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ATOLL RESEARCH BULLETIN 

NO. 449 

SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: 

AN ANNOTATED BIBLIOGRAPHY 

BY 

T.W. SCHMIDT AND L. PIKULA 

ISSUED BY 

NATIONAI, MUSEUM OF NATURAL HISTORY 

SMITHSONIAN INSTITUTION 

WASHINGTON, D.C., U.S.A. 

OCTOBER 1997

SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: AN 

ANNOTATED BIBLIOGRAPHY 

Thomas W. schmidt1 and Linda pikula2 

ABSTRACT 

Dry Tortugas National Park, located 110 km west of Key West, Florida, is an elliptical, 

atoll-like, coral reef formation, approximately 27 km long and 12 km wide with shallow 

water depths ranging from 12-20 m in channels between reefs. In 1935, the area was 

designated Fort Jefferson National Monument, the World's first underwater National 

Park unit. Central to the area is Fort Jefferson, America's largest coastal nineteenth 

century masonty fort. In 1992 it was re-designated Dly Tortugas National Park. 

Because of the islands' unique location, the first tropical marine biological laboratory in 

the Western Hemisphere was established on Loggerhead Key by the Carnegie Institution 

of Washington, Washington, D. C. Following the closure of the Tortugas Laboratory in 

1939, aperiodic marine biological assessments have been conducted in response to manmade 

and natural environmental perturbations. This annotated bibliography is an 

attempt to provide researchers and resource managers with access to the rapidly 

accumulating body of information on the park's natural resources. A total of 424 

references (published and unpublished) on scientific studies in, (and what later became) 

Dry Tortugas National Park were annotated and indexed according to major scientific 

topics. Studies from a wider area were included if they also sampled in Dry Tortugas 

National Park. 

BACKGROUND 

Seven small islands composed of coral reefs and sand in the eastern Gulf of Mexico, 

approximately 110 km west of Key West, Florida comprise Dly Tortugas National Park 

(Fig.). The Tortugas, an area known for its bird and marine life and shipwrecks, are an 

elliptical, atoll-like, coral reef formation, approximately 27 km long and 12 km wide 

with water depths ranging from 12-20-m in channels between reefs. 

1 South Florida Natural Resources Center, Everglades National Park, 40001 State 

Road 9336, Homestead, Florida 33034, U.S.A. 

2 NOAA Regional Libray, 4301 Rickenhacker Causeway, Miami, Florida 33 149, 

U.S.A. 

Manuscript received 2 September 1997; revised 2 October 1997

The Dly Tortugas, discovered by the Spanish explorer Ponce de Leon in 1513 and named 

The Turtles, Las Tortugas, were soon read on early nautical charts as "Dry Tortugas" to 

indicate they lacked fresh water. Central to the area and located on Garden Key is Fort 

Jefferson, America's largest coastal nineteenth century masonry fort. Work was begun in 

1846 and continued for thirty years but was never finished. As part of the United States 

coastal fortification buildup after the War of 1812, Fort Jefferson was considered critical 

for protecting Gulf trade and ports (Murphy, 1993). 

Following the Fort's use as a military prison during the Civil War (where the infamous 

Dr. Mudd was imprisoned after President Abraham Lincoln's assassination in 1865), and 

its abandonment by the Army in 1874, the area was proclaimed a wildlife refuge in 1908, 

to protect sooty tern rookeries from egg collectors. In 1935, the area was designated Fort 

Jefferson National Monument, the World's first underwater National Park unit. In 1992 it 

was redesigned Dry Tortugas National Park to preserve and protect both historical and 

natural features. 

Early descriptive observers of Dsy Tortugas natural resources include Louis and 

Alexander Agassiz during the 1850's, and the research vessel Blake in 1877 and 1878. 

Their visits resulted in a detailed map of the islands, and a description of benthic marine 

communities by Agassiz in 1888. 

In 1903, Alfred G. Mayer, under the auspices of the Carnegie Institution of Washington, 

recommended that a tropical marine biological research laboratory be established at the 

Tortugas (as opposed to other Caribbean sites) because of their isolation from 

continental land masses, lack of commercial fisheries, lush reefs, clear waters and 

proximity to the Gulf Stream. In 1904, Mayer selected Loggerhead Key as the site for 

Carnegie's Tortugas Marine Laboratory, the first tropical marine laboratory in the 

Western Hemisphere (Fig.2). Following the closure of the Laboratory in 1939, relatively 

few investigations were conducted in the Tortugas until the National Park Service (NPS) 

began in 1975, a series of cooperative, bench-mark studies to evaluate long-term changes 

in marine resources in combination with the earlier Carnegie Laborato~y studies. Since 

the initial Tortugas Reef Atoll Continuing Transect Studies (TRACTS) work of 1975-76, 

aperiodic biological assessments have been conducted in response to man-made and 

natural environmental perturbations. 

PURPOSE 

The primary purpose of this annotated report is to provide researchers and resource 

managers with a readily accessible document on the rapidly accumulating body of 

information on the natural resources of the Dry Tortugas. With the recent 

implementation of the Florida Keys National Marine Sanctuary, adjacent to the Park's 

boundary, there is a dire need for a scientific database that is centrally located, coherently 

organized, and directly related to the future and ongoing management and regulation of 

marine resource activities.

No complete bibliography of the scientific studies on the park's marine and terrestrial 

natural resources has been undertaken. In this report we have attempted to list published 

and unpublished reports from many fields which we feel will be useful as a starting point 

for natural science studies to be conducted at the Dry Tortugas for decades to come(Figs.2&3), 

METHODOLOGY 

The present bibliographic database containing 424 references was compiled using 

PROCITE software, and covers the period of approximately 1878-1996, with the 

exception of one report dated 1820. Most 1996 papers were listed through August. 

Arrangement is alphabetical by senior author and title. Entries are numbered in 

sequence, and each includes a complete bibliographic citation with abstract or summary 

While some attempt has been made to achieve uniformity in style, in many cases the 

terminology, spelling, capitalization, and phraseology of the original author or abstractor 

have been retained. Abstracts obtained from the Carnegie Institution of Washington's 

publication citations were adapted from the author's summaries of results. This 

bibliography includes books, book chapters, scientific articles, theses and dissertations, 

workshop and conference proceedings, reports, and government publications. No attempt 

was made to include articles from newspapers or popular boating or sport magazines 

Several maps and charts are cited, however. 

Research citations were indexed by broad fields of study, specialty sub-headings, and by 

both senior and joint authors. Each citation is listed under as many subject headings as 

is appropriate for the cited article. This cross indexing system was constructed using 

PROCITE. 

The geographic boundaries for citations in this bibliography are that the work was done 

either completely or partially within the Park boundary, which is the 60' contour line 

(Fig.4). Recently, a few studies were undertaken adjacent to or on the Park boundary, 

they were included also. Although all Carnegie Institution of Washington published 

studies were included if they were conducted at the Dry Tortugas, those studies that were 

conducted and identified as solely in the "Gulf Stream" or at satellite marine laboratories 

in Jamaica, Trinidad, Puerto Rico, Bahamas, or in the tropical Pacific were not included 

A broad range of marine and terrestrial topics were found, including vegetation, marine 

algae, invertebrates, sea water composition, and geology. The major topics were subdivided 

into those specialty areas that are shared most often among the studies examined. 

For example, sea water composition was sub-divided into salinity and temperature, the 

parameters measured most often. In many cases, inclusion or exclusion of a given 

reference within a major topic area or specialty sub-heading was a subjective decision.

LITERATURE SEARCHED 

The bibliographic search was predominantly done at South Florida Natural Resources 

Center Library, Everglades National Park, Homestead, Florida, and at the NOAA 

Regional Library, Miami, Florida. No starting date was established for the references in 

this compilation. The senior author began assembling marine archival materials (e.g., 

raw data sheets, correspondence, maps, etc.) and published and unpublished research 

results from major scientific studies conducted by NPS scientists and contractors 

working at the Tortugas. Pertinent record files were also searched at Dry Tortugas 

National Park 

The Park Library contains a complete 35 volume set of the Carnegie Institution of 

Washington's Tortugas Laboratory Papers. Volumes 1-6 were titled "Papers from the 

Tortugas Laboratory of the Carnegie Institution of Washington" (1908-19 14), while 

volumes 7-14 were "Papers from the Department of Marine Biology of the Carnegie 

Institution of Washington" (1915-1926), Volumes 15-29 were issued as "Papers from 

Tortugas laboratory of Carnegie Institution of Washington" (1928-1936) and volumes 

30-35 were titled "Papers from Tortugas Laboratory" (1936-1942). Each volume was 

given a separate publication number by the Carnegie Institution. 

The Year Book Series of the Carnegie Institution, which contains annual summaries by 

individual investigators on observations and results obtained during their visits to the 

Tortugas, were searched at the University of Miami's Richter Library, Coral Gables, 

Florida. The Richter Library contains volumes 1-12, and 20-32. Copies of annual 

investigator reports for volumes 13-19 and 33-39 were obtained from the Carnegie 

Institute of Washington, Washington, D.C. In nearly all cases it was found that the 

principal investigators published their final evaluations and conclusions in "Papers", 

while the Year Book contained mostly duplicative, preliminary, or unsubstantiated 

observations. For these reasons, and due to time constraints, we decided to cite only 

Year Book contributions for investigators who did not complete and publish their 

conclusions in the "Papers" series. For example, W. H. Longley published in both 

series, but is only cited in this bibliography under "Papers" (however, his Year Book 

citations can be found in the "literature cited" section of his contributions to "Papers"). 

S. Yamanouchi, however, only published in the Year Book, and is cited here as such. 

We searched documents regarding the Tortugas Laboratory during two visits to the 

Carnegie Institution in Washington D.C. Work conducted at the Tortugas Laboratory 

has been published in a wide range of journals. For example, a list of scientific writings 

produced by activities at the Laboratory during Mayer's directorship can be found in 

Papers Tortugas Laboratory 19:80-90. Many publications continued to appear in the 

literature following the closure of the Laboratory in 1939. 

On-line database searches were conducted during 1993-96 at the NOAA Miami Regional 

Library. Subject index terms such as coral reef, geology, vegetation, marine algae, fish,

etc., were used to search on a variety of DIALOG electronic databases including the 

following: BIOSIS PREVIEWS, Dissertation Abstracts, Oceanic Abstracts, 

SCISEARCH, Ei Compendex, INSPEC, and GEOBASE. These individual CD's 

were also searched: Aquatic Sciences and Fisheries Abstracts, Life Sciences Collection, 

Earth Sciences, GeoRef, NTIS (National Technical Information Service), GPO 

(Government Publications Office), and the OCLC (Online Computer Library Center) online 

catalog. 

Pertinent theses, dissertations, and journals identified in the abstracted literature were 

obtained via interlibrary loans. 

This current compilation undoubtedly does not list all available literature that might be 

useful in conducting research, monitoring, and resource management of the park's natural 

resources. There may be as many as 100 additional scientific papers generated from the 

Carnegie era. We would greatly appreciate additional references to the Tortugas 

literature, and if a sufficient number of additional articles become available, we will 

produce an addendum to this report. 

ACKNOWLEDGMENTS 

This report has benefited from the help of many people over the past 6 years. The 

original project was prompted and supported by Dr. Michael Soukup during his tenure as 

Director of the South Florida Natural Resources Center. Wayne Landrum, Facility 

Manager and Carolyn Brown-Wiley, Chief Ranger at Dry Tortugas National Park 

provided logistical support and took special interest in providing guidance to the 

pertinent files at Fort Jefferson. Ray Bowers, John Strom, and Pat Craig of the Carnegie 

Institution in Washington, D.C. permitted us to search their Tortugas Laboratory files, 

assisted in duplicating activities, and provided insightful discussions and original 

photographs of the Marine Laboratory. 

We thank George Stepney and Maria Bello of NOAA's Regional Library in Miami for 

acquiring many interlibrary loans. Special recognition goes to the staff of the South 

Florida Natural Resources Center including Marnie Lounsbury for photocopying and 

collating much of the Carnegie texts, Barry Wood who produced the map figures, and 

Mario Alvarado who expertly produced the author and subject indexes using PROCITE. 

Dr. William B. Robertson, Jr., United States Geological Survey/Biological Resources 

Division contributed numerous references and provided encouragement during the 

earliest stages of the project 

Valuable comments were provided by Elaine Collins of the NOAA Central Library, 

Silver Spring, MD and Bob Hamre, former technical editor for the US Forest Service, 

assigned to the Beard Center under the NPS "Volunteer-in-Parks." program.

Finally, our thanks to Carol Watts, Chief of the NOAA Libraries and Information 

Science Division, Janice Beattie, Chief of NOAA Libraries Public Services Division, Dr. 

Tom Armentano, Chief of Biological Resources, South Florida Natural Resources 

Center, and Dr. Caroline Rogers, United States Geological Survey/Biological Resources 

Division for their financial support and encouragement.

ANNOTATED BIBLIOGRAPHY 

1. Agassiz, A.. 1888. The Florida reefs. Three cruises ofthe United States Coasts and Geodetic 

Steamer 'Rlake' in rhe Gulf of Mexico, in the Caribbean Sea ar~d along the coast of the 

United States,froln 1877 to 1880., V. I, Chapter 3, pages 52-92. Houghton, Mifflin and 

Co. New York. 314 pp. 

While the Steamer "Blake" was n~ostly involved in deep water dredging operations in the 

Gulf of Mexico and the Caribbean Sea (1877-78) a five-week visit to Fort Jefferson (Dry 

Tortugas) provided the author with an opportunity to work in a laboratory-like situation, to 

examine carefully the topography of the different groups of corals characteristic of the 

Florida reefs and to give an extended account of the Florida reefs in a special chapter of 

this book. The Tortugas, as described by Agassiz, form the most recent of the cluster of 

the Florida reefs, and it is here where he begins a topographical sketch of the Florida reefs 

from the Tortugas to Cape Florida. 

2. -- 1888. The Tortugas and Florida Reefs. Memoirs of the America11 Academy 0fArt.s and 

Sciences, Philndelphia. Centennial ed., V.11, : 107- 132. 

V.11 entitled "Explorations of the surface fauna of the Gulf Stream, under the auspices of 

the United States Coast Survey". 

Agassiz reports on the formation of the Florida Reefs, commenting on the theories of 

Darwin, LeConte, and Hunt on this subject. At the time of this article it was believed that 

the elevation of the Florida Plateau, from Cape Florida southward to the Dry Tortugas and 

the Yucatan Banks, was based on the accu~nulation of coral sands, as well as animal debris 

brought to it by great Atlantic equatorial oceanic currents, the Gulf Stream and prevailitlg 

winds, and at which time, reef-building corals could flourish and a reef would be formed. 

They speculated that corals could not thrive below 6 or 7 fathoms, because siltation ooze 

would sink to the bottom and choke the corals. Coral reef formations were assumed to be 

established near strong equatorial currents which were suppliers of food for the reefbuilding 

corals. It was assumed that corals grow towards the surface as fast as the ooze 

deposited has closed up the circulation of the lower levels. At the time of Darwin little was 

known of limestone deposits formed by the accuoiulation of animal decay. Thus 

explanations of reef formation other than elevations as a result of submerged mountains 

and subsidence were not investigated. The Tortugas Reefs probably are newly developed, 

as they have not been above the sea long enough to have received the flora and fauna 

characteristic of the Keys north of Key West. 

3. Andres, B. A. 1991. Migration of sharp-shinned hawks in the Dry Tortugas, Florida USA. Wilsort 

Br~llerin 103, no. 3: 491-93. 

Some species of hawks have been found to make long water crossings during migration. 

One of the species, the sharp-shinned hawk Accipiter striatus, seldom undertakes water 

crossings of >I25 km. However, large numbers of sharp-shinned hawks are observed 

every fall in the Florida Keys including the Dry Tortugas, where water crossings are quite 

common. No information, however has been gathered concerning their migration after 

reaching the Tortugas. Based on wind speed, wind direction, and binocular observations 

made on Garden Key of six species of hawks (sharp-shinned hawks dominated the 

observations), it was found that sharp-shinned hawks are deliberately initiating an overflight 

across the Gulf of Mexico directly to Central America.

4. Austin, 0. L. Jr., W. B. Robertson Jr. and G. E. Woolfenden. 1972. Mass hatching failure in Dry 

Tortugas sooty terns, Sterna fuscata. Pruceedings of the h~ternutional Ornithological 

Congress. 627. Netherlands. 

The author attributes a mass hatching failure among 50,000 pairs of sooty terns (Sterna 

firscara) nesting on the Dry Tortugas to damage caused by sonic booms from low-flying 

military aircraft. Theoretically, eggshells and embryonic tissues should withstand pressures 

much greater than those generated by even the most intense sonic booms. 

5. Bailey, E., C. E. Woolfenden and W. B. Robertson Jr. 1987. Abrasion and loss of bands from Dry 

Tortugas sooty terns. Journal of Field Ornithology 58, no. 4: 413-24. 

During the past 25 years more than 400,000 sooty terns (Sterna,fuscata) have been banded 

at Dry Tortugas, Florida, with size 3 aluminum bands of several different alloys. Based on 

large samples of bands removed from the terns, regression lines were established for each 

of four alloys. Differences in the slopes of the regression lines for certain of the four alloys 

demonstrated differences in rates of abrasion. Band loss was evident for bands of the 

fastest abrading alloy (2-SO) that were carried more than 20 yrs by terns banded as chicks 

because all band weights fell above an extension of the regression line. For this alloy, the 

plots of weight loss showed that band loss becomes significant at 86% of original weight. 

Bands of alloy 2-SO began reaching 86% of original weight at age 14 when placed on 

adults and 20 when placed on chicks. The regression lines for the other 3 alloys suggest 

that loss is likely after 17-28 yrs for bands placed on adults and after 20-25 years for bands 

placed on chicks. Band loss probably occurs through abrasion of the inner surface, which 

increases the inner diameter until the band can slip over the toes. Any gap that develops 

would hasten loss. 

6. Bailey, P. L. 1938. Regeneration in sabellids. Carnegie instirution of Washi~~gton, Year Book 37: 

84-85. 

Sabellid worms collected in the moat on Garden Key proved to be suitable for fixation 

techniques needed to conduct various regeneration experiments to determine the effects of 

chemical solutions on the cells. 

7. Baker, B. 1994. Partitioning the National Marine Sanctuary. Bioscience 44, no. 7: 497. 

A management proposal is described to establish five zones in the Florida Keys National 

Marine Sanctuary: replenishment reserves, sanctuary preserve areas, rcsearcli-only zones, 

wildlife management zones, and special-use zones. The Sanctuary encircles the Florida 

Keys, including the Dry Tortugas, for 2800 square nautical miles. 

8. Ball, S. C. 1918. Migration of insects to Rebecca Shoal Light-Station and the Tortugas Islands, with 

special reference to mosquitoes and flies. Papers Tortugas Laboratory 12: 193-212. 

Carnegie Institution of Washington Publication Number 252. 

The circumstances which suggested the desirability of such investigations were the 

repeated experiences of Dr. Mayer and other scientists at Tortugas, Florida, in connection 

with the occurrence there of mosquitoes. These insects were abundant on Loggerhead Key 

only after northerly winds of several hours' duration, under conditions favorable to their 

migration from the mainland of Florida. Rebecca Shoal light-station was chosen as the 

study site, because of its isolation from the mainland and other keys, and because of its 

freedom from all except easily controllable breeding-places for mosquitoes. It was found 

that large numbers of mosquitoes and house-flies are carried by northerly and southerly 

winds to Rebecca Shoal light-station and the Tortugas Islands from Florida and Cuba. 

Easterly winds bring a few of these, as well as smaller numbers of blow-flies, horse-flies, 

and gnats from islands east on the Florida Reef. Occasionally Odonata, Neuroptera, and 

Lepidoptera are carried by the winds to these parts of the reef. Sarcophagidae breed in land 

crabs at Tortugas .

9. Ballantine, D. L. 1996. New records of benthic marine algae from Florida. Gulfof Mexico Science 

1: 11-15. 

Seven species of benthic marine algae are newly reported from the Dry Tortugas, Florida. 

These are Halirnedu hurnrnii Ballantine (Chlorophyta), Audouit~ella ophioglos.sa 

Schneider, Botr-yocladia uymei Ballantine, Cizarnpia vicllardii Kutzing, Mor~osporus 

irzdicus Borgesen, Hypoglossu~n rhizo~~hornm Ballantine et Wynne, and Rhodogorgort 

ramosissirna Norris et Bucher (Rhodophyta). Mor~osporus irldicus is reported for the first 

time from the Atlantic Ocean. The Dry Tortugas represents the northern distributional 

limit for the remaining species reported, except Audouirlella ophioglossa and Botryocladia 

uytzr~ei. 

10. Bartsch, P. 1919. "The bird rookeries of the Tortugas." Stnithsonian lnstiruriorl Ar~rzual Reporf for 

1917,2512. Smithsonian Museum. 

The author states that the most interesting island of the Tortugas group is Bird Key (circa 

1908). Of the 32,810 birds listed for the islands, 31,200 center about that Key. A 

numerical listing of the summer birds is given. These rookeries were first brought to the 

attention of ornithologists by John Audubon in his ornithological biographies. He gives an 

account of a visit in May 1832. A first list of birds observed in the Dry Tortugas is given 

by W.E.D. Scott in his paper on birds observed during parts of March and April 1890. 

Drs. John B. Watson and K.S. Lashley of Johns Hopkins University made an extensive 

study of the wild bird colonies there, hoping to throw light on the homing instinct. The 

article ends with an extensive listing of bird sightings in the Tortugas up to 1919. 

11. -- . 1920. Experiments in the breeding of Cerions. Papers Torrugas isrboraror}~ 14: 1-55. 

Carnegie Institution of Washington publication Number 282. 

Breeding experiments were conducted to determine if various forms of Cerior~ colonies 

were fixed forms, that is, will generations yield the same mode in measurement, or will 

changes in the local environment from season to season affect the developing organisms to 

such an extent as to produce an unending serics of slight variations? Introduced forms 

were placed where native species existed. Colonies of these land snails were planted on 

Keys in the Dry Tortugas in 1914, 800 in 1915 on Loggerhead Key, and a third planting in 

1916 of 8,317 specimens. A comparative anatomical discussion of the five species of 

Cerion involved in the breeding experiments is given. 

12. ----. 1915. Report on the Bahalna Cerions planted on the Florida Keys. Papers Torrugas 

Laboraror)' 8: 203-12. 


A study is made of the two races of Bahama Ccrior~ transplanted to the Florida Keys in 

1912. The conditions of the Ceriorl colonies are described.. Illustrations show the extent of 

the changes between the first generation and the parent generation. Changes that have taken 

place in the second generation in shell, color and sculpture are discussed. 

13. ---- , 1919. Results in Ceriorz breeding. Proceedings of rhe Biological Society of Waskbzjiton, 32 

Jourr~al Washir~gron Academy of Scier~cc 9:657 (abstr.) 

A short account is given of Dr. Paul Bartsch's report on the breeding of Cerions 

transplanted from Andros Island in the Bahamas to the Dry Tortugas. 

14. ---- 1916. Visit to the Ceriorz colonies in Florida. Smitksonian Explorarions 66, no. 17: 41-44 

The author visited the Dry Tortugas through the auspices of the Carnegie Institution and 

the U.S. National Museum to observe the transplanted Bahamian Cerion colonies. He 

reported finding many adult specimens of the first Florida generation. No adult second 

generation specimens were found. Four hybrid specimens between the native Cerion 

incanum and the transplanted Bahama stock were obtained.

15. Bellow, T. and C. Winegarner. 1975. Nesting of brown pelicans Pe1icanu.r occidermlis on the Dry 

Tortugas, Florida. Florida Field Naturalisr 3, no. 2: 47-48. 

On 14 June 1974 on Bush Key, Dry Tortugas, Florida T. Bellow and C. Winegarner found 

5 Brown Pelican nests about 12 feet above ground in the white mangroves (Lagw~cularia 

racernosa) along the north shore. Nineteenth-century records of pelicans breeding on the 

Dry Tortugas are ambiguous. It appears that a few pairs did breed on the Tortugas in the 

mid-1800's. but by late in the century none did so. This record is the first reported nesting 

of this species in the 20th century on these ornithologically well-known islands. Three of 

the nests found in 1974 contained 2 eggs each, one nest was empty, and the fifth was not 

checked. 

16. Bellow, T. H. 1979. A cardinal at the Dry Tortugas, Florida. Florida FieldNafuralisf 7, no. 2: 31. 

The southern range of the nonmigratory cardinal (Cardinalis cardinalis) extends through 

the Florida Keys, but is considered rare in Key West. An April observation of a cardinal 

on Garden Key represents the second published record of this species at the Tortugas. 

17. Bennett, F. M. Commander. 1909. A tragedy of migration. Bird-Lore 11: 110-113. 

On, April 14, 1909 a violent storm bit the Florida Keys, including the Dry Tortugas. An 

apparent bird migration was in progress at the time of the storm. On April 20th the author 

went to the Dry Tortugas and observed hundreds of dead birds, and tens of thousands of 

injured and exhausted birds. A listing of the types of birds observed is given. 

18. Berrill, N. J. 1938. Budding in polystryelid ascidians. Carrqie lr~stirrrrior~ of Washington, Year 

Book 37: 85. 

The area of the budding rudiment relative to the size of the parent zooid was closely related 

to the size of the parent zooid, and the general nature of the colony. 

19. Blinks, L. R. 1926-1929. Electrical conductivity in Vulonia. Carnegie Institution of Washington, 

Year Book. 

Note published as follows: 1926, v. 25, p. 240; 1927, v. 26, p. 217-18; 1928, v. 27, p. 270- 

71; 1929, v. 28, p. 280. 

This study makes use of the good supply of Vulonia at Tortugas for studies on the 

variability of electrical resistance in protoplasnl. Causes of uncertainty are discussed. 

20. Bohnsack, J. A,, D. E. Harper and D. B. McClellan. 1994. Fisheries trends from Monroe County, 

Florida. Bulletin ofMari~ie Science 54, no. 3: 982-101 8. 

Fishing is an important activity in the Florida Keys National Marine Sanctuary (FKNMS). 

Concern exists that excessive fishing could be deleterious to individual species, disrupt 

marine ecosystems, and damage the overall economy of the Florida Keys. We examined 

data from commercial, recreational, and marine life fisheries in Monroe County, Florida. 

Invertebrates comprised the majority of commercial landings. In 1992, the total reported 

commercial landings were composed of 54% invertebrates (4.09 x I0 kg) 28% reef fishes 

(2.19 x 10 kg), and 21 % non-reef fishes (1162 x 10 kg). In the recreational headboat 

fishery, reef fishes accounted for 92% of 0.107 x 10 kg average total annual landings from 

the Dry Tortugas and 86% of 0.201 x 10 kg landed from the Florida Keys since 1981. 

Average annual landings for other recreational fisheries were estimated at 1.79 x 10 kg for 

reef fishes (45%) and 2.17 x 10 kg for non-reef fishes (55%) from 1980 through 1992. 

Estimated landings from the Dry Tortugas did not show distinct trends and were highly 

variable. Finer resolution of catch and effort data are needed, especially for recreational 

fisheries. Landings for some species varied greatly over time. The most conspicuous 

declines were for pink shrimp, combined grouper, and king mackerel, while the most 

conspicuous increases were for amherjack, stone crab, blue crab, and yellowtail snapper. 

Landings of spiny lobster have remained constant. Fisheries closed to harvest included

queen conch, Nassau grouper, jewfish, and stony corals. Effective fishing effort has 

increased over time with more participants and more effective fishing technology. Since 

1965, the number of registered private recreational vessels has increased over six times, 

while the number of commercial and headboat vessels has remained stable. The number of 

management actions have continually increased and become more restrictive with increased 

fishing effort. Comparison of fisheries was complicated because different fisheries 

targeted different species and different sized organisms. Also, landings were sometimes 

reported by numbers and sometimes by weight. Measures of reproductive value and 

spawning potential are suggested as useful parameters for comparing effects of different 

fisheries. The new FKNMS provides a unique opportunity to shift management emphasis 

from a species approach to an ecosystem and habitat based approach. 

21. Bortone, S. A,, P. Rebenack and D. M. Siegel. 1981. A comparative study of Diplectrum formosum 

and D, bittatum (Pisces: Serranidae). Florida Scienrist 44, no. 2: 97-103. 

Specimens of the simultaneously hermaphroditic fish species Diplectrum formosum, the 

sandperch, and D, bivittatum , the dwarf sandperch, were collected near the Dry Tortugas, 

Florida, by means of shrimp trawl during December 1976. Stomach contents of 326 D. 

for-mosurn (100 empty) and 325 D. bivittaturn (131 empty) revealed little or no differences 

in their food habits relative to number and volume of food items, size of food items or the 

contribution, in grams each food item makes to each fish. Both species primarily 

consumed amphipods, shrimp, crabs, fish, and polychaetes. Temporally, both species fed 

at the same 2 diurnal periods. Species were collected sympatrically hut there were areas 

where each species dominated in relative abundance. 

22. Boschma, H. 1929. On the postlarval development of the Coral Maeandra aerolata (L.). Papers 

Tortugas Laboratory 26: 129-47. 


During six weeks in July and August 1925, the author studied Maeandra areolata for 

researches on the food of reef-corals at the Carnegie Laboratory in the Tortugas. Many of 

the colonies contained ripe larvae and the author reared these for the study of their 

development. The author concludes the development of the endotentacles which appear 

constantly in two successive groups of three, resembles in some way the facts recorded by 

de Lacase Duthiers (1872) in Actinia mesembryanthemum, some stages of which show a 

marked tri-radial arrangement of the tentacles. The data in the literature on the 

development of other coral polyps seem to prove that this successive development of the 

endotentacles in two groups is an exceptional case. The bilateral arrangement of the septa 

in the oldest stages is in accordance with that found by Duerden (1904) in Siderastrea. As 

in the majority of corals in which the young stages are known, the septa in Maeandra 

develop in two cycles, first the six endosepta and soon afterward the six exosepta. 

23. Bowles, A. E., F. T. Awbrey and J. R. Jehl. 1991. Effects of high-amplitude impulsive noise on 

hatching success: a reanalysis of the sooty tern incident, HSD-TR-91-0006. BBN 

Laboratories, Inc., Canoga Park, California. 

This article attempts to refute the Austin article which attributed a mass hatching failure 

among 50,000 pairs of sooty terns (Sterna fuscata), who had nested on the Dry Tortugas to 

sonic boom damage from military aircraft. Theoretically, eggshells and embryonic tissues 

should withstand pressures much greater than those generated by even the most intense 

sonic booms. An experiment was conducted to test whether impulsive noise could be 

responsible for the hatching failure. Four pest control devices were exploded near chick 

eggs in various states of development: 20 chicken and 20 quail eggs. The mean peak flat 

sound pressure level 177.3 db re 20 upa; mean CSEL of 139; mean frequency 620 Hz. No 

cracking damage similar to that of the Dry Tortugas eggs occurred. Hatch rates and 

weights between control and exposed embryos were not significantly different.

24. Bowman, H. H. M. 1918. Botanical ecology of the Dry Tortugas. Papers Tortugas Laboratory 12: 

109-38. 


As the name of these islands indicates, their vegetation is characteristically xerophytic, 

although the rainfall is sufficient to assure the plants tile necessary amount of water. The 

plants are very interesting when a close study is made of their individual characteristics. 

The opportunity for such study was given the writer during the summers of 1915 and 1916, 

while pursuing another line of botanical research at Loggerhead Key, where a marine 

laboratory is maintained. The Tortugas are really the westernmost of all the Florida Keys, 

but are more detached from them and have different geological and botanical aspects. 

Species distributional maps were created for each of the eight islands of the Tortugas Atoll. 

In this treatment of the species in the Tortugas it has been aimed to give some idea of the 

character of the dry-climate plants inhabiting these islands, their distribution, and 

particularly the changes which have occurred on the various keys since Lansing's 1904 

survey, with an attempt to analyze the reasons for such changes. Notes on the marine 

ecology of the Tortugas also are presented, along with descriptions of dominant 

submergent vegetation. 

25. Boyden, A. 1934-1939. Serological study of the relationships of some common invertebrata. 

Carnegie lnstitutior~ of Washb~grotr, Year Book. 

Note: published as follows: 1934, v. 33, p. 248-49; 1936, v. 35, p. 82; 1939, v. 38, p. 218. 

Preliminary results obtained through the study of the antigens collected from various major 

groups of animals at Tortugas were summarized. Blood relationships within Moilusca and 

Crustacea were emphasized. 

26. Bradbury, R. C. 1992. First Florida record of variegated flycatcher E~~tpido~~omus-varius at Garden 

Key, Dry Tortugas. Florida Field Naturalist 20. no. 2: 42-44. 

The variegated flycatcher occurs throughout most of Soutli America east of the Andes. The 

species migrates northward between September and February after breeding in the middle 

and southern part of the continent. It winters in the Guianas, northern Brazil, Venezuela, 

Colombia, and eastern Peru. This article describes observations of a variegated flycatcher 

in Florida, representing the first record in Florida and tlie third in the United States. 

27. Breder, C. M. Jr. 1934. On the habitats and development of certain Atlantic Synentognathi. Papers 

Tortugas Laboratory 28: 1-35 (issued Dec. 1932). 


In this paper data presented are intended to form a basis for further inquiry into the 

comparative development and life habits of the Synentognathi, which includes the familiar 

Belonidae (needlefish), Hernirarnphidae (halfbeaks), and Exocoetidae (flying fish). The 

data on which the present paper are based represent some field studies and laboratory work 

on material gathered in the Dry Tortugas, Florida during May and June 1929 The feeding 

habits, leaping, and flight during the presence and absence of light, eye specialization, 

enemies, and ontogeny and phylogeny are discussed. A new species, Strongylura lo~~gleyi 

is described. A key to the Tortugas Synentognathi is provided, along with tables, beak 

measurement and eye development. The Exocoetidae form the major item of diet of a 

variety of sea birds, about the Tortugas at least. Young Exocoetidae pass the most 

dangerous part of their day when the sun is low, at which time they are unable to see their 

predacious enemies coming from below because of light conditions. The eyes of Belonidae 

are provided with elaborate equipment to protect them from the brilliance of their 

environment. 

28. Breder, C. M. Jr. and J. E. Harris. 1936. Effect of light on orientation and stability of young 

plectognath fish. Papers Torrugas Laborator)' 29: 23-36 (issued Nov. 1935).


Under certain circulnstances some plectognath species will I-espond to a strong beam of 

light by violent gyrations. This was first observed by Breder (1929) at the Tortugas 

Laboratory. The niechanism by which these movements are effected, their relationship to 

the intensity and duration of the stimulus, and the disappearance of the phenomenon with 

advancing age of the animal give rise to a number of interesting problems concerning the 

action of the receptor-effector system in these fish. This paper is an attempt to explain this 

feature of fish behavior. It was found that small specimens of Moitocaitthus and 

Lacrophr-ys, if exposed to a beam of light, frequently exhibit somersaulting or rotational 

movements of great rapidity. Somersaulting is produced by passing the locomotor waves 

in opposite direction along the dorsal and anal fins. The rotational movement is 

accompanied by the deflection of the dorsal and anal fins to the opposite sides of the body, 

the direction of motion of the undulations being usually antero-posterior in both fins. 

Various combinations of fin and tail movements may occasionally give other twisting 

gyratory movements. The primary response to light is always an attempt at reorientation of 

the animal so that the light is incident upon the dorsal surface. In fish kept in complete 

darkness and "sensitized" by repeated stimuli, gyrations may continue after the light has 

been removed, and even mechanical stimuli may initiate similar paroxysms, the 

equilibrating system apparently being more or less permanently deranged. Specimens of 

Monocanthus over 50 mm. in length do not usually display this behavior, and species other 

than plectognaths show it very feebly or not at all. The integration of gravitational stimuli 

into the behavior pattern is apparently not perfectly attained until a comparatively late stage 

in development, and light is the primary orienting factor. The gyrations are apparently due 

to instability, consequent upon overcorrection. 

29. Brinley, F. J. 1937.1938. Studies on the implantation of embryonic fish tissue, with notes on the 

spawning habits and development of four species of fish. Carnegic Irutitution of 

Washingtorz, Yeur Book. 

Note: published as follows: 1937, v. 36, p. 86; 1938, v. 37, p. 86-7. 

Livers and spleens were transplanted from embryos of the hard head shiner to other 

embryos of the same age. No apparent effect on the host was noticed. Eggs of 

Por~~acentrus and parrot fish were collected for observation, along with nurse shark 

embryos. Additional work was performed on the origin of muscular movement in these 

species. 

30. Brooks, H. K. 1962. Reefs and bioclastic sediments of the Dry Tortugas (abs.). Geological Society 

ofAmerica. Special Papcr 73: 1-2. 

Many miscomprehensions exist relative to origin of the Florida Reef track and, in 

particular, its southwatern extremity-the banks, shoals, and reefs known as the Dry 

Tortugas. They are not an atoll as stated by Vaughan (1914). The component 

physiographic features rise from a shallow limestone platform 80 to 100 feet below sea 

level. Relief features are banks and shoals of bioclastic sands. Their genesis and 

circulation distribution are related to the prevailing seasonal storm patterns. Large patches 

of Acropora cervicor~trrs (Lamark) are widely distributed through the area in water less 

than 60 feet deep. Live coral on these patches is sparse. Proliferation of the staghorn 

corals is slow, but cumulative growth has produced a magnitude of skeletal remains. The 

coralla are preserved and are ultimately indurated into a porous rocky mass by the luxuriant 

growth of Lirhoharrtnioit and its cognate encrusting associates. The shallow reefs of 

Garden and Loggerhead Keys, populated by calcareous algae, alcyonarians, and 

scleractinians, etc., originate upon a foundation of the remains of these organisms. This 

can be seen where erosion in surge channels has exposed the underiying materials.

31. Brooks, W. K. 1908. Salpafloridian (Apstein) Part I1 in the Pelagic Tunicata of the Gulf Stream. 

Papers Tortugas Laborator-y 1: 75-89. 


This rare Sulpa about which little is known, has been noted in this paper. Mature 

specimens of both stages of Salpa were found, in May 1906, on the surface in the vicinity 

of the Marine Biological Laboratory at Tortugas, Florida; and an opportunity was afforded 

to study and sketch them while alive, and thus to make additions to, and some slightcorrections 

of, the count of the species.. 

32. Brooks, W. K and C. Kellner. 1908. On Oikopleura tortugensis, n.sp. a new appendicularian from 

the Tortugas, with notes on its embryology in Part IV, The Pelagic Tunicata of the Gulf 

Stream. Papers Tortugas Laborator)~ 1: 73-95. 


This species was found in abundance near the Marine Laboratory. The specimens are from 

5 to 8 mm. long and occur in great swarms at the depth of 5 to 6 fathoms. A description of 

the species is provided. 

33. Brown, D. E. S. 1935. Cellular reactions to high hydrostatic pressures. Carnegie institution of 

Washington, Year Book 34: 76-77. 

Physiological studies were carried out on the muscles of crabs and fish collected in deep 

(100 fathoms) and shallow water of the Tortugas. 

34. Brown, W. Y. and W. B. Robertson Jr. 1975. Longevity of the brown noddy. Bird-Banding 46, no. 

3: 250-251. 

Despite its abundance and pantropical range, little published information exists on the 

longevity of the brown noddy (Anous stolidus). Woodward (Atoll Research Bull, 164: 

280,1972) reported a maximum known survival of I0 years for brown noddies banded as 

adults on Kure Atoll , Hawaii. Brown noddies on Manana Island, Oahu, Hawaii (A. s. 

pileatus) and the Dry Tortugas, Florida (A. s. stolidus), are among the few populations that 

have been banded over a period long enough to provide quantitative data on longevity. 

Twelve of the brown noddies banded on Manana before 1948 were recaptured dead or 

alive before 1960, the longest interval from banding to recapture being 13 years. On 23 

May 1972 Brown recaptured on Manana a brown noddy that had been banded there as a 

juvenile on 12 June 1947, 25 years earlier. 

35. Bullington, W. E. 1940. Some ciliates from Tortugas. Papers Tortugas Laboratory 32: 179-221 

(issued Sept. 1940). 


During the summers of 1930, 1931, and 1935, during a special study of spiraling in certain 

species of ciliates at Tortugas, there appeared in the author's cultures from time to time 

many other species which seemed to be new or little known. There are now fifteen species, 

either new to science or Little known, about which it is believed sufficient information is 

available to justify their description or redescription. Five of the fifteen species have 

previously been described, but none of them is well known. Ten were described as new. 

They were characterized by amazing shades of color, yellow and red predominating. The 

species here discussed and described constitute only a few of those which have been seen 

at Tortugas at one time or another, but these are all the author feels justified in discussing, 

at the present time, with the information at hand. 

36. Burkenroad, M. 1929. Studies upon plankton and the mechanism of sound production in 

Haemulidae. Carnegie institution of Washington, Year Book 28: 283-90. 

Daily tows were made from May 31 to August 19 near Loggerhead Key. The variety of

species and numbers of individuals found disputed the notion that the Tortugas region 

"once noted for the variety and richness of its floating life, has gradually become in recent 

years an almost desert sea." 

37. Caira, J. N. and M. H. Pritchard. 1986. A review of the genus Pedibothrium Linton, 1909 

(Tetraphyllidea Onchobothriidae) with a description of two new species and comments on 

the related genera, Pachybothriurn Baer and Euzet. 1962 and Balanobothriurn Hornell, 

19 12. Journal of Parasitolog). 72, no. 1: 62-70. 

A review of the genus Pedibothrium Linton, 1909 is based on type and voucher specimens. 

The type species, Pedibothrium globicephalum Linton, 1909 is redescribed. Descriptions 

of Pedibothriurn brevispirte Linton. 1909 and Pedibothrium longispine Linton, 1909 are 

emended. Two new species are described, the generic diagnosis is emended, and a key is 

provided. 

38. Calder, D. R. 1992. Similarity analysis of hydroid assemblages along a latitudinal gradient in the 

Western Atlantic. Canadian Journal of Zoology 70, no. 6: 1078-85. 

Shallow-water (0.100 m depth) hydroid faunas reported from 26 locations along the 

western North Atlantic coast between the high Canadian Arctic archipelago and the 

Caribbean Sea were compared. Species numbers varied widely between locations, but 

were highest in the tropics and subtropics, lowest in arctic and subarctic waters, and 

intermediate in mid-latitudes. Percentages of species producing free medusae were lowest 

in high latitudes, intermediate in low latitudes, and highest in mid-latitudes (especially in 

estuaries). In a numerical analysis, sin~ilar hydroid faunas were identified at locations (i) 

between the high Canadian Arctic islands and the Strait of Belle Isle off western 

Newfoundland; (ii) between the Gulf of St. Lawrence and Chesapeake Bay; (iii) between 

North Carolina and southeastern Florida (south as far as St. Lucie Inlet), and including the 

northern Gulf of Mexico; (iv) in the Caribbean Sea, together with Dry Tortugas and the 

oceanic island of Bermuda. The greatest change in hydroid species composition along the 

coast appeared to occur around Cape Hatteras. 

39. Carrier, J. C., H. L. Pratt Jr. and L. K. Martin. 1994. Group reproductive behaviors in free-living 

nurse sharks, Ginglymostorna cirrarurn. Copeia 3: 646-56. 

Mating events of the nurse sharks were observed in a nine-day period in the Dry Tortugas 

islands. There were four stages of mating: precoupling, coupling, positioning and 

alignment, and insertion and copulation. Films were made of four of the mating events. 

Seminal fluid released into the water was obtained following one of the copulations. It 

showed the presence of free, nonpackaged sperm cells. Of the fifty mating events observed, 

ten of these involved multiple males attempting to copulate with single females. 

40. Carrier, J. C. and H. L. Pratt Jr. 1997. Habitat management enclosure of a nurse shark breeding and 

nursery grounds. Fisheries Research (In press). 

Based on nurse shark breeding studies conducted at Dry Tortugas, a sanctuary for nurse 

shark reproductive and nursery activities is being established at Dry Tortugas National 

Park. 

41. Cary, L. R. 1915. The Alcyorturia as a factor in reef limestone formation. Proceedings of the 

Natiortal Academy of Science 1: 285-89. 

In many areas of the Floridean-Antillean region, Gorgonaceae rather than stony corals 

make up the most characteristic feature of the lime-secreting organisms permanently 

attached to the bottom. In this paper, data are presented on the amount of material 

contributed to reef formation by gorgonians. Three factors were taken into consideration: 

spicule content (the amount of lime held as spicules in the colonies), distribution of 

gorgonians on the Tortugan reefs (the bulk of the gorgonians on any reef area) and

disintegration of the coenenchyma of the colonies and the addition of their spicules to the 

reef building materials. Using line surveys and the weight and percentage of spicules in the 

colonies, it was found that the amount of lime held as spicules in the tissue of living 

gorgonians per acre of reef area is 5.28 tons. Next to the destruction of the colonies by 

wave action (storms), the greatest mortality of the colonies is from overgrowth of tissues by 

other organisms. The destruction of Tortugan gotgonian colonies was nearly complete in 

the hurricane of October 1920. It has been estimated that nearly one-fifth of the gorgonian 

colonies are destroyed annually. 

42. -- . 1918. The Gorgonaceae as a factor in the formation of coral reefs. Papers Tortugas 

Laboratory 9: 341-62. 


An important constituent of the limestone of coral reefs is the calcium carbonate secreted in 

the skeletal structures of Anthozoa and marine calcareous algae. Representatives of the 

Hydrozoa were important reef formers in past geological epochs, but in the formation of 

modern reefs they constitute a minor factor. Representatives of the Anthozoa, the stony 

and flexible corals, are among animals the only important agents in the formation of the 

modern reefs. The results of this study show that over large reef areas, in the Tortugas at 

least, the gorgonian fauna is by far the most important element contributing to the 

formation of reef limestones. The amount of spicules in the tissues of gorgonian colonies 

would average at least 5.28 tons to the acre for all of the reefs in the Tortugas group. The 

figures given represent only a potential contribution to reef formation but a study of the 

normal cycle of changes in the gorgonian fauna of this region has shown that at least a fifth 

of this amount of calcium carbonate, as spicules, will be added to the reef limestones 

annually. 

43. ------, 1934. Growth of some tissues of Ptychodera bahamensis in vitro. Papers Tortugas 

Laborator), 28: 195-213. 


Nearly all refinements and expansions of the technique of tissue culture have taken place 

with wann-blooded animals as the experimental material. This line of development has, no 

doubt, been followed because of its possible medical application. Technical difficulties 

extending this method to invertebrates, where the necessary asepsis is more difficult to 

attain, have also played a part. The writer developed a technique which was successfully 

applied to some tissues of eleven species of marine animals belonging to seven phyla. In 

all cases, both migration and cell multiplication were obtained. Two organisms seemed to 

offer particularly favorable material for tissue culture. One was the gastropod Astroea 

longispina; the other was the enteropneustan Ptychodera bahamensis. This being the 

most convenient material with which to work, investigations in 1932 were confined to the 

tissues of this species alone. The technique of a method, using either hexyl-resorcenol or 

ultraviolet radiation in amounts harmless to the tissues, for growing in vitro the cells of 

~narine invertebrates is described. Because of their structure, members of the 

Enteropneusta lend themselves especially well to the obtaining of explants composed of 

one or of several types of tissue. The growth and reproduction of cells from the caecal 

portions of the intestine are recorded in detail. The changes undergone by muscle cells 

when removed from the body of an animal show a characteristically reversible series of 

stages peculiar to this type of cell. The bearing of the observations on Ptychodera cells to 

broader problems of cytology is considered 

44. ------- . 1915. The influence of the marginal sense organs on functional activity in Cassiopea 

xamachana. Proceedings of the NationaIAcademy of Science 1: 61 1-16. 

The influence of sense organs (nervous system) on the rate of regeneration was examined 

at the Dry Tortugas using the disks of the rhizostomous medusa Cassiopea, which can be

separated from the oral arms and kept in dishes of seawater for an indefinite period. Pairs 

of disks were examined from which all of the thopalia were removed, while from the other 

equal amounts of tissues were removed from the bell margin between the thopalia. In all 

instances, the disks where the half on which the thopalia remained regenerated at a more 

rapid rate than the inactive half. Other experiments focused on influence of sense organs 

on the rate of metabolism as measured by production of carbon dioxide. Carbon dioxide 

produced was always greater for the normal disk containing sense organs. It was 

concluded by the author that in this type of experiment there is some other form of 

metabolic activity which is of greater importance as a source of COz and which is more 

directly under the influence of the sense organs than is the activity of the muscular system. 

45. . 1916. The influence of the marginal sense organs on the rate of regeneration in Cassiopea 

xatnachana. Journal of Experiinental Zoology 21, no. 1: 1-3 1. 

Studies were conducted on accepting the view of the direct or indirect influence of the 

nervous system on regeneration in Cassiopea xamacharza collected from the Fort Jefferson 

moat at Dry Tortugas. Experiments conducted to determine the influence of sense organs 

on the rate of regeneration were inconclusive, when testing entire disks with sense organs 

removed, compared with specimens where the sense organs remained because of wide 

differences in physiological activity between different individuals. Half disks with sense 

organs regenerated more rapidly than those half disks without sense organs. Other 

experiments involving electrical stimulus by induction shocks on disk halfs, with and 

without sense organs, indicated regeneration is faster in the activated half, than from the 

inactive disk. These experiments indicate the rate of regeneration is simply one expression 

of the general metabolic activity of an animal, and as such is subject to the influence of the 

nerve centers, as are many of the functional activities. 

