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185
CAUDAL FIN SKELETON OF THE LATE CRETACEOUS LAMNIFORM SHARK,
CRETOXYRHINA MANTELLI, FROM THE NIOBRARA CHALK OF KANSAS
AbstractThe caudal fin morphology of the Late Cretaceous lamniform shark, Cretoxyrhina mantelli (Agassiz),
was previously inferred from scale morphology, which suggested that it was capable of fast swimming. A specimen
from the Niobrara Chalk of western Kansas is described here and offers new insights into the morphology of the
caudal fin of the taxon. The specimen preserves the posterior half of the vertebral column and a series of hypochordal
rays. These skeletal elements exhibit features suggesting that C. mantelli had a lunate tail and a caudal peduncle with
a lateral fluke. The specimen also supports the idea that the body form of C. mantelli resembled that of the extant
white shark, Carcharodon carcharias (Linneaus). Given a total vertebral count in Cretoxyrhina mantelli of about
230, this specimen suggests that the transition between precaudal and caudal vertebrae was somewhere between the
140th and 160th vertebrae. The estimated total body length of the specimen described here ranges from 640 cm to 700
cm, marking the largest C. mantelli individual estimated to date. New skeletal data from the specimen further supports
the view that C. mantelli was an active shark capable of fast swimming.
INTRODUCTION insights into the shape of the sharks tail. The purpose of this paper is 1) to
describe the morphology of the specimen, and 2) to discuss the caudal fin
Cretoxyrhina mantelli (Agassiz) was a Late Cretaceous lamniform morphology of C. mantelli and its paleoecological significance. For com-
shark that lived in CenomanianCampanian seas worldwide, including the parative purposes, fossil specimens in the following institutions are referred
Western Interior Seaway of North America (e.g., Cappetta, 1987; Siverson, to in this paper: Sternberg Museum of Natural History, Fort Hays State
1992, 1996; Shimada, 1997d). The species is represented chiefly by its University (FHSM), Hays, Kansas; and the vertebrate paleontology collec-
teeth, but some reasonably complete skeletons of the species are known tion of the University of Kansas Museum of Natural History (KUVP),
from the late Coniacian Santonian portion of the Smoky Hill Chalk Mem- Lawrence.
ber of the Niobrara Chalk in western Kansas (Shimada, 1997b). Those
skeletal remains suggest that large individuals of C. mantelli measured about SYSTEMATIC PALEONTOLOGY
5 to 6 m in total length and possibly had a body form similar to the modern
great shark, Carcharodon carcharias (Linnaeus) (Shimada, 1997b). The Class Chondrichthyes
fossil record demonstrates that Cretoxyrhina mantelli fed on large marine Subclass Elasmobranchii
vertebrates (e.g., teleosts, sea turtles, mosasaurs, and plesiosaurs: Shimada, Order Lamniformes Berg, 1958
1997c; Shimada and Everhart, 2004; Shimada and Hooks, 2004; Everhart, Family Cretoxyrhinidae Glikman, 1958
2004, 2005a) and possibly scavenged bloat-and-float carcasses of fully
terrestrial vertebrates (e.g., nodosaurid dinosaurs: Everhart and Hamm, Genus Cretoxyrhina Glikman, 1958
2005). Cretoxyrhina mantelli (Agassiz, 1843)
Despite the wealth of information about the paleobiology of
Cretoxyrhina mantelli (e.g., Shimada, 1997a, 1997b, 1997c), there are MaterialCMN 40906 (Figs. 15), a string of caudal and poste-
rior precaudal vertebrae with hypochordal rays and placoid scales.
still many unresolved questions. The morphology of its caudal fin is one
such gap in our knowledge. The previously suggested morphology, and the Horizon and localityThe specimen was collected by G. F.
