((Roeody/Us Porosus) from the Philippines

AcknowledgmeIlIS.-We would foremost like to thank Jorge Lar­ collaboration reveals encouraging status for the severely deplet­ gaespada for his dedication to this work. We gratefully thank Osa ed population of hawksbill turtles Eretmochelys imbricata. Oryx Conservation for providing key provisions for the study, Osa Adven­ 44:595-601. turas, EI Tigre Fund, Walter Aguirre Aguirre, Juan Diego Morales Cam­ --, R. L. LEWISON, I. L. YA"JEZ, B. P. WALLACE, M. j. LILES, W. J. NICHOLS, bronero, Marvin Villalabos Palma, and Dan Hughes for field support, A. BAQUERO, C. R. HASBIlN, M. VASQUEZ, j. UIlTEAGA, AND J. A. SEMINOFF. and Jorge Cortes, Pilar Bernal, and Didher Chacon for valued insigh t. 2012. Shifting the life-history paradigm: discovery of novel habitat This project was funded by a Greg Gund Memorial Fellowship. use by hawksbill turtles. BioI. Lett. 8:54-56. HAZEL, I., I. R.IAWLLR, lIND M. HAMANN. 2009. Diving at the shallow end: green turtle behavior in near-shore foraging habitat. I. Exp. Mar. LITERATtmE CrrED BioI. EcoJ. 371 :84-92. MANCINI, A., AND V. KOCH. 2009. Sea turtle consumption and black ALVARADO-DIllS, J., ,11'0 L. FIGUEHOA. 1990. The ecological recovery of sea market trade in Baja California Sur, Mexico. Endang. Species Res. turtles of Michoacan, Mex.ico. Special attention: the black turtles, 7:1-10. Chelollia agassizii. Final report 1989-1990, U.S. Fish & Wildl.ife MARQUEZ, R., C. S. PENIII'LORES, A. O. VILLANUEVA, AND I. F. DIAZ. 1982. A Service, Silver Spring, Maryland. 139 pp. model for diagnosis of populations of olive ridJeys and green tur­ AMOROCHO, D. E, F. A. Anm:u-GHoBoIs, P. H. DUTION, AND R. D. REINA. 2012. tles of West Pacific tropical coasts. [n K. A. Bjorndal (ed.), Biology Multiple distant origins for green sea turtles aggregating off Gor­ and Conservation of Sea Turtles, pp. 153-158. Smithsonian Insti­ gona Island in the Colombian Eastern Pacific. PLoS ONE 7:e31486. tution Press, Washington, D.C. --, AND R. D. REINA. 2007. Feeding ecology of the East Pacific green MORALES- RAMfHEZ, A. 20 II. La diversidad marina del Golfo Dulce, Paci­ sea turtle Chelonia mydas at Gorgona National Park, Colombia. fico sur de Costa Rica: amenazas a su conservacion. Biocenosis Endang. Species Res. 3:43-51. 24:9-20. BESSESEN, B. 2012. Geospatial and behavioral observations of a unique QUESADA-ALPfzAR, M. A., I. CORTEs-NIlJ';Fz, I. I. ALVARADO, AND A .c. FON­ x.anthic colony of pelagic sea snakes, Pelamis platurus, residing in SECA. 2006. Caracteristicas hidrograficas y bio16gicas de la zona Golfo Dulce, Costa Rica. Herpetol. Rev. 43:22-26. marino-costera del Area de Conservacion Osa. Serie Tecnica: BIOHNDAL, K. A. 1980. Nutrition and grazing behavior of the green tur­ Apoyando los esfuerzos en el manejo y proteccion de la biodivers­ tle, Chelonia mydas. Mar. BioI. 56: 147-154. idad tropical. The Nature Conservancy, San jose, Costa Rica. 79 pp. CHM:(lN, D., D. ROltIS, A. BARASH, AND C. QUESADA. 2011. New Pacific RICHARD, I. D., AND D. A. HUGHES. 1972. Some observation of sea turtle green turtle foraging ground at Dulce Gulf, south Pacific coast of nesting activity in Costa Rica. Mar. BioI. 16:297-309. Costa Rica. 31" Annual Symposium on Sea Turtle Biology & Con­ SEMINOFF, I. A., A. RESENDIZ, AND W. I. NICHOLS. 2002. Diet of the east Pa­ servation, San Diego, California, poster presentation. cific green turtle, Chelonia mydas, in the central Gulfof California, COHTI'S, J. 200 I. Requiem for an eastern Pacific seagrass bed. Rev. BioI. Mexico. J. Herpeto!. 36:447-453. Trop. 49:273-278. SENKO, I., V. KOCH, W. M. MEGILL, R. R. CARTHY, R. P. TEMPLETON, AND W. I. --, ,1:>10 E. SALAS. 2009. Seagrasses. [n I. S. Wehrtmann, and J. Cor­ NICHOLS. 2010. Fine scale daily movements and habitat use of East tes (eds.), Marine Biodiversity of Costa Rica, Central America, pp. Pacific green turtles at a shallow coastal lagoon in Baja California 119-122. Monographiae Biologicae, Springer Science, Berlin. Sur, Mexico. I. Exp. Mar. BioI. Ecol. 391:92-100. DEVAUX, B., AND B. DEWETIEH. 2000. On the Trail of Sea Turtles. Barron's, SVENDSEN, H., R. ROSI.AND, S. MYKING, I. A. VARGAS, O. G. LiltINO, AND E. Hauppauge, New York. 128 pp. I. ALFARO. 2006. A physical-oceanographic study of Golfo Dulce, DRAKE, D. 1996. Marine turtle nesting, nest predation, hatch frequen­ Costa Rica. Rev, BioI. Trop. 54:147-170. cy, and nesting seasonality on the Osa Peninsula, Costa Rica. Che­ TOFT, R. 2009. Osa: Where the Rainforest Meets the Sea. Zona Tropical Ion. Conserv. BioI. 2:89-92. Publications, Costa Rica. 222 pp. GilOS, A. R., E A. ABREU-GnoBoIs, J. ALFARO-SHIGUETO, D. AMOROCHO, R. WEHHTMANN, I. S., I. COIlTES, ,~ND S. ECHEVERR(A-SAENZ. 2009. Perspectives ARAUZ, A. BAQUERO, R. BRiSE/w, D. CHACON, C. DUENAS, C. HASBON, M. and Conclusions. [n I. S. Wehrtmann and I. Cortes (eds.), Marine LILES, G. MARIONA, C. MUCCIO, J .P. MUNOZ, W. J. NICHOLS, M. PEJ"iA, J. Biodiversity of Costa Rica, Central America, pp. 521-527. Mono­ A. SEMINOFI', M. VASQUEZ, J. URTL-IGA, B. WALLACE, I. L. YA.~EZ, AND P. graphiae Biologicae, Springer Science, Berlin. ZARATE. 2010. Signs of hope in the eastern Pacific: international

