Evidence of powered flight in Volaticotherium

antiquum
Abstract

Volaticotheres are a clade of eutriconodont mammals composed of Volaticotherium antiquum,

Ichthyoconodon jaworowskorum, Argentoconodon fariasorum, Jugulator amplissimus and
Triconolestes curvicuspis. V. antiquum bears a relatively well-preserved holotype specimen, displaying
several adaptations interpreted as indicating a gliding lifestyle, such as a patagium, long limbs and a
specialized femur adapted to deal with flight stresses. A. fariasorum displays similar post-cranial
features, thus also indicating aerial locomotion. However, volaticotheres differ from every other
known gliding synapsid clade in being carnivorous, cosmopolitan and relatively long lived. The femur
speciations seen in V. antiquum and A. fariasorum are unmatched by any gliding mammaliaform,
seeming hyperspecialized in comparison, and restrict terrestrial locomotion. Finally, I. jaworowskorum
occurs in a marine environment, its teeth showing few signs of degradation through aquatic
transportation, implying a close location to where the animal died. These are all signs that
volaticotheres were not gliders, but most likely active, powered flyers, rendering them a fourth
lineage of flying vertebrates alongside bats, birds and pterosaurs. Examinations and comparisons of
the V. antiquum holotype and its humerus with other mammals and quadrupedal flying vertebrates -
the eutriconodonts Jeholodens jenkinsi and Spinolestes xenarthrosus, the colugo Galeopterus
variegatus, the metatherians Petaurus breviceps and Sinodelphis szalayi, the rodents Glaucomys
volans and Anomalurus beecrofti, the bats Myotis myotis and Mystacina tuberculata and the
pterosaurs Anurognathus ammoni and Darwinopterus modularis - indicate a proportionally robust
upper arm, which may vindicate its status as a vertebrate capable of true powered flight.

Introduction

Volaticotherium antiquum is the type species for Volaticotheria, a clade of Mesozoic (Toarcian to
Cenomanian) mammals which was first erected with its description (Meng 2006). Another species,
Ichthyoconodon jaworowskorum, was described earlier in 1995 (Sigogneau-Russell 1995), but its
holotype was rather incomplete, composed of a pair of molar teeth. Nonetheless, the close
relationship between both taxa was recognised early on, the molar teeth being only distinguishable in
Ichthyoconodons less recurved cusps (Meng 2006). Soon, another taxa, Argentoconodon fariasorum
and Jugulator amplissimus, were recovered as related to Volaticotherium and Ichthyoconodon as well
(Gaetano 2011), as a sister-taxon to Volaticotherium and as an outgroup to the rest of the clade
respectively. In this arrangement, these mammals were recovered within the triconodontid subfamily
Alticonodontinae, and thus Volaticotheria was renamed Volaticotherini. However, soon after another
study recovered this clade as outside of the rest of Triconodontidae, and added Triconolestes
curvicuspis as a potential member (Averianov 2011). Most recently, Volaticotherium and
Argentoconodon were recovered as a clade outside of Triconodontidae (Martin 2015), lending
legitimacy to the use of Volaticotheria as the term to describe this clade over Volaticotherini.

Volaticotheres, like most Mesozoic mammals, are primarily known from teeth, specifically their highly
distinctive molars, bearing backwardly-oriented recurved cusps (Meng 2006, Gaetano 2011,
Averianov 2011, Sigogneau-Russell 2016). However, Volaticotherium antiquums holotype, IVPP
V14739, is a more complete specimen, bearing the skull, left hindlimb, portions of the left forelimb
and tail, as well as soft tissue impressions of the animals pelage and skin. Most notable is the
presence of a patagium, proportionally rather extensive and larger than that of the Oligocene gliding
rodent (Meng 2006). Covered in fur, this patagium bears wrinkles, presumably as a result of folding,
and extends to the hindlimbs, sandwiching them, and to at least the base of the tail as an
uropatagium (Meng 2006).

