 76306); Cave Point, 1 (ASFS 10304); 4.8 mi. SW Cave Junction, 3 (ASFS V10379-81); Yamacraw Beach, 1 (ASFS V10359 ); 0.6 mi. NW Yamacraw Beach, 1 (ASFS V7241); 0.3 mi. E Nassau East, 1 (ASFS V10733); data from ANSP 26119, holotype of A. a. oligaspis included in analysis; Eleuthera, Rock Sound, 8 (UMMZ 115617); between Governors Harbour and Savannah Sound, 1 (ASFS V6803); Great Exuma, 3.2 mi. NW George Town, 7 (ASFS V7028-30, ASFS V7063-66); Bahama Sound, southwest of The Forest, 3 (ASFS V7097-99); Long Island, 4.2 mi. S Adderleys, 1 (ASFS V10824); Deadman’s Cay Settlement, 2 (AMNH 76307-08 ); Clarence Town, 4 (MCZ 38016, MCZ 42288-90); Cat Island, Bennetts Harbour, 1 (AMNH 76309).
Hapirat NOTES
Anolis angusticeps is not so often encountered in the field as ‘are many less cryptic anoles. For this reason, its habitat has not been well defined. Collette (1961) recorded A. angusticeps as a primarily tree trunk frequenter in a wooded area near La Habana, and Ruibal (1964) considered it to be a lizard of “open habitats: fence posts, rocks, palm trunks, ...” Others (Alayo, 1955; Buide, 1966) have noted its occurrence in coastal sea grape (Coccoloba) situations. Hardy (1967, p. 30) stated that most of the type series of A. a. paternus was taken in grassy meadowland; a few more were found on the trunks of royal palms. In the Bahamas, Oliver (1948) observed A. angusticeps relatively high (6 to 25 feet) in light-barked trees.
Our own observations in Cuba encompass all of the above ob- servations for that island. Specimens on which habitat notes were taken were found on shrubs and grass (2), rocks (4), fence post in pasture (1), ground (1), trees, including coastal Coccoloba and Terminalia (3). On the Isla de Pinos one specimen was found on a shrub in a pasture.
In the Bahamas A. angusticeps appeared to us to be a more confirmed tree dweller. Most of our specimens were taken at night while they slept on small diameter branches and twigs rela- tively high off the ground (6 to 12 feet) in well developed
66 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
coppice growth (Bahaman coppice at best is distinctly lower than most wooded situations in Cuba). During the day one was taken on a barbed-wire fence (on the wire) adjacent to woods on New Providence and another on a sea grape tree near the coast. On Eleuthera a single specimen was taken on a sloping tree trunk about eight feet above the ground (day). In the Berry Islands - (Frazers Hog Cay) three were observed in the tops of Bursera and Lysiloma saplings 8 to 9 feet above the ground during the heat of the day. On Long Island a single lizard was found in daylight about five feet above the ground on a small sapling in a patch of tamarind trees. However, Mr. Thomas W. Schoener, who observed and collected anoles on Bimini, was able with close observation to find angusticeps more commonly than have other observers. He also found that it occurred on vegetation near the ground as well as high in trees.
The habit of this anole of remaining motionless, often adyress- ing itself to its resting place in order to evade notice, has been re- marked upon (Buide, 1966; Garrido and Schwartz, MS; Collette, 1961). The often lichenate coloration and low profile of this lizard make it particularly difficult to see in such circumstances.
Anolis angusticeps sleeps lengthwise along a small branch, twig, or vine; the tail is extended posteriorly and the distal portion curled loosely about the supporting object. This sleeping posture is virtually identical to that of the recently described A. occultus of Puerto Rico (Thomas, 1965); the sleeping site of the two species is very similar, but A. angusticeps is broader in tolerance of size and kind of sleeping surface and height above the ground. (A. occultus, it should be noted, is an inhabitant of montane rainforest canopy in contrast to the more xenotopic angusticeps). In gross aspect (short tail and limbs, long body, large head) and general coloration the two species are remarkably similar. One of the more notable coloration resemblances is the presence of a dark lumbar spot in some color phases of both species.
We do not suggest that angusticeps and occultus are close rela- tives. Williams and Rivero (1965) have discussed the affinities of occultus, which does not seem to lie with any Antillean species. We merely wish to point out a remarkable similarity. As a matter of fact, the distantly related A. valencienni of Jamaica is also
SCHWARTZ AND THOMAS: Review of Anolis 67
of this adaptive style and is in proportions and coloration generally similar to angusticeps. It too has the habit of pressing closely to the substrate to escape notice (Underwood and Williams, 1959).
In summary we feel that A. angusticeps is a cryptic tree anole whose means of evading capture lies primarily in protective color- ation and slow movements (as opposed to a cursorial tree anole such as A. distichus, which depends much upon speed to evade capture and which is much more in evidence). The possibility that Cuban and Isla de Pinos A. angusticeps are less arboreal than the Bahaman ones needs further investigation. Part of the reason that A. angusticeps has not been more often observed high in trees in Cuba may be the difficulty of seeing the individuals at all at any distance in natural surroundings.
DISCUSSION
Anolis angusticeps is one of a group of reptiles (Sphaerodacty- lus notatus, Sphaerodactylus decoratus, Leiocephalus carinatus, _ Ameiva auberi) whose range is primarily Cuba and the Bahama Islands. According to Schwartz (1968b) these animals are part of a relatively recent group of invaders from Cuba whose Bahaman distribution is in most cases restricted to the islands of the Great Bahama Bank. Other Cuban amphibians and reptiles (Hyla sep- tentrionalis, Eleutherodactylus planirostris, Tarentola americana, Anolis carolinensis, Anolis sagrei, Typhlops biminiensis) show re- lated patterns of distribution.
From a Cuban center of origin, A. angusticeps reached both the Isla de Pinos and the Bahama Islands. Since some lizards from Pinar del Rio Province in western Cuba still show at least one A. a. paternus character (ventral keeling) to some degree, we consider that A. a. paternus has been only recently divided from A. a. angusticeps. Cuba and the Isla de Pinos are separated by the shallow (18 meters) Golfo de Batabano. It seems likely that, when the Isla de Pinos and Cuba were a single unified land mass (a condition which has probably occurred numerous times in the past, since even relatively slight fluctuations in sea level will unite the two land masses) a more or less continuous popula- tion of A. angusticeps occurred from the Isla de Pinos to Oriente Province. In such a population, the southwestern (Isla de Pinos )
68 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
lizards developed keeled scales; some scale characters in this con- tinuous population were clinal from southwest to east (snout scales between first canthals, postmentals, contact of supraorbital semi- circles). With the separation of the Isla de Pinos from Cuba, A. angusticeps of the latter have been separated from the remainder of the cline and their characters have been fixed. Remnants of the older association are still retained by A. angusticeps in Pinar del Rio.
On the Great Bahama Bank, the trend has been toward greater number of postmentals, greater number of scales in the caudal verticils, slightly higher number of loreals, and more scales be- tween the first canthals. Of these, the last is most conspicuous. Some trends in the Bahaman populations have occurred in situ on these islands; the high incidence of ventral keeling in Eleuthera A. a. oligaspis is a concrete example. There are also variations in some scale characters (scales between the first canthals, scales be- tween the semicircles and the interparietal) which occur geo- graphically in a rather haphazard manner. These differences be- tween the Great Bank populations of A. angusticeps suggest that an active differentiation has taken place on several of the islands, but in no case in our opinion has this differentiation gone so far as to be acknowledged nomenclatorially.
It is instructive to compare A. angusticeps with another recent anoline invader of the Bahamas—Anolis distichus from Hispaniola. Not only has this species evolved a series of subspecies on its parent island (whereas A. angusticeps has not), but A. distichus has a series of six Bahaman subspecies. The ranges of the two lizards are comparable, except that A. distichus has reached two islands (Rum Cay, San Salvador) which are off the Great Bank and which together are inhabited by one of the six Bahaman races. The subspecies of A. distichus are easily characterized by head scalation, dewlap color, and body pattern and color. The differen- ces between A. distichus and A. angusticeps may be attributable to more ancient arrival in the Bahamas of A. distichus. There is also the possibility, supported by some evidence in other species, that a cryptic, bark anole (as opposed to more cursorial ones) is less liable to great color variation, presumably because of selective pressure to maintain the successful protective coloration.
SCHWARTZ AND THOMAS: Review of Anolis 69
LITERATURE CITED
AuAyo DatMau, Pastor. 1955. Lista de los reptiles de Cuba. Museo Charles T. Ramsden, Universidad de Oriente, mimeographed, pp. 1-29°7 pls.
Barspour, THomas. 1937. Third list of Antillean reptiles and amphibians. Bull. Mus. Comp. Zool., vol. 82, no. 2, pp. 77-166.
Bume, Mario S. 1966. Reptiles de la Peninsula Hicacos. Poeyana, ser. A, no. 21, pp. 1-12.
CoLLeTTe, Bruce B. 1961. Correlations between ecology and morphology in anoline lizards from Havana, Cuba and southern Florida. Bull. Mus. Comp. Zool., vol. 125, no. 5, pp. 137-162, 6 figs.
Garripo, ORLANDO H., AND ALBERT SCHwaRTz. MS. Aves y reptiles de Cayo Cantiles (Archipiélago de los Canarreos, Cuba).