Herpetological Relliew, 2012,43(4),541-546. ij) 20 12 by Society for (he S(udy of Amphibians and Repliles Here be a Dragon: Exceptional Size in a Saltwater Crocodile ((roeody/us porosus) from the Philippines ...... Crocodiles (Reptilia: Crocodylia) are generally regarded as the ADAM R. C. BRITTON* largest living reptiles by mass (Britton 2003). Exceptionally large Big Gecko Crocodilian Research, PO Box 1281, Howard Springs, individuals regularly attract mainstream attention, often en­ Northern Territory 0822, Australia; hanced by unrealistic and exaggerated accounts of their size and Research Institute ofEnvironment and Livelihoods, behavior. Actual evidence on the upper size limit of crocodiles Charles Darwin University, Darwin, Northern Territory 0909, Australia is lacking and romantic accounts and misleading photographs ROMULUS WHITAKER are often the only remains. Skulls found in museums and private NIKHIL WHITAKER collections provide a tantalizing glimpse into the size of their Madras Crocodile Bank Trust, Post Bag No.4, Mamallapuram 603104, former owners, yet total lengths are usually not available, the Tamil Nadu, India techniques used to obtain them are not known, and the verac­ *Corresponding author; e-mail: [email protected] ity of such measurements is questionable {Greer 1974; Whitaker

