CSIRO PUBLISHING www.publish.csiro.au/journals/wr UlildlifeResearch,2009,36,637-645

Habitat use by green turtles (Chelonia mydas) nesting in Peninsular : local and regional conservation implications

Jason Paul van de MerweA.B,G, Kamarruddin IbrahimC,D, Shing Yip Lee A andJoan Margaret WhittierE,F

AGriffith School of Environment and Australian Rivers Institute, Griffith University (Gold Coast), Griffith University, QLD 4222, Australia. Bpresent address: Centre for Marine Environmental Research and Innovative Technology, City University of Hong Kong, Hong Kong SAR, China. CTurtle and Marine Ecosystems Centre, Department of Fisheries Malaysia, , , Malaysia. Dpresent address: Marine Park Department of Malaysia, Ministry of Natural Resources and Environment, Federal Government Administration Centre, Putrajaya, Malaysia. ESchool of Biomedical Sciences, University of Queensland, St Lucia, QLD 4067, Australia. Fpresent address: School of Medicine, Locked Bag 24, University ofTasmania, Sandy Bay, Tas. 7005, Australia. GCorresponding author. Email: [email protected]

Abstract Context. Many green sea turtle (Chelonia mydas) populations are declining worldwide owing to their susceptibility to human impacts in the marine environment. Identifying the habitats used throughout different lifecycle stages is therefore important for managing the interactions between turtles and humans. Aims. To identify the habitat utilisation of a C. mydas nesting population in during breeding, inter-nesting, migration and foraging lifecycle stages. Methods. Satellite telemetry was used to track the movement ofthree C. mydas nesting females and one adult male from the Ma'Daerah rookery (Peninsular Malaysia). Key results. The male and female turtles remained within 30 km ofthe nesting beach during the breeding and inter-nesting periods, which includes habitat beyond the 'no trawl zone' designed to protect turtles in this area. Following the breeding season, the tracked turtles migrated up to 1955 km to four ditTerent foraging grounds in Vietnam, Indonesia, Peninsular Malaysia and Bomeo Malaysia. During foraging, turtles occupied areas threatened by human activities such as fishing and pollution. Conclusions. The habitats used by the Ma'Daerah C. mydas population during breeding are outside current local protection zones and extend into unprotected international waters during migration and foraging. Implications. Identification of habitats used by C. mydas populations is a critical element of management and conservation of this endangered, migratory species. Our study highlights the need to increase offshore protection around Ma'Daerah during the nesting season. Furthermore, this study has identified the countries within South-east Asia that Malaysia must cooperate with to ensure effective management ofthis C. mydas population. This infonnation is particularly relevant to sea turtle conservation and management in regions like South-east Asia, where many coastal countries occupy a small geographical area.

Additional keywords: breeding, foraging, migration, satellite telemetry.

Introduction result, many sea turtle populations are declining worldwide Sea turtles inhabit a range of oceanic and neritic environments and all species (except the flatback (Natator depresslls), which throughout their lifecycle (Miller 1997). Within these habitats, is listed as 'Data Deficient') have been listed as 'Endangered' sea turtles are exposed to anthropogenic impacts, such as or 'Critically Endangered' by the International Union for incidental capture in fishing gear, boat strikes and harvesting Conservation of Nature (IUCN 2007). Sea turtles are also (Hutchinson and Simmonds 1992; Poiner and Harris 1996). As a listed in Appendix 1 of the Convention of International Trade

© CSIRO 2009 10.1071/WR09099 1035-3712/09/070637 638 Wildlife Research J. P. van de Merwe et al.

