Scientific paper

Osteometry and size reconstruction of the Indian and Pacific Oceans’

Euthynnus species, E. affinis and E. lineatus (Scombridae)
by

Anaïs Marrast* (1) & Philippe Béarez (1)

Abstract. – Two neritic species of scombrids (Euthynnus affinis and E. lineatus) from the Indo-Pacific and the
Eastern Pacific are today classed as commercially important. They have long been exploited and are common
finds on coastal archaeological sites. Size reconstruction from isolated bones is interesting for both biologists and
archaeologists. In archaeology, these studies make it possible to highlight fishing strategies. Therefore, we built
an osteometric model for these two species, using 31 specimens of E. affinis (FL: 274 mm to 828 mm, W: 305 g
to 8674 g) from the Sultanate of Oman and 26 specimens of E. lineatus (FL: 294 mm to 614 mm, W: 481 g to
4200 g) from Ecuador. For E. affinis, the length-weight relationship is W = 1E-05 FL3.0682, and for E. lineatus,
the relationship is W = 2E-05 FL2.9578, with r2 higher than 0.98 for both species. For the osteometric model, we
used the neurocranium, premaxilla, dentary, anguloarticular, quadrate, hyomandibula, maxilla, opercle, anterior
and posterior ceratohyals, scapula and vertebrae. For each bone, we took between 2 and 5 measurements and
plotted the obtained values against the fork length. For all selected bones, we produce at least one regression
equation with a high r2 (> 0.9) that permits accurate estimates of the length and weight of Euthynnus individuals
© SFI
Submitted: 19 Oct. 2018
Accepted: 21 Feb. 2019 for both species.
Editor: E. Dufour

Résumé. – Ostéométrie et reconstruction de la taille des espèces du genre Euthynnus des océans Indien et Pacifi-
que, E. affinis et E. lineatus (Scombridae).
Key words
Euthynnus Deux espèces néritiques de scombridés (Euthynnus affinis et E. lineatus) de l’Indo-Pacifique et du Pacifique
Osteometry Est sont aujourd’hui considérées comme commercialement importantes. Elles sont exploitées depuis longtemps
Size reconstruction et sont souvent identifiées sur les sites archéologiques côtiers de cette partie du monde. La reconstitution de la
Allometry taille d’un poisson à partir d’os isolés est d’un grand intérêt pour les biologistes et les archéologues. En archéo-
Length-weight logie, ces études permettent notamment de renseigner les stratégies de pêches. Nous avons donc construit un
relationship modèle ostéométrique pour ces deux espèces, en utilisant 31 spécimens d’E. affinis (FL : 274 mm à 828 mm, W :
Ichthyoarchaeology 305 g à 8674 g) du Sultanat d’Oman et 26 spécimens d’E. lineatus (FL : 294 mm à 614 mm, W : 481 g à 4200 g)
d’Équateur. Pour E. affinis, la relation longueur-poids est W = 1E-05 FL3,0682, et pour E. lineatus, la relation est
W = 2E-05 FL2,9578, avec un r² supérieur à 0,98 pour les deux espèces. Pour le modèle ostéométrique, nous avons
utilisé le neurocrâne, le prémaxillaire, le dentaire, l’anguloarticulaire, le carré, l’hyomandibulaire, le maxillaire,
l’operculaire, les ceratohyaux antérieur et postérieur, la scapula et des vertèbres. Pour chaque os, nous avons pris
entre 2 et 5 mesures et représenté les valeurs par rapport à la longueur à la fourche. Pour tous les os sélectionnés
il y a au moins une équation de régression avec un r2 élevé (> 0,9) qui permet des estimations précises de la lon-
gueur et du poids des individus des deux espèces Euthynnus.

Found worldwide in tropical to temperate waters, the
scombrid genus Euthynnus is represented by three different
species: E. alletteratus (Rafinesque, 1810), the little tunny;
E. lineatus (Kishinouye, 1920), the black skipjack; and
E. affinis (Cantor, 1849), the kawakawa or mackerel tuna.
The first is present in the Tropical Atlantic and the Medi-
terranean, while the other two are present in the Tropical
Eastern Pacific and the Indo-Pacific, respectively. Euthyn-
nus species are epipelagic, essentially neritic fishes, which
occur in open waters but generally stay inshore. They have
a robust, elongated and streamlined body, and are known to
form large multi-species schools with other scombrids or
even other taxa. These schools reach between 100 and 5000
individuals (Collette and Nauen, 1983).
All three species are currently commercially impor-
tant for both industrial and small-scale fisheries, but they
also played an important role in ancient subsistence fisher-
ies. Evidence that these neritic species were consumed by
coastal populations is attested by their presence on many
archaeological sites around the world, such as in the East-
ern Pacific (Béarez, 1996; Martínez et al., 2009; Béarez et
al., 2012), the Western Atlantic (Wing, 2001), the Mediterra-
nean (Desse and Desse-Berset, 1994) and the Northern Indi-
an Ocean (Beech, 2004; Uerpmann and Uerpmann, 2003).
Apparently, Euthynnus affinis is less common in the Central
Pacific archaeological record, where it seems to be replaced
by the closely related skipjack tuna, Katsuwonus pelamis
(Linnaeus, 1758) (Lambrides and Weisler, 2017).
Despite the fact that Euthynnus species, especially
E. affinis, have commercial importance, only a few papers
provide information on their osteology or osteometry (Kishi-
nouye, 1923; Godsil, 1954; Mansueti and Mansueti, 1962),

