Sauropods were the largest animals ever to walk the earth, and evolved several specializations in their limbs in order to support their body mass. Their legs became columnar and their manual digits became reduced and encapsulated in tissue to form a single weight-bearing unit in the derived sauropods. A new three-dimensionally preserved cast of a sauropod manus, found in the Upper Jurassic Lourinhã Formation, Portugal, demonstrates not only the shape, but also the actual movements of the sauropod manus during the stride. The manus cast is 32 cm deep, and show the manus to be hoof-shaped and lacking any impressions of individual digits, except for digit I, the pollex. Well preserved striations from skin on the sides of the cast show that the manus was covered in rough, tubercular skin. The width of the manus cast is consistent from top to bottom, demonstrating that the manus was brought down and lifted vertically before any parasaggital movement of the upper limb took place.
Incorporating both ichnological and osteological
data is important when reconstructing extinct animals and formulating and testing hypotheses of
locomotory behavior and evolution. The discovery of well preserved fully articulated and conjugated sauropod manus specimens (Gilmore 1925, 1936), studies of osteology and functional morphology, and of numerous tracks and trackways (Lockley et al., 1994 and references therein) have demonstrated that at least neosauropod dinosaurs had a highly unusual manual structure (Fig.1). This comprises erectly positioned metacarpals, that were tightly interlocked proximally and formed a semicircle, as well as abbreviated phalanges that usually did not leave individual traces in even well preserved tracks. This has implications for sauropod locomotor evolution and functional morphology. In this paper we present new evidence for the shape and function of the sauropod manus, based on a three-dimensionally preserved cast of a sauropod manus from the Late Jurassic of
Sauropod manual morphology and evolution Sauropod dinosaurs were the largest terrestrial vertebrates in evolutionary history and thus faced severe problems with support of mass. The limb bones were sturdy and limb postures columnar, with inferred locomotory capabilities similar to elephants. The hind feet in all known taxa appear to have had a digitigrade posture with a posterior heel pad, as in elephants (Christiansen 1997). Forefoot postures and presumable functional morphology did, however, change markedly throughout evolution (Fig.1).
The most primitive sauropod known with a complete manus is Shunosaurus (Zhang 1988).
Shunosaurus has 3 carpals, decreasing in size mediolaterally, and the medial carpal is ﬂattened, although not block-like, having distinct articulating facets for the medial and lateral carpals (Zhang 1988, Christiansen 1997). The abbreviated phalanges (digital
formula 2-2-2-2-1) probably did not have much mobility, but carpal structure indicates more suppleness than in later forms (Christiansen 1997). Manual posture appears to have been semidigitigrade (Christiansen 1997, Wilson & Sereno 1998), and gently posteriorly concave. Omeisaurus also has 3 carpals (He et al., 1988, Christiansen 1997), and these were more block-like, showing greater size heterogeneity, the medial one being the largest. Lavocat (1955) reported on 3 carpals in a »Bothriospondylus« from Madagascar, but this ﬁnd has yet to be studied in detail. All later sauropods, Neosauropoda, (Wilson
& Sereno 1998) had but two, distinctly block-like carpals and highly abbreviated digits, albeit with a prominent ungual on digit I. Their metacarpals were maintained in an erect posture, tightly bound together proximally and formed a wide, U-shaped semicircle, as indicated by long, intermetacarpal articulating
facets (Christiansen 1997, Wilson & Sereno 1998),
fully articulated specimens (Gilmore 1925, 1936), and
numerous trackway ﬁnds. Some genera, such as
Janenschia, Brachiosaurus (Janensch 1922, 1961) and
Apatosaurus (Gilmore 1936) appear to have had only
one ossiﬁed carpal.
Titanosauriformes (Wilson & Sereno 1998) supposedly had a reduced ungual on digit I (Salgado et al.,
1997) but this character is ambiguous as its most
basal member, Brachiosaurus, did indeed have a
small ungual (Fig. 1, see also Upchurch 1994), but the basal titanosaurian Janenschia did not (Fig. 1, see also Janensch 1922, 1961). Supposedly Titanosauridae lacked not only a digit I ungual, but ossiﬁed manual phalanges altogether (Salgado et al., 1997), as evidenced by the absence of digits recovered with even well preserved specimens, such as Opisthocoelicaudia (Borsuk-Bialynicka 1977) and Epachthosaurus (Salgado et al., 1997). This graphically illustrates that the neosauropod manus was adapted for columnar support of mass, having little mobility and a limited contribution to forward propulsion relative to the hind limb (Christiansen 1997). The studies of osteology predict that sauropods initially retained some manual ﬂexibility and that sauropod evolution favoured support of mass over ﬂexibility. The manual morphology of neosauropods must have severely restricted mobility, and the manus appears to have been largely inﬂexible. The forelimb was essentially columnar during support of mass (Bonnan 2003),and the main propulsive force came from the hind limbs, unlike any extant quadrupedal animals, and probably early sauropods as well (Christiansen 1997). Neosauropods thus evolved not only a manual morphology that was different from more primitive forms, but also changed their locomotor mechanics.
