THE ANATOMICAL RECORD 296:709–722 (2013)

Anatomy and Age Estimation of an

Early Blue Whale (Balaenoptera
musculus) Fetus
RACHEL A. ROSTON,1 DAVID LICKORISH,2 AND EMILY A. BUCHHOLTZ1*
1
Department of Biological Sciences, Wellesley College, Wellesley, Massachusetts
2
Preclinical Development, Stryker Regenerative Medicine, Hopkinton, Massachusetts

ABSTRACT
The external anatomy of a 130-mm blue whale fetus (Balaenoptera
musculus) is described, and its internal anatomy is reconstructed nonin-
vasively from microCT scans. The specimen lies developmentally at the
junction of the embryonic and fetal periods. Similarly to the embryos of
many odontocetes, it lacks a caudal fluke and dorsal fin, but it also exhib-
its an elongated rostrum, resorbed umbilical hernia, partially exposed cor-
nea, and spatial separation of the anus and genitalia seen in early
odontocete fetuses. Dermal ossification of the cranial bones has begun,
but the endochondral skeleton is completely cartilaginous. The shape and
position of the maxilla suggest that the earliest stages of anterior skull
telescoping have begun, but there is no indication of occipital overlap pos-
teriorly. The nasopharynx, larynx, and heart already display the distinc-
tive morphology characteristic of Balaenoptera. This study develops a
model of body length changes during blue whale development by integrat-
ing the large International Whaling Statistics (IWS) database, historical
observations of blue whale migration and reproduction, and descriptions
of fetal growth trends in other mammals. The model predicts an age of 65
days postconception for the specimen. The early developmental milestones
of Balaenoptera mirror those of the odontocete Stenella to a remarkable
extent, but the first appearance of the caudal fluke and dorsal fin are
delayed relative to other morphological transitions. The accelerated pre-
natal growth characteristic of Balaenoptera occurs during fetal, not em-
bryonic, development. Anat Rec, 296:709–722, 2013. V C 2013 Wiley
Periodicals, Inc.

Key words: blue whale; fetus; mysticete; development; microCT

The development of a blue whale (Balaenoptera
musculus) from a zygote to a 7.5-m, 2,000–3,000 kg
neonate (Reidenberg and Laitman, 2008) is remark-
able for both its rapid increase in size and for the
reorganization of multiple mammalian anatomical sys-
tems for life in the water. Study of the transitions of
size and morphology during blue whale development
is limited by access to fetal specimens. Many thou-
sands of fetuses were recovered during the decades of
active commercial whaling, but very few were pre-
served for scientific examination. Embryos and small
fetuses are particularly rare, with the result that the
early ontogeny of the species is largely unknown.

Grant sponsor: NSF; Grant number: 0842507; Grant sponsor:
Wellesley College.
*Correspondence to: Emily A. Buchholtz, Department of
Biological Sciences, Wellesley College, Wellesley, MA 02481.
E-mail: ebuchholtz@wellesley.edu
Received 21 August 2012; Accepted 17 January 2013.
DOI 10.1002/ar.22678
Published online 28 February 2013 in Wiley Online Library
(wileyonlinelibrary.com).

C 2013 WILEY PERIODICALS, INC.

V
710 ROSTON ET AL.
Neonatal size is highly correlated with gestation
length in terrestrial mammals (Huggett and Widdas,
1951; Millar, 1981): taxa with larger neonates have lon-
ger gestation times. The same relationship has also been
documented in toothed whales (odontocetes), many of
which (e.g. Globicephala, Orcinus, Monodon, Physeter)
have gestation periods longer than a year (Frazer and
Huggett, 1973; Marino, 1997). Although specimens are
rare, the embryonic and early fetal anatomy of multiple
small odontocete genera (e.g. Lagenorhynchus, Stenella,
Phocoena) has received considerable attention over the
past century, and the bibliography of cetacean embryol-
ogy assembled by Thewissen and Heyning (2007) pro-
vides ready access to this literature. Developmental
milestones have recently been carefully documented for
size-graduated series of embryos and fetuses of the spe-
 erba et al., 2000; Thewissen
cies Stenella attenuata (St
and Heyning, 2007; Moran et al., 2011).
The development of baleen whales (mysticetes)
presents a strong contrast to the pattern of longer gesta-
tional periods for larger neonates. Observations of mysti-
cete reproductive anatomy and behavior (e.g. Hinton,
1925; Mackintosh and Wheeler, 1929) and analyses of
the size of fetuses recorded by whalers (e.g. Hinton,
1925; Risting, 1928; Mackintosh and Wheeler, 1929;
Laws, 1959; Frazer and Huggett, 1973; Rice, 1983; Reese
et al., 2001) have demonstrated conclusively that mysti-
cetes have relatively short gestation periods despite
their immense neonate sizes. Large neonate size is pri-
marily the result of the acceleration of fetal growth
instead of the elongation of the fetal developmental pe-
riod. Rates of fetal body weight increase that are more
than 250% those of terrestrial mammals and odontocetes
have been documented (Huggett and Widdas, 1951;
Frazer and Huggett, 1973, 1974), but the timing of accel-
erated growth relative to developmental milestones has
received little attention.
The preserved blue whale fetus studied in this project
provides the opportunity to document mysticete mor-
phology near the embryo/fetus developmental transition.
We describe the external anatomy of a very young blue
whale fetus and use microCT scanning to visualize fea-
tures of its internal anatomy without destructive sam-
pling. We use multiple data sources to construct a
preliminary model for the estimation of gestational age
in B. musculus based on body size, using congeneric
specimens to augment the very small database available.
This model allows a comparison of some of the late em-
bryonic and early fetal developmental milestones seen in
Balaenoptera with those previously documented in odon-
tocetes, and a preliminary evaluation of their timing
with respect to the onset of accelerated fetal growth.
MATERIALS AND METHODS
A southern blue whale fetus, Balaenoptera musculus
intermedia, American Museum of Natural History
(AMNH) specimen 63654, was collected by Messrs. Irvin
and Johnson at the Donkergat Whaling Station in Salda-
nha Bay, South Africa, almost certainly during the com-
mercial harvest of its mother. AMNH records indicate
that the listed collection date of June, 1922 is question-
able, an uncertainty reinforced by the rarity with which
small individuals of the species have been collected during
the austral winter. The specimen is now preserved in 70%
ethanol, and this may have been the case for most or all of
its 90 year history. Some shrinkage is likely to have
occurred as a result. The anteroposterior axis of the ani-
mal is twisted, making visualization of planar structures
difficult.
Photographs of the specimen were taken using a Leica
S6D stereoscope with mounted camera. Measurements of
external features were made using photographs and the
public domain software ImageJ to minimize damage from
handling. The young gestational age of the specimen, its
long preservation history, and the undesirability of using
contrast dyes limited the tissues that could be differenti-
ated using CT scans. MicroCT scanning was performed
on a Skyscan 1076 microCT (Bruker MicroCT, Belgium)
with the following scan parameters: 44 kV, 220 mA, 35 mm
resolution, 1.0 step over 180 . The length of the speci-
men dictated the use of four overlapping scans, which
were reconstructed and analyzed using the manufacturer
software programs NRecon, CTAn, and CTVol (Skyscan,
Belgium). Visual artifacts exist along the planar surfaces
at which these scans were combined. The data files were
converted to DICOM file format using manufacturer soft-
ware (Skyscan, Belgium) and further analyzed at Welles-
ley College using the imaging software OsiriX. The
generated images were colored arbitrarily and trimmed
digitally. Measurements of the internal anatomy were
made with the Osirix software from the scans
themselves.