Harpy, JERRY D., Jr. 1967. Geographic variation in the West Indian lizard, Anolis angusticeps, with the description of a new form, Anolis: angusticeps paternus, subsp. nov., from the Isle of Pines, Cuba (Reptilia: Iguanidae). Carib. Jour. Sci., vol. 6, nos. 1-2, pp. 23-31, 4 figs. : ;
Maerz, A.. aNnD M. Rea Pauxt. 1950. A dictionary of color. New York, McGraw-Hill Book Co., pp. i-vii, 1-23, 138-208, 56 pls.
OxiveR, JAMES A. 1948. The anoline lizards of Bimini, Bahamas. Amer. Mus. Novitates, no. 1383, pp. 1-36, 3 figs.
_ Rurpat, Ropotro. 1964. An annotated checklist and key to the anoline lizards of Cuba. Bull. Mus. Comp. Zool., vol. 130, no. 8, pp. 475-520, 18 figs.
SCHWARTZ, ALBERT. 1968a. Geographic variation in Anolis distichus Cope (Lacertilia, Iguanidae) in the Bahamas Islands and Hispaniola. Bull. Mus. Comp. Zool., vol. 137, no. 2, pp. 255-304, 4 figs., 2 pls.
—— 1968b. The geckos (Sphaerodactylus) of the southern Bahama Islands. Ann. Carnegie Mus., vol. 39, no. 17, pp. 227-271, 5 figs.
Tuomas, RicHarp. 1965. Field observations on Anolis occultus Williams and Rivero. Breviora, Mus. Comp. Zool., no. 231, pp. 10-16, 2 figs. THoMaAsS, RICHARD, AND ALBERT ScHwartTz. 1966. The Sphaerodactylus decoratus complex in the West Indies. Brigham Young Univ. Sci.
Bull., Biol. Ser., vol. 7, no. 4, pp. 1-26, 20 figs.
UNDERWOOpD, GARTH, AND ERNEST WituiaMs. 1959. The anoline lizards of Jamaica. Bull. Inst. Jamaica, Sci. Ser., no. 9, pp. 1-48, 6 figs.
WILLIAMS, ERNEST E., AND JUAN A. Rivero. 1965. A new anole (Sauria, Iguanidae) from Puerto Rico. Breviora, Mus. Comp. Zool., no. 231, pp. 1-9, 4 figs.
Miami-Dade Junior College, Miami, Florida 33167; Rt. 6, Box 382-Al, Tampa, Florida 33610.
Quart. Jour. Florida Acad. Sci. 31(1) 1968( 1969 )
A Mass inshore Movement of Fishes on the Florida Coast
CARTER R. GILBERT
SPORADIC mass shoreward movement of marine fishes and other animals is a fairly frequent phenomenon in certain parts of the world. Periodic fish kills have been reported from Walvis Bay, Southwest Africa (Copenhagen, 1953), and from Concepcion Bay, Chile (Falke, 1950). At both places these kills occur under spe- cific meteorologic and oceanographic conditions, which are coupled with the geography of the region concerned. In United States waters such movements are best known from Mobile Bay, Alabama (Loesch, 1960), where, in contrast to the situations in Africa and South America, mortality is infrequent. Although these move- ments are not common, they are by no means unusual; from 1946 to 1956, inclusive, 35 such occurrences were reported from Mobile Bay, varying in number from none in 1954 to ten in 1950. This phenomenon is locally termed a “Jubilee” or “Alabama Jubilee’, and when it occurs local residents congregate in large numbers to gather the animals so affected.
Large congregations of freshwater fishes have also occasionally been observed, in which neither spawning activity nor obvious physical barriers to migration appear to be involved. One such concentration, which occurred in a small stream near Gainesville, Florida, was investigated in March, 1963. Although the fishes were gathered in large numbers at the water’s edge, they other- wise did not appear to be in distress, and chemical analysis of the water failed to indicate anything abnormal.
At about 3:00 on the afternoon of 17 September 1962, a spec- tacular concentration of fishes was noted along the beach just south of Marineland, Florida, which is situated about 50 miles south of Jacksonville. This beach, which is composed basically of the quartz sand characteristic of the Florida upper east coast, contains a large concentration of coquina rocks at or near the water line, which, in turn, afford a favorable habitat for certain species of shore fishes. Although many fishes were found above the water line, most of the eels so stranded remained in their burrows and were still alive. What mortality had occurred apparently was due solely to prolonged exposure to the air, since those individuals still in the water did not appear to be in distress. A high percent-
GiutBertT: Inshore Movement of Fishes 71
age of the species involved usually live beyond the intertidal area (one species occurring as deep as 1500 feet), whereas others may live in shallow water but rarely are found in the ecological situation prevailing at Marineland beach. Furthermore, with one excep- tion the presumed non-resident species are burrowers, and belong to only three families (Bothidae, Ophidiidae, and Ophichthidae ).
Inasmuch as a “jubilee” has not, to my knowledge, previously been reported in the literature from the Atlantic coast of the United States, the surrounding circumstances seem worthy of dis- cussion. The prevailing meteorologic and oceanographic condi- tions during the Marineland and Mobile Bay jubilees are compared and are shown, for the most part, to be quite different. Although the causes of the Mobile Bay jubilees seem to be well understood, more information is needed before the Marineland phenomenon can be adequately explained. The usual habitats of the fishes col- lected at Marineland also are discussed, and an attempt is made to correlate these with possible movements of the various species.
Loesch (1960) presented a detailed discussion of mass shore- ‘ward movements of fishes and crustaceans in Mobile Bay. He showed that such movements occur as the result of a definite com- bination of conditions, which results in an inshore movement of water that is low in dissolved oxygen and which forces the animals ahead of it. He found that jubilees occur: 1) only in summer; 2) usually before sunrise; 3) usually when the wind on the previous day and during the jubilee is from an easterly direction (a change in wind direction will cause the jubilee to cease); 4) only on a rising tide (a change to a falling tide will stop the jubilee); and 5) when two water masses meet, with the saltier water invading during the jubilee. In addition, the concentration of fishes and invertebrates during a jubilee usually is on the east side of the bay; however, Dr. Herbert Boschung informs me that this may occasionally occur on the opposite shore. Presumably a west-shore jubilee results from the same combination of conditions described below, except that the wind direction is reversed.
Loesch determined that a jubilee results from the following sequence of events. A pocket of highly saline water is present in the deepest area of Mobile Bay, and this pocket is overlain by fresh water entering the bay from the Alabama River. In summer the heavier, salty water tends to lose oxygen and gain carbon
72 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
dioxide, from the combination of high temperature and accumu- lated organic debris. During the day plant life in this pocket of water produces oxygen and uses carbon dioxide, but at night this process is reversed. Normally, the oxygen-poor water remains in the deepest part of the bay; however, a gentle east wind, though of insufficient intensity to mix the water near shore, is nevertheless strong enough to push the surface water west and offshore. As the water moves offshore, it is replaced by the deeper, more saline water, which is pushed shoreward by a rising tide. When, under the conditions of east wind and rising tide, this body of deeper, oxygen-poor water moves very close to shore, the demersal animals crowd to the shore and a jubilee is in progress.
The concentration of fishes observed at Marineland differs from those from Mobile Bay in a number of ways. First, it occurred during the middle of the afternoon. Second, according to the tidal charts (U. S. Coast and Geodetic Survey, 1962), high tide around Marineland on 17 September 1962 occurred about 10:30 AM and low tide about 5:00 PM. Although these times are approximate for the Marineland area, the tide nevertheless was falling there around 3:00 PM. Third, the area where the jubilee was observed is not partially enclosed by land, as is Mobile Bay. Fourth, the jubilee does not seem to have been confined to the immediate area around Marineland, but was noted by John Taylor on the same afternoon just above Matanzas Inlet, about 4 miles to the north. Fifth, no unusual concentration of invertebrates was observed, ex- cept as noted below. Sixth, there are no major freshwater streams entering the ocean around Marineland and thus there is no meet- ing of large salt and fresh water masses. Seventh, no large quan- tity of organic debris, such as is present in Mobile Bay, is believed to be present off Marineland. Wind direction on the afternoon in question was not noted, although the day was very calm, and what wind there was presumably was blowing in from the ocean, i.e., from an easterly direction.
The only unusual invertebrates encountered were pteropod mol- lusks (Gastropoda: Opisthobranchia ), which were present in great numbers, not only at Marineland, but at least as far away as the St. Johns River, about 50 miles to the north (Ted Allen, in litt. to F. J. S. Maturo). Pteropods are pelagic, and sometimes are found in tremendous numbers along beaches in different parts of the world
GitBeRtT: Inshore Movement of Fishes 73
(Abbott, 1954, p. 292). As mentioned previously, the species of fishes believed to have moved into shore at Marineland are all demersal, as are those species involved in the jubilees in Mobile Bay. Although there may be a direct relationship between the unusual concentrations of pteropods and fishes, the ecological dif- ferences between these animals suggest that the two phenomena possibly are coincidental.
F. G. Wood, at the time Curator of Exhibits at Marine Studios, told me that he had seen only one other jubilee in the Marineland area, this having taken place during late summer of 1956. He said that the weather at the time had been hot and the seas very calm for several weeks previously; these conditions had also pre- vailed during early September, 1962. Thus, except for the simi- larity in wind direction, which in this case is probably coincidental, the only really basic similarities between the Marineland and Mo- bile Bay jubilees are the facts that all occurred during the summer and that the fishes involved are bottom dwellers.
Frederick H. Berry has suggested that the jubilee might pos- sibly be the result of deep (175-200 tathoms) subsurface waves over the slope zone off northeastern Florida. However, Paul Struh- saker, who has studied these waves, doubts that these phenomena are related. Struhsaker has also analyzed the bottom temperature data taken during the cruises of the research vessel Theodore N. Gill along the southeastern Atlantic coast, and he informs me (in litt.) that he can find no evidence of any conditions during late summer that might account for the jubilee.