(1) Archéozoologie, archéobotanique: sociétés, pratiques et environnements (AASPE), Muséum national d’histoire naturelle, CNRS,
CP 56, 57 rue Cuvier, 75005 Paris, France. [anais.marrast@mnhn.fr] [bearez@mnhn.fr]
* Corresponding author [anais.marrast@mnhn.fr]

Cybium 2019, 43(2): 187-198. https://doi.org/10.26028/cybium/2019-423-007

Osteometry of Euthynnus species Marrast & Béarez

Table I. – Description of the measurements illustrated in figure 2.

Measurement
Anatomical element Measurement description
number
ncr 1 Distance from the anterior part of the vomer to the posterior part of the basioccipital
Neurocranium / ncr ncr 2 Maximal width of the vomer
ncr 3 Maximal width between sphenotics
pmx 1 Length of the anterior dorsal process (without teeth)
Distance from the anterior tip of the pmx to the posterior base of the dorsal process
Premaxilla / pmx pmx 2
(without teeth)
pmx 3 Medio-lateral width at posterior level of the dorsal process
dn 1 Length of the dorsal dentigerous branch
Dentary / dn dn 2 Height of the symphysis
dn 3 Distance from the symphysis to the postero-lateral incisure
ang 1 Total length
ang 2 Distance from the dorsal curvature to the posterior part of the articular process
Anguloarticular / ang
ang 3 Medio-lateral width of the articular facet
ang 4 Total height of the articular
qd 1 Total width of the articular condyle
Quadrate / qd
qd 2 Distance from the articular condyle to the tip of the preopercular process
hm 1 Total height
Hyomandibula / hm hm 2 Greatest medio-lateral width at level of the opercular process
hm 3 Greatest distance between the sphenotic facet and the opercular process
mx 1 Total length
Maxilla / mx mx 2 Height of the main axis
mx 3 Greatest width of the anterior portion
op 1 Cranio-caudal length of the articular fossa
Opercle / op op 2 Greatest height
op 3 Height of the articular fossa
ach 1 Greatest cranio-caudal length
Anterior ceratohyal / ach
ach 2 Height of the external median bridge
pch 1 Greatest cranio-caudal length
Posterior ceratohyal / pch
pch 2 Greatest dorso-ventral height
sca 1 Distance between the scapular foramen and the articular facet
Scapula / sca
sca 2 Medio-lateral width of the articular facet
M1 Anterior height of the centrum, including the neural prezygapophyses
M2 Greatest width at level of neural prezygapophyses
First vertebra/ pc 1 M3 Length of the centrum
M4 Posterior height of the centrum
M5 Posterior width of the centrum
M1 Anterior height of the centrum
M2 Anterior width of the centrum
Vertebrae M3 Length of the centrum
M4 Posterior height of the centrum
M5 Posterior width of the centrum
M1 Height of the ural centrum
Hypural plate / hp M2 Width of the ural centrum
M3 Height of the triangular plate

while more information on their growth is available (e.g. often scale allometrically with total length. The reconstruc-
Landau, 1965; Mulhia-Melo, 1980; Valeiras et al., 2008). tion of fish lengths from isolated bones is significant for both
In fish, body shape, as well as body parts or organs, most biology and archaeology (Reitz et al., 1987). In biology, it

