AMER. ZOOL., 20:525-530 (1980) The "Lost Year" Question in Young Sea Turtles1 ROSS WlTHAM Florida Department of Natural Resources Marine Research Laboratory, 100 Eighth Avenue SE, St. Petersburg, Florida 33101 SYNOPSIS. An adequate understanding of young sea turtle dispersal patterns is necessary for effective management of threatened or endangered species. Such patterns are poorly understood, and the term "lost year" has been adopted to emphasize this gap in sea turtle life history information. Tag returns from pen-reared yearling sea turtles indicate ocean current dispersal. Evidence indicates hatchlings would be similarly dispersed by ocean currents. Feeding studies with tank-held animals suggest that food resources are available in ocean currents for long-term sea turtle survival. Green turtle (Chelonia mydas) growth appears slow in nature. INTRODUCTION 681). After reaching a weight between 200 The posthatchling period, or "lost year" and 900 g, turtles are released on beaches of sea turtle life history, after departure during daylight. Observations are made from natal beaches, is poorly understood. during turtle movement to the sea and for The turtles' coloration, small size, and agil- as long as possible after they start their offity make oceanic observations difficult. shore swim. Data acquisition during this extended RESULTS posthatchling period is further impeded Tag returns from pen-reared, yearling by lack of suitable, durable tags for hatchlings. However, in our continuing Jensen green turtle releases made in the 1960s inBeach study, tag returns from yearling dicated some long-term survival and green (Chelonia mydas Linne) and logger- movement (Witham and Carr, 1968; Carr head (Caretta caretta Linne) turtles are pro- and Sweat, 1969). More than 10,000 turviding data on oceanic survival, growth, tles have been released since large-scale efforts were begun in 1972; more than 90 and dispersal. tag returns have been received. Although some returns have been short-term recapMATERIALS AND METHODS Turtle eggs are collected from natural tures near release beaches, most have been nests on Florida's east coast, then incubat- long-term, long-distance returns (Table 1, ed and hatched in the laboratory. Hatch- Fig. 1). These data indicate slow oceanic lings are reared in facilities previously de- growth (Fig. 2). scribed by Witham (1976) and Witham and DISCUSSION Futch (1977). Additional facilities are also Numerous authors have suggested that provided at Sea World, Orlando (closed, artificial sea water system) and at a private ocean currents influence sea turtle disestate at Hillsboro Beach (open sea water persal. Deraniyagala (1939), in his zoogeographic discussion of testudinates, mensystem). Prior to release, all turtles are weighed tioned the seasonal clockwise and and measured, then tagged with self pierc- counterclockwise currents around Ceylon ing monel tags (National Band and Tag (Sri Lanka). Daniel and Smith (1947) comCompany,2 Style No. 4-1005, Size No. mented that for hatchling loggerheads, ". . . the current sense . . . and direction of movement . . . may be influenced by the 1 From the Symposium on Behavioral and Reproduc- water currents of the surf, water pressure tive Biology of Sea Turtles presented at the Annual factors, or other physical circumstances Meeting of the American Society of Zoologists, 27- . . . ." Carr and Ogren (1960) mentioned, 30 December 1979, at Tampa, Florida. "current sense" and "passive drifting" for 2 Use of product name does not indicate endorse- Caribbean green turtles. Little is known ment by the Florida Department of Natural Reabout the local currents which might affect sources Marine Research Laboratory. 