46. 1914. Observations upon the growth-rate and ecology of gorgonians. Papers Tortugas 



This report provides a record of observations extending over a 3-year period on the growth 

rate of Gorgoniaf7abellum and Plexauraflexuosa on the reefs around the Dry Tortugas, 

Florida. Ecological observations are supplemented by observations made in Jamaica. For 

effective attachment of the planule, the presence of depressions or cracks into which the 

planule could settle appears to be the most important factor. In comparison with young 

coral polyps the gorgonian colony has an obvious advantage, in that is most rapid growth is 

perpendicular to the surface, which permits its most rapidly growing part to secure food 

and oxygen. Wave action during very severe storms is by far the most destructive agent to 

which Gorgoi~in are subjected. It appears that the greatest destruction by storms comes 

from the tearing of the Gorgonia colonies from the substrate rather than laceration of 

tissue. 

47. 1917. Studies on the pliysiology of the nervous system of Cassiopea xamachana. Papers 

Tortugas Laborato,y 11: 121-70. 

Carnegie Institution of Washington Publication Number 25 1. 

In this paper are gathered the results of several distinct lines of experirnentation. They deal 

with some phase of the physiology of the nervous system of Cassiopea and represent 

portions of a general program of research on the nervous system of the lower animals. On 

account of its ability to live under adverse conditions and to withstand practically any type 

of operation, Cassiopea is an especially favorable form for experimentation and has been 

used as a subject for many researches. The experiments with entire disks, when the rates of 

regeneration of specimens on which the sense-organs remained are compared with those of 

specimens from which all sense-organs are removed, are inconclusive because of wide 

differences in physiological activity between different individuals. When we compare the

insulated halves of a disk, on one of which the sense-organs remain, while all of them have 

been removed from the other half, it is found that the half-disk with sense-organs always 

regenerates most rapidly. When all the sense-organs are removed from a disk and the 

halves insulated, the regeneration is faster from the activated than from the inactive halfdisk. 

These experiments indicate that the rate of regeneration is simply one expression of 

the general metabolic activity of an animal, and as such is subject to the influence of the 

nerve-centers, as are many other functional activities. Briefly summarized, the results of the 

observations made on the starved Cassiopea are as follows: In general the smaller 

Cassiopea loses relatively more in weight than does the larger Cassiopea. The percentage 

of water found in the entire body is nearly the same in all sizes of Cassiopea. The 

nitrogen-content of the entire body is higher in the small than in the larger Cassiopea. 

However, the absolute amount of nitrogen found in the starved Cassiopea is considerably 

higher than in the normal having the same bodyweight. The loss in weight of the different 

parts in the starved Cassiopea remains the same proportionately to those in the normal 

Cassiopea. 

48. -. 1918. A study of respiration in Alcyonaria. Papers TortugasLaborafory 12: 185-91 


Although the respiration of many species of invertebrates has been studied, the only 

references to that of Alcyonaria are those given by Montuori (1913), who studied two 

species, Alcyomeum pallidurn and Gorgonia cavolirrii. In these experiments the total 

weight of the colony was taken as the basis of comparison without taking into account the 

proportion of inert skeletal material- the spicules in the first species and the spicules and 

chitinous axis in the latter. The observations recorded were made as part of a study of the 

ecological factors determining the distribution of Alcyonaria on the coral reefs of southern 

Florida. All the species of the genus Gorgonia and the closely related Xiphigorgia, which 

have as a group the highest rate of respiration, are next to Rriareun~ the most resistant to 

increased temperature. Taken all together these observations indicate that some other 

factor is the controlling agency in the ability of a marine organism to withstand high 

temperatures. The acidity of the water at the close of the heat experiments was always 

greater than in respiration experiments carried on at 27.5" C. This may be only an 

expression of the abnormality of their metabolism at high temperatures, or have a causal 

relation to the death of the organism. 

49. Cate, C. N. 1978. New species of Ovulidae and reinstatement of Margovulap)wlina (A. Adams, 

1854) (Gastropods). Nautilus 92, no. 4: 160-167. 

Eight species of living Ovulidae are described as new, and the species M. pyr-ulina is 

reinstated. The 8 new species are listed as follows: Prionovolva castanea from the Gulf of 

Oman; Aperiovula restudiarza from Mukaishima, Japan; Prin~ovula sarttacaroliuensis 

from Mozambique; P. uvula from Moreton Bay, Queensland, Australia; Creuavolva 

periopsis from Java, Indonesia; Speculata adveuafrom off Sand Key, Florida; Cyphorna 

rhomba from Fort Lauderdale Reef, Florida; and Psudocyphoma gibbulurn from off the 

Dry Tortugas Islands, Florida. 

50. Chambers, E. L. 1937. The movement of the egg nucleus in relation to the sperm aster in the seaurchin, 

L,ytechinus varigeatus. Carnegie Instirutior~ of Washington, Year Rook 36: 86-87. 

With the aid of a camera, 30-second observations on the positions (rate and direction of 

movement) of the egg nucleus and sperm aster of Lytechiuus were made. 

51. Child, C. A. 1992. Shallow-water Pycnogonida of the Gulf of Mexico. Men~oir:~ of the Hourglass 

Cruises 9, no. 1: 1-86. 

This paper treats 11 species in 8 genera of the Pycnogonida that were collected during the 

Hourglass Cruises, a sampling program conducted on the central West Florida Shelf for 28

months during 1965.1967. Five benthic stations in depths from 6 to 72 m were sampled 

monthly with dredges and trawls among each of two transects. Treatments of 20 more 

species in 6 additional genera from other shelf collections are also included to offer a 

comprehensive survey of species (a total of 31 species in 14 genera) known from the 

continental shelf of the Gulf of Mexico, excluding the Dry Tortugas and the Florida Keys. 

Three of these species were previously unreported from the Gulf. Two new species, 

Ascorhynchus crenatum and A. horologiiam, are described from the Hourglass material, 

and an additional new species, Anoplodactylus dauphinus, is described from the other 

material. Artificial taxonomic keys are provided for all Gulf of Mexico families and 

species, and checklists are provided for all species known or expected to occur in the Gulf. 

All species are diagnosed and illustrated. and their distributions are given. Only four 

species were taken during the Hourglass Cruises with sufficient frequency to allow analysis 

of their distributions and abundances. 

52. Clapp, R. B. and W. B. Robertson Jr. 1986. Nesting of the masked booby Sula dactylatara on the 

Dry Tortugas, Florida. The first record for the contiguous United States. Colonial 

Waterbirds 9, no. 1: 113-16. 

In both 1984 and 1985 masked boobies (Sula dacfylatra) attempted to nest on sandy islets 

at the Dry Tortugas, Florida. Nesting attempts failed because the nest sites were washed 

away by summer storms. It seems likely that this species will eventually nest there 

successfully and will establish a small breeding population. This is the first documented 

nesting by this species in the contiguous United States. 

53. Clark, H. L. 1919. The distribution of the littoral echinoderms of the West Indies. Papers Tortugas 



The purpose of this investigation was to determine if the distribution of littoral 

echinoderms varied among various northern West Indian islands bounded by Bermuda to 

the North, Tobago to the South, and the Tortugas to the west. Five classes of echinoderms 

are discussed, including Comatulidea (feather-stars). Asteroidea (sea-stars), Ophiuroidea 

(brittle-stars), and Echinoidea (sea urchins). The number of species and numbers of 

individuals of the classes are discussed. Of the island areas investigated, the Tortugas 

appears to be the richest in the number of species found with 76 littoral echinoderms, with 

70 available to collect by hand. The sea urchins, Echinometra lacunfer were reported to be 

excessively abundant on many reefs, actually occurring by the thousands. The number of 

echinoid species found throughout the region and Florida are compared with comments 

provided on the origin of echinoids in the West Indies. Next to brittle-stars, holothurians 

were considered to be the most abundant of the littoral echinoderms. 

54. Clark, L. B, and W. N. Hess. 1942. Swarming of the Atlantic palolo worm,.Leodice fucata (Ehlers). 

Papers Tortugas Laboratory 33: 21-70 (issued Oct. 1940). 

Various organisms show reproductive activity coinciding with the lunar cycle. At the Dry 

Tortugas, swarming observations were recorded during 1937-39 on the Atlantic palolo 

worm, in association with the quarter-moon phase. Other important factors determining the 

time when the worms reproduce, include the maturity of the animal and the amount of 

water turbulence. It was concluded that: (1) the stimulating effect of the first-quarter moon 

is less than that of the third-quarter. (2) the worms increase in sensitivity to the stimuli 

inducing the swarming as they become sexually mature, and (3) wave action and water 

turbulence above a certain level induced by an 8 mph wind decreases or prevents swarming 

at the Dry Tortugas.

Cole, L. J. 1906. Ant Studies. Carnegie Institution of Washington, Year Book 5: 110. 

To investigate the biology of ants at the Tortugas, specimens were collected, and 

observations were made. 

Collie, M. R. 1979. A Sabine's gull at the Dry Tortugas. Florida Field Naturalist 7, no. 2: 28. 

A photographic observation was made by the author during August 1978 of a Sabine's gull 

at Garden Key, Dry Tortugas. There are four other records of this species in Florida along 

the Atlantic coast, however this sighting represents the first record of the Sabine's gull at 

the Tortugas. 

Colman, J. 1931. The superficial structure of coral reefs: animal succession on prepared substrata. 

Carnegie lnstitutior~ of Washington, Yearbook 30: 395 

Plant and animal successions were examined on concrete cubes planted in the water at 

three sites: Fort Jefferson moat, an iron wreck east of Loggerhead Key, and northwest of 

loggerhead Key. Also, a detailed ecological survey of Long and Bush Keys was made. 

Conger, P. S. 1924-1935. Diatom studies. Carrtegie Instirution of Washington, Year Book. 

Note: published as: 1924, V. 23, p. 220; 1925, v. 24, p. 221; 1926, v. 25, p. 240; 1927, 

v.26, p. 220; 1928, v. 27, p. 271; 1929, v.28, p. 283; 1930, v. 29, p. 323. 

Diatom studies were conducted in association with A. Mann. Narrative same as in 

reference no. 21 8. 

Conklin, E. G. 1908. The habits and early development of Lir~erges mercurius. Papers Tortugas 

Labor-atorj 2: 153-70. 


The jellyfish, Liner-ges rnercurius, Scyphomedusa, was investigated at Tortugas and Nassau 

Harbor. The sudden appearance in great numbers of this species at Tortugas was noted, 

followed by their rapid disappearance. Normal movements of the medusae are described 

as well. Other phases of early development, including egg-laying, egg structure, 

maturation and fertilization, first cleavage, second, and later cleavages, and blastula, 

gastula, and planula are described. Experiments on isolation of blastomeres and 

centrifugalized eggs are presented. The organization of the egg of Lir~erges and mechanics 

of cell division are described. 

-. 1908. Two peculiar actinian larvae from the Tortugas, Florida. Papers Tortugas 

Laborato~y 2: 171-86. 

Carncgie Institution of Washington Publication Number 103. 

During middle-of-the-day sampling tows at the Tortugas in May 1905, two peculiar larvae 

were collected. They did not undergo metamorphasis in aquaria. Natural history notes on 

living larvae are provided. Based on literature description, they were probably Zoanthidae 

of the Order Hexactinia. A band of strikingly brilliant, locomotor cilia was noted as most 

peculiar for these larvae. Their size, shape, and coloration are described. Yellowish-green 

symbiotic algae occur in both types and are hypothesized to be associated with their 

metabolism and play an important role in their nutrition. The morphology and histological 

character of these two types are similar but minor differences are described. Although 

these types have only been collected a few times world-wide, they are not considered rare 

at Nassau and the Dry Tortugas. 

Coonfield, B. R. 1940. Chromatophore reactions of embryos and larvae of Por~lacentrus 

leraco.stictras. Papers Torf~~gas Laboratory 32: 169-78 (issued Sept. 1940). 


Interest in the origin of the changing color patterns of fishes and certain other vertebrates, 

together with certain questions that have been raised by the work of investigators in their

study of this problem in embryos of vertebrates, prompted an investigation of the color 

mechanism in developing fishes. Both embryos and larvae of Pomucetrrriis leucosficf~rs, 

which is found in abundance in the Dry Tortugas, were used in this study. It was 

concluded that melanin granules migrate within the melanophores of Pornacentrus embryos 

as soon as these pigmentary bodies are completely formed. The melanophores of embryos 

a few hours of age contract in response to pressure applied with forceps and to a 

temperature of about 8'C. The melanophores of a majority of developing embryos, from 

their beginning up to a few hours before hatching, are found to be in a stellate state 

regardless of whether these young are over a white or a black background or are in total 

darkness. A few hours before hatching, the melanophores contract when the embryos are 

over a white background, expand when they are over a black background, and contract 

when they are in total darkness. This response continues in these young on through the 

hatching period and for a few hours after hatching. Larvae of two or more days after 

hatching do not show any conclusive response to different backgrounds or to the absence 

of light. The eyes of these young fish are believed to have no function in controlling their 

melanophore responses. The evidence is in favor of the release of a horn~one within the 

capsule just before the embryos hatch. This agency either permits or directly causes the 

melanophores to respond to various environments. 

62. -. 1940. The chromatophore system of larvae of Crarigorf artriillafus. Pq~er.s Torfugas 

Laboratory 32: 121-26 (issued May 1940). 


The ability of certain animals to imitate the color in their background is so striking that is 

has received the attention of investigators for a considerable period of time. This feature 

has been observed principally in fishes, amphibians, and reptiles of the vertebrates, and in 

crustaceans of the invertebrates. This paper adds to this field of study the results of 

observations on the reactions of the chromatophore system of larvae of Craf~gon 

armillarrrs. The erythrophores of normal larvae of Crangotf armillatus react as follows 

according to different backgrounds: the pigment is dispersed when the animals are kept in a 

white illuminated bowl. The erythrophores of enucleated specimens show the following 

conditions when subjected to different backgrounds: the pigment becomes concentrated 

when the specimens are kept over a white illuminated background; the pigment is 

dispersed when these specimens are subjected to black illuminated bowls. The time 

required for concentration of pigment in the erytlsophores is much longer than that 

required for its dispersion. Ablation of the eyes permits the erythrophores to react directly 

to stimulations caused by different backgrounds 

63. CoutiCre, H. 1910. The snapping shrimps (Alpheidae) of the Dry Tortugas, Florida. Proceedi~igs of 

the United States Notional Museufn 37: 485-87. 

The Alpheidae collected by Dr. McClendon at the Tortugas in 1908 are discussed. The 

Alpheidae are referable to eight different forms, including one new species and one new 

subspecies: Alpheus formosrrs Gibbes. Alpheus crisfulil,-011s Rathbun and Alpheus 

armillatus H. Milne-Edwards. 

64. Cowles, R. P. 1908. Habits, reactions, and associations in Ocjpodu arernaria. lJuper,~ Tortugas 



On Loggerhead Key, investigations were made on the behavior of 0c)podu arer~mria. It 

was found that adult ghost crabs build two kinds of burrows. One consists of a single 

tunnel extending down in the sand for 3 to 4 feet. The other is similar, except that it is 

shorter and has a passage branching off from it, which is used for escape. Young ocypodas 

make short burrows, only a few inches long, which often extend vcrtically downward. 

Breeding in the region of Loggerhead Key probably occurs in the spring and early summer.

Ocypoda is a scavenger and a cannibal. The eyes do not seem to play an important role in 

the detection of food, but they undoubtedly lead individuals to objects which may be food. 

That Ocypoda is stimulated by odors was not conclusively shown, but certain experiments 

point strongly in that direction. The eyes are highly developed, so far as crustacean eyes 

are concerned; they are quite sensitive to large differences in the intensity of light; they do 

not react to different colors; they aid much in the search for food, in the detection of 

enemies, and in the accuracy of locomotion. Ghost crabs probably do not have vision such 

as that of the human eye, nor do they see the color and finer characters of the surface of an 

object, but they undoubtedly see its outlines and possibly some of the more evident 

irregularities of the surface made evident by differences in lighting. The color-pattern seen 

through the carapace of Ocypoda changes in intensity under different conditions of 

temperature and light. In the absence of light when the temperature is anywhere between 

22" C. and 45" C., and undoubtedly when it is even lower or higher, a light coloration 

occurs. Generally in diffuse light and even direct sunlight a dark coloration appears, 

provided the temperature is not too high. Usually at low temperatures, not above 3S°C., a 

light coloration is the rule, and it occurs independently of the intensity of light. At high 

temperatures, above 35" C., a light coloration is the rule, and it occurs independently of the 

intensity of light. No indication of audition was observed in Ocypoda. The so-called 

"auditory organs" are equilibrating organs. Ocpoda has a stridulating ridge on the palm of 

its large chela. 

65. --- . 191 1. Reaction to light and other points in the behavior of the starfish. Papers Tortugas 

Laborator), 3: 95-1 10. 


Experiments were designed to test the reactions of starfish to light. Two species were used 

Echinaster crassispinu and Astropecten duplicatus. Both are migratory and are found in 

open waters over sandy bottoms, in areas generally exposed to light. In the Tortugas 

laboratory, starfish were placed in aquaria wooden boxes and tested for movement up in 

response to light, inclines, vertical walls, and tilted floors. Every specimen reacted 

positively, moving toward bright light. Even with eye-spots removed, movements towards 

light are positive, but not as quick as in normal individuals. When tested in different 

degrees of water temperature the reaction to light was positive at ordinary temperatures. 

Quality of light was tested using various color screens (UV, violet, blue, green, yellow, 

orange and red) in a box with closed and open ends. The source of light was sunlight. In a 

series of 10 tests with varied orientation and handling, in nearly every test starfish moved 

toward light without hesitation. 

66. Criales, M. M. and T. N. Lee. 1995. Larval distribution and transport of penaeoid shrimps during 

theTortugas Gyre in May-June 1991. Fishery Bulletin 93: 471.82. 

As part of the Southeast Florida and Caribbean Recruitment (SEFCAR) project, penaeoid 

shrimp larvae were collected during the spring and summer cruise of the RV Longhorn in 

the Lower Florida keys and Dry Tortugas from 29 May to 30 June 1991. Larvae of the 

pink shrimp, Penaeus duarurum, and the rock shrimp, Sicyonia sp., were distributed 

inshore close to the Dry Tortugas Grounds adjacent to the boundaries of Dry Tortugas 

National Park, whereas larvae of the oceanic shrimp Solenocera sp. showed mainly an 

offshore distribution. Significant concentrations of Solenocera sp., Sicyonia sp. and P. 

duorarum larvae at the Tortugas transect in early June were found within and above the 

seasonal thermocline, while the cold cyclonic Tortugas Gyre was intensively developed. 

For Solenocera sp., which spawn on the outer ridge of the gyre followed by onshore 

Ekman transport. Penaeus duorarum, which spawn in the shallow Tortugas Grounds, 

showed a mode of zoea 11-111 progressing to postlarvae I at the Tortugas Grounds during 

the 15 days in which the drifter Halley recirculated in the interior of the Tortugas Gyre. 

Retention of P. duorarum larvae by the internal circulation of the gyre at the spawning

grounds may be an important mechanism for local recruitment of these shrimp to the 

nursery ground of Florida Bay, Everglades National Park. 

67. Cushman, J. A,. 1922. Shallow-water Foraminifera of the Tortugas region. Papers Tortugas 



The paper gives the results of a study of collections made in the waters about the Tortugas 

Laboratory of the Carnegie Institution of Washington. Collecting was done largely from 

the boats, the most satisfactory method that was used with the Darwin. Collecting in the 

moat at Fort Jefferson in shallow water on Long Key, as well as on the reefs and flats, was 

done by hand. The Tortugas region presents an ideal spot for studying the shallow-water 

tropical Foraminifera of this particular region. It is removed from influence of shore 

conditions; the water is at all times warm and pure, so that ecological conditions that are 

present are constant. The twenty stations from which bottom samples were studied in the 

preparation of this paper, together with collections from reef flats and from the eel-grass, 

give a considerable range of conditions. The only stations at which Rotalia was found are 

two in the moat at Fort Jefferson and the other in a very shallow lagoon at Long Key, 

where the water was warm at low tide in June. By most authors, these specimens would 

ordinarily be referred without question to Rotalia beccarii (Linnaeus). There are 

differences from northern material, and in probability the Tortugas specimens belong to 

different species. On the banks of dead coral whicli become exposed at spring tides, great 

masses of attached Foraminifera develop. Of these, the most abundant is Homotrema, 

which makes an appreciable contribution to the mass of material. With it, in crevices of the 

dead coral, was a new species of Haliphysema. On the eel-grass (Posidonia), which forms 

in shallow water inside the reef, there is Iridiu, Planorbulina, Discorhis, Orbitolites, with a 

peculiar miliolid which spreads over the surface. The mass of these must add appreciably 

to the amount of carbonate of lime added to the bottom. The forms are rapid in their 

growth, as the leaves of Posidonia are quickly covered in their growth by Foraminiferea 

and other encrusting animals. 

68. Cutright, P. E. 1937. Studies on the development of the dorsal spine of sting rays. Carnegie 

Irrstiturion of Wazhington, Year Book 36: 90. 

Th~s report describes the collection of southern sting rays for a histological examination of 

the stinging mechanism. 

69. Dall, W. H. 1889. Reports on the results of dredging, under the supervisioti of Alexander Agassiz, 

in the Gulf of Mexico 1877-78 and in the Caribbean Sea (1879-80), by the U.S. Coast 

Survey Steamer "Blake":, Lieutenant Commander C.D. Sigsbee, U.S.N., and Corntnander 

J.R. Bartlett, U.S.N. Commanding. XXIX- Report on the rnollusca . Part I1 Gastropoda and 

Scaphoda. Bulletin of the Museu?n of Compamtivr Zoology at Hanard College 18: 1-492, 

with thirty one plates. 

This listing of Mullusca collected by the "Blake" is supplemented by the southern 

dredgings of the U.S. Fish Conlmission Steamer "Albatross" and other inaterial collected 

from the region. A systematic description and account of the gastropods and scaphopods is 

given and illustrated. Non~enclature is discussed and rectified in several cases. 

70. Darby, H. H. 1934. The rncchanisms of asymmetry in the Alpheidae. Paperx Tortugas I~boratory 

28: 347-61 (issued Feb. 1934). 


In 1901, Przibram reported a series of striking experiments on the regeneration of chelae of 

Alpheus dentipes, A. p1at)~rkynchras and A. ruber. There is a pronounced quantitative and 

qualitative difference between the right and left chelae. One chela is several times as large 

as the other. Pr~ibrarn showed that if the snap-claw is removed, at the next molt the pinch-

claw is changed into a snap-claw and a new pinch-claw is regenerated on the stump of the 

old snap-claw. This unusual reversal of asymmetry was confirmed by Wilson (1903) and 

Zeleny (1905). It was also shown that if both chelae are removed at the same time, they 

regenerate in their original positions. These experiments seem to indicate that the final 

degree of morphological expression of the gene may in certain cases depend on the 

environment. In Crango17 ar~nillarus, an asymmetrical individual, it is symmetrical ten 

days later. The morphological expressions of the gene are so concrete that it is difficult to 

realize that the gene may also be the controlling agency in the production of definite 

chemical substances, whose presence is manifested only by their physiological reactions. 

Environment from that point of view can quite easily be thought to control the amount of a 

substance produced. Changes niight well be induced by radiation, due to the ionization of 

the cell. Crangon has shown itself to be an organism in which studies on development as 

an expression of the activity of the gene can be undertaken with some hope of success. The 

nature of the regeneration of chelae in two members of the family of Alpheidae has been 

studied; in particular, in Crar~gor~ armillarus. It has been shown that at certain stages in 

the development of the chelae, a state is reached that permits the determination of which 

side is to have the large chela, or snap-claw. Equal chelae have been produced 

experimentally and are of three varieties: (I) both small (pinch-claws); (2) both large (snapclaws); 

(3) both intermediate. 

71. -- . 1940. Symmetry in normally asynnnetrical crustacea. Papers Tortugas Laboratory 32: 61- 

64 (issued Oct. 1939). 


A symmetrical specimen of Crungon armillatus was found in nature with two snap claws. 

These claws differed in no way from snap claws produced experimentally and reported 

previously. 

72. Darby, H. H., E. R. F. Johnson and G. W. Barnes. 1937. Studies on the absorption and scattering of 

solar radiation by the sea: spectrographic and photoelectric measurements. Papers 

Torrugas Labor-atot-y 31: 191-205 (issued Oct. 1936). 

Carnegie Institution of Washington Publication Nurnber 475. 

A considerable amount of work has been done in recent times on the penetration of 

radiation into sea and lake water. The importance of this work, in such matters as plant and 

animal metabolism and undcr-water photography, is obvious. The amount and spectral 

distribution of scattering, and the penetration of the ultraviolet component, are two phases 

of the subject which have received scant attention. These studies were made from the 

yacht, Elsie Fenbnore at the Tortugas Laboratory. A comparative study has been made of 

two methods of evaluating the transmission of various wave lengths of light through sea 

water: (1) photometry by means of photoelectric cells, and (2) photographic 

spectrophotometry. Bertel's observation that ultraviolet light penetrates a considerable 

distance into the sea has been confirmed. The extent of penetration is greater than would 

be expected from the laboratory data of Hulbert and Sawyer. A rough evaluation of the 

transmissive exponent from 4500A to 3250A was made, which indicates the magnitude of 

the disagreement. Scattering was found to be selective, becoming greater with decreasing 

wave length. The spectral distribution of scattered radiation is indicated. The importance 

of these observations for biological systems is outlined. 

73. Davis, G. E. 1977. Anchor damage to a coral reef on the coast of Florida. Biologicul Con.servation 

1 1 : 29-34. 

Twenty percent of an extensive staghorn coral Acropora cer-vicorrris has recently been 

damaged by boat anchors in Fort Jefferson National Monument, Dry Tortugas, Florida. It 

is suggested in this article that this type of damage may occur in other coral reef sanctuaries

unless anchor-sensitive areas are identified and closed to anchoring. Alternatively, 

mooring buoys should be provided by sanctuary managers. 

74. --- , 1982. A century of natural cl~ange in coral distribution at the Dry Tortugas, Florida USA. A 

comparison of reef maps from 1881 and 1976. Hulletirl of Marine Science 32, no. 2: 608- 

23. 

Changes in coral reef structure and composition at Dry Tortugas, Florida were compared 

over a 95-year interval from benthic maps prepared in 1881 and 1976. Living hermatypic 

corals occupied less than 4% of the 23,000-hectare area mapped, and showed little change 

in area during the interval between maps. However, major changes in coral species 

distributions and reef types were apparent. In 1976, a lush 220-hectare Acropora 

cervicornis reef occupied what had been octocoral dominated hard bottom in 1881. The 

44-hectare swath ofA. palninfa on the reef crest in 1881 was seduced to two small patches 

totaling less than 600 rnZ in 1976. More than 90% of the extensive thickets of A. 

cervicornis at Dry Tortugas were killed during the winter of 1976-77, apparently as a result 

of thermal shock. These changes in coral distribution and abundance demonstrated the 

natural dynamic nature of coral reefs, and showed the important role occasional short-term 

extreme climatic events can play in shaping coral reef structure and species distribution. 

The importance of protecting living corals and the value of ecosystem level sanctuaries as 

dynamic standards are discussed. 

75. -. 1977. Effects of recreational harvest on a spiny lobster, Panulirus nrgus population. 

Bulletin of Marine Science 27, no. 2: 223-36. 

A commercially unfished population of Panulirius argus was studied in Fort Jefferson 

National Monument at Dry Tortugas, Florida, from April 1971 to July 1975. For 29 months 

all harvest was prohibited, then an experimental sport harvest (hand caught by recreational 

divers) was allowed in 50% of the area for a period of 8 months, followcd by 16 months of 

complete protection for assessment of recovery. Data on the size, abundance, and natural 

history of the lobsters were collected using SCUBA, and commercial trapping techniques. 

A total of 4,257 lobsters, with a mean carapace length of 101 mm, was tagged and released 

at Dry Tortugas. The existence of a resident adult P. arg~rs population was demonstrated 

by the recovery of all recaptured lobsters (7.3%) with 10 km of their respective capture 

sites up to 104 weeks after release. Immediately following the experimental sport harvest, 

the population in the sport harvested area showed a 58% reduction in trap catch rate and 

dispersed to 42% of its pre-harvest lair occupancy density, while the population in the 

unharvested control area remained essentially unchanged. The catch rate in the sport 

harvested area recovered to 78% of its pre-harvest level after 1 year of complete protection 

from harvest, and the lair occupancy rate recovery was 71% after 16 months of post harvest 

protection. The pre-harvest standing crop was estimated at 58.3 kglha, wet weight. 

76. -. 1977. Fishery Harvest in an Underwater Park. Proceedings, Third lnfeniationul Coral Reef 

Symposium, 605-8 no. 2. RSMAS, Univ. Miami, Coral Gables, Florida. 

There is a potential conflict between park management for preservation of maximum 

species richness and fishery harvest in parks. The recreational harvest of spiny lobster, 

Par~ulirrrs argus, at Ft. Jefferson National Monument, Dry Tortugas, Florida, dcmonstrates 

the nature and extent of the conflict. An eight-month-long diver harvest, limited by a daily 

bag limit of two lobsters, reduced thc previously unfished population by 585 and 

significantly altered the local lobster distribution. Growth and natural recruitment did not 

restore the population to its 58.3 kglha pre-harvest level, even after 16 months with no 

additional harvest. The trophic status of spiny lobsters as high level carnivores and current 

ecological theory combined with the harvest impact observed at Dry Tortugas suggests that 

community structure and species richness would be si~nificantly altered by the harvest.

77. ------ . 1975. Minimum size of mature spiny lobsters, Panulirus argus, at Dry Tortugas, Florida 

USA. Trarisactioris of the American Fisheries Society 104, no. 4: 675-76. 

Of 1,594 female spiny lobsters examined during April 1973.1975 at the Dry Tortugas, 55% 

were bearing eggs (berried). The specimens ranged in carapace length from 39 mm to 140 

mm. No berried females were round with carapace lengths less than 78mm. Maturity was 

reached by one half of the females in the 86-95 mm size class. The current minimum legal 

size for sport and commercial lobster fishing in Florida is 76-mm carapace length. 

78. ---. 1974. Notes on the status of spiniy lobster.^, Pa~iulirus argus, at Dry Tortugas, Florida, 

SUSF-SG-74-201. State University (Florida) System. Sea Grant Program. Publ.. 

Until mid-1971, sport harvest of spiny lobsters, primarily Pariulirus argus, was permitted 

in the 19,000 hectares underwater preserve created in 1935 which included the Dry 

Tortugas atoll. At that time there was a two lobster per person per day limit. Few visitors 

reached the isolated atoll during the first 20 years, with an average of some 1,200 people 

per year. Annual visitation increased to over 21,000 in the late 1960's and early 1970's. 

Concern was expressed for the protection of the quality and quantity of the lobsters found 

in the area. The primary objective of the study was to assess the impact of human harvest 

on a natural unperturbed lobster population. 

79. -- . 1981. On the role of underwater parks and sanctuaries in the management of coastal 

resources in the southeastern United States, Et~vironmental Cortservation 8, no. 1: 67-70. 

Aquatic resources in parks and reserves are not as adequately protected as comparable 

terrestrial resources. Thus the values of protected aquatic ecosystems as standards for 

comparison, reservoirs of genetic materials, and 'emotional' reserves, are apt to be greatly 

diminished. Even seemingly static ecosystems such as coral reefs are dynamic, changing 

dramatically in response to natural short-term environmental variations. Such ecosystems 

require protected natural areas as dynamic standards that will allow distinctions to be 

drawn between effects of exploitation or pollution and normal variation. Furthermore, 

fisheries harvests may reduce the size at which exploited species mature, and reduce the 

amount and variability of genetic material produced by exploited populations. The seven 

underwater parks or sanctuaries established since 1935 (Dry Tortugas) in Florida and the 

US. Virgin Islands exhibit wide variations in the degree of protection accorded to aquatic 

resources, a range being apparent from nearly complete protection in the first parks to be 

established to virtually no protection at all in the recently established parks. The 

consequences of permitting consumptive uses of aquatic resources in parks and reserves 

need to be objectively evaluated. Unless these consumptive uses are severely curtailed or 

eliminated, the primary values of the parks and reserves may never by realized. 

80. -, 1980. Spiny lobster series. Gary E. Davis (ed.), 27 pgs. American Fisheries Society: 

Bethesda, MD. 

This series of papers regarding spiny lobster management represents the efforts of a broad 

cross section of the scientific fisheries community. Not only is there a diverse array of 

disciplines from biochemical genetics to ecology and economics, but nearly every source 

of research endeavor is represented. Members of two federal agencies (National Marine 

Fisheries Service and National Park Service), a state agency (Florida Department of 

Natural Resources), a public university (University of Florida), a private university (Nova 

University), and a private company (Science Applications, Inc.) have combined their 

efforts on a common subject that has already spawned thousands of scientific papers and 

countless popular articles. 

81. Davis, G. E. and J. W. Dodrill. 1989. Recreational fishery and population dynamics of spiny 

lobsters, Panulirus argus, in Florida Bay, Everglades National Park, 1977.1980. Bulletirt 

of Marine Science 44, no. 1: 78-88.

84. --- . 1942. The ecology of the vegetation and topography of the Sand Keys of Florida. Pal~ers 

Torrugas Laborator)' 33: 1 13-95 


This is a study of the vegetation and some of the physiographic features of about thirty 

islands of the Florida Keys, in an area extending west from Key West, Florida, and 

including the Dry Tortugas Keys. About thirty islands of the Florida Keys beyond Key 

West were investigated during the summer of 1940 and winter of 1942, and to some extent 

during the summers of 1937 and 1938. These studies were concerned with the topography 

and vegetation of these small, relatively isolated, and partly tropical islands. These islands 

are here termed the Sand Keys because most of the parts above high tide are composed of 

coarse calcareous sands, and also because this name was used by Willspaugh (1907). A 

few of the Marquesas and Tortugas Kcys have changed a great deal. The strand areas on a 

number of islands seem to be increasing at the expense of the mangrove swamps. The 

mangrove swamps have spread over wide areas in some instances and seem to be aiding in 

building up the islands. Most of the constructional processes are, however, due to maritime 

factors such as the ocean currents and tides. This paper is also a part of a series of studies 

of the plant ecology of southern Florida. This and the author's study of mangrove 

vegetation together describe most of the coastal and insular vegetation of that region. 

85. Davis, R. A. Jr. and C. W. O'Neill. 1979. Morphodynamics of East Key, Dry Tortugas, Florida. in 

Guide to Sedirnentationfor the Dr)' Tortugas,Fort Jeffcrsor7 National Monumerrr Florida 

Soutkcust Geological Socieij Puhlicatio~i 21: 7-13, 

East Key is comprised wholly of biogenic sand and fine gravel. It lacks beachrock or 

bedrock which may act as a stabilizing agent such as on Loggerhead Key. During the past 

two centuries, maps and charts documented the size, shape, and location of East Key. The 

Key moved in a generally southeasterly direction across the shallow carbonate bank. East 

Key was preserved, unlike some other islands, because of its easterly position with respect 

to the deep lagoon. Those islands west of the lagoon moved easterly and disappeared into 

the lagoon (O'Neill, 1976). East Key has decreased markedly in size during its 

southeasterly movement. In addition there appears to be a change in morphology which is 

related to seasonal changes in predominant wind direction. 

86. de Laubenfels, M. W. 1936. A discussion of the sponge fauna of the Dry Tortugas in particular and 

the West Indies in general, with material for a revision of the Families and Orders of the 

Porifera. Paperx Tortugas Lahoratorp 30: 1-225. 


Sponge specimens were collected ncar the Dry Tortugas by scientists affiliated with the 

Carnegie Institution of Washington, or working at tlie laboratory maintained on 

Loggerhead Key. These were sent to the U.S. National Museum to be studied by the 

author. The West Indian region has long been known as one of the richest collecting 

grounds for sponges in the world, and the Dry Tortugas offers a representative sample of it. 

The author identified several new fanlilies and in many cases proposed new names for 

families already in use. Representatives of each of the species discussed in this paper have 

been deposited in the United States National Museum. Each new species is described in 

detail. 

87. ------ . 1934. Physiology and morphology of Porifera exemplified by lotrochora bir-otulata Higgin. 

Paper-s Tor-tugas Laboratory 28: 37-66. 


The experimental work upon which this article is based was carried on during the summers 

of 1927 and 1928 at the Tortugas Laboratory. A taxonomic description of the sponge was 

provided. It was found that a hyaline ground mass or slime plays a very important role in 

the life of Iotrochota and perhaps numerous other sponges, Judging only by items visible

in living Iotrochotu cells, which were kept track of by conspicuously colored inclusions, 

new sponges resulted from disassociated cells without intermediate differentiation and 

respecialization. Reproductive bodies (gemmules) seem to result in Iotrochota by the 

migration together of cells previously specialized for the purpose. Bispecific 

conglomerations could be secured between lotrochota and other species, and these 

remained alive for two weeks or more, but whatever cell motility occurred within them 

tended toward the ultimate segregation of the two species after somewhat the manner in 

which animal gratings finally terminate. Amebocytes of lotrochota sometimes ingested 

flagellates which subsequently appeared as intracellular inclusions, and perhaps became the 

symbionts whereby there occurred a certain amount of photosynthesis, the existence of 

which was indicated by experimentation. 

88. 1953. Sponges of the Gulf of Mexico. Hulletir~ of Marine Science of the Gulfand 

Caribbean 2, no. 3: 511-77. 

In 1948, a collection of sponges was made by the Marine Laboratory of the University of 

Miami in the eastern Gulf of Mexico. Twenty-two stations were studied, at depths from 6 

to 20 meters, in the area between Dry Tortugas and the northeastern part of the Gulf. The 

collection comprises 52 species in 41 genera, all within the class Demospongea. Of these 

11 species are new. Additional description is provided for a number of species. An 

analysis of the sponge collection by stations is included. 

89. de Renyi, G. S. 1934. Studies of nerve cells of invertebrates. Carnegie Institution of Washington, 

Yea!- Rook. 33: 250. 

The nerve tissue (neuroplasm) of gastropods (Strombus gigas), Apljsia protea, Olivia 

litterata, Cypraea exanthetnu, Casio cameo), decapods (Panulirus argu,s, Crangon 

arnzillatus, Ocypodn albicnns), and he~nichordates (Ptyhodem bahamensis) were studied. 

The neoplasm of the Ciastropoda and Hetnichordata exhibited viscosity, and a certain 

degree of elasticity, whereas decapodean neuroplasrn was liquid. 

90. Detlaun, M. F. 1987. "The distribution and molecular characterization of dissolved DNA in aquatic 

environments." University of South Florida. Ph.D. Dissertation 

The distribution of dissolved DNA in oceanic, estuarine and freshwater environments in 

southwest Florida and the Gulf of Mexico was determined by using a method for the 

measurement of dissolved DNA based on the fluorescence of Hoechst 33258-DNA 

complexes. Oceanic concentrations of extracellular DNA ranged from 0.2 to 19 

decreasing as a function of distance from the shore and depth in the water column. 

Samples of the mucus-rich coral surface microlayer (CSM) collected on reefs in the Dry 

Tortugas had dissolved DNA concentrations from 1.8 to 11.7 times that in the overlying 

water. Estuarine concentrations, measured at three stations in Tampa Bay, FL over a 15- 

month period, followed the seasonal trend in concentrations in offshore environnients, 

while variations in the estuary were significant, with tnaxitnutn concentrations in nighttime 

samples. Although concentrations of dissolved DNA in the eutrophic Alafia River were 

generally higher than those in the oligotrophic Crystal River, values as low as 1.14 were 

measured in the Alafia. A wide range of molecular weights (determined by agarose gel 

electrophoresis) was found for extracellular DNA concentrated from various aquatic 

environments. These results indicated that dissolved DNA is in a size range sufficient to 

contain gene sequences, which may be important in natural transformation of microbial 

populations. A model system for probing extracellular DNA from aquatic environments 

was developed using the plasmid containing the herpes simplex thymidine kinase (TK) 

gene. Plasmid DNA and the TK gene fragment added to artificial seawater were 

concentrated and labeled TK to establish percent recovery and detection limits for the 

method. The degradation of plasmid DNA added to a natural seawater sample was 

monitored over a 36 h period by probing with the TK gene probe. Intact plasmid was

detected for up to 4 h and DNA hybridizable to the TK probe was detected for up to 24 h. 

These methods were used to probe for the TK gene in environmental samples of 

extracellular DNA. Hybridization to the TK probe was detected in both freshwater and 

estuarine samples. 

91, Dinsmore, J. J. 1972. Sooty tern behavior. Bulletin of the Florida State Museum of Biological 

Science 16, no. 3: 129-79. 

A four-year study of the breeding behavior of sooty terns (Sterna fuscata) was made at 

Bush Key, Dry Tortugas in the southeastern Gulf of Mexico. The results are compared 

with the behavior of other terns and the differences discussed, particularly in regard to the 

pelagic environment the sooty tern inhabits. sooty terns have a lower clutch size, longer 

period of development of the chick, and first breed when older than most other terns, many 

of which feed in marshes and coastal waters. These characteristics of sooty tern breeding 

biology are similar to those of many other pelagic birds. A distant food supply and high 

adult survivorship apparently have contributed to these differences from other terns. 

92. Dinsmore, J. J. and W. B. Robertson Jr. 1972. Sooty tern feeding on moths. The Auk 89, no. 2: 

440. 

While banding sooty terns (Sterna fuscata) at Bush Key, Dry Tortugas, Florida on June 28, 

1970, an adult tern regurgitated two moths 1.5 to 2 cm long together with several 

unidentified fish. The moths were identified to the family Noctuidae. Although the food of 

sooty terns at the Dry Tortugas has not been studied in detail, sizable collections of food 

regurgitated show that this population feeds on fish and squid. In 13 years of tern banding, 

this is the first time an insect has been found as part of the sooty tern's diet . 

93. Dole, R. B. 1914. Some chemical characteristics of sea water at Tortugas and around Biscayne Bay, 

Florida. Papers Tortugas Laboratory 5: 69-78. 


The chemical tests at Tortugas were performed by the writer in June 1913, in the Marine 

Biological Laboratory, Tortugas, Florida, for the primary purpose of ascertaining what 

soluble effect, if any, carbon dioxide in sea-water might have on coral and other deposits of 

calcium carbonate. The tests of waters from Biscayne Bay were made to ascertain the 

differences in concentration of sea-water in the bay and the diluting effect of Miami River. 

The salinities of the three samples taken outside the reefs agree closely with each other and 

with the salinity of Gulf water at Tortugas, Florida (36.01 ppt), which is somewhat greater 

than that of standard ocean water (35.02 ppt.). The water in the south part of the bay is 

somewhat more concentrated having salinities of 36.73,36.64, and 36.64 ppt., respectively. 

This evidence that the water in this part of the bay is concentrated by evaporation during its 

retention in the shallows serves further to indicate that circulation there is not very rapid 

and that the greater bulk of the water inside the keys is not thoroughly mixed or shifted by 

the tides. Sample 1 has a salinity obviously higher than the pure water of Miami River 

alone may be expected to have, and represents admixture with bay water: carbonates are 

absent from it, but bicarbonates are much higher than in the normal drainage from the 

Everglades and may be attributed to reaction of the carbon dioxide that the river water 

carries. 

94. Domeier, M. L. Speciation in the Serranid fish Hypoplectrus. Bulletin of Marine Science 54, no. 1: 

103-41. 

Research was conducted to determine the species status of individual color morphs of 

fishes in the genus Hypoplectrus (family Serranidae). Crossing two morphs of 

Hypoplectrus (H. unicolor x H. gerna) in the laboratory produced an F1 generation with an 

intermediate phenotype to that of the parental types. This intermediate morph cannot be 

assigned to any known morph and is thus termed a hybrid. Individuals of several

Hypoplectrus morphs were found to select only individuals of the same morph as a mate 

when provided a choice. Individual fish can sometimes be forced to mate with an 

individual of a different morph by not providing a choice of mates. The occurrence of 

hybrids was found to be low in the field, corresponding to the low occurrence of mixed 

matings in the field. Some differences in distribution were found between the different 

hamlet morphs. The new data provided by this study, which includes specimens collected 

from the Dry Tortugas, indicate that the different color lnorphs warrant full species rank. It 

is hypothesized that speciation in Hypoplecturs was driven by the rise and fall of sea level 

during the last ice age. 

95. Donaldson, H. H. 1916. Experiment on the feralization of the albino rat. Calrregie lrrrtitution of 


Domesticated albino Norway rats were released on East Key to determine if changes in 

brain weight occur over successive generations in a wild state. Since the rats were 

unmarked, it was impossible to ascertain if differences in weight were from new breeds or 

from animals in the original colony. 

96. Doyle, W. L. 1936. Cytology of Valortia. Papers Tortugas Laboratory 29: 13-21 (issued Nov. 

1935). 


For a number of years algal cells with large vacuoles have been the subject of research on 

the permeability of the plasma membrane. Prominent among forms investigated is 

Valonia. This paper describes the cytology of Valonia ventricosa and Valonia macrophysa 

with particular emphasis on structures of significance in physiological investigations. The 

cells were collected on Bush Key Reef and from the moat at Fort Jefferson and kept in 

finger bowls in the laboratory. The morphology of the various structures in the cytoplasm 

of Valorria macrophysa and V. verttricosu is described. The plastids produce starch and 

lipoid granules and are sufficiently numerous as to constitute two-thirds-of the volume of 

the cytoplasm. There are approximately three hundred nuclei per square millimeter of cell 

surface in the coenocytes. Mitosis is intranuclear. In the development of the rhizoidal 

hapteron cells of the aplanospores, the mitocho~idri arise from plastids of the coenocyte in 

which the aplanospores were formed. The large central vacuoles of the coenocyte arises by 

fusion of small vacuoles formed in the cytoplasm. Double vital staining of artifact 

vacuoles is noted. 

97. --- 1940. The structure and composition of Valortiu vcntricosu. Papers Tortugas Laboratory 

32: 143-52 (issued Sept. 1940). 

Carnegie Institution of Washington Publication Number 571 

The physiology of Valonia has been dealt with extensively by numerous authors. Cells 

were collected from Long Key and the adjacent reef and brought to the laboratory, where 

they were kept in large glass jars of sea water which was changed daily. Measurements 

have been made of the relation of the volume and thickness of the cytoplasm and cell wall 

to the size of the coenocyte. The specific gravities of various cell constituents and of cells 

of various sizes have been measured. Frorn a consideration of the results presented it 

would appear that the level of metabolic rate in Valorlia is of a low order, but not 

necessarily of a different order of magnitude from that of the barley-root and potato-slice 

systems. 

98. Doyle, W. L. and M. Metcalfe Doyle. 1940. The structure of zooxanthellae. Papers Torrugas 

Laboratory 32: 127-42 (issued May 1940). 


The structure of the zooxanthellae in various invertebrate reef organisms under various 

conditions was investigated at the Dry Tortugas in 1934, 1935 and rechecked in 1939. Ten

species of corals and foraminifera were studied in their living conditions, as well as after 

fixation. Zooxanthellae in foraminiferans, were examined for the effects of light in normal 

gas tensions, in increased carbon dioxide tensions, and on specimens in oxygen and 

hydrogen; while in corals the comparative cytology of zooxanthellae was studied. For the 

large heads of the Orl~icella (Madrepora), the amount of light present at the top and 

bottom of the corals determined the natural variations in the amount of calcium oxalate 

crystals in zooxanthellae. Increased levels of crystals were found at the bottom in 

darkness, while no crystals were found at the top. Similar results were found in 

foraminiferans. The converse is true for the amount of starch present. The zooxanthellae 

in corals under the most intense natural light conditions contains little starch, but abundant 

oil droplets. It was concluded that, overall, for the greater part of the day, the 

zooxanthellae, as well as the corals, are in need of oxygen. 

99. Drew, G. H.. 1914. On the precipitation of calcium carbonate in the sea by marine bacteria, and on 

the action of denitrifying bacteria in tropical and temperate seas. Papers Torrugas 

Luboraro~y 5: 7-45. 


The investigations described in this paper were made in the summers of 1911 and 1912 

under the auspices of the Carnegie Institution of Washington. The intent was to study the 

action of marine denitrifying bacteria in tropical seas. The discovery that these denitrifying 

bacteria also possess the power of precipitating calcium carbonate from soluble calcium 

salts present in sea-water has overshadowed the primary object of the work. The 

observations so far available are few, and the area they cover too small, to attempt to make 

broad generalizations. However, it can be stated that the very extensive chalky mud flats 

forming in the neighborhood of the Florida Keys are now being precipitated by the action 

of the bacterium calcis on the calcium salts present in solution in sea-water. The 

investigatioti can at most be considered to offer a mere indication of the part played by 

bacterial growth in the metabolism of the sea. To obtain a real insight into the question, it 

would be necessary to make more extensive bacterial and chemical observations in 

tropical, temperate, and arctic waters, to study the bacteriology of other areas where 

calcium carbonate is being precipitated from the sea, and to make further investigations in 

the laboratory into the chemistry of the reactions that can be brought about by various 

species of marine bacteria. 