Herpetological Review 43(4),2012 542 ARTICLES

and Whitaker 2008). Limited verification is possible using exist­ were found fastened around the crocodile's upper jaw. Dozens of ing formulae for predicting total length (TL) from skull measure­ people were reportedly required to haul the crocodile onto the ments such as dorsal cranial length (DCL; e.g., Verdade 1999; bank where his jaws and legs were bound using ropes (E. Elorde, Webb and Messe11978; Wu et al. 2006). However, such formulae pers. comm.). Key PWRCC staffwho were not present on the cap­ are typically derived from a subset of possible size ranges, and ture night provided additional guidance by mobile phone. Over become increasingly inaccurate if applied outside those ranges 100 people were required to haul the crocodile by cart along the to the largest examples of a species. Not only do crocodilian body river bank to the nearby village and a floating pontoon had to be parts show allometric change with increasing size (Gans 1980), constructed to float the crocodile across the creek to access the the rate and direction of change is not constant (Hall 1985; Hall only available road. The crocodile was then transferred to nearby and Portier 1994) and there is increasing variability particular­ holding facilities and now resides at Bunawan Eco-Park and Re­ ly near the maximum upper size limit (Whitaker and Whitaker search Center. 2008). As there is a dearth of data available from thorough and There was considerable international interest in the total systematic measurements of very large crocodiles, our ability to length of Lolong. The crocodile which killed Rowena Romano predict TL from other variables including DCL is compromised was estimated by a witness to be "30 ft" (9.1 m) in length, whereas near the upper limit. Such information is useful across a number the crocodile seen hunting carabao near the village shortly be­ of disciplines including paleontology, conservation and popula­ fore capture was suspected to be closer to "16 ft" (4.9 m) long (E. tion management (Brochu 2001; Schmidt-Nielsen 1984). Elorde, pers. comm.). His true size only became apparent during We present accurate measurements for an exceptionally capture. Media reports of his length were inconsistent, ranging large Saltwater Crocodile (Crocodylus porosus Schneider 1801) from 6.1 to 6.4 m (20-21 ft). It was not known at the time how that was captured alive from Mindanao in the Philippines. We accurately the crocodile had been measured, what method had compare these measurements with those of other exceptionally been used, or whether the figure had been reported correctly. large crocodiles and discuss factors responsible for exceptional However, photographic and video evidence showed his size to be size. exceptional. This was sufficient to attract serious interest in veri­ Background.-The capture of an exceptionally large male fying the measurement and Natural History New Zealand dis­ C. porosus from a small creek not far from Bunawan in the Agu­ patched one of the authors (AB) to visit Bunawan and indepen­ san del Sur province of Mindanao in the Philippines made in­ dently measure the crocodile where it would be witnessed and ternational headlines on 3 September 2011. This crocodile was documented. Funding was provided by National Geographic. captured after a two-year effort to remove the animal thought responsible for the death of 12-year-old Rowena Romano in Lake MATEHIALS AND METHODS Mihaba in Agusan Marsh on 7 March 2009 (Sistante 2009) and later in mid-2011 the unreported disappearance of a fisherman Capture and restraint.-One of the authors (AB) measured close to Bunawan village (R. Sumiller, pers. comm.). An experi­ Lolong in his enclosure at the Bunawan Eco-Park on 9 November enced member of the Palawan Wildlife Rescue and Conserva­ 2011 at 1400 h local time. Measurement conditions were consid­ tion Center (PWRCC), Ernesto "Lolong" Canete, died of a heart ered ideal. Lolong was lying relatively straight on the level con­ attack in late August 2011 shortly before the crocodile's eventual crete floor of a recently-drained pool. A combination of chemi­ capture. The crocodile was named "Lolong" in his memory (R. cal and physical restraint was employed to facilitate accurate Sumiller, pers. comm.). measurement, to improve safety for personnel, and to ensure Lolong's capture was coordinated by wildlife specialist Ron­ the crocodile's welfare. Chemical restraint was achieved using nie Sumiller and other PWRCC members. Steel nooses baited pancuronium bromide, the dose (2 x 2 mg, 40 minutes apart) in­ with Carabao (Bubalus bubalis carabanesis) meat were deployed tended to provide temporary partial immobilization (after Bates hanging from trees along the creek. The first four nooses were et al. 2004), plus diazepam (10 mg, single dose) to provide tem­ destroyed by the crocodile's struggling before a fifth, thicker­ porary mild sedation. Both drugs were injected 1M into opposite cabled noose was successful. The remains of the other nooses sides of the tail. As Lolong was outside the size range of croco­ diles for which either drug had previously been tested, we used TMII.F I. Measurements oflolong. See Materials and Methods for the highly conservative dose rates-the minimum required dosage difference between Method Aand Method B. Method Ais considered to permit safe handling without compromising handler welfare. the principal measmemenl. All measurements are illustrated in Fig. Ventilation rate and eye blink response were monitored through­ 2. All length measurements are taken from tip of premaxilla (snout). out the procedure. All width measurements are taken at the widest point. Induction took two hours, at which point the crocodile was Measurement (mm) Method A Method B capable only of lethargic movements of the head, limbs, and tail tip. At this point standard physical restraint was employed: the Dorsal craniallenglh (DCL) 700 694 head was secured with a controlling rope noose on the upper jaw, a second noose around both jaws to close them remotely Snout-eye length (SEL) 496 495 through tightening, and thick damp towels over the eyes to re­ Maximum head width (MHW) 450 450 duce visual reactions. At this point a five-person team engaged Maximum cranial width (MOV) 228 228 the crocodile from behind, applying downward pressure to the Inter-orbital width (lOW) 84 84 top of the head, the pectoral and pelvic regions, and the body Cranial height (CHj 363 363 and tail, while lifting each limb from the ground to limit trac­ Snout-pelvis length (SPL) 2851 2844 tion (Fig. 1). The jaws were secured shut using duct tape, with Snout-scute junction (S5]) 4982 4949 the nostrils free to permit normal breathing. At this point, han­ Total length (TL) 6170 6095 dling and close-quarters measurements were considered safe. Measurement activities lasted approximately one hour, after