RESULTS
External Anatomy
AMNH 63654 displays moderate cranial and caudal ven-
tral curvature (Fig. 1A), complicating evaluation of its body
length. Crown-rump lengths (CRL) are typically used for
cetacean embryos, and facilitate comparisons with terres-
trial mammals, for which they are used throughout gesta-
tion. However, the gradual reduction of ventral curvature
during early cetacean development makes location of the
rump and measurement of CRL ambiguous in larger fetal
specimens. Total length (TL), measured from the tip of the
rostrum to the notch of the fluke, is a more consistent mea-
surement after relaxation of curvature. AMNH 63654 has
a greatest linear dimension of 68 mm, an approximation of
CRL, and a TL of 130 mm.
Curvature of the neck orients the head at approxi-
mately 40 relative to the main body axis. The elongate
rostrum is clearly demarcated from the forehead. The
anterior end of the lower jaw is bent ventrally at an
angle inconsistent with the curve of the rest of the lower
jaw and there is a tear in the neck, indicating secondary
damage to the fetus. The tongue is visible within the
mouth. The external nares lie at an angle of 65 to each
other in the posterior portion of the rostrum (Fig. 1D).
The head posterior to the external nares is indented,
reflecting the presence of a large fontanel in the skull
roof. The eyes (Fig. 1B) bulge laterally and are dark
gray in color. The eyelids are fused at both inner and
outer margins, with a small linear gap in the center.
The skin is beige in color, probably reflecting both the
early developmental age of the fetus and desquamation
from long-term preservation in ethanol. There is no visi-
ble external auditory meatus.
The forelimbs project posteroventrally at approxi-
mately 55 to the main body axis. They are
 BLUE WHALE FETAL ANATOMY 711

Fig. 1. AMNH 63654, Balaenoptera musculus intermedia. (A) right lateral composite image of the entire
specimen; (B) detail of right eye; (C) detail of right forelimb; (D) rostrum and external nares in dorsal view;
(E) posteroventral midline showing the umbilicus (left), genital tubercle (center), and anus (right); (F) ventral
midline of distal tail. Scale in A 5 10 mm; scale in B–F 5 3 mm.

asymmetrical, with a flipper instead of a paddle outline.
The outlines of the cartilaginous radius and ulna are
visible through the skin (Fig. 1C); the hand is clearly
hyperphalangeous, although the phalangeal formula
cannot be determined with confidence. There are no
hind limb rudiments. Laterally, posteroventral ridges
indicate the course of the 10 most posterior ribs. The
ventral extremity of the last rib lies dorsal to the umbili-
cus. The stub of the umbilicus is oval in cross section,
and there is no trace of an umbilical hernia.
The anterior margin of the genital tubercle lies 6.5
mm posterior to the base of the umbilicus (Fig. 1E). The
tubercle is approximately 3 mm in length, with a
rounded apex that is deflected posteriorly and slightly to
the left. AMNH 63654 lies at the lower end of the size
range of individuals that can be sexed by external mor-
phology (Amasaki et al., 1989), and the genital tubercle
can best be described as undifferentiated. The center of
the small circular anal depression lies 5.2 mm posterior
to the base of the genital tubercle. A very subtle genital
raphe runs between the two structures in the ventral
midline. Mackintosh and Wheeler (1929) identified the
distance from the anus to the center of the reproductive
aperture as the sole trait that varied systematically
between the sexes in fetal and postnatal blue whales.
This difference was also described for B. acutorostrata
fetuses by Amasaki et al. (1989) and supplements geni-
tal morphology as a method for the prediction of fetal
sex. The anal-reproductive distance of AMNH 63654 is
intermediate between those typical of later male and
female B. musculus fetuses, again supporting a designa-
tion of its sex as indeterminate. The distal tail is dorso-
ventrally deep and laterally compressed, but there is no
lateral expansion of the fluke (Fig. 1F).
Raw and proportional dimensions (% of TL) of key
anatomical features of AMNH 63654 are presented in
Table 1. Dimensional comparisons are provided by pub-
lished measurements of a single unsexed 45 cm Balae-
noptera sulfurea (5 B. musculus) fetus collected by R. C.
Andrews (cited by Schulte, 1916) and by the mean val-
ues of multiple South Georgian B. musculus fetuses in 1
m size bins reported by Mackintosh and Wheeler (1929).
As noted by Mackintosh and Wheeler (1929), the most
marked feature of these proportions is their stability
across almost the entire prenatal growth period. Propor-
tional stability during fetal development has been
reported for other Balaenoptera species as well. Schulte
(1916) compared fetuses of B. borealis from 37.5 to 283
cm in length, finding “no progressive change of propor-
tion during the period represented, save in the breadth
of the flipper, which progressively diminishes.” Zemskiy
(1950) also found “proportional growth” in girth, skull,
umbilical and anal measurements in a study of B. physa-
lus fetuses (N 5 65) from 49.5 to 565 cm in length. These
results also agree with the more qualitative statement of
Hinton (1925) that the “general form” of blue whale
fetuses “is perfected” at a length of about 50 cm.
In contrast, measurements from the tip of the rostrum
to the blowhole and eye represent smaller proportions of
TL in AMNH 63654 than in any of the other size catego-
ries, indicating that the rostrum length must still
increase relative to the rest of the face after 13 cm TL.
 712

TABLE 1. Comparison of key body dimensions of Balaenoptera musculus fetuses as raw linear measurements (AMNH 63654) and as percentages
of total body length (TL) (AMNH 63654 and comparative specimens)
AMNH 63654 M&W 1929, 100 cm size bins
A/Sch. 1916
13 cm 13 cm $ 45 cm 0–100 100–200 200–300 300–400 400–500 500–600 600–700
raw data % TL % TL % TL % TL % TL % TL % TL % TL % TL
Snout tip to blowhole 1.3 10.0 14.9 – 5# 14.45 7# 14.76 3# 15.36 1# 15.16 – –
5$ 12.69 4$ 14.90 6# 15.00 – 6# 15.52 1$ 13.46 1$ 14.30
Snout tip to center of eye 1.6 12.3 19.3 – 5# 18.33 8# 18.66 3# 18.65 1# 18.74 – –
5$ 17.49 4$ 19.03 7$ 18.52 – 6$ 18.26 1$ 16.23 1$ 17.78
Fluke notch to post. – 5# 27.30 6# 27.36 3# 26.76 1# 26.76 – –
margin of dorsal fin 5$ 27.37 4$ 26.69 7$ 27.21 – 5$ 26.80 1$ 25.00 1$ 27.46
Fluke notch to anus 34.7 – 5# 30.67 7# 31.71 3# 31.43 1# 30.11 – –
5$ 30.47 4$ 31.62 7$ 31.26 – 6$ 31.52 1$ 34.03 1$ 29.84
Fluke notch to umbilicus 48.3 – 5# 46.44 8# 46.67 3# 45.47 1# 45.68 – –
5$ 46.36 4$ 47.58 7$ 47.61 – 6$ 46.33 1$ 50.00 1$ 43.65
Anus to reproductive aperture 4.5 3.5 – 5# 5.66 8# 5.88 3# 5.97 1# 6.11 – –
ROSTON ET AL.