of Endangered Species of Wild Fauna and Flora (CITES). areas (Bowen et al. 1992; Fitzsimmons et al. 1996). This Understanding habitat utilisation of sea turtles is therefore implies that each breeding C. mydas population must be important for managing and conserving endangered populations. treated as an independent management unit when considering The Ma'Daerah Turtle SanctuaJ)', on the east coast of threats and conservation. In addition, effective management of Peninsular Malaysia, supports a large breeding population of a breeding population requires knowledge of the threats at green sea turtles (Chelonia myda.v). However, this population the different foraging grounds and along the migration routes. has declined by >SO% since the 1950s, primarily owing to the This is complicated by the fact that foraging areas can be long collection of eggs for human consumption, nesting beach distances from the breeding areas and are often located in other development and the incidental capture of turtles in fishing countries with varying conservation ethics, regulations and gear (Ibrahim 1994). The current management of this resources. population involves the use of hatcheries on the nesting Satellite telemetry has previously been used to track the beaches to protect the eggs from collection and a 'no trawl movement of C. mydas in foraging habitats and along zone', which extends 5.5 nautical miles (-10 km) offshore migration routes (see Tables I, 2). However, there is limited from the nesting sites (Fisheries Act 1985, Malaysia) to information in the literature on the habitats used by C. mydas protect turtles in the marine environment. However, there is for breeding (courtship and mating) or the areas used by nesting currently no information on habitats used by this C. mydas females between laying subsequent clutches during the nesting population during breeding, inter-nesting, migration and foraging. season (inter-nesting habitat). FUlihermore, comprehensive C. mydas adults spend the majority of their lives in neritic studies encompassing all of the important habitats of a single foraging habitats, such as coastal seagrass meadows and nesting population are limited. The aim ofthe present study was reefs (Musick and Limpus 1997; Bjomdal et al. 2005). Sexually therefore to investigate the breeding, inter-nesting, migration mature C. mydas migrate seasonally to breeding grounds, which and foraging areas used by the Ma'Daerah C. mydas nesting are in their natal areas (Meylan et al. 1990; Bowen et al. 1992). population. This new information will assist in the management Females can remain in the nesting area for up to 6 months while and conservation of this endangered C. mydas population. they lay I-11 clutches at 2-week intervals (Miller 1997). During their reproductive years, C. myda.\· adults show strong fidelity to Materials and methods these foraging and breeding sites, which can be up to thousands of kilometres apart (Carr and Carr 1972; Limpus et al. 1992; Satellite transmitter attachment Lohmann et al. 1997). C. mydas individuals from a single Between 20 August and 9 September 2005, satellite foraging area may migrate to several different breeding areas, transmitters were attached to three C. mydas females nesting and a breeding population is often populated by individuals at the Ma'Daerah Toole Sanctuary (4°32'17"N, 103°2S'14"E) from a wide range of foraging areas (Liew et al. 1995; Cheng and one adult male captured in the cooling tanks of the YTL 2000; Godley et al. 2002; Seminoffet al. 200S). Therefore, threats Corporation's power station (4 C 35'26"N, J03°27'7"E) in Paka, to C. mydas in a foraging area can potentially affect several Terengganu, Peninsular Malaysia (Table 3). Kiwi Sat (0.5 W) different breeding populations and a single breeding population Platform Terminal Transmitters (PTTs) powered by two can be influenced by threats in several different foraging areas. lithium C cells (Sirtrack, Havelock North, New Zealand) were The strong fidelity to natal rookeries results in genetically attached to the carapace using methods modified from distinct populations being established around major breeding Balazs et al. (1996).

Table 1. Summary of the distance (km), days tracked and speed (km h-') of Chelonia mydas post-nesting migrations, determined by satellite tracking studies N/A, not available

Nesting site n Distance Days Speed Reference

Ascension Island 5 1792-2346 33-47 1.9-2.8 Luschi et al. (1998); Hays et af. (1999) Ascension lslanda 2 N/A N/A 2.2-2.3 Hays et al. (2001 b) Japan 4 966--1962 28-42 1.4-2.1 Hatase et al. (2006) Taiwanb 8 193-1909 7-57 1.2-2.8 Cheng (2000) Hawaii/Samoa 5 110~1750 26-45 1.6-1.9 Balazs et ai. (1994) Hawaii 3 830-1260 22-26 1.6--2.0 Balazs (1994) Hawaii" 3 105~1200 23·30 1.7-1.9 Balazs and Ellis (2000) Costa Rica 10 4101470 1127 0.9-2.5 Troeng et ai. (2005) b d Cyprus • 6 3222199 843 1.62.1 Godley et al. (2002) Galapagos Islandsd 12 12171912 2652 1.12.2 Seminotf et al. (2008) Peninsular Malaysiab 5 669-1744 13-30 1.7-2.6 Liew et al. (1995) Peninsular Malaysia 4 310-1955 15-45 0.9-2.4 Present study

aMale C. mydas only. ~ot all migrations completed to foraging areas. clncludes male C. mydas. dOverall inter-nesting, migration and foraging movement. Conservation implications of green turtle habitat use Wildlife Research 639