525 526 ROSS WlTHAM AZORES VIRGINIA;CT| N. CAROLINA" S. CAROLINA ATLANTIC OCEAN FIG. 1. Capture locations of pen-reared green turtles, Chelonia mydas, released by the Florida Department of Natural Resources. Numbers refer to the number of turtles captured in each area. the Tortuguero nesting colony (Carr et ai, 1978). Hughes (1974), Hughes and Richard (1974), Balazs (1976), and Pritchard (1976) all discussed ocean current transport. Witham (1976) argued strongly for ocean current transport of young sea turtles and suggested that such currents might also be important to adult sea turtle dispersal. Observations of thousands of yearling sea turtles, released on beaches and allowed to crawl to the sea, indicate that turtles orient themselves toward light until they reach the surf (Daniel and Smith, 1947; Ehrenfeld, 1968; Mrosovsky, 1972). Upon reaching surf with sufficient wave action, turtles unhesitatingly swim seaward. When waves are very small or absent, the turtles hesitate, and swimming appears non-directed. A wave orientation response has also been observed by students from the Florida Institute of Technology (FIT) (personal communication). Frick (1976) reported that hatchling green turtles swam at an average 1.57 km/ hr when moving offshore. Prang (1976), while studying swimming turtle energetics, X" " x*x 12 14 16 18 WEIGHT (Kg) 20 22 24 a6 28 30 FIG. 2. Wild growth (weight) of 31 pen-reared green turtle yearlings released by the Florida Department of Natural Resources. 527 YOUNG SEA TURTLE SURVIVAL, GROWTH, DISPERSAL TABLE 1. Released and captured data for pen-reared, yearling turtles tagged and released on Florida beaches.* Released Tag no A451 A498 A681 A1037 A1195 A1216 A1319 A1395 A1573 A1631 A1672 A1698 A1993 A1999 A2155 A2250 A2316 A2480 A2575 A3147 A3291 A3554 A3566 A3626 A3643 A3667 A3756 A3770 A3798 A3903 A3903 A3903 A3909 A3915 A3939 A4045 A4049 A4125 A4125 A4160 A4591 A4621 A4681 A4958 A5053 A5485 A5664 A5719 A5856 A5973 A6083 A6097 A6110 A6146 A6232 A6281 A6316 A6340 A7020 Date 31 May 72 9Jun72 4 Apr 72 19Sep72 24 Apr 73 25 Apr 73 16 May 73 16 May 73 17 May 73 23 May 73 6Jun73 6Jun73 18 Mar 74 18 Mar 74 27 Mar 74 27 Mar 74 27 Mar 74 17 Apr 74 29 May 74 13 Mar 75 21 Mar 75 2 Apr 75 2 Apr 75 16 Apr 75 16 Apr 75 16 Apr 75 16 Apr 75 16 Apr 75 16 Apr 75 25 Apr 75 9 Feb 76 23 May 76 25 Apr 75 25 Apr 75 25 Apr 75 2 May 75 2 May 75 2 May 75 7 Oct 77 2 May 75 10Jun75 29 Jul 75 29Jul75 21 Jan 76 21 Jan 76 26 Mar 76 31 Mar 76 31 Mar 76 14 Apr 76 14 Apr 76 3Jun76 3Jun76 3Jun76 3Jun76 25Jul76 25 Jul 76 25 Jul 76 25 Jul 76 7 Apr 77 Capture N Lat W. Long. 27°15' 27°15' 26°30' 25°40' 27°10' 27°10' 25°40' 25°40' 26°30' 27°15' 28°30' 28°30' 26°30' 26°30' 25°40' 25°40' 25°40' 27°50' 28°30' 27°15' 26°30' 27°15' 27°15' 25°40' 25°40' 25°40' 25°40' 25"40' 25°40' 26°05' 17°30' 17°30' 26°05' 26°05' 26°05' 27°50' 27°50' 27°50' 34°45' 27°50' 27°15' 27°05' 27°05' 25°40' 25°40' 26°30' 27°15' 27°15' 27°15' 27°15' 27°05' 27°05' 27°05' 27°05' 27°15' 27°15' 27°15' 27°15' 25°40' 80° 10' 80° 10' 80°05' 80° 10' 80° 10' 80°10' 80°10' 80°10' 80°05' 80° 10' 80°35' 80 o 35' 80°05' 80°05' 80°10' 80°10' 80°10' 80°25' 80-35' 80°10' 80°05' 80°10' 80° 10' 80°10' 80°10' 80°10' 80-10' 80°10' 80°10' 80-05' 88°10' 88°10' 80°05' 80°05' 80°05' 80°25' 80°25' 80°25' 76°25' 80-25' 80°10' 80°10' 80-10' 80°10' 80°10' 80-05' 80°10' 80°10' 80°10' 80°10' 80°10' 80°10' 80-10' 80-10' 80°10' 80°10' 80°10' 80°10' 80-10' Date Sep 76 May 79 25 Sep 78 26 Oct 72 7 Feb 76 15 Apr 76 19Nov75 25 May 73 23 Mar 78 24 Jan 77 23 Jul 77 30 Oct 76 23 Jan 78 20 May 76 20 Jan 77 10 Jan 75 28 Mar 74 15 Jan 77 15 Feb 78 28 Aug77 27 Apr 76 2 Apr 77 11 Nov 76 5 May 75 19 Dec 75 22 Apr 75 7 May 75 20 Jan 76 10 Oct 76 9 Feb 76 23 May 76 15 Jul 76 22 Nov 76 27 Dec 76 28 Jan 77 3Jun77 15 Sep 76 7 Oct 77 28 Nov 77 24Jul77 20 Jan 77 9 Nov 77 7Jun77 8 Feb 76 26 Jan 76 6Jun79 28 Dec 77 11 Oct 77 8 Dec 78 5 Apr 79 12Jul78 12 May 77 24 Feb 78 15 May 79 Aug 78 6 Feb 77 10 Nov 77 26 Aug 78 11 May 77 N Lat. W. Long. 32°40' 32°45' 34°45' 25°10' 26°35' 22°50' 25-10' 25°40' 21°40' 28°45' 25-35' 26°35' 27°35' 32°40' 28-40' 10-30' 25-40' 18°15' 26°30' 33-50' 16°00' 27°40' 35°10' 25-40' 22°50' 25-40' 26°25' 25°05' 35°40' 17-30' 17-30' 17°30' 21°40' 17°30' 24°55' 17°05' 37-00' 34°45' 29°55' 27°20' 27O55' 33°50' 29°55' 27-15' 25°10' 26°30' 28°40' 35°05' 26°30' 27°40' 27°45' 19-35' 18-10' 13-10' 24°50' 39°25' 25°00' 19°35' 