5$ 2.34 4$ 2.13 7$ 2.67 – 6$ 2.69 1$ 3.08 1$ 1.90

Flipper, tip to axilla 10.9 8.4 11.1 – 4# 11.01 8# 11.43 3# 11.91 1# 12.21 – –
5$ 9.48 4$ 10.81 7$ 11.65 – 5$ 13.09 – 1$ 13.02
Flipper, ventral border length 11.3 8.7 – 4# 15.78 5# 17.25 3# 17.81 1# 17.26 – –
5$ 12.74 4$ 16.07 6$ 16.16 – 5$ 18.45 1$ 16.73 1$ 19.52
Flipper, greatest width 3.4 2.63 4.0 – 4# 3.71 7# 4.30 3# 4.62 1# 4.42 – –
5$ 3.78 4$ 3.93 6$ 3.90 – 6$ 4.50 1$ 4.62 1$ 4.76
Flipper, notch to tip 9.6a – 3# 11.01 6# 13.77 3# 12.58 1# 12.63 – –
3$ 9.45 4$ 11.57 5$ 13.26 – 3$ 13.43 1$ 11.55 1$ 10.79

AMNH 63654 and the female specimen collected by Andrews (cited by Schulte, 1916, p. 398) are single specimens; values from Macintosh and Wheeler (1929) are
means of sexed fetuses in 100 cm size bins collected in South Georgia. Specimen count by sex is indicated for each character. All measurements are in cm. A/Sch,
Andrews in Schulte, 1916; M&W, Mackintosh and Wheeler, 1929.
a
The “tip to tip” value for fluke width cited by Schulte has been halved to generate a notch to tip value.
 BLUE WHALE FETAL ANATOMY
Fig. 2. Cranial anatomy of early Balaenoptera fetuses. (A, B) dermal
ossifications of B. musculus AMNH 63654 in lateral (A) and dorsal (B)
view. Offsets in the mandible and maxilla are artifacts of CT data set
suturing. (C) interpretation (dotted lines) of B. musculus AMNH 63654
dermal and endochondral skull components in dorsal view. (D, E) skull
Similarly, all three forelimb proportions (length, ventral
border length, width) are smaller in AMNH 63654 than
in any of the larger individuals, indicating that the fore-
limb must grow proportionally faster than TL in the
same 13–45 cm TL window.
Skeletal Tissues
Portions of multiple dermal bones of the skull, includ-
ing the mandibles, maxillas, premaxillas, palatines, pter-
ygoids, frontals, parietals, and squamosals are present as
irregularly-shaped ossifications (Fig. 2A,B). Surface tex-
ture is highly irregular, and a zone of lower density (prob-
ably mesenchyme) surrounds each ossification, indicating
that these elements are still in the process of formation.
This suite of dermal bones is similar to that of a 90-mm
Balaenoptera physalus fetus (M€ uller, 1971) and of a Car-
negie Stage 20 odontocete, Stenella attenuata (Moran
et al., 2011), but considerably more restricted than that of
a 230-mm B. acutorostrata specimen figured by Eschricht
(1849 and Fig. 2D,E). The mandible is dense and well
developed, already exhibiting a coronoid process. The
premaxilla, maxilla, and mandible are both larger and
more representative of their adult shapes than the fron-
tal, parietal, and squamosal. The frontal and squamosal
713
of a 230 mm B. acutorostrata fetus in lateral (C) and dorsal (D) view,
modified from Eschricht (1849); Scale bars 5 10 mm. Abbreviations:
Exo, exoccipital; Fo, fontanel; Fr, frontal; Ip, interparietal; Md, mandi-
ble; Mx, maxilla; Pa, parietal; Pal, palatine; Pmx, premaxilla; Pt, ptery-
goid; So, supraoccipital; Sq, squamosal.
lack the lateral expansion typical of adult skulls. The low-
density tissue between the parietals is loosely organized
and unossified, making it impossible to address the issue
of the presence or form of the interparietal (Ridewood,
1923; Koyabu et al., 2012). A large dorsal fontanel, seen
most clearly in sagittal section, lies between the develop-
ing frontals.
The chondrocranium is completely unossified. Resolu-
tion of its elements is complicated by their incomplete
formation and by the similarity of their optical density
to that of the presumed mesenchyme surrounding the
dermal bones. Positional information allows identifica-
tion of basioccipital, basisphenoid, and exoccipital con-
densations. The supraoccipital is not separately defined,
nor are ossification centers of the anterior braincase.
The lack of endochondral ossification in AMNH 63654 is
consistent with the entirely cartilaginous braincase
described in a 105-mm B. acutorostrata specimen by
Burlet (1914). In contrast, the 230-mm B. acutorostrata
fetus described by Eschricht (1849, and Fig. 2D,E) exhib-
ited well-defined occipital elements, and Schulte (1916)
reported ossification centers in the basioccipital, basi-
sphenoid and exoccipitals in a 375-mm B. borealis fetus.
The distinctive telescoping of mysticete skulls is the
product of rostral and caudal elongation of the maxillae
714 ROSTON ET AL.
Fig. 3. Relative anteroposterior centrum length (CL) of vertebrae of
Balaenoptera fetuses and adults. (A) B. musculus fetus (AMNH 63654)
and adult (USNM 124326). (B) B. borealis fetus (Schulte, 1916) and
adult (USNM 236680). Centrum lengths are standardized to the first
thoracic vertebra. C, cervical, T5 thoracic, L, lumbar, Cd, caudal.
and premaxillae as well as the anterior movement of the
occipital and parietals over the orbit (Miller, 1923). In
the telescoped skull, the maxillae, premaxillae, nasals,
frontals, and parietals overlap and occur in a single
transverse cross-section at the dorsal apex of the skull.
In AMNH 63654 the maxilla and premaxilla are elon-
gated but do not overlap the frontal posteriorly, indicat-
ing that the anterior component of braincase telescoping
is at a very early stage. Posteriorly, the supraoccipital is
not yet recognizable as a discrete cartilaginous conden-
sation, and there appears to have been no anterior
movement of the occipital unit relative to the parietals
and/or frontals. These observations suggest temporal
precedence of anterior relative to posterior components
of the telescoping process. This precedence is more
clearly observable in the 230-mm B. acutorostrata fetus
(Eschricht, 1849), in which the maxillae overlap the
frontals, but a fully defined interparietal still lies ante-
rior to the supraoccipital. Maxilla/frontal overlap also
occurs in the 375-mm B. borealis fetus in which anterior
movement of the occipital complex over the interparietal
is marginal (Schulte, 1916). More extensive supraoccipi-
tal / parietal overlap occurs in a B. sibbaldii [5 B. mus-
culus] cranium 40 cm in length (Smets, 1885), and the
frontals are overlapped by both the maxillae and the
parietals in a 193-cm B. musculus fetus (Ridewood,
1923).
The ribs, girdles, and appendages of AMNH 63654 are
entirely cartilaginous, lacking even incipient ossification.
Dense cartilaginous condensations at the distal edges of
the vertebral centra and of proximal portions of the neu-
ral arches and transverse processes (but not the neural
spines) allow determination of anterior column curva-
ture and vertebral dimensions. The cervical column
displays strong ventral curvature, reflecting the ventral
position of the head. The atlas and axis are indistin-
guishable from each other, but the remaining cervical,
thoracic, and lumbar vertebrae are separable. The verte-
bral formula of the anterior column is C7 T16 L14. In
cross-section, individual vertebral shapes already mimic
those of the adult B. musculus column (Fig. 5C), also
noted by Schulte (1916) for the 375-mm B. borealis fetus.