Table 2. Summary of foraging home ranges of Chelonia mydas in previous studies SCL, straight carapace length

Site Turtles Mean Foraging area Reference 2 SCL (cm) range (km )

Mosquito Lagoon, FL. USA 9 <65 0.48~5.06 Mendonya (1983) Kaneohe Bay, HI, USA 12 51.3 2.62 ±0.96" Brill et al. (1995) South Padre Island, TX, USA 9 34.5 0.22-3.1 I Renaud et al. (1995) Repulse Bay. Australia 10 105.4 0.848.50 Whiting and Miller (1998) Gulf ofCalifornia, Mexico 12 66.7 5.8439.08 Seminoffet al. (2002) Palm Beach, FL, USA 6 36.7 0.695.05 Makowski et al. (2006) Peninsular Malaysia 3 96.8 24-3575 Present study

aMean (±s.d.) movement of individuals from the point of release (km).

Table 3. Morphology and nesting details of the Chelonia mydas that were satellite tagged in AugustlSeptember, 2005 at the Ma'Daerah Turtle Sanctuary, Tcrengganu Malaysia N/A, not available

Turtle A Turtle B Turtle C Turtle D

Date captured 20/8/05 20/8/05 28/8/05 9/9/05 Sex Female Female Female Male Curved carapace length (cm) 100 95 lOS 90.5 Curved carapace width (em) 94.7 90 95 77.7 Mass (kg) liS 110 124 75 Previous nests 25/6/05 (104) No record 3/6/05 (90) N/A Date (# eggs) -6/7/05a 15/6/05 (91) -1817105a -28/6/05a a -2917105 1017105 (95) -9/8/05a 22/7/05 (97) 4/8/05 (98) -16/8/05"

aNesting dates estimated based on recorded nests and the II·· IS day inter-nesting period for this population ofC. mydas.

At the time of transmitter attachment, morphological removed from the dataset owing to their unreliability (Hays et al. measurements were taken according to Bolten (1999), and 200Ia). Route maps and movement within nesting and foraging nesting records for the season were consulted to determine grounds were generated using the Satellite Tracking and Analysis any previous nesting activity of the tracked turtles Tool (Coyne and Godley 2005) and Maptool (SEATURTLE. (see Table 3). Of the three nesting females, two had nested ORG 2002). A speed filter of 5 km h-I between successive previously (A and C) and one had no nests recorded for this points was applied to eliminate any biologically unreasonable season (B). Although Tmile A had laid only one previous data. Minimum convex polygons, joining the outermost nest, recorded on 25 June 2005, the II to IS day inter-nesting points of an individual's foraging, breeding or inter-nesting interval for this population (K. Ibrahim, pers. comm.) suggests range, were generated with the Animal Movement extension that this individual laid a total offive clutches before the date of for ArcView 3.2 (ESRI, Redlands, CAl. As well as providing transmitter attachment. Similarly, Turtle C had most likely laid location data, these transmitters also recorded sea surface two more clutches between the five clutches recorded before temperature (±1°C). Mean temperatures were calculated for the date oftransmitter attachment. Although there was no record breeding, migration and foraging habitats of each turtle. of Turtle B nesting before transmitter attachment, it is possible that this individual had previously nested undetected in the area Results this season. Location data from the satellite transmitters identified inter-nesting and breeding movement, post-nesting Tracking and mapping migration and movement within home foraging grounds Location information for each turtle was obtained from the Argos (Table 4). Two of the turtles (A and C) began post-nesting satellite system, a joint venture between the Centre National migration immediately following transmitter deployment. The d'Etudes Spatiales (CNES, the French space agency), the remaining two turtles (B and D) remained within 30 km of the National Aeronautics and Space Administration (NASA, USA) nesting beach for 41 and II days, respectively (Fig. I), before and the National Oceanic and Atmospheric Administration beginning post-nesting/breeding migration to foraging areas. (NOAA, USA) (Argos 1996). Position fixes were received in During the inter-nesting period, the nesting female (B) laid seven classes (3, 2, I, 0, A, B, Z) and classes 0, Band Z were three more clutches at Ma'Daerah (Table 3) and remained 640 Wildlife Research J. P. van de Merwe et al.