Herpetological Review 43(4),2012 ~ ;:l ~ ~" :< ~ ~ >­ ~

riG. I. Lateral view of Lolong under restraint during measurement procedure.

(a) Tal •

..... - - I ~v ~-.'.', \V' _ _ "'1""'10 I ' ~ -t , : :, :, . . , ). , , , SPl (B) .., : TaLI (B) j Tal2 (B) : :.. -:.. -:.. -: '. : : _ : ,al27 (A) : , . , , , , , : '

(b) ... MHW(NB) • ... Mew {AlB) .,

... Del (82 •

n :r:: ,.~ OJ o n r - .,..,.e::;:----- _ >

FIG. 2. Top-down and profile diagrams of entire crocodile (al and head (b) illustrating measurements taken using Method A(Al and Method B (Bl. DCL = dorsal cranial length; SEL = snout-eye length; MHW = maximum head width; MCW = maxi­ mum cranial width; row = inter-orbital width; CH = cranial height; SPL = snout-pelvis length; TaL = tail length; TaLl = an­ tprior tail length; TaL2 = posterior tail length; SPL+TaLl = snout-scute junction (SS]); SPL+TaLl +TaL2 = total length (TL).

Herpetological Review 43(4), 2012 which all physical restraint was carefully removed using estab­ with TaL to TL ratio of 1:1.9. Snout-scute junction (SS)) was 4.98 lished safety protocols. A reversal agent for pancuronium bro­ m (16.34 ft), therefore we calculated anterior tail length (Tall) as mide (e.g., neostigmine methylsulphate) was not used and the 2.13 m (6.99 ft) and posterior tail length (TaL2) as 1.19 m (3.90 ft). drugs were allowed to metabolize naturally while the crocodile The ratio TaL2 to TaL is 1:2.8. Measurements using Method Bare was under observation. Lolong was exhibiting normal behavior provided for comparison (Table 1). All measurements differed by six hours post-injection, moving around his enclosure with ease. less than 1% between Methods Aand BexceptTL, which differed Measurement.-Lolong's head and tail were straightened by 1.2%. Lolong's net mass (accounting for the trailer weight) carefully before any measurement took place. He maintained was 1075 kg (2370 Ibs), this posture throughout the procedure. Measurements concen­ General body condition was good, with normal muscle tone trated on head dimensions and body length variables that were and fat deposits typical of a large wild crocodile. Age cannot be unlikely to be influenced by body condition in any way (Figs. determined by physical examination, but there were no obvious 2a, 2bl. unlike for example belly width. All measurements were signs of senescence. Age was estimated subjectively at approxi­ taken using a flexible but non-stretchable steel tape with a mil­ mately 50 to 60 years based on indicators such as level of skull limeter scale. Conversion into other units (e.g., feet) was done rugosity, health of teeth alveoli, height of dorsal scute keels, and after the fact. The tape was held taut for each measurement and overall appearance, which can be compared with prior observa­ where necessary straight wooden rods held perpendicular to the tions. Very little historical scarring was found on the skin, and no ground were used as boundary markers for specific measures limbs, toes, or parts of the tail were missing, which was consid­ (e.g., maximum head width; Fig. lJ. ered unusual for such a large wild crocodile. The ventral surface There is no clear agreement on a "standard" method for mea­ was not examined. Recent superficial injuries evident in a hand­ suring crocodilian dimensions. We generally followed methods ful of locations (pads of feet, flanks, dorsal osteoderms, snout) previously published for C. porosus (Webb and Messel 1978) appeared to have been caused by rope abrasion incurred while slightly modified based on our experience on practical tech­ struggling during capture and transport. All recent injuries ap­ niques with wild crocodiles. We used DCL specifically for "head peared to be healing normally; almost all scab tissue had fallen length" to avoid any confusion with skull measurements that off, revealing fresh, non-pigmented or partially-pigmented skin have occasionally confused mandible length with head length tissue. (Whitaker and Whitaker 2008). Skull widths and heights were DISCUSSION taken at the maximum points. Snout-pelvis length (SPL, Fig. 2a), to the posterior margin