The following points, then, can be made. 1) The fact that the two jubilees observed in the Marineland area both occurred during late summer, following a period of hot, calm weather seems significant. 2) In all likelihood the Marineland jubilees resulted from a chain of as yet undetermined events that were triggered by the prevailing weather conditions. 3) Possibly the fish (and ptero- pods?) were driven toward shore by an oxygen-poor, carbon- dioxide rich water mass, of undetermined origin, moving inshore from deeper water. But if so, 4) the apparent absence of any unusual concentration of bottom-dwelling invertebrates is puzzling. A satisfactory explanation for these phenomena is yet to be found.
Inasmuch as the fishes obtained during the jubilee included a
74 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
mixture of resident and non-resident forms, a review and dis- cussion of the various species encountered seems pertinent.
I. Resident forms commonly encountered over an open bottom:
Trachinotus carolinus (Linnaeus ) Trachinotus faicatus (Linnaeus ) Menticirrhus littoralis (Holbrook ) Umbrina coroides Cuvier Eucinostomus argenteus Baird
II. Resident species inhabiting rocky shore, frequently in inter- tidal areas:
Centropristis striatus striatus (Linnaeus ) (young only ) Rypticus maculatus Holbrook (young only ) Anisotremus surinamensis (Bloch) (young only ) Scorpaena plumieri Bloch (young only )
Abudefduf saxatilis (Linnaeus) (young only ) Abudefduf taurus (Miller and Troschel) (young only ) Labrisomus nuchipinnis (Quoy and Gaimard ) Hypleurochilus geminatus (Wood)
Hypsoblennius hentz (Lesueur )
Gobiesox strumosus Cope
The presence of young only of the first six species above prob- ably is related to the lack of living space for adults, as well as the fact that spawning likely occurred offshore, and the eggs, or young, later drifted in and were deposited among the rocks. In some cases spawning may have occurred well to the south, and the eggs or young were carried north by the Gulf Stream. These species are free swimming, and, with the possible exception of Abudefduf, probably move offshore as they reach a larger size. Except for Centropristis striatus, all continue to live in rock or reef areas as adults.
The last four species never swim free of the rocky substrate, and, as a result, spend their postlarval life in a very limited area. All are found in shallow situations, and seldom occur in water much deeper than 10 or 26 feet. The populations of Labrisomus nuchi- pinnis, Hypleurochilus geminatus, and Hypsoblennius hentz, how- ever, could be a mixture of resident and immigrant individuals, in- asmuch as these species have pelagic larvae. Gobiesox strumosus does not have pelagic larvae, and is the species least likely to be affected by recruitment from the outside. It is unlikely that any
Gmupert: Inshore Movement of Fishes 75
of these four species move through open areas after having passed the larval stage.
A subsequent collection at the Marineland locality the follow- ing summer failed to turn up Anisotremus surinamensis or Rypticus maculatus; however, inasmuch as two other species (Lutjanus gri- seus and Diplodus holbrooki) not taken in the September, 1962, collection were found, this may not be meaningful. Although Courtenay (1967, p. 274) reports that most specimens of Rypticus maculatus he examined were collected at depths of from 15-50 fathoms, I have found this species to be fairly common within a few feet of shore around Anna Maria Island, Manatee County, Florida, in the eastern Gulf of Mexico (UF 10888, 10905, 10942; a total of 15 specimens, collected in July, 1963).
III. Probable non-resident species, thought to have moved in from deeper water:
Centropristis philadelphicus (Linnaeus ) Ophidion grayi (Fowler)
Rissola marginata (DeKay )
Syacium papillosum (Linnaeus ) Citharichthys spilopterus Ginther Etropus microstomus (Gill) Ophichthus ocellatus (Lesueur ) Ophichthus gomesi (Castelnau ) Ophichthus melanoporus Kanazawa Letharchus velifer Goode and Bean Bascanichthys scuticaris (Goode and Bean)
With one exception, all of the above species are frequently en- countered in shallow water, and perhaps are more common in close to shore than is generally realized. The species of Ophidiidae (genera Ophidion and _ Rissola) and Ophichthidae (genera Ophichthus, Letharchus, and Bascanichthys) live in the substrate and come out into the open only at night (Starck and Davis, 1966, pp. 317 and 342; and personal observation), with the cusk eels (ophidiids), in particular, moving free of the burrow. The snake eels (ophicthids) burrow well down into the substrate, and this, in company with their slender bodies, probably accounts for their relative scarcity in trawl collections. The geographic and bathymetric distributions of many species of Ophichthidae is con- sequently poorly known. The use of rotenone poisons has resulted
76 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
in sizeable collections of certain species (e.g., Bascanichthys scu- ticaris) occurring in very shallow water. This would suggest that many of the deep-water species probably are more common than collections indicate. Although the ophidiids and flatfishes live in the substrate, they do not live in deep burrows, as do the ophich- thids; consequently, they are much more easily collected by trawls and dredges.
Of the above species, the only ones with which I have had extensive experience are Citharichthys spilopterus (family Bothi- dae) and Bascanichthys scuticaris (family Ophicthidae). The for- mer is very common in estuarine situations, and frequently is found in completely fresh water. The latter is common around Cedar Key, Florida, on the Gulf coast, where it occurs in oyster beds or in closely similar situations in water a few inches deep.
The record of Ophichthus melanoporus is easily the most inter- esting and significant of those listed above. This species was de- scribed by Kanazawa (1963) from five specimens taken at a depth of 250 fathoms off Andros Island (Bahamas) at COMBAT station 448. The present record, which is the first for United States waters, suggests that the single specimen either is a stray from deeper water, or that the geographic and bathymetric distribution of the species is much more extensive than is presently realized. In view of our imperfect knowledge of the distribution of many species of Ophichthidae, the latter possibility certainly cannot be ignored. Nevertheless, if O. melanoporus should occur regularly within several hundred feet of the surface along the east coast of Florida, it is surprising that the species has never been found in the hundreds of collections made at various depths from this area.
The specimen of Ophichthus melanoporus was collected by John Taylor on the beach just north of Matanzas Inlet, under the same conditions described for the Marineland locality. Several other eel specimens also given to me by Mr. Taylor are of the same species found at Marineland. Inasmuch as Matanzas Inlet is only a short distance away and the ecological conditions were the same as those encountered at Marineland, the specimen of O. melanoporus is discussed concurrently with those taken at the latter locality.
Several subsequent rotenone collections at the site of the jubilee, both in and around the coquina rocks as well as over the open sand
SILBERT: Inshore Movement of Fishes a
bottom, failed to reveal any flatfish, cusk-eels, or snake eels. Al- though this does not necessarily mean that these fish were not present there at the time the collections were made, it neverthe- less is suggestive. Furthermore, the habitat from which Basca- nichthys scuticaris was collected in the eastern Gulf of Mexico is somewhat different from that encountered at Marineland. Finally, the collection of the specimen of Ophichthus melanoporus, which otherwise has been found only in 250 fathoms of water, further indicates that some movement into shallow water has occurred.
ACKNOWLEDGEMENTS
I wish to thank the following individuals: Dr. C. Richard Robins, Institute of Marine Science, University of Miami, for iden- tifying some of the cusk-eels (ophidiids) reported upon and for reviewing and commenting upon this manuscript; Frederick H. Berry, Tropical Atlantic Biological Laboratory, U. S. Fish and Wildlife Service, Miami, Florida, and Paul Struhsaker, University of Hawaii (formerly U. S. Fish and Wildlife Service, Brunswick, Georgia), for comments relating to possible causes of the Marine- land jubilee; Dr. Archie F. Carr, University of Florida, for review- ing this manuscript; Dr. E. Lowe Pierce, University of Florida, for information on tidal changes on the east coast of Florida and also for reviewing this manuscript; Dr. F. J. S. Maturo, University of Florida, for information on the pteropods encountered during the jubilee; John Taylor, U. S. Fish and Wildlife Service, St. Peters- burg Beach, Florida, for the donation of several eel specimens collected during the jubilee at Matanzas Inlet and for information pertaining to this collection; F. G. Wood, Jr., U. S. Naval Missile Center, Point Mugu, California (formerly of Marine Studios, Ma- rineland), for information on previous Marineland jubilees; and Dr. Herbert T. Boschung, Jr., University of Alabama, for additional information on the Mobile Bay jubilees.
LITERATURE CITED Appott, R. Tucker. 1954. American seashells. D. Van Nostrand Co., Inc., New York, N. Y., pp. vii-xiv, 3-541.
CorpENHAGEN, W. J. 1953. The periodic mortality of fish in the Walvis region. South African Jour. Sci., vol. 49, pp. 330-331.
78 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
CourTNEY, JR., WALTER R. 1967. Atlantic fishes of the genus Rypticus (Grammistidae). Proc. Acad. Nat. Sci. Phila., vol. 119, no. 6, pp. 241-293.
FaLkE, H. 1950. Das Fischsterben in der Bucht von Concepcion ( Mittel- chile). Senckenbergiana, vol. 31, pp. 57-77.
KANAZAWA, RopertT H. 1963. Two new species of ophichthid eels from the western Atlantic. Proc. Biol. Soc. Wash., vol. 76, pp. 281-288.