100. Dustan, P. 1985. Comn~unity structure of reef-building corals in the Florida Keys , Carysfort Reef, 

Key Largo, and Long Key Reef, Dry Tortugas. Atoll Research Rullerin 282-292: 1-29. 

This communication is the result of two parallel studies on the distribution of reef-building 

corals on Carysfort Reef, Key Largo and Long Key Reef, Dry Tortugas. The aim of the 

projects was to characterize the species composition of reef-building corals from the 

northern and southernmost localities of the Keys, and through comparison attempt to 

identify the impact of man on the reefs in the Key Largo area of the northern Florida Keys. 

101. Dustan, P., W. Jaap and J. Halas. 1976. The distribution of members of the Class Sclerospongiae. 

Lerhaia 9, no. 4: 419-20. 

The Sclerospongiae play an important and sometimes major role in the construction and 

infilling of reefs in tropical waters. Modern sclerosponges are limited to dark, quiet, 

sediment-shaded habitats. This study describes the distributions of sclerosponges in the 

Bahamas and the Florida Reef Tract. The sponges were found in the Grand Bahamas. 

After extensive SCUBA diving in Pennekatnp Park and the Dry Tortugas, no 

Sclerospongiac were found. Cold water temperature, or alternatively few, if any larvae to 

colonize the reef tract are possible explanations for the lack of Sclerospongiae in the 

Florida Reef Tract.

102, Edmondson, C. H. 1908. A variety of A~risonerna vitrea. Puper-s Tortugas Laboratory 1: 191. 


Notes are provided on the protozoan, Anisonema. Anisonema vitren (Dujardin) is a 

flagellated protozoan, elongate-oval in form, the anterior end broadly rounded, the 

posterior more acutely rounded. Aniso~tenta vitrea is distinguished from other species of 

the genus by eight furrowed surfaces extending in a slightly spiral manner from one end of 

the body to the other. During the summer of 1906, while working on marine Protozoa at 

the Tortugas, Fia., the author studied a form considered as a variety of the above species 

entitled Anisorlema vitreu (Duj.) var. pentagona. A description of the difference between 

the species and variety is presented 

103. Erseus, C. and M. R. Milligan. 1988. A new Buthydrillus oligochaeta Tubificidae from the eastern 

Gulf of Mexico. Bulleti~? of Marine Scie~ice 42, no. 2: 292-95. 

Bathydrilus natabilis is described from 4-58.5 meter depths off Crystal River and Dry 

Tortugas in the easterii Gulf of Mexico. The species is characterized by large, finely 

pectinate, penial setae in segment I I and entally curved, single-pointed, sperinathecal setae 

in segment 10 which distinguish it from all congeners. 

104. Farfante, 1. P. 1980. A new species of rock shrimp of the Genus Sic),onia penaeoidea, with a key to 

the western Atlantic species. Proceedings of the Biological Society of Washington 93, no. 

3: 771-80. 

Sicyor~ia algae, new species, ranges from Dry Tortugas Islands, Florida, to Surinam. It 

differs from Sic)~o~iia fypica (Boeck, 1864), its closest western Atlantic relative, in 

possessing sublateral carinae on the carapace, and in lacking posterior pleural sulci on the 

first three abdominal somites. Also distinctive are the sharply pointed, mesially directed, 

distomesial projections of the petasme in the male, and in the female the pair of long, 

slender spines on sternite XI and rounded posterolateral processes of the median plate of 

sternite XIII. A key to the western Atlantic species of Sicyonia is supplemented by 

synopses of their geographic and depth ranges which include many extensions of 

previously known limits. 

105. Feinstein, A. A. A. R. Ceurvels R. F. Hutton and E. Snoek. 1955. "Red tide outbreaks off the 

Florida West Coast." Report to the Florida State Board of Cortservatio~~ of Marine 

Loborutrories 

A compilation of reports of red tide on the west coast of Florida from 1844 to January, 

1955 is given. Also included are two working diagrams of incidence of red tide, 

suggesting that red tide occurs more frequently in the months of August through January, 

and that individual red tide outbreaks are part of larger outbreaks, which seem to move 

from south to north, and summer outbreaks appear to originate mostly north of Venice, 

winter and spring outbreaks further south. FulTher data are required to give complete 

support. If this is substantiated, control may be exerted by action in a limited focal area or 

areas of origin. Otherwise, the problem of control may be of the greatest difficulty, since it 

will require action over a much wider area. 

106. Field, R. M. 1919. Investigations regarding the calcium carbonate oozes at Tortugas, and the beach 

rock at Loggerhead Key. Carrzegie lnstitatio~r of Washington, Yeur Book 18: 197-98. 

Calcium carbonate accumulations in the shallow lagoons and channels between the reef 

flats were examined to ascertain their origin. Carbonate ooze hardens rapidly when 

exposed to air and when flooded with saltwater, mud-cracked z,ones can be formed similar 

to those in the geologic record, as in the Stones River limestone formation. An account is 

given on the origin of the "beach rock found between the high and low water marks on 

Loggerhead Key.

1920. Origin of "beach rock" (coquina) at Loggerhead Key, Tortugas (abs.). Bulletin of the 

Geological Society of America 31: 215. 

A study was made to discover the origin of the "beach-rock" or cemented shell-sands 

which occur between high and low tides. By means of a stand-pipe and pump, it was found 

that during heavy rains a shell key acts like a reservoir, and the meteoric water dissolves 

CaCO, on its way through the loose shell sands. The ground water was found to contain 40 

per cent more CaCol in solution, or colloidal suspension, than the normal sea water. This 

concentrated solution of CaCoi has a strong cementing value, and is probably an important 

factor in the formation of the "beach-rock" where the ground water flows out through the 

beach sands, between tides. 

Fisk, E. J. 1976. Black phoebe sighted at Dry Tortugas. Florida Field Naturalist 4, no. 2: 39. 

An observation of a black phoebe on Loggerhead Key, Dry Tortugas on April 13, 1976 is 

recorded. This is the fourth sighting and only spring record for Florida of a black phoebe. 

Gauld, G. 1820. Art accurate chart of the Torfugas and Florida Ke)'s or Martyrs, surveyed by 

George Gauld, A.M. in the years 1773, 1774, 1775. London, W. Faden. 

Flrst nautical chart of the Dry Tortugas is produced. 

Gee, H.. 1934. Lime deposits and the bacteria. I. Estimate of bacterial activity at the Florida Keys. 

Papers Tortugas Laboratory 28: 67-82 (issued Dec. 1932). 

Carnegie lnstitution of Washington Publication Number 435. 

Aerobic organisms were collected from the Florida Keys. Viable counts indicate that open 

areas are only thinly populated with these forms, but that sheltered areas may permit 

increased activity. Conditions in the mud are such as to favor the growth of anaerobes. 

There is a possibility that specific groups, such as the purple sulphur organisms are at work 

in addition to the conventional aerobes. 

Gee, H. and C. B. Feltham. 1934. Lime deposition and the bacteria. 11. Characteristics of aerobic 

bacteria from the Florida Keys. Papers Tortugas Laboratory 28: 83-91 (issued Dec. 1932). 


General bacterial conditions at the Florida Keys during the 1930 season have been 

discussed by Gee (1932). There was reported a collection of 138 representative aerobic 

organisms recovered from the water and mud of Bird Key harbor between Bird and Garden 

Keys, of the Marquesas lagoon, and of one vertical one in the vicinity of the Gulf Stream. 

Preliminary examinations were made of them at the Tortugas laboratory. The strains were 

found to be Gram-negative rods, ammonia-producing, and possibly fermenting. The 

collection was subsequently studied exhaustively at the Scripps Institution during the 

winter of 1930-31. When freshly isolated, these bacteria displayed considerable variation 

in size, in colony features and color, and in their degree of physiological activity. 

Gersh, 1. 1935. Studies on the anterior pituitary gland of the nurse shark. Carnegie li~stitution of 

Washington, Year Book 34: 8 1. 

Experiments were planned on the nurse shark to determine which of the activities of the 

anterior pituitary gland are referable to the eosinophile cells. 

Gilmore, R. G. and R. S. Jones. 1988. Li~)ograrnn~aJlavescens, a new grammid fish from the 

Bahama Islands with descriptive and distributional notes on L. evides and L. anuhantoides. 

Bulletin ofMarine Science 42, no. 3: 435-45. 

In 1981, dredge collections made north of the Dry Tortugas by Continental Shelf 

Associates under contract with the Bureau of Land Mangernent documented the first 

continental record of L. ui~ubanmides.

114. Ginsburg, R. N. 1953. Beach rock in south Florida. Journal of Sedimentary Petrology 23: 89-92. 

The rapid intertidal lithification of beach deposits in the coral seas has received the 

attention of numerous investigators. Study of beach rock from the Dry Tortugas shows that 

the aragonite cement is precipitated from the sea water remaining in the beach sands at low 

tide. High temperatures, rate of beach drainage, and the permanence of the beach control 

the localization of beachrock. The recognition of beachrock in the fossil record is briefly 

discussed. 

115. Goldfarb, A. J. 1913. Changes in concentration of sea-water and their influence upon regeneration. 

Proceedings of the Society for Experimental Biology and Medicine 10, no. 3. 

The regeneration under changed densities of sea water was observed under conditions that 

endured the elimination of uniformity of associated factors such as size of medusae, 

volume, surface and depth of solutions, extent of injury, level of amputation, temperature, 

crowding, aeration, etc. Dilutions were made with water containing a known quantity of sea 

salts, and concentrated solutions were made by slow evaporation, which corrected certain 

errors in previous experiments. Results were compared with those of Loeb. It was found 

that both the hydroid Eudendrium of Woods Hole as well as Cassiopeia of Dry Tortugas 

differed radically from the Loeb experiments, in respect to the range of solutions in which 

animals lived or regenerated, the optimum solutions, the normality of the regenerated parts 

and the character of the curve. It is stated that Loeb's curve probably is limited to 

Tubularia of Naples, and does not represent the behavior of organisms to changes of 

density of sea water, and that the differences in the behavior of these three organisms can 

hardly be correlated with the differences in concentration of the sea water in which they 

normally live. 

116. -- . 1914. Changes in salinity and their effects upon the regeneration of Cassiopea xurnachona. 

Papers Tortugas Labor-atory 6: 83-94. 


Cassiopea xamachana, a large scyphomedusa, is very abundant in the very shallow waters 

of the moat at Fort Jefferson, Dry Tortugas, Florida. The present report considers to what 

extent changes in salinity influence regeneration in Ca.rsiopea, and the results of the 

investigation are compared with those previously obtained with the hydroid Eudendrium 

ramosum of Woods Hole, Massachusetts and with the observations of Loeb with the 

hydroid Tubularia of Serino Bay, Italy. The object of this investigation was to ascertain to 

what extent changes in salinity affected Cassiopea xamachana normally subject to 

relatively great variation in the concentration of the sea-water, and to compare the results 

with those of the hydroid Eudendrium and the hydroid T~~bularia. The following variable 

factors were uniform for the series: size of medusae; volume, surface, and depth of the 

solutions; extent of injury; level of amputation; temperature; crowding. Injurious or other 

variable factors were guarded against. Cassiopea lived in solutions ranging from 40 to 153 

per cent sea-water solutions. Regeneration occurred in solutions containing 50 to 133 per 

cent sea-water. Normal regeneration of the arms occurred within much narrower range, 

namely 75-105 per cent. Beyond these limits regeneration was atypic . 

117. -- . 1918. Effects of aging upon germ cells and upon early development. Part 11. Biological 

Bulletirl 34, no. 6: 372-409. 

In a previous preliminary experiment it was shown that freshly liberated eggs of different 

females of three different species of sea urchins (Toxopneustes and Hipponoe collected 

from the Tortugas, and Arbacia from Woods Hole, Mass.) varied in respect to size, jelly 

layer, membrane formation, and cleavage. In this paper the same technique and the same 

three species of sea urchins were used to determine the physiologic condition of the germ 

cells, and then determine the nature of the changes in the eggs as they became increasingly 

overripe. As eggs in good physiologic condition aged, their volume increased until they

ecame srnaller than the norm. Eggs in poor condition were reduced in size., in all three 

species, there was a loss in jelly layer with age, depending on the condition of the egg. In 

all three species, as the eggs aged, tlie membrane appeared closer to the surface, becoming 

thinner until none was formed. The rate of decrease in cleavage with age was greater in 

Toxopneusre.~ arid Nijpnoe than in Arhacia. Overall, the change in size, jelly membrane, 

and cleavage with aging of germ cells are accurate, convenient and corroborative indices 

of physio-chemical and ~norphologi changes in the eggs as they age, and afford convenient 

measures of loss in vitality, or physical deterioration. 

1 1 8. ------ . 1914. Experimentally fused larvae of echinodernls with special reference to their skeletons. 

Papers Torruga.! Laboratory 6: 103-21. 


The early work of Loeb, Morgan, and llerbst on the production of multiple embryos from a 

single egg suggested the reverse experiment of grafting or reuniting several fertilized eggs 

into on embryo. In 1912, the writer repeated these experiments with tlie American form 

Arbacin punatulara and succeeded only after slightly modifying Driesch's method. 

Subsequently, in tlie performance of other experiments, it was discovered that eggs could 

be agglutinated and fused quite as readily by a very different method, which was not only 

simpler but free of certain objections that might be urged against previously known 

methods. The new method consisted in using an isotonic or slightly hypotonic NaCl 

solution diluted with varying quantities of sea-water. 

119. -- . 1913. The influence of the central nervous system in regeneration of an annelid worm. 

PI-oceedirigs of the Societ)'for Experimental Biology and Medicine 10, no. 3. np. 

(No abstract availablc). 

120. ------- 1914. Regeneration in the annelid worm Amphinson~a pacrfica, after removal of the central 

nervous system. 1'uper.s Tortugas Laboratory 6: 95.102. 


In a previous publication, tlie writer found that the head of the earth-worm Lumbricus was 

regenerated in the entire and permanent absence of the nerve-cord from the amputated 

region. The marine annelid worm Atnphblorna pacifica readily regenerated a head at all 

levels except the distal eighth of the worm. Regeneration may be prevented by a severe 

injury, either to the digestive tract or to the central nerve system; the greater the injury the 

more likely will regeneration be inhibited. Many pieces did not regenerate after removing 

the alimentary tract from five or more segments nearest the amputated level. Many pieces, 

about one-third, failed to regenerate after removing the nerve-cord by the forceps, i.e., with 

little injury to adjoining tissues. All failed to regenerate after removing tlie nerve-cord by 

the "window" method. The operated worms were examined in serial sections. In one group 

a regenerated nerve-cord connected the regenerated "brain" and commissures with the old 

intact nerve-cord. In a second group the regenerated nerve-cord approached and in 

instances reached the amputated level, yet no head was formed. In a third group, the nervecord 

liad regenerated, but several segments nearest the amputated end were yet without any 

nerve-cord or ganglia. These worms nevertheless had regenerated a head with its typical 

brain and nerve-cornmissures. 

121. -. 1917. Variability of eggs and sperm of sea-urchins. I'uper:! Tortuga.~ Lnborato,y 11: 71-87. 


A clear uoderstanding of the variability in normal fresh eggs and sperm is necessary in 

order to appreciate and to evaluate the changes that take place in oversipe germ-cells. This 

paper deals exclusively with the qualitative and quantitative differences of such freshly 

collected sea-urchin eggs and sperm arid with the differences in their early development.

122. ----. 1917. Variability of germ cells of sea urchins. Proceedi~~gs of the Natior7al Academy of 

Science 3: 241 -45. 

Three different species of sea urchins (Toxopneustes and Nippo~zoe collected at the Dry 

Tortugas, Arbacia collected at Woods Hole, Massachusetts) were used to determine the 

normal variability of sea urchin germ cells. Having determined the optimum and constant 

conditions of germ cells, studies were conducted to examine variations in size and shape of 

eggs, the jelly layer of eggs, membrane formation, and cleavage among tlie three species. 

Amazingly large variations were found in fresh germ cells among species, thus suggesting 

that anlong other investigators of the varying behavior of thc eggs, a large part of the 

variation was probably due to the physiologic conditions of the eggs which these 

investigators used. 

123. Goodrich, H. B. 1935. Color patterns in fish. Carnegie Ir~stitution of Washingron, Year Book 34: 81. 

Studies were carried out to investigate internal conditions which may control the 

development and maintenance of color patterns in fish by transplanting scales and tissues 

from one type of pigment area to another. 

124. Gordon, M. 1933. The internal pigment systems of fishes. Cnmegie 111stitution of Washington, Year 

Rook 32: 268. 

The internal pigmentary systems of major taxonomic groups were examined. Halichores 

bivittatus and Lutjanus griserrs showing possibly neoplastic growths were collected for 

study. 

125. Goy, 3. W. 1982. West Indian Stenopodidae. 2. Occurrence of Kichardbla spi~iicb~cra Crustacea, 

Decapoda, Stenopodidea off the Dry Tortugas. Blclletii7 of Marine Science 32, no. 1: 344- 

47. 

An examination of Kichardi~irr spir~icirlcta collected by W.L.Schrnitt in August of 1932 is 

made. It is concluded that this specimen is truly K. rpbiici~icta, that this is the sixth known 

specimen of the species, and the first record of the genus in the Western Atlantic. The 

occurrence suggests that the genus occurs at sllallower depths than those recorded in 

previous literature. 

126. Grave, C. 1934. Tlie Rotryllras Type of Ascidian larva. Paprs Tortups Laboratory 28: 141-56 

(issued Dec. 1932). 


R-ee-swimming larvae of at least three well-defined types arc found in life cycles of 

ascidians: one, characteristic of species of Molgraln (Crave '26) and related genera, that 

has one sense organ only, a statolith, in its sensory vesicle. The nerve cord lies in a middorsal 

position above the notochord, the caudal fin is expanded vertically in the median 

plane and adhesive papillae are lacking. In the text the structural organization of a type of 

larva characteristic of species of Rotr)'llus and related genera is described. The body is egg 

shaped, its depth being approximately the same as its width Three conical sensory 

papillae arranged in the form of an equilateral triangle a]-e borne at the anterior smaller end 

of the body, two located on either side of the median plane dorsal to the central body axis, 

one in tlie median plane ventral to the central axis. The same gross parts found in the 

central nervous system of larvae of other types are present. The anterior end of the visceral 

ganglion bends to the right and expands to form the sensory vesicle, which, in contrast with 

that of larvae of other types, does not project to the level of the dorsal surface of the body 

but retains an interior position relatively far below the surface. 

127. - . 1936. Metamorphosis of ascidian larvae. Paperx Tortuga,s idmratory. 29: 209-91 (~ssued 

Dec. 1935). 

Carnegie Institution of Washington Publication Number 452.

The studies of metamorphosis of larvae of ascidians were made during the summers of 

1927, 1930, and I933 at the Tortugas Laboratory with the purpose of finding methods of 

accelerating and controlling metamorphosis and thus of discovering something of thc 

fundamental nature of the internal mechanism involved and the environmental conditions 

with which it is causally related. The observations made in the course of this investigation 

are interpreted as follows: The ascidian larva is a dual organism, the action system of the 

larva'being quite separate from the action system of the ascidiozooid. Metamorphosis 

advances by three stages; (a) changes in the adaptive responses of the larva to light and 

gravity; (b) the attachment of the larva to the surface of some foreign object; (c) the 

disruptive phase during which the entire larval action system is destroyed. Swimming 

activity causes the production and concentration of some metabolic product in the larval 

tissues that is essential to the induction of n~etamorphosis. The presence in the larval 

tissues of metabolic products resulting from swimming is not alone sufficient to induce 

metamorphosis, but another substance with which this metabolic product may react is 

equally necessary. The great variability of ascidian larvae of the same species in the 

duration of their free-swimming period is apparently due to variability in the time of 

formation of the susceptibility substance and hence to the time of differentiation of the 

larval organ that produces it. Metamorphosis may be induced artificially by diverse 

chemical and biological substances placed in sea-water with groups of larvae in lactic acid. 

Metamorphosis is rapidly and consistently induced in the larva of Phallusia nigra. The 

activating agents extracted from the fresh ascidian tissues that were so specific in their 

effects may also be endosymes of a highly speciali7.ed kind, each found only in a single 

species of ascidian. The mechanism of metamorphosis is comparable in its organization to 

that of development of an egg, which also may be activated by numerous and diverse 

chemical and physical agencies. 

128. Grave, C. and P. A. Nicoll. 1940. Studies of larval life and metamorphosis in Ascidia nigra and 

species of Polyandrocarpa. Papen Tortugas Laboratory 32: 1-46 (issued Oct. 1939). 


Experimental studies made during the summer of 1933 (Grave, 1936) demonstrate that seawater 

extract of pharyngeal, atrial, or mantle tissues of adult Ascidia rzigra is effective in 

inducing 100 per cent metamorphosis in groups of Ascidia larvae within 3 hours after 

hatching and that similar extract of tissues of Polyandrocarpa induces 100 per cent 

metamorphosis in groups of Polyandrocarpa larvae within 42 minutes after liberation 

from the parent colony. These observations led to a search during the summers of 1935 

and 1936 for a specific chemical substance in the tissues of these ascidians having the 

properties required for the rapid acceleration of the process of metamorphosis. An account 

of the methods and results of this work is given in this paper. It was found that the amino 

acids l-histidine, leucine, glycine, cysteine, and d,l-alanine, in the form received from the 

laboratories in which they were prepared, accelerated metamorphosis in groups of larvae of 

both types. A sea-water extract of free-swimming larvae or of late embryonic stages of 

Ascidia has the same accelerating effect on metamorphosis of Ascidia larvae as an extract 

made from tissues of the adult ascidian. Heating adult Ascidia tissue or releasing distilledwater 

extracts of the tissue for several hours does not destroy the accelerating substance. 

Non-toxic concentrations of copper, iron, and aluminum salts induce early metamorphosis 

to a marked degree. The duration of the free-swimming period of Ascidia larvae is longest 

at the beginning of the breeding season of the species and becomes gradually shorter as the 

season advances. 

129. Gudger, E. W. 1921. Notes on the morphology and habits of the nurse shark, Ginglymostrorna 

cirratuni. Copeia 98: 57-59. 

A physical description of the nurse shark as observed by the author for several summers in 

the southern Florida Keys and Dry Tortugas is given.

130. -. 1929. On the morphology, coloration, and behavior of seventy teleostean fishes of 

Tortugas, Florida. Papers Tortugas Laborator)' 26: 149-204. 


In the course of work at the Tortugas, 70 teleosts, belonging to 28 families, have been 

studied. Habits have been recorded herein that stand out prominently to the taxonomist. 

First, basing thc classification of tropical fishes on coloration is a very dangerous thing. 

Most of the fishes in Tortugas have from two to five color phases in life and, even when 

studying the fish in a state of comparative quiet in an aquarium, it is very difficult to 

determine which is its normal color. When a fish dies, its color changes either entirely or in 

its intensity, so that the coloration of the dead fish is markedly different from that of the 

live fish. It is equally dangerous to describe and classify a tropical fish from a single 

specimen, since these fishes are so very variable in the number of fin rays, in the relative 

proportions of the body, in scale count, and in the many details which help to distinguish 

one species from another.. 

131. -. 1918. On the use of the diving helmet in submarine biological work. Atnericart Museuni 

Journal 18: 135-38. 

The use of the diving helmet for research at the Dry Tortugas was initiated in 1915 by 

Longley and Carey, for fish observations and photography. Its use was declared new for 

underwater work. However, such is not the case. The use of the helmet alone replaced 

cumbersome diving suits (scaphanders) used by the commercial spongers out of Tarpon 

Springs, Florida, and early workers on the construction of the overseas railroad, The 

Florida East Coast Railway Extension from Homestead to Key West, Florida. The diving 

helmet in biological workdates back to around 1845, when M. Milne-Edwards conducted 

bottom surveys off the coast of Sicily. In 1679, pressurized air was first supplied to 

Borelli, who attached a simple air compressing pump to a leather diving helmet. These 

devices are all refinements of the crude diving lielmets used back in ancient times by 

Alexander the Great, while recording plant and animal observations. These are some of the 

earliest underwater biological observations ever recorded. The earliest account of any type 

of diving apparatus is found in Aristotle and dates back to about 1000 B.C. 

132. ---- . 1918. Spltyrae~ia barracuda; Its morphology, habits, and history. Papers Tortugas 



This article provides a general description of the great barracuda, Splzyraena bnrt-acuda 

made at the Tortugas Marine Laboratory, based on local collections and an examination of 

12 large individuals using lengthlweight measurements, color and markings, jaws and teeth. 

internal organs, foods and feeding, and manners of breathing. Additional information is 

presented on their habits, how they may be caught, and parasites. An interesting historical 

side of the paper compiled from around the world includes verbatim quotes and 

descriptions of their great size, ferocity, fossil forms, nomenclature, habitats, and food 

poisoning in man. Accounts of their poisonous flesh in tile West Indies date as far back as 

1667. Largest sizes of West Indies individuals approach 8-10 feet in length, with some 

highly "dubious" reports of specimens reaching sizes of 18-20 feet in length. 

133. - 1913. Uterine gestation in the nurse shark, Gittgl)'mosrutila cirrarutn. Journal of the Elisha 

Mitchell Scientific Society 29: 8. 

Also, in Science, 1913, v.37, p.993. 

The breeding habits and embryology of this shark were studied at the Tortugas Laboratory 

in the summer of 1912. A brief account was published in the Year Book for 1912, p. 148- 

150.

134. Halley, R. B. and R. P. Steinen. 1979. Groundwater observations on small carbonate islands of 

southern Florida. In Guide to sedinier~rario~~ for the Dr): Tortugas. Compiler R. B. Halley, 

p. 82-89. Tallahassee, Florida: South East Geological Society Publication. 

Observations are reported on the unusual hydrology of Loggerhead Key, a sandy key in the 

Dry Tortugas in comparison with observations on Cluett Key, a mud key which lies 200 km 

NE of Loggerhead in western Florida Bay. The ground water of Loggerhead and Cluett 

Keys differs significantly from the surrounding sea water, despite the relatively small size 

of the island. Climate alone does not determine the character of these ground waters; for 

example, Loggerhead Key is underlain by less saline ground water than Cluett Key despite 

the fact that it receives less rainfall. Ground water under such small islands such as these is 

formed from topography, sediment character, vegetation, and many more parameters that 

are themselves interrelated. They conspire to form ground water that not only differs from 

sea water, but also can react with the island sediments to change the character of the 

ground water. In this manner, island ground waters serve as geologic agents, hastening the 

alteration of marine carbonate sediments to limestone and dolomite. 

135. Hanlon, R. T. and R. F. Hixon. 1986. Behavioral associations of coral reef fishes with the seaanemone 

Co~~dylacris gigar~tea in the Dry Tortugas, Florida USA. Brrllerin of Marble 

Science 39, no. 1: 130- 134. 

Over 30 small West Indian reef fishes dwell within the tentacular sphere of anemones, 

mainly to avoid predation. Most species swim carefully to avoid the stinging tentacles, but 

some species also have a physiological adaptation (skin mucus alteration) that allows them 

to be in full and vigorous contact with the tentacles in a manner similar to Indo-Pacific 

anemonefishes such as Amnm~iprion, Da,sc~llu.s and Prem~ras. The authors report herein six 

species of reef fishes that are facultative associates of the sea anemone Co1zdy1acti.s 

gigantea (Weinland) in the Dry Tortugas Islands. The fishes were not found associated 

with other anemones. One species, Lnhriso~nra.~ gol~io, is a new record of a fish with both 

the behavioral and physiological adaptations to dwell unharmed among the stinging 

tentacles of Condylactis gigm~tea. 

136. Hargitt, C. W. 191 1. Cradactis ~,ariabilis: An apparently new Tortugan Actinian. Papers Tortugas 

Lnborator): 3: 49-53. 


The author believes this species of actinian has never before been described, and names it 

variabilis. The specimens seem to have the capacity to move about more or less freely, 

and the frond-like organs situated about the margin of the oral disk and outside the outer 

cycloe of tentacles aid in such movement. The color is pale olivaceous-green to brownish; 

tentacles somewhat lighter; foliose organs darker, even brownish, with flake-white pads. 

The body is highly contractile, with a weak or diffused sphincter. The reproductive season 

seems to be in the spring and early summer. The habitat is chiefly in holes, crevices, or 

similar secluded places in the coral reefs or about the shoals where protection is available. 

137. Harrington, B. A. and J. J. Dinsmore. 1975. Mortality of transient cattle egrets at Dry Tortugas, 

Florida. RidBariding 46, no. 1: 7-14, 

This article examines the idea presented by Browder (1971) that cattle egrets pass through 

the Dry Tortugas with seasonal regularity, and that large numbers die after landing. This 

study concludes that regular spring movement occurs with niany egrets stopping at the 

island, and that niany of the egrets that stopped apparently died from starvation, especially 

in late June and in early July. The mortality in 1968 was higher that in 1970. 

138. Harris, .I. E. 1937. The mechanical significance of the position and movements of the paired fins in 

the Teleosti. Papers Torrugas Lnborafory 31: 171.89 (issued Oct. 1936). 


In the course of the evolution of the modern teleostean fish, a series of fairly well-defined 

changes has taken place in the body form and in the shape and position of the fins. The 

present paper discusses the mechanical factors concerned in the evolution of the teleost 

type of fish. A comparison of this type with the dogfish suggests that the development of 

an air bladder has been the primary factor involved in the change in general body form. 

The reduction in specific gravity of the fish, consequent upon this primary change, has 

removed the need for a lift force on the body during free swimming. The asymmetrical 

(heterocercal) tail has therefore disappeared. For the same reason, the pectoral fins are no 

longer needed as elevating planes, and become free to move up toward the mid line of the 

body to act as brakes in stopping and turning movements. The forward motion of the pelvic 

fins is a mechanism for producing a balanced vertical force and a balanced pitching 

moment. These fins are normally used in conjunction with the pectorals. The independent 

movements of the pectoral fins are then discussed. All types of movement so far observed 

are variations on a fundamental form, in which the metrachronal oscillation of the fin rays 

generates an undulating fin surface. The observed variations in form of the fin beat can be 

produced by varying the phase difference between the beat of successive rays, and also by 

making the oscillation of the fin ray asymmetrical. The characteristics of the pectoral 

musculature associated with such variations are pointed out, and illustrated by reference to 

a number of fish types. 

139. Hartman, C. G. 193 I. The hypophysis of fishes. Carnegie Ir~sritutior~ of Washington, Year Book 30: 

381-82. 

Studies on the influence of the hypophysis on menstruation and various forms of uterine 

bleeding in sharks were carried out. 

140. Hartmeyer, R. 191 1. Polycistor (Eudistoma) tna)reri nov. sp. from the Tortugas. Papers Tortugas 

Laborarory 3: 89-93. 


A new species Polycitor (Eudisrotna) ma)'eri, a new ascidian collected in 1907 at the 

Tortugas is described as the largest and most beautiful ascidian of the Tortugas. It was 

collected in the deeper water of the Southwest Channel near Loggerhead Key, on sandy 

bottoms, where it is abundant. The color is pale yellow, with a reddish or violet tint. From 

the western Atlantic only five species of this genus have been described, and all of these 

are mentioned by Van Name from the Bermudas, but all these species have four rows of 

stigmata in the branchial sac and are in many other respects quite different from this 

species. 

141. ---- . 1908. Reisebilder aus Westinidien mit besonderer Berucksichtigung der korallenbildungen. 

Deutsch. Gessel. Fur Volkstutnlich Natuir-kunde or Sattw Title in Meet-eskunde Jahrg. 3, 

Heft 2, 40 Pp 3, no. 2: 1-40. (No abstract available) 

142. Harvey, E. N.. 191 1. Effect of different temperatures on the medusae, Cassiopea, with special 

reference to the rate of conduction of the nerve impulse. Paper.s Torrugas Laboratory 3: 

27-39. 


During the sumnier of 1909 a study was made of the effects of water temperatures on the 

nerves and muscle tissue of Cassiopea. Temperatures in the moat at Fort Jefferson ranged 

from 27°C to approximately 32-33°C. Activity limits and thermal death points of nerve and 

muscle were measured. It was found that ncrve conduction rates fall off in rate with rise of 

temperature to a definite maximum, similar to that for enzyme action and for other life 

processes.

143. ------ . 1914. The relation between the rate of penetration of marine tissues by alkali and the 

change in functional activity induced by the alkali. Papers Torrugas Laboratory 6: 131-46. 


The present study, made at Tortugas in the summer of 191 1, is a continuation of 

permeability investigations undertaken at Columbia University in 1910 to 191 1. The 

author's aim has been twofold. First, to compare the permeability of the cells and tissues of 

salt-water organisms with those of fresh-water forms. Second, to determine the relation 

between the rate of penelration of the alkali and the appearance of structural or functional 

changes in the cell. The author thinks that the presence of a sufficient number of OH ions 

within the egg may aid in breaking down the granules and that this breaking down increases 

also the degree of swelling of the egg. Cytolysis in Holorhuris appears to be largely of this 

type, since NaOH enters before the increase in volume begins. From this point of view 

both theories of cytolysis contain an element of truth. Swelling of marine eggs is due both 

to an increase in permeability of the surface and also to the breakdown of lipoid or protein 

granules within. The latter tends to increase the swelling pressure or the osmotic pressure 

of the egg, but is secondary to the increase in permeability of the surface. 

144. ------ . 1921. Studies on bioluminescence XIII: Luminescence in the Coelenterates. Biological 

Bulletin 61: 280-287. 

(No abstract available). 

145. ------. 1923. Studies on bioluminescence. XV. Electroreproduction of oxyluciferin. Journal of 

Ge~~eral Phjsiology 5: 275-84. 

This work was on the light-producing reaction in the luniinous crustacean, Cypridina. 

Oxyluciferin may he reduced to luciferin at cathodes when an electric current is passed 

through the solution, or at cathodes formed by metal couples in solution, or at cathodes of 

oxidation-reducation cells of the NaCl - Pt - Na2S type. It is also reduced at those metal 

surfaces (Al, Mn, Zn and Cd) which liberate nascent hydrogen from water, although no 

visible hydrogen gas separates from the surface. Molecular hydrogen does not reduce 

oxyluciferin even though very finely divided, but will reduce oxyluciferin in contact with 

palladium. Palladium has no reducing action except in the presence of hydrogen, and 

apparently acts as a catalyst by virtue of some power of converting molecular into atomic 

hydrogen. Conditions are described under which a continuous luminescence of luciferin 

can be obtained. This luminescence may be used as a test for atomic hydrogen. It is 

suggested that the steady luminescence of bacteria is due to continuous oxidation of 

luciferin to oxyluciferin and reduction of oxyluciferin to luciferin in different parts of the 

bacterial cell. 

146. Hatai, S. 1916. Changes in the chemical composition of starving Cussiopea xamachana. Ccrrnegie 

lr~stitution of Washingion, Year Book 15: 206-7. 

Studies were conducted on chemical changes occurring in Cassiopea during starvation. 

The constancy of water content suggests that Cassiopea is largely a jelly-like mass, and 

remains so throughout its life. In contrast, mammalian body-water content varies by age. 

147. ------- 19 17. On the conlposition of Cassiopea xamachana and the changes in it after 

starvation. Papers Torrugas Lnboraroq l I: 95-109. 


For this study eight freshly caught normal Cassiopea, having different body weights, were 

subjected to starvation by placing the animal in filtered sea water. The results were as 

follows: 1. In general the smaller Cassiopea loses relatively more weight than the larger. 2. 

The percentage of water found in the entire body is nearly the same in all sizes of 

Cussiopea. However, the values of water content in the starved appear to be slightly higher 

than those found in the normal Cassiopea. 3. The nitrogen content of the entire body is

higher in the smaller than in the larger Cassiopea. 4. The absolute amount of nitrogen 

found in the starved Cassiopea is considerably higher than in the normal having the same 

body weight. It was noted that although high when compared with the normal, equal in 

weight to the starved animal, it is very low for the initial body weight of the starved animal. 

This shows that the nitrogen also has been consumed during the period of starvation. 5. 

The nitrogen contents for the different parts of the body are similar in their relations to 

those found in the normal Cassiopea. 6. The loss in weight of the different parts is of such 

a character that their proportions in the starved remain similar to those in the normal 

Cassiopea. 

148. - 1917. On the composition of the medusa Cussiopea xa~nachama. Proceedir~gs of the 

National Academy of Scier~ce 3: 22-24. 

In this study, an examination was made of three different parts of Cassiopea, mouthorgans, 

umbrella, and velar margin to determine whether starving specimens lose weight 

uniformly, or whether the loss is dissimilar in the three parts. Results indicated that the 

smaller Cassiopea loses relatively more weight than does the larger Cassiopea. The 

percentage of water is similar through the entire body, the nitrogen content is higher in the 

smaller than the larger individuals, and nitlogen is much higher in the starved Cassiopea 

than in the normal specimen with the same body weight. Results are compared with 

Mayer's experiments, which showed nitrogen loss to be constant during the entire period of 

starvation. Differences may be due to the size of animals used in his studies, as larger 

individuals show little variation in nitrogen loss, whereas small Cassiopeas show large 

variations in nitrogen loss due to body size. 

149. Hayes, F. R. 1932. Nitrogen in echinoid ontogeny. Carnegie lrlstituiion of Washington, Year Book 

31: 284-85. 

The chemical embryology of the echinoid egg was investigated, as well as variations in two 

sources of energy available in the egg: protein and lipins. 

150. -- . 1934. Variation in size and in nitrogen requirements during early development of the seaurchin, 

Echinomrera lacumer. Papers Tortugas Laboratory 28: 181-93 (issued Mar. 1933). 


After the penetration of a spermatozoon, the developing echinoderm egg receives nothing 

from the outside except water and salts, until the comparatively advanced larva begins to 

eat. The morphological phenomena of ontogeriy can be brought about only by the 

expenditure of energy, which must come from materials already present in the egg at the 

time of fertilization. The problems of chemical embryology include (a) a determination of 

the amount of energy required to produce structural changes, and (b) an investigation of the 

chemical transformations taking place. The work here reported deals with a certain phase 

of the chemical embryology of a common tropical sea-urchin, Echinomerra lacunter. 

Studies of the first 24 hours of development of the eggs of this form were carried on during 

the summer of 1932 at the Tortugas Marine Station of the Carnegie Institution. Eggs of the 

sea-urchin, Echinometra lacunte~; were concentrated with a hand centrifuge and then 

diluted with 500 times their volunie. Analyses of primary amino nitrogen groups and of 

total nitrogen were made, and the ratio of the former to the latter calculated. From 4 hours 

onward the ratio of primary amino groups to total !nitrogen increases. This does not mean, 

however, a synthesis of the former at the expense of the latter, but rather that in the 

combustion which provides the developing embryo with energy, some source of 

nitrogenous fuel other than NH2groups is being used. There is a marked loss in the 

quantity of nitrogen per egg during the period of development succeeding the first four 

hours. One million eggs contain some 13 milligrams of nitrogen, of which about 28 per 

cent is in the form of NH2 groups.

151. Heard, R. W. and D. G. Perlmutter. 1977. Description of Colomasrixjuniceae, new species. A 

commensal amphipod (Gammaridea Colomastigidae) from the Florida Keys, U.S.A. 

Proceedings of the Biological Societ)' of Washington 90, no. 1: 30-42. 

During November of 1968 and 1973 and June of 1970 more than 100 specimens of an 

undescribed commensal amphipod belonging to the genus Colon~astix Grube, 1861 were 

collected from loggerhead sponges, Spheciospongia vespuria (Lamarck), in the lower 

Florida Keys. Additional specimens of this new species, collected from Dry Tortugas, 

Florida were borrowed from the Division of Crustacea of the U.S. National Museum of 

Natural History for examination. 

152. Helwig, E. R. 1933. Regeneration in lotrocl~ota birotulata (Porifera). Carnegie institution of 

Washington, Yeur Book 32: 271-73. 

The development and formation of cells over time was examined, from cross-sections made 

from the branches of the sponge. lotrochora bii-otulara. 

153. Hendee, E. C. 1931. Formed components and fertilization in egg of the sea-urchin Lytechinus 

variegatus. Papers Tortugas Laborator); 27: 99-105. 


This investigation of the eggs of Lytechims variegatus collected during the summer of 

1925 at the Tortugas was undertaken to determine if any substance of the egg was it~volved 

in fertilization. Certain cytoplamic substances (macrosomes, hyaloplasm, chondriosomes, 

fat droplets, and extra-nuclear basophilic granules) were demonstrated both before and 

after fertilization. Lipid granules, present in the mature unfertilized egg, disappeared upon 

fertilization. 

154. Hendrix, S. A. and R S. Britman. 1995. NOx variation in the southeastern Gulf of Mexico, Florida 

Scieiltisr 58, no. 3: 292-97. 

An automated system capable of providing speciation and concentration information for 

several atmospheric NOx compounds was used to obtain diurnal and location variation data 

during a five-day research cruise in the southeastern Gulf of Mexico approximately one 

mile west of Fort Jefferson, Dry Tortugas between May 18 and May 22, 1987. Speciation 

of these nitrogen compounds was achieved by selective preconcentration onto a series of 

chemically coated glass hollow tubes. Analysis was performed by thermally desorbing the 

collected analytes into a chemilun~inescence detector providing sub parts-per-billion level 

determination. 

155. Hess, W. N. 1937. Reactions to light in Pr)chodem hahmnensis. Papers Tortugas Luborato,)r 31: 

77-86 (issued Aug. 1936). 


Little attention has been given to the study of light reactions in any of the Enteropneusta, 

and nothing is known, apparently, concerning the distribution or even the existence of 

photoreceptors in this important group of animals. The purpose of this investigation was to 

continue work on reactions to light and the photoreceptors in animals, using at this time a 

more highly evolved species than the earth-worm on which the earlier work was done. 

During the study, Pt)choderu hahen~ensis responded negatively to ordinary intensities of 

light. The movements of Pt)'chodera, when exposed to light were slow and deliberate and 

there was little evidence of trial and error movements. The entire surface of the body was 

sensitive to light, the most sensitive regions being on the proboscis and collar. Removal of 

different parts of the body involving the central nervous system caused little if any decrease 

in the percentage of negative responses to light. The reaction time of the proboscis was 

greatly increased when it was removed from the rest of the animal. This is taken to 

indicate that the central nervous system fu~~ctions to speed up responses greatly, but is not 

essential for responses. Removal of the proboscis together with the basal peduncle makes

it impossible for the animal to orient when stimulated by light. This would seem to suggest 

that the peduncle contains a coordinating center for certain bodily movements, or that the 

animal has been rendered incapable of orienting, due to removal of that portion of the body 

containing most of the notochord. 

156. -. 1940. Regional photosensitivity and photoreceptors of Crangon arn1illatus and the spiny 

lobster, Par~ulirus argrrs. Papers Tortugas Lrrboratory 32: 153-61 (issued Sept. 1940). 


Crayfish from which the eyes have been removed are sensitive to light in the region of the 

sixth abdominal segment, but no responses occurred when other regions were illuminated. 

Tile discovery that freshly molted Crangori ar~i~illatus are sensitive to light in other regions 

of their bodies, in addition to the sixth abdominal segment, led to this investigation. Results 

of this study indicated that Crangon nrmillufus is usually sensitive to light in many regions 

of its body, irrespective of how much time has elapsed since the last molting period. 

Freshly molted spiny lobsters (Panirlirirs argusj are sensitive to light in many regions of 

their bodies. Old spiny lobsters, with hard exoskeletons, from which the eyes have been 

removed are usually not sensitive to light of the intensity used in these experiments. The 

margins of the uropods of freshly molted Crangon nrmi1lafu.r and spiny lobsters are not 

sensitive to light, but the basal two-thirds of these appendages are sensitive to light. Adult 

Crarigorl armillurus and recently molted spiny lobsters react when illuminated from above 

after the sixth abdominal ganglion has been shielded by black cardboard and also after the 

ventral nerve cord has been cut between the fifth and sixth abdominal segments. This 

shows that photosensitivity in theseeyeless animals is not limited to the sixth abdominal 

ganglion. Newly hatched Crangon arrnillatus with normal eyes swim toward the light with 

their caudal ends foremost irrespective of the number of abdominal segments that have 

been removed. Crangon arrnillatus and spiny lobsters from which the eyes have been 

rcnioved do not usually orient to light, but respond by random movements. When their 

bodies are heavily pigmented, or if they are in poor physical condition, they do not respond 

at all. However, if they do respond their responses are usually much slower than those of 

freshly molted animals in good physical condition. The sixth abdominal segment of these 

eyeless spiny lobsters and crayfishes is the most photosensitive region of their bodies. 

However, in Crangon aunillatus and the American lobster Hoinarus arnericanus all the 

abdominal segments appear to be equally sensitive to light. On the basis of regional 

photosensitivity of the uropods, it seems probable that the cell bodies of the neurons which 

connect with the peripheral spines are sensitive to light and hence function as 

photoreceptors. 

157. Hoffman, W., and Jr. and P. C. Patty W. B. Robertson. 1979. Short-eared owl on Bush Key, Dry 

Tortugas, Florida. Florida Field Natriralist 7, no. 2: 29-30. 

The short-eared owl (Asioflamn~eus) is an uncommon but regular winter visitor to Florida. 

This record represents the second summer record of Asioflantrneus in Florida, and the first 

record for the Dry Tortugas. The authors suggested tliat the bird in question had been on 

Bush Key for some time, subsisting on the abundant tern chicks. 

158. Holmes, C. W. 1984. Carbonate fans in the Florida Straits. Society of Econonlic Paleontologists and 

Mineralogists At~iiual Meeting (Abstracts) 1: 39. 

No abstract available. 

159. Hooker, D. 191 1. Certain reactions to color in tlie young loggerhead turtle. Papers Tortugas 

Lrrborator): 3: 69-76 and illustrations. 


During the summer of 1907 observations and a series of experiments were made on the 

habits and early life history of young loggerhead turtles, which identified reactions to color

and geotropism as the determining factors for the causes of young hatchlings to reach the 

water. Based on daylnight experiments on Loggerhead Key, hatchlings did not orient 

towards the sun or the odor of the water, but exhibited positive phototropism by responding 

to large surfaces of light of low intensity. After entering the water, the animal swam out to 

sea apparently attracted by the darker blue of the deeper water. Young turtles displayed 

positive geotropism when all possible negative geotropic reactions had been exhausted. 

160. Hopkins, D. L. Locomotionlphysiology of marine amoebae. Carnegie lr~sfirutiorz of Washington, 

Year Rook. : 

1929,v.28,286-288: 1930, v.29,335-337. 

The chemical and physical factors in the locomotion of marine amoebae collected from the 

tidal pools at Tortugas and cultured in the laboratory were examined. Relationships 

between sea-water salt and locomotion were determined by concentration and dilution. 

Highest rates of locomotion were found in normal sea-water and could be a useful criterion 

in classification and determining physiological condition in amoebae. 

161. Jaap, W. C. 1985. An epidemic zooxanthellae expulsion during 1983 in the Lower Florida Keys 

coral reefs: hyperthermic etiology. Proceedings of the Fifth Internutior~al Coral Reef 

Symposium , 143-48. Moorea, French Polynesia: Antenne Museum-Eph6. 

Extensive reef coral zooxanthellae expulsion occurred from Key Largo to Dry Tortugas, 

Florida, during September 1983. Coral bleaching was intensive between Pelican Shoal and 

Sand Key Reef off Key West. Coral discoloration extended to depths exceeding 14 m but 

was especially severe in shallow (1-2 m) spur and groove habitats. Approximately 75.95% 

of all Millepora cotnplanata and Pa1)dtou caribaeorum were bone white, but most 

colonies remained viable. Affected M. complanata (bladed fire coral) retained the ability 

to inflict pain from dactylzooid nernatocysts. Some individuals (5 to 10%) had fine algal 

growth indicating death on all or parts of their skeletons. Although 15 species of 

cnidarians, principally Scleractinia, were affected, some species (Madracis mirabilis, 

Porites porites, Montastra~ cavernosa, Dendrogyru c)~lindrus ) appeared to be immune. 

A quantitative sample at Eastern Sambo Reef on 6 October documented 11 species and 209 

colonies; M. cor~~planuta comprised 32.5% of all colonies. Transmission 

eletromicrographs did not reveal epidemic pathogenic organisms in affected coral tissues. 

Warm, calm weather prior to the expulsion was conducive to elevated seawater 

temperature. A seawater thermograph deployed off Marquesas Key recorded temperatures 

of 32.3 degrees C. during the period. 

162. ---- , 1980. Stony coral community structure at Long Key Reef, Ft. Jefferson National 

Monument, Dry Tortugas, Florida. (abs.). Florida Scientist 43 (Suppl. 1). 