188 Cybium 2019, 43(2)

Marrast & Béarez Osteometry of Euthynnus species

Table II. – Biometric information concerning Euthynnus affinis and Euthynnus lineatus specimens (TL, FL and SL in mm; P in g).
Species Number in collection TL FL SL W Species Number in collection TL FL SL W
Euthynnus affinis MNHN-ICOS-01132 889 828 790 8674 Euthynnus affinis MNHN-ICOS-00984 319 288 275 343
Euthynnus affinis MNHN-ICOS-01131 842 806 772 8091 Euthynnus affinis MNHN-ICOS-00983 308 274 263 305
Euthynnus affinis MNHN-ICOS-01130 795 721 685 6420 Euthynnus lineatus MNHN-ICOS-01608 666 614 578 4200
Euthynnus affinis MNHN-ICOS-00320 776 725 682 6600 Euthynnus lineatus – 619 565 530 3250
Euthynnus affinis MNHN-ICOS-00239 765 714 690 5950 Euthynnus lineatus MNHN-ICOS-01630 580 525 490 2517
Euthynnus affinis MNHN-ICOS-00319 722 679 654 4900 Euthynnus lineatus MNHN-ICOS-01609 565 525 493 2500
Euthynnus affinis MNHN-ICOS-1433 690 592 630 4330 Euthynnus lineatus MNHN-ICOS-01629 510 466 440 1723
Euthynnus affinis MNHN-ICOS-00295 678 631 605 4050 Euthynnus lineatus – 510 444 420 1750
Euthynnus affinis MNHN-ICOS-00273 671 622 600 4030 Euthynnus lineatus MNHN-ICOS-01618 506 455 430 1457
Euthynnus affinis MNHN-ICOS-00238 660 611 590 3600 Euthynnus lineatus – 500 438 414 1716
Euthynnus affinis MNHN-ICOS-00296 640 587 567 3150 Euthynnus lineatus MNHN-ICOS-01612 495 435 412 1394
Euthynnus affinis MNHN-ICOS-00297 636 588 565 3000 Euthynnus lineatus MNHN-ICOS-01625 495 444 420 1450
Euthynnus affinis MNHN-ICOS-00298 626 575 553 2850 Euthynnus lineatus MNHN-ICOS-01627 470 423 400 1335
Euthynnus affinis MNHN-ICOS-00272 616 552 552 2788 Euthynnus lineatus MNHN-ICOS-01624 450 396 375 1129
Euthynnus affinis MNHN-ICOS-00994 610 539 521 2670 Euthynnus lineatus MNHN-ICOS-01626 445 407 385 1143
Euthynnus affinis MNHN-ICOS-00299 609 560 539 2950 Euthynnus lineatus MNHN-ICOS-01628 430 385 365 1005
Euthynnus affinis MNHN-ICOS-00271 584 543 530 2446 Euthynnus lineatus MNHN-ICOS-01620 425 380 360 944
Euthynnus affinis MNHN-ICOS-00300 578 525 507 2500 Euthynnus lineatus MNHN-ICOS-01621 418 380 360 886
Euthynnus affinis MNHN-ICOS-00986 570 510 489 2102 Euthynnus lineatus MNHN-ICOS-01611 413 374 355 883
Euthynnus affinis MNHN-ICOS-00107 564 519 499 2000 Euthynnus lineatus MNHN-ICOS-01623 412 364 345 828
Euthynnus affinis MNHN-ICOS-00301 536 497 481 1940 Euthynnus lineatus MNHN-ICOS-01622 401 359 340 727
Euthynnus affinis MNHN-ICOS-00311 531 497 477 1983 Euthynnus lineatus MNHN-ICOS-01619 390 348 330 750
Euthynnus affinis MNHN-ICOS-01103 513 486 472 1797 Euthynnus lineatus MNHN-ICOS-01613 360 326 310 646
Euthynnus affinis MNHN-ICOS-00985 495 423 406 1334 Euthynnus lineatus MNHN-ICOS-01614 350 316 300 550
Euthynnus affinis MNHN-ICOS-00312 492 455 438 1484 Euthynnus lineatus MNHN-ICOS-01610 346 311 295 481
Euthynnus affinis MNHN-ICOS-00249 467 431 414 1151 Euthynnus lineatus MNHN-ICOS-01615 340 316 300 558
Euthynnus affinis MNHN-ICOS-00960 425 383 368 998 Euthynnus lineatus MNHN-ICOS-01617 330 294 280 535
Euthynnus affinis MNHN-ICOS-00956 398 359 349 694 Euthynnus lineatus MNHN-ICOS-01616 320 294 280 524
Euthynnus affinis MNHN-ICOS-00955 393 353 342 703

allows the estimation of prey size in the diet of predators (e.g.
tunas, billfishes, sharks). In archaeology, length reconstruc-
tions are important for the study of ancient fisheries since they
allow the estimation of the fish biomass represented in the
site: human population consumption, information about the
fish sizes targeted (juveniles/adults) and fishing gear design
(e.g. mesh size of fishing nets, size of fish hooks). Size recon-
struction also allows the perception of changes in fish catches
through time, and inferences about the evolution of fishing
techniques or the status of the exploitation of the species to
be made (Reitz et al., 1987; Thieren and Van Neer, 2016;
Prestes-Carneiro and Béarez, 2017; Lidour et al., 2018).
The study of the relationships between body part lengths
or between length and body weight is of particular value in
fishery management (Ricker, 1958), ecological studies (Kul-
bicki et al., 2005) or body size reconstruction from isolated
parts (Casteel, 1974; Lidour et al., 2018).
In this study, we present the relationship between select-
ed fish bone measurements and fish length for Euthynnus
affinis and E. lineatus. We focused on these two species from
the Indian and Pacific Oceans, because of their importance
in this area both for modern and ancient fisheries (e.g. Béa-
rez and Lunniss, 2003; Rohit et al., 2012); the third commer-
cially important species is excluded because information on
osteometry in E. alletteratus is already available (Desse and
Desse-Berset, 1994).
Euthynnus affinis is found throughout the Indo-Pacific
Ocean, from East Africa, the Red Sea and the Persian Gulf
to Hawaii, Polynesia. Its maximum fork length is 100 cm,
with a maximum weight of 13.6 kg, but the average length
is about 60 cm (Collette and Nauen, 1983). Unlike other
tunas, which can resist temperatures down to 10°C, it is
always found in warm waters, between 18° to 29°C (Brill,
1994). In the Arabian Sea, the species reaches between 50
and 65 cm in its third year of age, and spawning is observed
all year round with peaks during June and October (Rohit et
al., 2012). Euthynnus lineatus lives in the Eastern Pacific,
along the coast of Western America, from California to Peru.

