Marine Turtle Newsletter Issue Number 151 October 2016

Leatherback hatchlings emerging from the wooden incubation device after 60 days. See pages 6-8. Photo by: Grande Riviere Nature Tour Guide Association, GRNTGA

Editorial Fibropapillomatosis Tumors at Honokowai: Underwater Observations with Potential Broad Application...... P Bennett & U Keuper-Bennett

Articles Artificial Incubation Trials of Leatherback Eggs at Grande Riviere Beach, Trinidad, West Indies (Phase II)...... R Shoy The effect of invertebrate infestation and its correlation with loggerhead sea turtle (Caretta caretta) nest success in Laganas Bay, Zakynthos, Greece...... AJ Andrews et al. Assessing the Impacts of Hatcheries on Green Turtle Hatchlings...... CM Balsalobre & I Bride Evidence of Turtle Poaching On Agalega, Mauritius...... I Webster et al. Hatching of Eggs Rescued from a Green Turtle Involved in an Automobile Collision...... I Kawazu et al. Northernmost Records of Hawksbill Turtle Nests and Possible Trans-Atlantic Colonization Event...... SA Finn et al.

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Marine Turtle Newsletter No. 151, 2016 - Page 1 ISSN 0839-7708 Editors: Managing Editor: Kelly R. Stewart Matthew H. Godfrey Michael S. Coyne The Ocean Foundation NC Sea Turtle Project SEATURTLE.ORG c/o Marine Mammal and Turtle Division NC Wildlife Resources Commission 1 Southampton Place Southwest Fisheries Science Center, NOAA-NMFS 1507 Ann St. Durham, NC 27705, USA 8901 La Jolla Shores Dr. Beaufort, NC 28516 USA E-mail: [email protected] La Jolla, California 92037 USA E-mail: [email protected] Fax: +1 919 684-8741 E-mail: [email protected] Fax: +1 858-546-7003

On-line Assistant: ALan F. Rees University of Exeter in Cornwall, UK

Editorial Board:

Brendan J. Godley & Annette C. Broderick (Editors Emeriti) Nicolas J. Pilcher University of Exeter in Cornwall, UK Marine Research Foundation, Malaysia

George H. Balazs ALan F. Rees National Marine Fisheries Service, Hawaii, USA University of Exeter in Cornwall, UK

Alan B. Bolten Kartik Shanker University of Florida, USA Indian Institute of Science, Bangalore, India

Robert P. van Dam Manjula Tiwari Chelonia, Inc. Puerto Rico, USA National Marine Fisheries Service, La Jolla, USA

Angela Formia Oğuz Türkozan University of Florence, Italy Adnan Menderes University, Turkey

Colin Limpus Jeanette Wyneken Queensland Turtle Research Project, Australia Florida Atlantic University, USA

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Marine© Turtle Marine Newsletter Turtle No. Newsletter 151, 2016 - Page 1 Editorial: Fibropapillomatosis Tumors at Honokowai: Underwater Observations with Potential Broad Application

Peter Bennett & Ursula Keuper-Bennett 3-24 Reid Drive, Mississauga, ON L5M 2A6 Canada (E-mail: honu@turtles.org, [email protected])

Forward. Decades of research and dozens of science publications turtle population. Such worries, while fully justified at the time, have have provided useful insights into the tumor-forming disease failed to come true and likely never will. Indeed, two of the world’s known as fibropapillomatosis (FP) affecting primarily green turtles, most historically FP-afflicted green turtle populations (Hawaii and Chelonia mydas. Yet the cause, cure, and manner of transmission Florida) have shown phenomenal long-term growth to the present of this affliction remain substantially unresolved. The leading day. This fact lends credibility to the Bennett’s observations and etiological candidate continues to be an alpha-herpes virus, parsimonious reasoning. That is, that the spread of FP is exacerbated but how and where is it spread, and under what environmental in dense green turtle populations, and deterred by lower densities. conditions? The answers to date remain inconclusive and Thus, while some of the material in this essay has become outdated elusive. But might important information available in plain view with progress in the research concerning fibropapillomatosis, and on the world wide web for the past 20 years have gone unnoticed or also with our own continuing education about the disease, this essay ignored? On 17 December 1995 Canadian naturalists Peter Bennett remains valuable. and Ursula Keuper-Bennett composed the following essay based on My congratulations to the Marine Turtle Newsletter for ensuring a 8 summers of meticulous underwater observations of green turtles place in the published literature so the Bennett’s essay can seriously off West Maui in the Hawaiian Islands. Their insightful thought- be taken into account by FP historians and researchers of the future. provoking article was prominently posted where it remains today The original essay can be found here: on one of the world’s first web sites devoted exclusively to sea www.turtles.org/tumoursa.htm turtles, Turtle Trax (www.turtles.org). When the essay was written George Balazs - 992 Awaawaanoa Place Honolulu, Hawaii 96825 FP seemed to signal the eventual collapse of the Hawaiian green USA (E-mail: [email protected])

“Pollution, right?” That is the first reaction of nearly everyone who We have therefore spent considerable time trying to make sees our images of tumored turtles. If only it were that simple. sense of what we have seen and learned. From our observations No one really knows what causes turtle tumors. Although and the inquiries they prompted, we think we have a plausible scientists have made progress towards understanding explanation of the dynamics of the disease and its transmission. fibropapillomatosis (FP), the cause is still uncertain. Until someone Certainly it explains our observations and many of the points finds the cause, no one can know how tumors spread. This makes raised by researchers in the papers we’ve studied. it difficult - but not impossible - to know what to do about the We have no doubt that our explanation is incomplete and disease. probably wrong in some areas. Nevertheless, it does fit the facts We reached this conclusion after reading many scientific papers, we have gathered from various papers, as well as our personal consulting Internet sources, and soliciting information through observations underwater. We believe that if we are correct, we can email from marine turtle specialists, virologists, government identify potential hot-spots, where the incidence of tumors will be officials, and fellow divers. exceptionally high (>50%). Note that we did not say fellow scientists. Scientists we are not. In this essay, we show how our thinking arose from our We are turtle-watchers. We have eight summers of observations observations and inquiries about our Honokowai dive site, and in the waters of Honokowai, West Maui, Hawaii, USA. Over that explain why we believe it can explain aspects of the development of time, we have done nearly 1000 dives averaging about 75 minutes the FP tumors worldwide. each, with 35-40 of those minutes spent actually watching turtles. Some premises: We have made over 90 hours of videotape and taken over Seaweeds, like any algae, respond to nutrients (Brodie 1995). If 2500 photographs. We have paid close attention to watching and you allow extra nutrients into a shallow coastal area, one where documenting the changes in the environment. We have developed they will not be quickly diluted and carried off, you will get a reliable method of identifying individual turtles (>100 so far) increased seaweed growth. allowing us to track the changes in their condition from year to Sea turtles, like other animals, tend to cluster at places where food year. is most plentiful. Green sea turtles feed primarily on various Why we wrote this. Since we first discovered the turtles of seaweeds (Balazs 1980). It seems likely that if you find an area Honokowai and their tumors, we have frequently been asked of abundant seaweed, you will find more turtles than you would to explain the problem. When we tell people that science is still in areas with less seaweed. s e a r c h i n g for the answers, we are next pressed for our opinion. Because we now maintain Turtle Trax, a webpage dedicated to When the turtle population density increases, there is more contact spreading information about marine turtles, we felt a need to make between individual turtles, and each turtle has contact with a sure that whatever opinions we expressed were solidly based. larger number of other turtles. This provides more opportunities Marine Turtle Newsletter No. 151, 2016 - Page 1 Figure 1. Turtles Clustering: This picture shows the end of Hilu’s return from a trip to the surface for air. He could have picked an empty spot anywhere in the area, but he chose instead to land literally on top of two other turtles. This emphasizes the affinity turtles have for the company of other turtles, and the resulting close contact. It is this social nature that we believe ultimately leads to a much higher potential for the spread of FP tumors.

for the spread of communicable disease (Herbst 1995).FP tumors erosion control. At roughly the same distance to the south, there are almost certainly spread by a virus. Although the virus has not is the Honokowai Stream Channel, also concrete. Runoff from yet been isolated, FP is virus-induced in other animals (Herbst pineapple and sugar cane fields collects in these channels and et al. 1995). Herbst (1995) and others have strongly implicated a until recently, was discharged right into the ocean. viral cause and are working towards conclusive proof. Just over 1 km south and about 500 m from the shoreline, the Viruses can be spread through mechanical vectors (P. Young, pers. Lahaina Wastewater Reclamation Facility uses injection wells to comm.). When turtles gather in sufficient numbers, some fish dispose of sewage effluent. Into holes ranging from 55-70 m deep, will become cleaners (Losey et al. 1994). Each one is a potential contained only by concrete casings extending approximately vector (Losey et al. 1994). 45% of their depth, Maui County pumps effluent at a rate of 15-25 million liters/day. Maui, of course, is a volcanic island, with plenty Below we examine how these points apply to the turtles of of porous rock and lava tubes. Where has this effluent been going? Honokowai. No one knows, but it seems extremely unlikely that none of it ever Nutrient input. Around 500 m to the north of where we watch these reached the ocean. turtles, there is the Mahinahina Channel, part of the Honoloua The waters of West Maui near Honokowai are shallow and Watershed Project. This is a concrete channel built for flood and do not appear to lend themselves readily to self-cleansing. When

Figure 2. Suspected Mechanical Vectors: The cleaners in this image, goldring surgeons (Ctenochaetus strigosus), are clearly grazing on the tumors. Much more likely suspects, however, are saddleback wrasses (Thalassoma duperrey), such as the one in the upper left portion of the photo. Four Spot, the turtle in this picture, was free of tumors in 1992. Marine Turtle Newsletter No. 151, 2016 - Page 2 Tropical Storm Dora passed by on 22 July 1993, about two hours of found year after year. Every summer, we also identify perhaps half heavy rain sent countless liters of runoff down the two concrete a dozen summer visitors, turtles that we see daily but for a single channels. This runoff was characterized by silt that left a muddy band summer only. Finally, each year we identify another ten or so along the coastline, stretching 100 m and more from the waterline. that are probably transient, because we see them once or at most, Despite wave and current action, this band persisted until well into half a dozen times. November 1993, providing a graphic demonstration of how long Turtles get together... The Honokowai turtles turn out to be social it can take to disperse material that enters the ocean in this area. creatures. They seem to like being together. A turtle arriving at a site No one tested this runoff, but because the channels both reach that is already occupied by another turtle frequently approaches the far back from the shoreline into agricultural areas, it is hard to original occupant. Often, there is physical contact. It is not unusual believe that this mud was not loaded with nutrients above and for the new arrival to settle down so that both turtles are touching beyond its natural levels, which already would be high enough in some manner. to disrupt the underwater environment. ...and are joined by cleaners. When enough turtles cluster in this This is only the most obvious example of what has been happening way, a symbiosis develops with some of the resident fauna. Fish to the waters of Honokowai. In 1989 and 1991, there were enormous discover that turtles are a source of food (Losey et al. 1994). blooms of Cladophora sericea, a seaweed usually found in deeper Some fishes will groom turtle shells, and others will look water. Since 1989, hooked red Hypnea musciformis has been for and remove such parasites as small barnacles. Some fishes establishing itself in the same area, and now is a permanent feature will even switch to feeding primarily through cleaning. A turtle along Honokowai, piling up on shore and floating in huge mats. cleaning station will evolve with several cleaner species, each of Underwater, a former garden of low corals has become choked and which represents a possible mechanical vector for the spread of overgrown with seaweeds. The northern portion of our dive site no any FP virus. longer has living corals in the 4-6 m depths. As early as 1985, Balazs In Honokowai, we see more and more species engaging in et al. documented that the southern portion of our dive site already cleaning. Some of them started because the tumors often host had dense algae growth (Balazs et al. 1985). This suggests that the parasites. At Honokowai, we have long observed that saddleback area was already receiving significant nutrient input well before the wrasses and Hawaiian spotted tobies bite at FP tumors, presumably blooms of 1989 and 1991. because the tumors are infested. In 1995, for the first time, we A bountiful foraging area. Honokowai has more seaweed in its observed long-nosed butterflies (Forcipiger flavissimus) cleaning waters than do the areas both to the north and south. Because turtles, a behavior that has continued. We believe it is significant there is so much food readily available, turtles should be more that this cleaner fish targets tumors. common at Honokowai than on the neighbor reefs. This has, in Suspects for mechanical vectors. None of the tumor-cleaners fact, been our experience. Communications with others who dive at Honokowai bite exclusively at tumors. From the beginning, West Maui tend to confirm this, although no one has undertaken we have observed this type of cleaner biting at the eyes of turtles. a thorough turtle census. The behavior occurs whether the turtle has tumors or not. One We might not be able to say exactly how many there are, but by possibility is that the cleaners are attracted to the greyish-white observing the behavior of the Honokowai turtles, we have learned mucous matter in the posterior corners of turtles’ eyes. a number of things about them. For example, we know that there is In other words, cleaners often first bite at tumors, then move on a resident population numbering roughly 30 individuals that can be to bite at the eyes of a healthy turtle. If it is possible that the FP virus

Figure 3. This picture was taken at a place we call The Graveyard. It is a long finger-like depression that drops about 1.5 m below the surrounding floor, bottoming out at about 8 m below the surface. Most of the time there is a mat of algae accumulating here, nearly deep enough to cover a 75 cm turtle. In this mat, we usually see two or three turtles. Every year, they are among the worst tumor cases we see, hence the reason for choosing its name. We speculate that the sickest turtles gather here because the area has abundant seaweeds and is shallow, so they expend as little energy as possible to get food and air. Marine Turtle Newsletter No. 151, 2016 - Page 3 Figure 4. When we first began watching turtles in 1989, we observed that goldring surgeons (C. strigosus), saddleback wrasses (T. duperrey), and whitespotted tobies (Canthigaster jactator), were the cleaners. By 1991, we began seeing eight- lined wrasses (Pseudocheilinus octoaenia) in black ring and millet seed butterfly fish (Chaetodon miliaris) participating in cleaning. This summer (1995), for the first time we saw long-nosed butterflies (Forcipiger flavissimus), engaged in cleaning activity. Unlike the other cleaner species, which we noticed at work no matter where the turtles were resting, we saw this change in the long-nosed butterflies at only one specific site. About half a dozen of these fish concentrated almost exclusively on tumors whenever the opportunity arose. Purple ring = goldring surgeons (C. strigosus); Red ring = saddleback wrasses (T. They clearly preferred tumors over duperrey); Yellow ring = whitespotted tobies (Canthigaster jactator); Black ring their usual foraging among the = eight-lined wrasses (Pseudocheilinus octoaenia); White ring = millet seed corals. This picture is interesting butterfly fish (Chaetodon miliaris); Green ring = long-nosed butterflies (Forcipiger because all six of the most common flavissimus) cleaners are present, each circled in a different color. is spread mechanically, we present some prime suspects, along with pers. comm.). We do not think it is a coincidence that it is a site with some incriminating circumstantial evidence: in almost all of the high tumor rates (Herbst 1994). turtles of Honokowai, we have documented tumors in the posterior When we check environmental data about coastal regions, corners of the eyes. With rare exception, this location is the first to we find that the kind of nutrient overload seen at Honokowai have visible tumors. is becoming common. For example, when writing about the Cleaners that bite tumors often snap at any other prominent white special aspects of Australian eutrophication, Brodie notes that this feature on a turtle’s body. We have learned to anticipate tumors on is a growing problem worldwide (Brodie 1995). As the number of any turtle subjected to these cleaning attempts. affected areas increases, so should the clustering of turtles. This FP is probably caused by a virus. In Honokowai, over 75% of the would lend itself to epidemic outbreaks of a virus anywhere that turtles we see from year to year have contracted FP. It is hard to find turtles gather in groups large enough to trigger cleaning symbiosis. a Honokowai turtle without tumors. Within 5 km north and south of The effects of site fidelity. In our experience, Honokowai turtles Honokowai, it is hard to find a turtle with tumors. If tumors are caused exhibit extremely high site fidelity. Not only do they remain in the by a communicable virus, this is the expected pattern (Herbst 1994). same area year after year, they like to settle in almost precisely Because turtles concentrate at Honokowai, the virus will spread the same spot on the reef. Honokowai has turtles that have migrated quickly through those turtles. to the French Frigate Shoals to nest and returned, to be found at the FP tumors do not appear to be a new phenomenon. Herbst same two or three favorite resting spots - within a meter! (1994) has reported suggestions that the disease has always In an area with easily obtained food, turtles also have much been present at some low level. If something encourages more time to spend resting. Our experience is that turtles seek turtles to gather at greater densities than before, there is increased each other out, so they tend to rest close together. This leads to opportunity for epidemic patterns to emerge (Herbst 1994). cleaning symbiosis and the possibility of mechanical vectors. We point out that nutrients introduced in certain types of coastal We think this can explain why a high percentage of the waters cause increased seaweed growth, an attractor for turtles. We residents at an infected site will have tumors, while turtles faithful therefore expect to find high tumor rates at this kind of site. to sites a few km away will likely be healthy. The cleaner species we suspect of being vectors are all reef fish that do not ordinarily Applying these premises to other sites. For example, the have much range. Sebastian Inlet area of the Indian River Lagoon system in Florida Unfortunately, a small percentage of the turtles we see annually has also been subjected to high nutrient input and has experienced turn out to be there for only a single summer. We believe this high seaweed growth (Woodward-Clyde Consultants et al. 1994). is long enough to become infected. If these turtles are residing We are told that it has a dense green turtle population (D.A. Bagley, elsewhere the next summer, the potential exists to spread the Marine Turtle Newsletter No. 151, 2016 - Page 4 disease and eventually give rise to another “hot spot.” eutrophication. Regardless of the cause of the ample food source, we The Honokowai cycle. When people assume that pollution is believe it has a high potential to be another tumor “hot-spot.” If the causing turtle tumors, we believe they are correct, but probably not food supply holds, the turtles will remain in the area, the population in the way they think. In summary, this is the cycle we see: density will increase, and there will be a tumor epidemic. A coastal area gets overloaded with nutrients. Right now, we can say, “Don’t put nutrients in the ocean, it’s probably not good for the turtles.” We’d like to be able to say with Seaweed growth increases proportionately. confidence, “Don’t put nutrients in the ocean, it’s definitely not Turtles are attracted by the excess of food. good for the turtles.” Sooner or later, an infected turtle arrives. BALAZS, G.H. 1980. Synopsis of Biological Data on the Green Turtles like to gather. Turtle in the Hawaiian Islands. NOAA-NMFS Tech. Memo. When turtles gather, cleaning symbiosis starts. NOAA-TM-NMFS-SWFC-7. 141pp. Cleaners learn that tumors host edible parasites. BALAZS, G.H. 1985. Behavioral changes within the recovering Cleaners bite tumors and then healthy turtles (particularly the Hawaiian green turtle population. In Keinath, J.A., D.E. Barnard, eyes). J.A. Musick & B.A. Bell (Comps.). Proceedings of the 15th More turtles become infected. Annual Symposium on Sea Turtle Biology and Conservation. More cleaners discover tumors are a food source, an escalating NOAA-NMFS Tech. Memo. NOAA-NMFS-SEFSC-387. pp. spiral. 16-21. This cycle is based primarily on observations at Honokowai BALAZS, G.H., R.G. FORSYTH & A.K.H. KAM. 1985. and how we fit them into what we have learned from research Preliminary assessment of habitat utilization by Hawaiian green papers on the disease. We do not know what occurs underwater at turtles in their resident foraging pastures. NOAA-NMFS Tech. Indian River or other tumor sites. We have not, however, found any Memo. NOAA-TM-NMFS-SWFC-71. 107pp. material that contradicts our thinking, while the concept does seem BRODIE, J. 1995. The problem of nutrients and eutrophication to fit the sites that we have information from. in the Australian marine environment. In Zann, L.P. & D.L. This does not mean we are right. We advance this idea in hope Sutton (Eds.). State of the Marine Environment Report, Technical that someone can either show us where it is wrong, or supplement Annex 2: Pollution. Great Barrier Reef Marine Park Authority, it with additional information. Townsville, Australia. pp. 1-29. http://hdl.handle.net/11017/236 The value of the Honokowai observations. We have tried to HERBST, L.H. 1994. Fibropapillomatosis of marine turtles. Annual assemble a reasonable explanation that is not in conflict with known Review of Fish Diseases 4: 389-425. facts, while offering explanations for our observations and those of researchers examining this problem. HERBST, L.H., E . R . JACOBSON, R. MORETTI, R. BROWN, If this explanation can withstand scrutiny, we can use it as J.P. SUNDBERG & K.A. KLEIN. 1995. Experimental a powerful argument in favor of reducing and even eliminating transmission of green turtle fibropapillomatosis using cell-free nutrient input into oceans that are home to green turtles. It is tumor extracts. Diseases of Aquatic Organisms 22: 1-12. important to understand that even if there are no turtles in an area, it HERBST, L.H. & P.A. KLEIN. 1995. Green turtle fibropapillomatosis: does harm to let nutrients into the water, since the area will soon challenges to assessing the role of environmental cofactors. begin to attract turtles. Areas with a low population density and Environmental Health Perspectives 103 (supp 4): 27-30. healthy turtles will become depleted. The nutrient-heavy area will LOSEY, G.S., G.H. BALAZS & L.A. PRIVITERA. 1994. Cleaning see a population density increase with a high potential for a local symbiosis between the wrasse, Thalassoma duperry, and the green tumor epidemic. turtle, Chelonia mydas. Copeia 1994: 684-690. th At the 15 Annual Sea Turtle Symposium, Balazs (1995) WOODWARD-CLYDE CONSULTANTS, MARSHALL reported that numerous Hawaiian green turtles had recently begun MCCULLY & ASSOCIATES, INC. & NATURAL SYSTEMS foraging at a site on the Kona Coast that had not previously ANALYSTS, INC. 1994. Preliminary Water and Sediment Quality been heavily populated. He reported that park personnel observed Assessment of the Indian River Lagoon, Indian River Lagoon. scores of turtles feeding, an obvious indication that the area Final Technical Report. National Estuary Program, Melbourne, has abundant seaweed growth. This is a strong indicator for FL. 212pp. www.sjrwmd.com/indianriverlagoon