Sternberg from the Niobrara Chalk (Upper Cretaceous: see Hattin, 1982)
estimated starting point of the caudal fin in C. mantelli (Shimada, 1997b),
were based on a series of assumptions. Shimada (1997b) demonstrated in western Kansas. It was sold to the Geological Survey of Canada, from
which the museum originated, in 1912.
that C. mantelli had keeled placoid scales with an average interkeel dis-
tance of approximately 45 microns. Because this interkeel distance is com- Because exact biostratigraphic data were not available to constrain
the age of the specimen within the Niobrara Chalk, chalk samples were
parable to that in scales of some extant fast-swimming sharks, Shimada
(1997b) inferred that C. mantelli was also capable of fast swimming. This taken from the matrix for stratigraphically diagnostic foraminiferans. Our
result shows that four taxa dominate the assemblage. Heterohelix globulosa
evidence, combined with the fact that C. mantelli had a conical head, led
Shimada (1997b) to consider that the species had a stout fusiform body (Ehrenberg), with a Campanian Maastrichtian range, is the most abun-
dant. The other key taxa are: Archeoglobigerina cretacea (dOrbigny),
with a lunate caudal fin for efficient hydrodynamic propulsion. Given that
the fossil species had a total vertebral count of approximately 230, Shimada with a range from middle Coniacianearly Maastrichtian; Rugoglobigerina
rugosa (Plummer), known from the early Campanian; and Whiteinella
(1997b) suggested that the caudal fin of C. mantelli could have begun at
about 133rd vertebra, on the basis of comparisons with some extant fast- centennialensis Frerichs, which occurs in late Santonianearly Campanian
strata (Pessagno, 1967; Frerichs et al., 1975; Frerichs, 1979). The pres-
swimming lamniform sharks with lunate tail fins (e.g., lamnids: Lamna
Cuvier, Isurus Rafinesque, and Carcharodon Smith). ence of R. rugosa, which makes up about 10% of the individuals in our
sample, strongly suggests that the Cretoxyrhina specimen came to rest on
The Canadian Museum of Nature (CMN) in Ottawa, Ontario, houses
a putative Cretoxyrhina mantelli specimen, CMN 40906, which consists the chalky ocean bottom in the early Campanian. The shark specimen is
thus from the upper part of the Smoky Hill Chalk. If so, whereas C. mantelli
of a vertebral column and some additional skeletal elements (Fig. 1). CMN
40906 is noteworthy because of its large size and features that provide new is known from early Campanian deposits (e.g., Siverson, 1992), CMN
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FIGURE 1. Photograph of CMN 40906, string of vertebrae with hypochordal rays of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 30 cm).
FIGURE 2. Line drawing of CMN 40906 (cf. Fig. 1), string of vertebrae (v) with hypochordal rays (hcr) of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale
bar = 30 cm).
FIGURE 4. Close-up view of putative caudal fin base, showing connection between vertebral column and hypochordal rays in CMN 40906 (scale bar = 10 cm; cf. Figs. 1,
2).
FIGURE 5. Placoid scales (four examples) of Cretoxyrhina mantelli (Agassiz) from CMN 40906 (scale bar = 50 micronsm). Sample locations are relative to vertebral
positions in Figure 2. A, Scale from dorsal to v24: top, apical view (anterior to the top); bottom, anterior view. B, Scale from above, or ventral to v66: left, oblique view
(anterior to the top left); right ,anterior view. C, Scale location same as A: lateral view (anterior to the left). D, Scale location same as B: apical view (anterior to the right).
188
FIGURE 6. Example of articulated vertebrae of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 10 cm; v 73v82 in FHSM VP-2187: see Shimada,
1997b). A, vertebrae in right(?) lateral view (note that this surface was against ocean floor when the shark skeleton was buried); B, vertebrae in dorsal(?) view (note post-
burial distortion that resulted in lateral flattening of vertebrae).
FIGURE 8. Tentative skeletal reconstruction of 6-m-long Cretoxyrhina mantelli (Agassiz) based on available fossil record (Shimada, 1997b; this study). Solid lines =
known parts; broken lines = inferential parts.
preserved posteriormost hypochordal rays are short, whereas the rays be- maximum possible angle observed in extant sharks (see Thomson, 1976).
tween them are markedly elongate. In addition, the base of the anteriormost Therefore, CMN 40906 further strengthens the idea (Shimada, 1997b)
hypochordal rays in the specimen occurs immediately anterior to the sharp that the body form of Cretoxyrhina mantelli (Fig. 8) resembled that of
bend (at an angle of ca. 45) that is observed in the vertebral column. If one extant lamnids, including the white shark, Carcharodon carcharias. We
considers the sharp bend at face value as a dorsally directed bend of the assume that Cretoxyrhina mantelli also had a caudal peduncle with a lat-
vertebral column that constitutes the axis of a tail, this skeletal organization eral fluke.