Combined with proportionally long limbs, dorsoventrally flattened tail and a peculiar femoral head
that restricts rotational movement in exchange for resisting flight stresses, Volaticotherium antiquum
was understood to be a gliding animal, an exceptional example in that it pre-dated both marsupial
gliders and bats for at least 70 million years and demonstrated an extent of the ecological diversity of
mammals in the Mesozoic (Meng 2006). Later on, a few post-cranial remains of Argentoconodon
fariasorum were uncovered, including a rather similar femur, indicating that it too was a glider
(Gaetano 2011). Other volaticotheres are so far known only from their molars, but they are
speculated to have similarly been gliders as well (Gaetano 2011).

However, volaticotheres are different from other gliding mammals in a variety of aspects. For
instance, like most eutriconodonts they are carnivorous; not only do they share adaptations to
carnivory seen in other eutriconodont mammals (Sigogneau-Russell 2016), Volaticotherium was
compared to insectivorous bats (Meng 2006), Jugulator was noted as having specialised in
proportionally large prey (Cifelli 1998) and in a study about mammalian jaw proportions
Volaticotherium has fallen with insectivores while Argentoconodon has fallen with
carnivores/omnivores (Grossnickle 2013). Meanwhile every other known clade of gliding synapsids is
predominantly herbivorous (Jackson 2012, Luo 2017), as gliding has evolved as means to cover larger
distances in search of static food objects (Jackson 2012).

Likewise, volaticotherians are among the most cosmopolitan of the Mesozoic mammals, with
Volaticotherium ocurring in Asia, Ichthyoconodon in Africa, Jugulator and Triconolestes in North
America and Argentoconodon in South America. The latter in particular bears mention as being not
only the only South American eutriconodont beside Condorodon (Gaetano 2012), but also among the
earliest known eutriconodonts (Gaetano 2011, Sigogneau-Russell 2016), suggesting that this clade
dispersed across the globe very quickly. They are also comparably long lived, with the youngest
representative, Jugulator, dating to the Cenomanian, at least 80 million years after the oldest
representative, the clearly aerial Argentoconodon, which contrasts most gliding mammal groups as
rather localised and brief lived clades (Jackson 2012).

Finally, Ichthyoconodon occurs in marine deposits, its teeth notably lacking degradation from aquatic
transportation, meaning that they have at most only been carried for a short distance at sea
(Sigogneau-Russell 1995). It was originally suggested to have been a piscivore (Sigogneau-Russell
1995), an interpretation that, in the light of incomplete remains and the presence of confirmed
aquatic eutriconodonts (Chen 2015) could still hold some legitimacy. However, the lack of
unambiguous speciations towards piscivory, coupled with the sheer rarity of the taxon, deems it more
likely that it wasnt an aquatic mammal.

The combined factors of carnivory, cosmopolitan distribution and appearance in marine deposits
without long aquatic transportation are highly atypical of gliding and terrestrial mammals, but are
consistent with truly volant animals. Indeed, volaticotheres closely parallel chiropterans in terms of
their rapid cosmopolitan expansion, predominantly carnivorous niches and otherwise poor fossil
record (Eiting 2009). Thus, rather than being passive gliders, these mammals were instead capable of
true powered flight, joining bats, birds and pterosaurs as a fourth lineage of winged vertebrates.

As the Volaticotherium holotype remains the most complete volaticothere fossil to date, it is pivotal in
understanding of the clades potential volancy. Several aspects of its description are consistent with
an interpretation as a powered flyer: its dentition is compared to that of insectivorous bats, as is its
body weight of 10g-1.5kg (Meng 2006), which could imply a similar lifestyle as an aerial insectivore.
Likewise, its membrane is noted as being proportionally large and possibly incomplete (Meng 2006),
which could be indicative of a conventional wing. Finally, its hand is noted as being incomplete and
bearing unusual phalanges (Meng 2006), a subject that will be discussed in greater detail on the
discussion segment.