Thus, neosauropods could have touched the ground
and subsequently lifted their forefeet in a near vertical manner, with the main ﬂexion mobility being in
the elbow, very different from extant animals, even
elephants (Gambaryan 1974, Christiansen 1997).
The new three-dimensionally preserved manus cast
New evidence for the shape and function of the
sauropod manus comes from a deep, three-dimensionally preserved, natural cast of a sauropod manus
found in the Upper Jurassic (Tithonian-Kimmeridgian) Lourinhã Formation, central west Portugal,
during ﬁeld work the summer 2003. The Lourinhã
Formation is part of the Lusitanian Basin and consists of approximately 140m of terrestrial sediments,
deposited during the initial rifting of the Atlantic in
the Kimmeridgian and Tithonian. The sediments
consist mainly of thick red and green clay layers,
interbedded with massive ﬂuvial sandstone lenses and
heterolithic horizons. The sandstone lenses appear as horizontally extensive, lenticular beds; some are traceable for several kilometres along the sections exposed along the coast. The sandstone lenses have been interpreted as distal alluvial fan facies originating from periods of extensive faulting (Hill 1989).
The Late Jurassic sediments of Portugal have yielded a rich dinosaurian fauna, including numerous sauropod remains as well as extensive ﬁnds of tracks and trackways. The ﬁrst dinosaur fossils were collected in 1863 in Porto das Barcas, near Lourinhã. Since then,
at least ﬁve sauropod forms have been recognized in Portugal: Lusotitan atalaiensis, Camarasaurus sp., Apatosaurus sp., Lourinhasaurus alenquerensis and Dinheirosaurus lourinhanensis (Antunes & Mateus 2003).
The new track was found in-situ in the coastal cliffs west of Lourinhã and is preserved as a ﬁne-grained sandstone cast in a layer of silt and clay. The cliff section at the site is vertical due to continuous coastal erosion. Additional tracks from the trackway are still buried in the cliff section and are inaccessible on account of approximately 20 metres of verburden.
Several less well-preserved casts of tracks have been found under similar circumstances in the coastal cliffs in the Lourinhã area.
The cast was collected and is stored at Museu da Lourinhã (ML 965). The depth of the cast is 32 cm, measured from the original tracking surface, and it is 45 cm wide. A horizontal fracture approximately in the middle of the cast caused it to split during
excavation. When viewed in horizontal section, as allowed by the crack, the cast revealed a distinct semi-lunate shape where the whole front of the manus is smooth and semi-circular, lacking any indication of free digits, except from the posteriomedially orientated pollex claw, which is present as a narrow triangular protrusion along the side of the cast
(Fig. 2). The sides and front of the manus cast are covered with prominent, vertical, 2–10mm wide striations ﬂanked by 1–2mm deep furrows, originating from the movement of a rough and tubercular skin through the sediment (Fig. 3).
Evidence from well preserved Middle Jurassic sauropod tracks and trackways (Santos et al., 1994) show the shape of the sauropod manus to be semilunate and lacking impressions of free digits except for digit I. Trackways from the Upper Cretaceous of Bolivia, purported to be titanosaurid, display evidence of separate, albeit very abbreviated manual digits (Lockley et al., 2002). These, however, appear more like depressions in the bottom of the tracks and do not seem to affect the outline shape of the manus
imprint. Additionally, the impressions of digit I in these tracks indicate a very short pollex claw.
In the cast described in this study, the impression of the pollex claw is proportionally smaller relative to manus size than in similar sauropod tracks from the Middle Jurassic of Portugal (Santos et al., 1994). This shows that the trackmaker possesed a relatively
small pollex claw, consistent with the brachiosaurid
manual morphology (Fig. 1, see also Upchurch 1994).
The abundance of brachiosaurid material found in
the area (Antunes & Mateus 2003) together with the
manus shape, suggests a brachiosaurid origin of the
track. Because the cast was found as a single ﬁnd,
and the circumstances prohibited excavation of
further tracks from the trackway, it is not possible to
refer the track to a wide, intermidate or narrowgauge trackway.