Marine Turtle Newsletter No. 151, 2016 - Page 5 Artificial Incubation Trials of Leatherback Turtle Eggs at Grande Riviere Beach, Trinidad, West Indies (Phase II)

Rachael Shoy1,2 1Institute of Marine Affairs, Hilltop Lane, Chaguaramas, Trinidad, West Indies; 2Current address: #7 Happy Hill Park, Gasparillo, Trinidad and Tobago, West Indies (E-mail: [email protected])

French Guiana, Suriname and Trinidad and Tobago, situated in the housed five wooden containers constructed with pitch pine lathes eastern Caribbean, support the largest regional nesting population of dimension 91.4 cm x 61 cm x 91.4 cm and were lined with fine of leatherback turtle (Dermochelys coriacea) (Eckert 2006; Dutton mosquito mesh. Sand was collected from an area above the high et al. 2013). In Trinidad, the main nesting beaches are located on tide where leatherbacks usually nest. The sand was hand-sieved with the north and east coasts and include Grande Riviere, Matura and fine wire mesh and wooden handles, and used to fill the wooden Fishing Pond, all of which are designated Prohibited Areas by the containers. Sieving allowed for the removal of all stones and debris Forestry Act, Chapter 66:01 (Lee Lum 2005). This means that during (including past seasons’ egg shells) that might impact incubation. the March to August turtle nesting season, permits are required to An artificial ‘boot-shaped’ nest chamber was excavated inside the enter these areas and a tour guide must accompany those wishing sand filled containers down to a depth of 70 cm in keeping with the to view nesting turtles. Grande Riviere Beach is the highest density average depth of a leatherback clutch (Chacon & Machado 2005). nesting beach in Trinidad with up to 300 leatherbacks coming up to The chamber was made only when the eggs were collected for nest in one night during the peak season of April/May (Livingstone relocation. Once an egg clutch was deemed at “high risk of being 2006; Eckert 2013) and annually this results in incubating nests laid lost” to natural phenomena, biometric data of the nesting female earlier in the season being excavated by later arriving females (Lee were collected she was also checked for flipper tags and for fisheries- Lum 2005). Egg clutch loss also results from unpredictable erosion related injuries. Her eggs were collected by carefully catching them due to natural beach processes and river shifting events. The Grande in plastic bags (with no seam at the bottom of the bag, to prevent Riviere Nature Tour Guide Association (GRNTGA), a community the bag from giving way) placed directly below the cloaca. The bag based organization, has repeatedly advocated for the relocation was then closed to prevent heat loss (Chacon & Machado 2005). of eggs to a beach hatchery due to the dynamic nature of Grande To avoid contamination, eggs were singly and carefully placed Riviere Beach. However, setting up an actual hatchery enclosure for in the artificial egg chamber using gloved hands and in the same the purpose of egg incubation in the sand medium directly above order as they were laid; that is, larger fertile eggs followed by the the high tide line where leatherbacks usually nest is considered a smaller shelled albumin globs/yolkless eggs. The clutch size was risky proposition due to erratic seasonal erosion events, which are simultaneously determined. Approximately halfway through the characteristic of this beach. relocation, HOBO (UA-001-64) temperature loggers were placed To address the matter of egg loss, the Institute of Marine Affairs with the clutches to automatically record hourly temperature for (IMA) in consultation with GRNTGA developed a novel incubation the duration of the incubation periods. The sand removed to create device for leatherback eggs. The project was conducted in phases: the chamber was then used to cover over the nest in the wooden Phase I was an investigation into the feasibility of utilizing wooden container. containers to incubate leatherback sea turtle egg clutches (Jobity et al. The five clutches were relocated to the wooden containers within 2014), which gave consideration to the three main factors that must two hours of oviposition, to minimize any movement-induced be optimized for sea turtle egg incubation: temperature, moisture and gas exchange (Koch et al. 2007). In Phase I, temperatures within sand-filled wooden containers were recorded and compared to the thermal tolerance range for leatherback embryos (Ackerman 1997), which was estimated at 25-35 °C. The study concluded that the temperature ranges within all three sand-filled wooden containers were 24.7-29.9 °C, 24.3-32.1 °C and 24.7-29.9 °C; all fell within the thermal tolerance range for leatherbacks (Ackerman 1997). Phase II of the project involved artificial incubation trials with leatherback egg clutches (Fig. 1) during the 2013 nesting season. Five leatherback clutches that were deemed “high risk of being lost” to natural phenomena (e.g., erosion and/or inundation) were relocated to five sand-filled wooden containers (Jobityet al. 2014). Only five nests were permitted to be relocated because, at the time of the study, leatherback turtles were considered Critically Endangered by the International Conservation Union (IUCN 2014); they are now globally listed as Vulnerable. The objective of this phase was to determine whether the sand-filled wooden containers were capable Figure 1. Experimental hatchery enclosure on Grande of incubating leatherback sea turtle eggs. The experimental hatchery Riviere Beach. Marine Turtle Newsletter No. 151, 2016 - Page 6 the end of the incubation period and after -61.054308 W hatchling emergence, hatchling biometric data (curved carapace length, width and weight) were collected and nest excavations were conducted. For excavations, nest contents for both natural and artificially incubated nests were characterized according to (Bell et al. 2006, Hall & Parmenter 2006). The hatching success rate for all nests was calculated using the following formula: Hatching Success Rate (%) = (Total Number of Empty or Hatched shells) / (Total Number of Eggs Incubated) x 10.828922 N 100 (Herrera 2006). Unhatched eggs were also staged according to Chacon et al. (2008). Generally, the project was conducted during the wet season and the average daily temperature for the period (n = 94) was 27.7±4.2 °C (range 24.1 ±0.1 °C to 30.3 ±6.1 °C). The mean temperatures for the two natural nests that were not lost to erosion were 31.1 ±0.2°C and 27.2 ±0.6°C; the hatching success rate was 0%. Average temperatures over the incubation Figure 2. Map of Grande Riviere Bay showing the location of the experimental period for artificially incubated nests (n = 5) hatchery and natural nests. Map credit: Hamish Asmath - IMA was 30.2 ±0.8 °C; the mean hatching success mortality (Limpus et al. 1979). The wooden containers were covered rate was 25.1% (range 12.7-31.8; n = 639 eggs). The average with mosquito mesh to prevent any infestation by maggots and worldwide hatching success rate for leatherbacks is 50% (Rafferty to hold hatchlings safely in place before subsequent release. The et al. 2011). Total hatchling production from the artificial incubation normal incubation period for leatherback turtles is 60 days (Herrera trial was 156 turtles. Fig. 3 outlines the mean ambient temperature 2006) and the expected date of emergence was calculated with inside the hatchery enclosure and the mean temperatures for natural hourly monitoring approximately 2-3 days prior to the expected and artificially incubated nests, with respect to the known thermal date of emergence. The incubation period was defined as the time tolerance range for sea turtle embryos. The average weight of between oviposition and the emergence of the first hatchling. hatchlings that emerged from the trials from was 49.3 g (range 47.3- For comparison to the hatchery nests, five natural nests were 52.1; n = 15) and the average carapace length was 6.1 cm (range selected and data were collected in a similar manner to the artificially 6.0-6.3; n = 15). According to Rhodin (1985) leatherback hatchlings incubated nests. Unfortunately, three of the natural nests were lost weigh approximately 30 grams and measure 6 cm in carapace length. to erosion. Fig. 2 shows the location of the experimental hatchery, A suitable gaseous, hydrous (moisture) and thermal (temperature) the natural nests and the “at risk” nests that were relocated. At nest environment is essential for the development of sea turtle

Figure 3. Comparison of ambient conditions at hatchery. Mean temperature in natural nests, Mean temperature in hatchery nests and thermal tolerance range for leatherback incubation (lower and upper limit, from Ackerman 1997). Marine Turtle Newsletter No. 151, 2016 - Page 7 embryos (Koch et al. 2007). Throughout the incubation period the COUTOU, J., S. RAJH, N. STEWART & A. WATSON. 2013. temperature for both the natural nests and the five hatchery nests Bacterial flora identified from Leatherback turtle (Dermochelys fell within the thermal tolerance range (25-35 °C) for sea turtle coriacea) eggshells and nest sand at Grande Riviere beach, embryos (Ackerman 1997). Trinidad. The School of Veterinary Medicine, Faculty of Medical Our results indicate that the low hatching success rates for both Sciences. UWI, Trinidad and Tobago Trinidad and Tobago. 62 pp. natural and hatchery nests were because of the high percentages of Unpublished manuscript. undeveloped and unhatched eggs. Only one natural nest showed DUTTON, P.H., S.E. RODEN, K.R. STEWART, E. LA CASELLA, high late stage embryonic mortality. In leatherbacks, low hatching A. FORMIA, J.C. THOME, S.R. LIVINGSTONE, S. ECKERT, success rates may be caused by embryonic mortality, which is D. CHACON-CHAVERRI, P. RIVALAN & P. ALLMAN. 2013. poorly understood (Bell et al. 2003). Studies have also shown that Population stock structure of leatherback turtles (Dermochelys undeveloped eggs may possess an embryo that may be invisible to coriacea) in the Atlantic revealed using mt DNA and microsatellite the naked eye. Another factor that may have contributed to the low markers. Conservation Genetics 14: 625-636. hatching success rates in both natural and artificially incubated nests ECKERT, S. 2013. An assessment of population size and status of was the presence of microorganisms in the nest environment. Sand Trinidad’s leatherback sea turtle nesting colonies. WIDECAST from areas above the high tide was used to fill the wooden containers Information Doc. No. 2013-01. 14pp. and that may have had a high bacterial load. A study conducted by Coutou et al. (2013, unpublished) during the 2011-2012 nesting ECKERT, S.A. 2006. High-use areas for Atlantic leatherback sea season found that a host of bacteria existed in leatherback eggshells turtles (Dermochelys coriacea) as identified using telemetered and in the nest sand on Grande Riviere Beach. location and dive information. Marine Biology 149: 1247-1257. Overall, our results revealed that the containers have the potential HALL, S.C.B. & C.J. PARMENTER. 2006. Larvae of two signal fly to incubate leatherback turtle eggs. However, further studies must be species (Diptera: Platystomatidae), Duomyia foliate McAlphine conducted to determine whether an intervention such as artificial egg and Plagiostenopterina enderleini Hendel, are scavengers of sea incubation is required for leatherback sea turtles nesting on Grande turtle eggs. Australian Journal of Zoology 54: 245-252. Riviere Beach. Recommendations for further studies include: 1) HERRERA, A.E. 2006. The effects of management methods on sex investigation of the overall hatching success rate (%) of leatherback ratio and hatching success of leatherback turtles (Dermochelys natural beach nests and 2) evaluation of the spatial and temporal coriacea). Conservation Biology Centre for Ecology and distribution of leatherback sea turtle nests and estimation of the Conservation, UK.27pp. extent of egg loss by natural phenomena over the nesting season. JOBITY, A.M.C., R. SHOY & J. ALEMU. 2014. Artificial Acknowledgements. The author thanks: The Forestry Division incubation trials of leatherback turtle eggs at Grande Riviere of Trinidad & Tobago for permission to relocate leatherback egg Beach, Trinidad, West Indies. Marine Turtle Newsletter 142: 3-6. clutches on Grande Riviere Beach, The Green Fund Executing KOCH, A.U., M.L. GUINEA & S.D. WHITING. 2007. Effects Unit of Trinidad and Tobago (GFEU) for funding this study, of sand erosion and current harvest practices on incubation of Grande Riviere Nature Tour Guide Association for assisting with the flatback sea turtle, Natator depressus. Australian Journal of the project’s activities, Professor Indar Ramnarine, Dr. Ann Marie Zoology 55: 97-105. Jobity, Hamish Asmath, Christopher Alexis, Addison Titus and Kamau Downes. LEE LUM, L. 2005. Beach dynamics and nest distribution of the leatherback turtle (Dermochelys coriacea) at Grande Riviere, ACKERMAN, R.A. 1997. The nest environment and embryonic Trinidad. Revista de Biologiá Tropical 53: 239-248. development of sea turtles. In: Lutz, P.E. & J.A. Musick (Eds.). Biology of Sea Turtles. CRC Press, Boca Raton, Florida. pp. LIMPUS, C.J., V. BAKER & J.D. MILLER. 1997. Movement 83-103. induced mortality of loggerhead eggs. Herpetologica 35: 335-338. BELL, B.A., J.R. SPOTILA, F.V. PALADINO & R.D. REINA. LIVINGSTONE, S.R. 2006. Sea Turtle Ecology & Conservation on 2003. Low reproductive success of leatherback turtles, the North Coast of Trinidad, West Indies. PhD Thesis, Division of Dermochelys coriacea, is due to high embryonic mortality. Environmental & Evolutionary Biology. University of Glasgow. Biological Conservation 115: 131-138. 284pp. CHACON, D. & H.L. MACHADO. 2005. Anidacion de RAFFERTY, A.R., P. SANTIDRIÁN-TOMILLO, J.R. SPOTILA, Dermochelys coriacea en Playa Gandoca, Informe Temporada F.V. PALADINO & R.D. REINA. 2011. Embryonic death is 2005. Projecto de Conservacion de Tortugas Marinas, Talamanca, linked to maternal identity in the leatherback turtle (Dermochelys Caribe Sur, Costa Rica. coriacea). PLoS ONE 6 (6): e21038. doi: 10.1371/journal. pone.0021038. CHACON, D., B. DICK, E. HARRISON, L. SARTI & M. SOLANO. 2008. Technical manual of management and conservation of RHODIN, A.G. 1985. Comparative chondro-osseous development marine sea turtles in Central America. San Jose, Costa Rica. 51pp. and growth of marine turtles. Copeia 1985: 750-771.

Marine Turtle Newsletter No. 151, 2016 - Page 8 The effect of invertebrate infestation and its correlation with loggerhead sea turtle (Caretta caretta) nest success in Laganas Bay, Zakynthos, Greece

Adam J. Andrews1, Andrew C. Smith1, ALan F. Rees2 & Dimitris Margaritoulis2 1Anglia Ruskin University, Cambridge, CB1 1PT, UK (E-mail: [email protected], [email protected]); 2ARCHELON, Solomou 57, GR104-32 Athens, Greece (E-mail:[email protected], [email protected])

Loggerhead sea turtle (Caretta caretta) nests are vulnerable to relative threat of this source of predation of sea turtle eggs (Bolton predators and scavengers, including invertebrates (Paris et al. 2002). et al. 2008). Dipteran larvae (Phoridae and Sarcophagidae) have been found to At the rookery level, infestation may be high, with reports of infest loggerhead and green sea turtle (Chelonia mydas) nests both 90% (Lopes 1982) and 84.6% (Hall & Parmenter 2006) of nests in northern Cyprus (Broderick & Hancock 1997; McGowan et al. being infested. However, at nest level, infestation is typically much 2001a), and Australia (Hall & Parmenter 2006), green sea turtle lower, e.g., 10.6% (Broderick & Hancock 1997), 0.8% (McGowan nests in Costa Rica and Mexico (Fowler 1979; Lopes 1982), as well et al. 2001a) and 3.6% (Katılmış et al. 2006) of eggs within a nest as hawksbill (Eretmochelys imbricata) (Bjorndal et al. 1985) and being infested. In terms of nest success, Gautreau (2007) noted that leatherback sea turtle (Dermochelys coriacea) nests in Costa Rica it was not significantly lower for infested leatherback nests in Costa (Gautreau 2007). In the Mediterranean, coleopteran larvae were Rica, as did Bolton et al. (2008) for spiny softshell turtles (Apalone found to infest loggerhead nests in Turkey (Baran & Türkozan 1996) spinifera) in Canada. Infestation is generally not considered a threat along with Muscidae larvae (Türkozan 2000; Katılmış et al. 2006; to nest success (McGowan et al. 2001a; Hall & Parmenter 2008). Katılmış & Urhan 2007a), Acarina, Nematoda and Oligochaeta However, invertebrate predation was linked to a 30% reduction in (Baran et al. 2001; Özdemir et al. 2006; Urhan et al. 2010). There green sea turtle hatching success in Mexico (Lopes 1982), and a is currently no information on invertebrate infestation of sea turtle lower success of green, loggerhead and flatback Natator( depressus) nests laid in Greece despite the significance of the nesting population nests in Australia (Hall & Parmenter 2006) and Nile Soft-shelled at Zankynthos (Margaritoulis 2005). turtle (Trionyx triunguis) nests in Turkey (Katılmış & Urhan 2007b). Invertebrates are known to feed on weakened or dead hatchlings To date, the influence of invertebrates on the success of loggerhead (Fowler 1979; Lopes 1982), empty egg shells (Baran et al. 2001; nests has been little studied, with the only available data from Bolton et al. 2008; Urhan et al. 2010), yolk, and dead tissue (Hall Moulis’s (1997) study on the impact of fire ants (Solenopsis spp.) & Parmenter 2006; Katılmış & Urhan 2007a; Hall & Parmenter on nests laid in the USA. 2008), although they can attack viable hatchlings (Lopes 1982; McGowan et al. (2001b) concluded that three main factors McGowan et al. 2001a; Özdemir et al. 2006; Gautreau 2007) and affected infestation; nest depth, distance to the high water mark, damage intact eggs (Donlan et al. 2004; Özdemir et al. 2006; Urhan and the duration of hatchling emergence. The depth of nests was et al. 2010). There is debate as to whether lower hatching success found to be the most important factor relating to dipteran infestation observed in sea turtle nests with invertebrates can be attributed to (McGowan et al. 2001b; Bolton et al. 2008). Özdemir et al. (2004) the presence of the invertebrates, or is simply a case of nests with also reported that the duration of hatchling emergence influenced reduced hatching success having a greater likelihood of infestation infestation. The most important factor that influenced infestation by because they contain more decaying matter. Understanding the coleopteran larvae, however, was the position of nests in relation to impact of invertebrates is important in understanding the overall vegetation (Donlan et al. 2004; Özdemir et al. 2006). The aims of this study were to identify the relationship between invertebrates and the success of loggerhead sea turtle nests, to understand the factors affecting invertebrate prevalence, and to determine whether invertebrates are acting as scavengers or predators in nests. We also present the first data on the extent of infestation present in Laganas Bay (Zakynthos), one of the largest loggerhead rookeries in the Mediterranean (Margaritoulis 2005). Data were collected between 05 August - 03 September 2013. We sampled 106 loggerhead nests between East Laganas and Kalamaki beaches, located within the National Marine Park of Zakynthos, Greece (37.73° N, 20.93° E, Fig. 1). See Margaritoulis (2005) for a full description of the sample sites. Nests were excavated ≥14 days after first hatchling emergence, to ensure that natural incubation or emergence was undisturbed, in accordance with ARCHELON (the Sea Turtle Protection Society of Greece) and National Marine Park of Zakynthos (NMPZ) Figure 1. Sample sites; East Laganas beach (LAG) and protocol. The majority of nests (75) were excavated 14 days after Kalamaki beach (KAL) in Laganas Bay, Zakynthos, Greece. first emergence, 31 were excavated between 16-38 days after first emergence. Marine Turtle Newsletter No. 151, 2016 - Page 9 Nests were excavated following standard practices (Broderick Specimens were identified to family level using keys (Unwin 1984; & Hancock 1997; McGowan et al. 2001a). Once the top egg was Chinery 1993; Pearce & Waite 1994; Gibb & Osteto 2005; Checklist exposed, distance from sand level to the top egg was measured, of the Collembola of the World. Available from: www.collembola. and from the top egg to the nearest vegetation; this was recorded org), with the aid of a low-powered microscope. Statistical analyses as “0 m” if roots were present. The interval between first hatchling were performed using SPSS (v. 20.0, IMB Corporation, Armonk, emergence and excavation was also noted. New York, USA). The α level used was 0.05. One-way chi-squared Observations were made for each egg within a nest, with eggs / tests were used to investigate associations among invertebrates, the hatchlings being removed in order they were found. Hatchlings were stage of development and position within nests. Mann-Whitney treated as the final stage of egg development. Egg position within the U tests were used to compare infested and non-infested nests. A nest was recorded with the uppermost numbered 1, then 2, 3, and so Wilcoxon signed-rank test was used to compare top and bottom on. Each egg was categorized into one of the following categories: halves of nests. The influence of ecological factors on infestation hatched (≥50% empty shell), non-viable (unhatched with no sign was investigated using Generalized Linear Models with a negative of embryological development), dead or alive embryo (placed into binomial model and log link, with data rounded to the nearest integer subcategories of early, middle, late stages of development), dead or for these analyses. This model removed the need to transform the alive pipped hatchling with shell, and dead or alive hatchling. Eggs data (O’Hara & Kotze 2010). The ecological factors tested included: were inspected for infestation; defined by at least one invertebrate distance to vegetation, nest depth (sand level to top egg), clutch size, larva or adult inside, or on, the egg, or the presence of puncture number of dead (hatchlings and embryos), number of non-viable holes in the egg shell - these had perforated edges, and were too eggs, and the interval between initial nest hatching and excavation. small to be caused by the pipping process (Fig. 2). Standard success (%) was calculated as The total clutch count was divided by two and individually [((hatched eggs + hatchlings) / total clutch) x 100]. numbered eggs were allocated to the top or bottom half as they would Embryonic success (%) was calculated as have been found on excavation; if the clutch count was uneven, [((hatched eggs + hatchlings) / viable eggs) x 100]. a middle egg was discarded from analyses to avoid bias when The average number of days between observed first hatchling comparing halves of the same nest. The number of invertebrates emergence and nest excavation was 15 ±0.37 SEM, range: 14-38 infesting each egg was counted or estimated in instances of heavy days). Of the 106 loggerhead nests examined, 44 (41.5%) were infestation (>50 individuals). Adult specimens were preserved in infested by at least one invertebrate group. Infestation was greater alcohol and larvae were transported to be raised to adults. Following on East Laganas beach (46.9%, n = 31), compared with Kalamaki inspection and sampling, eggs were returned to the egg chamber beach (32.5%, n = 13). Of all eggs sampled (n = 10,223), 3.1% were and re-buried to minimise the attraction of predators and scavengers infested. Nine invertebrate taxa were recorded infesting nests (Table to nearby nests. 1). Sarcophagidae (Diptera) (97.5% larvae, 2.5% adults) (Fig. 2) Invertebrates were raised to adulthood for identification, and were the most prevalent, found in 25.5% of nests and had a strong 2 preserved using standard practice (McGowan et al. 2001a). significant association with the top half of nests (X 1 = 93.633, n