Anteroposterior centrum length (CL) is shortest in the
mid cervical vertebrae, increases in the anterior thora-
cics, and plateaus throughout the posterior thoracic and
lumbar vertebrae. Standardization of CL to the first tho-
racic vertebra in fetal and adult individuals indicates
that AMNH 63654 lacks the progressive elongation of
lumbar CL found in an adult blue whale (National Mu-
seum of Natural History (USNM) 124326) (Fig. 3A), and
typical of all known adult mysticetes (Woodward, 2006).
A comparison of CL data reported for the 385-mm B.
borealis fetus by Schulte (1916) shows a similar differen-
tial in fetal and adult lumbar CL patterns (Fig. 3B). A
pattern of progressive lumbar centrum elongation with
ontogeny has also been reported in neonates and juve-
niles of Caperea marginata (Buchholtz, 2011), and may
be typical of mysticetes in general.
Soft Tissues
The nasopharynx is largest at its posterior junction
with the esophagus and trachea, narrowing anteriorly
until it widens again at the origin of the Eustachian
tubes (Fig. 4A,B). The proximal ends of the Eustachian
tubes exit dorsolaterally (Fig. 4B), passing through the
still incompletely ossified pterygoids. Anteriorly, the
paired nasopharyngeal ducts are laterally compressed,
running parallel to each other over the dorsal surfaces
of the palatines and palatal portion of the maxilla to-
ward the external nares. About two thirds of the dis-
tance in their course they outpocket dorsally to cover
the complex surface of the nasal (ethmo-)turbinals, mir-
roring the osteological and histological structures
reported in other fetal (Beauregard, 1883; Weber, 1904;
Olschl€ager, 1989) and adult (Olschl€ager, 1989) Balaenop-
tera individuals.
The structure of the larynx closely follows that
described for postnatal mysticetes by Reidenberg and
Laitman (2007). The laryngeal aditus (Fig. 4C) pockets
into the floor of the pharynx ventral to the cervical ver-
tebrae. The ventral laryngeal sac communicates with
the more dorsal laryngeal lumen across the distinctive
mysticete U-fold, oriented parallel to the body axis and
to airflow. The laryngeal cartilages present in adult mys-
ticetes (the medial epiglottal, thyroid, and cricoid, and
the paired arytenocorniculate complex) (Reidenberg and
Laitman, 2007) are barely distinguishable on the basis
of density from the surrounding soft tissues. Of these,
the arytenocorniculate is most clearly defined; the loca-
tions of the others are inferred from the location and ori-
entation of the U-fold and other soft tissues.
Cartilaginous tracheal rings are visible posterior to the
larynx and anterior to the bronchial split on the dorsal
surface of the trachea. The tracheal bronchus, typical of
cetartiodactyls (Slijper, 1979; Kida, 1998) branches from
the right side of the trachea slightly less than 1 mm an-
terior to the tracheal bifurcation near the end of the la-
ryngeal sac (Fig. 4D).
 BLUE WHALE FETAL ANATOMY
Fig. 4. Respiratory anatomy of AMNH 63654. (A) Nasopharynx in
dorsal view with adjacent dermal ossifications; (B) nasopharynx in lat-
eral view; (C) larynx in left lateral view; (D) tracheal bronchus in coro-
nal view. Abbreviations: ACC, arytenocorniculate complex; C7,
cervical vertebra 7; CrC, cricoid cartilage; Es, esophagus; Emt, ethmo-
turbinals; Ep, epiglottis; ET, Eustachian tube; H, hyoid cartilage; LA,
The position and size of the major internal organs are
visible in 3D reconstruction (Fig. 5), but resolution of
their internal structure is limited. The lungs are the
least clearly resolved, and appear as anteroposteriorly
elongate areas of low optical density. Their greatest dor-
soventral dimension (4.1 mm, right lung) occurs at the
sharp border of the mediastinal and hepatic surfaces,
which reflect the convexities of the heart and the liver,
respectively. The heart is irregularly triangular in sagit-
tal section, with long basal (dorsal), diaphragmatic (pos-
terior), and costal (anteroventral) surfaces (Fig. 5A–C).
Its long axis lies at an angle of about 40 to that of the
vertebral column. Although the ventral curvature of the
column makes this measurement somewhat arbitrary,
this orientation of the heart is somewhat less “vertical”
than that reported for older Balaenoptera fetuses
(Schulte, 1916; Walmsley, 1938). The ventricles are large
thin-walled elliptical spheres that lie adjacent to, but
715
larygean aditus; LL, laryngeal lumen; LS, laryngeal sac; Md, mandible;
Mo, mouth; Mx, maxilla; Npd, nasopharyngeal duct; Pal, palatine; Ph,
pharynx; Pt, pterygoid; SP, soft palate; T1, thoracic vertebra 1; T, tra-
chea; TB, tracheal bronchus; TC, thyroid cartilage; TR, tracheal rings,
U, U (vocal) fold. Inferred locations of cartilages are indicated with
dotted lines. Scales of all images 5 2 mm.
impinge only minimally upon, each other (Fig. 4C), con-
sistent with the independence of the ventricular walls
described in a 143-cm B. physalus fetus (Walmsley,
1938). Each ventricle is capped and overlapped by an
optically denser anterodorsal mass that is likely a com-
posite of atria, major vessels, and the thymus. Posteri-
orly, the course of the abdominal aorta can be traced
past the level of the renal arteries, where it runs paral-
lel to the paired posterior vena cavae.
The diaphragm is inferred to lie at the boundary
between the thoracic and abdominal cavities, although
its structure is clearly resolvable only at its ventral
insertion on the body wall. The diaphragm’s shape is
highly convex anteriorly, reflecting the extension of the
liver anterior to the posterior margins of the lungs and
heart. The liver is large and dense, occupying the major-
ity of the preumbilical abdominal cavity. It reaches fur-
ther posteriorly on the right than on the left side, in
716 ROSTON ET AL.
Fig. 5. Visceral anatomy of Balaenoptera musculus fetus AMNH
63654. (A) Right lateral 3D reconstruction; (B) schematic diagram of
anatomy inferred from CT reconstruction. Arrows indicate planes of vis-
ual artifacts from scan suturing, (*) indicates plane of image C; (C) ante-
rior view of cross-section of image B; (D) diagram of ventral dissection of
the heart and thymus of a 143-cm B. physalus fetus, modified from
Walmsley (1938). Abbreviations: A, aorta; EN, external nares; F, fontanel;
agreement with the observations made on the fetal liver
of B. borealis (Schulte, 1916). The stomach is masked by
the liver, although the location (but not the structure) of
the intestines is identifiable posteroventral to its caudal
border. The elongate kidneys are very nearly symmetri-
cal in size and shape, although both Schulte (1916) and
Daubt (1898) described the right kidney as lying some-
what more anteriorly in larger B. borealis and B. muscu-
lus fetuses, respectively. This discrepancy may reflect in
part the inability to resolve the kidneys and adrenals in
AMNH 63654. The margins of the kidney1adrenal unit
extend nearly to the vertebral border of the diaphragm
anteriorly, and past the umbilicus posteriorly. The lobu-
lar organization of the kidneys (“groups of renculi” of
Schulte, 1916) is faintly visible.
DISCUSSION
Estimation of Gestational Age
The inability to track a single blue whale pregnancy
from conception to birth imposes severe limitations on
fetal gestation age estimates. Nevertheless, patterns of
mating and migration provide some basic guidelines.