LorscH, Harotp. 1960. Sporadic mass shoreward migrations of demersal fish and crustaceans in Mobile Bay, Alabama. Ecology, vol. 41, no. 2, pp. 292-298:
Starck, IJ, WaLTER A., AND WILLIAM P. Davis. 1966. Night habits of fishes of Alligator Reef, Florida. Ichthyologica, the aquarium journal, vol, 38) no, 4) pp. olls-3oG6) -
U. S. DEPARTMENT OF COMMERCE, Coast AND GEODETIC SuRVEY. 1962. Tide tables - high and low water predictions, east coast of North and South America including Greenland. U. S. Govt. Printing Office, Washington, D. C., pp. 1-283.
Florida State Museum, University of Florida, Gainesville, Florida 32601.
Quart. Jour. Florida Acad. Sci. 31(1) 1968( 1969 )
Two Birds New to the Pleistocene of Reddick, Florida
RICHARD BREWER
In examining material collected from the Pleistocene fossil beds of Reddick, Marion County, Florida, I encountered elements of two taxa not previously reported from this deposit. The taxa are the family Ardeidae, represented by a nearly complete left cora- coid (University of Florida no. PB 9039) and the Mississippi kite, Ictinia misisippiensis, represented by a complete right tarsometa- tarsus (PB 9032).
The heron coracoid (from Site C of Hamon, 1964) resembles that element of Hydranassa tricolor, Leucophoyx thula, and Florida caerulea. The fossil bone is long in relation to the size of the upper part of it. The ratio between the distance from the notch of the procoracoid to the upper end and the distance from the notch of the procoracoid to the internal distal angle was 0.292 in the fossil. For the three contemporary species, means and ex- tremes for the same ratio were
H. tricolor (7 specimens ), 0.333 (.320-.360 ) L. thula (16 specimens ), 0.319 (.297-.346 ) F. caerulea (12 specimens ), 0.309 (.293-.327).
On this basis, the probability that the fossil specimen is from a population with the same mean as the sample of H. tricolor (stand- ard deviation, 0.0134) is less than 0.05. The fossil is somewhat larger than any available specimen of these three species. Its length is 41.7 mm measured from the internal distal angle. Means and extremes for the three contemporary species were
H. tricolor, 38.1 (36.8-40.2) ie thula, 31.6 (34.7-41.3) Hencaermled, 31-6 (35.4-39)7 )..
The fossil is not similar to Palaeophoyx columbiana McCoy, reported from the Pleistocene of Florida (McCoy, 1963). The coracoid of P. columbiana is longer with a very long, slender “shaft” and a high scapular facet.
The fossil kite (from Site B) differed from Ictinia plumbea and resembled I. misisippiensis in having the inner proximal fora- men at the base of the calcaneal ridge rather than medial to it,
80 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
in having the inner calcaneal ridge straight rather than curved laterally, and in having the wing of the trochlea for digit 2 rela- tively narrow. The Mississippi kite is known from _ prehistoric sites in Ohio and Ilinois (Brodkorb, 1964), but this appears to be the first record of its occurrence during the Pleistocene.
Other specimens examined, representing species already re- ported from Reddick (Brodkorb, 1957; Hamon, 1964) were a complete right ulna of Spatula clypeata; the distal portion of a right femur of Coragyps occidentalis; complete left and right femurs, a complete left tibiotarsus, and a complete right humerus of Colinus suilium; and a nearly complete left coracoid of Protocitta dixi.
The avifauna of Reddick identified to species now totals 64, of which 13 species are extinct.
I gratefully make the following acknowledgments: to Western Michigan University for the granting of a sabbatical leave which made this work possible; to the Department of Zoology of the University of Florida for use of its facilities; and to Pierce Brod- korb for allowing me to study the material and for a great deal of help while studying it.
LITERATURE CITED
Bropkors, Prerce. 1957. New passerine birds from the Pleistocene of Reddick, Florida. Jour. Paleontology, vol. 31, pp. 129-138, 1 text- fig., 20 pl.
——. 1964. Catalogue of fossil birds. Part 2 (Anseriformes through Galliformes). Bull. Florida State Mus., vol. 8, pp. 195-335.
Hamon, J. Hiri. 1964. Osteology and paleontology of the passerine birds of the Reddick, Florida, Pleistocene. Florida Geol. Surv. Geol. Bull, no: 445 210 pp:,, figs 13:
McCoy, Joun J. 1963. The fossil avifauna of the Itchtucknee River, Florida. Auk, vol. 80; pp. 335-351, fig. 3:
C. C. Adams Center for Ecological Studies, Department of Biology, Western Michigan University, Kalamazoo, Michigan.
Quart. Jour. Florida Acad. Sci. 31(1) 1968(1969 )
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Quarterly Journal of the — Florida Academy of Sciences
Vol. 31 June, 1968 No. 2
CONTENTS
Comets, superstitions, and history Duane Koenig 81 Amphioxus in Old Tampa Bay, Florida Gideon E. Nelson 93
Reproduction and ecology of the longnose killifish Robert A. Martin and John H. Finucane 101
An extinct Pleistocene owl from Cuba Pierce Brodkorb 112 The bone-eating dog, Borophagus diversidens Cope Walter W. Dalquest 115
Nesting status of the brown pelican in Florida in 1968 Lovett E. Williams, Jr., and Larry Martin 130
Hippoboscid flies from cattle egrets in central Florida John B. Funderburg, Margaret L. Gilbert, and Ernest L. Bostelman 141
Nitrate and ammonia in rumen of steers fed millet D. T. Buchman, R. L. Shirley, and G. B. Killinger 143
Oyster shell as roughage replacement in cattle diets T. A. Dunn and J. F. Hentges 150
JUL 2 1969 CIBRARIED
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of the FLORIDA ACADEMY OF SCIENCES
Vol. 31 June, 1968 No. 2
Comets, Superstitions, and History
DUANE KOENIG
THe day the Ikeya-Seki “hairpin” comet reached perihelion, October 21, 1965, the Miami Herald ran an article (p. 6A) entitled, “Nobody's Afraid of Comets. ... Anymore.” Broadly speaking that was true. From earliest times comets, meteors, and eclipses have been both of interest and concern to human beings. Of these, comets have attracted the most attention. One astronomy manual asserts, “More excitement has been caused by comets than by any other objects that appear in the sky. Battles have been stopped in mid-career, proclamations have been issued, whole populations have been thrown into panic, kings have abdicated from their thrones, men have died of fear” (Bernhard, Bennett, and Rice, 1948).
Comets have provided subject matter not only for astrologers, astronomers, philosophers, and historians, but also for poets, novel- ists, and playwrights. Mention of these astral visitors occurs in Homer, Vergil, Ovid, Plutarch, Tacitus, Juvenal, and Claudian of antiquity, Rabanus Maurus, Abélard, and Aquinas of the Middle Ages, and Spenser, Tasso, Defoe, Pope, Thomson, Byron, Tennyson, and Tolstoy of the modern period, and elsewhere. The name “comet” has been used for ships, trains, airplanes, cars, newspapers, race horses, taverns, filling stations, hotels, jazz bands, and cleans- ers. Comets have appeared in Christian art as emblems of Christ- mas and in heraldry as figures on escutcheons. The old boy scout “weather” merit badge displayed a comet. The Italians used to sell comete at church doors on saints’ days. These were cakes made with honey and sliced nuts. Americans say, “He rose like a comet,” when referring to someone who has achieved success quickly. Greeks say, “He disappeared like a comet,” when referring to some-
82 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
one who has vanished unexpectedly. Tibetans speak of an event being rare as a comet at noon. There is at least one comet joke. (What is a star with a tail? Mickey Mouse! ).
Though five or more telescope comets may be noted annually, on average fifteen to twenty comets per century are visible to the naked eye. Perhaps four or five are outstanding. A few are bright as or brighter than Venus and a very few such as Ikeya-Seki can even be seen in the daytime. Little wonder then that many super- stitions have arisen about the influence of comets on human affairs, particularly during the many millenniums before invention of the telescope and before enlightened acceptance of comets as natural law-abiding heavenly bodies. Walter Winchell summed up the ob- vious in 1964, “A comet is always more exciting than a fixed star” (Miami Herald, March 30, 1964, p. 5B).
The story of comets has several facets. These mix fact and legend. Almost always in classical times comets were regarded as portents, generally as warnings of dire events, but now and then as harbingers of happy things. Comets, meteors, and stars signal- ized birth of gods, heroes, and prophets. Heavenly lights were vis- ible for Krishna, Abraham, Moses, Lao-tzu, Buddha, Aesculapius, Christ, some of the Caesars, and Mohammed. The medieval John of Damascus appeared to think the Star of the Magi was a comet (Pingre, 1783-1784; Hellman, 1944). Old convictions continued in the Christian era. To these were added patristic pronounce- ments stating comets were threats of an angry God to the wicked, or signs of the Last Judgment to the righteous. Such concepts were held by Christians of the Middle Ages and by both Catholics and Protestants after the Reformation. Incidently, Pliny the Elder ex- plained in his Natural History that “comet” comes from Greek and Latin words for “hair,” because of the resemblance between a comet and a woman’s hair streaming in the wind.
Astronomers have been able to describe comets scientifically in the past three hundred years. With two exceptions popular atti- tudes have become less apprehensive. The notion gained ground for a long period that a comet might hit the earth or sun causing terrestial devastation. Conceivably a comet might bring in its tail cyanogen gas, asphyxiating populations. Nearly ninety years go Jules Verne plotted a collision between earth and comet in a novel called in English, Off on a Comet! Also there was specula-
KOENIG: Comets and Superstitions 83
tion comets were inhabited or that meteors (presumably cometary debris ) first carried life to earth.