Table 4. Inter-nesting, breeding, post-nesting and foraging ground movement details ofthree female (A, Band C) and one male (D) Chelonia mydas released from the Ma'Daerah Turtle Sanctuary, Terengganu, Malaysia N/A, not available

Turtle A Tuttle B Tuttle C Turtle D

Breeding/Nesting Nesting date (# eggs) Nile 4/9/05 (118) Nile N/A 18/9/05 (93) 30/9/05 (96) Days in breeding area N/A 41 N/A 11 2 Area occupied" (km ) N/A 78.5 N/A 398 Mean SSTb (s.d.) N/A 29.2 (0.8) N/A 28.5 (0.4) Locations per day" N/A 0.8 N/A 2.9 Migration Distance travelled (km) 1955 1330 1590 310 Days travelled 45 31 28 15 d I Ave. Speed (km h- ) 1.81 1.79 2.37 0.86 Final destination Sabah, Borneo Malaysia Thousand Islands, Indonesia Ly Son, Vietnam Pemanggil Island, Malaysia Mean SSTb (s.d.) 29.8 (0.8) 303 (0.5) 29.2 (0.8) 29.1 (0.4) Locations per dayc 1.4 1.4 4.6 1.9 Foraging Days foraging N/Af 16 35 43 2 Area occupied" (km ) N/A 91 24 3575 Mean SSTb (s.d.) N/A 30H 27.3 (0.6) 29.3 (0.4) Locations per dayc N/A 0.6 1.6 1.8

"Minimum convex polygons, joining the outermost points ofan individual's range. "Sea surface temperature (DC). cMean number of Argos location hits (classes 3, 2, I, A) received per day. dCalculated as the total distance divided by number ofdays travelled. "Turtles A and C began post-nesting migration immediately following transmitter attachment. f'rurtle A had not reached foraging area before transmitter stopped transmitting,

103°24' 103°28' 103°32' 103°36' 103°40' 103°44'

4°40'

4°36'

4°32'

4°28'

5km

Fig. J. Breedinglinter-nesting habitat used by the adult male, Tuttle D (0)and a nesting female, Turtle B C...) Chelonia mydas near the Ma'Daerah Turtle Sanctuary, Terengganu, Malaysia. Dashed line indicates the outer limit ofthe 'no trawl' zone. Inset map: Peninsular Malaysia. Conservation implications of green turtle habitat use Wildlife Research 641 within 10 km of the coastline. The male turtle (D) Discussion remained further than 8 km from the coastline in an area SatelIite telemetry of C. mydas at the Ma'Daerah Turtle of 320 km2 (Fig. 1). Sanctuary has identified some important habitats used by Each of the four turtles migrated towards a different this population in Peninsular Malaysia and the surrounding foraging ground within South-east Asia (Fig. 2) and the South-east Asian region. Although the male (D) was not distance, duration and speed of migration varied between observed mating after release, its curved carapace length individuals (Table 4). The male turtle (D) migrated at a (90.5 em) was within the range of adult males observed 1 much slower speed than the females (0.86 km h- ), and breeding in other C. mydas populations (Limpus 1993; there was a distinct loop in its migration route, which was Balazs and ElIis 2000). Furthermore, its subsequent 310-km not observed for the females. The straight-line movement of migration to a known foraging area indicated that this male Turtle A when the PTT stopped transmitting indicated was most probably in the area for breeding. The adult male that this individual was likely to be still migrating and had (D) spent the majority of time outside the current 'no trawl not reached its foraging ground (see Fig. 2). However, zone', which extends 5.5 nautical miles (~1O km) offshore the haphazard movements of turtles B, C and D within (Fisheries Act 1985, Malaysia). This indicates that this shallow coastal habitats at the end of migration routes C. mydas nesting population may be exposed to threats such indicated that these individuals had reached their foraging as incidental capture in fishing gear and boat strikes during grounds (Figs 3-5). this period. These results can be used to recommend the extent

102° 105° 108° 111 ° 114° 117°

•••••• A -8 --c 15°

12°

go

6°

3°

0°

500 km

Fig. 2. The post-breeding migration ofthree Chelonia mydas females (A, Band C) and one C. mydas male (D) released from the Ma'Daerah Turtle Sanctuary, Terengganu, Malaysia. 642 Wildlife Research J. P. van de Merwe et al.

