REP, INT, WHAL. COMMN 46, 1996 409 ScraTaNP2 Osteological Comparison of Bryde’s Whales from the Philippines with Specimens from Other Regions William F, Perrin Southwest Fisheries Science Center, PO Box 271. La Jolla, California 92038, USA Ma, Louella L., Dolar Marine Laboratory, Siliman University, Dumaguete 6200, Philippines and Center for Marine Biotechnology ‘and Biomedicine, Scripps Institution of Oceanography, University of California at San Diego, La Jolta, California 92093, USA Eduardo Ortega Mindanao State University ~ Higan Institute of Technology, ligan City, Phitippines ABSTRACT Results of «comparison of sie a indicated by sizeof skull and postcranial skele Bryde's whales from the Pilippines ( tvth smaller sample sizes for oval component, most salvage from waste heaps at whaling sites in Purllacan end Lils, Boke ‘nd Camiguin Ctra, Mindanao) with that of spesiacs from Myanmar (ormey Burn California ae cons ental South Pace, the India Ocean, Curagao, Venez KEYWORDS: BRYDE'S WHAL TAXONOMY; STOCK IDENTITY INTRODUCTION "The purpose of this study was (0 address the question of, stock identity of Bryde’s whales (Balaenoptera edeni) curently under exploitation in the Philippines (Dolar etal, 1994). ‘The existence or possible existence of geographic or ‘microallopatric ecological forms of varying body size has been described for Bryde's whales from Japan (Omura, 1977) and South Attica (Bost, 1977). Andersen and Kinze (1993) recently described Bryde’s whales from Thailand that Were similar to the relatively small coastal form in South, African waters (Best, 1977). It thus appears that body size ‘may be an important characteristic in defining stocks of Bryde’s whales. Length data are not available for the whales taken in the Philippine indigenous whale fishery. The fishery is not officially monitored. The whales are butchered in remote sites and the meat distributed for sale and consumption within the country (Dolar er al., 1994), However, we collected and measured whale skulls and disassociated bones tepresenting whales taken over several years for comparison with similar data in the literature and for specimens in other collections, MATERIALS AND METHODS ‘Skulls were collected from Pamilacan Island and Lila, Bohol fand at Sawang-Sawang (Sagay) on Camiguin Island, Mindanao (Fig. 1) from 1989 t0 1992 (Dolar et al., 1994). In 1993 and 1994, a concerted effort was made to collect the remaining skulls and bones at the three sites. ‘The material has been placed in the marine mammal collection of the Marine Laboratory of Silliman University, In addition. we Tria, Singapore, South Afsica, apa the with a9 hypothesis of mall size i the ppt indigenous i the Philippine { NORTH PACIFIC: SOUTH PACIFIC; INDIAN OCEAN; ANATOMY/MORPHOLOGY: igen 12° qe 10° Py 8° 122° 423° —=«124° «125° «126°. Fig. |. Sites where Bryde’s whale bones were collected ‘examined one skull with rami and two isolated rami in the collection of the Museum of Silliman University: these were collected at one of the whaling sites in about 1980. ‘The collected material includes: 23 skulls or basicranias 47 rami (24 left, 20 right and 4 unexamined or unassignable to side); approximately 350 vertebrae (271 sufficiently to allow measurement) including 10) atlases. and410 PERRIN wai: OSTEOLOGICAL COMPARISON OF BRYDE'S WHALES, hbumeri; 18 radii: 17 ulnae; 7 scapulae and a large body of skull and vertebral fragments, The material thus represents at least 24 whales and probably represents more. Local residents at Lila reported that a large uuckload of bones (mainly skulls) had been sold to a speculator for bone meal several yeurs ago, 0 itis likely that the collected vertebral ‘and appendicular remnants are from more whales than represented by the 23 skulls and basicrania. Many of the bones swere damaged during the butchering process. by beach erosion and/or by lengthy emersion in a fresh-water ditch (at Lil ‘The comparative base of published ostcological data is small (11 shales) and for some whales only fragmentary ddara_have been published. These include: the holotype specimen, a physically immature (Some thoracic epiphyses slill unfused to centra) 37ft (L1.3m) whale of unknown sex that stranded in Myanmar (formerly Burma) (Anderson, 1878);43.911¢13.2m) physically immature female from the fishery in South Africa (Linnberg, 1931); a physically rmauure whale of unknown sex and lengtt that stranded near Singapore (lunge, 1950); two additional South African fishery specimens and an Indian stranded specimen, all of, unknown or unstated sex, Tength and maturity (Andrews, 1919; Junge, 1950); a 4Sit (13.7m) immature female from the Japanese coastal fishery (Omura, 1959}, a physically ‘mature stranded specimen of unknown sex and length from Curagao (Soot-Ryen, 1961); a stranded specimen of unknown sex and length and unstated maturity. from Veneeuela (Cagnolaro and Notarbartolo-di-Sciara, 1979), anda 14,7m physically mature female and a 14.2m female of unknown maturity from Japanese fisheries in the central South Pacific and Indian Ocean, respectively (Omura et af 1981) In addition, we have included @ male of unknown length ‘and maturity’ that stranded in Corona del Mar, Orange ‘County, California and is now in the Los Angeles County Musetim of Natural History (LACMB84024).Itwas identified based om its baleen. Although unpublished specimens exist in other museums, ¢.g. the US National Museum, we were unable to examine them. We did not include in our comparative base a balacnoplerid ftom Florida reported by Miller (1924) as B. borealis and re-assigned to B. edeni by Mead (1977) based primarily on the characteristics of the baleen. A. Characteristic of the lumbar vertebrae (neural processes + sag : $a ca ve A Fig SP ee a Siiiahn Z., saves BS * | physically immature} 3 {= physically mature JA maturty unknown le holotype (immature) leaning posteriorly as much as 45 degrees) present in the holotype of B. eden? and in all other referred specimens {including ours) is absent in the Florida specimen. We are convinced that Miller was correct in his original identification of the specimen as B. boreatis. In this reaffirmation of Miller's identification we concur with Omura eal, (1981), who based his opinion on characteristics of the nasal bones. We took measurements of the Philippine and Californian specimens based on those of Omura (1959), Omura et al (1981) and Pertin (1975) and as suugested by 1.G. Mead (pers. comm.). In this analysis, only measurements of length and width are used for the skull (see Cummings, 1985, for illustrations of the Bryde’s whale skull); a future analysis of osteological variation will incorporate more skull _measurements (and more specimens). Most of the Philippine skulls lack the premaxillac Because of this, we used the length of the skull from the tip ‘of the left maxilla (right, if left damaged) to the posteriormost extension oF the occipital condyles as our basic index of skull length. When this measurement was not given in the published data, we calculated it from the published photograph or drawing RESULTS ‘The Philippine skulls are all smaller than the other 12 (Fig. ‘Unfortunately, none are known to be from physically mature whales, s0 this is not conclusive evidence that the species is smaller in the Philippines than elsewhere. The sample for skull width alone is larger (Fig. 3), and the largest specimen, from Camiguin Island, is assumed (0 be al least OTHER REGIONS (11) Mm physically mature maturity unknown |/ immeture 4 PHILIPPINES (17) 75 150 200 250300 360 Length of skull from tip of left maxillary (em) ‘ig. 2. Scalteple af masiemam (quamesal wiih on length of skull Frou tipof lett msillary for 9 skulls of Baleenoptera eden om tbe Philippines ad 12 skull rom other regione: SA, South Africa: MY. Myanmar larmesky Bust thoenype specimen: CU, Curagan: SU, Singapore: CA, California; VE, Venezuela IN, Ladi 10, lads ZZ poy 80 100 120 140 160 180 ‘Skull width (cm) Fig. 3. Frequency disrhatons of skull (squamosal ith for (7 shall ‘oF Balaenoprerd went Irom the Phlsppnes and LI fom other otype specimen ind Assumed sexuallyREP. INT. WHAL. COMMN 46, 1996 au sexually mature because it was accompanied by a stnall calf {also captured). This specimen is smaller than the three from ‘ther regions known to be physically mature. The holotype, from Myanmar, was of similar width and known to be physically immature, Again, whilst these data are also suggestive of small size inthe Philippine whales, they are not conclusive, ‘The sample size for ramus length for the Philippines is ceven larger, representing at least I7 whales (Fig. 4). None is as large as the smallest specimen from a known physically mature whale from elsewhere, but again, data on physical ‘maturity are lacking for all of them. ‘The vertebral data show a similar pattern, and here some ‘of the Philippine data are for adult specimens (vertebrae with epiphyses fused (0 the centra). The 48 Philippine thoracic vertebrae with fused epiphyses (from an unknown number of whales) are on average narrower than thoracic vertebrae from the physically mature South Pacific specimen (Fig. 5) ‘The modal size for the entire sample of both mature and immature vertebrae is smaller than for the combined sample {or three whales from other regions (South Pacific, India and Japan). The samples for length of the vertebral centrum (Fig, {6) are larger, and the Philippine sample (38 adult vertebrae} is modally tothe eft ofthe overall sample from other regions (five whales, from the central South Pacific, Curegso, India, Japan and Venezuela). The Philippine sample for width of the centrum (Fig. 7) is the largest (237. from an unknown number of whales, including 38 with fused epiphyses). The ‘mode is again well below that for the whales from other regions (Myanmar ~ the immature holotype, central South Pacific — physically mature and Japan ~ immature), The sample of whales from other regions is small, but the pattern is unlikely to be due ¢o chance alone; the vertebral centrum is almost certainly smaller in the Philippine whales, Other regions (16/11) maturity unknown, 720-160 200 240-260 320 -3a0 Ramus