Comets were recorded early in both the East and the West. Chinese and Japanese thought they saw likeness between a comet's tail and the bundle of twigs in a broom. The motion of a comet's tail across the constellations was similar to a broom over a floor. Comets were “besom stars.” One in 524 B.C., traveled eastwards towards the Milky Way. A Chinese officer declared, “This is a broom to sweep away the old and give us new. God often makes such signs. The feudal princes will suffer calamities by fire.” Per- sians and Koreans viewed comets as of evil nature and often an- nouncing war with the country in whose direction the tail pointed. Hindus felt they were disruptive of life. South American Indians deemed them objects of terror. Some North American Indians imagined they were spirits of departed chiefs (Encyclopaedia of Religion and Ethics, 1908-1927).
In the West the “blazing star” of 43 B.C. which occurred at the time of the funeral games for the murdered Julius Caesar, was considered a celestial chariot to carry heavenward the shade of the dead statesman. Augustus Caesar held this a good omen and in a temple at Rome instituted comet worship. Comets had marked earlier civil war between Pompey and Caesar, and afterwards, the deaths of Claudius and Nero, and the capture of Jerusalem by Titus. A 1910 satirist depreciating slow appearance of Halley's comet— the most celebrated in all history—wrote,
Where is that fiery ‘dagger in the sky’ That could so thrill the ancients and bamboozle ’em, That once (unless the annal-mongers lie) Spoke far from comfortably to Jerusalem (Punch, or London Charivari, 1910).
The Emperor Vespasian was cautioned about a comet. He con- tended the bearded star did not concern him because he was bald. It threatened his neighbor, the king of the Parthians, who was hairy. Withal Vespasian shortly died (Suetonius). The reasonable explanation for these apparent coincidences is simple. When it is remembered that four hundred comets were listed before the tele- scope and well over a thousand since, it is patent one was always at hand when a portent or memorial was needed. Post hoc, ergo propter hoc.
84 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Aristotle taught three centuries before Christ that comets caused severe winds and hot spells (Aristotle). Pliny emphasized their powers and described comets shaped like javelins, daggers, casks, torches, horses’ manes, and goats’ hair (Pliny the Elder). Men perennially debated whether comets were true celestial bodies, very possibly harmless, or were earthly exhalations, below the moon, and hence dangerous to life. Seneca, a contemporary of Pliny, sug- gested, “Men will some day be able to demonstrate in what region comets have their paths, why their course is so far removed from the other stars, what is their size and constitution” (Pingre, 1783- 1784; Hellman, 1944; Encyclopaedia Britannica, 1946; White, 1955; Sambursky, 1956).
While the Bible does not mention comets directly, Luke XXI:25 reads, “And there shall be signs in the sun, and in the moon, and in the stars; and upon earth distress of nations, with perplexity; the sea and waves roaring.” Comets might be celestial signs. Halley's comet rose in 837 and frightened Charlemagne’s son, Louis the Pious. He summoned an astrologer and asked what the “blazing star’ meant? Misfortune was predicted. Louis tried to avert this by special prayers, fasts, and church building. He died three years later and medieval chroniclers pointed to correlation.
The gravity given to comets may be illustrated. Andrew D. White, American university president, historian, and diplomat of the past century, judged the story of comets a worthy topic for a presidential address at the American Historical Association’s second annual meeting in 1885 (White, 1887). Fear of comets paralyzed self-help, he said, among people in face of untoward happenings, encouraged fanaticism, and strengthened ecclesiastical and political tyranny. He adduced Shakespeare’s familiar remark, “When beg- gars die, there are no comets seen; the heavens themselves blaze forth the death of princes” (Shakespeare).
Halley’s comet became conspicuous in 1066 about the time of Edward the Confessor’s death and William of Normandy’s invasion of England. Matilda, wife of Duke William, and her ladies repre- sented the comet on the Bayeux tapestry, which memorialized the operation (Fig. 1). The comet reappeared in 1145 accompanied by a great earthquake. In 1222 it foreshadowed demise of Philip Augustus. Albertus Magnus once was questioned why comets sig- nified wars and deaths of potentates? He answered, “But the death
Koenic: Comets and Superstitions 85
Oe aogit Ab OUy ne CoN ul
KY Ly,
Yon sK) ry (( Vy
Fig. 1. Halley’s comet in the Bayeux tapestry.
of kings is noticed more because of their fame, since their periods have greater planetary dignity, and so greater significations are re- ferred to them.” The waning Middle Ages brought additional sur- mise. Geoffrey of Meaux believed comets like the one of 1315 produced impure, corrupt, and infected blood. This occasioned melancholy, choler, and inordinate appetite. Henry of Hesse, hav- ing in mind the comet of 1368, denied arrival of a comet was prog- nostication of any future happenings (Pingré, 1783-1784; Humbert, 1948; Chambers, 1910; Thorndike, 1950).
Of all the legends about comets which have crept into literature, the most persistent has to do with Halley’s in 1456. The Ottoman Turks had captured Constantinople from the Byzantines three years previously. They were now advancing up the Danube and be- leaguring the city of Belgrade. Pope Calixtus III was on the papal throne. He sent forces to aid the Christians against the Moslems. Just as the two armies were approaching climax of battle, the comet appeared impressively, its tail 60° long (Ikeya-Seki’s tail was ex- ceptionally long also, fourth greatest on record). Some commenta- tors relate that the pope issued a bull excommunicating the Turks and the comet. Others say he called for prayers for salvation of the faithful, “From the Devil, the Turk, and the Comet, good Lord,
86 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
deliver us.” Abraham Lincoln supposedly credited the pope’s bull against the comet. So did the World Almanach for 1910 (Pingre, 1783-1784; White, 1887; Zwack, 1910; Enciclopedia Cattolica, 1948- 1954). Careful scrutiny of this romance was undertaken by Jesuit Johann Stein. He demonstrated that the myth of the bull and/or the prayer against the comet originated at least a generation after Calixtus IIT (Stein, 1909; Flammarion, 1904).
The surgeon Ambroise Paré was particularly distressed by the comet of 1528. He spoke of it as a bearded star of terrific aspect that alarmed the world and burned the heavens like “a great and gory sword.” Paré went on to report this comet caused some people to fall sick because it was so horrible. Some died of fear. It was of excessive length, the color of blood. His description was Apoca- lyptic. “At summit of it was seen the figure of a bent arm holding in its hand a great sword as if about to strike. At the end of the point were three stars. On both sides of the rays of this comet were seen a number of axes, knives, blood-colored swords, among which were a number of hideous human faces with beards of brist- ling hair” (Howe, 1902). Most comets are silvery grey, but some are accounted yellowish, yellow, or ruddy.
Even as Paré conjured up terrors—it was three hundred years later a Frenchman would dare say of a comet supposed to foretell his own death, “Ah, messieurs, la cométe me fait trop d honneur — two royalties took fright and another did not. Louise of Savoy, mother of King Francis I of France, heard of a comet. She assumed it indicated her end. Though in good health she retired to her bed and in three days died. Mayhap the comet of 1556 determined the Emperor Charles V to finish his round of abdications and retire to a Spanish monastery. British Queen Elizabeth I behaved different- ly. She was staying at Richmond when a comet was sighted. Nerves steady, she ordered curtains and windows opened. She said, “The dyce are throwne’; and she placed her confidence in God (Scott, 1890; Brand, 1900; Elson, 1910).
Little surprise that amulets were worn against comets. It is claimed the comet of 1577 led King Sebastian of Portugal to cross to Africa and his destruction. Cattle plague followed a comet in 1597. Shakespeare (as noted) and Milton entertained antique views. The former began a line in King Henry VI, Part 1, “Comets, importing change of time and states, . . .” and the latter in Paradise
KOENIG: Comets and Superstitions 87
Lost referred to a comet that “from its horrid hair / Shakes pesti- lence and war” (Shakespeare; Milton).
Invention of the telescope in Holland, 1608, and its early em- ployment by Galileo made accurate search of the sky possible. Comets became less vexing. Burton’s Anatomy of Melancholy in 1621 questioned, “whether there be generation and corruption, as some think, by reason of aetheral comets” (Burton, 1859). Comets were supra-lunary. Still 1664’s comet introduced rebellion in Cey- lon and coinage of “comet dollars” in the Germanies. These last intrigue collectors today. A medal of 1680 had inscription, “The star threatens evil things: Only Trust! God will make things turn to good” (Encyclopaedia of Superstitions, Folklore, and the Occult Sciences of the World,. 1903; Chambers, 1910; Webb and Espin, 1911). Obligingly several hens at Rome laid eggs with design of the 1680 comet on the shells. Two Englishmen first offered scien- tific and accurate explanations for comets. Isaac Newton published his famous work on gravitation, Principia Mathematica, 1687, and advised that comets like planets were obedient to universal attrac- tion. They were guided by the same laws. Edmund Halley through careful observation and calculation computed the orbit of the comet of 1682, the one now named after him. He predicted its return in roughly seventy-five years or 1758-1759. The comet came back as Halley promised (albeit he did not live to see it), being discovered in Saxony on Christmas of 1758 (Annual Register, 1759). As for Lexell’s comet, 1773, a Parisian scholar noted, “There were not wanting people who knew how well to play upon the super- stitions of people by turning to their advantage the alarm inspired by the portentous body and selling places in paradise at a very high rate’ (Elson, 1910; White, 1955).