Cybium 2019, 43(2) 189

Osteometry of Euthynnus species
Figure 1. – General view of Euthynnus
affinis (A) and Euthynnus lineatus (B)
and some selected vertebrae: eight first
precaudals (lateral vieuw) and nine first
preurals (lateral and dorsal views).
Its maximum fork length is 84 cm, with a maximum weight
of 9 kg, but the average length is about 60 cm (Collette and
Nauen, 1983). Along the coast of Central America, spawning
season occurs from October to June (Schaefer, 1987).
Material and methods
For this study, we analysed 31 specimens of Euthynnus
affinis from the Gulf of Oman and 26 specimens of E. ������
linea-
190
Marrast & Béarez
tus from Ecuador, Tropical Eastern Pacific. In order to attain
the most representative samples, and to avoid the need
for extrapolation, specimens were collected from within
the widest size range possible. For all specimens, the total
length (TL), fork length (FL), and standard length (SL) were
recorded to the nearest millimetre (mm), and the total fresh
weight (W) was recorded to the nearest gram (g) (Tab. I).
Their complete skeletons were prepared in the Muséum
national d’Histoire naturelle in Paris, where they are now
stored (Tab. II).
Cybium 2019, 43(2)
Marrast & Béarez Osteometry of Euthynnus species

Figure 2. – Description of the measurements used (see Tab. I).

Cybium 2019, 43(2) 191

Osteometry of Euthynnus species
In order to obtain reliable size reconstructions, our
osteometric model was based on the allometric principle,
which gives the best predictive model (Teissier, 1948; Cas-
teel, 1974; Reitz et al., 1987). Indeed, allometry takes into
account the fact that different parts of the body may have
distinct growth rates, which is a common feature in fish. The
length-weight relationship is represented by a power function
of the type: W = aFLb (Teissier, 1948; Le Cren, 1951), where
W is the total weight of the fish (g), FL is the fork length
(mm), “a” is a constant and “b” is the allometric coefficient.
The length-length relationships are expressed as FL = aBMb,
where BM is the bone measurement. The accuracy of the
equations was evaluated by the coefficient of determination
(r2) and the standard error of estimate (SEE).
Figure 3. – Length-length and length-weight relationships for Euthynnus affinis (left) and Euthynnus lineatus (right).
192
Marrast & Béarez
Among archaeological material, depending on preser-
vation, bones can be severely fragmented, with only their
strongest parts, mostly articular joints, surviving. For this
reason, some bone measurements were taken on parts select-
ed for their good preservation, and several flat bones (e.g.
preopercle, subopercle, interopercle), which do not preserve
well, were not considered in this study. According to our
archaeological observations and research on the bones most-
ly used in biometric studies (Desse, 1984), we decided to
focus our study on the bones more easily assigned to species:
premaxilla, dentary, maxilla, opercle, quadrate, anguloartic-
ular, ceratohyal, hyomandibula and neurocranium (Fig. 1).
We then added the scapula because this bone is often well
preserved among scombrids in an archaeological context.
Cybium 2019, 43(2)
Marrast & Béarez
For each cranial bone, we took between 2 and 4 measure-
ments (Tab. I, Fig. 2), based on previous works by Morales
and Rosenlund (1979) and Desse (1984). All the measure-
ments were taken with a digital caliper (0.01 mm). Osteolog-
ical nomenclature followed Dye and Longenecker (2004).
For the vertebrae, the problem of identifying their posi-
tion along the spinal column was overcome by using the
Global Rachidian Profile (GRP) (Desse et al., 1989). Since
the last precaudal and first caudal vertebrae are very similar,
GRP allows one to select parts of the spinal column where
the vertebral diameter does not vary much, which means
that any vertebra in the segment could give the same size
reconstruction. The GRP can also be used to estimate the
Minimum Number of Individuals (MNI) of Euthynnus in
fish bones assemblages. In order to construct this profile, we
took 5 measurements on each vertebra from modern speci-
mens, for which all the vertebrae were conserved with their
original rank.
Otoliths were not studied, as they are very small and are
generally not recovered, either in archaeological material or
stomach contents.
All the measurements were plotted against the FL of
individuals (in mm). We chose FL instead of TL because
Euthynnus species have a strong lunate tail that makes meas-
uring the total length difficult, and because FL is the most
frequently used length in tuna-like fisheries.
Figure 4. – Global rachidian profiles of Euthynnus affinis (MNHN-ICOS-00983: SL = 263 mm, FL = 274 mm, W = 305 g; MNHN-
ICOS-00986: SL = 489 mm, FL = 510 mm, W = 2102 g; MNHN-ICOS-1130: SL = 685 mm, FL = 721 mm, W = 6420 g) and Euthynnus
lineatus (dotted line, PB-6664: SL = 493 mm, FL = 525 mm, W = 2500 g).
Cybium 2019, 43(2)
Osteometry of Euthynnus species
Results
The specimens of Euthynnus affinis have a FL ranging
from 274 mm to 828 mm, and a weight ranging from 305
g to 8674 g (Tab. II; Fig. 3). The specimens of Euthynnus
lineatus have a FL ranging from 294 mm to 614 mm, and a
weight ranging from 481 g to 4200 g (Tab. II; Fig. 3).
The two species are allopatric except for a few stray
specimens, and hence should not be confused. However, they
can also be separated by the spots and lines on their body or
sorted by differences in their vertebrae; Euthynnus lineatus
always has a marked hyperostosis in its 5th and 6th preural
vertebrae (Béarez et al., 2005). Both Euthynnus species can
be differentiated from the closely related Katsuwonus pela-
mis thanks to the presence of lines on the abdomen, and oste-
ologically on the basis of their vertebrae. Katsuwonus pela-
mis has precaudal vertebrae with thinner and more elongate
holes over the median ridge on the lateral parts of the centra
than Euthynnus species.
The relationships between lengths or between length and
weight given by the power regression equations are highly
significant for both species. Most determination coefficients
(r2) are over 0.9, scoring slightly higher in Euthynnus affinis.
These significant correlations between the skeleton parts and
the size and weight of individuals allow reliable reconstruc-
tions of life size. Some measurements are, however, less sig-
nificant; for E. affinis, this is the case for the M2 of the DN
(0.83), and the M3 of the PU1 (0.81); and for E. lineatus, the
193
 Table III. – Allometric relationships between fork length and bone measurements for Euthynnus affinis (FL = aBMb). Number of specimens: 30.