length (cm) Fig. 4, Fregueney ditibucons of rani length fr 33 ram from at east 17 skull (39-17) of Baluenopter eden from the Pilippines and 16 from 11 (16/11) om other regions. H = holotype specks, of Other regions (27/3) = am physically mature immature * Philippines (48) am opiphyses fused oh untused 26 42~«S8 7480108122 \Wiatth of thoracie vertebrae (cm) Fig, 5. Froqlency distriutions of width of thoracic venetra for 48 entanse of Balaenoptera cent from. the Philipines and 27 ‘ertbrse fom 3 (27/3) spectuene rom other regions: central South Pacific (physisliy mature) and Indio and Japan (physically Sf other regions (1285/5) | mmm prysically mature 7 } zzimaturty unknown immature 15 \5 10 Philippines (38) iopiphyses fused 4 8 «12 «16 20 24 2B 82 Length of centrum (em) Fig,6. Frequency distributions of length of cesium for 38 vertebra Tihoracks, mits and caudaly wath Interal processes) of Salueneptera eden rom the Plipiaes and 125 versa for ive Specimens (139/5) from otter regions: central South Pact and Coragvoapysically mature), Indi ad Japan (physially imate) fel Veneeuela(seusee of physical mate unreported).412 PERRIN et ai: OSTEOLOGICAL COMPARISON OF BRYDE'S WHALES BC omer regions (77/9) |. mmphysically mature “TC immature Philippines (237) mm epiphyses fused untusea 40. 20 So 4182228 Width of centrum (cr) Fig. 7. Frequency distributions of widt of centrum foe 237 vercbnie Tinomaics, Tumors and. ceudals with Iter processes) of Bolaenepiera eden Som the Philipines and 77 from 3 whales (GhiA} trom ener gions: central South Pacific (physically mature) snd Mysemar and Japan (physically imate Other regions (10/3) mim physically mature maturity unknown immature Philippines (6/23) Im opiphyses fused ountused 14 18 2226 «30a BAO Length of humerus (em) Fig, 8, Frequency distibutions of length of © homes fom ‘pocimens (@/=8) of Balaenoprere eden! from the Pilippnes and ‘en ues frm 9 specirnens (10%) Trem other regions A similar pattem is evident for the lengths of humerus and radius (Figs § and 9), although the Philippine samples are very small DISCUSSION ‘The data are consistent with an hypothesis of relatively small size of Philippine Bryde's whales compared to those from ther regions. Before a firm conclusion can be drawn, however, data for Philippine specimens of a known desree of Other regions (10/7) mmm physically mature maturity unknown immature Philippines (15/28) Giepiphyses fused] ‘ountusea u aaa SBC Length of racius (ern) Tig, 9, Praquency distibutions of length of 18 sai from specimens (15/8) of Balernoptens eden fom the Philipines nd {fom 7 specimens 17) fom ater tegions physical maturity must be collected. This will only. be possible through systematie monitoring and sampling of the indigenous whale fishery, and we recommend that this be carried out ‘The affinities of the holotype specimen (fron Myanmar) are not clear. It was nearly physically mature (only a few vertebrae with unfused epiphyses, according to Anderson, 1878) and its skull was of a width similar to that of an assumed adult Philippine whale (Fig. 3). Myanmar, of course, is close to Thailand, where small Bryde’s whales have been reported (Andersen und Kinze, 1993), and the Burmese whale conceivably could have been from the same population as the Thai whales. Thus this population may be fof a small coastal form, rather than the larger form predominantly exploited in past fisheries in other regions, and this may be the same form that occurs inthe Philippines The number of Bryde’s whales from other regions represented in world collections is extremely small and inadequate for effective population analyses. We recommend that every effort be made to collect full daa for all stranded Bryde”s whales, and, if feasible, the entire skeleton for physically mature whales should be analysed (maturity can be determined by examination of the state of {usion of vertebral epiphyses (othe centra inthe thoracic and anterior lumbar region). Efforts should also be made to determine sex, through genetic techniques if nevessasy. If scientific or ‘commercial whaling of Bryde’s whales recommences, similar data and samples should be collected ‘or the whales taken: the colleetion of adult skulls (or atleast data on physical maturity) should be a prerequisite part of any such program,REP. INT. WHAL, COMMN 46, 1996 413 ACKNOWLEDGEMENTS LG, Mead of the Division of Mammals of the US Museum of Natural History, Washington, D.C., provided invaluable ‘advice on how to go about measuring baleen whale skulls; be also reviewed the manuscript. JE. Heyning allowed us ‘access to the specimen at the Los Angeles County Museum of Natural History and reviewed the manuscript. Many people helped us collect the bones atthe butchering sites and transport them back to the laboratory: IML. Tan, JS. Leatherwood, C:L. Hill, D. Inocencio, R. Binamira, Jr, A.A. Yapiinchay, E. Himoya, I. and J. Tan, R. Villabona, C. Libosada, R.A. Alejandro, R. Raymundo, M.N. Alava, E. Capuli, D. Wade, C, Dareza, M. Nievales, H. Senekof, D. 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