Southern blue whales typically mate and give birth in
low latitudes during successive austral winters, but
spend large parts of the intervening year in Antarctic
feeding grounds during the austral summer (Hinton,
1925; Yochem and Leatherwood, 1985; Sears and Perrin,
2008). This migratory behavior appears to be the domi-
nant factor in constraining the gestation period to a year
GI, gastrointestinal organs, indet.; GT, genital tubercle; K1Ad, kidney 1
adrenal gland; L, liver; LL, left lung; LV, left ventricle; LLT, left lobe of
thymus; RA, auricle of right atrium; RA1T, right atrium 1 thymus; RL,
right lung; RV, right ventricle; RLT, right lobe of thymus; R1, rib 1; T12,
centrum of thoracic vertebra 12; U, umbilicus. Scale of A and B 5 5 mm;
scale of C 5 2.5 mm; scale of D 5 20 mm.
or less and therefore dictating extremely rapid increases
in body size during most of fetal development. Important
analyses of fetal growth in southern blue whales have
been published by Hinton (1925), Risting (1928), Mack-
intosh and Wheeler (1929), Laws (1959), and Frazer and
Huggett (1973). In estimating the gestational age of
AMNH 63654, we rely on the foundation established by
these previous studies and also integrate early embry-
onic and fetal growth data from comparative terrestrial
mammals, the odontocete Stenella, and other mysticete
cetaceans. The following five assumptions, each treated
separately below, are critical to our estimate:
1. The average size of a full term southern blue whale
fetus is 7.5 m.
2. The average length of gestation is 11.5 months, or
350 days.
3. Increase in TL is linear in the late fetal period, and
continues at the same rate as that observed during
the documented mid fetal period.
4. On day 30 of gestation, the TL of a blue whale
embryo is approximately 10 mm.
5. TL growth rate increases gradually and progressively
during late embryonic and earliest fetal development.
Hinton (1925), Frazer and Huggett (1973), and Rei-
denberg and Laitman (2008) have all estimated the aver-
age southern blue whale total body length at birth as 7.5
m, and that value is adopted here. It falls between the
estimates of Laws (1959, 7.0 m) and Risting (1928, 8.1
m), while still accommodating the observation of an
 BLUE WHALE FETAL ANATOMY
Fig. 6. Hypothesis of prenatal growth in the southern blue whale.
(A) Fetus size by day of 13713 fetuses from the IWS database; (B)
number of IWS specimens by day and mean fetal TL calculated at
5 day intervals. The regression of fetal size on day of year for the
entire data set is superimposed (black line); (C) a model of prenatal
growth incorporating data from comparative taxa for earliest embry-
onic growth rates (Period 1), a gradual increase in growth rate and
size modeled with a cubic spline in the data gap between embry-
occasional full term fetus as large as 8.48 m (Risting,
1928). We also adopt the consensus estimate of a gesta-
tion period of 11.5 months or 350 days (Hinton, 1925;
717
onic and early fetal estimates (Period 2), IWS data for the mid-fetal
period (Period 3), assumed continuing linear growth for the late fetal
period (Period 4), and a short period of stasis in TL before birth
(Period 5). The estimated point of transition from embryonic to fetal
growth is based on the interpretation of AMNH 63654 as Fetal
Stage 20, the first fetal stage in odontocetes (Thewissen and Heyn-
ing, 2007). See the text for a discussion of the assumptions used in
model construction.
Mackintosh and Wheeler, 1929; Laws, 1959; Reidenberg
and Laitman, 2008). These two assumptions overwhelm
all of the others in importance, as they demand an
718 ROSTON ET AL.
extremely rapid growth rate over a relatively short ges-
tation period.
The extensive International Whaling Statistics (IWS)
database documents the date of mother’s capture and
total body length of fetuses recovered during the years
1925–1956. Each whaling season extended for a variable
period from late October to early April during the austral
spring and summer, crossing the calendar year boundary.
Embryos, very small fetuses, and very large fetuses are
largely but not entirely absent, reflecting their typical
temporal occurrence outside the whaling season window.
This data set has been variously used by earlier authors
(e.g. Naaktgeboren et al., 1960; Frazer and Huggett,
1973) who generated curves of mid-fetal growth very simi-
lar to that presented here. Our work differs primarily in
the way that the mid-fetal data is placed within the
framework of the entire gestational period.
We compiled IWS data for the years 1925–1950 into a
single data set and sorted by day of the year, using Jan-
uary 1 of year one as day 1 and January 1 of year two
as day 366 (Fig. 6A). The number of specimens per day
of the year peaks in the middle of the whaling season
(Fig. 6B). Variability in the TL of fetuses collected on a
given calendar date is substantial. Minor sources of vari-
ability likely include differences in maternal health and
fetal growth rates, as well as methods of specimen collec-
tion and measurement. For example, it is clear from
data clumping that the sizes of larger fetuses were
lumped into size categories when collected by whalers
(see also Frazer and Huggett, 1973). Several workers
(e.g. Hinton, 1925; Risting, 1928; Laws, 1959) have also
suggested that very small fetuses were almost certainly
overlooked during whaling operations, skewing data
towards high values in early gestation. The largest con-
tributor to variability in size, however, is almost cer-
tainly the wide window for conceptions as documented
for both blue whales and fin whales by many workers
(e.g. Burfield, 1912; Hinton, 1925; Risting, 1928; Laws,
1959; Naaktgeboren et al., 1960). Because we cannot
know conception dates or the number of specimens over-
looked, we work with this flawed but invaluable data
base with caution. Days with fewer than 10 data points
have been discarded, leaving a data set that consists of
13,713 fetuses collected over a period of 164 calendar
days (20 October of Year 1 to 1 April of Year 2). The
regression line of that data set (y 5 2.67x – 737.25,
r2 5 0.41) estimates that during that period (period 3 of
our model, see below), fetal TL increases from 45 to 480
cm at the rate of 2.67 cm/day, Fig. 6B). As a TL of 45 cm
far exceeds the size of any embryo, and a TL 480 cm is
far below that of neonates, the specimens represented by
the IWS data must be from the middle of the fetal
growth period.