Sauropod tracks usually fall into one of two categories: either a »narrow-gauge« type in which the respective tracks from the left and right sides intersect the midline of the trackway, or a »wide-gauge« type, where the imprints lie further away from the midline (Farlow 1992). The wide-gauge trackways ﬁrst occur in the Middle Jurassic Galinha tracksite in Portugal (Santos et al., 1994) and the Middle Jurassic Ardley Quarry in England (Day et al., 2002, 2004) and become dominant up through the Cretaceous
(Wilson & Carrano 1999). The temporal distribution and analyses of anatomy (Wilson & Carrano 1999) support a titanosaurian origin for the widegauge trackways. Based on evidence from the Galinha and Ardley Quarry tracksites, Day et al. (2004)
suggests four different morphologies of sauropod trackways: (1) narrow-gauge trackways showing a well-developed pollex claw impression, supposed to be of non-titanosauriform origin; (2) intermediategauge trackways with either well-developed or reduced pollex claw impression, formed by brachiosaurs or the most basal titanosaurs; (3) wide-gauge trackways characterized by reduced pollex claw impression and formed by basal titanosaurs like Janenschia and (4) wide-gauge trackways showing no indication of manual digits, formed by advanced titanosaurs. The shape and dimensions of the herein described cast of a manus track is constant from top to bottom, demonstrating the absence of a horizontal component in the lifting of the manus. Even the width of the pollex claw impression is constant from top to bottom of the cast, showing that the manus was brought down vertically, maintained vertical in the weightsupporting phase and subsequently lifted clear of the sediment before any parasaggital movements of the
The depth and shape of the present cast yields evidence about the locomotor mechanics of the particular sauropod that made the track. When tracks are emplaced in deep substrates, any horizontal component of the foot movements during the stride will result in an elongate morphology of the tracks, as demonstrated by Upper Triassic theropod tracks from East Greenland (Gatesy et al., 1999). In two of the Middle Jurassic sauropod trackways described from the Ardley Quarry, England, the manus impressions occurred as elongated disturbed structures indicating the manus was dragged through the mud during the stride (Day et al., 2004). On the contrary, deep well preserved, Upper Cretaceous, sauropod tracks from Humaca, Bolivia are 15-20 cm deep, and the manus tracks appears as steep-walled cavities with no sign of deformation from horizontal movements (Lockley et al., 2002). Upper Jurassic tracks from track level 5 (Lockley et al., 1994, Meyer et al., 1994), Cabo Espichel, Portugal, have sauropod manus tracks in excess of 20 cm depth, again appearing as steep-walled cavities with no indication of a horizontal component in the stride (J.M. pers. obs.). These examples from tracks reﬂects an evolution in sauropod locomotor mechanics from the Middle Jurassic tracks from the Arden Quarry (Day et al., 2004), where the manus is dragged forward, to the Upper Jurassic (Lockley et al., 1994, Meyer et al., 1994) examples, the Upper Cretaceous examples (Lockley et al., 2002), and the herein described cast, where the manus is emplaced and lifted vertically clear off the ground.
Eusauropods, like Shunusaurus from the Middle Jurassic had a semi-digitigrade manus stance (Christiansen 1997, Wilson & Sereno 1998) and are likely trackmakers to the manus tracks who show a forward drag of the foot, while the Neosauropods,
which appear from the Upper Jurassic and onward, has the more erect and reduced manus, likely to have made the tracks where the manus is lifted vertically clear off the ground.
The striations from the skin tubercles preserved on the sides of the cast of the manus track are similar to, although larger than, striations known from theropod and ornithopod tracks (Gatesy 2001, Currie et al., 2003) and from tracks of the extant emu (Milàn in press). This demonstrates uniformity in the pedal skin texture between a variety of non-avian and avian dinosaurs.
Based on the evidence herein presented, the brachiosaurid manus should be reconstructed with digits II to Vencapsulated by tissue to form a semilunate unit, with only a short digit I, the pollex, free and the skin texture roughly tubercular (Fig. 4).
A newly found three-dimensionally preserved cast of a manus track, probably from a brachiosaurid, demonstrates that the manual digits of advanced sauropod dinosaurs were encapsulated in tissue to form a hoof-like unit. Only digit I, the pollex, was separated and had a short triangular claw. The cast shows that the manus was brought down and lifted
vertically without any parasaggital components of movements before it was lifted well clear of the ground, and thus the entire forelimb retained its columnar form during the weight supporting phase, as predicted from anatomy, and here conﬁrmed by a well preserved ichnofossil.