One thing “blazing stars” benefited was wine production. The phrase vin de la cométe in some French-English dictionaries is trans- lated, “wine of 1811.” The grapes gathered in 1811 were unusual in quality and quantity. The port wine was the best tasted in dec- ades. Other years that provided exceptional wines also had worth- while comets, 1826, 1839, 1845, 1852, 1858, 1861, and 1882. Allega- tions were made that the comet of 1811 blinded flies and that chil- dren born under a comet would have a hard life. It was unlucky to start a business venture when a comet was in the heavens. Ru- mors circulated that the Great Lakes and the fjords of Norway had
88 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
been excavated by the comet of 1811 on a previous visit. Later the meteor crater of Arizona was imputed to a small comet (Reddall, 1889; Proctor, 1926; Bernhard, Bennett, and Rice, 1948; Hoyle, I).
The 1811 comet was a striking double-tailed affair perceptible all fall. The credulous linked it with the fortunes of the Emperor Napoleon who bestrode Europe. The astronomer Charles Messier had discovered a comet on August 8, 1769. Bonaparte was born seven days after and always talked about “my star,” a kind of pro- tecting genie. Though cynical about human beings, he paid some heed to the empyrean. Public opinion was divided as to the comet. Did it augur success in the coming Russian campaign or Napoleon’s downfall? In 1815 the Emperor was on St. Helena dictating his memoirs.
Comet hunting proved a delightful pastime in the eighteenth and nineteenth centuries. Women often had comets to their credit. Caroline Herschel, sister of the famous astronomer William Her- schel, beginning in 1786 discovered eight comets. Messier found eighteen. A certain Jean Louis Pons who started his career as doorkeeper of the observatory at Marseilles, turned up thirty-seven betweeen 1801 and 1827. A Nantucket girl, Maria Mitchell, located a comet in 1847. She eventually became professor of astronomy at Vassar (Howe, 1902; Chambers, 1910; Proctor, 1926; Larousse du XX° Siécle, 1928-1932).
The comet of 1832 scared people into selling their goods and crowding churches, but the French astronomer Arago proved to his satisfaction then that comets do not influence weather. Regarding collision, Percival Lowell portrayed a comet as a “bag full of noth- ing.” John Herschel said comets were of such airy nature that a comet, tail and all, could be packed in a portmanteau. Rober Ball agreed, “A rhinoceros in full charge would not fear collision with a comet” (Ball, 1915; Pickering, 1953).
If more evidence was needed, small stars could be seen through the nucleus of a comet of 1903. This was far cry from Moliere who had composed verses in the seventeenth century about a comet shattering the world like smashed glass. There were no noteworthy fin de siécle “blazing stars.” However there was interest generated when the public learned Halley’s was anticipated for 1909 or 1910.
All the old coincidences were racked up by one Edwin Emerson
KoENIG: Comets and Superstitions 89
who authored a catchpenny booklet, Comet Lore: Halley's Comet in History and Astronomy (1910). He was excitable as Pare. “Among all the stars known in astronomy, the periodically returning Comet, now known as Halley’s Comet has the most baleful record.” Emer- son went back to 11 B.C. to discuss the perils attendant on each visit of the comet. He cited the Bible and Church Fathers. When he got down to 1910 he made the comet responsible for panic in Mex- ico and massacre in China. He presaged collision, fire, or poison gas. Conveniently for Emerson’s audience, King Edward VII died in the spring of 1910 ( Proctor, 1926).
The Nation of London challenged such extravagance. “The list of notable events in mundane history of which Halley's comet was witness is not a very convincing one’ (1910). It went on to quip that Parisians were planning comet parties with men in blue eve- ning dress and ladies in gowns the color of the firmament. If the comet were bringing the world to end, the Rue de la Paix would be advertising the latest in ascension robes. Party favors would be miniature Gabriel's horns in gold (Cowell, 1910). Punch spoke of skirt length fashions. “Meanwhile the tail of the comet is said by some observers to be getting smaller. Apparently the comet has now approached sufficiently near to the earth to see that long trains are no longer worn” (1910).
Oldsters like to volunteer reminiscences albeit the most danger- ous man to the historian is he who argues, “I was there.” Sharpers sold pills to ward off cometary poison. A few souls hid in wells, supplied caves with provisions, or stored oxygen bottles. Scientists had grown weary reminding the public the earth had passed safely through the tail of the 1861 comet (Zwack, 1910; Chambers, 1910; Elson, 1910; White, 1955). It could again. The comet was closest May 19-20, 1910. Some recollect a faint haze, probably confusing Halley’s with the earlier more impressive “daylight comet” (Ray- mond A. Lyttelton, oral communication). Curiosity was so general that when an astronomer visited a state prison, he found the inmates had constructed fourteen homemade telescopes. These consisted of rolled newspapers stuck together with breadcrust paste and fitted with spectacle lenses. The prisoners had pointed them through the bars to view the comet ( Brashear, 1925).
Thereafter no comets of distinction rose until 1965 though as- trologers liked to point out the births of both President Roosevelts
90 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
were announced by “blazing stars.” A Sunday columnist occasion- ally managed a few lines with the stale intelligence that Mark Twain was born at the time of Halley’s comet in 1835 and died when it came back. One sour note was sounded, 1955, by the inquisitive Cambridge scientist Fred Hoyle. He squared the circle. “It is an old superstition that the appearance of comets in the sky presages disaster. Perhaps the old superstition was right” (1955).
A few years back readers might ask newspapers whether uniden- tified pips on British radar screens or “phantom” jets over Italy might be ascribed to comets, whether tire blowouts or elevator stallings were comet inspired. Yet journalists mentioned no disquiet over Ikeya-Seki. Astronomers went about their business with tele- scope and camera. They did likewise when the “cigar-shaped” Mitchell-Jones-Gerber comet was sighted in early July, 1967. No one conjectured on either instance comet-brought catastrophe.
Jesuit Horatio Grassi forecast today’s consensus three and a half centuries ago. “Have you seen the comet with its terrifying tail? Behold how with its fearsome beard it is carried sky-high. But no longer need you fear the stellar body with its menacing rays, nor is there harm in those stars which delight us all by their appear- ance’ (Drake and O'Malley, 1960).
ACKNOWLEDGMENTS
The writer wishes to express appreciation to University of Michi- gan Library, University of Miami Library, Memorial University of Newfoundland Library, Florida Atlantic University Library, and New York Public Library.
LITERATURE CITED
ANNUAL REGISTER. 1759. J. Dodsley, London, vol. 2, p. 91.
ARISTOTLE. 1923. Meteorologica. Translated by E. W. Webster, Oxford Uni- versity Press, London, bk. 1, chap. 7, lines 19-20.
BALL, W. VALENTINE (ed.) 1915. Reminiscences of Sir Robert Stawell Ball. Little, Brown, and Co., Boston, 407 pp.
BERNHARD, HusEerRT J., DororHy A. BENNETT, AND Hucu S. Rice. 1959. New handbook of the heavens. New American Library, New York, 272 pp.
BRAND, JoHN. 1900. Observations on popular antiquities. New ed. Chatto and Windus, London, 807 pp.
BRASHEAR, JOHN A. 1925. Autobiography of a man who loved the stars. Houghton Mifflin Co., Boston, 407 pp.
KOENIG: Comets and Superstitions 91
Burton, Rosert. 1859. The anatomy of melancholy. Veazie, Boston, 3 vols.
Carus Puinrus Carcmius SEcuNbus (Pliny). 1938. Natural history. Translated by H. Rackham. W. Heinemann, Ltd., London, bk. 2, chap. 22, lines 89-90; bk. 2, chap. 23, lines 90-92.
CHAMBERS, GEORGE F. 1910. The story of comets. Clarendon Press, Oxford, 272 pp.
CoweELL, P. H. 1910. Halley’s comet. Contemp. Rev. (Horace Marshall and Son), London, vol. 97, pp. 580-588.
DRAKE, STILLMAN, AND C. D. O’MALLEy (translators). 1960. The contro- versy on the comets of 1618: Galileo Galilei, Mario Guidicci, and Jo- hann Kepler. University of Pennsylvania Press, Philadelphia, 380 pp.
Extson, Henry W. 1910. Comets: their origin, nature, and history. Sturgis and Walton, New York, 54 pp.
EMERSON, Epwin. 1910. Comet lore: Halley’s comet in history and astron- omy. Schilling Press, New York, 144 pp.
ENCICLOPEDIA Catrorica. 1948-1954. Ente per lEnciclopedia Cattolica e per il Libro Cattolico, Vatican City, vol. 3, p. 394.
ENCYCLOPAEDIA BriIrANNiIcA. 1949. Encyclopaedia Britannica, Inc., Chicago, vol. 6, pp. 100-104.
ENCYCLOPAEDIA OF RELIGION AND Evruics. 1908-1927. Charles Scribner’s Sons, New York, vol. 12, p. 78.
ENCYCLOPAEDIA OF SUPERSTITIONS, FOLKLORE, AND THE OCCULT SCIENCES OF THE WorLpD. 1903. J. H. Yewdale and Sons Co., Chicago and Milwau- kee, vol. 2, pp. 1001-1002.
FLAMMARION, CAMILLE. 1904. Astronomy for amateurs. ‘Translated by Frances A. Welby. D. Appleton and Co., New York, 345 pp.
Gaius SUETONIUS TRANQUILLUS (Seutonius). 1914. Lives of the Caesars. Translated by J. C. Rolfe. W. Heinemann, Ltd., London, bk. 8, chap. 2online A:
HELLMAN, C. Doris. 1944. The comet of 1577. Columbia University Press, New York, 488 pp.