Figure 2. Sarcophagidae penetrating dead hatchling (a), punctured no visible embryo egg (b), Sarcophagidae present in non-viable egg (c), and Nematoda spp. present in late stage embryo egg (d). Photographed by Adam J. Andrews. Marine Turtle Newsletter No. 151, 2016 - Page 10 Infested Infested eggs Invertebrate Infested % nests eggs in top in bottom individuals Invertebrates nests sampled half of nest half of nest per egg Sarcophagidae (Diptera) 27 25.5 113 7 13.8 Punctured eggs 22 20.8 79 5 n/a Tenebrionidae (Coleoptera) 6 5.7 4 2 2 Elateridae (Coleoptera) 5 4.7 6 2 3.2 Nematoda spp. 4 3.8 5 11 5.9 Isotomidae (Collembola) 3 2.8 21 33 12.5 Formicidae (Hymenoptera) 3 2.8 5 20 4.7 Histeridae (Coleoptera) 2 1.9 1 1 2 Table 1. Invertebrate groups Scarabaeidae (Coleoptera) 1 0.9 0 2 2 (including punctured eggs) Rhinotermitidae (Isoptera) 1 0.9 0 1 1 observed in infested nests.

= 120, P < 0.001). Only a single live hatchling was categorised as 69.4% vs. 62.5%) (Mann-Whitney U test: U = 1108.5, n1 = 44, n2 = infested, where Formicidae numerously covered the hatchling on 62, P = 0.101) (Fig. 3: a)., which was also the case for embryonic its (assumed) journey to the surface. success (mean = 85.2% vs. 84.9%) (Mann-Whitney U test: U =

Dead hatchlings were the most frequently infested stage of 1320.5, n1 = 44, n2 = 62, P = 0.780) (Fig. 3: b). development, with 32.5% of samples infested (Table 2). Infestation Within infested nests, the proportion of infested eggs was occurred in 4% of non-viable eggs. Puncture holes were observed significantly greater in the top half of the nest (mean = 9.8% vs. in 1.8% (181) eggs and hatchlings, in 46.4% (84) of these cases, 3.2%) (Wilcoxon signed-rank test: T = 811, n = 106, P < 0.001). no invertebrates were present. Punctured eggs had a significant Up to 42.5% of the eggs in the top half of a nest were infested, 2 association with the top half of nests (one-way chi-square: X 1 = compared to a maximum of 33.3% in the bottom half of nests, 2 66.18, n = 85, P < 0.001), and with non-viable eggs (X 5 = 164, n = where infestation was more variable (Fig. 4). For Nematoda spp., 84, P < 0.001). Six eggs infested by Sarcophagidae were punctured, Isotomidae, Formicidae, Histeridae and Scarabaeidae, observed and one egg by Tenebrionidae. Sarcophagidae were the only group infestation was greater in the bottom half of nests. to penetrate hatchlings, and were significantly associated with them There was a significant negative correlation between a nest’s 2 (X 4 = 157.167, n = 120, P < 0.001). distance to vegetation and the proportion of infested eggs (GLZM(b): 2 There were more late stage embryos infested (3.1%) than any X 1 = 10.181, n = 106, P = 0.001) (Fig. 5: a) and between a nest’s 2 other stage of dead embryo eggs (Table 2.). Only 1.4% of hatched depth and the proportion of infested eggs (GLZM(b): X 1 = 106.561, eggs were infested, although this was a large number of eggs (Table n = 106, P < 0.001). There was a significant positive relationship 2 2). Nematoda spp. (X 2 = 21.125, n = 16, P < 0.001), Isotomidae between clutch size and the proportion of infested eggs (GLZM(b): 2 2 2 (X 2 = 80.333, n = 54, P < 0.001), and Formicidae (X 3 = 11.960, n X 1 = 18.459, n = 106, P < 0.001) (Fig. 5: c) and between the number

= 25, P = 0.008) were significantly associated with hatched eggs. of non-viable eggs and the proportion of infested eggs (GLZM(b): 2 In this study there was no significant difference between the X 1 = 11.061, n = 106, P = 0.001) (Fig. 5: e). success of non-infested nests compared to infested nests (mean = The proportion of infested eggs was not significantly correlated

with either the interval between hatching and excavation (GLZM(b): 2 Eggs Eggs infested/ X 1 = 1.317, n = 106, P = 0.251) or the number of dead in a nest 2 Stage of Development infested stage (%) (GLZM(b): X 1 = 0.012, n = 106, P = 0.913). Hatched (empty egg shells) 93 1.4 In this study, nests were excavated 14 days (or longer) after the first hatchling emergence. However, in the majority of other studies Non-viable 96 4 nests were excavated much sooner (within 24 hours (Acuña-Mesén Early 0 0 & Hanson 1990; Gautreau 2007; Hall & Parmenter 2008): within Middle 1 1.9 48 hours (McGowan et al. 2001a; McGowan et al. 2001b): within seven days after hatchling emergence (Katılmış et al. 2006; Özdemir Late 35 3.1 et al. 2006; Hall & Parmenter 2008; Urhan et al. 2010)). In just one Pipped 2 4 other study (Baran & Türkozan 1996) were nests excavated 14 days Dead Hatchling 90 35.2 after the first hatchling emergence. In the majority of cases, studies with a short interval to excavation noted lower levels of infestation. Live Embryo 0 0 However, since McGowan et al. (2001b), Gautreau (2007) and Live Hatchling 1 1.3 Hall & Parmenter (2008) suggest that infestation occurs during and Total 308 shortly after hatchling emergence, the interval between hatchling emergence and nest excavation should not significantly influence Table 2. Observed stages of development infested by the degree of infestation observed. We found that at least after a invertebrates. Marine Turtle Newsletter No. 151, 2016 - Page 11 Figure 3. Boxplots of nest success (%) (a) and embryonic success (%) (b), displaying differences between non-infested and infested nests. T bars = minimum and maximum values, excluding outliers. Outliers (circles) display values between 1.5 and 3 interquartile ranges from the 25th and 75th percentiles. Mean value excludes outliers.

14-day delay after emergence, there was no significant increase in nests, reported infestation levels are lower. Just 3.1% of all eggs likelihood of infestation with increased days between emergence and hatchlings sampled during this study were infested, similar to and excavation. previous studies which reported mainly dipteran infestation, e.g., Most sea turtle clutches contain embryos that fail during 0.5-0.8% to 2.1% (Baran et al. 2001; McGowan et al. 2001a), with development (Gautreau 2007). The decomposition of necrotic the exception of 10.6% from a relatively small sample reported by matter has been associated with invertebrate infestation (Fowler Broderick & Hancock (1997) for loggerhead and green sea turtle 1979; McGowan et al. 2001a; Bolton et al. 2008). Therefore most nests. nests can be expected to contain at least a few invertebrates. This Sea turtle nests, from deposition to post hatchling emergence, will cause the proportion of infested nests in a rookery to be high contain a range of potential food sources, intact viable and non- when an “infested” nest is defined by a single invertebrate infesting viable eggs at various developmental stages to live and dead a single egg/hatchling (the case in all studies mentioned). However, hatchlings, and post-hatch egg remnants. The number of invertebrate when infestation is measured as the number of infested eggs within taxa, from nine taxa (families/ orders), recorded in the excavated nests is similarly broad. Some invertebrates, notably Isotomidae, Nematoda spp., and Rhinotermitidae are known only to feed on readily available decomposing matter (Elke & Sybilla 1995; Myles 1997; Nicholas & Hodda 1999), whilst others such as Sarcophagidae are capable of puncturing eggs and predating on embryos and hatchlings (Lopes 1982). In the interest of conservation, it is necessary to understand the cause of puncture damage and predation of hatchlings in order to act against any threat posed. However, the only invertebrate group observed attacking a live hatchling was Formicidae, and in only one instance. Formicidae, in all cases fire ants, have been reported to negatively affect hatchlings of a range of species and locations (Allen et al. 2001; Paris et al. 2002; Wetterer et al. 2007). Although, Formicidae capable of stinging hatchlings are not present in Europe (Katılmış & Urhan 2007b). In previous studies (Baran & Türkozan 1996; Baran et al. 2001; Katılmış et al. 2006; Özdemir et al. 2006; Bolton et al. 2008) greater numbers of punctured eggs were found where invertebrates were present, therefore damage was linked to mainly tenebrionid larvae Figure 4. Boxplots displaying differences between the top and, in some cases, dipteran larvae (Acuña-Mesén & Hanson 1990; and bottom half infestation (%). T bars = minimum and McGowan et al. 2001a; Özdemir et al. 2006; Bolton et al. 2008). Our maximum values, excluding outliers and extremes. Outliers results showed 1.8% of eggs and hatchlings observed with puncture (circles) display values between 1.5 and 3 interquartile ranges holes, indicative of predation, which is far fewer than that reported from the 25th and 75th percentiles. Extremes (asterisks) by previous studies, e.g., 11.0% (Urhan et al. 2010), 8.2% (Özdemir display values greater than 3 interquartile ranges from the 25th and 75th percentiles. Mean value excludes outliers. et al. 2004) and 3.6% (Katılmış et al. 2006). In these studies, the Marine Turtle Newsletter No. 151, 2016 - Page 12 Figure 5. Scatterplots with trend line displaying correlation between infestation (%) and distance to vegetation (R2 = 0.035) 2 2 2 (a), nest depth (R = 0.077) (b), clutch size (R = 0.081) (c), number of dead (R = 0.8628-5) (d), number of non-viable eggs 2 2 (R = 0.066) (e), and interval between hatching and excavation (R = 0.7861-4) (f). damage was attributed to coleopteran larvae. However, in the current The finding that overall nest success was not significantly lower study, Sarcophagidae, which previous studies have reported to infest in infested nests is in agreement with the findings of Gautreau (2007) relatively low numbers of eggs (Lopes 1982; Broderick & Hancock and Bolton et al. (2008) but in contrast to those of Lopes (1982, 1997; Donlan et al. 2004; Gautreau 2007; Bolton et al. 2008), were cited in Broderick & Hancock 1997), who reports a 30% decrease in more closely associated with the punctured eggs, since they were nest success. However the sample size and method from this latter both observed in greater numbers than Tenebrionidae and infested study are unknown. That infested nests did not have a significantly a similar range of egg stages to those punctured. In agreement lower embryonic success rate is in agreement with Hall & Parmenter with other studies (Baran et al. 2001; Bolton et al. 2008; Hall & (2006). As the embryonic success of Greek loggerhead sea turtle Parmenter 2008), Sarcophagidae were associated with the top half nests was not significantly influenced by invertebrate infestation, of nests, as were punctured eggs. This increases the likelihood that it suggests that the majority of eggs infested were either those that Sarcophagidae were the cause of egg perforation, and that they were non-viable or had already hatched. Therefore the number of accessed nests by burrowing down to them. In our study, invertebrate viable eggs affected was small, indicating that the main food source abundance was not great enough to exploit the entire nest as a food within nests is not necrotic tissue, but non-viable eggs. Although, source, which may be why infested eggs were in the top half of nests we acknowledge that hatchlings may have been infested whilst alive (McGowan et al. 2001a; Katılmış et al. 2006; Bolton et al. 2008). and died before sampling was carried out. Although we cannot state with certainty that Sarcophagidae were While live hatchlings were found inside nests, only one was not responsible for the death of viable eggs or live hatchlings, their recorded as infested. Even considering the late excavation of nests trophic niche is normally that of a scavenger rather than a predator, in this study, it may be assumed that if live hatchlings were present feeding on, and breeding in, decaying animal or plant matter (Hall in infested nests, more observations would be made of invertebrates & Parmenter 2008). Therefore it is unlikely that Sarcophagidae feeding on, or at least attempting to predate them. Although several pose a threat to otherwise viable eggs. If invertebrates were able to studies have recorded invasion of live hatchlings, this has never penetrate viable eggs, infestation events approaching 100% would exceeded more than a few hatchlings in a nest (Fowler 1979; be commonly observed (Bolton et al. 2008). The greatest infestation McGowan et al. 2001a; Paris et al. 2002; Özdemir et al. 2006; in a single nest found during this study was 20.8%. However, in Gautreau 2007; Holcomb & Carr 2011). Therefore, excavating at Turkey, Özdemir et al. (2004) found that up to 88.4% of loggerhead hatchling emergence with a risk to disturbing still incubating eggs eggs may be infested, suggesting that although not often the case, may not be worthwhile considering the low risk posed to hatchlings extreme infestation events can occur. Our results suggest that in by invertebrates. Greece invertebrates principally act as scavengers, have little effect Several studies have suggested that invertebrates are only aware on nest success and pose no threat to the conservation of the species. of nests during the period of hatchling emergence. Disturbance of Marine Turtle Newsletter No. 151, 2016 - Page 13 sand by the hatchlings is thought to advertise nest position through for granting a field permit to conduct the research (Ref. No. the release of olfactory cues, thus attracting invertebrates to feed 1606/02.08.2013) and ARCHELON (the Sea Turtle Protection on the decaying matter (McGowan et al. 2001b; Gautreau 2007; Society of Greece), specifically the leadership team of 2013 at Bolton et al. 2008; Hall & Parmenter 2008). This may explain why a Zakynthos. We thank the ARCHELON volunteers who aided in the delay in excavation did not influence infestation, because hatchling data collection process. emergence had already ceased. In contrast, other studies found ACUNA-MESEN, R.A. & P.E. HANSON. 1990. Phorid fly larvae the timing of nest excavation to influence the level of infestation as predators of sea turtle eggs. Herpetological Review 21: 13-14. (McGowan et al. 2001a; Gautreau 2007). Our results suggest that ALLEN, C.R., E.A. FORYS, K.G. RICE & D.P. WOJCIK. 2001. hatchling emergence was more closely related to the infestation Effects of fire ants (Hymenoptera: Formicidae) on hatching sea of nests rather than the decomposition of matter, although more turtles and prevalence of fire ants on sea turtle nesting beaches study is needed to verify this during the first two weeks after first in Florida. Florida Entomologist 84: 250-253. hatchling emergence. The depth of nests had a significant influence on infestation rates, BARAN, I. & O. TURKOZAN. 1996. Nesting activity of the which agrees with the findings from two main studies (McGowan loggerhead sea turtle (Caretta caretta) on Fethiye Beach, Turkey. et al. 2001b; Hall & Parmenter 2008). Chemical odour traces Chelonian Conservation & Biology 2: 93-95. likely lose potency as they permeate up through the sand column BARAN, I., A. OZDEMIR, C. ILGAZ & O. TURKOZAN. 2001. (McGowan et al. 2001a). Therefore deeper nests are more difficult Impact of some invertebrates on eggs and hatchlings of loggerhead for invertebrates to locate and infest. In agreement with the current sea turtle, Caretta caretta, in Turkey. Zoology in the Middle East study, Hall & Parmenter (2008) reported that as clutch size increased, 24: 9-17. so did the infestation level. This suggests that invertebrates are BJORNDAL, K.A., A. CARR, A.B. MEYLAN & J.A. MORTIMER. attracted to decaying matter as larger clutches represent a greater 1985. Reproductive biology of the hawksbill, Eretmochelys food source. However, contrary to several studies (McGowan et imbricata, at Tortuguero, Costa Rice, with notes of the ecology of al. 2001b; Gautreau 2007; Hall & Parmenter 2008), the number of the species in the Caribbean. Biological Conservation 34: 353-368. dead in this study did not significantly affect infestation levels. This BOLTON, R.M., R.J. BROOKS & S.A. MARSHALL. 2008. may be because invertebrates are attracted by decaying non-viable Opportunistic exploitation of sea turtle eggs by Tripanurga eggs (Broderick & Hancock 1997; Acuña-Mesén & Hanson 1990; importuna. Canadian Journal of Zoology 86: 151-164. Saumure et al. 2006; Holcomb & Carr 2011). Therefore infested nests were those with a larger number of non-viable eggs, and a BRODERICK, A.C. & E.G. HANCOCK. 1997. Insect infestation larger clutch size (Katılmış & Urhan 2007a; Bolton et al. 2008). of Mediterranean sea turtle eggs. Herpetological Review 28: We found that as the number of non-viable eggs increase, so did 190-191. the level of infestation. CHINERY, M. 1993. Insects of Britain and Northern Europe (Collins While other studies noted predominantly dipteran infestation Field Guide) 3rd Edition. Cambridge University Press, Cambridge, (McGowan et al. 2001b; Hall & Parmenter 2006; Bolton et al. Cambridgeshire, UK. 2008), distance to vegetation significantly influenced infestation DONLAN, E.M., J.H TOWNSED & E.A. GOLDEN. 2004. in the current study. This suggests that these invertebrates as well Predation of Caretta caretta (Testudines: Cheloniidae) eggs as coleopterans may be associated with vegetation (Katılmış et al. by larvae of Lanelater sallei (Coleoptera: Elateridae) on Key 2006; Özdemir et al. 2006; Katılmış & Urhan 2007a). Because Biscayne, Florida. Caribbean Journal of Science 40: 415-420. coleopterans have a greater ability to cause damage to nests ELKE, M. & H. SYBILLA. 1995. Soil micro-arthropods (Acari, (Katılmış et al. 2006), nest relocation and hatchery position may be Collembola) from beach and dune: characteristics and ecosystem considered for sites with abundant Coleoptera if a large proportion context. Journal of Coastal Conservation 1: 77-86. of nests are laid close to vegetation. The infestation rates at nest level for Zakynthos were similarly FOWLER, L.E. 1979. Hatching success and nest predation in the low when compared with previous studies (Baran et al. 2001; green sea turtle, Chelonia mydas, at Tortuguero, Costa Rica. McGowan et al. 2001a; Katılmış et al. 2006). It is likely that Ecology 60: 946-955. invertebrates infested nests shortly before, or during hatchling GAUTREAU, S. 2007. Dipteran larvae infestation of leatherback emergence, with invertebrates acting as scavengers, feeding mainly sea turtle (Dermochelys coriacea) nests on Gandoca Beach, on non-viable eggs or dead hatchlings. Infestation appears to be more Costa Rica. M.Sc. Thesis. The University of Guelph, Ontario, closely related to hatchling emergence than the amount of decaying Canada. 101 pp. matter, as such, we did not detect an influence in delaying excavation GIBB, T.J. & C. OSTETO. 2005. Arthropod Collection and >14 days on infestation rates. More work is needed to accurately Identification; Laboratory and Field Techniques. London define the arrival times of invertebrates, as this remains poorly Academic Press, London, UK. understood. Our study suggests that invertebrates were attracted HALL, S.C.B. & C.J. PARMENTER. 2006. Larvae of two signal to nests with a lower hatching success, and were not the cause of fly species (Diptera: Platystomatidae),Duomyia foliate McAlpine reduced success. It is thus suggested that in the Mediterranean, and Plagiostenopterina enderleini Hendel, are scavengers of sea invertebrates pose little threat to sea turtle nests. turtle eggs. Australian Journal of Zoology 54: 245-252. Acknowledgements. Thanks are extended to the Research Ethics HALL, S.C.B. & C.J. PARMENTER. 2008. Necrotic egg and Subcommittee of Anglia Ruskin University who approved hatchling remains are key factors attracting dipterans to sea turtle the research, the NMPZ (National Marine Park of Zakynthos) Marine Turtle Newsletter No. 151, 2016 - Page 14 (Caretta caretta, Chelonia mydas, Natator depressus) nests in MYLES, T. G. 1997. Comparison of the penetrability of smooth and Central Queensland, Australia. Copeia: 75-81. crushed sand by subterranean termites (Isoptera: Rhinotermitidae). HOLCOMB, S.R. & J.L CARR. 2011. Infestation of a naturally Sociobiology 30: 295-304. incubated nest of the alligator snapping turtle (Macrochelys NICHOLAS, W.L. & M. HODDA. 1999. The free-living nematodes temminckii) by the Phorid fly Megaselia scalaris. Southwestern of a temperate, high energy, sandy beach: faunal composition Naturalist 56: 427-429. and variation over space and time. Hydrobiologia 394: 113-127. KATILMIS, Y., R. URHAN, Y. KASKA, E. BASKALE. 2006. O’HARA, R.B. & D.J. KOTZE. 2010. Do not log-transform count Invertebrate infestation on eggs and hatchlings of the loggerhead data. Methods in Ecology and Evolution 1: 118–122. sea turtle (Caretta caretta), in Dalaman, Turkey. Biodiversity and OZDEMIR, A., O. TURKOZAN, C. ILGAZ & R. MARTIN. 2004. Conservation 15: 3721-3730. Nest site factors and invertebrate infestation of loggerhead sea KATILMIS, Y. & R. URHAN. 2007a. Physical factors influencing turtle nests. Israel Journal of Zoology 50: 333-340. Muscidae and Pimelia sp. (Tenebrionidae) infestation of OZDEMIR, A., C. ILGAZ, Y. KATILMIS & S.H. DURMUS. 2006. loggerhead sea turtle (Caretta caretta) nests on Dalaman Beach, Invertebrate infestation of Caretta caretta nests at Fethiye beaches, Turkey. Journal of Natural History 41: 213-218. Turkey. Pakistan Journal of Biological Sciences 9: 507-513. KATILMIS, Y. & R. URHAN. 2007b. Insects and mites infestation PARRIS, L.B., M. LAMONT & R.C. CARTHY. 2002. Increased on eggs and hatchlings of the Nile soft-shelled turtle (Trionyx incidence of red imported fire ant (Hymenoptera: Formicidae) triunguis) in Kurkurtlu Lake (Dalaman, Turkey). Zoology in the presence in loggerhead sea turtle (Testudines: Cheloniidae) nests Middle East 40: 39-44. and observations of hatchling mortality. Florida Entomologist LOPES, H.S. 1982. On Eumacronychia sternalis Allen (Diptera, 85: 514-517. Sarcophagidae) with larvae living on eggs and hatchlings of the PEARCE, M. J. & B. S. WAITE. 1994. A list of termite genera east Pacific green sea turtle. Revista Brasileira de Biologia 42: (Isoptera) with comments on taxonomic changes and regional 425-429. distribution. Sociobiology 23: 247-263. MARGARITOULIS, D. 2005. Nesting activity and reproductive SAUMURE, R.A., A.D. WALDE & T.A. WHEELER. 2006. Non- output of loggerhead sea turtles, Caretta caretta, over 19 seasons predatory fly larvae (Delia platura: Anthomyiidae) in the nest (1984-2002) at Laganas Bay, Zakynthos Island, Greece: The of a northern map turtle (Graptemys geographica). Chelonian largest rookery in the Mediterranean. Chelonian Conservation & Conservation & Biology 5: 274-275. Biology 4: 916-929. TURKOZAN, O. 2000. Reproductive ecology of the loggerhead MCGOWAN, A., A.C. BRODERICK, J. DEEMING, B.J. GODLEY (Caretta caretta) on Fethiye and Kizilot beaches Turkey. & E.G. HANCOCK. 2001a. Dipteran infestation of loggerhead Chelonian Conservation & Biology 3: 686-692. (Caretta caretta) and green (Chelonia mydas) sea turtle nests in UNWIN, D.M. 1984. A Key to the Families of British Coleoptera. northern Cyprus. Journal of Natural History 35: 573–581. Cambridge University Press, Cambridge, Cambridgeshire, UK. MCGOWAN, A., L.V. ROWE, A.C. BRODERICK & B.J GODLEY. URHAN, R., Y. KATILMIS & M. YUKSEL. 2010. Invertebrate 2001b. Nest factors predisposing loggerhead sea turtle (Caretta infestation in loggerhead sea turtle (Caretta caretta) nests, in caretta) clutches to infestation by dipteran larvae on northern Dalyan, Turkey. Munis Entomology and Zoology 5: 982-984. Cyprus. Copeia: 808-812. WETTERER, J., L.D. WOOD, C. JOHNSON., H. KRAHE & S. MOULIS, R.A. 1997. Predation by the imported fire ant Solenopsis( FIRCHETT. 2007. Predaceous ants, beach replenishment, and invicta) on loggerhead sea turtle (Caretta caretta) nests on Wassaw nest placement by sea turtles. Environmental Entomology 36: National Wildlife Refuge, Georgia. Chelonian Conservation & 1084-1091. Biology 2: 433-436.