of the hind legs joining the pelvis, was Lolong's TL, at 6.17 m (20.24 ft), makes him the largest wild used as a substitute for snout-vent length (SVL) because it was crocodile ever captured alive. The TL measurement taken by Ed­ highly impractical to roll such a large crocodile onto its back and win Elorde shortly after capture (6.4 m, 21 ft) was slightly higher there were concerns about the welfare implications of doing so than the one obtained here. Although measurement inaccuracy with such a heavy crocodile on concrete. Previous comparisons may be a factor, it is also likely a result of the crocodile's pos­ (A. Britton, pers. obs.) suggest that SPL and SVL differ by less ture when it was strapped to a cart with the tail hanging onto the than 1%. ground. The weight of the tail muscle would be sufficient to pull To address possible disagreement over the most suitable vertebrae apart, enough to account for most of the additional 20 method for body length measurements, two methods were used cm (7.9 in) length, underlining the need for measurements to be for comparative purposes (Table I, Fig. 2a). Method A used a taken with a crocodile lying on flat, level ground. steel tape along the dorsal midline from snout tip to tail tip fol­ There are a number of reports ofwild-caught crocodiles with lowing the slope of the skull and latter part of the tail. Method greater TL, but these are either difficult to verify or are from in­ B used the horizontal distance from snout tip to tail tip exactly complete specimens (e.g., dried skins; Whitaker and Whitaker parallel to the ground. In practice Method B gave fractionally 2008). While it is improbable that Lolong is the largest C. porosus shorter lengths than Method A (by around 1% in large individu­ ever found, it is disappointing that the evidence for larger speci­ als) and yet Method A is more frequently used (A. Britton and R. mens is so incomplete. This evidence comes almost entirely from Whitaker, pers. obs.l. simply because it is more practical espe­ skulls residing in museums and personal collections around the cially with large individuals that are difficult to move. Measure­ world. Data from their original owners are often lacking, with TL ments using Method A are therefore more relevant because they either not recorded or impossible to verify, and even the country allow more meaningful comparison with measurements of other of origin is doubtful in some cases. However, it is possible to use large crocodiles. It should be noted that Lolong's tail tip did not known relationships between each DCL and TL to estimate the appear to be damaged. total length of the crocodile from which each skull was derived. Lolong's mass was recorded at a nearby truck weigh-bridge Unfortunately, the only thorough analysis to date for C. porosus during transport to his captive holding facilities on 3 September uses data from relatively small hatchlings up to sub-adults and 2011. As it was logistically impractical to verify his mass during the formulae were obviously never intended to be extrapolated our visit, his reported mass on that date is provided here. There for much larger animals (Webb and MesseJ 1978). It is appar­ is no reason to believe that rigor in this simple measurement was ent from the limited data available that skull shape is subject to lacking and we considered it valid. greater variation with increasing TL, DCL becoming a smaller proportion of TL with increasing size. The DCL:TL ratio for RESULTS smaller crocodiles has long been thought to be around 1:7 (Bel­ lairs 1970; Greer 1974; Wermuth 1964) but this would under-es­ Using Method A, Lolong's total length (TL) was 6.17 m (20.24 timate Lolong's length by over a meter (3.3 ftl. The DCL:TL ratio ft; Table lJ, with a dorsal cranial length (DCL) of 700 mm (27.6 for large C. porosus is closer to 1:9 (Whitaker and Whitaker 2008). in). Ratio of DCL to TL was 1:8.8. Snout-pelvis length (SPL) was Therefore, caution must be used when estimating TL based on 2.85 m (9.35 ft). Therefore tail length (TaL) was 3.32 m (10.89 ftl. DCL of existing museum skull specimens.