106"32' 106°36' 106'40' 106"44' -5°38' ~ Pcdamaran Island Thousand Islands (INDONESIA) ,, , •, " , ,f JAVA ~/ -5"'40' C7 '''II SEA • ,,",/' , ~ ,J>o"'---~\ , ' _... ':--1;'- ~-,-/---

Kolok Island " ,,' 17" '" ~'

.~---_... _... - CD -5"44' 3km

Fig. 3. SatelJite position data ofa Chelonia mydas adult female (B) within a foraging ground in the Thousand Islands (Indonesia), folJowing post-nesting migration from the Ma'Daerah Turtle Sanctuary, Terengganu, Malaysia. General area corresponds to the end ofthe migration route ofTurtle B (see Fig. 2).

r--_1..;;0..;.9°..;;0..;;.6'__~~__1..;;0:.:.9-=°0..;;.8_'__~ 1..;;0..:..9_01..;;.0_' 1;.:;.;;09°12' Dao Ly Son Marine Protected Area (VIETNAM)

15°24'

15 0 23'

150 22'

SOUTH CHfNA 15°21' SEA 2km

Fig. 4. Satellite position data ofa Chelonia mydas adult female (C) within a foraging ground near Ly Son Island (Vietnam), folJowing post-nesting migration from the Ma'Daerah Turtle Sanctuary, Terengganu, Malaysia. General area corresponds to the end ofthe migration route of Turtle C (see Fig. 2).

of offshore protection required for adequate management of behaviour because C. mydas males are known to migrate at the Ma'Daerah C. mydas breeding population. comparable speeds to females following breeding (Hays et at. Post-nestinglbreeding movement of C. mydas males and 200Ib). females has identified important migration routes from the The migration to different foraging grounds within South­ Ma' Daerah Turtle Sanctuary to foraging grounds in South­ east Asia has conservation implications for this Peninsular east Asia. Migration speed and distance travelled from the Malaysian C. mydas breeding population. The Thousand Ma'Daerah rookery to foraging grounds was quite variable Islands (foraging ground for Turtle B) lie directly to the north among individuals, although within the ranges observed for ofthe Indonesian capital city, Jakarta, and are under threat from other C. mydas populations (Table 1). Interestingly, the sediment and land-based pollution generated by a human male turtle (D) migrated at a slower average speed population of >20 million (UNESCO 1997). Similarly, Ly Son I (0.86 km h- ) and there was a distinct loop about halfway Island (foraging ground of Turtle C) is 25 km off the central through migration (see Fig. 2). This was unexpected Vietnamese coast and, although a Marine Protected Area, the Conservation implications of green turtle habitat use Wildlife Research 643

104°00' 104°20' 104°30' 104°40' Pemanggil Island (MALAYSIA)

2°50' SOUTH CHINA SEA ,, ,,

/

10km 2°00'

Fig. 5. Satellite position data of a Chelonia mydas adult male (D) within a foraging ground near Pemanggil Island (Malaysia), following migration from the Ma'Daerah Turtle Sanctuary, Terengganu, Malaysia. General area corresponds to the end of the migration route ofTurtle 0 (see Fig. 2). marine environment is under threat from destructive fishing range of the largest C. mydas foraging ranges previously practices and agricultural activities on the island and adjacent recorded (Seminoff et al. 2002), Turtle B and the adult male mainland (ADB 1999). Furthermore, a similar study in 1993 and (D) of this study had unusually large foraging ranges 1994 on C. mydas nesting at the nearby Redang Island rookery (Table 4). Previous studies found the home ranges of foraging (-150 km away) indicated there are at least three other foraging C. mydas adults to be associated with small abundant patches of grounds that would be within the migration range of the macroalgae and seagrass food resources. Furthermore, the small Ma'Daerah breeding population and outside the direct control areas used during feeding have been attributed to maximising ofPeninsular Malaysian authorities (Liew et al. 1995). A larger efficiency of energy expenditure and maintaining familiarity of scale satellite telemetry study would therefore be required to surroundings to assist in avoidance of predators and identify further foraging sites and highlight the areas and environmental extremes (Makowski et at. 2006). However, countries in the region that Peninsular Malaysian authorities comparison to these previous studies should be made with would need to collaborate with to effectively manage this caution owing to the considerable differences in turtle size, C. mydas population. duration of tracking and physical characteristics of the Satellite telemetry has also identified the habitat utilisation foraging areas. of C. mydas individuals within their foraging grounds A study on the C. mydas foraging population in the Gulf following migration. As with inter-nestinglbreeding of Mexico suggested that the larger home ranges observed movement, the male (D) occupied a much larger area during were due to the large area of this site and scattered distribution 2 2 2 foraging (3575 km ) than the two females (24 km and 91 km ). of food resources (Seminoff et al. 2002). These factors could 2 Although the area covered by Turtle C (24 km ) was within the account for the large foraging areas observed in the present study. 644 Wildlife Research J. P. van de Merwe et al.