Marine Turtle Newsletter No. 151, 2016 - Page 15 Assessing the Impacts of Hatcheries on Green Turtle Hatchlings

Carmen Mejías Balsalobre1,2 & Ian Bride2 1Kosgoda Sea Turtle Conservation Project (E-mail: [email protected]); 2Durrell Institute of Conservation and Ecology (E-mail: [email protected])

At present, six of the seven marine turtle species are globally of the valuable energy reserves used to distance themselves from classified as likely to become extinct in the near future by the shore (Gyuris 1994). International Union for Conservation of Nature (www.iucnredlist. The aim of this study was to examine how hatchling retention org). One tool commonly utilized in sea turtle conservation is the might affect survivability by assessing different quality parameters translocation of eggs into hatcheries (Mortimer 1999). Hatcheries (body condition, crawling and swimming performance), in green are widely perceived as being beneficial in protecting eggs from turtles (Chelonia mydas). The results provide valuable information threats such as poachers, natural predators and environmental that can be used to improve practice in Sri Lanka and indeed, in pressures (Mortimer 1999). Additionally, this strategy can be used hatcheries worldwide, in respect to their contribution to sea turtle to promote ecotourism and thereby provide financial income for conservation. local people (Rajakaruna et al. 2013). However, hatchery-based The study was conducted in May-June 2015 at the Kosgoda conservation programs have also provoked debate about their Sea Turtle Conservation Project (KSTCP; 6°N, 80°E), one of the effectiveness because they may negatively affect turtle populations. seven hatcheries situated along the southwest coast of Sri Lanka Some of the potential dangers of hatcheries include: detrimental (Rajakaruna et al. 2013). Kosgoda is Sri Lanka´s second largest effects on embryonic development and hatching success (Pritchard rookery, and is visited by five species of sea turtle, including the 1980; Mortimer 1999); high rates of mortality caused by incorrect green turtle, which exhibits a year round high nesting frequency. The release methods (Mortimer 1999); skewed sex ratios due to the coastline of Kosgoda has a high presence of human activity, mainly thermal effect of specific environmental conditions (Morrealeet al. due to beach tourism, which can be a cause of severe disturbance 1982; Mortimer 1999); and detrimental effects on hatchling energy for in situ nests. Nests can also be affected by the presence of and behavior when they are retained in artificial tanks (Pilcher & animal predators and tidal inundation (Ekanayake et al. 2010). In Enderby 2001; van de Merwe et al. 2013). Consequently, moving Kosgoda, local villagers collect the freshly laid eggs at night from eggs to hatcheries is considered an option of last resort, when in situ the surrounding beaches and sell them to the hatchery owner to be conservation is not a viable option (IUCN 2005). Nevertheless, this reburied first thing in the morning in the incubation pens, where does not mean that hatcheries cannot make a positive contribution, as they are then protected until they produce hatchlings (Tisdell & their effectiveness relies on the way that they are managed (Tisdell Wilson 2005). In the case of KSTCP, hatchlings are kept in tanks & Wilson 2005). for two or a maximum of three days. After this retention, tourists In Sri Lanka, hatcheries have proliferated, primarily as an indirect and volunteers release them at sunset at 5-10 m from the tideline so consequence of the effects of the high human population density hatchlings can crawl down the beach and get to the sea. (Rajakaruna et al. 2013). There are few special protected areas for During this study 10 ex situ green turtles nests were visually sea turtles in the country, which makes in situ conservation difficult inspected for emergence and, before trials, all newly emerged (Hewavisenthi 1993). Under these circumstances hatcheries seem to hatchlings were captured and transported to seawater-filled holding offer the most suitable conservation strategy. Although it is claimed tanks (160 cm long × 135 cm wide × 100 cm high) where they could that the primary motive of most of the Sri Lankan hatchery owners freely swim. In order to assess crawling speed and swimming power is profit from tourism, there is also a general understanding on their stroke rate, hatchlings were divided into five groups according to part of the need for hatcheries in turtle conservation (Rajakaruna the time since emergence: just emerged (0 hr), 6 hr, 12 hr, 24 hr et al. 2013). According to Rajakaruna et al. (2013), the closure of and 48 hr after emergence, as hatchlings are usually kept for two hatcheries in Sri Lanka would be impractical, thus there is a need days in KSTCP. Each group was comprised of 3 randomly selected to improve the poor practices employed in most of them. hatchlings from each nest and they were marked on the carapace One of the hatchery practices identified as in need of improvement for identification and to avoid being selected twice. After each is the post-emergence handling of the hatchlings. In the wild, swimming and running trials, the hatchlings were weighed with hatchlings emerge from the nest and immediately crawl frenetically an electric balance (±0.01 g) and measured along their notch to tip to reach the sea. Once in the sea, hatchlings swim continuously, in straight carapace length (SCL) and straight carapace width (SCW) a state of energetic frenzy, in order to get away from the shore as using a Vernier caliper (±0.1 mm). An overall size index, similar quickly as they can. The frenzy period is characterized by rapid to that used by Ischer et al. (2009), was calculated by multiplying and effective power strokes interspersed by a less effective dog SCL by SCW. All procedures were carefully carried out while trying paddling swimming style and resting periods, which become more to diminish any procedural stress inflicted on study hatchlings. frequent as time passes (Wyneken & Salmon 1992). In most Sri Hatchlings were not fed at any point during the first 48 hr after Lankan hatcheries however, hatchlings are held for 1-7 days after emergence and were eventually released at the discretion of the emergence to provide a tourist attraction (Rajakaruna et al. 2013). hatchery owner. This retention may result in a disturbance of their natural behavior To test crawling speed, hatchlings were run along a 3 m raceway. so as to compromise their chances of survival by depleting some The raceway (3 m × 0.5 m) was located outside with natural light Marine Turtle Newsletter No. 151, 2016 - Page 16 conditions but permanently shaded and with a slight downward acclimatization, hatchlings were videotaped for one minute at the slope facing the sea in order to emulate natural conditions as much beginning (0 min), middle (30 min) and end (60 min) of the hour as possible. A dull light was also placed at the end of the raceway to trial. The videotapes were then played back at slow speed (25%) add another stimulus for the hatchlings to run in the right direction; and the power strokes manually counted during the minute interval. hatchlings naturally crawl towards the main light source they see Power stroke rate was calculated as the average of power strokes (Pilcher et al. 2000). Crawling speed was calculated (speed [v] = min-1 of the three replicates. distance [d] / time [t]) by timing how long each hatchling took to IBM SPSS Statistics v22 was used to analyze the data. As the data crawl the 3 m. As hatchlings were exposed to natural fluctuations did not conform to the assumptions of a normal distribution, the non- in temperature, the air temperatures at the time of trials were parametric Kruskal-Wallis H test (KW-H) was used to determine obtained from the daily weather data recorded by Freemeteo (http:// if there were differences in average power stroke rate, crawling freemeteo.com.lk). speed and body measurements over time. In the cases where the Swimming performance in each group was measured using KW-H test was significant a pairwise Mann-Whitney U test (MW- a method similar to the one described by Burgess et al. (2006). U) was executed to determine which groups exhibited significant Hatchlings were allowed to swim individually for one hour in tanks differences. Relationship between variables of morphology, (60 cm long × 42 cm wide × 36 cm high), filled with 30 cm of temperature and performance were investigated using a Spearman’s seawater at 30 ºC (the average water temperature from the tanks of rank order correlation (S-rho). Statistical differences and rejection KSTCP during this season). They were fitted with a Velcro harness of the null hypothesis were assumed if p< 0.05. that provided resistance for the turtles to swim against, but did not Hatchling mass was correlated with hatchling size index in impede motion, simulating the natural environment (Salmon & hatchlings from swimming (S-rho [rs]=0.761, p=0.001) and crawling

Wyneken 1987). The harness was connected to a monofilament (rs =0.654, p=0.001) trials. However, the only morphology condition nylon tied to another tense monofilament above in the center of the that showed a significant change during hours of retention was the tank. To reduce visual stimuli and induce unidirectional oriented size index (KW-H [H]=20.844, p=0.001) of hatchlings in swimming swimming (Salmon & Wyneken 1987), three sides of the enclosure trials. Further comparisons (MW-U) showed between which periods were covered with black plastic and a dim light was placed at the of retention the significant differences were found (Table 1). For remaining side. Hatchlings were allowed to swim freely in the example, the first significant change (MW-U [U] =232, p=0.001) in tanks, but the nylon monofilament length prevented them from size happened after 24 hr. As Fig. 1 indicates, there was an increase touching the sides or the bottom of the enclosure. After 1 min of of the median size index between hours of retention. After 24 hr (Median [min, max]=1924 [1556, 2208] mm2), the median size Groups 0 hr 6 hr 12 hr 24 hr 48 hr index of the hatchlings was 5% greater than that of newly emerged 0 hr 1 hatchlings (Median [min, max]=1827 [1635, 2128] mm2). After 48 6 hr 0.564 1 hr (Median [min, max]=1965 [1779, 2229] mm2), the median size 12 hr 0.668 0.976 1 index was 8% greater. A total of 150 hatchlings completed the crawling trials. The 24 hr 0.001 0.005 0.048 1 KW-H test showed that crawling speed decreased significantly 48 hr 0.001 0.001 0.015 0.478 1 (H=17.872, p=0.01) when hatchlings were retained in the tanks Table 1. Pairwise MW-U test of hatchling size index between for 48 hr. MW-U test (Table 2) specified the significant difference groups, using 95% confidence intervals, in swimming trials. between particular groups, with the first one happening after 24 hr. According to the median speed of each group (Fig. 2), hatchlings Groups 0 hr 6 hr 12 hr 24 hr 48 hr assessed after 24 hr (Median [min, max] 0.072 [0.03, 0.169] m s-1) 0 hr 1 ran 26% slower than newly emerged ones (Median [SE]=0.097 [0.041, 0.243] m s-1) and up to 27% slower when assessed after 48 6 hr 0.976 1 hr (Median [min, max]=0.071 [0.016, 0.118] m s-1). In addition, 12 hr 0.367 0.326 1 minimum and maximum values in Fig. 2 denote a general tendency 24 hr 0.012 0.019 0.008 1 of the hatchlings to run slower with hours of retention. No correlation was detected between weight and crawling speed (r =-0.062, 48 hr 0.005 0.004 0.004 0.745 1 s p=0.448) or between size index and speed (rs=-0.093, p=0.259). Table 2. Pairwise MW-U test of crawling speed between Air temperature also was found to have no significant correlation groups, using 95% confidence intervals. (rs=0.136, p=0.096) with crawling performance. Another 150 hatchlings participated in the swimming trials, where Groups 0 hr 6 hr 12 hr 24 hr 48 hr the application of the KW-H test to the resulting data indicated that 0 hr 1 power stroke rate decreased significantly (H=19.538, p=0.01) with 6 hr 0.128 1 increasing hours of retention. Subsequently, the MW-U test (Table 3) found that the differences were not significant between adjacent 12 hr 0.004 0.188 1 groups, suggesting a gradual change. According to the medians of 24 hr 0.002 0.121 0.859 1 the average power strokes for each group, there was a decrease in 48 hr 0.001 0.010 0.371 0.478 1 swimming performance through hours of retention (Fig. 2), e.g., [min, max]= [57,149] -1 Table 3. Pairwise MW-U test of power stroke rate between after 48 hr (Median 117 strokes min ), the groups, using 95% confidence intervals. power stroke rate decreased up to 16%. However, retention between Marine Turtle Newsletter No. 151, 2016 - Page 17 Figure 1. Body condition. Box and whiskers plot of the size index of green turtle hatchlings from swimming trials, during hours of retention (n=150). The boxes represent the 25%-75%, whiskers represent the minimum and maximum values, and lines in the box represent the median values of the distribution.