Stony coral populations at Long Key Reef were studied during summers of 197.5-76 under 

National Park Service sponsorship. Plotless line transects (13, 25 m L ) were sampled in 

depths of 0.5-21.3 m. Abundance, cover, and diversity were greatest in depths greater than 

8 m. Of 34 species encountered, only 23 were censused quantitatively. Montastraea 

atinularis contributed 20% of all colonies and 37% of cover. Species richness was highest 

(I 1) on transects in 7.6-12.5 m deptlis. Shannon-Weiner diversity values H' log SUB-2 

computed by transects for individual colonies ranged from 1 .O-3.0. Pielou's eveness (J') 

values ranged from 0.36.1.00. Community relationships based on Morisita index values 

detected an assemblage dominated by M. arznularis in 8-13 m and a M. cavernosa 

community in 18-21 m depths. Temporal comparison using the Morisita index revealed 

strong community stability during 1975-76. 

163. Jaap, W. C. and J. Wheaton. 1992. Summary of preliminary results, long-term ecological coral reef 

studies, Ft. Jefferson National Monument, Dry Tortugas. Prepared for the National Park 

Service Workshop, 28-30 April 1992, Miami, Florida.

Coral reefs exist over time scales of thousands of years. Processes of change in the 

geological-time context occur slowly, e.g. sea level change correlated with glacial and 

interglacial periods. The etiology of change is often poorly understood. For example, in 

1878 a perturbation identified as, "black water" decimated Acropor-u spp. at Dry 

Tortugas. Determining what black water was may never be known. Long-term ecological 

research seeks to uncover processes that occur slowly or in which effects lag years behind 

the causes. In the absence of long-term research, serious misjudgments can occur in 

attempts to manage the environment. The National Park Service was interested in 

developing a reef resource monitoring plan for Dry Tortugas reefs and collaborated with 

the Florida Marine Research Institution in a joint study of reef resources. The goals of 

these studies included testing methods, acquiring a data base on coral reef benthic and fish 

conlmunities to better understand the etiology of change, and isolating natural from 

anthropogenic changes. Five study sites were selected in 1989. Repetitive sampling was 

executed as precisely as possible using several different methods. These methods included: 

transect sampling, video sampling, quadrat sampling, photographic sampling, recruitment 

sampling, and environmental sampling. Results indicated that eleven octocoral, 22 

scleractinian, and one milleporan species were enumerated on transects, while quadrats 

indicated 29 octocoral, 26 scleractinian and I milleporan species over the study's three year 

duration. Octocorals were consistently most diverse at Pulaski Shoal (20-21 species). Only 

42 of 212 plates recruited scleratinian corals. This yielded an average of 0.35 recruit per 

plate. A total of 187 milleporan corals recruited to 212 plates for an average of 0.88 

recruit per plate (34.9Iniz). The only octocoral recruit recorded was the gorgonacean 

Briareurn asbestinunl, whose common name is corky sea fingers. 

164. - 1994. Summary of preliminary results, long-term ecological coral reef studies, Ft. Jefferson 

National Monument, Dry Tortugas. Bulletin of Marine Science 54, no. 3: 1-10. 

Narrative same as in reference no. 163. 

165. Jaap, W. C., J. L. Wheaton and K. B. Donnelly. 1990. Materials and methods to establish 

multipurpose, sustained, ecological research stations on coral reefs at Dry Tortugas. Diving 

for Science ... 1990. Proceedings of the Amo-icnn Academy of Ur~derwuter Science, Tenth 

A~nzuul Scie~ice Diving S)v~tposium, 193-203. American Academy of Underwater Sciences: 

Costa Mesa, California. 

Sustained research requires precise, repetitive data acquisition to accurately evaluate 

patterns of change in species abundance and community structure. Permanent reference 

markers are essential to resample stations over time. The methods described here use solid 

markers from which several sampling devices can be deployed. A hydraulic drill is used to 

core 18-in deep holes into rock. A square stainless steel stake is inset, aligned, and 

cemented into each hole. Quadrats, photogrammetric and video apparatus, and recruitment 

arrays are deployed on or in reference to the stakes. Transects are extended between 

stakes. The method is suitable for coral reef and other hard-bottom investigations. 

166. Jaap, W. C., J. L. Wheaton, K. B. Donnelly, B. J. Kojis and J. E. McKenna Jr. 1994. A three year 

evaluation of community dynamics of corals at Ft. Jefferson National Monument, Dry 

Tortugas, Florida. Bulletin of Marille Science 54, no. 3: 1077. 


167. hap, W. C., J. L. Wheaton, K. B. Donnelly, B. L. Kojis and J. E. McKenna Jr. 1993. A three-year 

evaluation of community dynamics of corals at Ft. Jefferson National monument, Dry 

Tortugas, Florida, USA. (abs.). Proceedings of the 7th lnternatior~ul Coral Reef 

Sym/~osiunl, page 164. Guam: University of Guam. 

A study to evaluate methods and begin a long-term ecological research program at Ft. 

Jefferson was initiated at five reef sites in May 1989. Benthos was mapped and

photographed wilhin quadrats (5 x 2.56 m2 per site). Attached biota and substrates were 

measured along 20- to 25-m transects (3 per site). Recruitment arrays were constructed of 

PVC pipe, flat stock, and ceramic tiles (10.8 x 10.8 cm) and were secured to the reference 

stakes. A carriage-mounted video camera, suspended on cables between two " T poles 

secured to the stakes, was pushed the length of a transect. Results implied relative stability 

of the reef co~nmunities over three years. Dominant biota as determined by abundance and 

cover remained sirnilar. Classification analyses of station time-series also corroborated 

relative stability. Recruitment of Millepora, Octocorallia, and Scleratinia was variable; 

most recruits were found in cryptic refuge. The heterogeneity, high-relief, and multilayered 

canopy of these coral reef habitats restricts the usefulness of medium and long 

distance (>1.5m) photography and video. We conclude that multiple sampling methods are 

superior to a single sampling procedure. 

168. Jaap, W. C., W. G. Lyons, P. Dustan and J. C. Halas. 1989. Stony coral (Scleractinia and 

Milleporina) community structure at Bird Key Reef, Ft. Jefferson National Monument, Dry 

Tortugas, Florida. Florida Marine Research Publication 46: 1-31. 

Stony coral co~nmunity structure at Bird Key Reef was investigated during 1975 using 30 

continuous 25-m line transects in depths of 0.5 to 21.3 m. Thirty-two species, 872 colonies, 

and 198 cm of coral cover were sauipled quantitatively, Most species, colonies, and live 

coral cover occurred seaward of 8-m depths on spur and groove substrate. Montastrea 

annularis, M. cavernosa, and Siderasrrea siderea constituted more than 50% of all cover. 

Species diversity (Shannon index, log sub (2)) ranged from 1.0 for individual transects. 

Diversity and envenness values computed from cover data were generally lower than 

values computed from abundance data, reflecting M. annularis dominance. Numerical 

community classification (Czekanowski's quantitative coefficient) revealed three groups 

and an ecotone, each related to depth and substrate: 9 transects in 1 to 6 m depths 

dominated by Porires asteraides and Diploria clivosa; an ecotone of 6 transects in 5 to 6 

In depths; 5 transects in 6 to 9 m depths dominated by S. siderca; and 10 transects in 8 to 

21 in depths dominated by M. anriu1ari.s. 

169. Jacobs, M. H. 1914. Physiological studies on certain protozoan parasites of Diadernu setosum. 

paper^ Torrugas Laboratory 6: 147-57. 


It has been shown by the author and others that different species of protozoa have certain 

physiological characteristics, often almost as stiking as their morphological ones, and 

which are probably of considerable significance in the interpretation of their habits of life 

and their relation to their environment. It occurred to the author to test a series of forms 

which naturally live under essentially the same environmental conditions, and which may 

be assumed to have done so for many past generations, in order to see whether they show 

greater likenesses than a number of forms selected at random, or whether each has 

preserved its individuality in spite of the similarity of its environment. The general results 

of the experiments performed show surprising differences in the resistance of the parasites 

of Diadema to various unfavorable conditions. In some cases the most resistant form may 

live several hundred times as long as the least resistant one. Comparing all of the results 

obtained, it is therefore seen that the similar habit of life of the four fornu in question has 

not brought about physiological similarity except in certain adaptive characters which are a 

sine qua non for continued existence in the same host (e.g. ability to resist the digestive 

juices of the latter, etc.). In other respects they are just as different as almost any four freeliving 

forms that might be selected and the evidence of these experiments shows that the 

physiological characters of an organism are not merely the result of its environment, but 

may be as fundaniental and characteristic as its morphological ones.

170. Jefferson, J. P. J. Y. Porter and T. Moore. 1879. On the destruction of fish in the vicinity of the 

Tortugas during the months of September and October 1878. Pmceedings of the U.S. 

National Museum, Smithsonian Institution Press 1: 244-46. 

The information in this report is relative to the die-off of large numbers of fish due to a 

black water event in the Gulf of Mexico during the months of September and October 

1878. 

171. Jennings, H. S. 1909. Behavior of sea-anemones. Journal ofExperirnenta1 Zoology 2: 447-72. 

The study of the behavior of sea anemones (Stoichactis helianthus and Aiptasia spp.) was 

made at the Carnegie Research Laboratory, Dry Tortugas using specimens collected in the 

shallow waters near Fort Jefferson. Changes in behavior due to varying states of 

metabolism for S, he1ianthir.s were examined using red meat, crab hard parts and filter 

paper as food. After satiation, food is rejected through various reactions dependent upon 

internal processes. Descriptions of food ingestion are described. For Aiptasia spp., 

experiments suggested that when the animals were hungry, they took both red and filter 

paper; when satiated they took neither. Other topics of study included climatization to 

stimuli (light), and reactions modified as a result of past experiences of the organism. 

Results are compared to other lower groups of animals. 

172. Jindrich, V. 1972. "Biogenic buildups and carbonate sedimentation, Dry Tortugas reef complex, 

Florida." Ph.D. Dissertation, Geology, State University of New York at Binghamton. 

The Dry Tortugas, a horseshoe-shaped complex of carbonate banks and coral reefs, is 

located at the southern terminus of the Florida limestone shelf. The complex rises to the 

surface waters from a drowned Pleistocene surface that forms a circular platform having a 

general depth of 17-21 m. Three basic biogenic buildups (facies) comprise the reef 

complex: 1) detrital lagoonal bank, 2) Montastr-ea reef bank and, 3) Acropora palmata 

reef. These facies lie adjacent to one another and are also present in vertical succession as 

individual growth stages of varying thickness and lateral extent. A zone of Acropora 

cer-eicornis is developed as a transition between the Montastrea and A. palmata growth 

stages. The present organic assemblages and topography bear evidence of dominantly 

lateral progradation and cu~nulative storm effects that are linked to the slow eustatic sealevel 

rise for the past several millennia. Long-continued storm degradation is manifested 

by 1) continuous removal of A. palmata and its replacement by storm-resistant coralline 

algae and Millepora sp. to produce truncated rocky surfaces, 2) abundant reef rubble, 3) 

erosion of spur-grooves, and 4) developnlent of intertidal rubble reef flats. Sediments 

ranging from cobble-sized rubble to medium silt are composed of Halinieda, coral and 

mollusc grains; coralline algae and foraminifers are present in minor amounts. Variations 

in texture and constituent particle composition are interpreted to be mainly a result of mode 

of sediment transport and effect of grain shape. Broadly-defined grain size populations 

produced by three modes of transport have characteristic assemblages of constituent 

particles. The populations include a gravel-sized surface creep population, sand-sized 

saltation population, and very fine sand-to silt-sized suspension population. Strong mixing 

occurs between the gravel and sand population on the storm-degraded shoals, and between 

the sand and silt population on the lagoon bottom. Sand flanking the reefs and reef banks 

shows mini~nun~ mixing hence good degree of sorting. Incongruous mixtures of the inplace 

fraction and varying proportions of the transported populations constitute detrital 

lagoonal banks as a substrate stabilized by seagrass and coral growth. The gravel-sand and 

sand-silt mixtures are related to deposition under highly variable energy conditions. 

Variability in energy conditions does not cause strong population intermixing on beaches. 

For the same reason, beach sediments show a high degree of sorting in all size grades from 

cobbles to fine sand.

173. Jones, N. 1938. Investigations on ascidians. Carnegie lrtstitution of Washirigtow, Year Rook 37: 84. 

The su~nmer of 1938 was devoted to the study of the structure, development, budding, and 

colony formation of Ecteinascidia tortugensis, a new ascidian species. The small ascidian, 

one of the commonest during the season, occurred in large numbers on the under sides of 

rocks just below low-water mark on both Bush Key and Long Key. Data report provided 

by Plough and Jones, pp 97-98. 

174. Jones, R. D. 1991. An improved fluorescence method for the determination of nanomolar 

concentrations of ammonium in natural waters. Limrlology and Oceanograpl~~ 36, no. 1: 

814-19. 

An improved fluorescence method is described for measuring nanomolar concentrations of 

NH, in natural waters. This method is based on the conversion of NH4 to NH, and 

subsequent diffusion of NH, across a microporous hydrophobic Teflon membrane into a 

flowing stream of 0-phthaldialdehyde reagent to produce a fluorescent adduct. The 

product is detected fluorometrically with a lower detection limit of better than 1.5 nM. Up 

to 30 determinations h-l can be made. The method works well in freshwater or salt water. 

Field tests of the method in the Dry Tortugas and Gulf Stream gave NH, concentrations 

that ranged from 18.0 nM in Gulf Stream waters to 2,254.7 nM in interstitial waters from 

coralline reef sands. The method can be used to measure near real-time NH, 

concentrations in situations where it was previously difficult or impossible. 

175. Jones, R. S. and M. J. Thompson. 1978. Comparison of Florida reef fish assemblages using a rapid 

visaal technique. Bulletin of Marble Scierfce 28, no. 1: 159-72. 

Species composition, species diversity, and relative abundance of 4 coral reef fish 

communities in John Pennekamp State Park, Key Largo, Florida, are compared with 4 

co~nmunities at Fort Jefferson National Monument in the Dry Tortugas using the speciestime, 

random-count technique. The technique is similar to species-area methods, but time 

replaces area. Fish communities at Pennekanip Park showed the highest overall number of 

species and scores (reflecting species abundance, and species diversity). Two artificial 

reefs (shipwrecks) included in the study both show closer relationships to adjacent reefs 

than to wreck-specific species. 

176. Jordan, D. S. 1904. Notes on fishes collected in theTortugas Archipelago (by Dr. Joseph C. 

Thompson). Rulletirz of the United States Fish Commissionfor 1902 22: 539-44. 

An additional sixteen species of fish are described for the Tortugas based on collections 

made by J.C. Thompson while on the northward cruise of the steamer Chesapeake. 

177. Jordan, D. S, and J. C. Thompson. 1905. The fish fauna of the Tortugas Archipelago. Bulletin ofthe 

United Strites Bureau of Fisheries for I904 24: 229-56. 

The shallow water fishes of the Tortugas, as suggested by A.G. Mayer to David Starr 

Jordan, are unsurpassed in variety and abundance anywhere along the Atlantic coast of the 

United States, and based on the nearness of the Gulf Stream and the winds and currents, 

pelagic fish from all over the Gulf of Mexico and the West Indies may be drifted by the 

Tortugas. Collections made by Thompson while on duty as a medical doctor at the Garden 

Key Naval Station resulted in an annotated fish list containing 218 species known to occur 

at the Dry Tortugas at the time. 

178. Jordan, H. E. 1908. The accessory chromosome in Aploprrs mayeri. Atlatomischer Anzeiger Hd 32: 

284-95. 

The purpose of this paper is to trace the accessory chromosome in the phasmid, Aplopus 

mayeri from material collected from Loggerhead key, Florida. The accessory chromosome 

appears in the resting stage of the secondary sperlnatogonia as a chromatin nucleolus 

characteristically close to the nuclear wall. Both the primary and secondary spertnatogonia

have a metaphase group of 35 chromosomes. The accessory chromosome can be traced as 

a specific structure from the resting stage of the last order of spermatogonia through all the 

various phases of synapsis and maturation, until it disintegrates in the head of the ripening 

spermatozoa. 

179. ---. 1917. Aortic cell clusters in vertebrate embryos. Pmceeding.~ of the National Academy of 

Science 3: 149-56. 

Aortic cell clusters are described among various animal groups (pig and chick) and 

compared to 12-day loggerhead turtle embryos and mongoose embryos. Various aspects of 

the hemogenic activity of embryonic endothelium are described consequent to the inherent 

capacity of endothelium to produce hemoblast, and not in connection with an associated 

toxic substance. 

180. ------ . 1917. Atresia of the esophagus in the embryo of the loggerhead turtle, Caretta caretta: A 

normal developmental condition. Papers Tortugas luboratory 11: 345-60. 


A series of 26 embryos of the loggerhead turtle were collected and used originally for a 

study of the history of the primordial germ-cells. It was noticed that the esophagus was 

solid for a greater or less extent, approximately from the point of origin of the respiratory 

anlage to its bifurcation into the bronchi, from the eleventh to the thirty-second day of 

incubation. Points of special significance in regard to this ~naterial are: (1) the relatively 

longer persistence of the occlusion than has yet been described for any other form; (2) the 

absence of contributory yolk in the stenosed area; (3) close relation to the point of origin of 

the respiratory anlage, which fact may disclose its possible functional significance 

181. ------ . 1917. Embryonic history of the germ-cells of the loggerhead turtle (Carettn caretfa). 

Papers Torlugas 1.uborufory 11: 3 13-44. 


The wide discrepancies in the published accounts of the origin and early history of the 

germ-cells in vertebrates provided the stimulus for this investigation. Twenty-five embryos 

of the loggerhead turtle (Carefta carerfa), ranging from the second day (5 somites, 2mm. 

length) to the thirty-second day of incubation, were employed in this investigation. Results 

indicate that the primordial germ-cells migrate during the second day from the yolk-sac 

endoderm, where they are widely scattered caudally, into the lateral border of the area 

pellucida on each side of the embryonic disk. The germ-cells migrate by amoeboid 

activity. The migration period is not sharply limited. A certain number of germ-cells 

migrate out of the regular germ-cell route and go astray. The total number of primordial 

germ-cells counted in a 12-day embryo is 352. Occasional cells may divide by mitosis, or 

undergo degeneration, at any stage of their history or at any point of the route. No germcells 

were found contributing to the formation of the Wolffian duct. The germ-cells do not 

differ from young somatic cells in the character of their mitochondria1 content. No 

transition stages between coelomic epithelial cells and germ-cells appear up to the 32-day 

stage. The evidence derived from a study of the Car-etta embryos is in complete harmony 

with the idea of a single uninterrupted line of sex-cells from primordial germ-cells to 

oogonia and spermogonia, and with the hypothesis of a vertebrate Keimbahn or continuous 

germinal path. 

182. -. 1908. The germinal spot in echinoderm eggs. Papers Tortiigas Laborarory 1 : 1-12. 


This paper reports the results of further studies of the prematuration stages of echinoderm 

eggs of additional species of echinoderms, a star-fish (Echinaster crassispirza), and a 

brittle-star (Ophiocorna pumila). I11 Echinaster crassispir~u the chromosomes are derived 

exclusively from the nucleolus. In Ophiocomu purnila the chromosomes arise exclusively

from the nuclear reticulum. In some species the chromosomes arise from a chromatinnucleolus, 

in others from a chromatic reticulum, and in still others in part from one source 

and in part from the other. The eggs of different forms differ in that some have only a 

chromatin-nuleolus, without distinct plastin ground-substance, resting in an achromatic 

nuclear reticulum (Echinuster); others possess both chromatin-nucleolus and plasmosome 

as well as a chromatic nuclear reticulum (Ophioconta); and still othcrs possess a double 

nucleolus (chromatin nucleolus and plastin ground-substance), with the chromoso~ne 

complex gathered in a mass in the achromatic reticulum (Asterias). The function of the 

germinal spot then appears, in part at least, to be that of a storehouse of material which is to 

contribute to the formation of the chromosomes . 

183. -- . 1917. The history of the primordial germ cells in the loggerhead turtle. Proceedings of the 

Nutiorla1 Academy of Sciotce 3: 27 1-75. 

This study attempts to trace germ cell history in the loggerhead turtle and compare it to 

observations for other vertebiates. Embryos were collected for study from specimens taken 

on Loggerhead Key, Dry Tortugas during the Summer of 1914. The germ cell history of 

Curettu is very similar to that first described for Chrysemys and to that described for 

dogfish. 

184. ---. 1917. The microscopic structure of striped muscle of Lirnulus. Papers Tortugus Luboratory 

l I: 273-90. 


The study of the skeletal muscles of 1.imulus was undertaken with two chief objects in 

view: to test a conclusion suggested by earlier studies on the intercalated disks of 

vertebrate cardiac muscle, namely, that these disks are properly interpreted as "irreversible 

contraction hands" and to seek additional evidence in further refutation of the recently 

revived hypothesis that striped niuscle can be interpreted in terms of "muscle-cells" and 

intercellular tnyofibrillae. It was found that both the skeletal and the cardiac ~nuscles of 

Limulrrs consist of trabeculae of finely granular sarcoplam. In cardiac niuscle the main 

trabeculae and their branches form a loose-meshed syncytium. Neither type of muscle 

contains mesophragmata. Very rarely an intercalated disk of the simple-comb type appears 

in the cardiac muscle. Both types are very similar in respect of the presence and 

arrangement, in the same phase of contraciton, of Q and J disks, and the telophragmata. 

The evidence is unequivocal against an interpretation of structure in terms of "musclecells" 

and intercellular myobibrillae. The nuclei of the growing muscles multiply by 

amitotic division. In essential structure the cardiac and skeletal muscles of Limulus are 

closely similar, indicating a close functional similarity. The structure serves, moreover, as 

a splendid illustration of the "law of biogenesis," in that it is practically identical with a 

stage in the early histogenesis of striped muscle of teleosts . 

185. --- . 1908. The relation of the tiucleolus to the chromosomes in the primary oocyte of Asteiias 

forbesii. paper:^ Tortugas Lnborato~); I : 37-72. 

Carnegie Institution of Washington I'ublication Number 102. 

The primary object of this investigation was was to contribute to the subject of the relation 

between nucleolus and chromosomes during maturation. In summary, synizesis occurs in 

the oocyte of the first order at the very beginning of the growth-period (size of nucleus 5 

microns). The growth-period is passed through rapidly. During the latter half of the 

growth-period all the chromatin, with the exception of what is held by the chromosomes, 

becomes stored in the enlarging nucleolus. The nucleolus consists of a plastin groundsubstance 

infiltrated and covered over with chromatin. The chromosomes do not arise out 

of the nucleolus. The number of chroriiosomes in the prophase of the first polar mitosis is 

18. They vary somewhat in size (onc is considerably larger than the rest), all have a 

characteristic dumb-bell shaped appearance, and some are clearly double (bivalent). The

two maturation divisions effect a double longitudinal fission of the original bilobed 

chromosomes. The reduced number of chrornosomes is again 18. Observations on 

Hippottoe escraler~ra agree in essential points with those made on Asreria forbesii and 

support the conclusions regarding the origin of the chroruosomes, the function of the 

nucleolus, and the reduction phenomena. 

186. ----. 1908. The spermatogenesis of Aplopus rnapr-i. Papers Tortugas Laboratory 1: 13-36. 


The object of the present investigation is primarily to trace the history of the accessory 

chroniosome through the various stages in the process of spermatogenesis in the phasmid 

Aplopus mayeri. The material upon which the investigation is based was obtained from the 

Loggerhead Key, Florida. Primary spermatogonia divide both mitotically and amitotically. 

In the latter instance cell-division is frequently not consummated and a bi- or multi-nuclear 

cell results. In the first order of the secondary spermatogonia the accessory chromosome 

appears in the resting-stage. During synapsis the accessory chromosome lengthens into a 

club-shaped structure attached by its lesser end to the presynaptic thread, undergoes partial 

longitudinal division, closes up again during the height of synapsis, and returns again to its 

previous characteristic form and location in the nucleus of the growing primary 

spermatocyte. The second ~iiaturation division is equational, effecting a longitudinal 

division of univalent chromosomes. The accessory also divides equationally in the cells 

containing this element. A dimorphism of spermatozoa results; the accessory chromosome 

possessed by onc-half probably represents a sex-determinant. The history of the accessory 

chromosome gives evidence that it at least possesses a strict ~norphological and probably 

also a physiological individuality. 

187. Kaas, P. 1972. Polyplacophora of the Caribbean region. P. Wagenaar and L. J. Van Der Steen 

Hummelinck, 1-162. Studies on the Fauna of Curacao and Other Caribbean Islands, ed. P. 

Wagenaar and L. J. Van Der Steen Hummelinck. The Hague: Martinus Nijhoff. 

This paper includes Tables of Distribution of Polyplacophora of the Caribbean . The 

author took into consideration the whole of the Florida coast as far north as Fernandina, E. 

Florida and the Keys, the Dry Tortugas, W. Florida, the Gulf of Mexico, and also the 

Bermudas. Thiele's description of his single 5.5 specimen of i.schnochirotz harnneyeri from 

Bird Key Reef is translated into English here by Kaas. 

188. Kale, H. W. 1985. Florida birds - Dry Tortugas. Florida Naturalist 58, no. 2: 6. 

A sighting of a great black-backed gull is made at the Dry Tortugas, and a scarcity of land 

birds is reported. 

189. Kellner, Carl. 1907. Embryology of the appendicularian, Oikoplerrra. Zoological Anzeige,; Bd. 31: 

May. 

The appendicularia of the Dry Tortugas specimens of Salpae were collected. 

Appeodicularia of the genus Oikopleura and their "houses" were examined and found in 

surface waters. Their anatomy and histology are described. 

190. Kille, F. R. 1936-1937. Regeneration in l~olothurians. Curnegie irtstiturion of Washirtjiton, Year 

Hook. 

Note: published as follows: 1936, v. 35, p. 85-86; 1937, v. 36, p. 93-94. 

Histological studies were conducted on sea-cucumbers of the genus Holothuria, to 

determine the manner in which the digestive system is reconstituted following autotomy by 

means of electrical stimuli. 

191. Kopac, M. J. 1936. Electrical resistance of Valor~ia. I. Changes in the resistance with time in 

impaled coenocytes. Papers Tortugas Laborato,y 29: 359-86 (issued Mar. 1936).


During the summers of 1933 and 1934 the author worked at the laboratory of the Carnegie 

Institution located on Loggerhead Key, Dry Tortugas, Florida. Several species of Valonia 

were found growing abundantly on the various coral reefs of the Dry Tortugas. A study of 

the electrical resistance of impaled Valonia coenocytes by using a technique more highly 

refined that that ernploycd by previous investigators was initiated. Although only V. 

verrtricosa was used in this study, the methods developed and used here may be extended 

to the study of other species of Valonia. Glass microcapillaries, with tips ranging from 

0.025 to 0.1 mnl, in diameter and filled with vacuolar sap, served as microsaltbridges 

leading from the vacuole to a calomel half-cell. A larger glass tube (the macrosaltbridge), 

filled with sea-water, was used as a saltbridge lcading from the sea-water surrounding the 

coenocyte to another calomel half-cell. The Valonia coenocytes were impaled on the tip of 

the microsaltbridge with the aid of a micromanipulator. It was found that coenocytes with a 

high chloroplastid density have a constant Rp several times higher than coenocytes with a 

low chloroplastid density. It is postulated that only the inter-chloroplastidal protoplasm is 

capable of conducting a current. The average initial Rp in type A punctures was 60 to 65 

per cent of the constant Rp. In some coenocytes a constant Rp was reached in a few 

minutes. This increase in Rp is caused largely be the redistribution of those chloroplastids 

around the microtip which were disturbed by the puncture. The average initial Rp in type 

B punctures was 2 to 3 per cent of the constant Rp. After the chloroplastids are 

redistributed in this hyaline zone, the disintegrated chloroplastids are extruded, and the tiny 

vauoles are eliminated, no further increase in Rp takes place. 

192. Kunkel, B. W. 1934. The selective action of certain adverse environmental conditions on the hermit 

crab (Clibanarius tricolor Gibbes). Papers Torrugus Laboratory 28: 215-44 (issued Aug. 

1933). 


The problem of selection is undoubtedly a very complex one. The clwacters which enable 

one organism rather than another to survive are difficult to ascertain; a favorable variation 

of one part may be accompanied by an unfavorable variation of another, so that selection 

may have no effect upon the first feature. The present study has to do with a phase of the 

selection problem which, on the whole, has received rather scant attention from 

investigators. The selective effect of certain adverse conditions on a population has been 

studied. The probleni is that of determining how a given species may respond to a change 

in environment, of determining the morphological difference between those individuals 

which succumb to a certain change in the normal environment and those which are able to 

withstand the change. The material upon which the present study is based was collected 

and the experiments were made at the Tortugas Laboratory. The small hermit crab 

Clibanarius tricolor Gibbes was selected for the experiments. 

193. Le Compte, M. 1937. Some observations on the coral reefs of the Tortugas. Carnegie Institution of 


Particular attention was paid to the distribution and adaptation of the corals on the reef 

west of Loggerhead Key. A baseline of about 2500 yards is verified, documenting large 

heads of Orbicella (Madrepora) anr~ularis, extensive growths of Acropom, and areas of 

gorgonians and algae. Beach rock development, coral feeding habits, and the effects of 

boring animals on corals are discussed. 

194. Leitch, James L. 1936. The water exchanges of living cells. 111. The application of a photographic 

method to the determination of the non-solvent volume of the eggs of Echinornetra 

lacunter. Papers Tortugns Laboratory 29: 349-58 (issued Mar. 1936). 


Photography has been applied to the study of living cells and tissues. Leitch raised the

question of the feasibility of a photographic method in the study of the osmotic behavior of 

cells. The present paper outlines such a method and discusses some of the factors involved 

in the study of the water exchanges of the eggs of the sea-urchin, Echinometra lacunter, 

using measurements of photographs of eggs at equilibrium in dilute sea-water solutions. It 

was shown that photography can be employed in the study of the water exchanges of living 

cells. The non-solvent volume of the eggs of Echinoinetra lucunter- is 36 per cent when 

calculated after from 60 to 90 minutes' exposure to experimental solutions. Longer 

exposures to the experimental solutions result in a higher non-solvent volume of 48 per 

cent which is associated with pronounced vacuole formation. The appearance of vacuoles 

after the attainment of the first equilibrium is discussed and three different explanations 

proposed. 

195. --- . 1937. The water exchanges of living cells. IV. Further studies on the water relations of the 

eggs of the sea-urchin, Echinornetra laciinter. Pupers Tortugas la born tor^ 31 : 53-70 

(issued July, 1936). 


The application of a photographic method to thc determination of the non-solvent volume 

of the eggs of the sea-urchin, Echir~on~etra lacunter, and also the effect on the non-solvent 

volume determinations of the length of time of inlmersion of eggs in diluted sea-water 

solutions, was demonstrated in another publication (Leitch,1936). The present paper 

considers the utilization of this photographic method for the study of the swelling and 

shrinking of the eggs of the same sea-urchin, the effect on the water relations of these cells 

of the time between that of spawning and that of introducing the eggs into the experimental 

solutions, and an analysis of equations which have been developed and applied by several 

investigators to explain the kinetics of water exchanges of living cells. Results showed that 

an analysis of the equations developed to interpret the kinetics of water exchanges of cells 

the so-called permeability "constants" are not constant for the eggs of Echirmnetra 

lacuntcr but vary with the dilutions of sea-water used and also with different intervals of 

time in the same dilution. The permeability constants for swelling and shrinking do not 

coincide, being between 0.250 and 0.650 for the former process and between 0.180 and 

0.580 for the latter. The introduction of the correction for the non-solvent volume into the 

equations does not produce a better agreement between the constants for the two processes. 

The use of the photographic method (Iaeitch,1936) is furthcr substantiated for the 

determination of non-solvent volu~nes and is extended to the study of the swelling and 

shrinking of eggs. Approximately two hours from the time of spawning the non-solvent 

volumes is greatly increased, from 30 to 53 per cent of the initial volume. There is a slight 

retarding effect on thc water exchanges of the eggs brought about by standing. The time at 

which the effect of standing at room temperature appears in the values of the non-solvent 

volume and rate of penetration of water is correlated with a sharp decrease in the 

percentage of development, a slight increase in the volume of the eggs and a cytolysis-like 

phenomenon which finally ends with the complete disintegration of the eggs. The 

production of fertilization membranes as a criterion of non-injury of the egg cell is shown 

to be inadequate and the percentage of development of normal larvae is urged as a better 

test of normality 

196. Lessios, H. A,, D. R. Robertson and J. D. Cubit. 1984. Spread of Diaderna mass mortality through 

the Caribbean. Scier~ce 226: 335-37. 

Populations of the ecologically important sea urchin Diadeii~ailtillarurn suffered severe 

mass mortalities throughout the Caribbean. This mortality was first observed at Panama in 

Ja~iuary 1983; by January 1984 it had spread to the rest of the Caribbean and to Bermuda. 

The sequence of mortality events in most areas is consistent with the hypothesis that the 

causative agent was dispersed by major surface currents over large distances. However, 

some of the late die-offs in the southeastern Caribbean do not fit this pattern. Several lines

of indirect evidence suggest that the phenomenon is due to a water-borne pathogen. If so, 

this is the most extensive epidemic documented for a marine invertebrate. 

197. Linton, E. 1908. Helminth fauna of the Dry Tortugas. I. Cestodes. Papers Torrugas Laboratory 1: 

157-90. 


This report is based on data collected at the Marine Biological Laboratory, Tortugas, 

Florida, June 30 to July 18, 1906. A list of the hosts which were examined for parasites, 

and a summary of the results of that examination, together with a few food notes are 

presented. A few extracts from notes made at the time the material was collected are 

presented. Acanthocephala are presented. The species found in the frigate mackerel was 

Echinorphynchus pristis. Few nematodes were found in the nurse-shark. New species of 

parasites are described. 

198. -- . 1910. Hellninth fauna of the Dry Tortugas. 11. Trematodes. Puper.s Tortugas Laboratory 4: 

11-98, 


The collection here described was made at the Marine Biological Laboratory, Tortugas, 

Florida, in the summers of 1906, 1907, and 1908. The fishes examined were from the 

shallow waters of the reef. The distribution of parasites together with food notes have 

already been published in the Year Book of the Carnegie Institution of Washington for the 

years above named. This paper includes a list of Tortugas trematodes and their hosts, key 

to the genera and species described, and descriptions of species, habitats, etc, including: 

trematodes from loggerhead turtlcs and from fish. 

199. -. 1907. Note on the habits of Fierasfer aflinis. American Naturalist 41, no. 481: 1-4. 

Observations of the Fierasfer affiriis entering its host, tail first are made. 

200. Lipman, C. B. 1929. The chemical composition of sea water. Papers Turtugas Laboratory 26: 249- 

57. 


In his studies on marine bacteria and related subjects, the author realized the need for more 

accurate and complete analytical data on the inorganic components of sea-water and 

determined to obtain them.. Two samples were analyzed from the Atlantic area and they 

were both from the Gulf Stream, and taken near Loggerhead Key in the Tortugas. The data 

render it clear that a large part of the ions important to algae are removed from solution in 

sea-water by a rise in pH of that medium, which is well within the range of daily rise in pH 

of sea-water carrying an active algal flora under the proper conditions of light and 

temperature. 

201. --- 1924. A critical and experimental study of Drew's bacterial hypothesis on CaCO? 

precipitation in the sea. Papers Tortugas Laborator): 19: 179-91. 


Based on a series of experiments to explain the precipitation of CaC03 in sea water, it was 

found that there are several ways to explain CaCO1, where it occurs in seawater, without 

introducing Drew's hypothesis or any other bacterial hypothesis. These explanations of the 

phenomenon seem adequate to account for the qualitative and quantitative differences in 

CaCO, as found under different conditions in seawater. Changes in water and air 

temperatures, and marine plant activity, which Drew and others clearly appreciated and 

understood, yet have introduced a purely gratuitous bacterial hypothesis based on what 

appears to be sound experinients.

202. --- . 1929. Further studies on marine bacteria with special reference to the Drew hypothesis on 

CaCo-, precipitation in the sea. Papers Tortugas Luboratory 26: 231-48. 


Bacterial populations in the open sea are very small. Mixed or pure cultures of organisms 

isolated from sea-water are incapable of precipitating CaCO-, from sea-water to which no 

salts have been added, or even in the presence of added KNO, Mixed or pure cultures of 

such organisms are incapable of precipitating CaC03 in a sea-water medium if KNO, and 

organic matter as the sugars or similar form free from calcium are added to the medium. 

Upon the basis of evidence in this and in earlier papers the Drew hypothesis is shown to be 

untenable, and at the very least uproved. This strong probability is reemphasized in the 

purely physical-chemical nature of CaCO, precipitation on a large scale in nature. 

203. Locker, S. D. A. C. Hine and E. A. Shinn. 1991. Sea level geostrophic current control on carbonate 

shelf-slope depositional sequences and erosional patterns, South Florida platform. AAPG 

Rulletirr 75, no. 3: 623. 

High-resolution seismic reflection profiles across the shelf-slope margin between the Dry 

Tortugas and Key West, Florida, indicate that sea-level fluctuations and the eastward 

flowing Florida Current are major controls on late Quaternary sequence stratigraphy. The 

study area, a transition zone between the open south Florida shelf and the lower Florida 

Keys islandlreef system, is typified by a shallow shelf with reef margin adjacent to a deeper 

lower-shelflslope. The lower-shelfislope is composed of stacked or prograding sequences 

that downlap and pinchout on the Pourtales Terrace. Strike oriented stratigraphic sections 

exhibit many sea-level controlled features such as lowstand erosion, transgressive 

unconformities, and highstand system tracs. Lowstand reefs, notches, or barriers are 

observed as deep as 150111 below present sea level. Depositional styles change along-slope 

from west to east. The western portion of the study area is characterized by thick, low 

amplitude prograding sequences related to abundant supply of sedi~nenthrough off-shelf 

transport during high sea-levels as well as along-slope reworking by the Florida current. 

Part of this section has been severely eroded by along-slope current producing localized 

cur and fill structures and widespread erosional unconformities. To the east, a thinner 

section of higb-amplitude reflections is common seaward of the lower Florida Keys reef 

tract system. Again, along-slope current erosion and winnowing of sediment supplied by 

the adjacent margin is evident. This study provides new cvidencc of how a strong 

geostrophic boundary current along with fluctuating sea levels have interacted to control 

depositional sequences on a carbonate slope in the FloridaIBahamas platform complex. 

204. Longley, W. H. 1917. Changeable coloration in Brachyura. Proceedings of the National Academy of 

Science 3: 609- 1 1. 

Studies on changes in the color of brachyuran crabs (Ocypoda and Callir~ectesp.) at the 

Marine Laboratory, and in the field on Loggerhead Key, Dry Tortugas, demonstrate 

adaptive coloration based on temperature variations and the color of the substratum upon 

which the specimen is resting. It is expected that in future studies the same general rules of 

adaptation for fishes will apply to crabs. 

205. -. 1918. Haunts and habits of tropical fishes. Ari~erica~~ Museurn Jourr~al 18: 79-88. 

Observations are recorded at the Dry Tortugas using underwater photography. Habits of 

the shallow water reef species were photographed in water less than 10 feet deep. 

Emphasis is placed on the biological significance of color in fishes with their surroundings. 

Fish color change may be evoked by offering them food by hand at different locations. 

The foods and feeding habits of reef fish are discussed in this report. 

206. - , 1918. Marine camoufleurs and their camouflage: the present and prospective significance 

of facts regarding coloration of tropical fishes. Smithsorrian Report (1920): 475-85.

Fish are used as an example of an animal which uses color patterns, not as a struggle for 

existence as hypothesized by Darwin, but as a means of expressing its biological 

significance by displaying a natural system of camouflage. Some colors in fishes are not 

changeable, but seem to be correlated to definite habits. In the case of those that are 

changeable, there is conclusive evidence that they are displayed under specific conditions. 

For example, transverse bands are shown when the species is inactive. However, upon 

movement the bands are replaced by stripes. 

207. - 1916. Observations upon tropical fishes and inferences from their adaptive coloration. 

Proceedirzgs of the National Academy of Science 2: 733-37. 

The conception that species have been multiplied by divergent evolution of related strains 

is based on many observations. If the Darwinian hypothesis is true, the character of 

organisms should be largely of an adaptive sort, but its adherents have been content to 

support this position by inputting utility to structure and habits. It has not been proved in 

fishes that some color combinations ward off enemies nor that pigmentation is functionally 

conspicuous. Many of the brightly colored fishes of the Tortugas have been studied to 

evaluate their coloration objectively. Most species exhibit countershading with darkest 

shading on the upper surface and lighter shading on the mid-ventral or lower line. Thirteen 

species of fish studied exhibit color changes based on their surroundings observed from 

boats or from the bottom using diving equipment and photography. Correlation of color 

with habitat has been documented. Some examples suggest that red fish are rarely seen 

during the day, gray fish with diurnal activity patterns are found near large coral heads, 

lighter blue fish are habitually found swimming well above the bottom in moderate depths, 

and those species largely found over grass beds are of green color or have a green color 

phase. As far as this class of animals is concerned, Longley postdates that there is no 

groutid for the belief that bright color is correlated any way with armament or 

distastefulness. Problems of mimicry resemblance are unresolved, however the 

observations presented in this abstract undermine many speculative explanations of animal 

coloration in terms of natural selection and replace them with something which may not be 

dismissed fro~n consideration. 

208. - 1936. Species studies and the species problem. American Narurulist 70: 97.109. 


209. ------ 1917. Studies upon the biological significance of animal coloration. I: The colors and color 

changes of West Indian reef fishes. Jonr-nal of Experimental Zoology 23: 536-601. 

Studies were carried out at the Dry Tortugas to determine the biological significance of 

changes in color of reef fishes. It was found that fishes are countershaded; color changes, 

which are common even among the most gaudy, tend to assimilate them with their 

environment; and in general, their colors repeat those of their surroundings. Specially 

defended types are not unlike others in pigmentation, nor inferior to them in their ability to 

effect adaptive color adjustments. Finally, there is no evidence that brightly colored 

species enjoy greater immunity from attack than their fellows, for they constitute a large 

proportion of the food and may be readily identified in the stomach contents of predaceous 

forms. These statements, which rest upon a great body of verifiable observations, are 

consistent with the Darwinian hypothesis, but inconsistent with the assumption that animals 

of high color possess more than minimal conspicuousness under natural conditions. They 

impel one to reject the hypotheses of warning and immunity coloration, signal and 

recognition marks, and sexual selection, at least in so far as they may ever have been 

supposed to apply to these forms. Upon the contrary, they confirm Thayer's conclusions 

regarding the obliterative function of color and pattern, emphasize the common occurrence 

of adaptive characters among animals, and suggest that their evolution has been guided 

throughout by natural selection.

210. -, 1917. Studies upon the biological significance of animal coloration. 11: A revisional 

working hypothesis of mimicry. American Naturalist 51: 257-85. 

In this report, various hypotheses proposed by the author and other investigators relating 

changes in animal coloration in relation to habits are discussed. The author postulates that 

bright colors of tropical fishes are correlated with the animal's habits from work achieved 

at the Dry Tortugas. Other coloration hypotheses are provided dealing with butterflies and 

lizards, as well as warning coloration in bright and dull-colored insects. These ideas 

submitted by the author constitute working hypotheses to be tested by other biologists. 

21 1. Longley, W. H. and S. F. Hildebrand. 1940. New genera and species of fishes from the Tortugas, 

Florida. Papers Tortugas Laboratory 32: 223-85 (issued Sept. 1940). 


Thirty new genera and species of fishes described in these pages resulted from studies 

carried on for many years, principally at Tortugas, Florida, by the late Dr. William H. 

Longley, whose untimely death occurred before he had fully completed a study of his 

collections and a manuscript embodying a complete account of his field observations. The 

present writer has made further studies of the Tortugas collections, and has endeavored to 

extract interesting facts from Dr. Longley's notes on those species not treated in his 

unfinished manuscript. 

212. Longley, W. H. and S. F. Hildebrand. 1941. Systematic catalogue of the fishes of Tortugas, Florida; 

with observations on color, habits, and local distribution. Papers Tortugas Laboratory 34: 

1-331. 


Observations on the fishes of the Tortugas Atoll were made by the senior author spanning a 

period of over 25 years. An inventory of 442 species is included, covering a wide range of 

habitats: bare sand, seagrass beds, coral reefs, channels between the keys, and deep waters 

over 600 feet in depth a few miles southward. Over 300 species were associated with coral 

reef habitat. This study represents the first fish survey conducted on the Florida Reef Tract. 

Field observations were made largely with the use of a diving helmet, which enabled the 

investigator to observe and photograph the fish in their native habitat, and to give 

information as to their behavior, feeding and habits, and especially adaptive coloration. 

Much of Longley's work is documented by the world's first underwater color photography. 

Following the death of Dr. Longley, Dr. Hildebrand undertook the editing and the 

completion of the manuscript. Material added by him bear his initials. 

213. Luck&, B.. 1937-1938. Studies on the tumors of fishes (of the snapper family Lutjanidae) . Carnegie 

Institution of Washington, Year Book. 

Note: published as follows;'l937, v.36, p. 98-99; 1938, v.37, p. 92-94. 

Certain kinds of tumors found on thirty nine fish belonging to several species of snappers 

closely resemble human neoplasms arising from nerves. They arise in the subcutaneous 

tissue and appear as flattened oval masses. No tumors of this kind were found on other 

species of fish. Epithelial growths were found in thirty specimens of Halichores radiatus, 

from a total of six thousand fish observations. Multiple papillomas of the skin and the eye 

were reported in a green turtle caught off Cape Sable. 

214. --- , 1942. Tumors of the nerve sheaths in fish of the snapper family (Lutjanidae). Archiwr of 

Pathology 34: 133.50. 

Fish of the snapper family Lutjanidae are corn~nonly afflicted with tumors which resemble 

the nerve sheath tumors of man called variously neurinoma, neurolemmoma, schwannoma, 

or neurofibroma. Neoplasms of this kind have been observed in 76 fish of three species, 

the gray snapper (Lutjunus griseus), the dog snapper (L, jocu), and the schoolmaster (L. 

apodus). Most of the fish were collected from the Dry Tortugas. Many other fish families

were examined, however no tumors of the kind were found. The tumors generally were 

found along the course of the subcutaneous nerves, particularly of the head and dorsal 

regions, as solitary or multiple, relatively large firm white masses. Like liuman neoplasms, 

the tumors of fish are usually composed of two kinds of tissue: one compact and richly 

fibrocellular; the other loose reticulated and poorly cellular. The component cells and 

intercellular fibers of the tumors appear to be essentially the same, and arranged in similar 

patterns, in fish and man. Unlike human tumors, the fish tumors, though well 

circumscribed, are usually not encapsulated. Nervc sheath tumors appear to be more 

common in certain fish species than in man. The frequency of occurrence of these tumors, 

which can be maintained for long periods in marine aquariums, renders them favorable 

material for studies of neoplasms. 

215. Luck&, B. and H. G. Schlumnberger. 1949. Neoplasia in cold-blooded vertebrates. Pl~ysiological 

Reviews 29, no. 2: 91-126. 

This review complements an earlier review to source material, abstracts of all the reports in 

the literature dealing with tumors in fishes, reptiles, and amphibians. In regards to fishes of 

the Tortugas, a review is presented on the senior author's work on tumors of nerve tissue, 

as described in Lucke (1942, reference no. 214). 

216. Lynts, G. W. 1968. Analysis of recent foraminifera1 fauna from the Dry Tortugas, Florida. (abs.). 

Geological Society ofAmerica Special Paper 101: 128-29. 

This analysis of total foraminiferal population is based upon 16 samples collected in 1960 

from the Dry Tortugas, Florida. Fourteen samples represent reoccupatiori of stations 

sampled by Cushman (1922) in his original description of the foraminiferal fauna. Q- 

modal factor-vector analysis indicates that the fauna is characterized by three assemblages 

(factors) which account for 89 per cent of the total information (sum of squares of all 

entries in data table). In general, these assemblages are characterized by a few dominant 

species. One of the assemblages, Assemblage 111, is directly related (r= +0.91 I) to depth 

of water. The total population of the 14 samples representing rcoccupation of Cushman's 

stations were compared with the total population indicated by Cushman (1922). 

Comparison was made using F-ratios and percentage of number of species occurring in 

both samples (Sc). F-ratios estimate degree of variation between samples, whereas Sc 

measures variation in species composition. F-ratios indicated that at seven of the 14 

stations there were significant differences in total population between the 1919 and 1960 

collections. Sc's ranged from 18.3 to 56.9 per cent and showed no relationship to degree of 

variation estimated by F-ratios. This variation in foraminiferal fauna between collections 

may represent either real changes in populations or apparent variations. If variations are 

real, they reflect changes in the ecosystem over the 41 years. If variations are apparent, 

they may represent either inability to resample microhabitats or vagaries in taxonomic 

discrimination. 

217. Lyons, W. G. 1980. Polyplacophora of Dry Tortugas Florida with comments on ischnochitor~ 

harrmeyeri. Bulletin ofthe Anlericnn Malacological Uhion, inc. 46: 34-37. 