Herpetological Review 43(4), 2012 The largest known C. porosus skull is housed at the Paris potentially the result of heterosis (or "hybrid vigor"; Shull 1948). Museum (MNHN PMP specimen #A11803 = old museum col­ In June 2012 Guinness World Records pronounced Lolong to be lection #7738) originally from Cambodia. It has a DCL of 760 the largest crocodile in captivity (Guinness World Records 2012). mm (29.9 in), making it 8.6% longer than Lolong's skull. If we To address the question of why extremely large crocodiles are apply a DCLTL ratio of 1:9 for this skull, TL is estimated at 6.84 so rare, it is instructive to look at factors responsible for growth m (22.4 ft) which is 11.3% longer than Lolong's TL. Although the and size. An early analysis suggested that males cease growth actual TL was never preserved, these figures strongly suggest a from 3.9 to 6.0 m (12.8 to 19.7 ft; Webb et al. 1978), yet it is widely nearly 7 m (almost 23ft) crocodile. We can compare this with known that environmental and social factors that influence ac­ another slightly smaller skull (currently in the private collection cess to required resources (e.g., desired temperatures, access to of Shivendra Narayan Bhanja Deo, the Yuvaraj of Kanika in Bhu­ water, access to basking sites, food availability and quality, and baneshwar, Orissa) of DCL 730 mm (28.7 in), originally from the social stress) can have a significant impact on health and growth Indian Bhitarkanika province, reported to have come from a 7 rates (Coulson and Hernandez 1983; Dalrymple 1996; Lang 1987; m (23ft) C. porosus. If those figures are true then DCL:TL ratio Webb 1985). Crocodilians in captive situations are highly sen­ would be 1:9.5, sufficiently high for minor skepticism but still sitive to stress and the availability of resources, which have a feasible. Applying the 1:9 ratio to the Bhitarkanika skull gives an marked effect on growth rate (e.g., Choudhury and Bustard 1983; estimated TL of 6.6 m (21.7 ft). The truth is unlikely to be far from Lance 1994; Smith and Marais 1994). We also know that genetic these figures and there is a strong sense that 7 m (23 ftl is likely factors playa significant role in post-hatching growth and be­ the maximum possible length for C. porosus. havior, as does the incubation environment (Allsteadt and Lang After Lo)ong, the best documented evidence of a record-sized 1995; Isberg et al. 2004). crocodile comes from Obo village on the Fly River in Papua New Growth rate in crocodiles is based on a von Bertalanffy Guinea (Montague 1983). This crocodile drowned in a fishing net growth curve, being fastest in juveniles and progressively slower set for barramundi fish and after 50 men hauled the crocodile in adults. The greatest proportion ofgrowth occurs in the first 15­ onto the bank they found an entire Rusa Deer (Cervus timoren­ 25 years, to around three-quarters of maximum size, although sis) carcass in the stomach. The crocodile's skin had already been faster-growing individuals appear to sustain rapid growth for removed and salted when Jerome Montague and one of the au­ longer (Webb et al. 1978). Slow-growing juveniles become small­ thors (RW) visited the village, but the skin plus decapitated head er adults, growth rates falling too low to ever achieve very large measured 6.2 m (20.3 ft). The authors considered this likely an sizes. In such individuals growth essentially stops in older ani­ underestimate considering possible shrinkage of the skin plus mals (Webb et al. 1978; Woodward et al. 1995). Conversely it is an incomplete tail tip, suggesting a TL closer to 6.3 m. The DCL the fast-growing juveniles that reach the largest adult sizes and of this crocodile was 720 mm (28.3 in), which at 6.2 m TL would grow for longer. It is incorrect to assume that extremely large indicate a DCL:TL ratio of I :8.6, or I:8.8 considering the likely 6.3 crocodiles are necessarily the oldest crocodiles. For example, the m TL. While not a complete or living specimen, this is still con­ hybrid Yai reached 6 m (19.8 ft) in less than 30 years and a pure­ sidered the largest C. porosus ever measured and documented. bred C. porosus ("Jaws") at Madras Crocodile Bank reached 5.13 There are several unverified reports of even larger wild croco­ m (16.8 ft) in 38 years. Most wild C. porosus take at least 15 years diles. the most popular