Furthermore, foraging areas could be influenced by the quality of Bjorndal, K. A., Bolten, A. B., and Chaloupka, M. Y. (2005). Evaluating seagrass habitats in these areas. Lowerdensity seagrass meadows trends in abundance of immature green turtles, Chelonia mydas, in the and high degree of patchiness would require more extensive Greater Caribbean. Ecological Applications 15, 304-314. doi: 10.1890/ movement within a foraging area. Turtle B was found to forage in 04-0059 the waters offtheThousand Islands in Indonesia that coveran area Bolten, A. B. (1999). Techniques for measuring sea turtles. In 'Research and Management Techniques for the Conservation ofSea Turtles, Vol. 4'. of-750 km 2 and the adult male (D) foraged around a large group (Eds K. L. Eckeli, K. A. Bjomdal, F. A. Abreu-Grobois and M. Donnelly.) ofislands just offthe east coast ofPeninsular Malaysia covering pp. 119-123, (IUCN/SSC Marine Turtle Specialist Group: 2 an area of4800 km . Although the specific location and quality of Arlington, VA.) the macroalgae and seagrass meadows are not known in these Bowen, B. W., Meylan, A. B" Ross, J. P., Limpus, C. J., Balazs, G, H., and locations, these island congregations probably provide shallow Avise, J. C. (1992). Global population structure and natural history ofthe coastal areas with suitable foraging habitat, separated by deeper, green turtle (Chelonia mydas) in ternlS of matriarchal phylogeny. less productive waters. These extensive ranges ofwell separated Evolution 46, 865-881. doi: 10.2307/2409742 food resources could account for the relatively large areas covered Brill, R. W., Balazs, G. H., Holland, K. N., Chang, R. K, C., Sullivan, S., and during foraging by the C. mydas adults in this study. Furthermore, George, J, C. (1995), Daily movements, habitat use, and submergence intervals of normal and tumour-bearing juvenile green turtles (Chelonia none of the literature to date has tracked the movement of mydas L.) within a foraging area in the Hawaiian Islands. Journal of C. mydas adult males in a foraging area. Without the energetic Experimental Marine Biology and Ecology 185, 203-218. doi: 10.1016/ demands of preparing for and performing nesting (Kwan 1994; 0022-0981 (94)00146-5 Hamann et al. 2002), adult males may be able to travel further in Carr, A., and Carr, M. H. (1972). Site fixity in the Caribbean green turtle. search ofhigher quality food sources. Ecology 53, 425-429, doi: 10.2307/1934228 Cheng, \.-J. (2000). Post-nesting migrations ofgreen turtles (Chelonia mydas) at Wan-An Island, Penghu Archipelago, Taiwan. Marine Biology 137, Acknowledgements 747-754. doi: 10.1007/s002270000375 Coyne, M. S., and Godley, B. J. (2005). Satellite Tracking and Analysis Tool The Department of Fisheries, Malaysia provided logistical support for (STAT): an integrated system for archiving, analysing and mapping attaching transmitters to C. mydas in Peninsular Malaysia and supplied animal tracking data. 301, 17. two of the PTT transmitters. Funding for the remaining two transmitters Marine Ecology Progress Series doi: I0.3354/meps301 001 and all the Argos satellite data was supplied by an Earthwatch/Vodafone Fitzsimmons, N. N., Moritz, C., Limpus, C. J., Miller, J. D., Parmenter, C. J., Foundation agreement. Thanks also to all Earthwatch field staffandvolunteers and Prince, R. (1996). Comparative genetic structure of green, on the 'Green Turtles ofMalaysia' project who helped with data collection. loggerhead and ftatback populations in Australia based on viable Special thanks to Michael Coyne from SEATURTLE.ORG for supplying user mtDNA and nONA regions. 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