In this study, I compare the humerus of Volaticotherium antiquum with that of other mammals as well
as that of two pterosaur taxa. As the humerus has been altered in pterosaurs and bats as a
consequence of quadrupedal launching (Mark Witton 2013), it should determine if volaticotheres are
similarly specialised, or bear similar deviations from the terrestrial mammal standard.

Materials and Methods

(A) Materials

Fossils include the holotypes of Volaticotherium antiquum, Spinolestes xenarthrosus, Jeholodens

jenkinsi and Sinodelphys szalayi and the more complete specimens of Darwinopterus modularis and
Anurognathus ammoni. Specimens from living species include museum specimens as well as other
public sources.

All animals with the exception of Galeopterus variegatus - included to represent more gliding taxa as
well as due to the purported similarities between colugos and chiropterans - fit within the 10g-15kg
range suggested for Volaticotherium antiquum. Both Jeholodens jenkinsi and Spinolestes xenarthrosus
are examples of eutriconodont mammals with well preserved forelimbs, the former an arboreal
species and the latter terrestrial. Sinodelphys szalayi is the sole non-gliding or flying therian, fairly
similar to Jeholodens due to shared arboreal habits but distinct enough to showcase differences
between eutriconodonts and therians. All other taxa are gliding or flying vertebrates.

The inclusion of Jeholodens jenkinsi and Sinodelphys szalayi is of particular interest because the
former has been compared to opossums (Qiang 1999), so any results from the comparison of both
taxa should weight on the description of eutriconodont forelimb anatomy.

Purposely avoided are birds and other theropods. As bipedal animals whose primary airfoil is
composed of dead tissue (scansoriopterygids nonwithstanding) and that rely on different muscle
groups that both bats and pterosaurs utilise, they might be red herrings or otherwise unsuited
comparisons.
(B) Methods

The humeri of all taxa will be illustrated and measured against each other. As nearly all animals except
Galeopterus variegatus share a similar body mass, differences in proportion are immediate. Specific
focus is given to the humeral head, which as the muscle attachment site as well as the centre of the
biomechanical forces in quadrupedal launch is the most prone to modification (Mark Witton 2013).

All specimens are presented dorsolaterally. Note that Volaticotheriums humerus is displaced, it is
presented as conventional expected by tetrapod standards, with the broader, flatter region assumed
to be the humeral head.

(C) Potential caveats

A concern frequently voiced when studying fossil specimens is whereas the bones have been altered
post-mortem. Volaticotherium antiquums holotype is indeed described as squashed (Meng 2006),
but x-radiographic scans, as well as the description of the animals tail and femur, suggest that the
presented shape is true to life and is not modified.

All other fossil specimens do not have this ambiguity, being exquisitely preserved and noted has
having proportions true to life (Qiang 1999, Martin 2015, Bennett 2007, Luo 2003)

Results

Volaticotherium antiquum

Jeholodens jenkinsi
 Spinolestes xenarthrosus

Galeopterus variegatus

Petaurus breviceps

Sinodelphys szalayi

Glaucomys volans
Anomalurus beecrofti

Myotis myotis

Mystacina tuberculata

Darwinopterus modularis
 Anurognathus ammoni

Of the taxa examined, Volaticotherium antiquum is most similar to Anurognathus ammoni, followed
close by Glaucomys volans. All three taxa bear robust humeri with deep humeral heads located
proximally on the humerus. Structurally, the head of Volaticotherium antiquum is slightly more similar
to that of Glaucomys volans, perhaps as expected from their shared mammalian ancestry. However,
proportionally it is much closer to Anurognathus ammoni, bearing similarly large humeral crests.

Bats and Darwinopterus modularis follow a similar trend. However, in the former the humeral joint,
while relatively expanded in regards to the rest of the shaft, is still negligibly small compared to the
rest of the humerus, and the humeral crests are located slightly more distally in Mystacina
tuberculata. By contrast, Darwinopterus modularis bears the largest humeral head, owing to the large
humeral crests observed in derived pterosaurs. The humeral crest is also located more distally than in
Anurognathus ammoni, Glaucomys volans and Volaticotherium antiquum.