Howe, HERBERT ALONZO. 1902. A study of the sky. Charles Scribner’s Sons, New York, 340 pp.
Hoye, Frep. 1955. Frontiers of astronomy. Harper and Brothers, New York, 360 pp.
HuMBERT, Pierre. 1948. Histoire des découvertes astronomiques. Revue des Jeunes, Paris, 272 pp.
LAROUSSE DU XX° SiEcLE. 1928-1932. Librairie Larousse, Paris, vol. 4, p. 289; vol. 5, p. 703.
Mitton, JoHN. 1941. The complete poetical works of John Milton. Ed. by Harris Francis Fletcher. Houghton Mifflin Co., Boston, pp. 138-307.
Natron. 1910. The comet. New York Evening Post Co., New York, vol. 90, pp. 504-505.
PICKERING, JAMES SAyRE. 1953. The stars are yours. Revised edition. Mac- millan Co., New York, 298 pp.
PincrE, ALEXANDRE Guy. 1783-1784. Cométographie: ou traité historique et théorique des cométes. L’Imprimerie Royale, Paris, 2 vols.
92 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
Proctor, Mary. 1926. The romance of comets. Harper and Brothers, New York, 210 pp.
PUNCH, OR THE LONDON CHARIVARI. 1910. London, vol. 138, p. 379, 401.
REDDALL, HENRY FREDERIC. 1889. Fact, fancy, and fable: a new handbook. A. C. McClurg and Co., Chicago, 536 pp.
SAMBURSKy, SAMUEL. 1956. The physical world of the Greeks. Translated by Merton Dagut. Routledge and Kegan Paul, London, 255 pp.
Scott, WALTER. 1890. The journal of Sir Walter Scott. Harper and Broth- ers, New York, 2 vols.
SHAKESPEARE, WILLIAM. 1958. The comedies, the histories, the tragedies. Ed. by Peter Alexander. Heritage Press, New York, 3 vols.
STEIN, JOHANN. 1909. Calixte et la cométe de Halley. Tipografia Poliglotta Vaticana, Rome, 40 pp.
THORNDIKE, LYNN. 1923-1958. A history of magic and experimental science.
Columbia University Press, New York, vol. 2, p. 459.
. 1950. Latin treatises on comets between 1238 and 1368. University
of Chicago Press, Chicago, 274 pp.
VERNE, JULES. 1878. Off on a comet! Translated by Edward Roth. Claxton, Remsen, and Haffelinger, Philadelphia, 472 pp.
Wess, THOMAS WILLIAM AND T. E. Espin. 1911. Celestial objects for com- mon telescopes. 5th ed. rev. Longmans, Green and Co., London, 2 vols.
WuitTE, ANDREW D. 1887. A history of the doctrine of comets. Papers Am.
Hist. Assn., G. Putnam’s Sons, New York and London, vol. 2, pp. 5-43.
. 1955. A history of the warfare of science and theology in Christen-
dom. George Braziller, New York, 2 vols. in 1.
WorLp ALMANACH AND ENCYCLOPEDIA FOR 1910. 1909. World Publishing Co., New York, 838 pp.
Zwack, GeorceE M. 1910. The return of Halley’s comet and popular appre- hensions. Dept. of the Interior, Weather Bureau, Manila Central Ob- servatory, Manila, 22 pp.
Department of History, University of Miami, Coral Gables, Florida 33124.
Quart. Jour. Florida Acad. Sci. 31(2) 1968( 1969 )
Amphioxus in Old Tampa Bay, Florida
Gmron E.. NELSON
THE presence of the amphioxus, Branchiostoma caribaeum, in Tampa Bay was documented in 1890 when A. A. Wright collected them near Port Tampa at the mouth of Old Tampa Bay. Wright (1890) reported that specimens were abundant and easily secured. Other workers subsequently collected specimens in the area. Re- cently E. L. Pierce (1965) again affirmed their great abundance by reporting an average of 183 per liter of sand near Gandy Bridge. Boschung and Gunter (1962) and E. L. Pierce (1965) investigated B. caribaeum from other coastal areas of Florida emphasizing the taxonomy and distribution of the organism.
The anatomy and embryology of Branchiostoma have been thoroughly studied, but little is known concerning its ecology. The only detailed ecological study on the group was made by J. E. Webb (1958) and by Webb and Hill (1958) on the African amphioxus Branchiostoma nigeriense. Their publication describes its distribution in a large lagoon, the duration of its embryonic and larval stages, growth rates in the adult, salinity tolerance, and relationship to the substrate. They did not emphasize population characteristics of B. nigeriense.
The purpose of the study described here was to determine some of the basic attributes of amphioxus populations in Old Tampa Bay. The study was designed specifically to determine whether B. caribaeum had a definite spawning time or spawned throughout the year, how the population changed seasonally as reflected by changes in length frequencies and sexual state; whether age classes could be distinguished by the length frequency data; and some idea of the density of the population. The study was conducted from September 1963 through August 1967. The months during which samples were collected are indicated on Figs. 1 and 2.
METHODS
The site of the investigation was Old Tampa Bay, an estuary on the west coast of Florida. This is a shallow body of water, with greatest depth of 16 feet, and about 14 nautical miles long by 7 miles wide. Salinity fluctuates between 18-24 parts per thousand,
94 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
DEC ‘63 N 340 X 46
DEC ‘65 N 227
SEPT ‘63
K 42
SEPT ‘64 N 104 X 41
(op) WY) q i($(|0 SEPT 66
N 115 S 5 X41 Se O < a 10 JAN ‘66
N 198 my ii
FZ 2 KX 47 ¥ eee Py tet = ZZ LJ O io OCT ‘66 10 JAN °67 oC N122 N 206 uJ 5 X44 = xX 51 @= _inl
10 NOV ‘63
N75 5 X 41 00 hd ,
10 NOV °65 10 FEB ‘66 N 344 N 155 2 X 43 5 X44 10 NOV ‘66 10 FEB ‘67 N143 N 118 5 X48 5 KX 50 20 30 40 50 60 20 30 40 50 60
LENGTH CEASS) sIN Mii
NELSON: Amphioxus in Tampa Bay 95
and the area is subject to daily tidal fluctuations. The Bay contains a number of sandy bottom areas suitable for amphioxus, inter- spersed with mud bottom areas and grass flats not inhabited by the animal.
Old Tampa Bay has been severely changed by man’s activities, mainly by dredging and the construction of three roadways across it. In addition the area receives heavy recreational use. Despite this drastic alteration of the environment, amphioxus remains abun- dant in Tampa Bay, even as Wright (1890) and Andrews (1893) reported many decades ago.
A general sampling survey of the Bay area indicated that a sandy strip along the southeast side of Courtney Campbell Park- way maintained a large population of lancelets and was convenient for sampling purposes. This area served as the collecting station for the study. It was usually visited at low tide to facilitate sampling. Approximately five liters of sand were shoveled into a plastic bucket and carried to the beach for screening. Small amounts of the sand were placed on the screen and washed with sea water. Specimens were picked from the screen and placed in gallon jars of water. Two sieving devices were used, a brass testing sieve with 0.98 mm mesh openings, and a home-made sieve con- structed with two layers of plastic screen having the equivalent of 1 mm openings. The latter device was easier to use and re- tained the smaller specimens about as well as did the brass screen. Both were used throughout the study.
Specimens obtained in the manner described above, about twenty per jar, were returned immediately to the University and placed in an air-conditioned laboratory. With no additional atten- tion these specimens survived for three to five weeks in the summer and for several months if collected in winter.
The individuals constituting each sample (100 or more) were measured while alive by placing them in a petri dish resting on a plastic, transparent millimeter ruler. Using a low magnification dissecting microscope, the specimen’s total length (from tip of the
Fig. 1. Monthly histograms for amphioxus samples collected from Septem- ber through February. Open bars represent individuals without gonads (less than 30 mm in length), or else containing empty gonads. Solid bars represent males and females with full gonads. Number of individuals in the sample and the mean length of members comprising the sample are given.
96
IN EACIa) CLASS
on xi 2D Snw NW
PIE IACIEIN Wf
QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
JUNE “65 N 102 X48
10f MAR ‘64 10 JUNE *66 FN 159 N 206 Sf =X 44 = 5 X45 on a4 10 MAR ‘67 10 JUNE ‘67 N 142 N 104 5 X 51 5 X 50 10 APR “64 10 JULY >66 N 246 N213 -) X 46 5 X43 10 PR ‘66
10 APR 67 N 105 5 R51
-~—--- —, fe
10 MAY “64
AUG 24°66 N 102 N 145 5 X 43 5 X 41 10 MAY 66 10 AUG 10 ‘67 E Ni99 3 N 98 oF X45 eel oy,
AUG 28 ‘67 N 88 X% 44
20 30 40 50 60 20 30 40 50 60
L-E NG ll” CAP ANSS SiN ane
NELSON: Amphioxus in Tampa Bay 97
rostrum to tip of the caudal fin) was recorded to the nearest millimeter. At the same time the specimen’s sex and degree of sexual maturity (full, empty, or partially empty gonads) was ob- served and recorded. The use of live specimens is a basic and crucial aspect of the study because the animal is semi-transparent while alive but becomes opaque when preserved. The opacity of the preserved specimens makes sex determination or determination of sexual development difficult. On living specimens these condi- tions are obvious at a glance.
The length frequency data and data concerning the sexual state of the individuals in a sample are summarized on the length- frequency histograms in Figs. 1-2. Conclusions concerning the population characteristics of B. caribaeum were based primarily on these histograms.