450 specimens arid 14 species of chitons were collectcd during 1978-79 from a station near 

Garden Key. Only Acarlthochitona sp. and Slenol~laxprrrpur-ascet~s were relatively 

common. 

218. Mann, A. 1936. Diatoms in bottoni deposits from the Bahamas and the Florida Keys. Papers Fr-orrl 

the Tortugas Lahorator)~ 29: 121-28. 

Note: This is Appendix 1 to Calcareous shallow water marine deposits of Florida and the 

Bahamas by EIdon Marion Thorp . 

The twenty-four samples of calcareous sand collected by Doctor Vaughn in 1914 between 

Cape Florida and Key West and at Tortugas were examined at the time these samples were

eceived. A list of stations at which diatoms were collected is given, including Tortugas, 

with a list of the diatoms found. Species of the genus Mastogloia are very abundant in 

these Florida samples, and the author has found them to be so in all collections from 

Florida Waters. In other parts of the world they are relatively much less abundant. 

219. Manter, H. W. 1942. Gasterostomes (Trematoda) of Tortugas, Florida. Papers Tortugas Laboratory 

33: 1-19 (issued June, 1940). 


A report of the Gasterostomatus trematodes collected in 1930, 1931, and 1932 is given 

here. Fifteen species are reported; nine are considered new. 

220. ------ . 1934. The genus Helicometra and related trematodes from Tortugas, Florida. Papers 

Tortugas Laboratory 28: 167-80 (issued Mar. 1933). 


Observations on the trematode genera Helicometra, Helicometrina and a new related form 

are based on material collected at the Carnegie Biological Laboratory at Tortugas, Florida. 

The genus HeIicometra is represented at Tortugas by three species, H. execta, H. torta, and 

H. fasiata. The characteristics of each of these are described. A key is given to the species 

of the genus. H. execta is recorded from 6 additional hosts, making a total of 10. A 

mutilated specimen of H, torta showed this species has little or no power of regeneration. 

Helicornetra fasciatn from three hosts at 50 to 60 fathoms is reported for the first time from 

America. Metacercariae of Helicometrina nirnia are described encysted in the muscles of 

the shrimps, Lysrnata intermedia and Crangon formosum. Cercariae from Columbella 

mercatoria identified as Cercaria J of Miller were found to encyst readily in the muscles of 

L)'nnata intermedia. Helicon~etrina parva, a new species is described . 

221. --- . 1934. Some digenetic trematodes from deep-water fish of Tortugas, Florida. Papers 

Tortugas Laboratory 28: 257-345 (issued Jan. 16, 1934). 


The parasitic fauna of ocean depths is practically unknown. Extensive fish population 

occurs at all depths, very little study has been made on the helminths of these fish. During 

the summers of 1930, 1931, and 1932 collections were made of parasites from fisli trawled 

from depths varying from 40 to 582 fathoms at Tortugas, Florida. Most of these hauls 

were made about 10 miles south of Loggerhead Key. Fish taken from these depths were 

commonly parasitized by helminths and especially by trematodes. It was found that the 

trematode fauna of the deep-water fish is practically as abundant and as varied as is the rich 

trematode fauna of the reef fish. A description is given of 49 species of trematodes 

collected from approximately 90 species of fish from depths of 40 to 582 fathoms. 721 

individual fish were examined. Approximately 80 per cent of the host species were 

infected with trematodes, a percentage comparable with the degree of infection found in 

fisli of shallow water. One new subfamily (of the family Heterophyidae), I I new genera 

and 33 new species are described. Seven species of trematodes, from deep water only at 

Tortugas, are identical with forms well known from northern regions. Studies from shallow 

water at Tortugas show practically no similarity to northern forms. The deep-water 

trematode fauna is more like the surface fauna of Maine, Great Britain or Norway than like 

the shallow-water fauna at Tortugas, only a few miles away. This tendency to resemble 

surface trematodes of cold-water regions suggests that temperature is an important factor in 

the distribution of marine fish trematodes. This study emphasized the fact that a gradient of 

changing environment (such as depth) is reflected not only in the free-living population of 

a rcgion but also in their parasites. 

222. Marsh, G. 1940. The effect of light on the inherent E. M. F. of Valor~ia verztricosa. I. Intensity and 

time relations. Papers Tortrqas Laborator)~ 32: 65-84 (issued Oct. 1939).


The interpretation of the electrical changes produced in green plants by light has been 

retarded by the confused nature of the published results. In order to interpret electrical 

changes in a tissue in terms of some underlying process it is essential to obtain precise 

information concerning the distribution of E.M.F. within the tissue and the conditions of 

summation of thc potentials of the individual cells included in the electrical circuit. The 

present paper reports the effect of visible light at known intensities upon the inherent 

E.M.F. of the coenocytic alga Valonia veniricosa. Results indicated that when the intensity 

of incident light is altered, the inherent E.M.F. of an impaled Valonia cell undergoes a 

characteristic cycle of change with definite time relations, following which a steady level is 

reached. The steady level of E.M.F, plotted against the logarithm of the light intensity rises 

from the dark potential along a sigmoid curve to a maximum at about 250 foot-candles, 

then descends along a similar curve toward the dark potential. The decline in potential with 

light intensity beyond the maximum was reversible. No injury was detected at any 

intensity. The effect of intermittent light with equal light-dark periods was similar to that 

of continuous light of half the intensity, save in one experiment, wherein the effect was 

similar to that of continuous light of increased intensity. It is concluded that the effect of 

light on the E.M.F. is due primarily lo the release of oxygen in photosynthesis. 

223. ------. 1940. The effect of light on the inherent E. M. F. of Valonia ventricosa. 11. The relative 

energy absorption spectrum. Papers Tortuga.~ Laboratory 32: 99-120 (issued May 1940). 


The interest in the relative effectiveness of different wave-length bands of visible light 

upon bioelectric potentials centers about the question of the nature of the agent in the living 

cell which absorbs the radiant energy, and its contribution to the electromotive mechanism. 

For the green plants three principal lines of evidence have been adduced to support the 

conclusion that chlorophyll is the photosensitive agent. Results indicated that the steady 

E.M.F. of impaled Valonia ventrico.sa illuminated with light of limited spectral 

composition was matched with white light. The ratio of the intensity of white to that of 

filtered light for an E.M.F. match was independent of the magnitude of the E.M.F. matched 

and of the absolute intensities. The relative energy absorption is shown to compare fairly 

well for the filter series with the relative absorption of chlorophyll mixtures over the same 

spectral range calculated from the determinations of the absorption coefficient published by 

Zscheile. It is concluded that chlorophyll is the photosensitive material absorbing the 

radiant energy responsible for the effect of light upon the protoplasmic E.M.F. in Valonia. 

The chlorophyll system is, therefore, an intimate part of the electromotive mechanism. 

224. - . 1937. Effect of temperature upon the inherent potential of Valonia. Papers Torrugas 

Laborarory 31 : 1 - 16. 


The effect of temperature upon the potential is of prime importance in the determination of 

the nature of the underlying electrochemical process. The electromotive force of a system 

in thermodynamic equilibrium (including the diffusion potential) is proportional to the 

absolute temperature (Q 10 of 1.04 or less within the biological range of temperatures). 

The E.M.F. found across the protoplasmic layer of Valonia is not a thermodynamic one. It 

is produced by an oxidation-reduction system in flux equilibrium at phase boundaries 

within the cell. The E.M.F. is not primarily determined by the external medium. The 

influence of the salt content of the sea-water upon the inherent potential is fundamentally 

no different from the influence of the composition of the medium upon any other biological 

process, as respiration, it-ritability, contractility, etc., where specific electrolytes in different 

proportions condition, but do not cause, the process.

225. Mast, S. 0. 191 1. Behavior of the loggerhead turtle in depositing its eggs. Papers Tortugas 



The nesting behavior of a single loggerhead turtle is described. 

226. Matthai, G. 1915. Preliminary report on the comparattve morphology of the recent Madreporaria 

around Tortugas. Carr~egie Institution of Washington, Year Rook 14: 209. 

General observations were recorded on the common corals of the Tortugas. The only 

species that extruded larvae was Favia fragum. 

227. Mayer, A. G.. 1908. The annual breeding swarm of the Atlantic Palolo. Pa11er.c Tortugas 



The habits of the "Atlantic palolo" are quite similar to those of the palolo worm of Samoa 

and the Fiji Islands. The worms are, however, specifically different, the Atlantic palolo 

being Ew~ice fucata Ehlers, and the Pacific worm E. viridis Gray. The annual swarming of 

the Atlantic palolo has been observed only at Tortugas, Florida, although the worm is 

abundant in the Bahamas and other parts of the West Indies. 

228. -- . 191 1. The converse relation between ciliary and neuro-muscular movements. paper.^ 

Tortugas Labor-atory 3: 1-25. 


A series of experiments on marine invertebrates were conducted beginning at the Tortugas 

Laboratory, and later at Woods Hole, Mass, and the New York Aquarium involving the 

effects of ions of blood salts, magnesium, sodium, calcium, ammonium, potassium, and 

hydrogen on neuro-muscular systems in relation to maintaining ciliary movements. In each 

case they are the exact apposite of their effects upon ciliary movements of invertebrates 

studied. Studies were carried out on invertebrate organisms abundant at Tortugas 

including annelid larvae, Lirii~rlus, veligers , actinian larvae, larvae of the Atlantic palolo 

worm, Eunice fucata, and ctenophores, Cassiopea. Preliminary reports oi the research 

were published in the Biological Bull., Woods Hole, v. 17 (341-342); in the Proceedings 

of the Sac. for Experimental Biology and Medicine, 1909, No. 7, (19-20), and in the 

Carnegie Year Book for 1909, p. 152. 

229. -- 1914. The effects of temperature upon tropical marine animals. Papers Tol-tugas 



Tropical marine animals commonly live within 5" C. of their temperature of maximum 

activity and within 10" to 15" C. of their upper death temperature. In marine tropical forms 

even a few degrees of heat or cold cause a marked depression in movement. In tropical 

Scyphomedusae this depression of movement appears to augment about as the square of 

the change in temperature from that of the optimum. Time is an iinportant factor in these 

experiments, for animals can withstand a higher degree of heat if the temperature be raised 

quickly than if it be raised slowly. It appears that the reef corals at Tortugas, Florida, live 

in water which is commonly within 10" C. of their upper death-temperature, and if the 

ocean were heated to 38' C. (100.4" F.) only one species, Siderastraea radians, could 

survive. Next to Siderastraea radians the most resistant coral is S. sideren. It is associated 

in its habitat with Orbicella. annularis one of the most sensitive of the reef corals, which is 

killed at 14.1" and 36.8' C. In general, however, the corals of the shallow-reef flats, such 

as Siderastraea radians, Porites furcata, and Maeandr-a areolata, are the most resistant 

both to heat and cold, whilc those of deep water, such as Madrcpora /~alnmta, Eusirnilia 

knorri, and Oculirla diffuusa, are the least resistant. As a result, we are led to conclude that 

were the water cooled by an exceptionally prolonged norther to 13.9' C. for 9 hours,

Siderastraea radians, S. siderea, and Maeaizdra areolata would survive without apparent 

injury while Porites furcata, P, clavaria, Maeandra clivosa, and Favia fragum would also 

survive, but with more or less injury. This temperature would be fatal to Orbicella 

annularis, Porites astraeoides, and Madreporn ~nuricatcr (cervicornis). 

230. -. 1922. Hydrogen-ion concentration and electrical conductivity of the surface water of the 

Atlantic and Pacific. Pupers Tortugas Laboratorj' 18: 61-85. 


The hydrogen-ion concentration of sea-water was determined by placing 0.4 c.c, of 0.1 per 

cent of the red dye thymolsuphonemphthalein in 70 per cent alcohol, in a test-tube of 

resistance glass, 24 mm. in caliber, then adding sea-water so as to make up 30 C.C. of 

solution. A series of such tubes, ranging from 7.95 to 8.3 pH, was standardized by 

Professor J.F. McClendon and presented to the author who restandardized these tubes at 

intervals of two years by comparison with determinations of pH made by a Leeds and 

Northrup potentiometer. In order to avoid writing negative exponents, the synlbol "pH to 

indicate the negative logarithm of the hydrogen-ion concentration was devised. Despite its 

artificiality, one soon finds that the pH system gives a clearer idea of the alkalinity or 

acidity of a solution than does a direct expression of the hydrogen-ion concentration. In 

testing water, pH 7 would indicate practical neutrality; pH above 7, alkalinity; and below 7 

acidity. The carbon-dioxide tension of the sea-water was calculated from the pH and the 

temperature by the method devised by McClendon, Gault, and Mulholland (1917, Carnegie 

Inst. Wash. Pub. No. 251, p. 36). McClendon found that the pH of sea-water falls 0.01 for 

I" C. decline in temperature. The salinity of the sea is expressed in grams of total salts per 

1,000 grams of sea-water, and was determined by the well-known method of using a 

standard AgN03 solution with K2CrO4 as an indicator, and testing against a sample of 

standard sea-water obtained from Professor Martin Knudsen. Upon being taken from the 

sea, the water was tested for temperature and pH, and a sample was preserved for 

determination of salinity. In connection with these tests of hydrogen-ion concentration, the 

electrical conductivity of the sea-water off Tutuila, Samoa and Tortugas, Florida was 

determined by Kohlrauscli's method. At Tortugas, Florida, the conductivity of sea-water 

having 20.06 grams of chlorine in 1,000 grams of water, corresponding to a salinity of 

36.24, was determined by the same apparatus, and with a portion of the same KC1 solution 

used in Samoa. In lagoons such as that of Tortugas, Florida, and in closed shallow areas, 

McClendon found there was a diurnal variation in the pH, the water becoming more 

alkaline by day and relatively acid during the night. This was attributed to the effect of 

photosynthesis by plant life, which is active in daylight but ceases during the night. Over 

shallow regions, where the water may become impounded in tide-pools at low tide, the 

effect of photosynthesis is often very marked, the pH changing greatly while the 

temperature may change but little. The rise in pH was due to the loss of Co2 resulting from 

photosynthesis. 

231. - . 1914. The law governing the loss of weight in starving Cassiopea. Papers Tortugas 

Laboraror)~ 6: 55-82. 


The medusae were always starved in the purest sea-water which was either dipped from the 

ocean in glass or canvas buckets or pumped into glass reservoir tanks through hard-rubber 

pipes by means of a hard-rubber pump. The medusae were starved side by side in one and 

the same glass aquarium, but when this was impossible the aquaria were of similar size and 

form and were placed side by side, so as to be subjected to similar environmental changes. 

The decline in weight of two normal medusae of Cassiopea xanlachanu starved each in one 

liter of sea-water, changed once in 24 hours, and kept in the diffuse daylight of the 

laboratory at Tortugas, Florida, from June 8 to 20, 1913. One medusa, A, was starved in 

sea-watcr which had been passed through two glass funnels each holding two sheets of

Chardin filter paper. The other medusa, B, was starved in sea-water, which, in addition to 

having been filtered through the Chardin Filters, was also filtered through a bacteria-proof 

porcelain filter. It appears that all food had been removed from the water by Chardin 

filters and the medusa in the bacteria-frce sea-water starved more slowly than the one in the 

sea-water which had not been passed through the porcelain filter. 

232. --- . 1908. Marine laboratories, and our Atlantic coast. The American Naturalist 42: 533-36. 

In this general article by Dr. Mayer concerning the importance of marine laboratories along 

our Atlantic Coast, tbe Tortugas is mentioned as having a certain remoteness from the 

busy world and consequent freedom from interruption peculiarly favorable to the conduct 

of research. 

233. -. 1915. The nature of nerve-conduction in Cassiopca. Proceedings ofthe National Academy 

of Science I: 270-274. 


234. ------ 1917. Nerve-conduction in Cassiopea xamachano. Papers Tor-tugas Laborafory 11: 1-20. 


By means of Professor E.G. Conklin and the authorities at Princeton the author used the 

facilities at the Biological Laboratory in Guyot Hall, where the kymograph records taken at 

Tortugas were studied and the results tabulated. The object of this research was to obtain 

an accurate quantitative determination of the rate of nerve-conduction in natural and in 

diluted sea-water at constant temperature, and also to estimate the effects of various 

artificial sea-water solutions containing all or some of the sodium, magnesium, calcium, 

and potassium cations of sea-water. The effects of temperature upon nerve-conduction are 

also of great importance. These studies were carried out in June and July 1916, upon 

Cossiopea xamachann, a thizostornous scyphomedusa which is abundant in the salt-water 

moat surrounding Fort Jefferson at Tortugas, Florida. In summary, nerve-conduction is due 

to a chemical reaction involving the cations of sodium, calcium, and potassium. 

Magnesium is non-essential. Observations do not support the "local action" theory of Lillie 

(1916). 

235. ---. 1918. Nerve-conduction in diluted and concentrated sea-water. paper,^ Torrugas 



Ring-shaped strips of subumbrella tissue of the scyphornedusa Cassiopea xamachana were 

deprived of rnarginal sense-organs and placed in concentrated sea-water in order to 

determine the effect of concentration of electrolytes upon their rate of nerve-conduction. 

Experiments made in 1917 upon Cassiopa found that the rate has increased while the 

electrical conductivity has diminished. The injurious effects of concentrated sea-water 

upon regeneration and growth have been studied by Loeb, and by Goldfarb (1914), and 

there is a general resemblance between their curves and those in this paper for the rate of 

nerve-conduction, excepting that for regeneration somewhat dilute sea-water seems to be 

more favorable than nor~nal sea-water, whereas in nerve-conduction the highest rate is 

obtained in slightly concentrated sea-water . 

236. --- . 1908. A plan for increasing theefficiency of marine expeditions, marine laboratories and 

our Atlantic coast. American Naturalist 42: 533. 


237. ---- 1914. The relation between the degree of concentration of electrolytes of sea water and the 

rate of nerve conduction in Cassiopea. Papers Torrugas Laboratory 6: 25-54. 


If sea-water be diluted with distilled water, or with a 0.9 molecular solution of dextrose, 

thus preserving its normal osmotic pressure but reducing the concentration of the cations of 

sodium, magnesium, calcium, and potassium, the rate of nerve-conduction increases as 

dilution proceeds, becoming most rapid in 90 per cent sea-water + 10 per cent distilled 

water or dextrose. The sodium cation is an active stimulant for nerve-conduction. 

Experiments with the magnesium calion show that it is not a stimulant for nerveconduction. 

In very slight excess the potassium cation produces a permanently stimulating 

effect, as does sodium, but in denser concentration it produces momentary stimulation of 

the rate of nerve-conduction followed by depression. In all essential respects the effects of 

potassium are similar in kind, but more marked in degree, to those of sodiunl. 

238. ---. 1908. Rhythmical pulsation in Scyphomedusae.11, Papers Torfirgas Laboratorj 1: 113-31. 


The following paper presents thc results of a continuation of studies, the first report of 

which appeared in publication No. 47 of the Carnegie Institution of Washington, 1906. 

The present paper aims to correct certain errors in the previous report, and to announce 

some new results. Conclusions presented suggested that sea-water is a balanced fluid 

neither inhibiting nor stimulating pulsation in Cassiopea xamachann. The stimulus which 

causes pulsation is due to the constant formation of sodium oxalate in the terminal 

entodermal cells of the marginal sense organs. This sodium oxalate precipitates calcium, 

as calcium oxalate, thus setting free sodium chloride and sulphate which act as nervous 

stimulants. Pulsation is thus caused by the constant maintenance at the nervous centers in 

the sense-organs of a slight excess of sodium over and above that found in the surrounding 

sea-water. In Cassiopea the pulsation-stimulus is conducted by the diffuse nervous network 

of the subumbrella, and is independent of the rnuscles which may or may not respond to its 

presence by contraction. Strong primary nervous and muscular excitement followed by 

exhaustion and sustained muscular tetanus is produced in Lepas or in Cnssiol~ea by a 

solutioti containing the amounts and proportions of NaCI+KCI+CaC12 found in sea-water. 

239. -- . 1900. Some medusae from the Tortugas, Florida. Bulletin of tlze Museuni of Conl~arntive 

Zoology at Hanard College 37, no. 2: 1-82 . 

This extensive article on the medusae of the Dry Tortugas includes an alphabetical list of 

species described, table showing the wide geographical range on some Tortugas Medusae, 

and comparisons of the Tortugas Fauna with that of Southern New England, the Tropical 

Atlantic, Fiji Islands and Tropical Pacific. 

240. -- . 1902. The Tortugas, Florida as a station for research in biology. Scierrce (Washingror~, 

D.C.) 17: 190-192. 

The advantages of establishing a tropical research marine biological laboratory in the Dry 

Tortugas over other Caribbean sites are discussed. The nearness of the Florida Current, an 

extension of the Gulf Stream, to the Tortugas is a distinct advantage over other sites 

because of its richness in pelagics, especially small juveniles and larvae during the summer. 

Other sites are richer in coral, such as Jamaica, but they are further from the Gulf Stream 

and are impacted by land runoff. 

241. ----. 1918. Toxic effects due to high temperature. Papers Tortugus Lnboratory 12: 173-78. 


The experiments cited below appear to indicate that death from high temperature may he 

due to the accumulation of acid in the tissues. Reef corals from Tortugas, Florida, were 

kept at a constant temperature in warm ocean-water for 60 minutes in a thermostat, in the 

dark the temperature remaining constant within about 0.1" C, throughout the hour. In this 

manner the temperature was found that is just sufficient to kill the coral. The results are as 

follows: Acropora muricata 34.7, Orbicella ar~nularis 35.6, Porites astraeoides 35.8,

Porites clavariu 36.4, Maeandra areolata 36.8, Porites furcata 36.85, Favia fragurn 

37.05, Siderastrea radians 38.2. It is apparent that those corals which live in cool, 

relatively agitated water, free from silt, are those that can not withstand high temperatures, 

whereas those which live in the hot, silt-laden shallows near shore are, generally speaking, 

forms which can resist high temperature. Favia fragurn is however an exception. It seems 

possible that death from high temperature may be due to the accumulation of acid (possibly 

H2C03) in the tissues, the rate of formation of this acid being related to the rate of 

metabolism of the tissues. Thus animals of the same class having a high rate of 

metabolism, as measured by oxygen consumption, are more sensitive to heat and to C02 

than those having a low rate of metabolism. 

242. --- 1922. The tracking instinct in a Tortugas ant. Pa11er.r Tortugas Labomtory 18: 101-7. 


Monomoriuni destructor Jerdon, a tropicopolitan ant of East Indian origin, was identified 

in Florida . It is a small, reddish-brown ant, a great pest in the wooden buildings of the 

Tortugas laboratory. These pests have killed rats confined in cages within 24 hours. The 

experiments described were made on the flat wooden floor of the laboratory. To attract the 

ants, a number of recently killed houseflies were impaled upon a pin. The pin with its lure 

of flies was then thrust into the floor in front of a foraging ant, which would often pass 

within 0.25 inch of the lure without perceiving the flies; but if its course were such that it 

came appreciably nearer than 0.25 inch, the ant suddenly turned toward the flies, and 

without apparent excitement appeared to "inspect" them, spending a half minute or more 

crawling over them and stroking them with its antennae. This "finder ant" soon leaves the 

flies without carrying off any piece of them, but instead of moving off in the erratic and 

tortuous path it was pursuing before it found the flies, it now goes in a fairly straight path 

toward some crevice in the floor, out of which there soon pours an excited swarm of its 

nest-mates, who proceed toward the flies in a fairly straight path. When an ant returns to 

the nest it pursues a fairly straight path which is more or less right in direction, but when 

the ant has gone the correct distance, it begins to wander in more or less tortuous courses 

until it finds the nest. 

243. McClendon, I. F. 1917. Diurnal changes in the sea at Tortugas, Florida. Proceedings of the National 

Academy of Scier~ce 3: 692. 

The only diurnal change noted in the Gulf Stream was a change in temperature of about 1 

degree C and the resulting change in oxygen tension. However, marked differences were 

found in temperature, pH, C02, and Oz concentration in shallow water where light could 

reach the bottom The temperature, O2 concentration and 0 2 tension were lowest and the 

C02 concentration and C02 tension highest at 5 A.M. The temperature, O2 concentration 

and O2 tension were highest and C02 concentration and C02 tension were lowest at 3 P.M. 

during July at Dry Tortugas. The magnitude and exact time of minima and maxima varied 

from day to day, and varied a great deal with station location. Studies on the effects of 

these changes on organisms were made. The limiting factor for plants was nitrogen, while 

the limiting factor for animals was food and oxygen. 

244. - . 1917. Effect of oxygen tension on the metabolism of Cassiopea. Proceedings of the 

Natior~al Academy of Science 3: 715-16. 

Experiments on the effect of oxygen tension on the metabolism of Cassiopea were carried 

out at the Tortugas Laboratory by using the umbrella of Cassiopea to maintain a layer of 

cells in seawater at 30 degrees C. The metabolism varied with oxygen concentration. This 

may be true for all animals, however there is a distinction between the metabolism of 

vertebrate cells and Cassiopea. Vertebrate cells give out lactic acid when asphyxiated 

whereas Cassiopea may remain without oxygen for seven hours without giving out COz or 

any other acid. Although details of the experiments will be published elsewhere, it was

concluded that changes in the threshold of stimulation of the respiratory and basomotor 

centers may affect metabolism in Inan and animals. 

245. -. 1917. The effect of stretching on the rate of conduction in the neuro-muscular network in 

Cassiopea. Proceedings of the National Academy of Science 3: 703. 

The experiments on Cassiopea collected at the Dry Tortugas, tend to support the 

conclusions reached by Carlson, that stretching the nerve does not change the rate of the 

nerve impulse, and that the conducting substance itself, can be stretched and relaxed. 

246. ---- . 1917. The equilibrium of Tortugas sea water with calcite and aragonite. Proceedings of the 

National Academy of Science 3: 612.19. 

This report provides information on the continuing controversy on the solubility of 

Calcium chloride in sea water. The precipitation of CaC03 at Tortugas was studied by 

T.W. Vaughan, R.B. Doyle, and G.H. Drew. Drew observed that denitrifying bacteria, 

Pseudornonas calcis obtained from sea water was capable of changing calcium nitrate to 

calcium carbonate in culture media, and supposes that a similar process occurs in seawater. 

This study attempts to determine the nitrates or nitrites. If the pH is maintained (by plants) 

at 8.2 at the Tortugas, the introduction of calcite crystals would result in a lowering of the 

calcium content of Tortugas seawater by about 4.5%. 

247. -. 1914. Experiments on the permeability of cells. Papers Torrugas Laboratory 6: 123-30. 


One of the most important steps in the analysis of life was the discovery of oxygen. Ever 

since that time it has been known that animals absorb free oxygen and give it out in a 

combined form. In this experiment three methods of procedure were followed: (1) the use 

of cell masses as partitions (on eggs of Lyreclzir~us); (2) the use of quantities of eggs 

suspended in a liquid medium (on eggs of Fundrrlus); (3) experiments on individual eggs 

(of Arbacia). The permeability of the egg to ions and perhaps some other substances 

increases on fertilization. The unfertilized egg is perhaps in a dormant condition and the 

increase in permeability probably allows a rapid interchange with the surrounding medium 

necessary for activity (development). Whereas this supposed significance of permeability 

has not been proven, the sea- urchin;^ egg is not an exception. The relation of permeability 

to oxidation can hardly be determined until more is known about the mechanism of animal 

oxidations. These seem to depend on structure since complete oxidations cease when 

structure is completely destroyed. Reference is made only to oxidations resulting in the 

formation of COz. Oxidizing enzymes such as tyrosinase, which are independent of 

structure, do not completely oxidize the substances acted on. 

248. - . 191 1. On adaptations in structure and habits of some marine animals of Tortugas, Florida. 

Papers Torrugas Lrrboratory 3: 55-62. 


This article discusses the habits of some marine animals of the Tortugas. Many of these 

animals were thigtnotactic and remained in glass tubes rather than in the open. They 

learried to find the tubes when removed from them. Such was the case with five species of 

the Alpheidae, one of the Pontoniidae, T)'pion rorrugae Rathbun, and Gonodactylus 

aertedii. All the anemones were thigmotactic on their bases. These same animals were 

heliotropic. The crustaceans were negativcly heliotropic and the anemones kept their bases 

from the light, while Cradactis im+dilis Hargitt hid all but the tips of the fronds and 

tentacles from the light. In removing its base from the light, Sroichacris helianthus, which 

lives on coral heads, makes snail-like movements similar to Metridiiam, while Cradactis, 

which lives in holes in decayed coral heads, crawls on its tentacles.

249. ---- 1918. On changes in the sea and their relation to organisms. Poper.s Tortugos Laborntot-y 

12: 213-58. 


The sea and air form the circulating media for the living organisms of the world. The local 

composition of the sea is distinctly affected by living organisms. The local changes in the 

composition of the sea are the subject of the present paper. These changes are due chiefly 

to organisms, but partly to nleteorological causes. The water evaporated is returned with 

addition of fixed nitrogel] from electric discharges or falls on the land and is returned with 

various salts, chiefly CaC03, and with fixed nitrogen and other products of organisms. 

Various seaweeds absorb COz thus leaving an excess of CaC03, which has a very low 

solubility and is cotlstantly being precipitated in certain warm seas, and is precipitated 

within the bodies of organisms in the surface waters of all seas. In working out the relation 

of H-ion concentration (pH) to the solubility of CaCOi in sea-water, it was found that all 

sea-water is supersaturated with CaCO,, and will lose sonle of it if shaken with calcite or 

aragonite crystals. The pH is influenced by plant and animal life and arises at Tortugas to 

8.35 during the day over well-lighted bottoms rich in vegetation, and falls to 8.18 during 

the night. It may be said, therefore, that conditions in shallow water over eelgrass or other 

seaweed or corals (with symbiotic algae) favor the precipitation of CaCO3. The question 

arises whether the occasional high pH of Tortugas sea-water is sufficient to explain the 

precipitation of CaC03. The author's experiment showed that if thc pH of sea-water should 

be maintained (by the action of plants) at 8.2 while it was agitated with calcite crystals, the 

loss of CaCO, would be about 0.001 N, or 0.0005 M, or 0.1 gram per liter. This would 

cause a deposit of 10 kg. per square meter of bottom in water 100 meters deep. This would 

cansc a lowering of the total calciu~n content of Tortugas sea-watcr by about 4.5 per cent. 

The actual precipitation of CaCO, was most noticeable in the Marquesas lagoon. At 4 

p.m., July 30, the pH was 8.46 and there was a precipitate of CaCO, coming down in the 

water and encrusting the eel-grass. 

250. -, 1917. The sta~idardization of a new calorimetric method for the detern~ination of the 

hydrogen-ion concentration, CO, tension and COz and 0, content of sea water, of animal 

heat, and of C02 of the air, with a sunnnary of similar data on bicarbonate solutions in 

general. Journal of IMiological Chemistry 30: 265-88. 

Experiments were conducted on pH, C02 tension, COz, and oxygen content of Dry 

Tortugas seawater and seawater from other oceanic areas using a Leeds & Northrup 

potentimeter and a 0.1 KCL calo~ncl electrode. It was found that neither the salinity nor 

the alkaline reserve in seawater of the tropical or temperate oceans cl~atige sufficiently to 

noticeably change the relation of pH to CO, tension, although the alkaline reserve does 

change sufficiently to affcct thc total C02 greatly. 

251. McClendon, 1. F., C. C. Gault and S. Mulhollantl. 1917. The Iiydrogeii-ion concentration, C02 

tension, and C02 content of sea water. Papers Tortugns LaDorarory 11: 21-69. 



252. Meeder, J. F. 1979. Corals and coral reefs of the Dry Tortugas, Florida. in Guide to sedimer~tntion 

for the Dry Tom~gas. R.B. Ilalley (Compiler), 46-47. S.E. Geological Society Pub. 

This paper presents a description of the corals from two localities in the Dry Tortugas, on a 

back reef environment of a fringing reef ncar Garden Key, and the second, a series of patch 

reefs off Loggerhead Key. The general setting, ecology, distribution, and types of corals 

are discussed for each locality. Forty-one of the forty-two specics of corals reported at the 

Tortugas are covered in a ficld key and latcr described in this paper.

253. Miller, H. M. Jr. 1926.1929. Behavior of trematode larvae. Carnegie Institution of Washington, 

Year Book. 

Note: Published as follows: 1926, v.25, p. 243-244: 1929, v. 28, p. 295. 

Anatomical/morphological descriptions of six larval trenieatodes infesting the mollusk, 

Cerithiuni litteratuni, taken from Bird Key Reef Porites beds, were provided. Percentages 

of occurrence of the 6 cercariae were given. The behavior of the members of this 

morphological group are described in detail, including aspects of their life history. 

254. Miller, R. A. and H. B. Smith. 1931. Observations on the formation of the egg of Echi11ornetra 

lacunrer. Papers Tortugas Laboratory 27: 47-52. 


The study of the ovaries of Echinori~erra was undertaken in the hope that by the use of 

some of the newer cytological methods, it might be possible to extend our knowledge of 

the processes of oogenesis, particularly of those concerned in the formation of yolk in the 

chinoid egg. This paper presents the observations that have been made, and the 

conclusions that have been drawn. It was found that in Echinonietra lancunter, 

undifferentiated cells along the wall of the ovary are uniform in appearance, although they 

are destined to develop into two entirely different kinds of cells. As development 

proceeds, some of the cells become oogonia, while others enlarge and disintegrate to form 

deutoplasmic bodies. These are more coarsely granular than was the cytoplasm from 

which they were formed. The granules exhibit different affinities for stains. Fully formed 

nutritive spheres are of two kinds, granular and non-granular. The former are composed 

entirely of cytoplasmic, or of both cytoplastic and nuclear material. The nutritive spheres 

group themselves around the oogonia in a follicular arrangement. Eventually the nutritive 

spheres enter the egg and disappear as such, forming the yolk content of the cytoplasm, 

which becomes homogeneous and evenly granular. The deutoplasmic granules are smaller 

and more diffuse in the mature eggs than in the oocytes. The nutritive spheres have been 

shown to be composed of phospholipins suspended in a homogeneous medium. It is 

probable that they are largely lecithin in content. Their origin is not known. 

255. Millspaugh, C. F. 1907. Flora of the sand keys of Florida. Coluri~bian Museun~ 118, no. (Bot. Ser. 

2): 193.245. 

A list of species and details of vegetation, as well as elevational descriptions and size 

dimensions of sand keys westward of Key West, including theTortugan group, were 

compiled by the author during the winter through spring of 1904. 

256. Mitchell-Tapping, H. J. 1981. Particle breakdown of recent carbonate sediment in coral reefs. 

Florida Scientist 44, no. 1: 21-29. 

Skeletal particles of the major components of the carbonate sediment of the reef shoal 

environment were examined using the scanning electron microscope. This examination 

revealed no set pattern of skeletal breakdown according to microarchitectural structure, as 

postulated by the Sorby principle, but that such a breakdown depends on mineralogical 

composition, wall thickness, grain size and pattern, density, and the amount of cementation 

and bloerosion. To investigate general particle-size abundances and deficiencies in 

carbonate sediment, samples were taken from the reef crest, back-reef-rubble and opensand 

ecozones of the reef shoal environments of sites from the Bahamas, Dry Tortugas, 

Lower Florida Keys, Grand Cayman Island and the US. Virgin Islands. Size analyses of 

these samples showed that the sediment is moderately sell-sorted, coarsely-skewed and 

leptokurtic. Although particle-size abundances (or modes) exist in each individual site, 

there is no particular particle-size abundance that is common to all the sites. It is inferred 

that the particle abundances (or modes) for each site are a product of the sorting potential 

of the wave energy and that this sorting potential is the major control of the breakdown of

sand-sized skeletal particles rather than the microarchitectural structure as proposed by the 

Sorby principle. 

257. Moritz, C. E. 1936. Embryology of the sea-hare, Aplysiu protea and of Cmrigon nrmillatus. 

Carnegie 111stitution of Washingioi~, Year Hook 35: YO. 

Observations from aquaria are recorded on the early development of Aplysiu, from embryo 

to 5 days beyond hatching. Adults were collected from the moat on Garden Key and Bird 

Key Reef. 

258. Multer, H. G. 1975. Field guide to some carbonate rock environments; Florida Keys and Western 

Bahamas. Fuirleigh Dickblson U~~iversii~~ 40: 175 pp. 

This report presents the most recent compilation on Holocence sediments in the western 

Bahamas and in the Florida Keys including the Dry Tortugas. Selected geologic literature 

pertinent to local environments is noted within the text, with full bibliographic citations 

following each subject area. Carbonate sand beach and beach rock environments from the 

Florida Keys, Bimini, and Loggerhead Key are compared. In summary, Holocene 

sediments of these areas today present a vast array of textures and constituent particles 

charactcristic of environments which have been subjected to fluctuating sea levels and 

storm action. Such data may be used to interpret ancient environments. 

259. ---- 1971. Holocenc cementation of skeletal grains into beachrock, Dry Tortugas, Florida. 

Curbonote Co~zent. 0. P. Bircker. Baltimore, Maryland: Johns Hopkins Press. 

A discussion is presented on the origin of beach rock at Loggerhead Key, Dry Tortugas. 

Present evidence suggests that cementation is due to alternate wet and dry salt water spray 

conditions with skcletal grains providing nuclei for precipitation from a supersaturated 

calcium carbonate solution. The limited ground water conditions and lack of grain solution 

for providing aragonite cement are two evidences in favor of the above cited evaporation 

origin for the cement in this rock. Ginsburg (1953) reached similar conclusions for beach 

rock in the same area. 

260. Murphy, L. E. 1993. Dry Tortugus Natioiml Park, Submerged Cultural Resources Assessnlent, L. E. 

Murphy. Submerged Cultural Resources Unit, Southwest Region, National Park Service, 

Santa Fe, New Mexico. 

This volume describes and assesses the known and potential archcological resources in 

Fort Jefferson National Monument, later redesignated Dry Tortugas National Park. The 

emphasis is on submerged cultural sites, particularly shipwrecks. The importance of 

linking the natural resources with submerged cultural resources is provided by identifying 

the biological influences on the cultural resources. The Dry Tortugas and South Florida 

geological development and environmental succession is summarized with focus on the 

postglacial development of the Florida-Reef Tract, depositional environments, coral reef 

and sand key development as well as Late-glacial and Postglacial succession of 

environments. An overview on the physical oceanography of the eastern Gulf of Mexico 

concentrating on the Dry Tortugas with emphasis on currents and climate that affect 

shipping vessel casualties and site preservation is provided. Recommendations for future 

research and resources management are given. 

261. Nance, J. M., E. F. Klima and F. J. Patella. 1986. Review of the Tortugaspb~k shrimpfishery from 

Mny 1984 to Decen16e1- 198.5, Galveston, Texas, NOAAlNMFS, Southeast Fisheries 

Center, Galveston, Texas. NOAA Tech. Memo. 177. 

Commercial pink shrimp fishing data from the Tortugas (Dry Tortugas Islands, Florida) 

fishery were reviewed for biological year 1984 (May 1984-April 1985) and the first 8 

months of biological year 1985 (May 1985-Deccmber 1985). Pink shrimp landings were 

just over 11.0 million pounds in biological year 1984 with 17,000 days of fishing

expended. This computed to a CPUE value of 643 pounds per day. Pink shrimp landings 

for biological year 1985 are estimated to be around 9 million pounds with 15,000 days of 

fishing expended. The predicted CPUE value for 1985 should be around 600 pounds per 

day. Biological year 1984 experienced two extended periods of pink shrimp recruitment 

into the Tortugas fishing grounds. 

262. O'Neill, C. W. 1976. Sedimentology of East Key, Dry Tortugas (abs.). Florida Scientist 39 (Suppl. 

1 ), Fortieth Annual Meeting of the Florida Academy of Sciences at Eckerd College, St. 

Petersburg, Florida March 18,19,20, 1975: 10. 

East Key of the Dry Tortugas rests on a large crescent shaped hank and oscillates about a 

stable core in response to seasonal variations. A simple strandldune plant co~nmunity is 

largely responsible for short term stabilization of this central core. Historical studies 

covering 200 years show that East Key varies in its bank position on long term basis and 

has in the past been of much greater areal extent. In addition, historical records show the 

Dry Tortugas group has decreased in extent from 11 keys to the present seven. This effect 

is thought to be due to a combination of eustaic sea level change and storm degradation. 

263. -, 1976. "Sedimentology of East Key, Dry Tortugas, Florida." University of South Florida. 

Ph.D. Dissertation. 

The Dry Tortugas platform is a complex of reefs, banks, and shoals which lie 65 nautical 

miles west of Key West, Florida. This reef platform most closely rescrnbles the resorbed 

reef of Maxwell's (1968) classification. A review of historical records covering a 463-year 

period indicates that the Dry Tortugas island group is undergoing progressive degradation 

and has been reduced from 11 rocky islets in 1513 to 7 at present. This reduction is 

thought to be due to the combined effect of episodic events and the Holocene 

transgression. East Key maintains a position on the windward segment of the Tortugas 

group. Short-term changes to the key are basically in response to regular seasonal variation 

and cause the island to oscillate to the north and south about a plant-stabilized core. Longterm 

changes reflect the effect of episodic events and entail inundation, major shifts 

inposition, and changes in the orientation of the key. Mean grain size of East Key 

sediments is near 0.900 or coarse sand. The sediments are nearly without exception 

moderately well sorted with an average value of 0.570. The mean skewness value is -0.22 

(coarsely skewed). Texture was compared to morphology. The most striking correlation 

occurred on the foreshore where there is a distinct tendency for sediments to become 

coarser moving seaward from the berm. The average percentage composition of East Key 

sediments, by constituent, was found to be 52 percent Nalbneda, 35 percent coral, 6 

percent Mollusca. 3 percent coralline algae, I percent Foraminifera, 1 percent echinoid 

fragments and 2 percent miscellaneous plus unknown. The most significant correlation 

between composition and ~norphology occurred on the foreshore, where percent Halimeda 

generally increased seaward from the berm. This general increase in percent Halbneda 

correlates with a tendency for sediments to become coarser moving seaward from the berm. 

264. Ogden, J. C., W. C. Jaap, J. W. Porter, N. P. Smith, A.M. Szmant and D. Forcucci. 1993. 

SEAKEYS: A large scale interdisciplinary study of the Florida Keys Reef Tract (abs.). 

Ptnceedings of the Seventh Internatio~~nl Coral Reef Synposiunt, v. 2. Mangilao, Guam: 

University of Guam Marine Laboratory. 

The SEAKEYS program has established a research framework which encompasses the 

large geographic and long time scales of natural marine processes and ecosystem variation. 

The core of the program is a series of instrumented, satellite-linked monitoring stations 

which span the 220 mile coral reef tract. Mesoscale physical oceanographic studies are 

concentrated in the major channels potentially linking Florida Bay and the population 

centers of the Keys with the reef tract. Simultaneously, nutrient studies are probing the 

possibility of sewage and agrichemical contamination, coniplicated by natural sources of

nutrients. A series of long-term photomosaic stations have tracked coral community 

dynamics for more than 5 years. The design of the SEAKEYS program may provide an 

example for long-term research on coral reefs elsewhere. 

265. Osburn, R. C. 1914. The Bryozoa of the Tortugas Islands, Florida. Papers Tortugas Laboratory 5: 

18 1-222. 


In the summer of 1908 the writer had the privilege of spending the month of June at the 

Carnegie Institution Laboratory of Marine Biology on Loggerhead Key of the Tortugas 

Islands. The entire period was devoted to a close search for the bryozoa inhabiting the 

shallow waters about the reefs, on the piles of the old government dock on Garden key, in 

the moat of old Fort Jefferson on the same key, and in dredging the shallow waters about 

the islands down to 22 fathoms. Comparatively little work has been done on the bryozoa 

of the Florida and West Indian regions. By comparing with lists from other regions where 

the bryozoa have been carefully worked, it will be seen that the bryozoan fauna of the 

Tortugas and of the Florida-West Indian regions is fairly rich in species and fairly 

representative of tropical and semi-tropical regions. 

266. Payne, F. 1937. Early development of Ptychodera bahamensis. Papers Torfugas Laboratory 31: 

71-76 (issued July, 1936). 


The author records some observations made on Ptychodera bahamensis during the summer 

of 1933 while working at the Carnegie Laboratory at Tortugas, Florida. The animals were 

found most abundantly on the coral reef near Fort Jefferson in water which at low tide was 

from one to two feet deep. The larvae described by Weldon, Morgan, and Stiansy and 

assigned to Pt)'chodera bahrnemis do not agree with the author's own observations of the 

larvae of Ptychodera bahamensis which have been followed from the fertilized egg. It 

seems conclusive that more than one species is involved or that errors have been made on 

the part of someone. Even though the author has not followed development through to 

metamorphosis, it seems clear that the tornariae, described by Weldon and Morgan are 

different from the tornaria of Ptychodera bahamensis, assuming his identification is 

correct. 

267. -- . 1933-1938. Embryology and cytology of the balanoglossid, Ptychodera bahamertsis.. 

Carnegie l~utitution of Washington, Year Book. 

Note: published as follows: 1933, v. 32, p. 277-78; 1938, v. 37, p. 84. 

Embryological work was conducted on the supposed protochordate, Ptychodera 

baharnensis, starting with the fertilized egg. Development was followed as far as the 

tornaria. 

268. Pearse, A. S. 1934. Animals in brackish water ponds and pools at Dry Tortugas. Papers Dry 

Torrugas Laboratory 28: 125-42 (issued Dec. 1932). 


During the summer of 1931 two ponds and three pools were studied on Long and Garden 

Keys, Dry Tortugas. Because of the remoteness of the Dry Tortugas, the animals which 

live in these isolated habitats are of particular interest. Results were as follows: Five 

brackish water pools and ponds were studied on Long and Garden Keys. These were 

variable in salinity and temperature, and limited in extent. They showed various stages of 

evolution from sea to fresh water and conditions of life in them were more or less severe. 

The animals in the ponds and pools were resistant to environmental variations. They lived 

in salinities between 0.6 and 6.4 per cent and endured temperatures above 42" C. Some of 

the small bodies of brackish water on Dry Tortugas contained curious mixtures of marine 

and fresh-water animals. Callinnassas, marine snails, mullets, and needle-fishes lived with

dragon-fly nymphs, water boatmen, surface bugs, aquatic beetles and midges. Lists of the 

aninials collected and observed in each pond or pool are given. Each pond or pool 

contained certain characteristic animals, which had become dominant in that particular 

habitat. As the small pools trended toward fresh water, insects were increasingly dominant 

in them. Insect populations show great pressure and spread into all available habitats. 

When small pools are cut off from the ocean and are gradually transformed into fresh-water 

habitats, insects are the pioneers and soon become dominant. 

269. - 1934. Freezing points of bloods of certain littoral and estuarine animals. Papers Tortugas 

Luboraror)' 28: 93-102 (issued Dec. 1932). 


It is well known that the bloods and body fluids of marine invertebrates and elasmobranchs 

have about the same osmotic pressure as the ocean water in which they live, though the 

salt content of such fluids is usually a little lcss than that of the surrounding medium. 

Those of teleost fishes (marine, fresh-water and land) and of fresh-water and land animals 

generally have osmotic pressures which are much below those of sea-water. Littoral 

crustaceans and fishes are of particular interest because they represent various stages of 

adjustment to life on land. At Tortugas, conditions are particularly favorable for the study 

of such adjustments for the littoral group. The observations cited in this paper appear to 

justify the conclusion that crabs and fishes which take up terrestrial life or air-breathing 

have bloods of lower osmotic pressures than coniparable marine or fresh-water animals. 

The attain~nent of land life by marine animals is apparently associated with a reduction in 

salinity and stabilization of the contents of the blood. 

270. ---. 1934. Inhabitants of certain sponges at Dry Tortugas. Papem Tortugas LaDol-utory 28: 117- 

24 (issued Dec. 1932). 


Many sponges are veritable living hotels. Their canals are densely populated with a variety 

of animals and some species have not been found elsewhere. During the summer of 1931 

the writer studied the animals which occurred in five species of sponges at Dry Tortugas. 

The number of animals which live in big sponges is enormous. Though each sponge 

appears to be occupied to its full capacity, the number of animals per cubic centimeter of 

sponge is apparently influenced by depth and the size of the sponge itself. In loggerheads 

there are relatively more guests in smaller sponges and in deep water. Arnong the species 

of sinall sponges there are striking differences in the number of animals present and these 

may be due to inherent qualities. For example Steriiatnmerlia foerirlu (Schmidt) contained 

very few animals, whereas the slightly smaller Spoi~gia officinalis L. was crowded. 

271. ---- 1918. Notes on certain amphipods from the Gulf of Mexico, with descriptions of new 

genera and new species. Proceedings of the 0.S. Natiortal M~r.retrrn 43, no. 1936: 369-79. 

This is a report on a portion of the amphipods from the Gulf of Mexico in the collection of 

the United States National Museum. The collections are from several sources and extend 

over a long period of years. The greater part of them had not yet been examined. Those 

described were taken chiefly by the steamers Fish Hawk and Albatross of the United States 

Bureau of Fisheries. 