0 (Median [min, max]=140 [99, 171] strokes min-1) and 12 hr (Median [min, max]=126 ± [76, 163] strokes min-1) was associated for a decrease of 10% in the median power stroke. Minimum and maximum values in Fig. 2 also denote a general tendency of the hatchlings to reduce their power stroke rate with hours of retention. Figure 2. Locomotor performance. Box and whiskers The S-rho test revealed a weak, statistically significant, negative plot of the crawling speed (upper panel) and power stroke correlation between size index and power stroke rate (r =-0.181, rate (lower panel) of green turtle hatchlings during hours s of retention (n=150). The boxes represent the 25%-75%, p=0.027). No correlation was observed between weight and power whiskers represent the minimum and maximum values, r stroke rate ( s=-0.035, p=0.673). and lines in the box represent the median values of the The only body condition that recorded a significant change across distribution. hours of retention was the size index of hatchlings after swimming trials. Hatchlings slowly increased their size during time of retention trials showed a significant change in size, and although statistically with the first significant change after 24 hr. After 48 hr, hatchlings’ significant, this change was relatively small and therefore unlikely median size index had increased 8%, from 1827-1965 mm2. Since to play a major role in regards to predation. hatchlings were not fed at any point during the first 48 hr and the Crawling performance decreased with time of retention. Air absorption of the residual yolk in reptiles is not likely to be directly temperatures during the speed trials ranged from 24-31ºC, though involved in the growth of the hatchlings (Kraemer & Bennet 1981; the most frequent temperature was 29 ºC. Even though temperature Radder et al. 2007), the most likely explanation for this increase has been reported to influence performance in turtles (Adamset al. in size is rehydration. Bennett et al. (1986) reported loggerhead 1989), in this case the time spent running in the trials, 12-18 s, was hatchlings (Caretta caretta) losing 12% of their weight due to not long enough for temperature to influence hatchling crawling dehydration in the process of emergence. Hatchlings can rehydrate performance. In addition, body condition showed no correlation by drinking water once they enter the sea, but it takes from 10-15 with hatchling crawling speed. Therefore, the observed change in days to recover their hatching weight (Bennett et al. 1986). In Sri crawling speed was most likely related to period of retention. In this Lankan hatcheries, hatchlings are placed in the tanks with marine study, the median hatchling speed was reduced by 26%, from 0.097- water from the moment they are collected from the nests, despite 0.072 m s-1, after 24 hr and 27%, from 0.097-0.071 m s-1, after 48 hr suggestions from hatchery management guidelines (Mortimer 1999), of retention. This decrease in crawling speed was not gradual as the which recommend that they should be kept inside a damp cloth sack first significant change was observed after the first 24 hr. However, in a cool dark quiet space. As hatchlings spent more time inside the as hatching usually occurs immediately after sunset, if hatchlings tanks, they rehydrated, increased their weight and therefore their are not released just after emergence, up to 24 hr will need to pass size, with hatchling mass showing a strong positive correlation with until the next release window, by which time the hatchlings will have size. However, no significant change was shown in weight, which already lost valuable running speed capacity. Releasing hatchlings may be due to the fact that the increase in size was relatively small during the morning is considered an improper method since it is and a significant change in weight perhaps required more hours of likely to decrease their chances of survival (Mortimer 1999). A rehydration. An increase in the body size of the hatchlings during previous study by van de Merwe et al. (2013) also investigated the retention might improve chances of survival, following the “bigger effect of time of retention in green turtle crawling performance. is better hypothesis” (Gyuris 2000). According to this hypothesis, Although their results coincide in terms of speed reduction, in their larger hatchlings are less susceptible to predation as they can avoid study the decrease in swimming speed was greater; after just 6 hr of gap-limited predators. However, only hatchlings from swimming retention the hatchlings’ mean crawling speed decreased by 50%. Marine Turtle Newsletter No. 151, 2016 - Page 18 One plausible explanation for this difference from the current study 2006). However, the decrease found over hours of retention in this is that retention conditions were not similar, as hatchlings were study was not as marked as that found by previous studies. For retained in nest netting during the entire period, thereby suffering example, the first significant decrease happened after 12 hr, while dehydration (van de Merwe et al. 2013), whereas in this experiment Pereira et al. (2011) reported a rapid decrease of power stroke rate they were kept in tanks. Rusli et al. (2015), who studied the effects of green turtles during the first 2 hr of swimming, followed by a of different incubation methods on locomotor performance, found slower decrease after 8-12 hr. Pilcher & Enderby (2001) found that that 48 hr retention in styrofoam boxes actually improved crawling from 4-6 hr of retention the hatchlings used at the end of the trials speed. However, this improved speed was approximately half that of exhibited a more erratic power stroke instead of a continuous one. hatchlings newly emerged from in situ and hatchery nests. So this In their experiment they quantified swimming speed and found a alternative method of incubation and retention would likely not be significant reduction after 3 hr of retention, and after 6 hr, a drop effective for Sri Lankan hatcheries, where speed would be reduced by over 12%. Although the results of the present study were not in comparison with newly emerged hatchings from ex situ nests. quite as pronounced, in terms of recommendations for hatchery In nature, hatchlings emerge from their nest and crawl rapidly management they do support previous findings. Moreover, because towards the sea where they disperse into offshore waters (Wyneken hatchlings need to be released after sunset, if they are not released & Salmon 1992). Their crawling performance is important as immediately after emergence, which usually occurs soon after they are exposed to predators during the beach running stage. sunset, 24 hr would need to pass until the next release when power Nevertheless, in captivity some hatcheries offer protection when stroke rate would have significantly declined. Having an energetic releasing the offspring (Mortimer 1999). In the observed case of and rapid swimming performance can be important for survival, as KSTCP, volunteers, tourists and owners protected the hatchlings hatchlings do not display any other predation avoidance mechanism from predators on their way to the sea. However, this practice may (Gyuris 1994). Consequently, hatchlings should be released right not be followed by hatcheries worldwide, especially during the after emergence to avoid this reduction in swimming performance. tourist off-season. In the case of hatchlings released and not protected Populations of the endangered green turtle in Asia are believed after hours of retention, a decrease in crawling performance is likely to have declined over the last decades, including Sri Lankan to affect their chances of survival in the wild. And even if they are populations (Shanker & Pilcher 2003). The highest mortality rate protected, retention is likely to affect their chances of being predated in sea turtles occurs during the first stages of their lives, between once they enter the sea. incubation, crawling to the sea and swimming away from shore Although some researchers have investigated swimming (Crouse et al. 1987). An experiment by Pilcher et al. (2000) found performance by employing direct measurements in the wild (e.g., hatchlings suffer 40-60% mortality within the first two hours in the Salmon & Wyneken 1987; Gyuris 1994; Pilcher et al. 2000), sea, but once they reach deeper waters this predation rate decreased this is difficult in terms of logistics. In the present study, indirect by two thirds. With such high levels of mortality, it is important that measures of swimming performance were used following a model hatchery management practice seeks to maximize the chances of similar to the one described by Burgess et al. (2006); hatchlings survival of the hatchlings by minimizing the depletion of the energy were tethered in tanks instead of using a raceway system as other they need for the frenzy swim. The present study further reinforces studies (e.g., Pilcher & Enderby 2001). It is important to consider the idea that time of retention has a negative impact on hatchlings, that tethering the hatchlings might affect their swimming behavior by reducing crawling and swimming performance. The reduction and therefore the resulting data might not accurately reflect their of swimming performance in this case can be considered the most behavior the under natural conditions. However, results of previous potentially significant outcome in terms of survival, as hatchlings studies with tethered hatchlings under experimental conditions cannot be protected while they swim to deep waters. recorded similar behavior to that found in their natural environment In addition, this retention may affect the natural migration of (Wyneken & Salmon 1992). Power stroke rate was assumed to be hatchlings. Okuyama et al. (2009) suggested that retention of a valid parameter for assessing swimming performance, as power hatchlings decreases their probabilities of experiencing the natural strokes generate the greatest swimming force and they have been migration of wild hatchlings. Releasing hatchlings offshore to found to be more than twice as effective as dog paddling (Ischer et reduce their mortality rate and minimize the effects of retention al. 2009). In this study power stroke rate suffered a gradual decrease on their migratory route is a practice carried out by some hatchery with increased retention time. Despite previous studies having found operators (Hewavisenthi 1993). In the past hatcheries have been a relation between body condition and swimming performance discouraged from using this strategy as it may disturb the imprinting (Burgess et al. 2006; Ischer et al. 2009), in the present study the mechanism of hatchlings, which may affect females in their return to correlation between swimming performance and size index was not the natal beaches for nesting (Pritchard 1980). However, according strong enough to explain the decrease in power stroke rate. It can to Lohmann & Lohmann (1996), sea turtles may be able to use therefore be assumed that the decrease in swimming performance the earth´s magnetic field to return to their natal nesting beaches was mainly due to retention time. using a bicoordinate magnetic map. Hence, offshore release may According to the data, after 12 hr there was a 10% drop in the be a good solution; nonetheless, further study is required. Another median power stroke rate, from 140-126 strokes min-1, and after strategy followed by hatcheries is to feed the hatchlings prior to 48 hr this drop increased up to 16%, from 140-117 strokes min-1. release (Rajakaruna et al. 2013). The effects of feeding on hatchling These findings seems to be in line with previous studies, where condition needs to be investigated, but this is complicated by the hatchlings gradually decrease their power stroke rate as they move likelihood that not all hatchlings will consume the same amount through the frenzy period, and when dog paddling and resting of food in these early stages. Therefore, it still seems that the best become more frequent (Wyneken & Salmon 1992; Burgess et al. practice would be to release hatchlings just after emergence. Marine Turtle Newsletter No. 151, 2016 - Page 19 The effectiveness of sea turtle hatcheries relies on improving KRAEMER, J.E. & S.H. BENNETT.1981. Utilization of current practices (Tisdell & Wilson 2005). This study provides posthatching yolk in loggerhead sea turtles, Caretta caretta. Copeia experimental evidence supporting the importance of releasing 1981: 406-411. hatchlings immediately after emergence. Hatcheries may be LOHMANN, K.J. & C.M.F. LOHMANN. 1996. Orientation and somewhat resistant to the idea of adjusting their ecotourism policies open-sea navigation in sea turtles. Journal of Experimental to maximize hatchling survival upon release, as this would mean Biology 199: 73-81. that the collection and release of hatchlings should take place in MORREALE, S.J., G.J. RUIZ, J.R. SPOTILA & E.A. STANDORA. the dark, which may be less appealing for tourists. However, it 1982. Temperature-dependent sex determination: current practices is possible to combine tourism and conservation, following best threaten conservation of sea turtles. Science 216: 1245-1247. practice guidelines (IUCN 2005), which recommend releasing at least 90% of the hatchlings from each nest immediately after MORTIMER, J.A. 1999. Reducing threats to eggs and hatchlings: emergence, and holding the remaining hatchlings as a tourist hatcheries. In: Eckert, K.L., K.A. Bjorndal, F.A. Abreu-Grobois attraction. Tourists could release these remaining hatchlings, despite & M. Donnelly (Eds.). Research and Management Techniques being fewer in number, the next day at the sunset. In this way, Sri for the Conservation of Sea Turtles. IUCN/SSC Marine Turtle Lankan, and turtle hatcheries worldwide, would be able to improve Specialist Group Publication 4. pp. 175-178. their contribution to sea turtle conservation whilst maintaining much OKUYAMA, J., O. ABE, H. NISHIZAWA, M. KOBAYASHI, needed tourist revenue. K.YOSEDA & N. ARAI. 2009. Ontogeny of the dispersal Acknowledgements. We thank Kosgoda Sea Turtle Conservation migration of green turtle (Chelonia mydas) hatchlings. Journal of Project (KSTCP), and especially D. Perera, (the director of KSTCP) Experimental Marine Biology and Ecology 379: 43-50. for allowing us to spend time at the hatchery and providing their PEREIRA, C.M., D.T. BOOTH & C.J. LIMPUS. 2011. Locomotor assistance for everything that we needed. We also thank the Durrell activity during the frenzy swim: analysing early swimming Institute of Conservation and Ecology for the financial support. behaviour in hatchling sea turtles. Journal of Experimental ADAMS, N.A., D.L. CLAUSSEN & J. SKILLINGS. 1989. Effects Biology 214: 3972-3976. of temperature on voluntary locomotion of the eastern box turtle, PILCHER, N.J. & S. ENDERBY. 2001. Effects of prolonged Terrapene carolina carolina. Copeia 1989: 905-915. retention in hatcheries on green turtle (Chelonia mydas) hatchling BENNETT, J.M., L.E. TAPLIN & G.C. GRIGG. 1986. Sea water swimming speed and survival. Journal of Herpetology: 633-638. drinking as a homeostatic response to dehydration in hatchling PILCHER, N.J., S. ENDERBY, T. STRINGELL & L. BATEMAN. loggerhead turtles Caretta caretta. Comparative Biochemistry 2000. Nearshore turtle hatchling distribution and predation. In: and Physiology A 83: 507-513. Pilcher, N. & G. Ismail (Eds.). Sea Turtles of the Indo-Pacific: BURGESS, E.A., D.T. BOOTH & J.M. LANYON. 2006. Swimming Research, Management and Conservation. Asean Academic Press, performance of hatchling green turtles is affected by incubation London. pp. 151-166. temperature. Coral Reefs 25: 341-349. PRITCHARD, P.C.H. 1980. The conservation of sea turtles: CROUSE, D.T., L.B. CROWDER & H. CASWELL. 1987. A stage- practices and problems. American Zoologist 20: 609-617. based population model for loggerhead sea turtles and implications RADDER, R.S., D.A. WARNER, J.J. CUERVO & R. SHINE. 2007. for conservation. Ecology 68: 1412-1423. The functional significance of residual yolk in hatchling lizards EKANAYAKE, E.M.L., R.S. RAJAKARUNA, T. KAPURUSINGHE, Amphibolurus muricatus (Agamidae). Functional Ecology 21: M.M. SAMAN, D.S. RATHNAKUMARA, P. SAMARAWEERA 302-309. & K.B. RANAWANA. 2010. Nesting behaviour of the green RAJAKARUNA, R.S., E.L. EKANAYAKE & T. KAPURUSINGHE. turtle at Kosgoda rookery, Sri Lanka. Ceylon Journal of Science 2013. Sea turtle hatcheries in Sri Lanka: Their activities and (Biological Sciences) 39: 109-120. potential contribution to sea turtle conservation. Indian Ocean GYURIS, E. 1994. The rate of predation by fishes on hatchlings Turtle Newsletter 17: 2-12. of the green turtle (Chelonia mydas). Coral Reefs 13: 137-144. RUSLI, M.U., J. JOSEPH, H.C. LIEW & Z. BACHOK. 2015. GYURIS, E. 2000. The relationship between body size and predation Effects of egg incubation methods on locomotor performances rates on hatchlings of the green turtle (Chelonia mydas): is bigger of green turtle (Chelonia mydas) hatchlings. Sains Malaysiana better? In: Pilcher, N. & G. Ismail (Eds.). Sea Turtles of the 44: 49-55. Indo-Pacific: Research, Management and Conservation. Asean SALMON, M. & J. WYNEKEN. 1987. Orientation and swimming Academic Press, London. pp.143-147. behavior of hatchling loggerhead turtles Caretta caretta L. during HEWAVISENTHI, S. 1993. Turtle hatcheries in Sri Lanka: boon or their offshore migration. Journal of Experimental Marine Biology bane. Marine Turtle Newsletter 60: 19-22. and Ecology 109: 137-153. ISCHER, T., K. IRELAND & D.T. BOOTH. 2009. Locomotion SHANKER, K. & N.J. PILCHER. 2003. Marine turtle conservation performance of green turtle hatchlings from the Heron Island in South and Southeast Asia: hopeless cause or cause for Rookery, Great Barrier Reef. Marine Biology 156: 1399-1409. hope. Marine Turtle Newsletter 100: 43-51. INTERNATIONAL UNION FOR CONSERVATION OF NATURE TISDELL, C. & C. WILSON. 2005. Do open-cycle hatcheries (IUCN). 2005. Marine Turtle Conservation Strategy & Action Plan relying on tourism conserve sea turtles? Sri Lankan developments for Sri Lanka. Colombo: Department of Wildlife Conservation 79. and economic-ecological considerations. Environmental Management 35: 441-452. Marine Turtle Newsletter No. 151, 2016 - Page 20 VAN DE MERWE, J., K. IBRAHIM & J. WHITTIER. 2013. Post‐ WYNEKEN, J. & M. SALMON. 1992. Frenzy and postfrenzy emergence handling of green turtle hatchlings: improving hatchery swimming activity in loggerhead, green, and leatherback hatchling management worldwide. Animal Conservation 16: 316-323. sea turtles. Copeia 1992: 478-484.

Evidence of Turtle Poaching On Agalega, Mauritius

Imogen Webster1, Adele Cadinouche2 & Annette Huggins3 1Mauritius Marine Conservation Society, c/o Mauritius Underwater Group, Railway Rd. Phoenix, Mauritius (E-mail: [email protected]); 2SEABIOME, Riviere Noire, Mauritius (E-mail: [email protected]) 3Whitstable, Kent, UK (E-mail: [email protected])

The remote islands of Agalega (10°25’S, 56°40’E), lie 990 km to last rookeries within the Mauritian EEZ (Mangar & Chapman 1996; the north of Mauritius, west of the Mascarene Ridge (Fig. 1). The Bourjea et al. 2008). In 2006, Tatayah & Griffiths (2007) confirmed islands are a dependency of Mauritius and are managed by the that at least two species of marine turtle regularly nest on Agalega: Outer Island Development Corporation (OIDC), a government- greens (Chelonia mydas) and hawksbills (Eretmochelys imbricata). parastatal organization. Currently there are no reliable nesting data There have been few scientific trips to Agalega, consequently the for Mauritius or its islands (Bourjea et al. 2008) and the population flora and fauna are still poorly known, and this is especially true and conservation status of both species is unknown (Mangar & for marine fauna. With increasing interest from commercial bodies Chapman 1996). During the 19th century, the area was heavily to develop tourism and other projects on the islands, it is important exploited for turtle shell, meat and eggs (Mangar & Chapman to document the biodiversity of the islands. In this context, in 1996). This resulted in the protection of all sea turtles in Mauritian June 2013, the Mauritius Marine Conservation Society (MMCS) waters since 1948 (Chapman & Swinnerton 1996), although there conducted an opportunistic expedition to Agalega via a routine was no legal enforcement until their inclusion in the Fisheries Act, supply ship voyage. The objectives were to record mega-fauna 1980 and most recently the Fisheries and Marine Resources Act, sightings during the boat trip to and from the island and, while 1998 (Koonjul 2009). there, to collect data on various wildlife activity including turtles, Nesting of marine turtles was considered common on the outer cetaceans and sea birds, in addition to documenting potential threats islands of Mauritius with Agalega and St. Brandon thought to be the to the island environment. Agalega is a coral island in the western Indian Ocean (Fosberg 50°0’ 55°0’ et al. 1983). It is composed of two islands, North (11.7 km2) and South Island (9.3 km2), separated by a shallow pass 1.5 km wide.

-10 °0’ North Island is long and narrow (12.4 km x 1.6 km), whereas South Island is rounder, measuring 6.6 km x 3.6 km (Fig. 1). Montagne d’Emmerez (15 m) on North Island is the highest point. The islands are bound by sandy beaches of 20-50 m width, and are surrounded by a narrow lagoon, 25-100 m wide (Cheke & Lawley 1983) with areas of both seagrass beds and reef, providing excellent habitat for foraging juvenile sea turtles as well as mating and nesting adults. During the voyage two observers recorded the position and species (where possible) of any turtle sightings. Throughout the -15 °0’ seven days on the islands any signs of turtle activity, including nesting attempts, tracks and sightings in the lagoon were noted. Additionally, evidence of poaching (e.g., bones, carapace, egg shells) was recorded while walking the beaches and sand roads of both islands. Global Positioning System coordinates of any activity were recorded and photographs taken. Where possible the curved carapace length (CCL) of dead animals was recorded to give an idea of life stage.

-20 °0’ No turtles were observed during the voyage. While on the island turtles were seen in the lagoon on both sides of North Island and an adult green turtle was sighted when it surfaced in the vicinity of Figure 1. Position of Agalega in the Indian Ocean relative the anchored ship off Port St. James while loading for departure. to Mauritius. Sightings occurred from the beach near the village of Vingt Cinq, Marine Turtle Newsletter No. 151, 2016 - Page 21 A B C

Figure 2. A. Palm leaves used as a working area. B. and C. Bones and carapace of poached animals. and another three individuals were seen in the lagoon on the west juveniles encountered were netted, with no consideration for sex coast during approximately four hours of opportunistic snorkelling and/or size of individuals caught. Furthermore, unlaid and laid eggs by two of us. Based on the size (<60 cm CCL), these turtles were are collected for consumption. If this is occurring at a subsistence juveniles and at least one was a green turtle. All displayed nervous level, it is expected that there would be limited impact on the general behaviour, moving away rapidly from the observers. turtle population as there are so few people living on the islands. A number of locations were identified where turtles had been However, it was suggested that poaching activity increased when the poached. Sites were characterized by palm leaves spread out on supply boat was due, implying that meat and products were being the ground to work on (Fig. 2a) and/or bone and carapace pieces in sold and sent back to the mainland. This represents a potential threat the area (Fig. 2bc). At some sites, the palm leaves were still green to the population if there is ongoing demand. No hawksbill shells and remains were decomposing, indicating recent kills. In total, were found. However, they are known to occur in the area and are remains of approximately nine individuals were found along with reportedly caught (Griffiths & Tatayah 2007). As the carapace of some eggshells. Sites were distributed around South Island with a this species is highly valued, we suspect it would be removed and single site identified on the southern end of North Island (Fig. 3). sold. Interviews conducted by Griffiths & Tatayah (2007) revealed However, this site appeared to have been used numerous times over that locals considered the hawksbill turtle common, more so than the long periods with both green palm fronds and bleached plastron green turtle, which were reported as being seen only occasionally. and carapace pieces scattered through the adjacent vegetation. The turtles of Agalega are under threat from numerous sources. Skull shape and scute patterns on the carapaces indicated all were These include marine debris in the form of plastic and nets and/ green turtles. Only four intact shells were recovered and the curved 56°36’ 56°40’ carapace length of these ranged from 92 -104 cm. This size range indicates that these were breeding individuals of green turtles as -10 °20’ females reach sexual maturity around 95-100 cm and males at ≤95 cm (Limpus & Chaloupka 1997). Our observations indicated juveniles use the lagoon area and poaching occurs on the islands, especially on South Island. Agalega has extensive, undeveloped coastline and a small human population (between 250-350); as such it is likely the islands represent an important habitat for marine turtle nesting (Griffiths &Tatayah 2007). However, no tracks were found on the beaches that were covered, but this could be due to the time of year rather than a -10 °24’ lack of nesting activity. In the Western Indian Ocean, green turtle nesting shows a variation in the peak of activity between islands (Dalleau et al. 2012) ranging from summer (January) on Tromelin to early winter (June) on Mayotte (Bourjea et al. 2007; Dalleau et al. 2012). While green turtles can nest throughout the year, hawksbill nesting peaks during the summer months (October - February) (Mortimer et al. 2011). Only juveniles were observed in the lagoon area. Agalega has extensive seagrass beds in the pass between the islands and around South island, as well as coral reef structure; this is ideal feeding habitat for both green and hawksbill turtles (Meylan -10 °28’ & Donnelly 1999). A concerned citizen reported that sea turtles coming to nest were regularly flipped on the beach by locals, animals mating in the lagoon would also be caught together, and, in addition, any Figure 3. Locations of turtle remains and evidence of poaching on Agalega. Marine Turtle Newsletter No. 151, 2016 - Page 22 or fishing gear (Griffiths & Tatayah 2007; Webster & Cadinouche Acknowledgements. Thanks to the Outer Islands Development 2013), direct hunting both in the sea and when they come ashore Corporation for the opportunity to visit the island, Captain and to lay eggs, as well as the poaching of turtle nests and habitat crew of the Mauritius Pride for support during observations on destruction from heavy use of the lagoon by local fishermen with the voyage and island Manager Mr. Carver, National Coast Guard damaging techniques including walking on the reef and dragging and Police and Mr. X for their local knowledge, comments and nets (Webster & Cadinouche 2013). A further threat involves attacks assistance while on the islands. by domestic but free ranging dogs on nesting females and their nests. BOURJEA, J., J. FRAPPIER, M. QUILLARD, S. CICCIONE, D. Our observations support those reported by Griffiths & Tatayah ROOS G. HUGHES & H. GRIZEL. 2007. Mayotte Island: another (2007) and show that there is turtle activity in this area but also important green turtle nesting site in the southwest Indian Ocean. that poaching is a continuing threat. The Mauritian laws protecting Endangered Species Research 3: 273-282. marine turtles prohibit the capture, killing or selling of marine BOURJEA, J., R. NEL, N.S. JIDDAWI, M.S. KOONJUL & turtles or their products. During Griffiths & Tatayah’s visit (2007), G. BIANCHI. 2008. Sea turtle bycatch in the WIO: Review, interviews with locals revealed that they knew turtle poaching was recommendations and research priorities. Western Indian Ocean illegal but continued with the activity due to the potential to earn Journal of Marine Science 7: 137-150. money. In addition, they said that there was limited enforcement and observations from our expedition show that this is still the case. CHAPMAN, R.E. & K.J. SWINNERTON. 1996. The Mauritius On mainland Mauritius, the outlook for turtles is increasingly Wildlife Fund St. Brandon Expedition: Marine Turtles. pp 1-7. positive with regular sightings of both hawksbill and green turtles CHEKE, A.S. & J.C. LAWLEY. 1983. Biological history of Agalega, (Webster 2013), the implementation of marine education projects with special reference to birds and other land vertebrates. Atoll in schools and a general increase in public awareness such as the Research Bulletin 273: 65-108. reporting of injured animals. In February 2014 there was a sighting DALLEAU, M., S. CICCIONE, J.A. MORTIMER, J. GARNIER, of a green turtle on one of the southern beaches (R. Mohit, pers. S. BENHAMOU & J. BOURJEA. 2012. Nesting phenology of comm.). However, the last confirmed successful nesting on the marine turtles: insights from a regional comparative analysis on mainland was in 2007 (Koonjul 2009). Based on observations green turtle (Chelonia mydas). PLoS ONE 7(10): e46920. reported here, the future of those turtles using the outer islands is FOSBERG, F.R., M.-H. SACHET & D.R. STODDART. 1983. of concern, especially as these islands are possibly some of the last List of the vascular flora of Agalega. Atoll Research Bulletin nesting sites in Mauritius. The level of poaching on the island needs 273: 109-142. to be determined and a baseline of the turtle population nesting there established. A number of alternatives could be attempted GRIFFITHS, O. & V. TATAYAH. 2007. Rapid survey of marine to combat this gap in knowledge. These could include a) training turtles in Agalega, Western Indian Ocean. Marine Turtle and employing locals in protection and monitoring, b) focused Newsletter 115: 14-15. education in schools and environmental awareness programs for KOONJUL, M.S. 2009. Green turtle nesting at Gris Gris beach in all island residents, including those posted from Mauritius, c) Mauritius. Indian Ocean Turtle Newsletter. 9: 24. regular patrolling and enforcement of Mauritian law protecting LIMPUS, C. & M. CHALOUPKA. 1997. Nonparametric regression these animals with appropriate level consequences to discourage modeling of green sea turtle growth rates (southern Great Barrier further poaching and d) possible warden posted during peak breeding Reef). Marine Ecology Progress Series 34: 23-34. seasons for both species. MANGAR, V. & R. CHAPMAN. 1996. The status of sea turtle Considering that there are already Coast Guard and Police conservation in Mauritius. In: HUMPHREY, S.L. & R.V. SALM regularly posted to Agalega, it should not be difficult to also have (Eds.). Status of Sea Turtle Conservation in the Western Indian a ‘nature warden’ posted on one or both islands, similar to that on Ocean. UNEP Regional Seas Reports and Studies No. 165. IUCN/ Round Island Nature Reserve with National Parks Conservation UNEP, Nairobi, Kenya. pp. 121-124. Services and Mauritian Wildlife Foundation. Alternatively, a local inhabitant could be trained to conduct all necessary enforcement MEYLAN, A.B. & M. DONNELLY. 1999. Status justification for activities. Enforcement of current laws is essential not only for listing the hawksbill turtle (Eretmochelys imbricata) as Critically the protection of the turtles but also the mangroves, birds, coconut Endangered on the 1996 IUCN Red List of Threatened Species. crabs and marine mammals found on and around the island. Should Chelonian Conservation & Biology 3: 200-224. a warden be posted or regular research visits implemented it would MORTIMER, J.A., J. CAMILLE & N. BONIFACE. 2011. provide the opportunity to undertake monitoring to examine the long Seasonality and status of nesting hawksbill (Eretmochelys term condition of both species and their habitats to quickly identify imbricata) and green (Chelonia mydas) turtles at D’Arros Island, and minimize threats. Mauritius currently has no reliable nesting Amirantes Group, Seychelles. Chelonian Conservation & Biology data (Bourjea et al. 2008) therefore Agalega provides an opportunity 10: 26-33. to address this gap. Turtle populations should be monitored using WEBSTER, I. 2013. Observations of green and hawksbill turtles methods employed in other areas of the Western Indian Ocean, on the southwest coast of Mauritius. Marine Turtle Newsletter specifically during peak nesting periods to determine abundance 138: 15-17. and nesting trends as this will provide maximum data while also WEBSTER, I. & A. CADINOUCHE. 2013. Agalega expedition assisting in reducing poaching activities. All details collected could report: summary of results with recommendations for then contribute to regional knowledge of turtle movement and management, research and monitoring. Report to the Outer population status within the Indian Ocean. Island Development Corporation. 27pp. Marine Turtle Newsletter No. 151, 2016 - Page 23 Hatching of Eggs Rescued from a Green Turtle Involved in an Automobile Collision