being a 7 m plus (over 23ft) C. porosus to reach 3.5 m (Webb and Manolis 1989). sighted within the Bhirtarkanika Wildlife Sanctuary in Orissa, In order to achieve maximum growth rates and size in the India, in 2006 (Whitaker and Whitaker 2008). However, this was wild, crocodiles require greater access to optimal environmen­ not a measurement but a size estimate taken from a boat and tal conditions particularly when young, access to all necessary regardless of the skill of the observers it cannot be compared resources when desired, and minimal growth-inhibiting stress. to a verified tape measurement, especially considering the un­ Given the role that stress plays in inhibiting growth, it is likely certainty inherent in visual size estimation in the wild (Bayliss that exceptional size can only be achieved when crocodiles do 1987). Another famous giant crocodile shot on the Norman Riv­ not encounter humans on a regular basis. Lolong, for exam­ er in Australia in 1957 was reported by the shooters to be over ple, appears to have spent his entire life in and around Agusan 8 m (approx. 26ft). While it seems likely that an exceptionally Marsh, an area that sees very low human traffic due to difficulty large crocodile was shot, no actual evidence was ever taken. For of access and where overall crocodile densities appear to be well a crocodile whose length exceeds that of any other record by a below carrying capacity (Pontillas 2000). The lack of scarring large margin, a high degree of skepticism is understandable and body injuries also suggests limited competition. Under such when bearing in mind the track record of inaccurate or exagger­ favorable conditions for a crocodile, individuals that may also ated size records (Greer 1974; Whitaker and Whitaker 2008). have been genetically predisposed for faster growth would have [n 2011 Guinness World Records announced that a 5.48 m (18 excellent opportunities to thrive and reach exceptional sizes. ft) C. porosus called "Cassius" at Marineland Melanesia on Green How important genetic factors are in influencing growth rates Island, Queensland, was the largest crocodile in captivity (Guin­ and maximum size in wild C. porosus remains unclear, although ness World Records 2011). Cassius was captured alive from the it is possible that extensive and Widespread hunting of popula­ Finniss River in the Northern Territory, Australia, in 1984, and tions until the 1970s may have eliminated genetic traits that had was reportedly missing a small amount of his tail tip. Lolong is 69 selected for exceptionally large crocodiles preferred by trophy cm (2.2 ft) longer than Cassius. The only other captive crocodile hunters, traits that may exist today only extremely rarely. that rivals Lolong for length is "Yai" at Samutprakarn Crocodile Farm, Thailand, who was measured in 2000 at 6.0 m (19.7 ft) with Acknowledgments.-AB thanks the following for the opportu­ a mass of 1114 kg (2455 lbs). While Yai will have grown slightly nity to visit Bunawan and measure Lolong: Office of the Mayor of in the last decade, it seems unlikely that he currently exceeds Bunawan (Mayor E. EIOI'de, R. Nuer). Palawan Wildlife Rescue and Lolong. Yai is also the result of artificial captive hybridization be­ Conservation Center (including R. Sumiller), Protected Areas and tween C. porosus and C. siamensis (Siamese Crocodile), his size Wildlife Bureau (including T. M. Lim, Dr. E. Toledo, J. de Leon),

Herpetological Review 43(4),2012 Natural History New Zealand (NHNZ), C. Meade, N. Javellana, K. a predictor of the sex and size of individuals. Herpeto!. Monogr. Siney, R. Taylor, B. Maddul, R. Ordona, and the communities of Ba­ 8:203-225. rangay Nueva Era, Barangay San Marcos, and Barangay Consuelo. ]SBERG, S., P. THOMSON, F. NICHOLAS, S. BARKER, lIND C. MORAN. 2004. Funding was provided by National Geographic Channel and NHNZ. Farmed saltwater crocodiles: a genetic improvement program. RW and NW thank the Trustees and Staff at Madras Crocodile Bank UnpubJ. report to the Rural Industries Research and Development Trust for support and assistance. We also thank Brandon Sideleau for Corporation Report 04/147, Australia. providing the artwork. All capture protocols and measurement pro­ LANCE, V. A. 1994. An overview of stress in farmed crocodilians. 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