272. --- 1929. Observations on certain littoral and terrestrial animals at Tortugas, Florida, with 

special reference to migrations from mal-ine to terrestrial habitats. paper:^ Tortugas 



In the past, various types of animals have migrated from the ocean into fresh-water streams 

or lakes, and from there gained a foothold on land. On the shores of all oceans, animals 

may be found which are partially adjusted to life in fresh water or on land. The Dry

Tortugas contain no fresh water, and therefore offer an excellent opportunity to study 

littoral animals which have no immediate contact with fresh-water habitats. In summary, at 

Tortugas certain reef, beach and land aniruals were studied with reference to migrations 

from sea to land. Hermit crabs which have become more or less adjusted to life on land 

show a progressive reduction in the number of gills. Crabs which have migrated landward 

show a progressive lessening of gill-volume. Beach animals which show any landward 

trend usually live longer when kept in air than when kept in fresh water. Animals which 

have attained some degree of ability to live on land have often also acquired a greater 

degree of ability to resist the extraction of constituents of body fluids into fresh water. 

Animals which migrate from the sea and become established on land do not do so on 

account of one "lure" or one "danger." Each habitat has certain advantages and certain 

disadvantages. A continually changing animal must continually make adjustments to a 

contitiually changing environment, and when it migrates to a new habitat, must make many 

compromises between new advantages and dangers, old necessities and new requirements, 

and old habits and new abilities. 

273. ----. 1934. Observations on the parasites and commensals found associated with crustaceans and 

fishes at the Dry Tortugas, Florida. Papers Tortugas Laboratory 28: 103-15 (issued Dec. 

1932). 


During the summer of 1931 the writer had opportunity to conduct post ~nortetns on various 

crustaceans at Dry Tortugas from June 3 to August 22. Dr. Waldo Schmitt furnished and 

identified many of the specimens. The following parasitic isopods were taken from fishes: 

2 Cyr~zothoa oesturm (L.) from the gill cavity of Caranx ruger (Bloch), July 26; 1 

Rociiteia sigrlata Schioedte and Meinert from the gills of Promicrops itaiara Lichtenstein 

in July and 1 from the gills of Lutiar~irs analis (Cuvier et Valiencennes), June 27; 8 

Excorallana tricornis (Hansen) from the nose of Prornicrops itaiara Lichtenstein, July, 

and 31 from the gill cavity of Epinepheltas moro (C. et V.). The occurrence of the 

parasites and commensals associated with crustaceans depends upon a variety of factors - 

host specificity; habitat; habits, structure and physiology of hosts and parasites, etc. In 

general the greatest number of species of parasites occurred in or near the littoral zone. 

However, great numbers of parasites per host were encountered among some land 

crustaceans. 

274. --- . 1929. Two new mites from the gills of land crabs. Papers Tortugas Laboratory 26: 225-30. 


During July and August 1928, mites were found at Dry Tortugas on the gills of the land 

hermit crab, Cenobita diogenes (Latreille), and on the Nassau crab, Gecarcinus lateralis 

(Freminville). These crustaceans visit the ocean only once each year when they hatch out 

their young. No mites were found on the gills of the ghost crab, Ocypoda albicans (Bosc), 

which often visits the ocean and bathes its gills. Mixed or pure cultures of bacterial 

populations are incapable of precipitating CaCOn in a sea-water medium if KNOl and 

organic matter as the sugars or similar forms free from calcium are added to the medium. A 

number of different forms of bacteria in the sea possess the power of precipitating CaC03 

in appropriate media containing a large excess of soluble salts, but only under such 

conditions. Among such organisms there is great variability to perform the task in 

question, depending on the composition of the medium. 

275. Perkins, H. F. 1908. Notes on the medusae of the Western Atlantic. Papers Tortugas Laboratory I: 

133-56. 


The Marine Biological Laboratory in the Dry Tortugas is well situated for the study of 

many of the lower marine animals, their behavior, and the conditions of life, particularly

the coelenterates. One quite unique feature occurs in the Tortugas in the presence of the 

old fortification and surrounding moat of Fort Jefferson. The moat affords remarkably 

favorable conditions for the growth and ~nultiplication of the lower forms of plants and 

animals sheltered by the sea-wall, its shallow water warmed by the sun and kept from 

stagnation by the agitation and partial change of the tides. The writer has for several years 

been interested in the causes of migration and segregation of Medusae and has had the 

privilege of examining specimens of this genus from Jamaica, and has studied the 

characteristics of the specimens found in the Baha~na Islands and at the Tortugas. There 

was less of peculiarity in all the surroundings, the temperature of the water, storm 

influence, and food supply being normal for the shores of coral islands. The only points of 

difference to be noted in the medusae are with reference to size and color-pattern. The 

main features of the two species, Cassiopea xamuchmla and Polyclonia frondosa are 

presented. 

276. Petrovic, C. A. and J. King Jr. 1973. Bird records from the Dry Tortugas. Florida Field Naturzllist 

1, no. 1: 5-8. 

During a visit March 25 to April 4, 1967 to the Tortugas, a total of 70 species of mostly 

land birds were recorded. Detailed observations on twenty specimens rarely seen at the 

Tortugas are presented. 

277. -- . 1972. Common elder and king rail from the Dry Tortugas Florida. Auk 89, no. 3: 660. 

The authors watched the early spring migration at the Dry Tortugas Islands, which lie in 

the Gulf of Mexico about 70 miles west of Key West, Florida, and recorded 70 species, the 

majority land birds. The two records reported here represent significant additions to the 

species known distribution . 

278. Phillips, A. H. 1917. Analytical search for metals in Tortugas marine organisms. Papers Tortugas 

Laboratory l I: 89-93. 


This study concerns tlie problem stated in Year Book no. 14, page 193 of the Carnegie 

Institution of Washington. A large number of specimens were collected to be analyzed for 

metals. The metals determined were iron, manganese, zinc, copper, and lead. For the 

determination of zinc, copper, and lead, when the dried material was sufficient, 20 grams 

were used as a sample; when it was not possible to use 20 grams, the results are all 

calculated to 20 grams.. 

279. -- 1922. Analytical search for metals in Tortugas marine organisms. Papers Tortugas 



Included in the material collected at the Tortugas and analyzed for metals (some of the 

results of which were reported in the annual report of the Carnegie Institution for 1917) 

was a brown spotted holothurian. Stickoprrs n~obii, which was analyzed by the methods 

there indicated. The element vanadium was found in the holothurian material and 

heretofore has never been reported from seawater. Vanadium has been reported from 

freshwater and in the blood of an acidian from the Bay of Naples. This vanadium content 

of the blood does not seem to be a characteristic of all acidians, as two other species from 

the Tortugas yielded no vanadium, neither did two other species of holothurians yield 

vanadium. Two species, a chordata and a echinoderm contained vanadium, indicating that 

other forms [nay use vanadium as an oxygen carrier in their vascular systems. The source 

of vanadium in sedimentary rock and coals has always been somewhat of a puzzle. It is 

possible that such forms as Stickopus mobii may concentrate vanadium and that in depth, 

could easily be fixed and held as a constituent of the sedimentary rocks thus formed.

Another possibility of the fixation of vanadium under the above conditions is the presence 

of hydrogen sulphide which is constantly liberated from muds of mangrove lagoons. 

280. -, 1918. A possible source of vanadium in sedin~entary rocks. Arnericar~ Journal of Science 

46: 473-74. 


281. Pichard, S. L. and J. R. Paul. 1991. Detection of gene expression in genetically engineered 

microorganisms and natural phytoplankton populations in the marine environment by 

messenger RNA analysis. Applied Em~ironmentul Microbiology 57, no. 6: 1721 -27. 

A simple method that combines guanidinium isothiocyanate RNA extraction and probing 

with antisense and sense RNA probes is described for analysis of microbial gene 

expression in planktonic population. Probing of RNA sample extracts with sense-strand 

RNA probes was used as a control for nonspecific hybridization or contamination of 

mRNA with target DNA. This method enabled detection of expression of a plasmidencoded 

neomycin phosphotransferase gene (nptII) in as few as 10 super(4) Vibrio cells 

per ml in I00 ml of seawater. We have used this method to detect expression of the 

ribulose-I, 5-hisphosphate carboxylas large-subunit gene (rbcI) in Syr~echococcus cultures 

and natural phytoplankton populations in the Dry Tortugas, Florida. During a 36-h die1 

study, rbcL expression of the indigenous phytoplankton was greatest in the day, least at 

night(1100,0300, and 0100 h, and variable at dawn or dusk (0700 and 1900 h). These 

results are the first report of gene expression in natural populations by mRNA isolation and 

probing. 

282. Pitts, R. F. 1936. Clearance values of sucrose and creatinin in the kidneys of the red grouper, 

Epinephelus striatus. Camegie instirutior~ of Washington, Year Book 35: 90-91 

Studies were made on the excretion of urinary creatine nitrogen from the red grouper, 

E/hephelus striarus. 

283. Plan Development Team, Reef Fish Management Plan South Atlantic Fishery Management Council. 

1990. The potential of marinefishery reserves for- I-eeffish management in the U.S. 

southern Atlantic, Coastal Resources Division. 

Marine fishery reserves (MFRs), areas with no consumptive usage, are recommended as a 

viable option for management of reef fisheries in the U.S. southern Atlantic region. MFRs 

are designed to protect reef fish stocks and habitat from all consumptive exploitation within 

specified geographical areas for the primary purpose of ensuring the persistence of reef fish 

stocks and fisheries. Fishery reserves are intended to protect older and larger fishes. This 

will benefit reef fisheries by protecting critical spawning stock biomass, intra-specific 

genetic diversity, population age structure, recruitment supply, and ecosystem balance 

while maintaining reef fish fisheries. The MFR concept is easily understandable by the 

general public and possibly more easily accepted than some other management strategies. 

Fishery reserves provide some insurance against management and recruitment failures, 

simplify enforcement, and have equitable impact among fishery users. Data collection 

needs solely for management are reduced and management occurs without complete 

information and understanding about evesy species and interaction. Use of fishery reserves 

will establish US. leadership in producing model strategies for cooperative international 

reef resource management in the Caribbean. Large resident fishes that wander out of 

reserved can help maintain certain trophy fisheries. MFR sites with natural species 

equilibrium will allow measurement of age, growth, and natural mortality for fisheries 

purposes and will provide a basis for other educational, economic, and scientific benefits. 

Because there is no fishing within MFRs, impacts of hook and release mortality are 

eliminated and the temptation for incidental poaching is reduced. A mixed management 

strategy is recommended where 20% of the shelf is MFR, while the remaining 80% is

managed for optimal yield by any of several traditional options. Coordinated fishery 

reserve efforts in state waters would enhance the benefits of MFRs. Obstacles to fishery 

reserves includc automatic resistance to new approaches in US. marine fisheries, 

opposition by some local special interests near proposed reserves, and uncertainty 

concerning the size, location, and number of reserves necessary to ensure persistence of the 

reef fish fisheries. The incentive for deliberate poaching may be increased within the 

reserves; thus, at-sea surveillance and enforcement may be necessary. New artificial reefs 

may be needed to replace those lost by inclusion within fishery reserves. Other fishery 

management plans should be coordinated to control trolling and other fishing activities 

within reserves that may impact reef fishes. The short-term impacts on total harvest caused 

by placing fishing habitat into fishing reserves should be compensated for by long-term 

fishery benefits. The Dry Tortugas is listed as a potential marine fishery reserve site. 

284. Plantier, T. L. 1988. "A comparison of reproductive success in early and late breeding sooty terns 

Sterna fuscata in the Dry Tortugas." MS.Thesis, Florida Atlantic University, Boca Raton. 

Evidence indicates that earlier-nesting birds are often older, choose preferred nest sites, 

and have greater reproductive success than those nesting later. The sooty terns at Bush 

Key appear to follow a similar pattern. The first birds arrive at the west end of the 

breeding grounds three weeks earlier than birds at the east end and behaviorally appear to 

be older and more experienced. The west birds settled in the more desirable habitats (the 

west end was cooler than the east end) and laid larger eggs, hatched larger chicks, enjoyed 

greater hatchability, fed their chicks at a lower frequency when they were young, and had 

greater reproductive success than birds in the east. This was accomplished through a 

combination of choosing physically and thermally more favorable habitat, which was more 

centrally located, being more persistent incubators and brooders, and, by nesting earlier, 

having larger, less-easily eaten chicks by the time avian predators arrived on the island. 

285. Plough, H. H. and N. Jones. 1940. Ecteinascidia tortrrger~sis, species Nova; with a review of the 

perophoridae (Ascidiacea) of the Tortugas. Papers Tortugas Lahornto!-y 32: 47-60 (issued 

Oct. 1939). 


During the season of 1936 at the Tortugas Laboratory the senior investigator undertook a 

study of the regeneration of pieces cut from the growing stolons of several species of the 

family Perophoridae (Ascidiacea Phlebobranchia), which grow in profusion at many places 

in the Tortugas area. A new member of the family Perophoridae is described and named 

Ecteinascidia tortugensis from its type locality, the Dry Tortugas Key, Florida. It is 

shorter than other Ecteinascidia, lies on the ventral side attached along the test, has the 

siphons on the dorsal side widely separated and opening in opposite directions, and 

possesses a marked secondary loop in the intestine. This species reaches sexual maturity 

early in July at the Tortugas, about two weeks later than E. conklini. A brief account of the 

development is given. The structure and growth habits of E. tortugensis indicate that it is 

intermediate between E. turbinata and Perophom. They suggest a relationship of the 

Perophoridae with the Ascidiidae. 

286. Porter, J. W. 1977. Pseudorca strandings. Oceans 10, no. 4: 8-16. 

This article provides information and observations surrounding the stranding of thirty false 

killer whales (Pseudorca crassidens) on the Dry Tortugas Islands near Florida in July 

1976. The herd appeared to be protecting an injured male, as evidenced by aspects of 

social behavior and agnostic behavior directed at sharks and the author. Among other 

suggestions, the author postulates that the injured male was unable to feed due to parasitic 

infestation of the ears and consequent impairment of echolocation, which caused the whale 

to beach in order to avoid drowning in its weakened state. Other strandings are discussed 

in light of the information obtained.

287. Porter, J. W., J. F. Battey and G. J. Smith. 1982. Perturbation and change in coral reef 

communities. Proceedings ojrhe Natior~al Academy oJScierzce 79, no. 5: 1678.8 1. 

Ninety-six percent of surveyed shallow-water Dry Tortugas reef corals died during the 

severe winter of 1976-77. Data from skeletal stains indicate that death occurred during the 

mid-January intrusion of 14 degree C water onto the reef. In deeper water, community 

parameters such as percent cover, species number, and relative abundance showed no 

significant change. However, an analysis of competitive interactions at the growing edges 

of adjacent colonies reveals a 70% reduction in space competition during this 

environmental disturbance. These results can explain high variability in the growth rate of 

Floridian reefs and demonstrate the importance of obtaining long-term spatial information 

to interpret successional dynamics of complex communities. 

288. Porter, J. W., 0. W. Meier, L. Chiang and T. Richardson. 1993. Quantification of coral reef change 

(Part 2): the establishment and computer analysis of permanent photostations in the Florida 

SEAKEYS survey (abs). Proceedings ($the Severith I~iternnrionul Coral Reef Syrnposirrm, 

v.1. Mangilao, Guam: University of Guam Marine Laboratory. 

Photostations in five of six locations in the Florida Keys reveal a decline of monitored 

coral reef resources during the 1980's when up to 40% of the coral died in some protected 

areas. Reductions in the number of species and extraordinary shifts in the pattern of 

species abundance occurred in addition to loss of live coral cover. While normally 

associated with catastrophic physical disturbances, this coral mortality occurred during a 

period without major hurricanes in Florida. Relocatable photostations reveal a multiplicity 

of causes for this decline. These include: (I) mortality due to "white band" and "black 

band" disease, (2) direct and delayed mortality from "coral bleaching," caused by 

abnormally elevated sea temperatures, (3) some mechanical damage, and (4) an increase in 

cover by algae. The establishment and sequential analysis of remote sensing data acquired 

from permanent photo-stations will be described in detail, as well as limits to the 

intcrpretability of these photogrammctric data. 

289. Potthoff, T, and W. J. Richards. 1970. Juvenile bluefin tuna, Tlzunnris thy~i~zus (Linnaeus), and other 

sconibrids taken by terns in the Dry Tortugas, Florida. Bulletirr ofMarine Science 20, no. 

2: 389-413. 

The identification and seasonal distribution of juvenile scombrids in the waters near the 

Dry Tortugas, Florida, are described. Specimens were collected (1960 through 1967) from 

regurgitated food of terns. Fishes identified were Thunrzus thyrinris, Thrrr~rrirs atlanticus, 

Euthynnus allefterafirs, Airxis spp, and Krrfsirwonur pelamis; sizes ranged from 24.146 

nim. standard length For the first time, juvenile bluefin tunas are reported in the Dry 

Tortugas region; their presence may indicate that spawning of the species takes place in 

the area. Idcntification methods are discussed, with special emphasis on features of the 

axial skeleton and the number of gillrakers over the ceratobranchial bone of the first gill 

arch. A method is presented for estimating the standard length of damaged specimens on 

the basis of the length of the vertebral column. 

290. Powers, P. B. A. 1933. Ciliates infesting the tortugas echinoids. Carriegie hrstitution of WashLtgton, 

Year Book 32: 278-79. 

Based on studies conducted on the ciliates of theTortugas sea-urchins, Echinoida, it was 

found that when sea-urchins were infested, they made excellent reservoirs for certain 

species of ciliates in which to conduct detailed studies of their internal morphology, 

cytoplasmic inclusions and neuromotor apparatus. 

291. -- . 1936. Studies on the ciliates of sea-urchins: a general survey of the infestations occurring in 

Tortugas Echinoids. Papers Torrugas Lnborafoq 29: 293-326 (issued Dec. 1935). 


With the accumulation of data concerning the ciliate infestations of the alimentary tract of 

echinoids, it became of increasing interest to have a complete record of the ciliates 

infesting sea-urchins about Tortugas. The writer spent the summer of 1933 at the Tortugas 

Laboratory. During this time twelve well-defined species of ciliates were found distributed 

among seven species of sea-urchins. The present paper gives a complete account of the 

general morphology of these ciliates, as well as a description of the various associations 

encountered. Twelve species of ciliates are described which infest the alimentary tract of 

seven species of sea-urchins from the region about the Dry Tortugas. Eight of the twelve 

are new. Only five of these twelve species show any marked host specificity. The 

remaining seven species all show a definite host preference. All of these ciliates are 

associated with their host when the latter are found occurring near the tide line. In seaurchins 

taken below a depth of ten fathoms, only four species of ciliates are found : 

Cryptochilidiurn bennudense, Anophry elongat, Cohrzilernbus coeci and form M. The 

nature of this infestation is one of endocommensalism, there being no present evidence to 

indicate pathogenic tendencies for any of its members. Data concerning the geographical 

distribution of the ciliates infesting sea-urchins from the localities of Beaufort, North 

Carolina, Bermuda and Tortugas have been summarized. It is suggested that the center of 

this infestation will be found in the sea-urchins from the region of the Lesser Antilles and 

that this infestation has been carried northward along with its host, through the agency of 

the Gulf Stream. 

292. Pratt, H. S. 1910. MonocotyleJloridur~u, a new monogenetic trematode. Papers Tortugas 


Carnegie Institution of Washington Publication Nuniber 133. 

The genus Monocotyle was established by Taschenber, in 1878, for a worm which he had 

found on the gills of the eagle-ray (Mj'liobatis aquila) at Naples, and which he named 

Mor?ocotyle ~nyliobatis. The only other known species is Mo~tocotyle ijimue, which was 

discovered in Japan in the mouth of Trygon pastinacea, and described by Goto in 1894. 

The worm herein described makes the third member of the genus and was taken from the 

gills of the whip-ray (Myliobatis fremirrvi!lei) in the Gulf of Mexico and studied at the 

Marine Biological Laboratory at Dry Tortugas, Florida. It differs in certain features from 

the two other species of the genus, but in the general shape and size of the body, the form 

and structure of the suckers, down to the smallest details, and the general arrangement of 

the genital organs it shows a close relationship to them, especially to M. ijirnae. 

293. -. 1910. Parallel transport in tropical trematodes. Science 31: 471-72. 

The digenetic trematodes, as well as other internal parasites, have probably in their phyletic 

history followed somewhat different rules of descent from those of other animals. The fact 

that they live inside of other animals and have a very co~nplex life history must affect their 

phyletic development, in that migrations are very much limited, and their structure is very 

uniform in the parasites themselves. It is probable that where there are apparently related 

species of digenetic trematodes living in widely separated localities, they possess the same 

or similar structural features. This does not necessarily indicate that there is a close genetic 

relationship between them. These facts are well illustrated by the several species of 

digenetic trematodes belonging to the genus Helicornetra, which were found in certain 

fishes of the Tortugas, Florida, and also occur in the Meditteranean Sea. The species of this 

peculiar genus are thus taken as an indication, not that they necessarily bear a close genetic 

relationship to one another, but that similar or identical environmental conditions exist for 

them in these places, so that they have come to possess in the course of time a structure so 

similar that they are included in one and the same genus. 

294. ------ 1916. The trematode genus Stepha~~ochasrnus Looss in the Gulf of Mexico. Parasitoiogy 

8, no. 3: 229-38.

Two species of the genus Ste~~ha~~ochasrili~.~ were found in fishes examined for parasites at 

the Dry Tortugas: S. casus Linton and S, sentus Linton. The anatomy of these worms have 

several interesting and unique features. S. cams Linton is described in this article. 

295. -------. 1913. The trematode parasites of the loggerhead turtle. Science 37: 264-65. 

The studies of trematode parasites of the loggerhead turtle in the Mediterranean and the 

Gulf of Mexico, Dry Tortugas are bricfly discussed and compared. Nineteen species 

occurred in the turtles of the Gulf of Mexico. Eight of these species also occur in the 

Mediterranean Sea. The most numerous trematode occurring in the Dry Tortugas is 

Cymorocarprrs rr,~dulat~r,s. A more detailed discussion is found in the Archives de 

Parasitologie article by Pratt. 

296. -. 1912. Trenlatodcs of the Gulf of Mexico. Verliandlrrnger~ des Vlll Internationale~z 

Zoologe11- Ko~~gresses, 780-781. Jena, G. Fischer. 

This is a discussion of the article written by Pratt in no. 133 Carnegie Institution of 

Washington, listing the trematodes collected at the Dry Tortugas and the hosts they live in. 

297. -. 1916. Trematodes of the loggerhead turtle (Caretta caretta) of the Gulf of Mexico. Archives 

Oe Parasitologie: 4 11-27, 

Five species of trematodes are discussed in this paper. Reference is made to the studies of 

Linton (1910), as well as studies made in the Mediterranean Sea. The five trematode 

species were found in turtles captured on Loggerhead Key in the Dry Tortugas: Wilderia 

elliptica, Pachypsolus tertius, and 1Utyditode.s secundirs, I'elsiochorus cyritbifunnis and 

Cymatocarpus undu1atu.s. 

298. Raim, A. W., W. Cochran and R. D. Applegate. 1989. Activities of a migrant merlin during an 

island stopover. Journal of Raptor Research 23, no. 2: 49-52. 

Activities of a radio-tagged merlin (Falco colunibarius) which was trapped and identified 

as an adult female by George Allex and Daniel D. Berger, were observed from 10-16 April 

1977 on Loggerhead Key. 

299. Reighard. J. 1908. An experimental field-study of warning coloration in coral-reef fishes. Papers 

Tortugas Lnboraror): 2: 257-325. 


This paper embodies a search for the biological significance of the conspicuousness which 

it attempts to show characterizes many of the coral-reef fish of the Tortugas region. After 

showing that this conspicuousness is not a secondary sexual character and that it serves 

neither for protective nor aggressive resemblance, its value as a warning character is 

subjected to experimental test. Experimental evidence is presented to show that the gray 

snapper, the connnonest predaceous fish, discriminates certain colors, forms associations 

with rapidity, and retains these for a considerable time (memory). If any of the coral-reef 

fishes possess a combination of consicuousness with such unpleasant attributes as render 

then unpalatable, the gray snapper should have learned to avoid them at sight and their 

conspicuousness would then have a warning significance. It is shown that when atherina, 

an inconspicuous fish which serves normally as the food of the gray snapper, is given an 

artificial warning color and at the same time rendered unpalatable, it is after a brief 

experience, no longer taken as food by the gray snapper. Artificially colored atherinas thus 

come to have a warning significance for the gray snapper and are avoided, even when not 

unpalatable, although normal atherinas are still readily eaten. The conclusion is thus 

reached that the existence of a warning coloration or of warning conspicuousness in coral 

fishes is easily possible. The conclusion is reached that the conspicuousness of coral-reef 

fishes, since it is not a secondary sexual character and has no necessary meaning for 

protection, aggression, or as warning, is without biological significance. The coral-reef

fishes have no need of aggressive i~iconspicuousness because their food consists of 

invertebrates, chiefly fixed. They have no need of protective inconspicuousness because 

the reefs and their agility afford them abundant protection. Selection has therefore not 

acted on their colors or other conspicuous characters, but these have developed in the 

absence of selection and through internal forces. They are the result of race tendency 

unchecked by selection . 

300. ------- 1907. The photography of aquatic animals in their natural environment. Uui/etiri of rlie 

United States Bureau of Fisheries 27: 41-68. 

This paper describes the photography of aquatic organisms in their native environment and 

under normal conditions by carrying the camera into the field. Photos and diagrams are 

provided for cameras and apparatuses that remain above the surface of the water and 

cameras that are submerged. 

301. Reynolds, J. E. I11 and I. C. Ferguson. 1984. In~plications of the presence of manatees, Triclzeclzus 

manatus near the Dry Tortugas Islands, Florida USA. Florida Scie~rtist 47, no. 3: 187-89. 

Two West Indian manatees (Trichechus manatus) were observcd 61 km northeast of the 

Dry Tortugas Islands, a location not normally considered to be part of the species range. 

When spotted, the animals were swimming in a soutwesterly direction, away from Florida. 

Observations such as this, of manatees far from freshwater, raise the question of whether 

manatees require regular access to fresliwater for osmoregulation, as suggested in the 

literature. 

302. Reynolds, J. E. I11 and J. C. Steinmetz. 1983. Dry Tortugas: products of time. Sea frontier.^ 29, no. 

2: 66-75. 

This article discusses the general formation of the islands and their history. 

303. Richards, 0 . W. 1934.1936. Growth studies in the aseidian, Phallrasia nigra, and hermit crab, 

Caenohita clypeatus. Carnegie Institution of Washington, Year Book 

Note: published as follows 1934, v. 33, p. 261; 1936, v. 35, p. 92. 

The early growth and development of the ascidian Phallusia nigro was recorded over time 

using motion and still photography. Claw size ratios to body size of the hermit crab. 

Coenohita clylmms, were examined by correlation metl~ods. 

304. Ricklefs, R. E. and S. C. White. 1981. Growth and energetics of chicks of sooty tern, Sterna fuscafa 

and common tern, Sterna Izirundo. Auk 98, no. 2: 363-78. 

The energy budgets of chicks of tlie common tern (Ster~m himtido) were measured on 

Great Gull Island, New York. Also measured were the sooty tern (S, fuscafa) on the Dry 

Tortugas. Florida The respiratory energy requirement was determined by measuring 

oxygen consumption in a closed system. The growth energy requirement was calculated 

from the lipid and protein contents of a series of chicks spanning the range between 

hatching and fledging. Energy budgets calculated for the two species differed in several 

ways. (I) Maintenance metabolism was lower in the sooty tern owing to its warm 

environment. (2) Sooty terns allocated more of their energy intake to lipid accumulation 

from an earlier age. (3) In the sooty tern, the allocation of energy to growth initially was 

high, but its absolute amount decreased steadily throughout the growth period. In the 

common tern, both growth and maintenance energy allocations increased rapidly during tlie 

first half of the development period. (4) In sooty tern chicks energy ~netabolis~n 

approached its maximum rate (135 kJlday) by the end of the first third of the development 

period, after which it leveled off. In the common tern, energy metabolism increased from 

about one-quarter of its maximum during the first five days after hatching to its maximum 

of 200kJlday during the third week of the postnatal development period. Although these 

observations support tlie hypothesis that slow growth in pelagic seabirds is selected to

educe the energy requirement of the chick, the energy budgets also suggest that a doubling 

of the growth rate by the sooty tern would increase the maximum energy requirement of the 

chick by only 20% and the total feeding requirement of the adult by only 5%. Moreover, 

the levels of water in muscles suggest that the sooty tern develops mature function earlier 

than does the common tern, which in itself might be sufficient to account for the slower 

growth of the first species. 

305. Ricklefs, R. E. and S. C. White-Schuler. 1978. Growth rate of the brown noddy on the Dry Tortugas 

Florida USA. Bird-Banding 49, no. 4: 301-12. 

Growth rates within seabird species can vary with locality, season, and year. In this study 

noddy tern checks captured on Bush Key, Dry Tortugas, June, 1972, were weighed and 

measured. Growth increments were used to calculate a composite wing length growth 

curve to estimate the ages of chicks. A logistic curve was fitted to describe the relationship 

between weight and age. Growth constants of the fitted curve (growth rate K = 0.153, 

asymptote A=160 grams, and age at inflection ti=14.0 days) were similar to values reported 

for the brown noddy on Kure Island and Manana Island, Hawaii. Also reported are outer 

primary and rectix lengths and body temperatures of nestlings and adults. 

306. Riley, G. A. 1938. Study of the plankton in tropical waters. Carnegie institution of Washington 

Year Book 37: 98. 

The small quantity of plankton in tropical waters as contrasted to higher latitudes is 

investigated, and when compared to a similar survey underway in Long Island, N.Y., the 

indication is that chlorophyll and plant pigments are one-twenty-fifth the amount found in 

New York. 

307. Riska, D. E. 1986. An analysis of vocal communication in the adult brown noddy, Anous stolidus. 

Auk 103, no. 2: 359-69. 

The author analyzed vocal signals of marked adult Brown Noddies (Anous stolidus) 

throughout their nesting season in the Dry Tortugas, Florida from 1979 to 1982. The basic 

unit of the adult repertoire is a wide-band click, less than 4 msec duration, ranging in 

frequency from 200 to 3,300 Hx. He identified nine temporal arrangements of these clicks, 

which form the notes of the calls. These calls differ little in frequency range, but they 

differ in the mean frequency of the most intense sound energy hand, in note duration, in the 

number of clicks per note, and in internote interval. These calls are used in different 

contexts, which sometimes overlap. Frequency, note duration, and length varied among 

individuals for some calls. No tonal elements characteristic of calls of brown noddy 

nestlings remain in the adult repertoire. 

308. - . 1986. "Communication behavior of the brown noddy (Anorrs siolidrrs) and sooty tern 

(Sterna fuscata), Dry Tortugas, Florida (vocalizations, laridae, signals, colonial, 

breeding)." Ph.D. Dissertations, University of California at Los Angeles. 

The basic unit of the adult repertoire is describcd as a wide-band click, less than 4 msec 

duration, in the frequency range 200 to 3300 Hz. Nine calls differ in temporal 

arrangements of clicks, mean frequency of the most intense sound energy band, note 

duration, number of clicks per note, and inter-note interval. Frequency, not duration, and 

inter-note interval do not differ between sexes. The nestlings of the brown noddy produce 

three structurally different vocalizations within one day after hatching. Postures of chicks 

and contexts in which these signals are used differ. The repertoire is composed of 

frequency-modulated tonal elements and broad-band bursts of sound with little 

resemblance to the adult repertoire. Juvenile bush-nesting noddies begin flying when 40- 

48 days old, after which they are still fed at their nests. Adult noddies accept a substituted 

nestling differing from their own in size, color and plumage stage, up to at least 20 days 

post-hatching. The adult Sooty Terns produce eight structurally different vocalizations,

and nestlings produce three, in the frequency range 300-7000 Hz. Postures differ for each 

call, but contexts in which these are used overlap. The range of frequencies in which 

young birds call extends higher than that of adults, but the frequency-modulated tonal 

elements characteristic of nestling vocalizations remain complex in the adults. 

309. Rivas, L. R. 1951. Preliminary review of the western North Atlantic fishes of the family 

Scombridae. Bulletirt of Marine Scierzce of the Gulfand Caribbean 1, no. 3: 213-30. 

This paper brings up to date the taxonomy of the western North Atlantic mackerels and 

tunas. In addition to a key to the genera and species, a complete synonymy, a diagnosis 

and pertinent comments are given under each species. 

310. Roberts, H. H., L. J. Rouse Jr., N. D. Walker and J. H. Hudson. 1982. Cold water stress in Florida 

Bay and northern Bahamas, a product of winter cold air outbreaks. Journal of Sedirner~tary 

Petrology 52, no. I: 145-55. 

During January 1977 three consecutive cold fronts crossed south Florida and the northern 

Bahamas which depressed shallow-water temperatures below the lethal limit for most reef 

corals. Digital thermal infrared data acquired by the NOAA-5 meteorological satellite, in 

situ water temperatures, and meteorological data were used to study the thermal evolution 

of Florida Bay and Bahatna Bank waters. The third and most important frontal system 

depressed Florida Bay water below 16 degrees C, a thermal stress threshold for most reef 

corals, for 8 days. Coral mortality at Dry Tortugas was up to 91 percent during the 1977 

event. Coral and fish kills were also reported from other parts of the Florida Reef Tract 

and northern Bahamas. Study results show that cold-water stress conditions can exist over 

vast shallow-water areas and have residence times of several days. 

31 I. Roberts, H., H. Lawrence, J. Rouse Jr. and N. D. Walker. 1983. Evolution of cold water stress 

conditions in high-latitude reef systems: Florida Reef Tract and the Bahama Banks. 

Caribbean Journal of Science 19, no. (1-2): 55-60. 

Thermal depression of shallow bank and bay waters accompanying the passage of severe 

cold fronts can stress high latitude coral reef systems, such as those of the Florida Reef 

Tract and nortliern Bahama Banks. Laboratory and field experiments suggest that 

sustained temperatures below 16 degrees C are detrimental to most reef-building corals. 

Time-series satellite imagery provides a data base for assessing the thermal variability of 

waters interfacing with reef systems. Digital thermal infrared data acquired by the NOAA- 

5 nieteorological satellite were used to study thermal evolution of Florida Bay and Bahama 

Bank waters during a succession of three cold-air outbreaks (January 1977). These studies 

indicate that the temperature of subtropical bank and bay waters is subject to depression 

below 16 degrees C accompanying the outbreak of unusually cold air. This superchilled 

water can have a residence time of days. The cooling process creates water masses that are 

out of density equilibrium with warmer ocean water. Offshelf movement of the cold, dense 

water occurs at particular sites, as shown by time-series satellite data. The absence of coral 

reefs opposite tidal passes in the Florida Keys is attributed to this process, which has 

probably limited development of the entire reef tract. 

312. Robertson, W. B. Jr. 1978. Species of special concern sooty tern. Birds 2, no. Edited by H. Kale: 

89-90. 

A description, range, and habitat of the sooty tern at-e given along with its life history and 

ecology at the Dry Tortugas. Its classification is based not on its abundance, but it is 

because the Dry Tortugas colony is a major Florida wildlife resource. Aside from the 

Tortugas no other location in Florida is suitable for sooty tern nesting. This colony affords 

a means of monitoring the general health of offshore Gulf waters of southern Florida.

313. -. 1964. The terns of the Dry Tortugas. Bulletin of the State Museum, Biological Science 8, 

no. 1: 1-95. 

New information from unpublished sources and from published records hitherto 

overlooked permit a re-evaluation of the history of the Dry Tortugas and of the terns that 

inhabit them. The geography and ecology of the 1 I keys that have variously comprised the 

group since it was first mapped in the 1770's are described and their major changes traced. 

The recorded occurrences of the seven species of terns reported nesting on the Keys are 

analyzed in detail. The sooty tern colony has fluctuated from a low of about 5,000 adults 

in 1903 to a reported peak of 190,000 in 1950; for the past four years it has remained 

steady at about 100,000. The brown noddy population, which reached a peak of 35,000 in 

1919, was reduced by rats to about 400 adults in 1938; it is in the neighborhood of 2,000 

today. A colony of 150 to 450 roseate terns has nested in most years from 1917 to the 

present. About 500 least terns nested regularly from 1918 to 1932, then unaccountably 

dwindled to a few pairs by 1937 and shortly afterward disappeared. Royal and sandwich 

terns nested abundantly in the mid-19th century, and a colony of royals may have existed as 

late as 1890. Both species are believed to have been extirpated from the Tortugas by 

egging. No verifiable evidence exists for the nesting of the common tern, which has been 

reported several times. The black noddy, first reported for the continental United States at 

Dry Tortugas in 1960, has been found there each summer since. 

314. -. 1969. Transatlantic migration ofjuvenile sooty terns. Nature 222: 632-34. 

From 1959 to 1968,70,000 adult and 130,000 juvenile sooty terns (Sterna fuscata) were 

banded at Bush Key, Dry Tortugas, Florida. By December 1968. 29 juveniles were 

recovered in West Africa. It appears that the primary biological function of the 

transatlantic migration is to avoid intraspecific competition and this adaptive value 

becomes evident when the migration of juveniles is seen in the context of the rigidly 

structured sooty tern population. It may be evidence for a successful evolutionary 

mechanism. 

315. Robertson, W. B. Jr. and B. Given. 1980. Ruddy quail dove Ceotrygon montanaagain at Dry 

Tortugas Florida USA. Florida Field Naturalist 8, no. I: 23-24. 

About noon on December 15, 1977, a cold day with sevcre northwesterly squalls, Given 

found and photographed a large, reddish dove on the second tier of Fort Jefferson, Dry 

Tortugas, Florida. This record is the fifth report of the species from Florida and the second 

from Dry Tortugas. 

316. Robertson, W. B. Jr. and C. R. Mason. 1965. Additional bird records from the Dry Tortugas. 

Florida Naturalist 38: 131-38. 

Sprunt (1962-63) summarized what was known about the occurrence of birds at the Dry 

Tortugas through the Summer of 1962. In this paper the authors report on recent bird 

records up to April 1965. Comments relate to 12 species new to the list or those known 

from either one or two records. Sprunt listed 227 species of birds for the Tortugas, the 

authors add 12 to bring the total to 239 species. 

317. Robertson, W. B. Jr. and L. C. Below. 1975. A red-headed woodpecker at Dry Tortugas. Florida 

FieldNaruralisr 2, no. I: 20. 

On May 5, 1973 Mr. and Mrs. G.H. Perbix of Cincinnati and Mrs. Below, members of the 

tern-banding party then at Dry Tortugas, visited Loggerhead Key and at once noticed an 

adult red-headed woodpecker (Melanerpes erythrocephalus) in the large Australian Pines 

(Casuarbia equi.setifolia) near the dock. We find only one other report of the red-headed 

Woodpecker at Dry Tortugas. Howell (1932:308) wrote that the species was unknown in 

the Florida Keys "...except for a single occurrence on the Tortugas - a bird seen there on a 

number of days early in June." The red-headed woodpecker is not known to occur outside

the United States but the present record inevitably raises the question: was the bird 

migrating across the Gulf or was it merely a vagrant? 

318. Robertson, W. B. Jr. D. R. Paulson and C. R. Mason. 1961. A tern new to the United States. Auk 

78: 423-25. 

This note provides a description of the black noddy, Anous tenuirostris collected at Dry 

Tortugas. This is the first of this species collected in the United States. Two specimens 

were taken from Bush Key during July 1960. The bird occurs nearly world-wide in the 

warmer seas, but is absent from most of the Atlantic Ocean north of the equator and most 

of the Caribbean Sea. 

319. Robinson, A. H. 1976. Marine, island and coastal parks in the United States National Park system: 

A review and progress report in 1975. lnternntional Conference on Marirze Parks and 

Reserves., pp. 226-27. Gland, Switzerland: IUCN. 

This paper provides a basic introduction to critical marine habitats and the planning and 

management of marine parks and reserves, including interpretation and environmental 

education in marine parks. Progress in the creation of marine parks and reserves is 

reviewed, and a special report on marine park systems in the Pacific region is included. 

Fort Jefferson National Monument is discussed as the first underwater preserve established 

in the United States. 

320. Schaeffer, A A. 1925. Experiments on the influence of temperature and dilute and concentrated 

sea-water on ameboid movement. Bulletin of tlze Ecological Society of America: 11. 

The reactions of various species of amoebas to different concentrations of sea-water have 

been used during the past several years at Tortugas as important aids in the identification 

and fixation of species. The rate of movement of several species was studied in various 

concentrations of sea-water indicating that the optimal concentration of sea-water is below 

the norm in every case when measured by the rate of cell-coordinated movement. 

321. -. 1926. Taxonomy of the amebas: with descriptions of thirty-nine new marine and 

freshwater species. paper.^ Tortugas Lnhoratory 24: 1-116. 


The purpose of this report is to set forth a description of 39 new species and 1 I new genera 

of amebas (Amoebaea), and to propose a preliminary system of classification of the 

amebas, based on their general morphology. Gencral observations on structure, 

physiology, distribution, and methods of investigation are provided. The changes of form 

which amebas undergo is a fundamental morphological characteristic of amebas, and forms 

the basis of a natural classification. For the purpose of quickly recognizing a species other 

characteristics are more valuable, such as the nucleus, vacuoles, crystals, resistance to 

dilutions and concentrations of sea-water, etc. A brief discussion of these characteristics 

with reference to specific descriptions is given along with colored drawings and 

photographic text-figures. 

322. Schtnitt, W. L. 1924-1932. Systematic-ecologic studies of the decapod crustacea. Carnegie 

Instiiurion of Washingion, Yearbook. 

Note: published as follows: 1924, v.23, p. 200-201; 1925, v.24, p. 230-231; 1930, v.29, p. 

143; 1931, v.30, p. 389; 1932, v.37, p. 279. 

Very striking color characteristicsivariations affecting chela, and often the appendages, are 

noted among snapping shrimp, S)walpkeus, and giant isopod crustaceans. Bathy~netric 

distribution of decapods are investigated. 

323. Schnell, G. D. 1974. Flight speeds and wing beat frequencies of the magnificent frigate bird. Auk 

91, no. 3: 564-70.

Wingbeat frequencies and flight speeds of magnificent frigatebirds were recorded with a 

Doppler radar in the Dry Tortugas, Florida. The flapping rate averaged 2.84 beats per 

second (SD 0.14) and was not significantly correlated with flight speed, providing further 

evidence that the birds' wingbeat frequency is essentially constant within species. The 

flapping rate is somewhat higher than predicted from the theory of mechanical oscillators 

when the distance from the end of the wing to the first articulated joint is used as an 

estimate for the average effective wing length. Flight speeds of birds in a flat calm 

averaged 22.55 mph. The highest average ground speed of 30.17 mph was obtained from 

frigatebirds flying in a 6 to 8 mph wind, and the lowest of 16.00 rnph for birds flying into 

the 65 tnph wind. Airspeeds were greater for frigatebirds flying into the wind than for 

those moving across or with the wind. 

324. Schreiber, R. W., W. B. Robertson Jr. and T. Bellow. 1976. Nesting of brown pelicans, Pelecarlus 

occiderltalir, on the Dry Tortugas, Florida. Florida Field Naturalist 3, no. 2: 47-48. 

On June 14, 1974 Bush Key, Dry Tortugas Ted Bellow and C. Winegarner found 5 brown 

pelican nests about 12 feet above ground in the white mangroves (Laguncularia racemosa) 

along the north shore. Nineteenth-century records of pelicans breeding on the Dry 

Tortugas are ambiguous. ... on the Tortugas (I 860) it thus appears that a few pairs did breed 

on the Tortugas in the mid-1900's, but by late in the century none did so. Our record is the 

first reported nesting of this species in the 20th century on the ornithologically well-known 

islands (Robertson and Mason, 1965). Three of the nests found in 1974 contained two 

eggs each, one nest was empty, and the fifth was not checked. 

325. Schroeder, P. B. and J. H. Davis. 1971. Ecology vegetation and topography of the Dry Tortugas 

updated lo 1970. Quarter1.y Jourriol of the Florida Academy of Scierlce (Suppl I): 12-13, 

The half-dozen islets of the Dry Tortugas have been ecologically studied periodically since 

the turn of the century. In November and February a year ago, a field party from the 

University of Miami made a topograpl~ical and vegeational study of several of these keys. 

The pertinent information gathered at that timc is now available and provides continuity 

with the studies of Millapaugh (1907). Bowman (19 18) and Davis (1942). The keys 

studied have changed fro~n barren coral and sand to substantial islets largely covered with 

vegetation. The configuration of one of these has been completely altered. All the keys 

have been changed considerably in shape. Vegetational communities have shown similar 

changes and maturity. Mangrove areas (red and white) have become established and 

enlarged. Australian pines and other exotics, introduced to Loggerhead Key, have spread 

over tnuch of the island and now are found on Bush Key. 

326. Scott, W. E. D. 1890. On birds observed at the Dry Tortugas, Florida, during parts of March and 

April, 1890. The Auk: A Quarterly Journal of Ontithology 7, no. 4: 301-14. 

The list of birds observed at the Tortugas includes eighty species, fifty seven of which were 

land birds. The author states that no land birds breed on any of the keys group, and that the 

stay of any land bird is of very short duration. 

327. Seaman, W. Jr. and D. Y. Aska. 1974. Research arid iujorrnation needs of the Florida spiny lobster 

fishery. State University System of Florida Sea Grant Program, Miami FL. 64 pgs. 

In response to a number of fishermen in South Florida, the State University System of 

Florida Sea Grant Program became involved in research on the spiny lobster, Panulirus 

argtrs. When additional research needs were expressed, Florida Sea Grant decided to 

become better informed on the subject, and evaluate its potential for service to the persons 

dependent on this fishery resource. A meeting of persons and organizations involved in the 

biology and/or utilization of the spiny lobster fishery in Florida was called to identify 

broadly the problems and information needs of persons dependent on the spiny lobster

esource, to assess existing sources of information and their possible applications, and to 

identify priorities and actions needed to resolve user problems. 

328. Shinn, E. A., J. H. Hudson, R. B. Halley and B. Lidz. 1977. Topographic control and accumulation 

rate of some Holocene coral reefs: South Florida and Dry Tortugas. Proceedi~~gs, Third 

lrlternatior~al Coral ReefSyn~posium, RSMAS, Univ. of Miami, Coral Gables FL. p. 1-7. 

Core drilling and examination of underwater excavations on 6 reef sites in south Florida 

and Dry Tortugas revealed that underlying topography is the major factor controlling reef 

morphology. Carbon-14 dating of coral recovered from cores enables calculation of 

accumulation rates. Accumulation rates were found to range from 0.38 1/1000 years in 

thin Holocene reefs to as much as 4.85 rnllOOO years in thicker buildups. Cementation and 

alteration of corals were found to be more pronounced in areas of low buildup rates than in 

areas of rapid accumulation rates. Acroy~ora palrnata, generally considered the major reef 

builder in Florida, was found to be absent in most reefs drilled. At Dry Tortugas, the more 

than 13-meter thick Holocene reef did not contain A. palmata. The principal reef builders 

in this outer reef are the same as those which built the Pleistocene Key Largo formation, 

long considered to be a fossilized path reef complex. 

329. Shinn, E. A. 1984? Geologic history, sediment, and geo~norphic variations within the Florida Reef 

Tract. Advances in reef sciences, abstracts and schedule ofpre.sentations: a joint meeting 

of the Atlar~tic Reef Committee and the International Society for ReefStudies , 113-14. 

Miami, Florida: University of Miami. 

A combination of core drilling, high resolution seismic profiling, and constituent particle 

analysis reveal these major aspects of Holocene reef development and sediment 

distribution within the Florida reef tract: (1 j reef distribution and shape are controlled by 

underlying Pleistocene limestone topography; (2) accun~ulation of sand and rubble occurs 

in forereef and backreef areas; and (3) conlposition of sediment and coral distribution are 

controlled by the reef tract trend relative to prevailing wind and exposure to Gulf of 

Mexico water. 

Ihinn, E. A,, B. H. Lidz, R. B. Halley, J. H. Hudson and J. L. Kindinger. 1989. Reefs of Florida 

and the Dry Tortugas: Miami to Key West, Florida, July 2-7, 1989 .28th International 

Geological Congress Field Trip Guidebook (American Geophysical Union), no. T176. 

Washington, D. C.: American Geophysical Union. 

This field guide concentrates on explaining the distribution of Holocene coral reefs, the 

relationship between topography and Holocene sea-level rise, and the compositional and 

thickness variation of sediments produced in and adjacent to the reefs. A discussion and 

speculation of the future of the reefs under a stable sea, and a lowered sea-level is included 

Also attached is a key to the Stony Corals of the Florida Keys and Dry Tortugas, a species 

list, illustrations of geologic cross-sections, aerial and underwater photographs of reefs and 

coral. 

331. Shoemaker, C. R. 1934. Two new genera and six new species of Amphipoda from Tortugas. Papers 

Tortugas labor at or)^ 28: 245-56 (issued Nov. 1933). 

Carnegie Institution of Wasliington Publication Number 435. 

New genera and species of Amphipoda are described from specimens collected at the 

Tortugas, including Socarnes cor~cnvus, Giranopsis torrrlgae, Hererophilar seclusus, 

Pontogeneia lor~gleyi, Ampithoe divur,sia, Leucorhoidespottsi (new species); and 

Heterophlios, Leucothoides (new genera). 