Rates of increase in TL after the documented mid fe-
tal period are unknown. They have been assumed to be
linear or nearly so in both fin and blue whales by many
(Hinton, 1925; Huggett and Widdas, 1951; Frazer and
Huggett, 1973; Naaktgeboren et al., 1960) but not all
(Laws, 1959, 1960) previous workers. In domestic ani-
mals (e.g. Huggett and Widdas, 1951; Evans and Sack,
1973; Noakes, 2001) and many odontocetes (Laws, 1959),
rates of fetal CRL (crown-rump length) or TL growth are
linear throughout almost the entire fetal growth period,
sometimes with the exception of a short period of
decreasing growth and/or stasis just before birth. These
observations provide a rationale for assuming the same
pattern in blue whales. An assumption of linear late fe-
tal growth is further supported by the general agree-
ment of the conception and parturition dates that it
dictates with those inferred from observations of ovar-
ian and testicular cycles (Mackintosh and Wheeler,
1929). Using this assumption, an additional 101 days
after the end of the period documented with the IWS
data base would be required for a fetus of 480 cm to
reach the estimated birth TL of 750 cm (Period 4 of our
model). The addition of an arbitrary very short (4 day)
period (Period 5) of growth stasis before birth places
the birth of a “typical” blue whale individual 105 days
later. It also locates July 15 as the date of a typical
birth and 350 days earlier, or July 31 of the previous
year, as the date of a typical conception. More critical
to this model, this reasoning suggests that the initial
growth of the average blue whale embryo from concep-
tion to a TL of 45 cm requires only about 82 days. Cal-
culation of the estimated gestational age in days (x) for
a fetus of given TL (y) is also possible without reference
to date of collection for fetuses in the 45–750 cm TL
range. The only correction factor necessary is the differ-
ence between the day of year and the typical day of con-
ception, or 211 days:
Estimated gestational age in days
5½ðTL in cm1737:25Þ=2:67 – 211
The very early embryonic stages of Balaenoptera are
almost entirely undocumented. Gill (1927) described a
6.5 mm B. musculus embryo, the smallest known mysti-
cete embryo, but did not report day of collection. This
specimen exhibits frontal and mandibular facial promi-
nences, forelimb buds, a cardiac bulge, and a spiraling
tail, all characteristic of Carnegie Stage 11/12 Stenella
embryos in the same size range (Thewissen and Heyn-
ing, 2007). This correspondence argues for conservation
of the earliest developmental sequences and embryo
sizes in odontocetes and mysticetes, and Gill (1927) ex-
plicitly extended these parallels to the chick, rabbit, pig,
and man. More recent studies indicate that this corre-
spondence in size and development extends to conceptus
age as well. Crown-rump lengths of 10 mm are reached
about 30 days after fertilization in humans (Pedersen,
1982; Wisser et al., 1994; DesC^oteaux et al., 2010),
domestic cattle (Noakes, 2001), and odontocete cetaceans
 erba et al., 2000; Thewissen and Heyning, 2007). The
(St
embryos of the much larger Asian and African elephants
reach 10 mm only between 60 and 82 days after concep-
tion (Hildebrandt et al., 2007), but elephants delay im-
plantation and thus elongate (instead of condense) the
gestation period (Hildebrandt et al., 2007, Drews et al.,
2008). Here we adopt the value of TL 5 10 mm as the
best available estimate for blue whale embryo size after
30 days of gestation (Period 1 of our model). We also test
the impact of this assumption (see below) by using alter-
native values of TL 5 5 mm and TL 5 20 mm at 30 days,
with the result that final age estimates for AMNH
63654 varied by less than a single day from those calcu-
lated using TL 5 10 mm.
Use of 10 mm as the TL at day 30 dictates that the
transition from slow early embryonic growth rates to
 BLUE WHALE FETAL ANATOMY
Fig. 7. Age estimates for early fetal specimens of Balaenoptera sp.
in the undocumented interval (Period 2) between early embryogenesis
and the IWS data set. Estimates are based on averages of CRL and
TL (two smallest specimens) and TL (six largest specimens) and an
assumption of gradual increase in the rate of growth from 30 to 82
cm, modeled with a cubic spline. Curves based on three estimates of
rapid mid fetal growth rates, and from 1 to 45 cm TL,
must occur during the very short period between day 30
and day 82, or about 52 days (Period 2). Lacking speci-
mens with both TL and date of collection data in this
gap, we assume a default gradual increase in the rate of
growth and predict conceptus size after a given number
of days of gestation using a cubic spline. We constrain
the spline to run through the day(x) and TL(y) coordi-
nates 30, 1 (estimated gestational age in days of a 1-cm
TL embryo based on comparative taxa, 5 x1, y1) and 82,
45 (estimated gestational age in days of a 45-cm TL fe-
tus at the beginning of the period documented by the
IWS database 5 x2, y2). We adopt values for the slopes at
these points based on an assumed linear growth rate
from conception to day 30 (slope 5 0.03) and the calcu-
lated linear trend of the IWS data from day 82 to day
245 (slope 5 2.67). The resulting spline allows assign-
ment of an estimated TL for fetuses between gestational
ages of 30 and 82 days. This reasoning predicts that
AMNH 63654, a fetus with TL5 13 cm, has a gestational
age of about 65 days. Alternative assumptions of 0.5 or
2.0 cm TL at day 30 change this prediction by less than
1 day (Fig. 7).
719
TL at 30 days of gestation (0.5, 1.0, and 2.0 cm) are presented.
Grayed lines represent periods of growth below day 30 (Period 1) and
after day 82 (Period 3) modeled with different assumptions. Specimen
diagrams, redrawn from the articles cited, have been resized to the
same scale and rotated to a consistent orientation. Scale bar 5 2 cm
for all specimens.
Comparison With Other Balaenoptera
Specimens
Previously described specimens of four species of
Balaenoptera are used to provide developmental context
for the morphology and estimated age of AMNH 63654.
The smallest of these overlap the size of the Stenella
embryos assigned to Carnegie Stages by Thewissen and
Heyning (2007), and the largest overlaps the low end of
the size range of the IWS fetal database for B. musculus.
This comparison assumes that morphology at a given TL
is similar across different species of the genus Balaenop-
tera during early development; the limited data avail-
able support this assumption. For example, a 38-mm
CRL B. physalus specimen (photo, Oelschl€ ager, 1989)
and a 40.3-mm CRL B. acutorostrata specimen (photo,
Amasaki et al., 1989) display the same head shape, rela-
tive rostrum length, eyelid aperture, and umbilical her-
nia size. Similarly, fetuses of B. borealis (photo, Schulte,
1916) and B. physalus (photo, Pilleri and Wandeler,
1962) of 375 and 380 mm TL display almost identical
external morphology, and B. musculus AMNH 63654 is
intermediate in both TL and morphology between the
720 ROSTON ET AL.
specimens of B. acutorostrata (Amasaki et al., 1989) and
B. physalus (Pilleri and Wandeler, 1962).
We calculate TL for the following comparative speci-
mens from rostrum tip to caudal midline tip using scales
published with the original figures. This procedure gen-
erates measurements from consistent landmarks across
all prenatal sizes, but overestimates TL in embryos and
small fetuses with marked ventral curvature. We predict
gestational age on the basis of TL for all of the speci-
mens, and also on the basis of CRL for the two speci-
mens with marked ventral curvature. Representatives of
each size group are diagrammed in Fig. 7.
1. B. musculus embryo (photo) figured by Oelschl€ ager
(1989) with stated CRL 5 31 mm, calculated TL 5 79
mm. Estimated gestational age based on CRL 5 51
days; estimated gestational age based on TL 5 59
days. Ventral flexion is marked in both the head and
the tail.
2. B. acutorostrata embryo (photo) figured by Amasaki
et al. (1989), with stated CRL5 40.3 mm, calculated
TL 5 86 mm. Estimated gestational age based on
CRL 5 54 days; estimated gestational age based on
TL 5 60 days. Ventral flexion is marked in the head
and muted in the tail.
3. B. acutorostrata embryo (photo) figured by Amasaki
et al. (1989), stated CRL 5 83 mm, calculated
TL 5 112 mm, estimated gestational age 5 63 days.
4. B. physalus early fetus (photo) figured by Pilleri and
Wandeler (1962). Using the published scale, the calcu-
lated TL 5 132 mm, predicting a gestational age of 65
days.
5. B. physalus fetus (photo) figured by Laws (1959) with
stated crown (not rostrum) - tail tip length of 18.5 cm
and calculated TL 5 244 mm. Estimated gestational
age 5 73 days.
6. B. acutorostrata fetus (drawing) figured by Eschricht
(1849). Based on the stated publication at “natural
size” and a vertex to fluke measurement of 8 inches
(Eschricht 1849), calculated TL 5 258 mm, and esti-
mated gestational age 5 73 days.
7. B. borealis fetus (photo) figured and minutely
described by Schulte (1916). Schulte (p. 395) cited two
TL values, 375 mm when the specimen is “straight”
but 450 mm when “the curvature of the dorsum is
included.” Citing preservational deformation, Schulte
used the smaller value in his proportional calcula-
tions. This smaller TL predicts a gestational age of 79
days, and the larger predicts 82 days.