Isao Kawazu1, Kunio Komesu2 & Muneyuki Kayo2 1Okinawa Churashima Foudation, 888 Ishikawa, Motobu, Okinawa 905-0206, Japan (E-mail: [email protected]); 2Sea Turtle Association of Japan, Nagaomotomachi 5-17-18-302, Hirakata, Osaka 573-0163, Japan

Female green turtles (Chelonia mydas) nest on the sand beaches to the scene for her rescue, she was already dead by the time we of Okinawa Island, Japan, usually during the spring and summer arrived (Fig. 3A). Parts of the internal organs spilled out through months (May - August; Uchida 1985; Kikukawa et al. 1999). Nest its split carapace, which was a horrible sight (Fig. 3A). However, sites selected by green turtles are most often on the backshore that it seemed that the turtle was still alive just before our arrival. After has uneven beach topography and permanently vegetated areas inspecting her tracks on the beach, we concluded that she did not (Hays et al. 1995; Turkozan et al. 2011). Approximately 2 months lay the eggs and that they were still in her oviducts. We proceeded after egg laying (i.e., nesting), the hatchlings emerge from their immediately to examine her reproductive organs (Figs. 3B and C). nests in a frenzy and immediately move toward the sea, swimming We rescued 83 eggs from her oviducts in the early morning of 17 actively offshore when they reach the water (Musick & Limpus August (Fig. 3D). Simultaneously, the reproductive organs (left and 1997). right ovaries and the oviducts) without the eggs were also removed, Kijyoka Beach is located in northwest Okinawa Island, Japan fixed in Bouin’s solution, and stored in 70% alcohol and kept as an (26°42′32′′N, E128°08′40′′E; Fig. 1). It is a nesting site of academic specimen at the Okinawa Churashima Research Center, loggerhead turtles (Caretta caretta), green turtles, and hawksbill Okinawa, Japan, for future analysis of oviduct structure. turtles (Eretmochelys imbricata) (Kikukawa et al. 1999; Komesu Miller (1999) categorized oviposited eggs of sea turtles as: 1) et al. 2016). However, the environmental conditions for nesting and hatching of sea turtles on this beach are unsafe. Permanent A vegetation grows wild in the hinterland of this beach (Figs. 2B and C), and the main road of Okinawa runs along the beach (Figs. 2A and C). In addition, the lighting along the main road illuminates this beach in the night, thereby inducing road kill of hatchlings (Komesu et al. 2016). Hatchlings are phototactic and thus attracted to light (Lohmann et al. 1997). On the night of 16 August 2015, a female green turtle arrived for nesting on Kijyoka Beach and mistakenly entered the main road. Unfortunately, a car collided with the turtle. Although we went B

C

Figure 1. A map of Okinawa Island and Kijyoka Beach of Figure 2. Photographs of the Kijyoka Beach where a female Ogimi-son. green turtle was killed in a traffic accident. Marine Turtle Newsletter No. 151, 2016 - Page 24 A B

C D

E F

Figure 3. Photographs of the female green turtle killed in a traffic accident (A and B), both oviducts (C), rescued eggs (D), the incubator used for hatching the eggs (E), and the hatchlings (F).

Marine Turtle Newsletter No. 151, 2016 - Page 25 normal eggs (of pliable shape, with albumen and yolk), 2) very Acknowledgments. We express our deep gratitude to the citizens small eggs (50% diameter of a normal egg, without yolk), and 3) of Kijyoka, Ogimi-son, Okinawa Prefecture, Japan, for providing odd-shaped eggs (unpliable shape, without yolk). Based on this us information about the traffic accident involving the green turtle. categorization, the 83 rescued eggs were classified as: 49 normal, 1 We also thank staff of the Okinawa Churashima Research Center very small, and 1 odd-shaped egg in the left oviduct and 31 normal for their help with the dissection of the green turtle and the artificial and 1 odd-shaped egg in the right oviduct (clutch size: 80 normal, 1 incubation of the rescued eggs. very small, and 2 odd-shaped eggs in both oviducts). The diameter HAYS, G.C., A. MACKAY, C.R. ADAMS, J.A. MORTIMER, of the 80 normal eggs was 45.8 ± 0.8 mm (mean ± SD; range: 43.5 J.R. SPEAKMAN & M. BOEREMA. 1995. Nest site selection - 48.0 mm). Eighty normal eggs were incubated at a uniform air by sea turtles. Journal of the Marine Biological Association UK temperature of 29 °C and controlled humidity of more than 90%, 75: 667-674. maintained by fresh-water spray, in an incubator (Autoelex Co., HIRTH, H.F. 1980. Some aspects of the nesting behavior and Ltd., Gimhae, Korea; Fig. 3E). reproductive biology of sea turtles. American Zoologist 20: After 54 days of incubation (on 10 October 2015), a pip was 507-523. confirmed, which continued until 14 October. After 58 days, 20 baby turtles hatched from the eggs (Fig. 3F; hatching success: 25%). KIKUKAWA, A., N. KAMEZAKI & H. OTA. 1999. Current status Their straight carapace length was 47.8 ± 2.0 mm (range: 43.6 - of the sea turtles nesting on Okinawajima and adjacent islands of 51.1 mm), and body mass was 26.0 ± 1.6 g (range: 23.3 - 28.5 g). the central Ryukyus, Japan. Biological Conservation 87: 149-153. After confirming that the baby turtles were in a frenzy (excitement KOMESU, K., M. KOGACHI, M. KAYOU & I. KAWAZU. 2016. phase), they were released at night on the Kijyoka shore where Record of a hawksbill turtle with regular reproductive cycle in their mother had the accident. In other words, the baby turtles were Ogimi of Okinawa Island, Japan. Umigame Newsletter of Japan released and journeyed into the ocean from the shore where they 103: 6-10. were meant to hatch. LOHMANN, K.J., B.E. WITHERINGTON, C.M.F. LOHMANN To the best of our knowledge, this case is the first report of & M. SALMON. 1997. Orientation, navigation, and natal beach hatching success of eggs rescued from the female green turtle killed homing in sea turtles. In: Lutz, P.L. & J.A. Musick (Eds.). The in a traffic accident. However, the rate of hatching success (25%) Biology of Sea Turtles, Volume 1. CRC Press, Boca Raton, FL. was lower than the rate of hatchling emergence observed in the pp. 137-163. wild, which usually exceeds 70% (Hirth 1980; Miller 1997). Miller MILLER J.D. 1997. Reproduction in sea turtles. In: Lutz, P.L. & (1999) categorized the different phases in embryo development J.A. Musick (Eds.). The Biology of Sea Turtles, Volume 1. CRC and defined the egg content as follows: 1) undeveloped (unhatched Press, Boca Raton, FL. pp. 52-81. eggs with no obvious embryo; 2) unhatched (unhatched eggs with obvious small embryo); and 3) unhatched term (unhatched, apparent MILLER J.D. 1999. Determining clutch size and hatching success. full term embryo in intact egg shell or pipped, with a small amount In: Eckert K.L., K.A. Bjorndal, F.A. Abreu-Grobois & M. of external yolk material). Based on this classification, there were Donnelly (Eds.). Research and Management Techniques for the 47 (58.8%) undeveloped eggs, 0 (0%) unhatched eggs, and 13 Conservation of Sea Turtles. IUCN/SSC Marine Turtle Specialist (16.2%) unhatched term eggs, out of the 80 rescued normal eggs. Group Publication 4. pp. 124-129. The number of undeveloped eggs (58.8%) was abnormally high, MUSICK, J.A. & C.J. LIMPUS. 1997. Habitat utilization and because the actual fertility of deposited eggs probably exceeded migration in juvenile sea turtles. In: Lutz, P.L. & J.A. Musick 95% (Miller 1997). We therefore believe that the shock from the (Eds.). The Biology of Sea Turtles, Volume 1. CRC Press, Boca car crash damaged to the eggs prior to embryogenesis, leading to Raton, FL. pp. 137-163. the low hatching rate. Moreover, the eggs may have suffered other TURKOZAN, O., K. YAMAMOTO & C. YILMAZ. 2011. Nest site damage during egg removal (approximately 2 hours), although we preference and hatching success of green (Chelonia mydas) and gave scrupulous attention to manipulation of the eggs. loggerhead (Caretta caretta) sea turtles at Akyatan Beach, Turkey. It is generally unusual to have succeeded in artificially hatching Chelonian Conservation & Biology 10: 270-275. eggs that were removed from a turtle’s body and it was that much UCHIDA, S. 1985. The sea turtles in the Nansei Archipelago Japan. more gratifying to release the hatchlings. However, the most Animals and Nature 15: 22-26. important action is the preservation of the shore environment so that sea turtles can safely lay their eggs. With lessons learned from this accident, we hope measures will be taken to prevent sea turtles from entering roads where there is heavy traffic alongside sea turtle nesting beaches.

Marine Turtle Newsletter No. 151, 2016 - Page 26 Northernmost Records of Hawksbill Sea Turtle Nests and Possible Trans-Atlantic Colonization Event

Sarah A. Finn1, William P. Thompson2, Brian M. Shamblin3, Campbell J. Nairn3 & Matthew H. Godfrey1 1Wildlife Diversity Section, NC Wildlife Resources Commission, Wilmington, NC USA (E-mail: [email protected]; [email protected]); 2Cape Hatteras National Seashore, National Park Service, Lighthouse Rd., Buxton, NC 27920 USA (E-mail: [email protected]); 3D.B. Warnell School of Forestry and Natural Resources, University of Georgia, Athens, GA 30602, USA (E-mail: [email protected]; [email protected])

The hawksbill sea turtle (Eretmochelys imbricata) is considered DNA extraction and genotyping protocols as previously described to be mostly tropical in its distribution in the Atlantic, Pacific and (Shamblin et al. 2011). Briefly, a single fresh egg is collected from Indian Oceans (Witzell 1983; Meylan & Redlow 2006). In the each nest. The egg is opened, its contents discarded, and the eggshell western Atlantic, this species can be found in waters between 30° is preserved in 95% ethanol prior to DNA extraction. Extracted N and 30° S, and regular nesting is more concentrated on beaches maternal genomic DNA is amplified at 16 microsatellite markers to between Cuba to the north and the state of Bahia in Brazil to the produce a unique genetic tag for each nesting female. Of the markers south (Prosdocimi et al. 2014). Occasional nesting (<4 per year) by currently used, there are five that can also identify hawksbill turtles: hawksbills in the state of Florida, USA, has been documented, with CcP1H11, CcP1G03, CcP5C08, CcP1F01, CcP2H12 (Shamblin et no known nests in the SE USA outside Florida, with the exception al. 2007; 2009; CJN and BMS unpublished data). For the eggshell of a single hawksbill nest found in Texas (Meylan & Redlow 2006). samples collected from the two clutches in Table 1, all five of In North Carolina, there have been only 10 reports of stranded these markers indicated that the alleles were pure hawksbill. Also, hawksbill sea turtles since stranding records were kept beginning in egghshell samples from both nests in Table 1 had identical genetic the mid-1980s; all were small juveniles (≤25 cm straight carapace fingerprints for the 13 loci available for comparisons, meaning that length), and all were observed between 2001-2009 (NCWRC the same individual laid both clutches. Sea Turtle Stranding and Salvage Network, unpublished data). In In order to assess possible nest beach origins for this female, contrast, stranded loggerhead, green, Kemp’s ridley and leatherback mitochondrial DNA from both samples was analyzed as previously turtles are found annually in North Carolina (www.seaturtle.org/ described, except using primer LCM rather than LTEi9 (Gorham strand). Of sea turtle nests found on North Carolina beaches, the et al. 2014). Briefly, an 817 base pair (bp) fragment of the majority are laid by loggerheads, with occasional nests laid by green, mitochondrial control region was amplified using primers LCM leatherback and Kemp’s ridley turtles (Bowen et al. 1994; Woodson and H950 and sequenced using LCM and an internal sequencing & Webster 1999; Rabon et al. 2003). primer, Cc271. Comparisons with reference sequences from During the second half of the 2015 nesting season in North Genbank confirmed that this female carried haplotype EiA48. To Carolina, two hawksbill nests were documented at Cape Hatteras date, this haplotype has not been found in any western Atlantic National Seashore, which is located on southern Hatteras Island in nesting populations, but it has been found in foraging populations. Dare County, North Carolina (Table 1). Both nests were initially It was previously reported in two of 18 juveniles foraging around recorded as being laid by loggerhead turtles, based on characteristics Ascension Island in the central Atlantic (Putman et al. 2014). An of the crawl left on the beach by the nesting females, and both analogous haplotype, EiBR7 (739 bp EiBR7 is inclusive within the clutches were relocated because they were laid in a zone that is prone EiA48 sequence), was detected in 6% of juveniles foraging around to daily overwash and erosion in the late summer and early fall. the Rocas Atoll and the Fernando de Noronha Archipelago off the However, through genetic analysis, we were able to conclusively northeastern coast of Brazil (Vilaça et al. 2013). The 380 bp version identify these two nests as hawksbill. of these haplotypes, EATL, is the dominant haplotype in eastern Eggshell samples were collected as part of an ongoing loggerhead Atlantic foraging aggregations, accounting for 68% of Cape Verde turtle genetic capture-recapture project aimed at identifying juveniles and 92% of Príncipe juveniles (Monzón-Argüello et al. individual females nesting north of Florida for the estimation 2010; 2011). EATL been reported in only a single individual in the of population size, clutch frequency, remigration intervals, and Northwest Atlantic, a juvenile foraging near Buck Island, United annual survival (www.seaturtle.org/nestdb/genetics). Eggshell States Virgin Islands (Bowen et al. 2007). The only known nesting samples were processed in accordance with the standard eggshell source in the Atlantic for EATL is the Príncipe Island rookery off

Clutch Date Emergence Date Laid Lat Long size emerged success Comments 7/13/2015 35.198 -75.724 96 9/10/2015 64% One dead hatchling in nest Nest washed out by storm on 9/4/2015 35.199 -75.721 206 N/A N/A 2015-09-28