Therian mammals aside from bats and Glaucomys volans all follow the same patterns, bearing small
humeral heads and shallow, slender crests, usually positioned more distally. Humeral shafts are
universally slender compared to other groups.

Eutriconodonts bear generally robust humeri compared to therians, but neither Jeholodens jenkinsi
nor Spinolestes xenarthrosus have large humeral heads. The former lacks a distinctive humeral head,
while the latter bears a small one, located distally.

Conclusion

Compared to other mammals, Volaticotherium antiquum bears a deep humeral head, large humeral
crests and a robust shaft. These features are fundamentally present in both pterosaurs and bats; they
are exaggerated in relation to bats, proportionally similar to those of anurognathids pterosaurs and
not as extreme as those of derived pterosaurs. It seems apparent, then, that Volaticotherium
antiquums features are consistent with those of true powered flyers.

The caveat is the humeral head of Glaucomys volans, which bears similar proportions, as opposed to
the less enlarged humeral heads of the other non-volant mammals. As these features are still less
pronounced than in Volaticotherium antiquum, I consider it closer functionally to flying vertebrates
than to Glaucomys volans. Instead, given how radically different this condition is from other gliding
mammals, I suggest further studies on Glaucomys volans biomechanics and behaviour.
Discussion

Volaticotheriums hand is described as incomplete (Meng 2006). We do know three metacarpals

with associated proximal phalanges; these are about as long as the former (and longer than their
pedal equivalents), bearing pronounced flexor sheath ridges and broadened distal ends. Hindlimb
phalanges also bear flexor sheath ridges, interpreted as granting a grasping function to the hindlimbs.
In the forelimbs, however, assuming these would not have also been used for climbing, they would
most certainly control the wing fingers, grasping the air much as modern bats do (Hedenstrm
2015). Bats thinned out their wing bones (Hedenstrm 2015), while volaticotheres, like pterosaurs
(Mark Witton 2013, Habib 2010), appear to have made at least their humerus more robust, meaning
that their flight style might have been a mosaic of the two.

Because Volaticotheriums hand is incomplete, any statements on the animals wing structure are
purely speculative. As the wing morphologies of birds, bats and pterosaurs are all radically different,
one would expect the volaticothere wing to be similarly unique. Here, I have proposed three
hypotheses:

(A) Chiropteran wing

Triconolestes curvicuspis (Cerri Thomas) and Ichthyoconodon jaworowskorum (Julio Lacerda) depicted
with wings superficially similar to those of bats.

In this scenario volaticotheres, like bats, elongated their phalanges to form the wing. As mentioned
before, Volaticotherium, like bats, has flexible digits, but the metacarpals and proximal phalanges are
not as elongated as those of bats. Nonetheless, it is possible that such wings would have have been
composed of elongated distal phalanges, which have so far not been preserved.

(B) Intermediary chiropteran wing

Argentoconodon fariasorum (Cerri Thomas) depicted with elongated wing fingers that still allow
terrestrial motion.
A novel idea is that volaticotheres, having only elongated their distal phalanges, could still use most of
the hand to walk and climb, as opposed to bats and pterosaurs that fold their wing fingers when
walking. In essence, this is a more specialized version of the condition seen in colugos, which have
elongated and webbed forelimbs but still use them to hang and climb.

(C) Styliform wing

Triconolestes curvicuspis (Julio Lacerda) and Ichthyoconodon jaworowskorum (Dylan Bajda) depicted
with extremely elongated, ossified styliform bones.