332. Silberman, J. D., S. K. Sarver and P. J. Walsh. 1994. Mitochondria1 DNA variation and population 

structure in the spiny lobster Panulirus argus. Marine lliology 120: 601-8. 

Adult spiny lobsters (Panulir-us argrrs) were collected from nine locations including the

Tortugas, throughout the tropical and subtropical northwest Atlantic Ocean and examined 

for mitochondria1 DNA (mtDNA) variation. 187 different mtDNA haplotypes were 

observed among the 259 lobsters sampled. Haplotype diversity was calculated to be 0.986 

and mean nucleotide sequence-diversity was estimated to he I .44%; both of these values 

are among the highest reported values for a marine species. Analysis of molecular variance 

(AMOVA) and phenetic clustering both failed to reveal any evidence of genetic structure 

within and among populations of P.argus The present data are consistent with high levels 

of gene flow among populations of P.argus resulting from an extended planktonic larval 

stage and strong prevailing ocean currents. 

333. Smayda, T. J., Y. Shimizu, C. R. Tomas and D. G. Baden. 1993. The influence of phosphorus 

source on the growth and cellular toxin content of the benthic dinoflagellate Prorocet~trum 

lirna. Toxic Phytoplar~kfon Blooms 111 The Sen., 565-70. 

The relationship between toxin content and nutritional status of the toxic marine 

phytoplankton species Proroco~trum lima was examined in a clonal culture isolated from 

the Dry Tortugas, Florida, grown with inorganic phosphate and glycerol phosphate 

enriched media. Growth, alkaline phosphatase activity and okadaic acid content were 

measured. Phosphate enriched cultures exhibited rapid growth rates(0.75 divld), moderate 

terminal densities of 134,779 cellslml and low alkaline phosphatase activity (200,000 cellslml, and had alkaline 

phosphatase activity an order of magnitude greater than those grown in inorganic 

phosphate. When comparing toxin levels at 20 and 30 days, cells grown on the organic 

phosphate enrichments had consistently higher per cell values (I 1.2 and 14.2 pglcell, 

respectively) than those with inorganic phosphate (7.5 and 8.9 pglcell), respectively). 

Phosphorus source effected growth, maximal densities, and okadaic acid content of P. 

lima. 

334. Smith, H. G. 1937. Contribution to the anatomy and physiology of Cassiopenfioi~dosa. Papers 

Torrrrgas Lnhorator-y 31: 17-52 (issued July 1936). 


This research was undertaken to extend our previous scanty knowledge on the physiology 

of feeding and digestion in the Scyphozoa. Cassiopea was selected as the experimental 

material for two reasons, it is a member of the Rhizostomeae in which the mode of feeding 

is particularly interesting owing to the sub-division of the mouth, while certain species, 

including the one studied, possess zooxanthellae. Other aspects of the structure and 

physiology of species of this genus have been extensively studied, notably at the Tortugas 

Laboratory, by Mayer and others. It was originally intended to work on C. xamachana, 

which was very abundant at one time in the moat at Fort Jefferson. Recent changes in the 

conditions in the moat, the result of silting up, have caused the complete disappearance of 

the species although from this locality. In the absence of this species, C. frondosa was 

investigated and this although less hardy than C. xumachar~a, proved satisfactory material. 

Experiments have been made on the effect of starvation in light and in darkness on the 

medusae. In light, specimens were kept alive for I5 days, and in darkness for 7 days. 

Numerous algae were ejected by way of the gastric filaments and plaited membranes at the 

base of the filaments and the medusae became brown in color. They also shrank 

considerably in size. The effect of the zooxanthellae on phosphorus excretion has been 

studied, the amount of phosphorus in the sea-water surrounding one specimen being 

reduced to zero within 24 hours. Finally, feeding, digestion and symbiosis in C. fror~dosa 

have been discussed. It has been suggested that the variation in pH in the coelenteron 

affects the activity of the jellyfish. The association with zooxanthellae is probably similar 

in nature to that which occurs in the Madreporaria.

84 

335. South Florida Area : Synthesis of avail. lble biological, geological, chemical, socioeconomic and 

cultural resource information. 1990. OCS Study, MMS 90-0019. U.S. Department of the 

Interior, Minerals Management Service, Atlantic OCS Regional Office. 

This study summarizes the available biological, geological, chemical, and socioeconomic 

information in south Florida in relation to the potential effects of offshore gas and oil 

exploratio~i and development. The synthesis will help Federal and state policy makers 

make informed decisions about future lease offerings and environmental restrictions on 

offshore oil and gas operations. The Dry Tortugas is included as part of the South Florida 

Reef Tract. In summary it would be very difficult to protect the mangroves, reefs, seagrass 

beds and their associated assemblages from large oil slicks. Severe weather would make it 

impossible. The Dry Tortugas experienced an oil spill from the beaching of Brother 

George in 1964. Birds were killed . Some coral may have been killed around the 

Tortugas from the 3,100 barrel spill, but it did not affect other areas further to the east of 

the site. If a large oil spill did occur here it would take 100+ years for the oldest coral 

heads to regrow and achieve the same level of pre-spill structural complexity. The effects 

of an oil spill on other flora and fauna of the Florida Reef Tract can only be guessed. 

336. Spence, J., and 0. W. Richards. 1940. Native cellulose in the ascidian Phallusia nigra. Papers 

Tortugas Laboratory 32: 163-67 (issued Sept. 1940). 


Many organic compounds of high molecular weight are readily identifiable from the 

characteristics of the X-ray diffraction diagrams. Cellulose and its derivatives have been 

extensively examined by X-ray diffraction methods in the search for a complete solution of 

the structure and crystallite arrangement of the cellulose molecule. Fron~ the analytical 

standpoint, X-ray diffraction diagrams not only confirm the initial chemical identification 

of cellulose by Schmidt (1845), but they also show the presence of crystallites and their 

orientation. The Phallusia rligra was collected in the moat of Fort Jefferson and the tunic 

was removed on return to the Tortugas Laboratory. The result, namely, the recognition of 

native cellulose and the preferred orientation of the crystallites in l-'hallusia nigra, is 

naturally anticipated from previous observations on other ascidian tests. This method 

provides a useful analytical "tool" for use in zoological investigation. 

337. Sprunt, A. Jr. 1963. Birds of the Dry Tortugas. Florida Naturalist: 22-26, 52-53. 

This is a continuation of the listing from the 1962 series on listings of birds of the Dry 

Tortugas. 

338. 1962. Birds of the Dry Tortugas 1857-1961. Florida Naturalist 35: 35-40, 82-85, 129-32. 

A brief discussion of the history of bird studies of the Dry Tortugas is given. Special 

attention is paid to the migratory birds passing through the Dry Tortugas in hope of 

sliedding light on trans-Gulf migration. 

339. -- , 1947. Blizzard of birds: the Tortugas terns. National Geographic Magazine February: 

213-30. 

This article gives a history of the Tortugas terns up to 1947. Boobies and noddies are 

included also. 

340. -. 1950. Bridled tern, Sterna a. Melanoptera, taken at Dry Tortugas. Auk 67, no. 4: 514 

This article provides an account of the first Stenla rnelanoptera recorded at the Dry 

Tortugas, and the fifth specimen recorded in Florida. 

341. -- . 1950. A list of birds of the Dry Tortugas Keys, 1857-1949. Florida Naturalist 23: 49-60, 

73-78, 105-1 1.

A listing of the birds of the Dry Tortugas is given. Land birds-pigeons through vireosincluding 

warblers through sparrows and water birds. 

342. -- . 1951. Some observations on the fall migration at Dry Tortugas. Auk 68: 218-26. 

The author arrived at tlie Tortugas following a hurricane August 26-27, which seemed to 

have no effect on the Tortugas, or the birds there. He found the birds to be in good 

physical condition, with no signs of exhaustion. Birds were tame and could be approached. 

A listing of the birds sighted is given. 

343. --- . 1948. The tern colonies of the Dry Tortugas keys. Auk 65: 1-19. 

Tlie first post-war (1945-46) status report on tern populations inhabiting the Keys of the 

Dry Tortugas is presented in this paper. A brief history on population counts dating back 

to 1832 by Audubon is given, as well as a description, mostly vegetative, on Keys utilized 

by terns for nesting activities. Tern springtime arrival and summer departure are discussed, 

along with numbers of eggs produced, nesting locations and tern behavior. Based on the 

square-yard unit system, it was determined that the population count for Sooties was 

97,200, while a count of 550 was found for the noddies by numbers of nests. The tern 

populations have suffered virtually no damage during the occupation of the islands by 

naval forces. Aside from weather, predation by natural enemies includes sand-crabs and 

man-0'-war-birds. The tern colonies appeared safe, but certain topographical changes, such 

as the recent increase in vegetation tilay be problematical. 

344. Stevenson, J. 0. 1938. The tern colonies of Dry Tortugas. Bird-Lore 40, no. 5: 305-9. 

This article describes briefly the history of the tern colonies of the Dry Tortugas. The 

author visited the Tortugas on May 24, 1937, two years after a hurricane swept tlirough tlie 

islands destroying Bird Key, the historic breeding grounds for thousands of sooty and 

noddy terns. 

345. Steward, F. C. 1940. The growth of Valonia vertrricosu J. Agardh and Vulonia ocellnta Howc in 

culture, with a note on the sap cotnposition of Valonia ocellata Howe at Tortugas. Papers 

Tortugas Laboratory 32: 85-98 (issued Oct. 1939). 


So much physiological work has been done using species of Valonia that their mode of 

development has special interest. Living material of V ventricosa and V. ocellata was 

collected at the Dry Tortugas, Florida. These species were chosen because of the 

difference in their morphology. Valor~ia venrricosa J. Agardh and V. ocellata Howe have 

been kept alive for over two years from their original collection. Vesicles of considerable 

size (V. verltricosa) and with all the characteristics of the plant in nature have been grown 

attached to a suitable substratum. The development of the vesicle and rhizoids from 

aplanospores is illustrated by a series of photographs. V verzrricosa also produces 

filaments which penetrate the substratum and from which close clusters of vesicles arise as 

they do in the normal habitat. The appearance of the aplanospore and growing vesicle 

between crossed Nicols is described and its bearing on the structure of the wall indicated. 

Valonia ocellata produces pear-shaped vesicles, cylindrical rhizoidal processes (which it is 

shown may become long and branched), and apparently proliferated masses composed of 

sriiall cellular segments. The growth and development of all these structures from 

aplanospores, or the product of "segregative division" have been observed and are 

recorded by photographs. 

346. Steward, F. C. and J. C. Martin. 1937. The distribution and physiology of Valortin at the Dry 

Tortugas, with special reference to the problem of salt accumulation in plants. Papers 

Tortugas Luborutor)~ 31: 87-1 10 (issued Oct. 1936). 


This paper presents the results of a survey, made during the summers of 1933 and 1934 of 

the physiological behavior of the two species of Valorlia which are most abundant at 

Tortugas, Florida. One may well ask what justification there can be for yet another paper 

on Valonia. Whatever the legitimate claims which may be made for such attention, they 

are somewhat counterbalanced by the inaccessibility of Vulonia, which has prevented that 

examination by a variety of investigators which is the best safeguard against overemphasis. 

Valor~ia rnacrophjsa occurs at Tortugas only in the moat of old Fort Jefferson. This 

organism demands complete protection from the effects of swell and surf. The growth 

obtained on a horizontal ledge is luxuriant; that on an inclined or vertical surface sparse 

and irregular. In the protected locations it demands, V. lllacrophj~su is exposed to and 

withstands, a wide range of light conditions and diurnal fluctuations in the composition of 

the external medium. Vulor~iu ~,entricosu is abundantly obtained on Bird Key Reef. The 

distribution of V. veritricosa is complementary to that of V. rnacrophysa, and the solution 

of the problem whether the species are distinct, raised thereby , must await adequate 

transplant experiments. The range of sap composition which V. verttrico,sa and V. 

macrophysa exhibit at Tortugas in sea water is described. Differences occur in the 

composition of the sap of V. macr-ophysn grown in different parts of the moat of Fort 

Jefferson. The principal causal factor appears to be the light condition which it obtains 

during growth. In general the conditions which produce the most abundant growth of V. 

macrophysa likewise produce the greatest concentration of potassium and lowest 

concentration of sodium. 

347. Stockard, C. R. 1908. Habits, reactions, and mating instincts of the "Walking Stick," Aplopus 

nmayeri. Papers Tortugus Laboratory 2: 43-59. 


This investigation of a protectively adapted insect is important to show definitely whether 

the actions of such an animal are coordinated with its protective structure. It is concluded 

that the habits of Aplopus maperi on its food-plant Strriana maritirna are as truly 

protectively adapted as is its singular stick-like appearance. 

348. -. 1911. The influence of regenerating tissue on the animal body. Papers Tortugas 

Laboratory 3, no. 41-48. 

Carnegie Institution of Washington Publication Nuniber 132. 

It is stated that when the adult animal body begins to regenerate new tissue in order to 

replace a lost part, or when abnormal secondary growths arise, the condition of growthequilibrium 

is disturbed and such a disturbance is followed by changes which affect the 

usual physiological condition of the body. The question as to whether the changes 

following normal regenerative growth are in any way similar to those effects resulting from 

malignant or abnormal secondary growths arises. 

349. ------ 1908. Studies of tissue growth. I. An experimental study of the rate of regeneration in 

Cassiapea xarna~zchuria (Bigelow). Pupers Tor-tugas Laboratory 2: 61 -102. 


The author responds to the studies of 7xleny (1903 and 1905) in which he suggested that 

the greater the degree of injury within limits, the morc rapid the rate of regeneration. 

Zeleny suggested that the animal with !he greater number of appendages removed might 

exercise the regenerating ones more than the animal with less: activity should increase the 

rate of regeneration in animals. The author tests the influence of rest and activity on 

regenerating tissues of medusa and finds no increase in the regeneration rate from activity. 

Rate of regeneration was also tested against food consumption, distance of cuts from the 

margin of the medusa disks, cuts from different parts of variously shaped surfaces, removal 

of oral epithelium of different sizes and at different distances, and the influences of 

changed chemical conditions on regeneration.

350. Stoddart, D. R. and F. R. Fosberg. 1981. Topographic and floristic change, Dry Tortugas, Florida, 

1904-1977. Atoll Research Bulletin 253: 1-56. 

Topographic and floristic surveys ofthe Dry Tortugas Keys in 1904, 1915, and 1937 have 

been used in discussions of the changing relationships between area and floristic diversity 

on small islands over time, and of the processes of colonization and extinction. It is shown 

that earlier topographic surveys are in general too unreliable to be so used. A list of Dry 

Tortugas plants, including all published records was as well as new collections made in 

1962 and 1977, is presented, together with maps of the keys made in 1977. The total flora 

of about 130 species includes at least 35 native species, including 5 species of sea-grasses 

and 4 species of mangroves. Introduced species are largely confined to the two largest 

islands, and the floras of the smaller keys are dominated by a small number of native 

species. 

351. Stone, R. G. 1931-1932. Effect of irradiation by radium upon regeneration in marine annelids. 

Carnegie Institution of Washinjiton, Yearbook. 

Note: published as follows: 1931, v. 30, p. 395; 1932, v. 31, p. 279. 

The effect of combined beta and gamma radiations upon regeneration in polychaetes is 

studied. Histological material is being used to determine the source of new tissue in 

regenerated segments and to discover what tissues are affected by radiation. 

352. -- 1934. Radium radiation effects on regeneration in Eur-atella chatnberl~t. Papers Tortugas 

Laboratory 28: 157-66 (issued Jan. 1933). 

Carnegie Institution of Washington Publication Nutnber 435. 

Regeneration in the polychaete annelids has been investigated in some instances but the 

histological changes are not so well known as in the oligochaetes. The influence of X-rays 

and radium upon regeneration in various animals has been demonstrated, but the 

polycheates ha\,e seldom been used in thesc investigations. It has been found that the 

effects of radiation are often limited to specific tissues; by reason of their greater 

susceptibility they may he injured or destroyed by the exposure. During the summers of 

1931 and 1932 the author was able to study the effects of radiation upon polychaete 

regeneration at the Tortugas Laboratory. In the sabellid Euratellu chan~berlin, posterior 

regeneration of abdominal segments is rapid and complete. Regeneration is inhibited by 

sufficient exposure to the beta and gamma rays of radium. Similar exposure to gamma rays 

alone has no effect upon the amount of regeneration. No structural changes were observed 

in the radiated worms to account for this change. It is suaested that ionization induced by 

the beta rays is responsible for the failure of regeneration 

353. ---- . 1936. Regeneration in the cirratulid Cii-rir~eris. Papers Tortugas Laboratory 29: 1-12 

(issued Nov. 1935). 


There has been considerable study of polychaete regeneration , but the observations are not 

as extensive as those among the oligochaetes. This investigation of Cirrineris was 

undertaken to determine the extent of segment replacement and the source of the new 

tissues. Material was secured at the Tortugas Laboratory. In summary a head region and 

six to seven segments posterior to it are regenerated when more than this number are 

removed. Posterior regeneration is rapid and complete; the approximate number of 

segments removed is replaced. Wound closure is effected in the same manner at both ends 

of Cirf-ineris. The edges of the everted intestine unite with the epidermis to close the body 

cavity. New nervous tissue arises by proliferation and inward migration of cells from the 

adjacent epidermis. The old nerve cells do not participate in regeneration. Material for 

regeneration of the intestinal lining arises by proliferation within the gut epithelium. 

Mesodermal structures regenerate from old mesodermal tissues. Replacement material is 

supplied by (a) nuclei and cytoplasm from lnuscles cells and connective-tissue elements

after degeneration of their differentiated cytoplasm; (b) peritoneal cells which furnish most 

of the new material. 

354. Stoneburner, D. L. and C. S. Harrison. 1981. Heavy metal residues in sooty tern, Sfenla fuscuta 

tissues from the Gulf of Mexico and North Central Pacific Ocean. Sciertce of the Total 

E~~viro~iri~ent 19, no. I: 51-58. 

Tlie comparison of mean cadmium, mercury and selenium concentrations in the eggs, 

feathers and body tissues of breeding sooty tern (Sterna fuscata) from the Dry Tortugas, 

Florida, and Lisianski Island, Hawaii, supports the hypothesis that a physiological 

mechanism exists which functions in tlie detoxification of heavy metals. The data collected 

from two geographically isolated populations of this pelagic bird indicate that the 

mechanism responds in a uniform manner to widely different environmental levels of heavy 

metals. The data and observations suggest that the mechanism evolved in response to 

natural fluxes of heavy metal concentratiotis in the marine ecosystem, not in response to 

recent injections of heavy metal laden industrial wastes. 

355. Stoneburner, D. L., P. C. Patty and William B. Robertson Jr. 1980. Evidence of heavy metal 

accumulations in sooty terns. Science of the Total Environme~lt 14, no. 2: 147-52. 

Sooty terns from the population that nests at Bush Key, Dry Tortugas, Florida, had 

substantial burdens of Cd, Hg and Se. Analysis of selected tissues, feces and eggs by 

neutron activation techniques showed highest levels of Hg in eggs, feathers, and blood; of 

Cd in kidney and bone; and of Se in kidney, liver, and feathers. The concentrations of Cd, 

Hg, and Se in the eggs suggests that the heavy metals are being transmitted to succeeding 

generations. The significance of the concentrations, their effect on the reproductive 

success of the population, and the question of whether or not the metals tmnsmitted to eggs 

represent "bio-niagnification" merit further work. 

356. Strom, R. N., R. S. Bramen, W. C. Jaap, P. Donan, K. B. Donnelly and D. F. Martin. 1992. 

Analysis of selected trace metals and pesticides offshore of the Florida Keys. Floridu 

Scientist 55, no. 1: 1-13. 

Trace metal and pesticide contents of sediments and producer and consumer organisms 

were analyzed from samples taken from eighteen stations off the Florida Keys from 

Biscayne National Park to the Dry Tortugas. Samples were analyzed for total mercury, tin 

(inorganic and organic), arsenic (inorganic and methylated), lead, copper, cadmium, and 

halogenated pesticides. Pesticide concentrations were below detection limits. In general, 

concentrations of trace metals increased from sediments to producers to consumers at each 

station. Though the concentrations tended to be low, some deviations were ascribed to 

liuman inputs. Fewer significant correlations were observed than expected, possible 

because of the dependence of tile uptake niechanism upon the ability of the system 

(sediment, producer, consumer) to remove trace metals from particular materials. Sponges 

have this ability and may represent a useful means of monitoring the quality of the 

environment on a sustained basis. The results are generally consistent with a relatively 

clean environnient with some localized anthropogenic effects. 

357. Stromsten, F. A. 191 1. A contribution to the anatomy and development of posterior lymph hearts of 

turtles . Papers Tortugcis Lahorator.,y 3: 79-87 

Carnegie 11dtution of Wuslzi~~gto~t PuOlicatior~ Number. 132. 

This article concludes that the development of the posterior lymph hearts of turtles is 

initiated by he vacuolation of tlie postiliac niesenchymal tissue during the middle and latter 

part of the second week of development of the loggerhead turtle. Tlie spongy tissue thus 

formed is then invaded by capillaries from the first two or three dorsolateral branches of 

the caudal portion of the postcardinal veins. The final stage in the development of the 

posterior lymph hearts is rcached by the dilation and confluence of these vcno-lymphatic

sinuses, from before backward, forming a pair of sac-like organs, each with a single central 

cavity. 

358. 1910. The development of the posterior lymph hearts of the Loggerhead turtle 

Thalassochelys caretta. Proceedings of the Iowa Academ), of Science 17: 227-28. 

Observations made on the lymphatic systems of turtles indicates their origin is more or less 

independent of the venous system. Later investigations confirm this view, suggesting that 

posterior lymph hearts of the loggerhead turtle are developed from embryonal cappillaries, 

which have been captured and modified by the mesenchymal spaces of the post-iliac 

regions of the body. 

359. Tandy, G. 1931. The superficial structure of coral reefs; plant succession upo~l prepared substrata. 

Carnegie I~istitution of Washington, Year Book 30, 32: 395; 265. 

Plant and animal successions were examined on concrete cubes planted in the water at 

three sites: Fort Jefferson moat, an iron wreck east of Loggerhead Key, and northwest of 

Loggerhead Key. 

360. Tartar, V. 1938-1939. Regeneration in the starfish Linckia and in the protozoan Cond)dostorna. 

Carriegie Institutiot~ of Washington, Year Book. 

Note: published as follows: 1938, v. 37, p. 99-102; 1939, v. 38, p. 230.31. 

Regeneration experiments were conducted on starfish with and without arms and isolated 

arms. Under normal conditions polarity of arms is not altered by isolation. Tube feet cell 

differentiation was examined in relation to color changes. In the ciliate, Cond)dostomu, the 

normal form and typical arrangement of cytoplasmic differentiations may easily be altered. 

361. Tashiro, S. 1914-1915. Further studies on COz in sea water and C02 production in tropical marine 

animals. Carnegie Institution of Washingrori, Year Book. 

Note: published as follows: 1914, v. 13, p. 170; 1915, v. 14, p. 217-19. 

Studies were conducted on the presence of "free CO2 in sea water." A rapid metl~od to 

estimate amounts of C02 produced in sea water by marine animals was devised. 

362. Taylor, J. B. 1981. Premetamorphic veligers of Fort Jefrerson Dry Tortugas, Gulf of Mexico, and 

Beaufort Inlet, North Carolina. Bulletin of the American Mulacology Union, Inc. 50: 29- 

30. 


363. Taylor, W. R. 1928. The marine algae of Florida, with special reference to the Dry Tortrrgas. 

Papers Tortugas Labor-ator)' 25: 1-219. 

Carnegie Institution of Washington I'ublication Number 379. 

The study of marine vegetation of the Dry Tortugas was originally undertaken to provide a 

simple check-list of algae of the islands for use of persons visiting the Carnegie Laboratory 

there, with a description of the more important ecological features and records of the 

locations where plants of experimental importance might be found. When it was discovered 

that information about Florida algae in general was scanty, the study extended to a 

thorough study of all available Florida material. Records of the occurrence of marine algae 

on the east coast of Florida and theFlorida Keys were collected. This is the first time, since 

Harvey, Farlow and Melvill, that an attempt was made to list completely the Florida algae. 

364. --- , 1925. The marine flora of the Dry Tortugas. Revue Al~ologiqrte 2: 113-35. 

The marine algae of the Dry Tortugas are listed, and a description of the distribution is 

given of the important types throughout the al:ea.

365. Teas, H. J. and P. B. Schroeder. 1971. Vegetation analysis in the Dry Tortugas by remote sensing. 

Quarterly Journal of the Florida Academy of Science 34, no. (Suppl 1): 13. 

Detailed ground truth observations were carried out on the four large islands in the Dry 

Tortugas using aerial photography and 12s image enhancement equipment. Several 

vegetation associations (strand-beach, strand-dune, strand-scrub) are distinguished and a 

number of plant species identified (Rhizophora, Lagurrcularia, Rursera, Conocarpus, 

Casuarina, Cocos, and Phoenix). 

366. Tennent, D. H. 191 1. Echinoderm hybridization. Papers Torru,gasLaborator). 3: 117-51. 


The Toxopneustes female x Hipponoc male and the reciprocal cross Hipponoe female x 

Toxopr~eustes male were easily made after allowing the eggs lo stand in sea-water for some 

hours before fertilization. In the embryos of both crosses made in ordinary sea-water, 

which was alkaline, the Hipponoe influence showed a tendency to predominate. It is 

suggested that the variations in the alkalinity of the sea-water, which have been brought 

about artificially, may correspond to normal seasonal changes. The results of this and of 

other investigations show species tendencies toward different grades of temperature and of 

alkalinity. The explanation of the preponderance of one character over another in 

echinoderm hybrids seems to lie in the reaction of the species toward a complex of factors. 

367. ---- . 1920. Evidence on the nature of nuclear activity. Proceedings of the National Academy of 

Science 6: 217-2 1 

The author describes the results of the Arbaciu eggs and other materials examined. 

Basophilic bodies found are not in the nature of chromidia, but are the result of indirect 

nuclear activity. 

368. --- . 1942. The photodynamic action of dyes on the eggs of the sea urchin, Lytechbw 

wriegatus. Papers Torrugas Laboratory 35: 1-153. 


The work recorded in this paper was begun as a study of the experimental modification and 

control of cell division in the egg of the sea urchin, Lytechb7us variegatus. Early in the 

investigation the photodynamic effects of the dye neutral red were found to be striking and 

it was decided to undertake a study of the effects produced by other dyes. Transmission of 

visible light by some of the filters was so low that the intensity of the light transmitted was 

not sufficient to produce photodynamic effect. With dyes that produced a photodynamic 

effect, irradiation of a solution of the dye resulted in the formation of a photocompound. 

This photocompound was the active agent in the production of the photodynamic effect. 

The threshold for violent surface reaction (blister cytolysis) of Lytechirlus eggs in l:150,000 

solution of neutral red in sea water lay at about 2500 foot-candles. From this point to 

about 4300 foot-candles violent surface reaction usually stood at about 2 per cent. Between 

7000 and 7500 foot-candles it increased to 20-25 per cent, and between 800 and 9500 footcandles 

it increased to 75-90 per cent. At intensities from 300 to 10,000 foot-candles there 

was a regular increase in the violence of the surface reaction and complete inhibition of the 

cleavage processes. Irradiation in some of the solutions of dye at temperatures above 32" C. 

resulted in injury from which the eggs did not recover. In blister cytolysis the formation of 

blisters starts at a single point. Adjacent blisters come into contact with one another until 

the entire surface is covered. The content of these blisters is liquid, is clear on the living 

egg, and in the fixed egg seems to be the same as the cytoplasm of the egg with all formed 

components removed. It is conceivable that these coniponents could be filtered out, but 

there is no evidence of the accumulation of granules at the point where the cytoplasm might 

have been extruded .

369. Tennent, D. H. and V. H. Keiller. 191 1. The anatomy of Pentaceros reticulatus. Pupers Tortugas 

Lnboratory 3: 113-16. 


This account is a description of the anatomy of Pentaceros reticulatus. Figures are used to 

illustrate the organs which are described. Those which seem of greatest interest are the 

intestinal caeca. These were found in some instances to be greatly distended, stimulation 

causing their contraction. In this behavior we support the idea of the analogy of the 

intestinal caeca of the starfish to the respiratory trees of the holothurian, an idea which has 

been based upon the similarity of position of these organs. 

370. Tennent, D. H., C. V. Taylor and D. M. Whitaker. 1929. An investigation on the organization in a 

sea-urchin egg. Papers Tortirgas Laboratory 26: 1-104. 


In this report the eggs of the sea urchin, L)'teclzinus, were studied from samples taken at the 

Tortugas. The differentiation of ectoderm-forming substance over the entire surface of the 

egg begins before fertilization by the exclusion of the endoderm-forming material from the 

superficial layers of the egg. The number and relative distribution of ~nicromeres is 

independent of the plane of section and of the size of the fragment. There is no localization 

of micromere-forming material. 

371. Tennent, D. H., M. S. Gardiner and D. E. Smith. 1931. A cytological and biochemical study of the 

ovaries of the sea-urchin Echi~ioinetra lacunter. Papers Tortragas Lnhorator)' 27: 1-46. 


The investigations upon which tl~is paper is based constitute a new method of attack on the 

problem of thc functional significance of chondriosornes, Golgi bodies and other 

"inclusions" in protoplasm. In 1926 a definite research progranm for histochemical and 

biochemical study of the eggs and ovaries of the sea-urchin Echinonietra lacunte~- was 

begun. Summarizing: analytical figures for the percentage of lipids and of glycogen are 

given. The amount of the latter, 12.42 per cent, and 12.72 per cent of the dried extracted 

tissue is high. In addition, the presence of cerebrosides and sphingomyelin are indicated. 

The lipid composition of this tissue seems to be complex. The unsaturation of several 

preparations used in the study of staining reactions was determined, to find out if there was 

any correlation between unsaturation and staining with omit acid (see Section I). On the 

whole the lipids are probably more unsaturated than similar preparations from mammalian 

tissues. 

372. Thiele, J.. 1916. Molluskenfauna Westindiens. Zoologisclw Juhrhucher Supplement 11: 109-32. 

A listing of the rnollusks of the West Indies is given and a preliminary catalogue of the 

shell-bearing marine mollusks and brachiopods of the southeastern coast of the United 

States. This article is in German. 

373. Thompson, M. J. and T. W. Schmidt. 1977. Validation of the speciesltime random count technique 

sampling fish assemblages at Dry Tortugas. PI-oceediqs of the Third I~~tenlatiot~al Cord 

Reef Sytnposiram, No. I:283-288. Miami, Florida: RSMAS, University of Miami. 

Ichthyofauna at four coral reef sites in Fort Jefferson National Monument, Dry Tortugas, 

are compared during summers 1975 and 1976. Samples were taken using the speciesltime 

random count technique, a newly developed visual censusing method based upon the rate at 

which species are encountered by a free swimming observer. Data were collected by 

diffcrent observers during the two years' sampling. Within nine fish families dominating 

the Tortugas ichthyofauna, the rank of five did not vary at all between 1975 and 1976 

sanlplings. Among the four families exhibiting changes in abundance, only the Serranidae 

showed a variation greater than 10.0%. The marked variation of 25.8% within this family 

is attributed to identification problems within the genus Nypoplectru,~. Overall numbers of

species and relative species abundances within each sampled coral reef area showed 

minimal variation between years. The species rank correlation coefficient (Spearman's r s) 

between two years of observations was 0.92. High correlation between results from two 

different observation teams shows the speciesltime random count technique to be a highly 

reliable method of comparing coral reef fish assemblages. 

374. Thorp, E. M., A. Mann, T. W. Vaughan and F. J. Haight. 1936. Calcareous shallow-water marine 

deposits of Florida and the Bahamas. With appendices: I.Mann, A. Diatoms in bottom 

deposits from the Bahamas and the Florida Keys; 2. Vaughn, Thomas Wayland. Current 

measurements along the Florida Coral Reef Tract with notes on current observations, 

Florida Keys, June, October, November, 1914. See separate entries for Appendices. Paperr 

Tortugas Laborafory 29: 37.143 (issued Dec. 1935). 


Determinations of the quantities of material derived from organic and inorganic sources 

have yielded the following results: Coralline algae, collectively, are shown to be the 

organic group that makes the largest contribution of organically secreted calcium 

carbonate. Next in order of magnitude are the mollusks, followed in descending order by 

foraminifera, madreporarian corals, alcyonarian spicules, worm tubes, crustacean 

fragments, and Bryozoa. The principal non-calcareous mineral is quariz. Quantitative 

counts of alcyonarian spicules show that they are relatively minor components of the 

sediments, being exceeded by madreporarian fragments in a ratio of about 2.5 to 1. 

Terrigenous minerals are remarkable scarce. A very small amount of volcanic glass and a 

few species of heavy minerals occur well distributed over the region. The sources of all the 

volcanic glass and some of the heavy minerals are thought to be distant and that they are 

wind blown. Coal and ashes brought from outside sources by human agencies have been 

introduced into the sediments of Tortugas lagoon, and, in smaller quantities, in a few other 

places. 

375. Tomas, C. R. and D. G. Baden. 1993. The influence of phosphorus source on the growth and 

cellular toxin content of the benthic dinoflagellate Proroco~trrrnz lima. Fqfh l~~ter~~arior~al 

Conference on Tmic Marine I'hgroplankton, 565-70. St. Petersburg, Florida: Florida 

Marine Research Institution. 

The relationship between toxin content and nutritional status of the toxic marine 

phytoplankton Prorocentrum limu was examined in a culture from the Dry Tortugas, grown 

with inorganic phosphate and glycerol phosphate. Phosphorus source affected growth, 

maximal densities, and okadaic acid content of Prorocenrnrm limn. 

376. Torrey, H. B.. 1927.1928, Effect of thyroxin on division rates of various cells. Carmgie ins~itutio~i 

of Wasl~ington, Year Rook. 

Note: published as follows: 1927, v. 26, p. 228-229; 1928, v. 27 , p. 287. 

Thyroxin depressed ccll division and differentiation in eggs of sea-urchin (Echinometra 

lacunler), ascidians (Phallusic~ nigra), and hydroids (Pen~~aria ticit-ella), collected at 

Tortugas. 

377. Treadwell, A. L. 1911. Eunicidae of Tortugas. Hulletin of the Ar~rerica~~ Museurn of Natur-a1 History 

30: 1-12. 

Systematic accounts of six species of polychaetous annelids are provided from specimens 

collected in the dead coral rock around Fort Jefferson during 1908. Some species are 

redescribed because of their earlier incomplete descriptions. Brief notes on their 

abundance and distribution are included. 

378. Treadwell, A. L. 1921. Leodicidae of the West Indian region. Papers Tortugas Lahomtor;)~ 15: l- 

131.


A systematic study based on specimens of the family Leodicidae is presented. Collections 

were made at the Dry Tortugas and Key West region of Florida, and in Bermuda, Porto 

Rico, Montego Bay, Jamaica, and Tobago. Collecting was done along shore or in 

comparatively shallow water. The Leodicidae are a well-defined family in which the most 

constant structures are internal rather than external. There is always a well-developed jaw 

apparatus, composed of bilaterally arranged series of chitinous plates developed in a 

pharyngeal pouch, and capable of protrusion for feeding purposes through the mouth. The 

structure of these jaws was used by Ehlers as a basis for classification, though the external 

organs are a more convenient means of recognition. 

379. 1917. Polychaetous annelids from Florida, Porto Rico, Bermuda, and the Bahamas. Papets 

Tortugas Laboratory 11: 255-68. 


This paper is a preliminary description of some new species belonging to the Polychaetous 

annelids, as well as new species of other families which have been collected incidentally in 

this work, including a new sabellid belonging to the collection of the American Museum of 

Natural History. 

380. Ubelacker, J. M. 1982. Review of some little-known species of syllids (Annelida: Polychaeta) 

described from the Gulf of Mexico and Caribbean by Hermann Augener in 1924. 

Proceedings of the Biological Societ)' of Washington 95, no. 3: 583-93. 

The types of six little-known syllid species described by Augener in 1924 from the Dry 

Tortugas, Florida, and from St. Thomas and St. Croix in the West Indies, were reexamined. 

Ha~~los.yllidesflorida~ta is a sexual form herein synonymized with it. Eusyllis anfillensis 

and Syllis (Typosyllis) tigrinoides are synonyms; the latter name is retained. Syllis 

(Tyl~o.s)'llis) fuscosuturata has previously been synonymized with Rrarzchiosylli.~ exilis 

corallicoloides and remains a valid species. 

381. Vaughan, T. W. 1914. The building of the Marquesas and Tortugas Atolls and a sketch of the 

geologic history of the Florida Reef tract. Papers Turftigas Laboratory 5: 55-67. 


The study of the geology and the geologic processes of the Florida reef tract, and especially 

of the Tortugas and the Marquesas, has been continued since 1910. It is now possible to 

outline the salient geologic episodes in the history of the entire Florida Reef tract and to 

Institution comparisons with other coral-reef areas. 

382. --- , 1910. A contribution to the geologic history of the Florida Plateau. Papers Tortugas 



This paper is the outgrowth of the author's association with two organizations, the United 

States Geological Survey and the Carnegie Institution of Washington. The author has 

visited all the principal keys between Miami and Key West, to collect and study bottom 

samples, particularly the deposits accumulating behind the keys, and to examine several 

important living coral reefs around the Tortugas. The scope of the paper was enlarged to 

trace the geologic history of the Floridian Plateau from Oligocene to Recent time. This is to 

be regarded as only a sketch of the geologic development of the Floridian Plateau, as many 

problems need solution and many phases of its history need further investigation. Perhaps 

its principal value may be in directing attention to some of the unsolved problems. It is 

necessary to know more accurately the amount of water discharged by the streams and the 

quantities of solids borne by then1 to the sea. The chemical processes of precipitation have 

not been sufficiently studied. There is also great need for more extensive studies of the 

marine bottom deposits within the 100-fathom curve. The deep wells recently put down on

Key Vaca, Big Pine Key, and Key West have given valuable data, but deep wells are also 

needed on the Marquesas, and the Tortugas, in order to discover what underlies the surface 

formations. It is hoped this paper may serve as a convenient summary of the present 

knowledge of the geologic history of this interesting region, perhaps present an 

interpretation somewhat different from those preceding, and be a stimulus to further 

investigation. 

383. --- . 1915. Coral-reefs and reef corals of the southeastern United States; their geologic history 

and significance. Rullerin of the Geological Societj' ofAmerica 26: 58-60. 

The geologic history of the extensive coral reefs of the southeastern United States and 

near-by West Indian Islands was outlined, and the bearing they have on the theory of coral 

reef formation was indicated. The author stated his conclusions regarding the Florida coral 

reefs as follows: (1) Corals have played a subordinate part, usually a negligible part in the 

building of the Floridean plateau; (2) every conspicuous development of coral reefs or 

reef corals took place during subsidence; (3) in every instance the coral reefs or reef corals 

have developed on platform basements which owe their origin to geologic agencies other 

than those dependent on the presence of corals. The conclusions in this report are 

summarized as follows: Critical investigations of corals as constructional geologic agents 

are bringing increasing proof that they are not as important as was believed. All known 

modern offshore reefs which have been investigated grow on platforms which have been 

submerged in Recent geologic times. No evidence has been presented to show that any 

barrier reef began to form as a fringing reef and was converted into a barrier by subsidence. 

There were platforms in early Teritiary time on the site of inany of the present-day 

platforms, and evidence has yet been adduced to prove long-continued, uninterrupted 

subsidence in any coral-reef area. The width of a submerged platform bordering a land 

area is indicative of the stage attained by planation movetncnt. The importance of coral 

reef studies to geology suggests they are only a conspicuous incident in time. 

384. -. 1936. Current lneasureinents along the. Florida coral reef tract. Paperr Tortngas Laboratory 

29: 129-41. 

Carnegie Institution of Washington Publication Number 451. Note: This is Appendix 2 to 

Calcareous shallow water marine deposits of Florida and the Bahamas by Eldon Marion 

Thorp. 

During June and July 1914, while studying the phenomena associated with the Florida 

Coral-Reef Tract the author initiated a series of current measurements by using Ekinan 

current meters. South of the Tortugas a non-tidal current toward the west is clearly 

indicated. The data here presented are inadequate for positive conclusions regarding the 

Counter Current. 

385. -. 1915. The geologic significance of the growth rate of the Floridian and Bahamian shoalwater 

corals. Journal of the Washingtoft Acadony of Scier~ce 5, no. 17: 591-600. 

The object of this investigation has been to aid in understanding the amount of work stony 

corals may do as constructional geologic agents, and especially in the formation of coral 

reefs. This subject needs to be studied from at least five different view points, e.g.: (1) the 

quantity of material contributed by corals and that contributed by other agents must be 

estimated and the respective proportions detern~ined; (2) in coral reef areas the ratio of the 

area covered by corals to that not covered by corals should be estimated; (3) the relations 

of coral reefs, continuity and discontinuity must be determined; (4) marine bottom deposits 

must be analyzed according to the source of the material, and the percentage of the calcium 

carbonate contributed by the differing agents estimated; (5) the rate of growth of corals 

needs to be known. There is no single formula for the growth rate of corals, as it varies by 

species and ecologic conditions. Observationslexperi~nents on the growth rates of Tortugas 

corals are as follows: (I) Colonies obtained from the planule whose history was known,

and were planted off the moat wall and on the NW side of Loggerhead Key; (2) Colonies 

cemented to tiles and planted at the same sites as above; Colonies naturally attached at sites 

described above. The reef species of greatest concern and importance is Orbicella 

(Montastrea) ai~iruluris followed by in importance, Muearldm sfrigoso, M. 

labyriut/~ormes, and Sideraswea siderea. The upward growth is critical of the massive 

heads Orbicella (Mor~tastreaJ anr!ularis which form the strong framework of the reef and 

averages 1 foot in 43.54 years or 7 mm /year and which might form a reef 150 feet thick in 

between 6500 and 7600 years. A table on the average annual growth rates of corals from 

the Florida region is provided. 

386. -- 1914. The platforms of barrier coral reefs. Bulletin ofthe America,! Geogruphical Society 

46: 426-29. 

The author states that there are three kinds of coral reefs: fringing or shore reefs which 

occur along the strand line, barrier reefs which occur at varied distances off shore and have 

lagoons from one to as inuch as forty fathoms depth between them and the strar~d line, and 

atolls, which are ring-like and enclose lagoons. As the relations of barrier reefs and atolls 

to the platforms on which they stand constitute the essential part of the theory of 

development of Recent reefs, the discussion of coral reef theory has been waged over the 

interpretation of these relations. The object of this paper is to point out the relations of 

barrier coral reefs to the last dominant change in position of the strand line and to indicate 

the organisms forming Recent barrier reefs have played in building the reef platforms. 

387. --- . 1914. Preliminary remarks on the geology of the Bahamas, with special reference to the 

origin of the Bahamian and Floridian Oolites. Papers Torrugas Luboratory 5: 47-54. 


The author presents a preliminary summary of the information compiled (to 1914). on the 

origin of calcium carbonate sediments in south Florida, and the Bahamas, using various 

hypotheses developed by the leading geologists of the time. These geologists included 

Alexander and Louis Agassiz, Sanford, and Drew who worked in the Dry Tortugas and 

believed the precipitation of calcium carbonate was due to the effects of denitrifying 

bacteria. 

388. - 1916. The results of investigations of the ecology of the Floridian and Bahamian 

shoalwater corals. Proceedings of the Nafioml Acadein)r of Sciences 2: 95-100. 

This paper presents a summary of the knowledge on the ecology of shallow water corals in 

the Florida-Bahamian region, with a detailed description of new infomation on food 

preferences of corals, and salinity and water tempesature tolerances, based on studies 

conducted at the Dry Tortugas and the upper Florida Keys area. Mayer, at the Tortugas 

Laboratory, found that temperatures of 13.9 'C would exterminate the principal Floridan 

corals; similar results were found for corals around Australia. Light experiments at Fort 

Jefferson suggested vigorous coral growth in well lit wharf areas, and little growth in piling 

areas of perpetual shading. Tests conducted at the Marine Laboratory suggested that corals 

could survive at salinities of 27-38 ppt, but not as low as I9 ppt. Other conditions 

necessary for vigorous coral growth are maximum water depths of 45 meters and rocky or 

firm bottoms, without silty deposits. The growth rate of corals was determined by planting 

planulae in the laboratory, by measuring colonies which bad been cemented to disks, and 

fixed on heads of stakes driven into the sea bottom. Measurements of colonies naturally 

attached were also made. Plantings at the Tortugas were made at Loggerhead Key, and 

around Fort Jefferson. The more massive the coral, the slower the growth; while the more 

rarnose (Acropora palmafa) and the more porous the skeleton, the more rapid the growth. 

The growth rate of the principal reef builders (massive corals) in the Florida region, 

Or6icella (Moura.rfreuJ a111ru1uri.r is from 5-7 mnl per year and would form a reef of 150

feet in thickness from 7620 to 653 1 years. A. palrnata may build a similar thickness in 

1800 years. 

389. --- . 1914. Sketch of the geologic history of the Florida coral reef tract and comparisons with 

other coral reef areas. Journal of the Washington Academy of Sciences 4: 26-34. 

The author presents the two hypotheses for the formation of atolls: one attributes atolls to 

the submarine solution of the interior of a mass of limestone; the other accounts for them 

by constructional agencies. The author believes the solution theory is disproved by a 

chemical examination of sea-water from a Tortugas lagoon. He believes the Marquesas 

and the Tortugas are constructional phenomena and owe their configuration to the 

prevailing winds and currents. 

390. -- , 1918. The temperature of the Florida Coral-Reef Tract. Papers Torrugas labor at or)^ 9: 

319-39. 


The temperature data presented were assembled primarily for their bearing on the effect 

temperature exerts on the bathymetric and geographic distribution of coral reefs. 

Temperature is also one of the most important factors in determining the geographic 

distribution of sea-level and near sea-level reefs. 

391. Vaughan, T. W., M. A. Goldman, J. A. Cushrnan, M. A. Howe and others. 1918. Some shoal-water 

bottom samples from Murray Island, Australia, and comparisons of them with samples 

from Florida and the Bahamas. Papers Torrugas Laboratory 9: 235-97. 


The present paper is a preliminary contribution to the study of the marine bottom deposits 

in three corel-reef areas: Murray Island, Australia; the Bahamas, and southern Florida. 

Mechanical analyses have been made of all samples except those obtained in 1915, and the 

results of the chemical analyses of a selected set are presented. An attempt has been made 

to outline a method of studying calcium carbonate bottom deposits, in the hope that 

progress may be made toward an adequate classification of such sediments. The Tortugas 

Lagoon samples are coarser than those in Marquesas Lagoon, and those from the latter 

locality are coarser than the Bahania sample from South Bight and the west side of Andros 

Island. Some terrigenous material, mostly quartz sand, is washed into Biscayne Bay, 

Florida, and into the sounds south of it, but otherwise practically none reaches the key and 

reef region. The Florida area is therefore a perfect example of limestones forming in shoal 

water near a land area which is not crossed by large streams. The Fe,O, content of the 

Florida samples seems somewhat higher, up to about 0.37 per cent, than that of the Bahama 

samples. Reconsideration of the evidence bearing upon the precipitation of CaCoi in 

tropical and subtropical waters and the possibility of its re-solution by ocean-water leads to 

the conclusion that precipitation is resulting from both organic and inorganic agencies, and 

that no appreciable re-solution is taking place in the water. 

392. Visscher, J. P. 1930-1931. Distribution of barnacles with special reference to behavior of larvae. 

Can~egie institution of Washington, Year Book 

Note: published as follows: 1930, v. 29, p. 346; 1931, v. 30, p. 397. 

A study of the barnacles was made, more than twenty species being found. Several species 

appear to be new to science. Behavior of larvae appeared to vary depending on habitat, as 

certain barnacles were found on crabs, others above the tide on pilings, still others only on 

coral and on the spiny lobster, Panulirus ar-gus. 

393. Vukovich, F. M. 1988. On the formation of elongated cold perturbations off the Dry Tortugas. 

Jourrial of Physical Oceanography 18, no. 7: 1051-59. 

The life cycle of a cold perturbation on the boundary of the Loop Current in the Gulf of

Mexico was studied over the period of 18 March to 22 May 1984, approximately a 60-day 

period. The study focused on the behavior of the surface and subsurface area of the cold 

perturbation as it moved along the boundary of the Loop Current. The area of the 

perturbation was defined by an alongflow-scale length, which is the scale length parallel to 

the unperturbed flow of the Loop Current, and the crossflow-scale length, which is the 

scale length perpendicular to the unperturbed flow of the Loop Current. 

394. Vukovich, F. M. and C. A. Maul. 1985. Cyclonic eddies in the eastern Gulf of Mexico. Journal of 

f'hysicol Oceanography 15, no. 1: 105.17. 