Table 1. Descriptive data of the two hawksbill sea turtle clutches laid in North Carolina in 2015. Marine Turtle Newsletter No. 151, 2016 - Page 27 the west coast of Africa (Monzón-Argüello et al. 2011), and some of is usually done following incubation, when nest contents are these females were carrying EiA48 specifically (Monzón-Argüello inventoried and species-specific patterns of carapacial scutes of pers. comm. 2016). EiA48 is also identical to the 766 bp EiIP-16 hatchlings remaining in the nest cavity can be discerned (Serafiniet (haplotype EiIP-16 is inclusive within EiA48), and this haplotype al. 2009). Although the nesting female who laid these two clutches has been recorded at low frequency in the Seychelles and Chagos was pure hawksbill, it is possible that she mated with a different Archipelago nesting populations that comprise the western/central species and produced hybrid offspring (Karl et al. 1995). In Bahia, Indian Ocean genetic stock (Vargas et al. 2016). Brazil, where both species nest, hybridization between hawksbill and Trans-Atlantic dispersal of oceanic juveniles was previously loggerhead turtles is common: approximately 40% of morphological invoked to explain the presence of neritic juvenile hawksbills hawksbills carry loggerhead mitochondrial DNA (Lara-Ruiz et al. carrying apparent Eastern Atlantic haplotypes on Western Atlantic 2006). Analyses of Brazilian hybrid sea turtle nuclear DNA, which foraging grounds (Bowen et al. 2007, Vilaça et al. 2013). This is inherited from both parents, confirm that these hybrids are fertile connectivity has been further substantiated by three tag returns for and capable of back crossing with both parental species (Vilaça et hawksbills tagged by Project TAMAR as juveniles in Brazil later al. 2012). Unfortunately, no hatchling or embryo samples were appearing along the west coast of Africa (Vilaça et al. 2013). Storm- collected from the hawksbill nests in North Carolina. Thus, it is forced dispersal of oceanic juveniles has been implicated to explain unknown whether the hatchlings were pure hawksbill or possibly the presence of juveniles outside of their normal distribution, and hawksbill x loggerhead hybrids. residence in these novel juvenile foraging areas may in turn affect The second clutch of hawksbill eggs was lost to severe erosion adult migratory pathways for these wayward turtles (Monzón- from a week-long northeastern storm with sustained 20-30 mph NE Argüello et al. 2012). Adult female hawksbill turtles typically winds, and beach front wave heights ranging from 4-8 feet. This, exhibit strong natal homing, so much so that rookeries on the in turn, kept park staff from employing further protective measures, leeward and windward coats of Barbados separated by less than and thus there was no chance to identify species by visual inspection 25 km are genetically distinct populations (Browne et al. 2009). of live or dead hatchlings/embryos. However, species identification Nonetheless, natal homing is not absolute, and female straying is was confirmed by analysis of maternal DNA found in fresh eggshell important for colonization of novel nesting habitats. Given the sum collected this nest. This exemplifies the power of this technique to of the evidence, it is plausible that the hawksbill female that nested in identify maternal identity of clutches lost during incubation because North Carolina, USA, originated from the nesting colony at Príncipe of environmental impacts and also to document nesting events by Island in the East Atlantic, representing a possible transoceanic rare species that might have otherwise been missed. colonization event by this species. However, not all hawksbill Acknowledgments. The NC Sea Turtle Project is supported by rookeries in the Atlantic have been genetically characterized, so the NC Wildlife Resource Commission Nongame & Endangered the possibility of nesting relocation within the Western Atlantic Wildlife Fund, the US Fish & Wildlife Service, the National Park cannot be ruled out. Service, Cape Hatteras National Seashore Resource Management Regardless of where the hawksbill female came from, these two staff, in addition to other federal, state, local, and private institutions nests are >800 km north of the nesting beaches in central Florida, and individuals. Funding for the genetic mark-recapture project which previously had been considered to be the northernmost nesting came from a NOAA Fisheries Species Recovery Grant. The authors area for this species in the western Atlantic (Meylan & Redlow thank Kelly Stewart for insightful comments and suggestions. 2006). Globally, these two nests are the furthest north (>35° north of BEYNETO, S. & E. DELCROIX. 2005. Underwater oviposition the equator) of any known hawksbill sea turtle nest (http://seamap. by a hawksbill turtle in Guadeloupe, French West Indies. Marine env.duke.edu/swot). Turtle Newsletter 107: 14. The two hawksbill nests in North Carolina were laid 53 days apart. The average inter-nesting interval for hawksbills is around 14 BOWEN, B.W., T.A. CONANT & S.R. HOPKINS-MURPHY. days (Kamel & Delcroix 2009), suggesting that this turtle may have 1994. Where are they now? The Kemp’s Ridley Headstart Project. laid up to three other clutches between the two nests documented Conservation Biology 8: 853-856. on Cape Hatteras. However, extensive sampling of all documented BOWEN, B.W., W.S. GRANT, Z. HILLIS-STARR, D.J. SHAVER, sea turtle clutches laid in Georgia, South Carolina, North Carolina K.A. BJORNDAL, A.B. BOLTEN & A.L. BASS. 2007. Mixed- and Virginia during 2015 did not reveal any other hawksbill nests stock analysis reveals the migrations of juvenile hawksbill turtles (CJN & BMS, unpublished data). Possible explanations for these (Eretmochelys imbricata) in the Caribbean Sea. Molecular missed nests include: the female migrated to and nested on a beach Ecology 16: 49-60. that is outside of the ongoing DNA capture-recapture study; the BROWNE, D.C., J.A. HORROCKS & F.A. ABREU-GROBOIS. turtle nested within the study area but the nesting crawls and eggs 2010. Population subdivision in hawksbill turtles nesting on were not sampled due to wind/rain that obscured them; the turtle Barbados, West Indies, determined from mitochondrial DNA expelled her eggs at sea (Beyneto & Delcroix 2005), or she nested control region sequences. Conservation Genetics 11: 1541-1546. only twice that year, 53 days apart. GORHAM, J.C., D.R. CLARK, M.J. BRESETTE, D.A. BAGLEY, Initially, when the nesting crawls were found, they were K.L. KESKE, S.L. TRAXLER, B.E. WITHERINGTON, categorized as being made by loggerheads. This is not surprising: B.M. SHAMBLIN & C.J. NAIRN. 2014. Characterization of both species have similar nesting crawls, and average egg diameter a subtropical hawksbill sea turtle (Eretmochelys imbricata) and average clutch size are similar for the two species (Hirth 1980; assemblage utilizing shallow water natural and artificial habitats van Buskirk & Crowder 1994). On nesting beaches used by both in the Florida Keys. PLOS One 9: e114171. loggerheads and hawksbills in the same season, species identification

Marine Turtle Newsletter No. 151, 2016 - Page 28 HIRTH, H. 1980. Some aspects of the nesting behavior and from North Carolina, with a summary of leatherback nesting reproductive biology of sea turtles. American Zoologist 20: activities north of Florida. Marine Turtle Newsletter 101: 4-8. 507-523. SERAFINI, T.Z., G.G. LOPEZ & P.L.B. DA ROCHA. 2009. Nest KAMEL, S.J. & E. DELCROIX. 2009. Nesting ecology of the site selection and hatching success of hawksbill and loggerhead hawksbill turtle, Eretmochelys imbricata, in Guadeloupe, French sea turtles (Testudines, Cheloniidae) at Arembepe Beach, West Indies from 2000-07. Journal of Herpetology 43: 367-376. northeastern Brazil. Phyllomedusa 8: 3-17. LARA-RUIZ, P., G.G. LOPEZ, F.R. SANTOS & L.S. SOARES. SHAMBLIN, B.M., B.F. FAITCLOTH, M.G. DODD, D.A 2006. Extensive hybridization in hawksbill turtles (Eretmochelys BAGLEY, L.M. EHRHART, P.H. DUTTON, A. FREY & C.J. imbricata) nesting in Brazil revealed by mtDNA analyses. NAIRN. 2009. Tetranucleotide markers from the loggerhead Conservation Genetics 7: 773-781. sea turtle (Caretta caretta) and their cross-amplification in other MEYLAN, A.B. & A. REDLOW. 2006. Eretmochelys imbricata marine turtle species. Conservation Genetics 10: 577-580. - hawksbill sea turtle. In: Meylan, P.A. (Ed.). Biology and SHAMBLIN, B.M., B.F. FAIRCLOTH, M.G. DODD, A. WOOD- Conservation of Florida Turtles. Chelonian Research Monographs JONES, S.B. CASTLEBERRY, J.P. CARROLL & C.J. NAIRN. 3: 105-127. 2007. Tetranucleotide microsatellites from the loggerhead sea MONZÓN-ARGÜELLO C., F. DELL’AMICO, P. MORINIÉRE, turtle (Caretta caretta). Molecular Ecology Resources 7: 784-787. A. MARCO, L.F. LÓPEZ-JURADO, G.C. HAYS, R. SCOTT, R. SHAMBLIN, B.M., M.G. DODD, K.L. WILLIAMS, M.L. FRICK, MARSH & P.L.M. LEE. 2012. Lost at sea: genetic, oceanographic R. BELL & C.J. NAIRN. 2011. Loggerhead turtle eggshells as a and meteorological evidence for storm-forced dispersal. Journal source of maternal nuclear genomic DNA for population genetic of the Royal Society Interface. 9: 1725-1732. studies. Molecular Ecology Resources 11: 110-115. MONZON-ARGÜELLO C., N.S. LOUREIRO, C. DELGADO, VAN BUSKIRK, J. & L.B. CROWDER. 1994. Life history variation A. MARCO, J.M. LOPES, M.G. GOMES & F.A. ABREU- in marine turtles. Copeia 1994: 66-81. GROBOIS. 2011. Príncipe island hawksbills: Genetic isolation of VARGAS, S.M., M.P. JENSEN, S.Y.W. HO, A. MOBARAKI, D. an eastern Atlantic stock. Journal of Experimental Marine Biology BRODERICK, J.A. MORTIMER, S.D. WHITING, J. MILLER, and Ecology 412: 345-354. R.I.T. PRINCE, I.P. BELL, X. HOENNER, C.J. LIMPUS, F.R. MONZÓN-ARGÜELLO, C., C. RICO, A. MARCO, P. LÓPEZ SANTOS & N.N. FITZSIMMONS. 2016. Phylogeography, & L.P. LÓPEZ-JURADO. 2010. Genetic characterization of genetic diversity, and management units of hawksbill turtles in eastern Atlantic hawksbill turtles at a foraging group indicates the Indo-Pacific. Journal of Heredity 2016: 199-213. major undiscovered nesting populations in the region. Journal of VILAÇA, S.T., P. LARA-RUIZ, M.A. MARCOVALDI, L.S. Experimental Marine Biology and Ecology 387: 9-14. SOARES & F.R. SANTOS. 2013. Population origin and historical PROSDOCIMI, L., I. BRUNO, L. DIAZ, V.G. CARMAN, D.A. demography in hawksbill (Eretmochelys imbricata) feeding and ALBAREDA & M.I. REMIS. 2014. Southernmost reports of nesting aggregates from Brazil. Journal of Experimental Marine the hawksbill sea turtle, Eretmochelys imbricata, in temperate Biology and Ecology 446: 334-344. waters of Argentina and evidence of hybrid origin supported by VILAÇA, S.T., S.M. VARGAS, P. LARA-RUIZ, É MOLFETTI, mitochondrial DNA analysis. Herpetological Review 45:1-5. S.C. REIS, G. LÔBO-HADJU, L.S. SOARES & F.R. SANTOS. PUTMAN, N.F., F.A. ABREU-GROBOIS, A.C. BRODERICK, C. 2012. Nuclear markers reveal a complex introgression pattern CIOFI, A. FORMIA, B.J. GODLEY, S. STROUD, T. PELEMBE, among marine turtle species on the Brazilian coast. Molecular P. VERLEY & N. WILLIAMS. 2014. Numerical dispersal Ecology 21: 4300-4312. simulations and genetics help explain the origin of hawksbill WITZELL, W.N. 1983. Synopsis of biological data on the hawksbill sea turtles in Ascension Island. Journal of Experimental Marine turtle Eretmochelys imbricata (Linneaus 1766). FAO Fisheries Biology and Ecology 450: 98-108. Synopsis 137, 78p. RABON, D.R., S.A. JOHNSON, R. BOETTCHER, M. DODD, M. WOODSON, H.M. & W.D. WEBSTER. 1999. Chelonia mydas LYONS, S. MURPHY, S. RAMSEY, S. ROFF & K. STEWART. (green sea turtle). Nesting distribution. Herpetological Review 2003. Confirmed leatherback turtle Dermochelys( coriacea) nests 30: 224-224.

Marine Turtle Newsletter No. 151, 2016 - Page 29 REPORT Meeting Report for the 2015 International Summit of Fibropapillomatosis: Global Status, Trends, and Population Impacts

Stacy Hargrove NOAA Fisheries SEFSC, 75 Virginia Beach Dr., Miami, FL 33149 USA (E-mail: [email protected])

The 2015 International Summit on Fibropapillomatosis (FP) was currently exceeding population growth rates in some intensively convened in Honolulu, Hawaii 11-14 June 2015. The purpose for monitored populations (e.g., Florida and Hawaii, USA and convening the 2015 Summit was to provide a forum to assess the Southern Great Barrier Reef stock Queensland, Australia) as status and trends of the disease globally and its demographic impact evidenced by increasing nesting trends despite the incidence of on sea turtles. To identify regions to be represented, the Steering FP in immature foraging populations. Committee conducted a Monkey Survey poll, and from the 47 6. Pathogens, hosts, and potential disease and environmental responses received, 6 broad regions of priority were selected and cofactors have the capacity to change; while we are having travel invitations extended to key scientists having longer-term data success now, there needs to be continued monitoring to detect and insights for each region. Important cornerstones for the 2015 changes in the distribution, occurrence, and severity of the Summit were two prior workshops held in Hawaii in 1997 and 1990, disease. 19 and 26 years ago. 7. While we do not have clear evidence to provide the direct link, Scientists invited to present at the 2015 Summit included: globally, the preponderance of sites with a high frequency of FP Alexandre Girard, Brian Stacy, Carlos Diez, Cecilia Baptistotte, tumors are areas with some degree of degradation resulting from Colin Limpus, Daniel Walsh, Jennifer Lynch, Karina Jones, Kyle Van altered watersheds. Watershed management and responsible Houtan, Llewellyn Ehrhart, Milani Chaloupka, Shawn Murakawa, coastal development may be the best approach for reducing the and Thierry Work. The participants engaged in discussions that spread and prevalence of the disease. resulted in the following conclusions: 8. Future research efforts should employ a multi-factorial ecological 1. Globally, FP has long been present in wild sea turtle populations approach (e.g., virology, parasitology, genetics, health, diet, - the earliest mention was in the late 1800s in the Florida Keys. habitat use, water quality, etc.) since there are likely several 2. FP primarily affects medium-sized immature turtles in coastal environmental cofactors involved in the expression of the foraging pastures. disease, which is still thought to be caused by a herpesvirus. 3. Expression of FP differs across ocean basins and to some degree 9. Minimum FP data collection in new areas should include: within basins. Turtles in the Southeast US, Caribbean, Brazil, individual identification (photo ID, PIT tags, etc.), standard and Australia rarely have oral tumors (inside the mouth cavity), measurements (length and weight), presence/absence of tumors, whereas they are common and often severe in Hawaii. Internal tumor severity, body condition, oral examination, method of tumors (on vital organs) occur in the Atlantic and Hawaii, but capture, and effort. only rarely in Australia. Liver tumors are common in Florida A workshop report including extended abstracts by each presenter but not in Hawaii. was published in August 2016. The report, Proceedings of the 2015 4. Recovery from FP through natural processes, when the affliction International Summit on Fibropapillomatosis: global status, trends, is not severe, has been documented in wild populations globally. and population impacts, can be accessed on-line at: 5. FP causes reduced survivorship, but documented mortality rates https://pifsc-www.irc.noaa.gov/library/pubs/tech/NOAA_Tech_ in Australia and Hawaii are low. The mortality impact of FP is not Memo_PIFSC_54.pdf or http://go.usa.gov/xDzWV

Marine Turtle Newsletter No. 151, 2016 - Page 30 OBITUARY Jack Bryan Woody (1932-2016)

Earl Possardt1 & Barbara Schroeder2 1U.S. Fish and Wildlife Service, Office of International Affairs, Falls Church, VA 22041 USA (E-mail: [email protected]); 2NOAA-NMFS, Office of Protected Resources, Silver Spring, MD 20910 USA (E-mail: [email protected])

Jack Woody, conservation hero, dear friend, and mentor to so many Among Jack’s greatest contributions to sea turtle conservation of us in the global sea turtle conservation community died in his was his leadership and collaboration with Mexican colleagues to save Albuquerque home on April 26, 2016. the Kemp’s Ridley from the very edge of extinction. Woody won the Woody, as he was fondly called, was an inspiration to many of us respect and admiration of the international marine turtle conservation in the sea turtle community. We remember his visionary, dynamic, community for his many conservation accomplishments and in 2013, and no nonsense leadership as the U.S. Fish and Wildlife Service’s the International Sea Turtle Society, representing the global sea turtle first National Sea Turtle Coordinator and the founder of the Service’s conservation community, expressed its appreciation for his efforts sea turtle program in the early 1980’s. During his long career with and accomplishments by recognizing him with its highest and most the USFWS (1969-1993) he initially worked with Native American esteemed honor: The Lifetime Achievement Award. tribes to advance conservation efforts on reservation lands and More than his many accomplishments, for those fortunate to be with various endangered species conservation projects such as the his friend or colleague, Woody was simply loved and admired for the Mexican wolf. He became fascinated and committed to sea turtle man he was. Well after his retirement from USFWS he encouraged conservation while working in Mexico early in his career and this and advised his sea turtle colleagues still on the front lines, while inspired him to establish a much needed sea turtle program within cheering our victories. We will miss his humor, authenticity, the USFWS. Woody focused urgently needed attention on critical goodness, and the unique determined spirit that conveyed confidence sea turtle conservation issues such as the looming extinction of the in the face of bureaucracy and other such impediments to sea turtle Kemp’s ridleys and the Mexican black turtle nesting population, conservation. and the unsustainable olive ridley harvest in Mexico. He fearlessly Perhaps Ken Dodd put it best in his tribute to Jack, concluding pushed for the National Marine Fisheries Service to mandate Turtle with these words: “As long as sea turtles crawl from the water, they Excluder Devices (TEDs) in the shrimp industry during the 1980 will sing his praise, even if there is a deafening silence and sadness and early 1990’s. Through Jack’s relentless and effective advocacy at his loss.” as well as vision and initiative he established the USFWS as a key force for sea turtle conservation domestically and internationally.

Jack Woody in Albuquerque, NM, looking out over his beloved Sandia Mountains. Photo by Ken Dodd.

Marine Turtle Newsletter No. 151, 2016 - Page 31 REPORT Meeting Report for the 1st Photo ID Workshop, 36th Annual Symposium on Sea Turtle Biology and Conservation, Lima, Peru, 29 February 2016

Stephen G. Dunbar1,2, Jillian Hudgins3 & Claire Jean4

1Protective Turtle Ecology Center for Training, Outreach, and Research, Inc. (ProTECTOR Inc.), Colton, CA, USA 92324 (E-mail: [email protected]); 2Marine Research Group, Department of Earth and Biological Sciences, Loma Linda University, Loma Linda, CA, USA 92350; 3The Olive Ridley Project, 8 Throton Ave., Lipson, Plymouth, UK, PL4 8RS (E-mail: [email protected]); 4Kelonia, Science Department, 46 rue du General de Gaulle, 97436 Saint-Leu, La Reunion, France (E-mail: [email protected])

At the 36th Annual International Sea Turtle Symposium on associated with both underwater and nesting beach photographs was Conservation and Biology (ISTS) held in Lima, Peru between 29 a recurrent theme among participants. February - 4 March 2016, the first sea turtle photo-ID (PID) workshop At the completion of the talks, participants were invited to join associated with the ISTS was held on Monday, 29 February at the one of three small discussion groups: those who were already using Universidad Científica del Sur. There were 41 participants in the PID in their projects; those wishing to begin PID in their projects; workshop, which was organized by Stephen G. Dunbar (workshop those who were interested in understanding more about the technical Chair), assisted by Jillian Hudgins and Claire Jean. and development of PID programming. One member of the IBEIS The workshop ran from 10:00 AM - 4:45 PM, with the Chair team was asked to join one of the three small discussion groups, welcoming participants, thanking the workshop sponsors, and while participants in each of these groups were asked to think about opening the session with comments on the history of sea turtle their “dreams and needs.” This was an opportunity for participants PID, which, according to George Balazs (pers. comm.), likely to seek advice on using PID in their projects, to think of what started with the work of Peter Bennett & Ursula Keuper-Bennett challenges they faced in implementing or using PID, to express ideas in the mid-1990s (Richardson et al. 2000). Workshop presentations they would like to see developed for PID users, and to provide a followed from Jillian Hudgins, Stephen G. Dunbar, Claire Jean, list of what would be most useful to them in a future PID system. Konstantinos Papafitsoros, Andy Estrada & Jess Williams, who all These ideas, challenges, dreams, and needs were recorded to be presented aspects of how PID was being used in their respective considered by the IBEIS team as they move forward in modifying sea turtle research projects. Workshop presentations recognized the the current zebra PID and Wildbook platforms, and developing the growing body of sea turtle PID literature (Schofield et al. 2004; turtle PID platform (Leslie et al. 2016). Schofield et al. 2008; Jean et al. 2010; Lloyd et al. 2012; Dunbar et al. 2014; Valdés et al. 2014; Dunbar & Ito 2015; Carpentier et al. 2016) that is helping to bring PID methods to the attention of Question the sea turtle research community as a valid way of monitoring 1 The PID workshop was informative and useful to me. populations over long periods of time. Common themes that came I learned several new things from the presentations out of the presentations were: the requirement for high quality 2 images repeatedly taken from approximately the same angles and made during the workshop. distances; the need to manipulate all photos with spots, lines, or 3 The presenters for the workshop were well chosen. polygons; the lack of analytical power within current PID programs; I believe that the workshop waswell organized and 4 the need for meta-data handling within PID systems; the lack of ran smoothly. connectivity between current PID programs which limits turtle From what I have learned, I would be interested in identifications across platforms; and how to engage the public to 5 increase the amount of high quality images from an area. including PID in my own sea turtle projects. Gilber Mechado presented material on the development and use From what I have learned, I believe a global PID 6 of the Pic4Turtle smartphone application for species identification system would be of benefit to my turtle projects. and sightings, followed by a combined presentation by Tanya I would be interested in attending another PID Berger-Wolf, Chuck Stewart & Jason Holmberg on the development 7 of the Image-Based Ecological Information System (IBEIS) workshop in the future. developed for PID and analyses of zebra photographs. While Berger- I would be interested in presenting my PID work at a 8 Wolf & Stewart presented an overview of the IBEIS zebra photo future PID workshop. ID program and some initial trials of the sea turtle PID system, I would like to talk more with one of the presenters or 9 Holmberg presented the photo database system called Wildbook. organizers about PID applications. The IBEIS team is currently working on modifying the zebra PID programming for use with sea turtle photos. The concept of a global Table 1. Likert-scale questions posed to participants as part sea turtle PID system with the ability to analyze large metadata sets of the post-workshop. Marine Turtle Newsletter No. 151, 2016 - Page 32 Response Quotation 1 “Let’s do this again!” 2 “More time for debate.” 3 “I would like to see more dmos of PID software during presentations” 4 “It would be helpful to receive more info on the speakers and forums prior to the workshop so that the questions can be covered.” 5 “I think a better place is needed to faciliate group discussions.” Table 2. All responses provided by participants for the Figure 1. Mean scores for each of the Likert scale workshop open-ended request for comments and suggestions. evaluation questions.