Most groups of gliding and flying vertebrates possess styliform bones, and in two particular instances,
in scansoriopterygids and flying squirrels, these are located on the wrists (Xu 2015). Thus, it is possible
that volaticotheres also had styliforms, and perhaps elongated and ossified them into forming wing
supports. This would be an exceptional case among flying vertebrates, since all other groups rely
primarily on their digits to support the patagium. The sheer lack of precedent, as well as indication
that Volaticoheriums phalanges were specialised for flight, renders this hypothesis relatively unlikely
for now.

No discussion on volaticothere wing structure would be complete without understanding how these
animals became volant. It is generally agreed that gliding does not lead to powered flight (Kaplan
2011), which is consistent with the observation that nearly all gliding mammals are herbivores and
thus potentially incapable of meeting the metabolic requirements for active flight (Jackson 2012, Luo
2017). Debates on bird flight origins have resulted in a myriad of possibilities from terrestrial hopping
(Garner 1999) to Wing-Assisted Incline Running (Dial 2003, Tobalske 2007), a debate that has not yet
been settled in any significant capacity. Much less has been done on the origins of pterosaur flight,
but the relationship between pterosaurs and Scleromochlus (Mark Witton 2013) and the fact that
basal pterosaurs have hindlimbs adaptations towards hopping (Mark Witton 2015) heavily suggests
that pterosaurs evolved from ground-dwelling hoppers. Speculations on bat flight origins have largely
been restricted to assuming that these animals evolved from gliders, but more recent studies have
instead suggested that, rather than undergoing a gliding stage, bat ancestors fluttered their forelimbs,
a motion still seen in juveniles bats that have not yet developed the ability to fly, and therefore it can
be assumed that bat flight was evolved through this unique method of parachuting rather than gliding
(Kaplan 2011).

Volaticotheres, lacking any sort of intermediate stages and forming an isolated clade compared to
other eutriconodonts (Averianov 2011, Martin 2015), and largely lacking in post cranial remains aside
from Volaticotherium and Argentoconodon, are in an even worse position. Based on Volaticotherium
antiquums pedal anatomy it was possibly arboreal, though whereas this is the ancestral condition as
seen in bats or a later developed exclusive to the clade in question is unknown. For now, an arboreal
ancestry can be presumed on the basis of Volaticotheriums grasping digits, on the characteristic
femur that restricts terrestrial motion and possible chiropteran-like wing anatomy, suggesting that
volaticotheres became airborne through fluttering as well.

Volaticotherium antiquum bears pelage on its wing membranes. While most bats and pterosaurs
possess naked wings, and there are likely aerodynamical advantages for this, some bats such as
Lasiurus species do have fur covering large parts of their wing membrane (Shump 1982) and at least
some anurognthid pterosaurs possessing pycnofibers trailing the edges of their wings (Mark Witton
2013), so there is precedent. In these cases, it is speculated that fuzz on the wing membranes might
diminish air turbulence and thus lower the sound of wing strokes, allowing the animal to pursue prey
undetected.

Volaticotheres would have co-existed with anurognathid pterosaurs. For example, Volaticotherium
antiquum would have co-existed with Dendrorhynchoides mutoudengensis and Jeholopterus
ningchengensis (Wang 2006). Both lineages occupied a similar ecological niche as nocturnal, aerial
insectivores, much as bats and nightjars and similar birds do today. Competition between both groups
hasnt been extensively studied, but it is believed that the birds cope at lower populational densities,
being unable to outmanoeuvre bats. As anurognathids vastly outnumber volaticotheres in the fossil
reccord, the reverse might have happened in the Mesozoic. However, while all known anurognathids
are aerial insectivores (Mark Witton 2013), volaticotheres display a wider ecological range, with
Jugulator amplissimus and Argentoconodon fariasorum being vertebrate predators (Cifelli 1998,
Grossnickle 2013) and probably capable of hunting anurognathids.

The extinction of volaticotheres, like that of modern eutriconodont mammals, is probably tied to the
mid-Cretaceous floral turn-overs, which caused several changes in insect and vertebrate faunas.
Anurognathids also disappear around the same time, though both groups could have survived as
Lazarus taxa, given their fossil fossil record.

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