8. B. physalus fetus (photo) figured by Pilleri and Wand-
eler (1962) with CRL listed as 31 cm, calculated
TL 5 380 mm, and estimated gestational age 5 79
days.
9. B. physalus fetus photographed, but not individually
described, by Pilleri and Wandeler (1962). The pub-
lished TL is 470 mm, estimated gestational age 5 83
days.
Transitions in development can be inferred from the
differences in morphology among these specimens. In
the smallest specimens, progressive changes with
increasing TL include the reduction of ventral curvature
of the head and tail, elongation of the rostrum, reduction
of the size of the umbilical hernia, decrease in the angle
between the forelimb and main body axis, and transition
from a paddle-shaped to a flipper-shaped forelimb. The
eyes are open at the smallest TLs, but are covered by
fused lids as size increases. In 58 and 61 mm specimens,
the urogenital sinus is confluent with the anus, but
Amasaki (1989) identified both urogenital/anal separa-
tion and made a definitive sex determination for a 112-
mm B. acutorostrata specimen.
In the mid-size range, AMNH 63654 (130 mm) exhib-
its modest ventral curvature and partially opened eye-
lids. Internally, dermal ossification has begun, although
the endochondral skeleton is completely cartilaginous.
The umbilical hernia is absent. The nasopharynx, lar-
ynx, and heart all exhibit anatomical specializations
characteristic of adult Balaenoptera. Anteriorly, poste-
rior elongation of the maxilla signals the earliest stages
of cranial telescoping. AMNH 63654 falls within the size
range (71–161 mm) and also meets the key anatomical
criterion (umbilical hernia retraction) of Fetal Stage 20,
defined as the first stage of fetal development by The-
wissen and Heyning (2007) in Stenella. The similarities
in size and external morphology between mysticetes and
Stenella argue for adoption of these criteria in mysti-
cetes as well, and identify AMNH 63654 as a very early
fetus. The timing of caudal fluke development is clarified
by comparison of AMNH 63654 to larger specimens.
Although only marginally longer than AMNH 63654, the
132 mm TL B. physalus TL specimen (Pilleri and Wand-
eler, 1962) has a minimally formed caudal fluke that is
longer than broad, and the fluke of the 244 mm B. phys-
alus specimen (Laws, 1959) is still of immature propor-
tions. All of these specimens lack a dorsal fin.
Each of the three specimens above 300 mm TL pos-
sesses a caudal fluke that mimics those of adults in pro-
portions and a dorsal fin. There is a progressive
reduction of the angle between the forehead and rostrum
and between the forelimb and the body axis (Pilleri and
Wandeler, 1962). Pigmentation has increased, the eyelids
are at least partially unfused, and sensory hairs occur
on the rostrum (Schulte, 1916; Pilleri and Wandeler,
1962). Dissection of the 375 mm B. borealis specimen
(Schulte, 1916) revealed partial endochondral ossifica-
tion of the skull, vertebral column, and long bones of the
forelimb, as well as telescoping of both anterior and pos-
terior cranial elements.
Comparison With the Developmental Sequence
of Stenella
The sequence of morphological transitions described
above suggests that developmental parallels between
mysticetes and odontocetes vary with age. Embryos of
Balaenoptera and Stenella are similar in morphology,
body size, and estimated age. For example, the shape
and differentiation of the hand, the shape and relative
length of the rostrum, and the structure of the umbilical
hernia are all comparable in Stenella (Carnegie Stage
18, Thewissen and Heyning, 2007) and Balaenoptera
(Oelschl€
ager, 1989) specimens with CRLs of 21–31 mm
and estimated gestational ages between 40 and 60 days.
These similarities are remarkable given the antiquity of
mysticete/odontocete common ancestry and the large dif-
ferences in both neonate and adult body size.
By the embryo/fetal transition, however, the effects of
accelerating growth in Balaenoptera are beginning to
appear. Although there are many parallels in size and
external anatomy, the first appearances of the caudal
 BLUE WHALE FETAL ANATOMY
fluke and dorsal fin are delayed in Balaenoptera relative
to Stenella (Carnegie Stage 17, 12–23 mm CRL). In addi-
tion, estimated gestational ages at this size and mor-
phology are somewhat shorter for Balaenoptera (65
days) than for Stenella (70–110 days, Moran et al.,
2011). Study of AMNH 63654 also indicates that distinc-
tive mysticete nasopharyngeal, laryngeal, and heart
anatomy are present internally. The remainder of early
fetal growth is characterized by continued dermal ossifi-
cation, the onset of endochondral ossification, caudal
fluke remodeling, dorsal fin formation, and skull tele-
scoping. These observations reinforce earlier suggestions
(e.g. Frazer and Huggett, 1973; Lockyer and Waters,
1986) that the accelerated growth responsible for large
mysticete neonate size occurs principally in the fetal,
and not the embryonic, phase of development.
ACKNOWLEDGEMENTS
We thank Darrin Lunde and Eileen Westwig of the
American Museum of Natural History for access to the
specimen and for searches of the museum documenta-
tion of its original collection. Charlie Potter of the
National Museum of Natural History provided access to
additional comparative fetal mysticete specimens. We
also thank Mei-Yee Chan of Wellesley College and Rich-
ard J. Cleary of Bentley University, who helped enter
and analyze the IWS data set, and Louis Buchholtz of
California State University, Chico, who set up the cubic
spline to estimate early fetal ages.
LITERATURE CITED
Amasaki HH, Ishikawa H, Daigo M. 1989. Development of the
external genitalia in fetuses of the southern minke whale, Balae-
noptera acutorostrata. Acta Anatomica 135:142–148.
Beauregard H. 1883. Recherches sur l’enc ephale des Balaenides.
Journal de l’anatomie et de la physiologie normales et pathalogi-
ques de l’homme et des animaux 19:481–516.
Buchholtz E. 2011. Vertebral and rib anatomy in Caperea margin-
ata: implications for evolutionary patterning of the mammalian
vertebral column. Marine Mammal Sci 27:382–397.
Burfield ST. 1912. Report of the committee appointed to investigate
the Biological Problems incidental to the Belmullet Whaling Sta-
tion. 82nd Report to the British Association on the State of Sci-
ence for 1912:145–186.
de Burlet HM . 1914. Zur Entwicklungsgeschichte des Walsch€ adels
III. Das Primordialcranium eines Embryo von Balaenoptera ros-
trata (105 mm). Morphologishes Jahrbuch 49:119–178.
Daubt W. 1898. Beitr€ age zur Kenntnis des Urogenitalapparates der
Cetaceen. Jenaische Zeitschrift f€ ur Naturwissenschaft 32:231–
313.
DesC^oteaux L, Colloton J, Gayrard V, Picard-Hagen N. 2010. Bo-
vine pregnancy. In: DesC^ oteaux L, Gnemmi G, Colloton J, editors.
Practical atlas of ruminant and camelid reproductive ultrasonog-
raphy. Ames: Wiley-Blackwell. p 81–98.
Drews B, Hermes R, G€ oritz F, Gray C, Kurz J, Lueders I, Hilde-
brandt TB. 2008. Early embryo development in the elephant
assessed by serial ultrasound examinations. Theriogenology
69:1120–1128.
Eschricht DF. 1849. Untersuchungen u € ber Die Nordischen Wall-
thiere. Vol. 1. Leipzig: L. Voss. 205p.
Evans HE, Sack WO. 1973. Prenatal development of domestic and
laboratory mammals: growth curves, external features and
selected references. Anatomy, Histology, Embryology 2:11–45.