194
Bone Measurement Equation r2 SEE Bone Measurement Equation r2 SEE Bone Measurement Equation r2 SEE
ncr 1 y = 5.2615x1.1057 0.9905 16.2846 M1 y = 93.054x0.8361 0.9782 23.3623 M1 y = 76.249x0.8353 0.9726 25.3686
ncr ncr 2 y = 48.424x0.9248 0.9896 16.8776 M2 y = 91.977x0.7935 0.9634 31.3574 M2 y = 81.646x0.7348 0.9837 17.0673
ncr 3 y = 9.0822x1.0276 0.9873 18.8479 pc 3 M3 y = 81.568x0.8904 0.9776 24.6110 pu 6 M3 y = 76.102x0.6699 0.9568 19.0342
pmx 1 y = 36.07x0.9489 0.9754 23.5420 M4 y = 95.817x0.8561 0.9821 20.2419 M4 y = 78.257x0.8467 0.8212 50.6276
pmx pmx 2 y = 37.19x1.0095 0.9590 29.4210 M5 y = 87.8x0.7867 0.9781 26.8131 M5 y = 85.039x0.7214 0.9583 27.3033
pmx 3 y = 196.25x0.9622 0.8804 47.3650 M1 y = 99.801x0.846 0.9774 26.1528 M1 y = 75.543x0.874 0.9221 40.7968
dn 1 y = 5.8451x1.1768 0.9824 21.7980 M2 y = 93.731x0.7664 0.9813 25.7346 M2 y = 88.874x0.7116 0.9597 28.5115
dn dn 2 y = 95.52x0.8378 0.965 30.1513 pc 4 M3 y = 74.105x0.9638 0.9794 23.7504 pu 5 M3 y = 79.894x0.6888 0.9295 38.5846
Osteometry of Euthynnus species

dn 3 y = 16.744x1.0738 0.9751 22.5286 M4 y = 95.142x0.8568 0.9796 22.4173 M4 y = 91.448x0.863 0.7991 58.6627