’ SPAWNING
As shown by the histogram data, spawning for B. caribaeum in Old Tampa Bay begins in late August. At this time large adults begin to appear with discharged gonads as shown in August 24, 1966, and August 28, 1967 (Fig. 2). This becomes even more evident during September (Fig. 1) and continues into December. In addition, some amount of spawning evidently occurs throughout the year because at least a few small specimens (less than 20 mm in length) are found nearly every month of the year.
Sexual maturity in B. caribaeum occurs at 30 mm in length. Specimens smaller than this often contained developing gonads, but the smallest undoubtedly mature individuals were 30 mm or longer.
Why reproductive activity begins in late summer is not clearly indicated since there are no sharp changes in salinity, temperature, light intensity, or tides at this time to act as a trigger. Carlisle (1951) found that the ascidians Ciona and Phallesia spawned when fed eggs and sperm of their own species. It is tempting to hypothesize this sort of mechanism operating in amphioxus. That
Fig. 2. Monthly histograms for amphioxus samples collected from De- cember through August. Open bars represent individuals without gonads (less than 30 mm in length), or else containing empty gonads. Solid bars represent males and females with full gonads. Number of individuals in the sample and the mean length of members comprising the sample are given. ss ae
98 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
is, as a portion of the population reaches sexual maturity during the summer, they spontaneously release eggs and sperm. These products are unavoidably ingested by other mature or near mature individuals to bring about a large scale or simultaneous spawning such as recorded for September (Fig. 1). Unfortunately, evidence available at this time does not support the hypothesis. As quoted from Bone (1958), “Ripe adults were dissected and the eggs and sperm mixed in Boveri dishes, or sperm was placed in the water containing ripe females and vice versa, but these experi- ments yielded no results.”
From September through December (Fig. 1) the population consists predominately of members which have either discharged their sexual products or are in the process of developing gonads.
After December a steady increase in individuals with full gonads is noted. This build-up continues through the spring and summer until the August spawning.
AGE CLASSES
A close examination of the histograms indicates that three age groups probably make up the population in Tampa Bay. This is especially evident in samples for September 1964, November 1963, December 1966, January 1964, February 1966, May 1964, July 1966 and 1967, and August 1966 and 1967. The following age classes are proposed on this basis:
1) First year class. This group consists of individuals hatched in late summer or early fall. They are evidently planktonic at first and_ start appearing in the sand when they are 10-15 mm in length. Specimens as small as 10 mm could probably wriggle through the openings of the sieve used in the study. However, samples of sand, treated with formalin, were carefully searched without success for specimens smaller than 10 mm. Some members probably attain 30 mm total length and sexual maturity during the year. Members of this age group are especially noticeable on histograms for September, October, and November (Fig. 1).
2) Second year class. This age group averages around 35 mm in the fall as shown on some of the September through December graphs. This portion of the population grows and matures sexually until it averages 40-45 mm in length during the summer, May-August (Fig. 2).
3) Third year class. This group peaks on the graphs around 50-55 mm with a few members reaching 60 mm in length. The mean class length shifts slightly downward in September and November. Presumably this year
NELSON: Amphioxus in Tampa Bay 99
class fluctuates with the removal of older members by death and addition of new members from the second year class. If these interpretations are correct, B. caribaeum does not exceed four years of age in Old Tampa Bay.
It is evident from the histograms that the amphioxus population varied somewhat in its length classes from year to year between 1963 and 1967. Some of the variation may be credited to sampling procedures and some to environmental influences. A severe freeze or the actions of drastic storms like hurricanes (one occurred during the period of the study) are influential events in the life of the shallow water organism. After allowing for this annual variation, there still remains a general picture of spawning during the fall months and maturation of age groups during the remainder of the year.
Densiry DETERMINATIONS
Attempts to obtain an accurate measure of density were not particularly successful. The number of animals per liter of sand varied from 0-15 depending on which portion of the sandbar was sampled. In an area four miles from my collecting site E. L. Pierce (1965) recorded an average of 183 specimens per liter in one series of dredge hauls. Boschung and Gunter (1962) estimate that B. caribaeum in Mississippi Sound seem to be numbered in billions. It is encouraging to encounter an organism that evidently is not threatened by man’s use and misuse of the environment.
SUMMARY
The amphioxus Branchiostoma caribaeum dwells in sandy areas of Old Tampa Bay in considerable numbers. The population spawns during the fall of the year. Sexual maturity is reached at about 30 mm in length. Length frequency data are interpreted as showing three age classes, namely: 1) less than one year old, specimens up to about 30 mm total length; 2) second year class, specimens 30-50 mm long; 3) third year class, 45-60 mm long.
Accurate density determinations were unsuccessful. The Bay appears to support large numbers of Branchiostoma in spite of intense human use of the area.
100 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
LITERATURE CITED
Anprews, E. A. 1893. An undescribed acraniate: Asymmetron lucayanum. Johns Hopkins Univ. Biol. Lab. Stud., vol. 5, no. 4, pp. 213-247.
BONE, QUENTIN. 1958. Observations upon the living larva of amphioxus. Pubbl. Staz. Zool. Napoli, vol. 30, no. 3, pp. 458-471.
BoscuHuNnc, HERBERT V. AND G. GunTeER. 1962. Distribution and variation of Branchiostoma caribaeum in Mississippi Sound. Tulane Stud. Zool., vol. 9, no. 5, pp. 245-257.
CaruisLeE, D. B. 1951. On the hormonal and neural control of the release of gametes in Ascidians. Jour. Exp. Biol., vol. 28, no. 4, pp. 463-472.
Prerce, E. Lowe. 1965. The distribution of lancelets (Amphioxi) along the coasts of Florida. Bull. Mar. Sci., vol. 15, no. 2, pp. 480-494.
Wess, J. E. 1958. The ecology of Lagos Lagoon. III. The life history of Branchiostoma nigeriense Webb. Phil. Trans. Royal Soc. London, series B, vol. 241, no. 683, pp. 335-353.
Wess, J. E. anp M. B. Hut. 1958. The ecology of Lagos Lagoon. IV. On the reactions of Branchiostoma nigeriense Webb to its environment. Phil. Trans. Royal Soc. London, series B, vol. 241, no. 683, pp. 355-391.
WricHt, ALBERT A. 1890. Amphioxus in Tampa Bay. Amer. Nat., vol. 24, p. 1085.
Life Science Building, University of South Florida, Tampa, Flor- ida 33620.
Quart. Jour. Florida Acad. Sci. 31(2) 1968 (1969)
Reproduction and Ecology of the Longnose Killifish
RoBERT A. MARTIN AND JOHN H. FINUCANE
Atmosr nothing is known about the reproductive behavior and early development of the longnose killifish, Fundulus similis (Baird and Girard). Breder and Rosen (1966) omitted the species in their comprehensive survey of literature on reproduction of fishes. Springer and Woodburn (1960) discussed courtship displayed by this fish in captivity. Simpson and Gunter (1956) noted that spawning occurred in the shallows of Copano Bay, Texas, during July but offered little descriptive data; no eggs were recovered after the presumed spawning had taken place. Male breeding colors were described by Joseph‘and Yerger (1956) and later by Springer and Woodburn (1960).
The ecology of F. similis in Tampa Bay, Florida, is incompletely known. Springer and Woodburn (1960) summarized their own and other ecological data. The junior author has contributed addi- tional data on life history from field collections of 1962 to supple- ment knowledge of the local distribution, seasonal occurrence, spawning, growth, and environmental tolerances of the species.
MATERIALS AND METHODS
Fish were preserved in a neutralized 10 per cent formalin solu- tion. Field specimens were measured to the nearest millimeter and aquarium specimens to the nearest 0.1 millimeter of standard and total lengths. Weights were recorded to the nearest 0.01 gram on a triple-beam chemical balance.
A breeding pair of Fundulus similis, a large gravid female (111.6 mm SL) and a mature male (46.3 mm), collected by seine in Boca Ciega Bay near Three Palms Point on March 30, 1967, was placed with other fish in a 30-gallon tank containing filtered sea water (see Table 1). When courtship began, the other occupants of the tank were removed; the parents were also removed after spawning to prevent them from eating the eggs.
An inspection of egg masses on the third day of development indicated that conditions in the original aquarium were not optimal for growth and survival. A number of eggs were then transferred
102 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES
to a 1,000-ml beaker containing dilute sea water (Table 1), where they remained until they hatched or were preserved.
Newly hatched fry were transferred to a 30 gallon aquarium and maintained on a diet of live brine shrimp and dry food (Tetra- marin ) [Reference to trade names in this publication does not imply endorsement of commercial products]. Water quality was similar
TABLE 1
Water quality in laboratory experiments
Laboratory equipment Salinity Temperature Oxygen
(%) (2c) (ppm ) pH Spawning aquarium 34.5 DAL) Sy 8.2 Hatching beaker 19.0 24.0-26.9 5.7-6.2 8.1 Rearing aquarium 20.4-22.8 24 .0-26.9 5.7-6.2 8.1
to that in the beaker. With the exception of salinity, the water conditions resembled those in the original aquarium where spawn- ing occurred (Table 1). Aquariums were supplied with air and contained silica sand, subsand filters, coral rocks, and plastic plants. Ecological studies consisted of monthly and bimonthly collec- tions of fish and water quality data at 18 sampling sites in 1962 throughout Tampa Bay (Fig. 1). All specimens were collected with a 70-ft nylon beach seine having a body and bag of %-inch stretch mesh. A 30-ft minnow seine was used at a few stations where bottom conditions did not permit the use of the larger net. Water