is remarkably similar to that of extant sharks with a high aspect ratio: i.e., a Quantitative studies of vertebral dimensions in modern sharks are
lunate (symmetrical) tail with large dorsal and ventral lobes (Fig. 7A). rare. However, inspection of published illustrations of caudal fins of extant
Thomson and Simanek (1977, table 3) listed Rhincodon Smith fast swimming lamniform sharks with lunate tails (e.g., Lamna in Mivart,
(Orectolobiformes), Cetorhinus Blainville, Lamna, Isurus, and 1879, pl. LXXV, fig. 1; Isurus in Garman, 1913, pl. 63, fig. 5: e.g., Fig.
Carcharodon (Lamniformes) as best representatives of this type of shark, 7A) shows that a marked reduction in vertebral size is present at the
with the caudal portion of the vertebral column bent at an angle of ca. precaudal-caudal transition. There are two specimens of Cretoxyrhina
=30. The caudal fin of other extant sharks is strongly heterocercal: i.e., an mantelli that preserve a series of vertebrae extending near the terminal
asymmetrical tail with a much smaller ventral lobe compared to the dorsal end: FHSM VP-2187 and KUVP 69102. A re-examination of vertebral
one (Thomson and Simanek, 1977). In these sharks, the vertebral column diameters in the posterior half of the vertebral column in these specimens
shows a weak bend or virtually no bend, and the length of the hypochordal (i.e., post-v110: Appendix 2) reveals that a marked decrease in the diam-
rays is more or less uniform across the tail (Figs. 7B, 7C). Therefore, CMN eter between v140 and v145 in FHSM VP-2187 (i.e., 6 mm drop in that
40906 supports the hypothesis that Cretoxyrhina mantelli had a lunate tail interval as opposed to 04 mm per drop every six vertebrae) and between
(Fig. 8). It is noteworthy that, because Rhincodon is not a lamniform (Shirai, v155 and v160 in KUVP 69102 (i.e., 8 mm drop in that interval as op-
1996), and because C. mantelli does not share an immediate common an- posed to 04 mm drop per every six vertebrae where distortion is mini-
cestry with Cetorhinus, Lamna, Isurus, or Carcharodon (Shimada, 2005a), mal). Therefore, we here suggest that the caudal fin of C. mantelli possibly
the similarity in caudal fin morphology between Cretoxyrhina mantelli occurred at an approximate range of v140v160.
and these extant sharks is interpreted to be a homoplasy due to convergent If one assumes that the range of v155v160, which would give a
evolution. However, we note that the caudal fin of C. mantelli possibly had conservative caudal fin size, is equivalent to that of v35v40 in CMN
a wider angle between the upper and lower lobes and a slightly shorter 40906 (where a marked decrease in vertebral size is observed: see above),
lower lobe (Fig. 8) compared to the caudal fin of those extant sharks (e.g., the total vertebral count of the shark individual (CMN 40906) was at least
Fig. 7A), because the hypochordal rays in CMN 40906 are largely directed 216. The number, 216, corresponds remarkably well to the estimated total
ventrally (Figs. 2, 4; as opposed to ventroposteriorly in those extant sharks). vertebral count of Cretoxyrhina mantelli, 230 (see Shimada, 1997b), by
We also note that, because the posteriorly located hypochordal rays appear considering the probable number of missing terminal vertebrae (i.e., 10
to be thinner and shorter as compared to those in Rhincodon, Cetorhinus, 25 vertebrae: see above).
Lamna, Isurus, or Carcharodon, the lobes of the lunate tail in Cretoxyrhina The largest measurable vertebra in CMN 40906 is v3, which has
mantelli could have been slightly narrower than the lobes in those extant a diameter of 88 mm. If v35 is assumed to be v140 in life, v3 would
sharks. then represent v108. In FHSM VP-2187, which is conservatively estimated
Thomson and Simanek (1977) found that extant sharks with a het- to be 500 cm TL, the diameter of v108 is 69 mm (Shimada, 1994, table 1).