ang 1 y = 5.4654x1.1571 0.9855 18.2458 M5 y = 94.939x0.7534 0.9607 25.1211 M5 y = 97.825x0.7378 0.9269 47.0356
ang 2 y = 22.201x1.0632 0.9875 17.4743 M1 y = 99.7x0.8387 0.9746 24.4229 M1 y = 70.821x1.007 0.8846 55.1539
ang
ang 3 y = 145.12x0.8451 0.9551 36.0657 M2 y = 88.715x0.7931 0.9862 20.8602 M2 y = 101.23x0.7337 0.9391 44.6035
ang 4 y = 34.909x0.9619 0.9789 24.2272 pc 5 M3 y = 71.441x0.9658 0.9765 24.0301 pu 4 M3 y = 79.489x0.7828 0.8773 54.9784
qd 1 y = 152.57x0.8209 0.9183 46.2016 M4 y = 96.084x0.8516 0.9854 19.5661 M4 y = 127.39x0.7367 0.6800 94.4206
qd
qd 2 y = 21.565x0.9942 0.9919 13.1806 M5 y = 88.869x0.7909 0.9814 26.6451 M5 y = 112.46x0.7452 0.9259 34.9156
hm 1 y = 10.867x1.0322 0.9906 16.1084 M1 y = 99.252x0.8386 0.9799 23.0957 M1 y = 68.921x1.1016 0.9193 41.2237
hm hm 2 y = 33.302x0.9556 0.9772 24.0559 M2 y = 93.264x0.7829 0.9876 22.5433 M2 y = 95.787x0.8284 0.9747 27.8959
hm 3 y = 67.172x0.9561 0.9489 36.5657 pc 6 M3 y = 70.966x0.9615 0.9825 19.3550 pu 3 M3 y = 84.676x0.895 0.9068 55.2000
mx 1 y = 8.5473x1.095 0.9878 15.9913 M4 y = 89.226x0.887 0.9794 22.6466 M4 y = 95.697x0.9144 0.9587 31.0050
mx mx 2 y = 139.72x0.8733 0.8526 53.9242 M5 y = 91.029x0.7948 0.9576 32.0832 M5 y = 92.624x0.8992 0.9552 37.1870
mx 3 y = 80.859x1.0276 0.9807 19.0873 M1 y = 98.073x0.8464 0.9732 25.2329 M1 y = 100.06x0.9266 0.9624 26.8710
op 1 y = 45.382x1.0117 0.9635 31.0683 M2 y = 89.005x0.8174 0.9855 22.4637 M2 y = 113.02x0.8969 0.8712 61.6445
op op 2 y = 45.382x1.0117 0.9635 129.4188 pc 7 M3 y = 68.36x0.9731 0.9815 22.8178 pu 2 M3 y = 188.3x0.7671 0.7413 80.4127
op 3 y = 45.382x1.0117 0.9635 127.8943 M4 y = 87.407x0.8916 0.9769 27.0491 M4 y = 125x0.7646 0.9185 24.8901
ach 1 y = 11.192x1.0903 0.9888 16.0867 M5 y = 90.845x0.7966 0.9611 28.4812 M5 y = 95.695x0.9457 0.9655 27.4408
ach
ach 2 y = 118.95x0.9348 0.9388 41.8872 M1 y = 87.091x0.8936 0.9742 32.2053 M1 y = 105.41x0.8941 0.9841 22.8050
pch 1 y = 29.724x1.015 0.9814 20.7912 M2 y = 86.755x0.827 0.9873 30.6535 M2 y = 117x0.9698 0.9076 44.9523
pch
pch 2 y = 41.277x0.9054 0.9720 27.8825 pc 8 M3 y = 70.606x0.9397 0.9859 26.6567 pu 1 M3 y = 204.59x0.8911 0.8136 60.6825
sca 1 y = 163.54x0.8013 0.9442 41.4283 M4 y = 78.462x0.9348 0.9803 33.4870 M4 y = 162.45x0.6503 0.8618 33.4825
sca
sca 2 y = 120.51x0.8895 0.8261 64.5534 M5 y = 91.675x0.7932 0.9826 34.8516 M5 y = 104.8x0.9435 0.9378 41.6108
M1 y = 61.202x0.8727 0.9756 21.5510 M1 y = 72.634x0.8616 0.9429 36.2072 M1 y = 120.61x0.8713 0.9857 18.6626
M2 y = 48.714x1.0239 0.9351 38.3012 M2 y = 79.641x0.7901 0.9752 27.1680 hp M2 y = 129.02x1.0121 0.9418 33.2386
pc 1 M3 y = 122.94x0.7589 0.9638 30.9592 pu 8 M3 y = 72.345x0.7577 0.9483 33.8715 M3 y = 36.528x0.8866 0.9929 10.8054
M4 y = 86.297x0.83 0.9885 16.7135 M4 y = 68.2x0.8654 0.9547 30.0470
M5 y = 95.078x0.8141 0.954 35.2393 M5 y = 77.557x0.7717 0.9832 20.3948
M1 y = 90.397x0.8127 0.9769 21.7212 M1 y = 76.065x0.8281 0.9812 24.1060
M2 y = 94.487x0.8063 0.9508 34.6229 M2 y = 75.238x0.7849 0.9833 21.2338
pc 2 M3 y = 85.87x0.8566 0.9712 26.7243 pu 7 M3 y = 73.197x0.7082 0.948 35.7684
M4 y = 88.639x0.8575 0.9752 23.2340 M4 y = 62.63x0.9041 0.9437 30.6240
M5 y = 83.534x0.8287 0.9749 28.3761 M5 y = 82.699x0.7303 0.9853 20.3260

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 Table IV. – Allometric relationships between fork length and bone measurements for Euthynnus lineatus (FL = aBMb). Number of specimens: 21-22.
Bone Measurement Equation r2 SEE Bone Measurement Equation r2 SEE Bone Measurement Equation r2 SEE
ncr 1 y = 6.4785x1.0382 0.9830 9.8153 M1 y = 80.503x0.8843 0.9523 16.1027 M1 y = 74.286x0.8253 0.9644 14.8095
ncr ncr 2 y = 52.438x0.8981 0.9818 11.6257 M2 y = 92.853x0.7676 0.9586 16.2237 M2 y = 77.152x0.7377 0.9591 15.1965
ncr 3 y = 16.363x0.8584 0.9208 28.4773 pc 3 M3 y = 74.23x0.9113 0.9756 13.0374 pu 6 M3 y = 66.189x0.6829 0.8704 27.9809
pmx 1 y = 34.318x0.9366 0.9617 14.3914 M4 y = 80.806x0.9178 0.9328 19.1812 M4 y = 72.887x0.8457 0.8861 30.7155