26-35' 79°50' 16-45' 76-25' 80°30' 77-50' 74°25' 80°30' 80°10' 71°45' 80°55' 76°40' 77°50' 80°35' 79°50' 80°45' 75-40' 80°10' 87°55' 77°05' 78°10' 86°00' 80°20' 75°45' 80-10' 74°25' 80-10' 80-05' 80°25' 75°10' 88°10' 88-10' 88°10' 71°20' 88°10' 80°40' 62°35' 76-05' 76°25' 81°20' 80°25' 82°50' 78°00' 81-20' 80°10' 80°25' 78°35' 80°45' 75°55' 78°35' 80°20' 80°25' 69°00' 87°55' 59°30' 76°20' 31-05' 77°55' 69°00' 80-05' Location S. Carolina Ilha da Madeira N. Carolina Turkey Pt. Bahamas Bahamas S. Biscayne Bay Virginia Key Middle Caicos Titusville area Bahamas Bahamas Vero Beach S. Carolina Mosquito Lagoon Colombia, S.A. Miami Beach Mexico Bahamas N. Carolina Honduras Vero Beach N. Carolina Miami Beach Bahamas Miami Beach Boca Raton Key Largo N. Carolina Belize, C.A. Belize, C.A. Belize, C.A. Turks-Caicos Belize, C.A. Long Key Nevis, W.I. Virginia N. Carolina St. Augustine Satellite Beach Clearwater N. Carolina St. Augustine Hutchinson Isl. Key Largo Bahamas Mosquito Lagoon N. Carolina Bahamas Vero Beach Sebastian Dominican Rep. Mexico Barbados Bahamas Azores Bahamas Dominican Rep. Lake Worth Beach ROSS WlTHAM 528 TABLE 1. Continued. Released Tag no A7027 A7045 A7054 A7157 A7182 A7206 A7257 A7485 A9062 A9222 A9249 Date 7 Apr 77 7 Apr'77 7 Apr '77 7 Apr 77 19 Apr 77 16 Apr 77 18 Apr 77 19 May 77 29 Aug 78 6 Feb 79 6 Feb 79 N. Lat 25°40' 25°40' 25°40' 25°40' 26°30' 26°05' 26°30' 26°50' 27°15' 27°15' 27°15' iCapture W. Long 80° 10' 80° 10' 80" 10' 80° 10' 80°05' 80°05' 80°05' 80°05' 80° 10' 80° 10' 80° 10' Date 21 Apr 77 25 Apr 77 12Jun'77 13 May 77 19 Apr 77 28 Mar 79 13 Jan 79 23 May 77 1 Sep 78 19 Mar 79 25 Feb 79 N. Lat. W. Long. Location 25°30' 25°40' 25° 10' 26°50' 26°30' 17°2O' 35°10' 27°15' 27°15' 28°25' 26°30' 80°20' 80° 10' 80°25' 80°05' 80°05' 62°45' 75°45' 80°10' 80° 10' 80°30' 80°05' Biscayne Bay Bal Harbor Key Largo Juno Beach Delray Beach St. Kitts, W.I. N. Carolina Port Salerno St. Lucie Inlet Coca Beach Delray Beach * No. A498 was loggerhead, Caretta caretta. The others were greens, Chelonia mydas. Witham and Futch (1977) reported 22 such returns for greens and their data are not repeated here. found yearling green turtles did not swim when currents were 0.14 m/sec (0.50 km/ hr) or less, and they became exhausted at current speeds of 0.35 m/sec (1.26 km/hr). An FIT report (unpublished) gave an average of 0.5 m/sec (1.8 km/hr) for yearling green turtles swimming through surf. Frick (1976), based on her observations of the "swim frenzy" of hatchlings, suggested they may swim steadily for 24 hr. This might result in hatchlings being further offshore than yearlings would be after 24 hr. Hughes (1974) estimated that hatchling sea turtles, entering the sea at a Caribbean site, would require 4 to 7 yr to circumnavigate the North Atlantic and return to their natal beach. Our data (Witham, 1976; Witham and Futch, 1977) indicate that such a circumnavigation could take less than 1 year in the North Atlantic gyre, which includes the Florida Current and Gulf Stream. The Florida Current, a northerly flowing current near the Florida east coast, reaches its maximum speed of about 20 km offshore of Dade County (Brooks and Niiler, 1977). The research submersible, Ben Franklin, drifted more than 2,596 km in 30 days while submerged in the Florida Current and Gulf Stream (Gatskell, 1973). The swimming behavior of young sea turtles makes them comparable to vessels. Therefore the long-range, rapid dispersal of hatchling turtles would occur when they entered these major currents; off some Florida east coast beaches, turtles would enter the current almost immediately. For example, hatchling turtles swimming from a Dade County beach would be deflected northward by the nearshore Florida Current (Dunlap and Shufeldt, 1969), and after 24 hr the turtles would probably be about 130 km north of the starting point (Fig. 3). Their longrange, rapid dispersal would continue until they left the major current. Dispersal patterns for other beaches would be influenced by local currents and the Dade County model should