erocercal ( vertebral bent) angle of ca. 30, not only have a well- If one assumes that the vertebral diameter in CMN 40906 has the same
developed ventral hypochordal lobe, but also a deep fusiform body with a size relation to the TL as the relationships between the diameter and TL in
conical head and a caudal peduncle bearing the lateral fluke. Thomson and FHSM VP-2187, CMN 40906 is 128% of FHSM VP-2187. This suggests
Simanek (1977) characterized such sharks as fast swimming pelagic that the shark individual represented by CMN 40906 was possibly 640 cm
sharks, typified by extant lamnids, such as the mako and white sharks. TL. Whereas this assumption of v35 representing v140 gives a conser-
The vertebral bend in CMN 40906 is about 45, which approximates the vative TL estimate for CMN 40906, one may assume v35 to be v155 for
191
a less conservative estimate, and in this case, v3 would represent v123 in mosasaurs, is energetically costly and requires efficient swimming capabil-
life. The diameter of v123 in FHSM VP-2187 is 63 mm (Shimada, 1994, ity. Thus, as suggested by Shimada (1997c), it is likely that C. mantelli was
table 1). Therefore, CMN 40906 is 140% of FHSM VP-2187, suggesting an active shark capable of fast swimming. Such information is important in
that the shark could have measured 700 cm TL. The range of 640700 cm understanding the paleoecological dynamics of the Cretaceous seas.
TL for CMN 40906 represents the largest Cretoxyrhina mantelli individual
estimated to date (cf. Shimada, 1997b). ACKNOWLEDGMENTS
The caudal fin is the primary locomotor structure in sharks. Al- We thank M. Feuerstack and C. Kennedy (CMN) for assistance with
though functional analysis of caudal fins is far from complete even for the specimen, and co-op students H. Bernatchez and M. Brire from lcole
living sharks (for review, see Lauder, 2000; Lingham-Soliar, 2005), ex- secondaire de Casselman for their help in locating isolated denticles and
amination of caudal fin morphology in extinct sharks is important because scale patches for study. A. Murray (CMN) took the SEM and overall speci-
it gives insights into their swimming capability, which in turn, as noted by men photos. We thank M. J. Everhart (FHSM) for his discussion on the
Motani (2002), has implications to behavior, physiology, and other aspects Niobrara stratigraphy. The senior author thanks the following present and
of their biology. The exact outline of the body and tail of Cretoxyrhina past FHSM and KUVP associates for allowing us access to comparative
mantelli (Fig. 8) remains inferential at the present time. However, it is Cretoxyrhina specimens in their care: J. Chorn, C. Fielitz, D. Miao, H.-P.
noteworthy that the notion of C. mantelli as a fast-swimming shark, Schultze (KUVP), and R. J. Zakrzewski (FHSM). Comments made on
supported here, agrees well with the inferred feeding behavior of the taxon earlier drafts by D. J. Ward (Kent, United Kingdom) and M. D. Gottfried
based on taphonomic evidence. In particular, a mosasaur vertebra speci- (Michigan State University) as well as reviews by J. I. Kirkland (Utah Geo-
men from Kansas shows signs of bone healing over an embedded logical Survey) and D. R. Schwimmer (Columbus State University) greatly
Cretoxyrhina tooth, suggesting the post-bite survival of the mosasaur indi- improved the quality of this manuscript.
vidual (Shimada, 1997c). Biting large, presumably active vertebrates, like
REFERENCES
Agassiz, L., 18331843 [1843], Recherches sur les poissons fossils (5 volumes): sharks from the Smoky Hill Chalk (Upper Cretaceous) of western Kansas: Ab-
Neuchtel, Imprimerie de Patitpierre, 1,420 p. stracts Kansas Academy of Science, v. 22, p. 3031.
Bardack, D., 1965, Anatomy and evolution of chirocentrid fishes: University of Hattin, D. E., 1982, Stratigraphy and depositional environment of Smoky Hill Chalk
Kansas Paleontological Contributions, v. 10, p. 188. Member, Niobrara Chalk (Upper Cretaceous) of the type area, western Kansas:
Beeson, E. and Shimada, K., 2004, Vertebrates from a unique bonebed of the Upper Kansas Geological Survey Bulletin 225, 108 p.
Cretaceous Niobrara Chalk, western Kansas: Journal of Vertebrate Paleontol- Lauder, G. V., 2000, Function of the caudal fin during locomotion in fishes: Kine-
ogy, v. 24, supp. to no. 3, p. 37A. matics, flow visualization, and evolutionary patterns: American Zoologist, v.