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Marrast & Béarez

pmx pmx 2 y = 37.78x0.9765 0.9625 14.4471 M5 y = 93.419x0.739 0.9681 14.4475 M5 y = 87.447x0.6819 0.9000 20.2352
pmx 3 y = 196.67x0.8737 0.7958 35.1222 M1 y = 87.856x0.8883 0.9561 16.5201 M1 y = 79.027x0.8177 0.9247 24.7902
dn 1 y = 7.8663x1.0757 0.9658 12.9910 M2 y = 95.764x0.7372 0.9814 12.5691 M2 y = 92.497x0.6573 0.9367 21.9593
dn dn 2 y = 98.44x0.7989 0.9381 18.1801 pc 4 M3 y = 66.936x0.9922 0.9728 12.5919 pu 5 M3 y = 51.701x0.803 0.9373 19.8445
dn 3 y = 27.315x0.8874 0.9263 19.3894 M4 y = 85.087x0.8881 0.9543 16.6341 M4 y = 72.841x0.9486 0.9172 22.9266
ang 1 y = 6.9066x1.0777 0.9828 9.5603 M5 y = 93.042x0.7459 0.9698 14.8547 M5 y = 84.831x0.7791 0.9145 26.0025
ang 2 y = 22.924x1.0359 0.9748 12.0622 M1 y = 91.387x0.8652 0.9636 14.7645 M1 y = 66.265x1.0079 0.8993 29.1562
ang
ang 3 y = 144.44x0.7959 0.9623 16.2330 M2 y = 94.873x0.7485 0.9853 10.8009 M2 y = 95.139x0.7369 0.8983 26.6779
ang 4 y = 32.717x0.9678 0.9617 14.9794 pc 5 M3 y = 65.317x1.0055 0.9562 17.6235 pu 4 M3 y = 52.582x0.9052 0.9035 28.4824
qd 1 y = 136.55x0.8703 0.9342 19.1446 M4 y = 89.463x0.8621 0.9579 15.0148 M4 y = 84.878x0.913 0.8516 29.6267
qd
qd 2 y = 24.759x0.931 0.9673 12.9685 M5 y = 85.373x0.8033 0.9835 10.7737 M5 y = 71.284x0.9427 0.9316 18.7833
hm 1 y = 12.386x0.9767 0.9805 10.5917 M1 y = 91.791x0.8561 0.9708 13.0997 M1 y = 58.598x1.1591 0.9269 21.7323
hm hm 2 y = 34.503x0.9241 0.9738 13.3588 M2 y = 91.835x0.7783 0.9728 13.3523 M2 y = 79.557x0.9064 0.9368 19.0983
hm 3 y = 71.948x0.8823 0.9275 20.5952 pc 6 M3 y = 70.199x0.9584 0.9735 13.1225 pu 3 M3 y = 76.141x0.8842 0.8284 35.5598
mx 1 y = 8.7044x1.07 0.9817 9.9004 M4 y = 86.939x0.8764 0.9623 15.6394 M4 y = 88.878x0.9504 0.9172 22.5869
mx mx 2 y = 136.68x0.8272 0.7420 39.0486 M5 y = 81.702x0.8407 0.9843 10.3154 M5 y = 72.424x1.0126 0.9621 15.1082
mx 3 y = 91.704x0.9445 0.9111 23.9390 M1 y = 95.315x0.8269 0.9626 15.9765 M1 y = 95.165x0.9491 0.9380 20.7788
op 1 y = 52.815x0.921 0.9326 19.2882 M2 y = 86.356x0.827 0.9852 9.8539 M2 y = 100.72x0.9405 0.8399 31.0818
op op 2 y = 12.756x0.9108 0.9781 12.0354 pc 7 M3 y = 63.899x0.9966 0.9766 11.4856 pu 2 M3 y = 181.57x0.7143 0.6140 52.5358
op 3 y = 108.09x0.8538 0.9479 17.8224 M4 y = 85.875x0.8646 0.9584 17.5928 M4 y = 75.509x1.0065 0.9619 15.2665
ach 1 y = 12.953x1.0286 0.9792 11.3792 M5 y = 82.357x0.8404 0.9740 12.4849 M5 y = 89.367x0.9649 0.9666 13.7251
ach
ach 2 y = 156.93x0.7096 0.8175 30.4365 M1 y = 87.744x0.8544 0.9650 16.7757 M1 y = 90.365x0.9767 0.9602 16.0636
pch 1 y = 29.943x0.9932 0.9788 11.1191 M2 y = 83.59x0.837 0.9771 12.3476 M2 y = 103.31x1.0684 0.8907 27.8399
pch
pch 2 y = 44.995x0.8551 0.9389 18.9375 pc 8 M3 y = 53.373x1.0592 0.9581 15.8354 pu 1 M3 y = 209.23x0.8565 0.8139 35.4401
sca 1 y = 147.72x0.8322 0.9376 20.8655 M4 y = 83.315x0.8679 0.9600 16.3713 M4 y = 97.379x0.8996 0.9361 19.9379
sca
sca 2 y = 139.65x0.8006 0.9365 19.3598 M5 y = 78.969x0.8562 0.9880 11.2349 M5 y = 103.16x0.9281 0.9457 19.7091
M1 y = 62.112x0.8419 0.9825 9.5511 M1 y = 56.832x0.9511 0.9612 15.4095 M1 y = 105.9x0.936 0.9620 15.8317
M2 y = 52.691x0.9536 0.9696 77.6146 M2 y = 77.57x0.7905 0.9649 14.2249 hp M2 y = 108.85x1.1331 0.9574 17.2272
pc 1 M3 y = 105.42x0.7853 0.9054 26.1891 pu 8 M3 y = 47.155x0.8947 0.9688 13.9942 M3 y = 33.944x0.9193 0.9769 12.3732
M4 y = 90.784x0.7953 0.9691 14.1715 M4 y = 70.661x0.8279 0.9224 21.9828
M5 y = 103.43x0.7355 0.9641 17.2075 M5 y = 81.658x0.7501 0.9667 13.2195
M1 y = 85.893x0.8222 0.9626 15.7454 M1 y = 72.173x0.8302 0.9650 16.0456
M2 105.46x0.7227 0.9655 15.2220 M2 y = 76.775x0.7781 0.9627 14.5925
pc 2 M3 y = 81.186x0.8322 0.9770 12.3854 pu 7 M3 y = 55.136x0.7909 0.9645 14.6216
M4 y = 77.802x0.9048 0.9600 14.4513 M4 y = 63.831x0.8812 0.9703 13.3459
M5 y = 91.489x0.7677 0.9647 15.5911 M5 y = 81.749x0.7184 0.9526 15.9994