Frazer JFD, Huggett ASG. 1973. Specific foetal growth rates of
cetaceans. J Zool Lond 169:111–126.
721
Frazer JFD, Huggett ASG. 1974. Species variations in the foetal
growth rates of eutherian mammals. J Zool Lond 174:481–509.
Gill EL. 1927. An early embryo of the blue whale. Trans R Soc
South Africa 14:295–300.
Hildebrandt T, Drews B, Gaeth AP, Goeritz F, Hermes R, Schmitt
D, Gray C, Rich P, Streich WJ, Short RV, Renfree MB. 2007. Foe-
tal age determination and development in elephants. Proc R Soc
B 274:323–331.
Hinton MAC. 1925. Reports on papers left by the late Major G. E.
H. Barrett-Hamilton relating to the whales of South Georgia.
Crown Agents for the Colonies, London, 1925:57–209.
Huggett ASG, Widdas WF. 1951. The relationship between mamma-
lian foetal weight and conception age. J Physiol 113:306–317.
Kida MY. 1998. Morphology of the tracheobronchial tree and the
route of the pulmonary artery in the fetal Minke whale (Balae-
noptera acutorostrata). Okajimas Folia Anatomica Japonica
75:251–258.
Koyabu D, Maier W, S anchez-Villagra MR. 2012. Paleontological
and developmental evidence resolve the homology and dual em-
bryonic origin of a mammalian skull bone, the interparietal. Proc
Natl Acad Sci 109:14075–14080.
Laws RM. 1959. The foetal growth rates of whales with special ref-
erence to the fin whale, Balaenoptera physalus Linn. Discovery
Reports 29:281–308.
Laws RM. 1960. Researches on the period of conception, duration of
gestation and growth of the foetus in the fin whale, based on data
from International Whaling Statistics. Comments on C. Naaktge-
boren’s, E. J. Slijper’s and W. L. van Utrecht’s Article. Norsk
Hvalfangst-Tidende 3:216–220.
Lockyer C, Waters T. 1986. Weights and anatomical measurements
of northeastern Atlantic fin (Balaenoptera physalus Linnaeus)
and sei (B borealis, Lesson) whales. Marine Mammal Sci 2:169–
185.
Mackintosh NA, Wheeler JFG. 1929. Southern blue and fin whales.
Discovery Reports 1:257–529.
Marino L. 1997. The relationship between gestation length, enceph-
alization, and body weight in odontocetes. Marine Mammal Sci
13:133–138.
Millar JS. 1981. Pre-partum reproductive characteristics of euther-
ian mammals. Evolution 35:1149–1163.
Miller GS, Jr. 1923. The telescoping of the cetacean skull. Smithso-
nian Miscellaneous Collections 76:1–68.
Moran MM, Nummella S, Thewissen JGM. 2011. Development of
the skull of the pantropical spotted dolphin (Stenella attenuata).
The Anatomical Record 294:1743–1756.
M€uller F. 1971. Comparative embryological investigation based on a
Balaenoptera physalus of 90 mm in length. Investig Cetacea
3:203–215.
Naaktgeboren C, van Slijper EJ. Utrecht WL. 1960. Researches on
the period of conception, duration of gestation and growth of the
foetus in the fin whale, based on data from International Whaling
Statistics. Norsk Havalfangst-Tidende 3:113–119.
Noakes DE. 2001. Development of the conceptus. In: Parkinson TJ,
England GCW, editors. Arthur’s veterinary reproduction and ob-
stetrics. Edinburgh: Saunders. p 57–68.
Oelschl€ager HA. 1989. Early development of the olfactory and ter-
minalis systems in baleen whales. Brain Behav Evol 34:171–183.
Pedersen JF. 1982. Fetal crown-rump length measurement by ultra-
sound in normal pregnancy. Br J Obstet Gynecol 89:926–930.
Pilleri G, Wandeler A. 1962. Zur Entwicklung der K€ orperform der
Cetacea (Mammalia). Revue Suisse de Zoologie 69:737–758.
Reese, CS, Calvin JA, George JC, Tarpley RJ. 2001. Estimation of
fetal growth and gestation in bowhead whales. J Am Stat Assoc
96:915–938.
Reidenberg JS, Laitman JT. 2007. Discovery of a low frequency
sound source in Mysticeti (Baleen whales): Anatomical establish-
ment of a vocal fold homolog. Anat Record 290:745–759.
Reidenberg JS, Laitman JT. 2008. Cetacean prenatal development.
In: Perrin WF, W€ ursig B, Thewissen JGM, editors. Marine Mammal
Encyclopedia. 2nd ed. Amsterdam: Academic Press. p 220–230.
Rice DW. 1983. Gestation period and fetal growth of the gray whale.
Report of the International Whaling Commission 33:539–544.
722 ROSTON ET AL.
Ridewood WG. 1923. Observations on the skull in foetal specimens
of whales of the genera Megaptera and Balaenoptera. Phil Trans
R Soc Lond B 211:209–272.
Risting S. 1928. Whales and whale foetuses. Conseil Permanent
International pour L’exploration de la Mer, Rapports et Proc es-
Verbaux des Reunions 1:1–122.
Schulte HW von. 1916. The sei whale (Balaenoptera borealis
Lessen). Anatomy of a foetus of Balaenoptera borealis. Memoirs
of the American Museum of Natural History New Series 1:389–
499.
Sears R, Perrin WF. 2008. Blue Whale, Balaenoptera musculus. In:
Perrin WF, W€ ursig B, Thewissen JGM, editors. Marine mammal
encyclopedia. 2nd ed. Amsterdam: Academic Press. p 120–124.
Slijper EJ. 1979. Whales. 2nd ed. London: Hutchinson.
Smets M, L’abbe G. 1885. Notes sur la t^
ete d’un foetus de Balaenop-
tera Sibbaldii (Gray). Annales de la Soci
et
e Scientifique de Brux-
elles 1885:187–204.
 erba O, Klima M, Schildger B. 2000. Embryology of dolphins:
St
staging and ageing of mammalian embryos and fetuses. New
York: Springer.
Thewissen JGM, Heyning J. 2007. Embryogenesis and development
in Stenella attenuata and other cetaceans. In: Miller DL, editor.
Reproductive biology and phylogeny of cetacea: Whales, dolphins
and porpoises. Enfield: Science Publishers. p 307–329.
Walmsley R. 1938. Some observations on the vascular system of a
female fetal finback. Contributions Embryol 164:107–178.
Weber M. 1904. Die S€ augetiere. Einf€
uhrung in die Anatomie und
Systematik der recenten und fossilen Mammalia. Jean: Gustav
Fischer Verlag.
Wisser J, Dirschedl P, Krone S. 1994. Estimation of gestational age
by transvaginal sonographic measurement of greatest embryonic
length in dated human embryos. Ultrasound Obstet Gynecol
4:457–462.
Woodward B. 2006. Locomotory strategies, dive dynamics, and func-
tional morphology of the mysticetes: Using morphometrics, osteol-
ogy, and DTAG data to compare swim performance in four species
of baleen whales. PhD thesis, the University of Maine.
Yochem PK, Leatherwood S. 1985. Blue whale, Balaenoptera muscu-
lus (Linnaeus 1758). In: Ridgway SH, Harrison R, editors. Hand-
book of marine mammals. Vol. 3. London: Academic Press. p 193–
240.
Zemskiy VA. 1950. Material for the study of the embryonic develop-
ment of Balaenoptera physalus L. [in Russian]. Bull Moscow Soc
Naturalists New Series 55:15–20.