Cold-domed cyclonic eddies juxtaposed to the cylconic shear side of the Gulf Loop 

Current are observed in si~nultaneously obtained hydrographic, current meter mooring, and 

satellite data as cold perturbations on the northern extreme of the current and grow either 

into a cold tongue or a quasi-stable meander off the Dry Tortugas, Florida. Areal 

shipboard surveys show closed isopleths of temperature and salinity, and surface 

geostrophic current speeds relative to 1000 db are in excess of 100 cm s super(-I). The 

diameter of the cold domes varied from 80 to 120 km. 

395. Wallace, W. S. 1908. A collection of hydroids made at the Tortugas during 1908. Carr~egie 

lr~stiruriorl of Wa.rhington, Year Hook 7: 136-37. 

At least fifty species of hydroids were collected. A tentative list of those identified is 

provided. 

396. Wartnian, W. 1929. Studies on Echinometra. Cnrnegie Institution of Washingtott, Year Book 28: 

277. 

(No data report provided). 

397. Watson, J. B. 1908. The behavior of noddy and sooty terns. Papers Torfugas Laboratory 2: 187- 

255. 


The work presented in this report is preliminary in nature. Following a general description 

of the two species, a geographical situation and present history of the tern colony at the 

Tolmgas is given All observations were recorded during the nesting season on Bird Key, 

a small coral island covered in part by bay cedar, mixed with cactus in the central western 

parts of the island, with little vegetation elsewhere. Observations on their foods and 

fceding habits indicated that the birds usually feed in groups, never swim nor dive, but 

ski111 along the surface picking up small fish being attacked by larger fish. Feeding 

distance from the Key was estimated between 4 and 10 knots. Mating has been suggested 

prior to arriving at the Tortugas, although some indications of sexual activity occurred for 

the noddies, but not the sooties. Noddies nest in vegetation, while sooties build nests in 

sand. Usually one, sometimes two eggs are laid, with a period of incubation for the noddy 

from 32 to 35 days. The parents alternately feed the young at intervals froni I to 4 hours; 

their general conduct does not greatly change at the arrival time of the young. However, 

two days after the arrival, the parents are more ferocious; both species return to normalcy 

as the chicks gain strength. The birds become exhausted caring for their young, and collect 

upon the beach for "sunning". Egg coloration tests indicated that neither species 

recognized its own egg. As for the noddy's nest environment, it could be disturbed without 

affecting the bird, as long as the egg position was not changed. Tests conducted using 

Porter's learning maze indicated that noddy's were slower than sooties, because of their 

longer standing time. Further maze tests using darkness and maze rotation were 

inconclusive. Other in captivity tests showed that the sooty is highly excitable and nervous, 

whereas the noddy is stolid and indifferent.

398. Watson, 1. B. and K. S. Lashley. 1915. Homing and related activities of birds. Papers Tortugas 

Laborutor). 7: 1-104. 

Carnegie Institution of Washington Publication Number 21 I. 

The present series of studies on the behavior of birds is a direct outgrowth of an 

investigation made on the noddy and sooty terns nesting on Bird Key, Tortugas, Florida. 

The homing "instinct" is the central topic in all the papers. In the 1907 investigation the 

fact appeared that terns possess a homing sense, behaving exactly as do homing pigeons 

when sent away from their nests and young. The 1907 investigation already referred to is 

concerned largely with instincts in terns-those of feeding, nesting, brooding, etc. In general 

the problems of proximate orientation are relatively simple and straightforward. On the 

island of Bird Key the terns make their adjustment to the nest, mate, young, etc., on the 

basis largely of visual habits. There is no evidence of any remarkable or unusual 

sensitivity, nor of the functioning of any hypothetical sense-organ. The present paper 

seems to call for a separation between proximate orientation and distant orientation. 

Mathematical considerations show that at such tlista~ices the goal can not possibly 

(directly) visually stimulate the bird, even granting absolute visual acuity and complete 

absence of haze, etc. This work has shown further, in the terns at least, that there is no 

special Spursinn-special tactual or olfactory mechanism situated in the nasal cavity which 

may function in homing. The task of explaining distant orientation is an experimental one, 

which must yield positive results as soon as proper methods are at hand. Two lines of 

investigation offer hopeful results: the rearing of homing pigeons in a cote, or the rearing 

of the birds in a wire-covered yard attached to a cote. We could tether individual birds to 

the top of the cote by cords which would permit a view only of the neighborhood 

immediately surrounding the cote. With these experiments upon homing, work upon the 

sensory equipment of the homing pigeon should be carried on. It is just possible that these 

aninials possess on certain parts of the body, tactual and thermal mechanisms which may 

assist them in reacting to slight differences in pressure, temperature, and humidity of air 

columns. The experiments and conclusions on homing proper can be found on pages 59 

and 60. These results, which not settling the question of the sensory mechanism by means 

of which the birds rcturn to the nests, do remove all doubts about the fact that the noddy 

and sooty terns can return from distances up to 1,000 miles in the absence of all landmarks. 

The problem of homing has thus become defined, and experimental work of a definite kind 

is needed for its solution . 

399. Wells, R. C. 1922. Carbon-dioxidc content of sea water at Tortugas. Puper,~ Tortujias Laboratory 

18: 87-93. 


It is generally considered that the carbon-dioxide content of sea-water may be increased by 

accessions from the air, by animal life, by the decay of organic matter in the sediments on 

the bottom or elsewhere, by the solution of carbonate rocks, by the contributions of rivers, 

and by gas vents beneath the sea. Sea-water may lose carbon dioxide to the air, to plants, 

and ill the formation of carbonate rocks and the carbonaceous parts of organisms. Mere 

evaporation and precipitation also alter the carbon-dioxide concentration somewhat if other 

conditions remain unchanged. The writer made determinations on sea-water from 

Tortugas, Florida, in June 1919 taken directly from the sea at various points about 

L.oggerhead Key, which reveal unmistakable diurnal variations. The water has sufficient 

contact with plants and sea-weeds to show the effect of photosynthesis on its C02content. 

There is a loss of COz by day and a gain by night. Plant life appears to be the chief agency 

in causing a daily variation in the C02 content. Determinations of C02 should probably be 

made soon after the time the samples are collected, on account of the possibility of the 

decay of organic matter, such as algae, in preserved samples. The average "excess base" 

found at Tortugas corresponds to a norinality of 0.00239. This titration includes 

everything that consunies acid; it represents chiefly bicarbonate, about 0.00183, some

carbonate, about 0.00041, and other substances that contribute to the alkalinity, about 

0.00015. The methods used in arriving at these figures were provided along with a record 

of determinations made at Tortugas and the relation between the carbon-dioxide content of 

the water and time of day. 

400. Westinga, E, and P. C. Hoetjes. 1981. The ititrasponge fauna of Sphecios[~ongia vesparia (Porifera, 

Dernospongiae) at Curacao and Bonaire. Marine Biology 62, no. (2-3): 139-50. 

The infauna of 35 individuals of Sphecio.r/~ongia vesparia (Lamarck,l814) of different 

volumes and frotn different sites and depths have been inventoried and compared. The 

number of sponge-inhabiting taxa is logarithmically related to sponge volume. Biomass 

and total number of the animals contained in the sponge are directly proportional to sponge 

volu~ne. Numerical and taxonomic composition of infaunas from different sampling sites is 

fairly constant. Biomass and total number of sponge inhabiting animals is not significantly 

different for any of the four sampling sites. Several taxa, however, are Inore abundant in 

sponges frotn one or more localities. The ratio of total biomass to total number of 

intraspotige fauna is found to be significantly smaller for sponges collected in deep water 

than in shallow water. Differences frotn and similarities with Pearse's results (1932,1950) 

on the infauna of the same sponge species at Dry Tortugas and Bimini are discussed. The 

relation of the number of contained taxa and the volume of a sponge is compared with the 

relation of island size and number of taxa present according to MacArthur and Wilson's 

island theory (MacArthur, 1972). Finally the erratic occurrence of some taxa as opposed to 

the highly regular occurrence of some other taxa is discussed. It is concluded that the 

composition of the sponge-infauna in specimens larger than 11 is highly constant and that 

the sponge-inhabiting fauna constitules an ecological community. 

401. Westrum, B. L. and P. A. Meyers. 1978. Organic carbon content of seawates from over three 

Caribbean reefs. Bulletir~ of Marbre Science 28, no. I: 153-58. 

Seawater samples from transects crossing three Caribbean coral reefs, including the Dry 

Tortugas, showed variations in concentrations of organic carbon. Total organic carbon 

increased substantially over two fringing rcef crests. ,Most of this increase occurred in the 

particulate fraction at the seaward edge of the crest but in the dissolved fraction at the 

landward edge. Back reef levels of total organic carbon were lower than those seaward of 

the reef. These observations support the hypothesis that organic carbon can be physically 

removed from the benthos at the turbulent reef crest and be subsequently utilized in 

hackreef areas. The reef-flat formation studied in January 1975 in the Dry Tortugas was 

situated off the western shore of Loggerhead Key. This study indicates that organic matter 

contributed at the crest is available as a resource to only a limited portion of the backreef 

comriiunity - that part located directly behind the crest. The observed decrease in TOC 

levels implies quick biological utilization or loss through physical processes. Thus, despite 

continual input, no net accumulation of organic matter occurs in the backreef area, and this 

region can be described as being relatively depleted in organic carbon. If large coral 

formations are present, as at the Dry Tortugas location, they can contribute organic matter 

to the surrounding seawater. 

402. Wheaton, J. 1980. Ecology of gorgonians (Octocorallia: Gorgonacea) at Dry Tortugas, Florida. 

Fiorida Scie~ttist, 43 (suppl. I), 20. 

This study reports the species composition and distribution of the gorgonian fauna of Long 

Key Reef, Dry Tortugas during tlie sunimers of 1975-1976. 23 species were recorded. 

Additional samples increased the number of species to 35. Most shallow reef gorgonians 

were Plexaura. 

403. Wheaton, J. L., W. C. Jaap, B. L. Kojis, G. P. Scbmahl, D. L. Ballantine and 1. E. McKenna Jr. 

1993. Transplanting organistiis on a damaged reef at Pulaski Shoal, Ft. Jefferson National

Monument, Dry Tortugas, Florida, USA: An experinlent to enhance recruitment. (abs.). 

Proceedings of the Seventh luteri~atiorml Coral Reef Symposium, p. 639. Mangilao, Guam: 

University of Guam. 

Grounding of the 475-ft. freighter, Mavro Vetranic, at Pulaski Shoal Reef, Dry Tortugas, 

on 30 October 1989, damaged 3,465 111' of reef surface. After one year, minimal 

recruitment of macrobenthos, principally the alga Dictyota, had occurred. An experiment 

was designed to test effects of adding relief and transplanting sponges, octocorals, and 

scleractinian corals on recruitment of biota to the damaged area. In Sept. 1991, one control 

and two experimental sites, each 9-m2, were selected, marked, mapped, and photographed. 

Large reef rocks were placed in one experimental plot to provide relief and refuge. More 

than 185 organisms (73 species of algae, Porifera, and Cnidaria) were transplanted and 

cemented into five of the nine square-meter subunits in the other experimental plot. 

Transplanting was labor intensive, requiring 64 man-hours to collect, move, and cement 

organisms. We then rephotographed and mapped the plots. Sites will be monitored to 

determine if recruitment of macrobenthic organisr~~s 

enhanced . 

404. Whitaker, D. 1926. Organization of echinoderm egg, and a measurable potential difference betuleen 

the cell interior and outside medium. Carmgie Institution of Washingtou, Year Rook 25: 

248-55. 

Egg development, investigated in the sea-urchin, Lytechi~~us, suggested that the 

differentiation of ectoderm begins before fertilization by the exclusion of the endodermforming 

substances from the superficial layers of the egg. Micromere-forming substances 

do not differentiate before fertilization. 

405. White, S. C., W. B. Robertson Jr. and R. E. Ricklefs. 1976. The effect of Hurricane Agnes on 

growth and survival of tern chicks in Florida. Bird-Banding 47, no. 1: 54-71. 

In June, 1972 Ricklefs and White were studying the energetics of nestling growth in sooty 

terns (Sterna fuscata) at the Dry Tortugas, when Hurricane Agnes passed west of the area. 

High winds, heavy rain, rough seas, and low temperatures prevailed for more than a week. 

Robertson worked in the colony from 28 June, about a week after the storm subsided, to 6 

July. It is reported here the effects of Huvricane Agnes on the growth and survival of 

young sooty terns and brown noddies (Anous stolidus ). 

406. Wichterman, R. 1942. Cytological studies on the structure and division of three new ciliates from 

the littoral earthworm of Tortugas. Papers Tortugas Lnboratory 33: 83-103 . 


During the summer of 1939 this study of Protozoa inhabiting the intestine of the littoral 

earthwor~n Po~~todrellrrs hcrinurlensis Beddard, was begun. The study revealed three 

previously undescribed ciliates: Hysrerociuitrr pontoddu, n. s.p.; Anoplophyra 

macro~~eucleata, n. sp.; and Maupasella leptas, n. sp. This paper describes the ciliates and 

gives an account of fission in each species. Of the 230 worms examined, 64% were 

infected with the ciliates. Generally a worm was parasitized with two different species. 

Observations on the length of life of the ciliates in seawater were recorded. Encystment 

was not encountered. The presence of stages in the life history of acephaline gregarines 

and nematodes was noted. 

407. --- . 1942. A new ciliate from a coral of Tortugas and its symbiotic zooxanthellae. Papers 

Tor-tirgas Laboratory 33: 105-1 I . 


A new ciliate was found on the coral Eurliciu crassa E, and H. and is described as 

Parnerrplofes tortuge~iesis, n. gen. and n. sp., and is placed in the family Paraeuplotidae, n 

fam. The coral it was found on is a member of the Alcyonaria fauna, and is commonly 

found in the Caribbean, and is abundant or1 the reefs of the Tortugas. The morphology of

the ciliate is discussed, as well as the presence of zoozanthellae. The question is posed 

"what is the nature of the syinbiosis between the protozoan and the zooanthellae it 

contains?". 

408. Williams, 0. L. 1932. Studies on the nematodes of Tortugas fishes. Carnegie It~siitution of 


Observations of more than 800 fishes representing about 175 species during the sulnmer of 

1932 demonstrate that the incidence of infestation with nematodes is lower in fishes of the 

Tortugas than in the cooler, shallow waters found farther north. 

409. Willier, B. H. 1936. A study of the early embryology of the Loggerhead sea turtle and of sharks. 

Cnrrtegie Institrrtior~ of Washington, Year Book 35: 92. 

The embryological development of the Loggerhead turtle (Carertu caretta) was examined 

from the time of egg laying to within a few days of hatching. Significant observations are 

presented. 

410. Wilson, C. B. 1936. Parasitic copepods from the Dry Tortugas. Papers 7i)rfugas Luboraiory 29: 

327-47 (issued Dec. 1935). 


Two collections of parasitic copepods contained in the present paper were made at the 

Marine Laboratory of the Carnegie Institution in the Dry Tortugas, involving the handling 

of a large number of the local fishes. In addition to the specific objects of investigation it 

was soon noted that the fish were more or less infested with parasitic copepods and 

isopods. Upon identification, seven of the species are new to science, and two others have 

been made the types of new genera. The other species have been obtained before either in 

the waters around the Dry Tortugas, the Bahamas, the Bermudas or the West Indies. 

41 I. Winegarner, C. E., W. B. Robertson and W. I-loffinan. 1984. Anoiis sageri sagen (brown anole) 

USA: Florida: Monroe Co: Dry Tortugas, Garden Key. Nerpeioiogical Review 15, no. 3: 

77-78. 

Three males and one female specimen were taken on a large pile of bricks and rubble just 

east of the moat surrounding Ft. Jefferson, April 8-10 1983. Population currently seems 

limited to this small portion of the island, so introduction may have been very recent. A 

construction barge moored adjacent to the collection site fiom October 1981 to June 1982 

possibly was a source of colonizing individuals. However the regular arrival of Park 

Service boats and private vessels are other possibilities. 

412. Wolfe, C. A. 1989. "Growth of the Brown Noddy (Anous srolidirs) in the Dry Tortugas (Florida)." 

Master of Science, Florida Atlantic University, Boca Raton. 

The author discusses the slow growth rate of the brown noddy nestlings in the Dry 

Tortugas as to what would be predicted based on adult body size and mode of 

development. This prolonged growth pattern is typical of tropical pelagic seabirds. An 

intraspecific cotnparison of growth rates among several populations of brown noddies, 

indicates that growth of body mass of the Tortugas noddies is significantly faster, the 

development period shorter, and the asymptotic size smaller than in Pacific populations. 

However, there were no differences among the populations in the rates of wing or culmen 

growth. The Bush Key nestlings appear to receive a higher quality diet that contains 

proportionally more fish, while Pacific nestlings receive substantial amounts of squid. The 

Pacific nestlings also seem to be subjected to a thermally more stressful microclirnate, 

which may necessitate the allocation of proportionally more of their total energy to 

thermoregulation and less to growth.

of ciliary currents. Not only is sediment removed very rapidly from the surface, but 

colonies can completely uncover theniselves within twelve hours after being buried in the 

sad. Unlike Fungia which uncovers itself by the exclusive action of cilia, M. areolata first 

distends the tissues with water. Distension for cleansing is essentially different from 

expansion for feeding. Planulation, so far as can be determined at present, has a lunar 

rhythm, culminating about the time of new moon. After an initial stage when upward and 

outward growth are about equal, outward growth predominates, an oval or rounded colony 

with a small basal attachment being finally produced. The stage at which detachment 

occurs must vary with environmental conditions. Colonies may be formed from a single 

planula or from the fusion of several. M, areolata is a species highly adapted for life on 

sand occupying in the Atlantic, the same habitat occupied by the Fungiidae in the Indo- 

Pacific. Adaptability in reef-building corals is discussed. 

419. --- . 1936. Studies on the biology of Tortugas corals. 11. Variation in the genus Siderastrea. 

Papers Tortugas Labornfop 29: 199-208 (issued Dec. 1935). 


This paper presents data on stony corals on the Tortugas reefs. The genus Siderastrea is 

represented at the Tortugas by the two species, S. siderea and S. radians. S, siderea forms 

larger rounded colonies which were not observed above the level of low-water springs. S. 

radians is essentially a shore-living species possessing the physiological adaptations 

characteristic of all shore-living animals. S. radians is capable of great modification both 

in the form of the skeleton as a whole and also in the size and shape of the corallites and in 

the number, slope and thickness of the septa. This species has been enabled, as a result, to 

occupy a variety of habitats, the extremes being represented by the surf region on the beach 

rock on the one hand, and by the still, sediment-laden water in the moat at Fort Jefferson on 

the other. The relation between form and environment in corals is discussed and the 

general conclusion reached that the great success of the Madreporaria is probably due to 

the presence of species highly specialized for a particular environment and also of others 

which can be modified for existence in a variety of different environments . 

420. -- . 1937. Studies on the biology of Tortugas Corals. 111. The effect of mucus on oxygen 

consumption. Pupe~r Tortugas Laboratory 3 1: 207-14 (issued Oct. 1937). 


Experiments are described which indicate that a large proportion of the apparent utilization 

of oxygen by corals is actually due to oxidation of mucus secreted by them during the 

course of the experiment. The amount of mucus varies greatly in different genera and may 

also be increased at certain times, e.g. during planulation. In view of this source of error it 

is i~npossiblc to accept their face value figures which claim to represent either the absolute 

or the comparative rates of respiration in different corals, or general conclusions which are 

based on these figures. Oxidation of mucus may be expected to affect the apparent rate of 

respiration in all aquatic animals which normally secrete mucus. 

421. Yonge, C. M. and H. M. Nicholas. 1940. Structure and function of thegut and symbiosis with 

zooxanthellae in Tridachiu crispata (Oerst.) Bgh. Papers Tortugas Lahomory 32: 287- 

301 (issued Sept. 1940). 


During the visit of the senior author to the Tortugas Laboratory in the season of 1934, a 

number of specimens of a very interesting and beautiful species of opisthobranch mollusk 

were collected a low tide on the surface of the reefs. Exatnination revealed the invariable 

presence of brown unicellular algae, or zooxanthellac, within their tissues. Tridachia 

c~ispata is an elysoid opisthobranch with the body extremely flattened dorsoventrally and 

extended into undulating body folds laterally and terminally. It occurs under stones on the 

reefs of the Torugas group and elsewhere in the West Indies. The feeding and digestive

systems are described. These have the typical elysoid structure with modifications, notably 

in the digestive diverticula, correlated with the excessive flattening of the body. Like the 

othcr members of tlie Elysiidae, T. crispata is a highly specialized herbivore. 

Zooxanthellae are habitually present in a restricted zone a short distance from the margiti 

of the body fold They occur freely within the connective tissue and increase by division. 

There is no evidence that the animal normally consumes them, but reasons are given for the 

suggestion that they may be of value to the animal by removing waste products of 

tnetabolisln produced within the body fold . 

422. Zeleny, C. 1907. The effect of degree of injury, successive injury, and functional activity up011 

regeneration in tlie scyphomedusan Cassiopea xamuchana. Jourr~al of Experimental 

Zoology 5: 265-74. 

This study is part of a series of experiments at the Dry Tortugas Marine Lab on the internal 

factors controllitig regeneration in Cussiopea and other forms, including the degree of 

injury and successive removal of a part and rhythmical pulsations of the disk. The removal 

of 6 to 8 arms constitutes the most favorable degree of injury for the regeneration of each 

arm. When comparing the rate of regeneration of disks, where the disk was made to 

pulsate rhythmically with cases without pulsation, there is no advantage in favor of the 

pulsa(ing ones, but rather a retardation. Other tcsts of successive injury upon regeneration 

were made on chelae of the gulf-weed crab, I'ortuttus sayi, which reveal that the second 

regeneration is greater than the first. However, when the age factor is removed the two are 

exactly alike. 

423. . 1908. Some internal factors concerned with the regeneration of the chelae of the gulf-weed 

crab (I'ortunirs sayi). Papers Torrugas L.uhorarory 2: 103-38. 


The primary object of the experiments described was twofold: the quantitative 

determination (I) of the effect of successive removal of an organ upon its power to 

regenerate and (2) of the character of the changes, if any, produced in the uninjured parts 

of the animal by such removals. It was found that (1) individuals of Portunus suyi with a 

cephalo-thoracic length between 3-9 and 14.5 mm, show but a slight correlation between 

the length of the molting period and the size or age of the animal. (2) The amount of 

regeneration of the right chela between the same limits of size is likewise but slightly 

correlated with the length of the molting period, but is very closely correlated with the size 

of tlie animal. (3) The specific amount of regeneration of the right chela increases slightly 

with increase in size or age of the anirnal.(4) The specific length of the left chela in 

uninjured individuals increases slightly with increase in size or age of the auimaL(5) The 

proportion between the amount of regeneration of a chela and the length of the chela in 

uninjured individuals of the same size is constant, uninfluenced by the size of the animal. 

(6) In single individuals the third regeneration is greater than the second and the second is 

greater than the first. (7) When the correction for change in the power of regeneration with 

size or age is made, it is found that successive removal neither retards not accelerates tlie 

regeneration of the right chela. (8) The right chela is slightly larger that the left in a great 

majority of the individuals. (9) The removal and regeneration of the right chela produces 

no change in the growth of the uninjured left chela. 

424. Zheng, W. and E. S. Van Fleet. 1988. Petroleum hydrocarbon contamination in the Dry Tortugas 

USA. Marine Pollutiotl Rulletirt 19, no. 3: 134-36. 

The present study extends a previous wol-k westward to the point where Florida Keys 

island chain intersects the Golf Loop Current. Since the Dry Tortugas are located in this 

unusual area, they provide an ideal location for examining the fate of petroleum discharged 

into the eastern Gulf of Mexico. Beach tar samples were collected along 1 m wide 

transccts at 18 stations according to tlie procedures established by CARIPOL, (1980). The

distribution of Dry Tortugas beach tar ranged from 0.6 g m super (-2) to 22.1 g m super (- 

2) dry weight with an average of 9.2 plus or minus 7.8 g tn super (-2). There appear to be 

no strong correlations between Dry Tortugas beach tar concentrations and either 

predominant wind direction or major Gulf Loop Current circulation patterns.

Author Index 

Agassiz, A., 1, 2 

Andres, B., A,, 3 

Applegate, R. D., 298 

Aska, D. Y., 327 

Austin, 0. L. Jr., 4 

Awbrey, F. T., 23 

Baden, D. G., 333,375 

Bailey, E., 5 

Bailey, P. L., 6 

Baker. B.,7 

Ball, S. C., 8 

Ballantine, D. L. 9,403 

Barnes, G. W., 72 

Bartsch, P., 10, 11, 12, 13, 14 

Battey, J. F., 287 

Bellow, T. H., 15, 16,324 

Below, L. C., 315 

Bennett, F. M., 17 

Berrill, N. J., 18 

Blinks, L. R., 19 

Bohnsack, J. A,, 20 

Bortone, S. A,, 21 

Boschma, H., 22 

Bowles, A. E., 23 

Bowman, H. H. M.. 24 

Boyden, A., 25 

Bradbury, R. C., 26 

Braman, R. S., 154 

Brainen, R. S., 356 

Breder, C. M. Jr., 27,28 

Brinley, F. J., 29 

Brooks, H. K., 30 

Brooks, W. K., 31,32 

Brown, D. E. S., 33 

Brown, W.Y.. 34 

Bullington, W. E., 35 

Burkenroad, M., 36 

Caira. J. N., 37 

Calder, D. R.. 38 

Carrier, J. C., 39, 40 

Cary, L. R., 41- 48 

Cate, C. N., 49 

Ceurvels, A. R,. I05 

Chambers, E. L., 50 

Chiang, L., 288 

Child, C. A., 51 

Clapp, R. B., 52 

Clark, H. L., 53 

Clark, L. B., 54 

Cochran, W. W., 298 

Cole, L. J., 55 

Collie, M. R., 56 

Colman, J., 57 

Conger, P. S., 58 

Conklin, E. G., 59, 60 

Coonfield, B. R., 61, 62 

CoutiCre, H., 63 

Cowles, R. P., 64,65 

Crialcs, M. M., 66 

Cubit, 1. D., 196 

Cushman, J. A., 67, 391 

Cutright, P. E., 68 

Dall, W. H., 69 

Darby, H. H., 70-72 

Davis, G. E., 73- 82 

Davis, J. H., 325 

Davis, J. H. Jr., 83, 84 

Davis, R. A. Jr., 85 

de Laubenfels, M. W., 86- 88 

de Renyi, G. S., 89 

Deflaun, M. F., 90 

Dinsmore, J. J., 91, 92, 137 

Dodrill J. W., 8 1 , 82 

Dole, R. B., 93 

Domeier, M. L., 94 

Donaldson, H. H., 95 

Donan, P., 356 

Donnelly, K. B., 366-168,356 

Doyle, M. M., 98 

Doyle, W. L., 96-98 

Drew, G. H., 99 

Dustan, P., 100, 101,163 

Edmondson, C..H., 102 

Erseus, C., 103 

Farfante, I. P., I04 

Feinstein, A. A,, 105 

Feltham C. B., 1 I1 

Fenirnore Johnson, E. R., 72 

Ferguson, John C., 301 

Field, R. M., 106, I07 

Fisk, E. J., I08 

Forcucci, D., 264

Fosberg, F. R., 350 

Gardiner, M. S., 369 

Gault, C. C., 251 

Gee, H., 110, 11 1 

Gersh, 1.. 112 

Gilmore, R. G., 113 

Ginsburg, R. N., I14 

Given, B., 316 

Goldfarb, A. J., 115-122 

Goldman, M. A., 391 

Goodrich, H. B., 123 

Gordon, M., 124 

Goy, J. W., 125 

Grave, C., 126-128 

Gudger, E. W., 129-133 

Haight, F. J., 374 

Halas, J. C., 101 

Halley, R. B., 134,328, 330 

Hanlon, R. T., 135 

Hargitt, C. W., 136 

Harper, D. E., 20 

Hasrington, B. A,, 137 

Harris, J. E 28, 138 

Harrison, C. S., 354 

Hartman, C. G., 139 

Hartmeyer, R., 140, 141 

Harvey, E. N., 142-145 

Hatai, S., 146.148 

Hayes, F. R., 149, 150 

Heard, R. W., 151 

Helwig, E. R., 152 

Hendee, E. C., 153 

Hendrix, S. A. 154 

Hess, W. N., 54, 155, 156 

Hildebrand, S. F., 211,212 

Hine, A. C., 203 

Hixon, R. F., 135 

Hoetjes, P. C., 400 

Hoffman, W., 157.41 1 

Holmes, C. W., 158 

Hooker, D., 159 

Hopkins, D. L., 160 

Howe, M. A., 391 

Hudson, J. H., 310,328,330 

Hutton, R. F., 105 

Jaap, W. C., 101, 161-168, 264,356,403 

Jacobs, M. H., 169 

Jefferson, J. P., 170 

Jehl, J. R., 23 

Jennings, H. S., 171 

Jindrich, V., 172 

Jones, N., 173,285 

Jones, R. D., 174 

Jones, R. S., 113, 175 

Jordan, D. S., 176, 177 

Jordan, H. E., 178-186 

Kaas, P., 187 

Kale, H. W., I88 

Keiller, V. H., 369 

Kellner C., 32, 189 

Kille, F. R., 190 

Kilma, E. F., 261 

Kindinger, J. L., 330 

King, J. Jr., 276,277 

Kojis, B. J., 167, 168, 403 

Kopac, M. J., 191 

Kunkel, B. W., 192 

Lashley, K. S., 398 

Le Compte, M., 193 

Lee, T. N., 66 

Leitch, J. L., 194, 195 

Lessios, H. A,, 196 

Lidz, B. H., 328, 330 

Linton, E., 197- I99 

Lipman, C. B., 200- 202 

Locker, S. D., 203 

Longley, W. H., 204-212 

LuckC, B., 213-215 

Lynts, G. W., 216 

Lyons, W. G., 163,217 

Mann, A., 218,374 

Manter, H. W., 219-221 

Marsh, G., 222- 224 

Martin, D. F., 356 

Martin, 5. C., 346 

Martin, L. K., 40 

Mason, C. R., 316,318 

Mast, S. O., 225 

Matthai, G., 226 

Maul, George A,, 394 

Mayer, A. G., 227-242 

McClellan, D. B., 20 

McClendon, J. F., 243-25 1 

McKenna, I. E. Jr., 167, 168,403

Meeder, 3. F., 252 

Meier, 0 . W., 288 

Meyers, P. A., 401 

Miller. 1-1. M. Jr., 253 

Miller, R. A., 254 

Milligan, M. R., 103 

Millspaugh, C. F., 255 

Mitchell-Tapping, H. J., 256 

Moore, T., 170 

Moritz, C. E., 257 

Mulholland, S., 251 

Multer, H. G., 258, 259 

Mumme, R. L., 415 

Murphy, I,. E., 260 

Nance, James M., 261 

Nicholas, H. M., 421 

Nicoll; P. A,, 128 

O'Neill, C. W., 85,262,263 

Ogden, J. C., 264 

Osburn, R. C. ,265 

Patella, F. J., 261 

Patty, P. C., 157, 355 

Paul, J. R., 281 

Paulson, D. R., 31 8 

Payne, F., 266, 267 

Pearse, A. S., 268-274 

Perkins, H. F., 275 

Perlmutter, D. G., 151 

Pctrovic, C. A., 276, 277 

Phillips, A. H., 278-280 

Pichard, S. L., 281 

Pitts, R. F., 282 

Plan Development Team, Reef Fish 

Management 

Plan, South Atlantic Fishery Management 

Council., 283 

Plantier, T. L., 284 

Plough, H. H., 285 

Porter, J. W.. 264,286-288 

Porter, J. Y., 170 

Potthoff, T., 289 

Powers, P. R. A., 290, 291 

Pratt, H. L. Jr., 39, 40 

Pratt, H. S., 292- 297 

Pritchard, M. H., 37 

Raim, A., 298 

Rebenack P., 21 

Reighard, J., 299,300 

Reynolds, J. E. 111, 301, 302 

Richards, 0 . W., 303, 336 

Richards, W. J., 289 

Richardson, T., 288 

Ricklefs, R. E., 304, 305,405 

Riley, G. A,, 306 

Riska, D. E., 307,308 

Rivas, L. R., 309 

Roberts, H H., 310, 31 1 

Robertson, D. R., 196 

Robertson, W. B. Jr., 4,5, 34,52,92, 157, 312- 

318, 324,355,405,411,413-416 

Robinson, A. H., 319 

Rouse, L. J. Jr., 310, 31 1 

Sarver, S. K., 332 

Schaeffer, A. A,, 320, 321 

Schlumberger, H. G., 215 

Schmahl, G. P., 403 

Schmidt, T. W., 373 

Schmitt, W. L., 322 

Schnell, G. D., 323 

Schreiber, R. W., 324 

Schroeder, P. B,. 325,365 

Scott, W. E. D., 326 

Seaman, W., Jr., 327 

Shimizu, Y., 333 

Shinn, E. A,, 203, 328-330 

Shoemaker, C. R., 331 

Siegel, D. M.., 21 

Silbennan, J. D., 332 

Stnayda, T. J., 333 

Smith, G. I., 287 

Smith, H. B., 254 

Smith, H. G., 334 

Smith, N. P., 264 

Snoek E., 105 

South Florida Area Study 335 

Spence, J., 336 

Sprunt, A. Jr., 337-343 

Steinen, R. P., 134 

Steinmetz, J. C., 302 

Stevenson, I. O., 344 

Steward, F. C., 345,346 

Stockard, C. R., 347-349 

Stoddart, D. R., 350 

Stone, R. G., 351-353

Stoneburner, D. L., 354,355 

Strom, R. N., 356 

Strornsten, F. A,, 357, 358 

Szmant, A. M., 264 

Tandy, G., 359 

Tartar, V., 360 

Tashiro, S., 361 

Taylor, C. V., 371 

Taylor, Jane B., 362 

Taylor, W. R., 363, 364 

Teas, H. I., 365 

Tennent, D. H., 366-371 

Thiele, J., 372 

Thompson, J. C., 177 

Thompson, M. J., 175,373 

Thorp, E. M., 374 

Tomas, C. R., 333,375 

Torrey, H. B., 376 

Treadwell, A. L., 377-379 

Yamanouchi, S., 417 

Yonge, C. M., 418-421 

Zeleny, C., 422, 423 

Zheng, W., 424 

Ubelacker, J. M., 380 

Van Fleet, E. S., 424 

Vaughan, T. W., 374,381-391 

Visscher, J. P., 392 

Vukovich, F. M., 393, 394 

Walker, N. D., 310,31 1 

Wallace, W. S., 395 

Walsh, P. J., 332 

Wartman, W., 396 

Watson, J. B., 397, 398 

Wells, R.r C., 399 

Westinga, E., 400 

Westrum, B.L., 401 

Wheaton, J., 164-168, 402, 403 

Whitaker, D. M., 371,404 

White, S. C. 305, 405, 415 

White-Schuler, S. C., 304 

Wichterman, R., 406, 407 

Williams, 0. L., 408 

Willier, B. H., 409 

Wilson, C. B., 410 

Winegarner, C. E., 15,411 

Wolfe, C. A,, 412 

Woolfenden, G. E., 4, 5,413-416

Subject Index 

Accipiter striatus, 3 

Acropora, 24 1 

Acropora cervicornus, 30,73, 74, 193 

Acropora palmata, 74, 172, 328, 388 

Actinian, 60, 136 

Alcyomeum, 48 

Alcyonaria, 48,407 

Alpheidae, 63,70, 248 

Alpheus, 63,70 

Amphipnds, 151,271,331 

Anchor damage, 73 

Animal succession, 57 

Anisonema vitrea, 102 

Annelids, 119, 120, 228, 351, 352, 353,377, 

378,379,380 

Anous stolidus, 34, 307, 308,405,412 

Ants, 55, 242 

Aplopus, 178, 186,347 

Aplysia protea, 257 

Ascidians, 18, 126, 127, 128, 140, 228, 285, 

303, 

336,376 

Asio flarnmeus 157 

Astroea longispina, 43 

Astropecten duplicatus, 65 

Atlantic palolo (Worm), 227, 228 

Atmospheric compounds, 154,249 

Audouinella ophioglossa, 9 

Audubon, 10 

Avicennia, 83 

Banded red knot, 4 13 

Barnacles, 392 

Bathydrilus oligochaeta, 103 

Beach rock, 106, 107, 1 14,259 

Beach tar, 424 

Belonidae, 27 

Bioluminescence, 144, 145 

Bird history, 338, 339, 343, 344, 397 

Birds,4,5, 10, 17, 108, 137, 157, 316,326, 335, 

337,338,339,341,342,344.413 

Black noddy, 313,318 

Black phoebe, 108 

Black water event, 163, 170 

Blood relationships, 25 

Bluefin tuna, 289 

Botanical ecology, 24 

Brackish-water ponds, 268 

Breeding, 11, 13, 14, 284, 308 

Briareum, 48, 163 

Brown anole, 41 I 

Brown noddy, 34,305,307,308,313,397,398, 

405,412 

Brown pelican, 15, 324 

Bryozoans, 265 

Bubulcus ibis, 415 

Calcium carbonate, 99, 106, 107, 201, 202,246, 

249,259,274,385,391 

Calidris canutus, 413 

Cardinal, 16 

Cardinal cardinalis, 16 

Cassiopea, 44,45,47, 142, 228, 237, 244,245, 

334 

Cassiopea xamachana, 116, 146, 147, 148, 231, 

234,235,238,275,334,349,422 

Cattle egrets, 137, 4 15 

Caulerpa, 41 7 

Cerions, 11, 12, 13, 14 

Chart, 109 

Ciliates, 35,290, 291, 360, 406, 407 

Clibanarius, 192 

Cnideria, 22, 41,42, 46, 48, 59, 98, 135, 146, 

147, 148, 161, 162, 168, 171, 172, 193,228, 

231,234, 235, 237, 241, 244,245, 334,349, 

395,402,403,418,419,422 

Coelenterates (Cnideria), 144, 275 

Color patterns, 61, 62, 123, 130, 204, 205,206, 

207,209,210,299 

Common tern, 304 

Condylactis gigantea, I35 

Conocal-pus, 83 

Coral bleaching, 161,288 

Coral growth, 335, 385, 388 

Coral mortality, 287, 288, 310,31 I 

Coral polyps, 22 

Coral reef, 1,2,57,73, 161, 162, 163, 165, 167, 

168,252, 264, 287,288, 310, 31 1, 359, 381, 

382,383,384,385,386,387,390,403 

Coml reef sampling, 165, 167,288, 359,403 

Coral reef structure, 74,79, 100, 168, 172, 335, 

385,387,389,391 

Corals, 98, 226,229, 24 I , 248,252, 287, 383, 

402,403,418,420 

Crabs, 33,7 1, 192, 204, 257,274,303,422,423 

Crangon armillatus, 62, 70, 71, 89, 156, 257 

Crustaceans, 66, 71,75, 78, 81, 89, 104, 125, 

145, 151, 156, 192, 204, 248, 261,274, 303, 

322,33 1,4 10,422,423 

Cultural resources, 260 

Currents, 384, 389, 393, 394,424 

Cytology, 33,43,50, 59, 87, 96,97, 98, 117,

Diplectrum vittatum, 21 

Diadema, 196 

Diatoms, 58. 218 

Diplectrum formosurn, 21 

Diporia clivosa, 168 

Diving helmet, 13 1 

DNA,90,332 

Dove, 3 15 

Echinaster, 65, 182 

Echinoderms, 53, 118, 121, 122, 149, 150, 153, 

182, 185, 190, 194, 195, 196,247,254,280, 

290,291,360, 366, 368, 369, 370, 371, 376, 

404 

Echinometra lacunter, 53, 150, 194, 195, 254, 

371,376 

Ecology, 83, 84 

Effect of light on organisms, 28,65,72, 155, 

156, 159,222,223,334,346,368 

Effect of photosynthesis. 230,246 

Effects of radiation, 35 I, 352 

Effects of temperature on organisms, 224, 229, 

234,241,244, 287, 310, 31 1, 388, 390, 393, 

394 

Empidonomus varius, 26 

Epinephelus, 282 

Epinephelus morio, 273 

Evolution, 138 

Exococetidae, 27 

Fish, 21.27, 28, 29, 33, 36, 39, 61, 68, 94, 112. 

113, 123, 124, 129, 130, 132, 133, 135, 138, 

139, 170, 176, 177,205,206,207,209,210, 

211, 212, 213,214, 215, 221, 273,282,283, 

289,299,309,373,408,410 

Fish communities, 175, 212, 283, 373 

Fish kill, 310 

Fish sampling techniques, 175,205 

Fish tumors, 213, 214, 215 

Flight speed (Birds), 323 

Florida Keys National Marine Sanctuary, 7.20 

Food habits, 21.92, 171, 205,397, 421 

Foraminiferae, 67, 216 

Frigate birds, 323 

Geologic history, 381, 382, 383 

Geology, 30,83, 106, 107, 114, 134, 172,203, 

256,258,259, 260, 262, 263,302,328, 329, 

374,381,382,383,385,391 

Ginglymostoma cirratum, 39, 112, 129, 133 

Gorgonacae, 41,42,46,402 

Gorgonia, 48,402 

Great black-backed gull, 188 

Growth, 303,304,305,346, 348,405, 

412 

Habits, 64, 132, 133, 248, 272, 273, 

275,347,398,418,419,421 

Haemulidae, 36 

Halichores, 213 

Halichores bivittatus, 124 

Halimeda, 263 

Hali~neda hummii, 9 

Hatching success, 23 

Hemirhamphidae, 27 

Histology, 190 

Holothuria, 190, 279, 280, 369 

Homing instinct, 10, 398 

House sparrow, 416 

Hydroids, 38, 116,376,395 

Hydrology, 134 

Hypoglossum rhizophorum, 9 

Hypoplectrus, 94, 373 

Insects, 8, 347 

Invertebrates, 6, 8, l I, 12, 13, 14, l8,22,25, 32, 

35, 35, 38,41,42,43,44,45,46,47,48,49, 

50, 51, 53,54,55,58,59,60,62, 63, 64. 65, 

66,67, 69,70,71,73,74,75,76,77,78, 80, 

81, 82, 86, 87, 88, 89,98, 100, 101, 102, 103, 

104, 117, 118, 119, 120, 121, 122, 125, 126, 

127, 128, 135, 136, 140, 142, 143, 144, 145, 

146, 147, 148, 149, 151, 152,.153, 155, 156, 

161, 162, 169, 171, 172, 178, 182, 183, 185, 

I86,187,189,l90,192,193,195,196,197, 

198, 199, 217,219,220,221, 227,229, 231, 

235,238,239,242,244,245,246,247,248, 

253,254,261,265,266,267,270,271,272, 

275, 279, 280,285, 290,291,293, 294,295, 

296,297, 303,322, 331,334,347, 35 1, 352, 

353,360, 366, 367,368,369, 376, 377, 378, 

379,380, 392, 395,400,402,408,4I8,419, 

421,422 

Iotrochota, 87, 152 

Ischnochiton, 187, 217 

Gastropods, 89

Larval shrin~p, 66 

Launcularia, 83 

Least terns, 414 

Leodice fucata, 54 

Limulus, 184, 228 

Linckia, 360 

Lipogramrna anabantoides, 113 

Littoral, 269, 272 

Loggerhead turtle, 159, 179, 180, 181, 183, 198, 

295,297,357,358,409 

Long Term Ecological Research (LTER) 163, 

167,225,264 

Longevity, 34 

Lutjanus griseus, 124, 214 

Lytechinus variegatus, 50, 153,247,368, 370, 

404 

Maeandra aerolata, 22, 229,241,418 

Maps, 1,74 

Manatees, 301 

Mangrove, 83,325, 365 

Marinc algae, 9, 19,96, 97, 191, 222, 223,224, 

345,346,363,364,417 

Marme amoebe, 160,320,321 

Marine bacteria, 99, 110, 11 1, 202 

Marine fishery reserves (MFR), 283 

Marinc Laboratory, 232,236, 240 

Marine parks, 79,319 

Masked booby, 52 

Mastogloia, 218 

Mating, 39 

Medusae, 44,45,228, 231, 234, 239, 275,349 

Merlin, 298 

Metals in organisms, 278, 279, 280, 354, 355, 

356 

Migration, 17 

Millepora complanata, 161, 172 

Mites, 274 

Mollusks, 11, 12, 13, 14,49,69, 89, 187, 217, 

253,372,392,421 

Monocotyle, 292 

Monosporus indicus, 9 

Montastrea annularis, 162, 168, 172, 241, 385 

Montastrea cavernosa, 168 

Morphodynamics, 85, 192 

Mosquitoes, 8 

Myliobatis, 292 

Nematodes, 408 

Neoplaslic growths, 124 

Nesting, 15, 324, 343, 397,414,416 

Nurse shark, 39,40 112, 129, 133, 197 

Ocypoda, 64 

Oikopleura, 189 

Oil spill, 335 

Ophicoma, 182 

Orbicella (Madrepora), 193,229, 24 1, 388 

Osmotic pressure, 269 

Owl, 157 

Panulirus argus, 75,77,78, 89, 156,327, 332 

Parasitic copepods, 4 10 

Parasitic isopods, 273,410 

Parasitic worms, 37, 197, 198, 199, 219, 220, 

221,253,292,293,294,295,296,297 

Pedibothrium, 37 

Penaeus duorarum, 66,261 

Pesticides, 356 

PH, 230,250,251,334 

Physiology, 47,97, 111, 135, 160, 169,334, 

345,346,357,358 

Phytoplankton, 333, 375 

Plankton, 306 

Plectognath, 28 

Plexaura, 46 

Pollution, 335, 424 

Polycistor, 140 

Polyplacophora, 217 

Pomacentrus, 29 

Pomacentrus leucostictus, 61 

Porites, 229, 241 

Porites asteroides, 168 

Portunus sayi, 422,423 

Promicrops (Epinephelus) itajara, 273 

Protozoan, 102, 160, 169,406,407 

Pseudocyphoma, 49 

Ptychodera, 43, 89, 155,266, 267 

Pycnogonida, 51 

Rails, 277 

Rat, 95 

Recreational headboat fishery, 20 

Red-headed woodpecker, 3 17 

Red tide, 105 

Reef fishes, 20 

Reef formation, 2,41,42 

Reefs, 30 

Regeneration, 6,44,45,47, 1 15, 1 16, 120, 152, 

190, 235,285, 348,349,351, 352, 353, 360, 

422,423 

Replenishment reserves, 7,20,283 

Rhizophora, 83 

Richardina spinicincta, 125 

Roseate terns, 3 13,4 14

Sabellids, 6, 352, 379 

Sabines's gull, 56 

Salinity. 93, 116 

Salpa floridian, 31 

Scleractinia, 161 

Scyphomedusa ijellyfish), 59,234 

Sea-birds, 4, 15, 23, 34, 52, 56, 91, 92, 188, 

284, 

298, 304,305,307, 308, 312, 313,318,323, 

324,140,341,354,355,397,398,405,412, 

414 

Sea-cucumber, 190 

Sea-level change, 203, 258, 262, 263 

Sea turtles, 159 

Sea-urchin, 50, 117, 121, 122, 150, 153, 

194, 195, 196, 247, 254,290,291,366, 

368,370,37 1,404 

Sea-watcr composition, 72, 90, 93, 115, 

174, 200, 201,202,230,235, 237, 238, 

243,246,249,250, 251,281, 320, 361, 

389,399,401 

SEAKEYS, 264 

Sedimentation, 85, 172, 256, 258, 259, 

262,263,329,374 

Sharks, 39, 139 

Sharp-shinned hawks, 3 

Sicyonia penaeoidea, 104 

Siderastera, 168,229,241, 385,419 

Snapping shrimps, 63, 322 

Sooty terns, 4,5,23,91,92, 284, 304, 

312,313.314,354,355,397,398,405 

Speculata advena, 49 

Spheciospongia vesparia, 151, 400 

Sphyraena barracuda, 132 

Spiny lobster, 75,76,77,78, 80, 81, 82, 

156,327,332 

Sponges, 86, 87, 88, 101, 151, 152,270,400, 

403 

Sport harvest, 75,76, 78, 79, 81, 82 

Stal.fish, 65, 360, 369 

Starvation, 231, 334,415 

Stephanochasmus, 294 

Sterna f~lscata, 4, 5, 91, 92, 284, 304, 314, 354, 

405 

Sterna hirundo, 304 

Sterna ~nclanoptera, 340 

Sting ray, 68 

Storms, 17,405 

Sula daccylata, 52 

Syllids, 380 

Synentogiiathi, 27 

Topography, 1,84,325,328,329,343,350 

Transatlantic migration, 314 

Trans~lantinrr orzanisms. 403 

Trichechus, 301 

Tunicata, 3 1, 32 

Valonia, 19,96,97, 19 1,222, 223,224, 345, 

346 

Vanadium, 279,280 

Variegated flycatcher, 26 

Vegetation, 24, 84,255, 325, 350, 365 

Vessel groundings, 335,403 

Vocal signals, 307, 308 

Worms, 6,54, 103, 119, 120, 155, 197, 198, 

220,253,292,293,294,295,296,297, 352, 

353,377,378,379,380,408 

Zooxanthellae, 98, 334,407,421

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