Berg, L. S., 1958, System der Rezenten und fossilen Fischartigen und Fische: Ber- 40, p. 101122.
lin, Hochschulbcher fr Biologie, 310 p. Lingham-Soliar, T., 2005, Caudal fin in the white shark, Carcharodon carcharias
Cappetta, H., 1987, Chondrichthyes II: Mesozoic and Cenozoic Elasmobranchii, in (Lamnidae): A dynamic propeller for fast, efficient swimming: Journal of Mor-
Schultze, H.-P., ed., Handbook of Paleoichthyology, Volume 3B: Stuttgart, Gustav phology, v. 264, p. 233252.
Fischer Verlag, p. 1193. Mivart, St. G., 1879, Note on the fins of elasmobranchs, with considerations on the
Compagno, L. J. V., 1990, Relationships of the megamouth shark, Megachasma nature and homologues of vertebrate limbs: Transactions of Zoological Society,
pelagios (Lamniformes: Megachasmidae), with comments on its feeding hab- London, v. 10, no. 10, p. 439484, 6 pls.
its: National Oceanic and Atmospheric Administration Technical Report, Na- Motani, R., 2002, Swimming speed estimation of extinct reptiles: Energetic approach
tional Marine Fisheries Service, v. 90, p. 357379. revisited: Paleobiology, v. 28, p. 251262.
Compagno, L. J. V., 1999, Endoskeleton, in Hamlett, W. C., ed., Sharks, Skates, Pessagno, E. A., Jr., 1967, Upper Cretaceous planktonic foraminifera from the West-
and Rays: The Biology of Elasmobranch Fishes: Baltimore, Johns Hopkins ern Gulf Coast Coastal Plain: Paleontographica Americana, v. 5, p. 242445.
University Press, p. 6992. Shimada, K., 1994, Paleobiology of the Late Cretaceous shark, Cretoxyrhina
Everhart, M. J., 2004, Late Cretaceous interaction between predators and prey. Evi- mantelli (Lamniformes: Cretoxyrhinidae), from Kansas [M.S. thesis]: Hays,
dence of feeding by two species of shark on a mosasaur: PalArch, Vertebrate Kansas, Fort Hays State University, 169 p.
Palaeontology Series, v. 1, p. 17. Shimada, K., 1997a, Dentition of the Late Cretaceous lamniform shark, Cretoxyrhina
Everhart, M. J., 2005a, Bite marks on an elasmosaur (Sauropterygia: Plesiosauria) mantelli, from the Niobrara Chalk of Kansas: Journal of Vertebrate Paleontol-
paddle from the Niobrara Chalk (Upper Cretaceous) as probable evidence of ogy, v. 17, p. 269279.
feeding by the lamniform shark, Cretoxyrhina mantelli: PalArch, Vertebrate Shimada, K., 1997b, Skeletal anatomy of the Late Cretaceous lamniform shark,
Paleontology Series, v. 2, p. 1424. Cretoxyrhina mantelli, from the Niobrara Chalk in Kansas: Journal of Verte-
Everhart, M. J. 2005b, Oceans of KansasA Natural History of the Western Inte- brate Paleontology, v. 17, p. 642652.
rior Sea: Bloomington, Indiana University Press, 322 p. Shimada, K., 1997c, Paleoecological relationships of the Late Cretaceous lamniform
Everhart, M. J., and Hamm, S. A., 2005, A new nodosaur specimen (Dinosauria: shark, Cretoxyrhina mantelli (Agassiz): Journal of Paleontology, v. 71, p. 926
Nodosauridae) from the Smoky Hill Chalk (Upper Cretaceous) of western Kan- 933.
sas: Transactions of the Kansas Academy of Science, v. 108, p. 1521. Shimada, K., 1997d, Stratigraphic record of the Late Cretaceous lamniform shark,
Frerichs, W. E., 1979, Planktonic foraminifera from the Sage Breaks Shale, Centen- Cretoxyrhina mantelli (Agassiz), in Kansas: Transactions of Kansas Academy
nial Valley, Wyoming: Journal of Foraminiferal Research, v. 9, p. 159184. of Science, v. 100, p. 139149.