195
Osteometry of Euthynnus species
Osteometry of Euthynnus species Marrast & Béarez

measurements with the lowest r2 are the M3 of the premax-
illa (0.79), and the M2 of the maxilla (0.74).
Taking all the M2 measurements of the vertebrae for
one individual, we built a GRP for three different speci-
mens of E. affinis (Fig. 4). We observed that the vertebral
diameters (M2) are very homogeneous among the last pre-
caudal and the first caudal vertebrae (rank 5 to 29). The pro-
file for E. lineatus is very similar (Fig. 4: dot-line) as is the
one for E. alletteratus (Desse and Desse-Berset, 1994: 72).
Note, however, that the diameter of the last caudal is higher
in E. lineatus, a fact that could be linked to the hyperostotic
condition of the preural vertebrae in this species.
The GRP allows to obtain a rather good estimate of the
length and weight of an individual, even when it is not pos-
sible to assign a precise rank to an isolated vertebra from the
median part of the backbone (Desse et al., 1989).
However, we have to contrast these results after observ-
ing the standard error of estimate (SEE), which indicates the
prediction error. Here, we have a large ranging of SEE val-
ues (Tabs III, IV). In E. affinis, the majority of the length-
length relationships presented SEE values oscillating around
37; for E. lineatus, the SEE values obtained were lower than
those obtained for E. affinis, around 18.
lated bones. Some differences in determination coefficients
indicate that some measurements are more appropriate than
others for size reconstruction. However, all selected bones
had at least one regression equation with a high r 2 (> 0.9)
that should permit accurate estimates of the length or weight
of Euthynnus individuals, and we recommend using the best-
fitted regressions whenever possible.
Similar work has previously been done by Desse and
Desse-Berset (1994) on E. alletteratus using dentary meas-
urements for size reconstruction of archaeological remains
from the Cape Andreas Kastros site (Cyprus). As they used
a linear model the results are not directly comparable, but
the GRP curve they present is very similar to the one of
E. affinis. Indeed, as all Euthynnus species have a very sim-
ilar shape it is likely that our models can also be used for
E. alletteratus.
The data presented here should facilitate reconstructions
of diet and feeding behaviour of piscivorous species, includ-
ing past humans, and help reconstruct fishing activities and
human impact on neritic scombrids at least at a regional
scale.

Acknowledgements. – We would like to thank Salem el-Ghazali,

Discussion and conclusion Khamis Nasser, Héctor Parrales, Cruz Elías Pincay, Enrique Toro,
Elena Maini and many other fishermen for their help in catching
the fish. Eric Pellé and the ‘Service des Préparations Ostéologiques
When we compare our length-weight relationship results et Taxidermiques’ of the MNHN for his help in the preparation of
with the previously published data, we can see a great simi- the skeletons. Thanks are also due to Gabriela Prestes Carneiro for
larity between the equations values (Tab. V). For E. affin- her advice, Alice Diaz Chauvigné, and Jill Cucchi for copy-editing.
is, our results are very close to those of Sivasubramaniam
(1966) and Silas (1967) for the Indian waters, with an ‘a’
value close to 0.00001 (10-5) and a ‘b’ value close to 3.06, References
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E. affinis Indian Ocean Morrow, 1954 0.000018 2.9630
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In this article, we have pre- E. lineatus Eastern Pacific Klawe and Calkins, 1965 0.000011 3.0817
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allow the estimation of the length E. lineatus Ecuador, Eastern Pacific Present study 0.000020 2.9982
and weight of E. affinis and E. lin- E. alleteratus Turkey, Mediterranean Sea Kahraman and Oray, 2001 0.000090 2.7256
eatus individuals from their iso- E. alleteratus Tunisia, Mediterranean Sea Hajjej et al., 2009 0.000025 2.9264

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