be modified to meet local conditions. It has been assumed that wild yearling green turtles weigh from 1 to 4 kg (Hirth, 1971), but no data from wild populations are available to support this assumption. Caldwell (1962) suggested that green turtles would require 8 (tropical waters) to 13 yr (temperate waters) to reach maturity. Our data (Fig. 2) indicate that pen-reared, yearling green turtles grow slowly after their release. These data suggest that 15 to 20 yr might be required for Western Atlantic green turtles to reach maturity. Balazs (1979) and Limpus (1979) estimated that Hawaiian and Australian green turtles might require as long as 50 yr to reach maturity. Captive feeding studies (Witham, 1977; Witham and Futch, 1977) have shown that green (Chelonia mydas), loggerhead (Caretta caretta), and leatherback (Dermochelys coriacea Linne) turtles grow well on a diet of one species of jellyfish, Cassiopea xama- YOUNG SEA TURTLE SURVIVAL, GROWTH, DISPERSAL 110- E CO o POSITION OF TURTLE AFTER EACH HOUR - ASSUMING NO CURRENT 40ESTIMATED POSITION OF TURTLE AFTER EACH HOUR 529 Chesnut (1974) reported on the ctenophores, jellyfish, and Portuguese man-ofwar in coastal and neritic waters of North Carolina. One widely distributed species of jellyfish, Cyanea capillata, reaches a bell diameter of 2 m with tentacles 36 m long (Halstead and Courville, 1965); one such jellyfish might supply food for many small turtles. Hirth (1971) reported that at an indeterminate age green turtles become primarily herbivorous, but adults continue to eat jellyfish. The diverse fauna associated with drifting sargassum (Weis, 1968; Fine, 1970; Ryland, 1974) might also provide supplemental food for turtles at sea. Young sea turtles would probably find adequate food resources for survival in ocean currents. Our data strongly suggest that the initial posthatchling period, "the lost year," is a period of oceanic existence, when turtles opportunistically use oceanic currents and food resources for dispersal and survival. ACKNOWLEDGMENTS 10- Mt, 50 60 OFFSHORE DISTANCE (Km) 70 Fic. 3. Hypothetical positions of a hatchling green turtle during 24 hours of easterly swimming from a Dade County, Florida, beach. chana Linne. The widespread oceanic distribution of cnidarians has been reported. Mayer (1910) commented, . . . I have frequently observed that they [cnidarians] are more abundant near the surface when the ocean has been perfectly calm for a few days than under ordinary conditions; for if they are undisturbed by the movement of waves, some of these forms, in common with many of the rarely seen ctenophorae and siphonophorae, will gradually rise until they become in great numbers over the unrippled surface. Piccard (1971), reporting on the drift cruise of the research submersible, Ben Franklin, observed salpas and jellyfish throughout the trip. Kennedy (1972) discussed the widespread surface distribution of Portuguese man-of-war (Physalia) in the Caribbean, Gulf of Mexico, and the Florida Current (Gulf Stream). Schwartz and This research has enjoyed long-term support from many organizations and individuals. To name them all would require space equal to a large portion of the paper. Their assistance has had many forms, including administrative and supervisory support, editorial assistance, egg collection, pen-rearing, and providing tag return data. REFERENCES Balazs, G. H. 1976. Green turtle migrations in the Hawaiian Archipelago. Biol. Conserv. 9:125— 140. Balazs, G. H. 1979. Growth, food sources and migrations of immature Hawaiian Chelonia. Marine Turtle Newsletter IUCN/SSC No. 10:1-3. Brooks, I. H. and P. O. Niiler. 1977. Energetics of the Florida Current. J. Mar. Res. 35(1): 163-191. Caldwell, D. K. 1962. Growth measurements of young captive Atlantic sea turtles in temperate waters. Los Ang. Cty. Mus. Contrib. Sci. 50. Carr, A., M. H. Carr, and A. B. Meylan. 1978. The ecology and migrations of a sea turtle colony. Bull. Am. Mus. Nat. Hist. 162(1). Carr, A. and L. Ogren. 1960. 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