At the completion of the workshop, participants were provided a JEAN, C., S. CICCIONE, E. TALMA, K. BALLORAIN & J. workshop evaluation form comprising nine questions using a Likert BOURJEA. 2010. Photo-identification method for green and scale of 1 to 6 (1 = strongly disagree; 5 = strongly agree; 6 = not hawksbill turtles - first results from Reunion. Indian Ocean Turtle applicable; see Table 1), and one open-ended question requesting Newsletter 11: 8-13. comments on the workshop and recommendations/suggestions LESLIE, A., C. HOF, D. AMOROCHO, T. BERGER-WOLF, J. th for the subsequent PID workshop at the 37 ISTS. Responses of HOLMBERG, C. STEWART, S.G. DUNBAR & C. JEAN. 2016. participants were analyzed and mean scores for each of the nine The Internet of turtles. State of the World’s Sea Turtles Report Likert scale questions calculated (Fig. 1). Table 2 shows all of 11: 12-13. responses from participants to the final, open-ended request for LLOYD, J.R., M.A. MALDANADO & R. STAFFORD. 2012. comments and suggestions. Methods of developing user-friendly keys to identify Green Sea In summary, the first sea turtle PID workshop was an opportunity Turtles (Chelonia mydas L.) from photographs. International for sea turtle researchers and conservationists to exchange ideas Journal of Zoology Article ID 317568, 7 pages, 2012. and methods on current PID platforms and uses, as well as to doi:10.1155/2012/317568 provide feedback to computer programmers who are developing a sea turtle PID platform that may potentially be accessible in the RICHARDSON, A., L.H. HERBST, P.A. BENNETT & U.K. future to a global network of users. It was recommended by many BENNETT. 2000. Photo-identification of Hawaiian green of the participants that another PID workshop, or a full session, be sea turtles. In: Abreu-Grobois, F.A., R. Briseño-Dueñas, R. convened at the 37th ISTS. Márquez-Millán & L. Sarti-Martínez (Comps.). Proceedings of the Eighteenth International Sea Turtle Symposium. NOAA Acknowledgments. We kindly thank WWF International, The Ocean Technical Memorandum NMFS-SEFSC-436, p. 249. Foundation, SWOT, Kelonia, and ProTECTOR Inc. who invested time and funding to support this workshop. We also thank each SCHOFIELD, G., K.A. KATSELIDIS, P. DIMOPOULOS & J.D. one of the presenters and all participants for being involved with PANTIS. 2008. Investigating the viability of photo-identification this workshop. as an objective tool to study endangered sea turtle populations. Journal of Experimental Marine Biology and Ecology 360: 103- CARPENTIER, A.S., C. JEAN, M. BARRET, A. CHASSAGNEUX 108. & S. CICCIONE. 2016. Stability of facial scale patterns on green sea turtles Chelonia mydas over time: A validation for the use of SCHOFIELD, G., K.A. KATSELIDIS & J.D. PANTIS. 2004. a photo-identification method. Journal of Experimental Marine Assessment of photo-identification and GIS as a technique to Biology and Ecology 476: 15-21. collect in-water information about loggerhead sea turtles in Laganas Bay, Zakynthos, Greece. In: Mast, R.B., B.J. Hutchinson DUNBAR, S.G. & H.E. ITO. 2015. Picture perfect: photography & A.H. Hutchinson (Comps.). Proceedings of the 24th Symposium for hands-off turtle monitoring In: Mast, R.B., B.J. Hutchinson & on Sea Turtle Biology and Conservation. NOAA Tech Memo P.E. Villegas (Eds.). The State of the World’s Sea Turtles Report. NMFS-SEFSC-567. p. 152. 10: 10-11. VALDÉS, Y.A., J.A. RICARDO, F.B. TRELLES & O. ESPADA. DUNBAR, S.G., H.E. ITO, B.K, S. DEHOM & L. SALINAS. 2014. First assay of photo-identification in marine turtles’ nesting 2014. Recognition of juvenile hawksbills Eretmochelys imbricata population. Revista de Investigaciones Marinas 34: 43-51. through face scale digitization and automated searching. Endangered Species Research 26: 137-146.

Marine Turtle Newsletter No. 151, 2016 - Page 33 TRIBUTE Jack Woody’s Long-Term Influence on Sea Turtle Conservation

Thane Wibbels Department of Biology, University of Alabama at Birmingham, 1300 University Blvd., Birmingham, AL 35294-1170 USA (E-mail: [email protected])

The sea turtle world experienced a great loss with the recent passing Government sent Jack Woody (along with a couple of body guards of Jack Woody, but his influence will continue for generations due to for his protection) to personally tell the local shrimping groups his accomplishments, leadership, and his inspiration of a whole new when and why they had to use TEDs. Jack Woody’s Herculean era of sea turtle conservationists. My memories of his personality efforts regarding the implementation of TEDs and establishment of and impact revolve around the near extinction and recovery of a binational effort at Rancho Nuevo cannot be overstated. Those the Kemp’s ridley, but that was just one of his many impacts on efforts stabilized the Kemp’s ridley when it was near extinction conservation. The Kemp’s ridley story is a classic example of his and initiated its recovery. And while he used the Kemp’s ridley to great influence on sea turtle conservation. He was a champion push through the implementation of TEDs, he fully understood that at understanding what needed to be done to save an endangered TEDs would begin enhancing the recovery of all sea turtle species species, and then achieving major goals by cutting though what throughout U.S. coastal waters. often appeared to be unsurmountable politics and red tape. As the The Kemp’s ridley was just one of many facets in the Jack Woody USFWS Chief of Endangered Species for the southwestern U.S., conservation legacy. Many of his other conservation achievements he heard about the plight of the Kemp’s ridley and as he put it “So have been mentioned in previous tributes, but I have one anecdote to I snuck down to Mexico to check out the situation first hand”. As emphasize the point. We were at Jack Woody’s house in Albuquerque the Kemp’s ridley was heading toward extinction, he was the one talking to him about the history of Kemp’s ridley conservation and who realized that a binational effort on the primary nesting beach at we eventually left through the garage. Behind the door to the garage Rancho Nuevo might be the only way to give the ridley a fighting was a wrinkled map of Mexico pinned to the wall with numerous chance for survival. He used his political savvy to significantly multi-colored push pins ranging from northern to southern Mexico enhance logistical support for Rene Marquez’s group conducting and from Baja California to the Yucatan. He said, “Hey, you might conservation at Rancho Nuevo. When it became clear that the high like this”, and then he said, “these are all conservation projects I mortality of juveniles and adults was preventing the recovery of the supported or helped initiate in Mexico” (Figure 1). That was classic ridley, Jack Woody used the near extinction of the ridley to force the Jack Woody: the map was behind the door to the garage along with implementation of turtle excluder devices (TEDs) in U.S. waters. the broom and step stool. Jack Woody was never out for notoriety, He spearheaded an effort by USFWS, NOAA Fisheries, and an he was out to save species from extinction and preserve the natural NGO (Center for Marine Education) for TED implementation which environment. His personality and attitude toward conservation was turned into a major political battle. When the long and intense battle contagious. Due to his achievements and inspiration, his influence was finally won and it was time to announce the implementation on sea turtle conservation will carry on for generations. regulations to the fisheries groups in the Gulf of Mexico, the U.S.

Jack Woody sketch based on a scene from Jack Woody beside his map of Mexico. His impact on the 1981 PBS documentary, “The Heartbreak conservation extended well beyond Kemp’s ridley conservation Turtle.” During the scene, he was expounding and the implementation of turtle excluder devices. He mentioned on the absolute necessity of saving species from that each of the push pins in the map represented projects that he extinction. (Sketch by Sarah Adkins, University had supported or helped initiate in Mexico. of Alabama at Birmingham). Marine Turtle Newsletter No. 151, 2016 - Page 34 RECENT PUBLICATIONS This section is compiled by the Archie Carr Center for Sea Turtle Research (ACCSTR), University of Florida. The ACCSTR maintains the Sea Turtle On-line Bibliography: (http://st.cits.fcla.edu/st.jsp). It is requested that a copy of all publications (including technical reports and non-refereed journal articles) be sent to both: The ACCSTR for inclusion in both the on-line bibliography and the MTN. Address: Archie Carr Center for Sea Turtle Research, University of Florida, PO Box 118525, Gainesville, FL 32611, USA. The Editors of the Marine Turtle Newsletter to facilitate the transmission of information to colleagues submitting articles who may not have access to on-line literature reviewing services.

RECENT PAPERS Boyle, M., L. Schwanz, J. Hone & A. Georges. 2016. Abdelrhman, K.F.A., G. Bacci, C. Mancusi, A. Dispersal and climate warming determine range shift in model Mengoni, F. Serena & A. Ugolini. 2016. A first insight into reptile populations. Ecological Modelling. 328: 34-43. M. Boyle, the gut microbiota of the sea turtle Caretta caretta. Frontiers in Univ Canberra, Inst Appl Ecol, Canberra, ACT 2601, Australia. Microbiology. 7: 1060. A. Mengoni, Univ Florence, Dipartimento (E-mail: [email protected]) Biol, Sesto Fiorentino, Italy. (E-mail: [email protected]) Braz, J.K.F.S., M.L. Freitas, M.S. Magalhaes, M.F. Anonymous. 2016. Conservation laws need reshaping to protect Oliveira, M.S.M.O. Costa, N.S. Resende, N.K. sea turtles. Marine Pollution Bulletin. 108: 1. Clebis, N.B. Silva & C.E.B. Moura. 2016. Histology and immunohistochemistry of the cardiac ventricular structure in the Anonymous. 2016. Mediterranean loggerhead turtles dying in green turtle (Chelonia mydas). Anatomia Histologia Embryologia. waters off the Middle East and North Africa. Marine Pollution 45: 277-284. J.K.F.S. Braz, Univ Fed Rio Grande do Norte, Dept Bulletin. 107: 4-5. Morphol, CP 1524,Campus Univ Lagoa, BR-59072970 Natal, Anonymous. 2016. Study proves removing beach debris RN, Brazil. (E-mail: [email protected]) increases sea turtle nests. Marine Pollution Bulletin. 108: 2. Brei, M. A. Perez-Barahona & E. Strobl. 2016. Baker, L., W. Edwards & D.A. Pike. 2016. Sea turtle Environmental pollution and biodiversity: Light pollution and sea rehabilitation success increases with body size and differs among turtles in the Caribbean. Journal of Environmental Economics and species. Endangered Species Research. 29: 13-21. D.A. Pike, Management. 77: 95-116. A. Perez-Barahona, INRA, Econ Publ Centre for Tropical Environmental & Sustainability Science, UMR210, Ave Lucien Bretignieres, F-78850 Thiverval Grignon, James Cook University, Cairns, Queensland 4870, Australia. France. (E-mail: [email protected]) (E-mail: [email protected]) Burns, T.J., H. Davidson & M.W. Kennedy. 2016. Large- Barco, S., M. Law, B. Drummond, H. Koopman, C. scale investment in the excavation and “camouflaging” phases by Trapani, S. Reinheimer, S. Rose, W.M. Swingle & nesting Leatherback Turtles (Dermochelys coriacea). Canadian A.S. Williard. 2016. Loggerhead sea turtles killed by vessel Journal of Zoology 94: 443-448. M. W. Kennedy, Univ Glasgow, and fishery interaction in Virginia, USA are healthy prior to Inst Biodivers Anim Hlth & Comparat Med, Coll Med Vet & Life death. Marine Ecology Progress Series. 555: 221-234. S. Barco, Sci, Graham Kerr Bldg, Glasgow G12 8QQ, Lanark, Scotland. Dept Biology & Marine Biology, University of North Carolina (E-mail: [email protected]) Wilmington, 601 S. College Road, Wilmington, NC 28403, USA. Caillouet, C.W. Jr., B.J. Gallaway & N.F. Putman. (E-mail: [email protected]) 2016. Kemp’s ridley sea turtle saga and setback: novel analyses Blackwood, K. 2016. Giving green sea turtles the green light. of cumulative hatchlings released and time-lagged annual nests Frontiers in Ecology and the Environment. 14: 179. in Tamaulipas, Mexico. Chelonian Conservation & Biology. 15: Block, B.A., C.M. Holbrook, S.E. Simmons, K.N. 115-131. C.W. Caillouet, Jr., Montgomery, Texas 77356 USA. Holland, J.S. Ault, D.P. Costa, B.R. Mate, A.C. Seitz, (E-mail: [email protected]) M.D. Arendt, J.C. Payne, B. Mahmoudi, P. Moore, Candan, O. & D. Kolankaya. 2016. Sex ratio of green turtle J.M. Price, J. Levenson, D. Wilson & R.E. Kochevar. (Chelonia mydas) hatchlings at Sugozu, Turkey: higher accuracy 2016. Toward a national animal telemetry network for aquatic with pivotal incubation duration. Chelonian Conservation & observations in the United States. Animal Biotelemetry 4: 6 DOI Biology. 15: 102-108. O. Candan, Ordu Univ, Fac Arts & Sci, 10.1186/s40317-015-0092-1. S.E. Simmons, Marine Mammal Dept Biol, Gen Biol Sect, 52200 Cumhuriyet Campus, Ordu, Commission, Bethesda, USA. (E-mail: [email protected]) Turkey. (E-mail: [email protected]) Boura, L., S.S. Abdullah & M.A. Nada. 2016. New Chapman, P.A., R.J. Traub, M.T. Kyaw-Tanner, H. observations of sea turtle trade in Alexandria, Egypt. Report to Owen, M. Flint, T.H. Cribb & P.C. Mills. 2016. Terminal MEDASSET, Mediterranean Association to Save the Sea Turtles. restriction fragment length polymorphism for the identification 27 pp. MEDASSET - Mediterranean Association to Save the of spirorchiid ova in tissues from the green sea turtle, Chelonia Sea Turtles, 1c Licavitou St., 106 72 Athens, Greece. (E-mail: mydas. PLoS One. 11(8): e0162114. P.A. Chapman, Veterinary- [email protected]) Marine Animal Research, Teaching and Investigation Unit, School

Marine Turtle Newsletter No. 151, 2016 - Page 35 of Veterinary Science, The University of Queensland, Gatton, Museo Zool, AP 70-399, Mexico City 04510, DF, Mexico. Queensland, Australia, (E-mail: [email protected]) (E-mail: [email protected]) Clusa, M., C. Carreras, M. Pascual, S.J. Gaughran, Foran, D.R. & R.L. Ray. 2016. Mitochondrial DNA profiling of S. Piovano, D. Avolio, G. Ollano, G. Fernandez, J. illegal tortoiseshell products derived from hawksbill sea turtles. Tomas, J.A. Raga, A. Aguilar & L. Cardona. 2016. Journal of Forensic Sciences. 61: 1062-1066. D. R. Foran, Potential bycatch impact on distinct sea turtle populations Michigan State Univ, Dept Integrat Biol, 655 Auditorium Rd,560 is dependent on fishing ground rather than gear type in the Baker Hall, E Lansing, MI 48824 USA. (E-mail: [email protected]) Mediterranean Sea. Marine Biology 163: 122. DOI:10.1007/ Fuentes, M.M.P.B., C. Gredzens, B.L. Bateman, R. s00227-016-2875-1 M. Clusa, Univ Barcelona, Dept Anim Biol, Boettcher, S.A. Ceriani, M.H. Godfrey, D. Helmers, Av Diagonal 643, E-08028 Barcelona, Spain. (E-mail: m.clusa@ D.K. Ingram, R.L. Kamrowski, M. Pate, R.L. Pressey hotmail.com) & V.C. Radeloff. 2016. Conservation hotspots for marine Collareta, A., M. Bosselaers & G. Bianucci. 2016. turtle nesting in the United States based on coastal development. Jumping from turtles to whales: a Pliocene fossil record depicts Ecological Applications DOI 10.102/eap.1386/abstract. M.M.P.B. an ancient dispersal of Chelonibia on mysticetes. Rivista Italiana Fuentes, Dept of Earth, Ocean and Atmospheric Science, Florida De Paleontologia e Stratigrafia. 122: 35-43. A. Collareta, Via State University, Rm 507 OSB, 117 North Woodward Avenue, Santa Maria 53, I-56126 Pisa, Italy. (E-mail: alberto.collareta@ Tallahassee, FL 32306-4320, USA. (E-mail: [email protected]) for.unipi.it) Fuentes, M.M.P.B. & V.S. Saba. 2016. Impacts and effects of Crear, D.P. D.D. Lawson, J.A. Seminoff, T. Eguchi, R.A. ocean warming on marine turtles. In: Laffoley, D. & J.M. Baxter LeRoux & C.G. Lowe. 2016. Seasonal shifts in the movement (Eds.). Explaining Ocean Warming: Causes, Scale, Effects and and distribution of green sea turtles Chelonia mydas in response Consequences. Full Report. Gland, Switzerland: IUCN. pp. 289- to anthropogenically altered water temperatures. Marine Ecology 302. (Address as above) Progress Series. 548: 219-232. D.P. Crear, College William & Fujisaki, I. & M.M. Lamont. 2016. The effects of large beach Mary, Virginia Institute Marine Science, POB 1346, Gloucester debris on nesting sea turtles. Journal of Experimental Marine Point, VA 23062 USA. (E-mail: [email protected]) Biology and Ecology. 482: 33-37. I. Fujisaki, Univ Florida, Ft da Silva, P.F., M.F. Chaves, M.G. Santos, A.J.B. Santos, Lauderdale Res & Educ Ctr, 3205 College Ave, Davie, FL 33314 M.D. Magalhaes, R. Andreazze & G.J.B. de Moura. USA. (E-mail: [email protected]) 2016. Insect infestation of hawksbill sea turtle eggs in Rio Grande Fukuoka, T., M. Yamane, C. Kinoshita, T. Narazaki, do Norte, Brazil. Chelonian Conservation & Biology. 15: 147-153. G.J. Marshall, K.J. Abernathy, N. Miyazaki & K. G.J.B. de Moura, Univ Fed Rural Pernambuco, Dept Biol, Recife, Sato. 2016. The feeding habit of sea turtles influences their PE, Brazil. (E-mail: [email protected]) reaction to artificial marine debris. Scientific Reports. 6: 28015. de Carvalho, R.H., N. Mamede, R.R. Bastos & B.M. T. Fukuoka, Atmosphere and Ocean Research Institute, The de Sousa. 2016. Attitudes towards conservation and fishing University of Tokyo, Kashiwa, Chiba, 277-8564, Japan. (E-mail: interaction with sea turtles in the southeast coast of Brazil. Ocean [email protected]) & Coastal Management. 127: 55-62. R.H. de Carvalho, Univ Fed Gonzalez Carman, V., A. Mandiola, D. Alemany, Juiz de Fora, Dept Zool, Inst Ciencias Biol, Cidade Univ, BR- M. Dassis, J.P. Seco Pon, L. Prosdocimi, A. Ponce 36036900 Juiz de Fora, MG, Brazil. (E-mail: robsonhc1000@ de Leon, H. Mianzan, E.M. Acha, D. Rodriguez, M. yahoo.com.br) Favero & S. Copello. 2016. Distribution of megafaunal Marcovaldi, M.A.G., M. Lopez-Mendilaharsu, species in the Southwestern Atlantic: key ecological areas A.S. Santos, G.G. Lopez, M.H. Godfrey, F. Tognin, and opportunities for marine conservation. ICES Journal of C. Baptistotte, J.C. Thome, A.C.C. Dias, J.C. de Marine Science. 73: 1579-1588. V.G. Carman, UNMdP, IIMyC, Castilhos & M.M.P.B. Fuentes. 2016. Identification CONICET, Mar Del Plata, Buenos Aires, Argentina. (E-mail: of loggerhead male producing beaches in the south Atlantic: [email protected]) Implications for conservation. Journal of Experimental Marine Grafeld, S., K. Oleson, M. Barnes, M. Peng, C. Chan, & Biology and Ecology. 477: 14-22. M.M.P.B. Fuentes, Florida M. Weijerman. 2016. Divers’ willingness to pay for improved State Univ, Dept Earth Ocean & Atmospher Sci, Tallahassee, FL coral reef conditions in Guam: An untapped source of funding 32306 USA. (E-mail: [email protected]) for management and conservation? Ecological Economics. 128: Eagle, L., M. Hamann & D.R. Low. 2016. The role of social 202-213. S. Grafeld, University of Hawaii - Manoa, Department of marketing, marine turtles and sustainable tourism in reducing Natural Resources & Environmental Management, 1910 East West plastic pollution. Marine Pollution Bulletin. 107: 324-332. Road, Honolulu, HI 96822 USA. (E-mail: [email protected]) M. Hamann, James Cook Univ, Coll Marine & Environm Sci, Hackenburg, D. 2016. It runs in the blood: How sea turtles Townsville, Qld 4811, Australia. (E-mail: mark.hamann@jcu. respond to interactions with fishermen. Coastwatch. Summer edu.au) 2016: 21-25. www.ncseagrant.ncsu.edu/coastwatch Flores-Villela, O., K. Adler & T.G. Eimermacher. Hargrove, S., T. Work, S. 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Post-nesting loggerhead turtle around dawn, returning to the sea on Iztuzu Beach, Dalyan, a Specially Protected Area for sea turtles in Turkey. For more detail, see Whitmore et al. MTN 50: 6-8. Photo: Matthew H. Godfrey.

Marine Turtle Newsletter No. 151, 2016 - Page 38 INSTRUCTIONS FOR AUTHORS

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