Frerichs, W. E., Atherton, C. C., and Shive, P. N., 1975, Planktonic foraminifera Shimada, K., 2005a, Phylogeny of lamniform sharks (Chondrichthyes:
from the chalky members of the Niobrara Formation, Centennial Valley, Wyo- Elasmobranchii) and the contribution of dental characters to lamniform system-
ming: Journal of Foraminiferal Research, v. 11, p. 294312. atics: Paleontological Research, v. 9, p. 55-72.
Garman, S., 1913, Plagiostoma (Sharks, Skates, and rays): Memoir of Museum of Shimada, K., 2005b, Skeletal and dental anatomy of the Late Cretaceous lamniform
Comparative Zoology, v. 36, p. 1528. shark, Cretalamna (=Cretolamna) appendiculata: Journal of Vertebrate Pale-
Glikman, L. S., 1958, [Rates of evolution in lamnoid sharks]: Doklady Akademia ontology, v. 25, supp. to no. 3, p. 114A115A.
Nauk S.S.S.R., v. 123, p. 568571. (In Russian) Shimada, K. and Cicimurri, D. J., 2005, Skeletal anatomy of the Late Cretaceous
Hamm, S. A., and Shimada, K., 2002, The first associated tooth set of the Late shark, Squalicorax (Neoselachii: Anacoracidae): Palaeontologische Zeitschrift,
Cretaceous lamniform shark, Scapanorhynchus raphiodon (Mitsukurinidae) v. 79, p. 241261.
from the Niobrara Chalk of western Kansas: Transactions of Kansas Academy Shimada, K. and Everhart, M. J., 2004, Shark-bitten Xiphactinus audax (Teleostei:
of Science, v. 105, p. 1826. Ichthyodectiformes) from the Niobrara Chalk (Upper Cretaceous) of Kansas:
Hamm, S. A., Shimada, K., and Everhart, M. J., 2003, Three uncommon lamniform Mosasaur, v. 7, p. 3539.
192
Shimada, K., Ewell, K., and Everhart, M. J., 2004, The first record of the lamniform Siverson, M., 1996, Lamniform sharks of the mid Cretaceous Alinga Formation
shark genus, Johnlongia, from the Niobrara Chalk (Upper Cretaceous), west- and Beedagong Claystone, western Australia: Palaeontology, v. 39, p. 813849.
ern Kansas: Transactions of Kansas Academy of Science, v. 107, p. 131135. Stewart, J. D., 1990, Niobrara Formation vertebrate stratigraphy, in Bennett, S. C.,
Shimada, K., and Hooks, G. E., III., 2004, Shark-bitten protostegid turtles from the ed., Niobrara Chalk Excursion Guidebook: Lawrence, University of Kansas
Mooreville Formation (Upper Cretaceous) of Alabama: Journal of Paleontol- Museum of Natural History and Kansas Geological Survey, p. 1930.
ogy, v. 78, p. 205210. Thomson, K. S., 1976, On the heterocercal tail in sharks: Paleobiology, v. 2, p. 19
Shirai, S., 1996, Phylogenetic interrelationships of neoselachians (Chondrichthyes: 38.
Euselachii), in Stiassny, M. L. J., Parenti, L. R., and Johnson, G. D., eds., Inter- Thomson, K. S. and Simanek, D. E., 1977, Body form and locomotion in sharks:
relationships of Fishes: San Diego, Academic Press, p. 9-34. American Zoologist, v. 17, p. 343354.
Siverson, M., 1992, Biology, dental morphology and taxonomy of lamniform sharks Welton, B. J. and Farish, R. F., 1993, The Collectors Guide to Fossil Sharks and
from the Campanian of the Kristianstad Basin, Sweden: Palaeontology, v. 35, p. Rays from the Cretaceous of Texas: Lewisville, Texas, Before Time, 204 p.
519554.
APPENDIX 1.Approximate diameter (in millimeters) of vertebral centra in CMN APPENDIX 2.Approximate diameter (in millimeters) of vertebral centra in two nearly
40906. Dash = vertebral centrum with severe damage or distortion. complete skeletons of Cretoxyrhina mantelli (Agassiz) (data based on Shimada,
1994, table 1). Value in parentheses = estimated value from adjacent centra; value
in brackets = unreliable measurements due to severe distortion of centra.