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Ulisse Aldrovandi (1522-1605): The Study of Trace Fossils During the

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 Ichnos, 16:245–256, 2009
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ISSN: 1042-0940 print / 1563-5236 online
DOI: 10.1080/10420940902953205
Ulisse Aldrovandi (1522–1605): The Study of Trace Fossils
During the Renaissance
Andrea Baucon
Geopark Naturtejo Meseta Meridional, Geology and Paleontology Office, Centro Cultural Raiano,
Idanha-a-Nova, Portugal
The Renaissance was a time of flourishing for literature, art
and science. A prominent figure in Renaissance science was the
Italian naturalist Ulisse Aldrovandi, considered today to be one of
the founding fathers of modern paleontology. Aldrovandi is known
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to have widely studied body fossils, but his work on trace fossils
has yet to be explored by science historians. This paper proposes
to critically analyze Aldrovandi’s approach to trace fossils, based
on his writings (most of which are still unpublished) and on the
illustrations that accompany them. In his studies, the Bologna-
born naturalist accurately describes such ichnological celebrities as
Gastrochaenolites and Cosmorhaphe and discusses a few theoretic
principles on the trace-making process. The study of the works of
Aldrovandi brings to light a fascinating and little-known source of
the history of ichnology.
Keywords Ichnology, Aldrovandi, Renaissance, Cosmorhaphe, Gas-
trochaenolites, History of Ichnology
INTRODUCTION
Ichnology is a rapidly evolving field of study and is
considered a relatively young scientific discipline. It is not an
easy matter to find historical references, either in vertebrate or
invertebrate palaeoichnology, before the 19th century. Although
trace fossils are sufficiently mentioned in folklore and popular
culture (Mayor, 2000; Mayor and Sarjeant, 2001; Mietto et al.,
2003; Carvalho and Cachão, 2005), only in the 19th century
can a definite, widespread scientific approach be detected.
According to Sarjeant (1987), the first scientific study of
tetrapod footprints dates back to around 1824 when Reverend
William Buckland described a trackway (Sarjeant, 1987; Mayor
and Sarjeant, 2001) that he had been given by Reverend Henry
Duncan of Ruthwell (Duncan, 1831).
Address correspondence to Andrea Baucon, Geopark Naturtejo
Meseta Meridional, Geology and Paleontology Office, Centro Cultural
Raiano, Av. Joaquim Morão, Idanha-a-Nova, 6060-101, Portugal.
E-mail: andrea@tracemaker.com
245
Shortly afterward came the studies of Sickler (1834) and
Kaup (1835) on Chiroterium, the discovery of the dinosaur
tracks known as the footprints of Noah’s Raven (see Lockley,
2002) and the famous studies of Hitchcock (Hitchcock, 1836,
1858; see Pemberton et al., 2007a) on New England ichnology
(see Sarjeant, 1987, for a more complete review of the history
of vertebrate ichnology).
The study of nonvertebrate trace fossils also coincides with
the 1800s and the dawn of ichnology. In his historical review
of ichnology, Osgood (1970) identifies three stages, the first
of which he calls the “age of fucoids.” During this period,
trace fossils were given a botanical interpretation, as exemplified
by the term “Fucoides” used to indicate branching, plant-like
structures such as Chondrites. The vast bibliography on the age
of fucoids includes “Flora Fossilis Helvetiae” (Heer, 1877), in
which the botanical interpretation is evident right from the title.
For a more complete historical and bibliographical review of
the age of fucoids, see Osgood (1970, 1975), Häntzschel (1975)
and Pemberton et al. (2007b).
The age of fucoids ended in 1881 (Osgood, 1970), with
the neoichnological experiments conducted by Nathorst (1881)
highlighting the similarities between “fucoids” and various
kinds of recent traces (this was the start of Osgood’s “period
of reaction”; an exhaustive essay regarding the experiments
of Nathorst can be found in Cadeé and Goldring, 2007).
Although initially received with heated opposition (Häntzschel,
1975, mentions De Saporta, Delegado and Lebesconte), this
approach cleared the way for the modern ideas on trace
fossils.
The historical review of ichnology clearly indicates the 19th
century as the dawn of the study of trace fossils. This gives rise
to a question: What was there before the dawn? What were the
origins of the study of trace fossils?
The present study starts from these questions and proposes to
bring to light one of the most obscure and compelling chapters
in the history of ichnology—the studies of Ulisse Aldrovandi
on trace fossils.
 246
ULISSE ALDROVANDI: BETWEEN LEONARDO
AND GALILEO
“[. . .] & anco la Giologia, ovvero de Fossilibus” (and also
Geology, that is, the Science of the things found underground”).
(Aldrovandi, 1603)
The word “geology” is one of the most conspicuous legacies left
us by Ulisse Aldrovandi. The term was coined by Aldrovandi
at the time he was drawing up his will (Aldrovandi, 1603;
see Vai and Cavazza, 2003, Vai and Caldwell, 2006). Not
only did Aldrovandi define the word geology—he made wide-
ranging studies on “fossilia,” meaning all that can be found
underground (rocks, minerals, fossils). Notwithstanding the
importance of his studies on “fossilia,” geology was only one
of Aldrovandi’s many interests; large portions of his works are
dedicated to zoology, botany and medicine. The amplitude of
his interests defines Aldrovandi as a true son of his time, the
Italian Cinquecento, when great minds pursued a universal,
all-encompassing knowledge (cf. Olmi, 1976).
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The Cinquecento (or the 16th century) was a period of
renewal for science, marked by the brilliant intuitions of
Leonardo da Vinci and enshrined in the modern scientific
method laid out by Galileo. Ulisse Aldrovandi appears as an
intermediate figure between Leonardo and Galileo, along the
ideal line of continuity linking the culture of the Middle Ages
to that of the Renaissance (cf. Olmi, 1976, p. 18). Aldrovandi’s
outlook is directed both to the future and to the past: while
anticipating certain aspects of the Galilean revolution, at the
same time he resorts to notions that are scientifically retardative
(Vai, 2003a; Vai, 2003b; Vai and Cavazza, 2006).
Among the elements of innovation is the importance of
direct experience, repeatedly underlined by Aldrovandi. In his
Discorso Naturale (Aldrovandi, 1565), written in vulgar Italian,
Aldrovandi explains his hands-on approach: “non iscrivendo
cosa alcuna che co’ propri occhi io non habbi veduto e con
le mani toccato et fattone l’anatomia,” or “writing only about
what I have seen with my own eyes and touched with my own
hands, and examined both externally and internally.” Aldrovandi
(1565) admonishes Aristotle for not having personally verified
the information collected (see Pattaro, 1981). This shows the
critical position assumed by Aldrovandi toward classical authors
and how he felt it necessary to ascertain the truthfulness of their
data.
Yet Aldrovandi often fails to comply with his own method:
he grounds his observations on the “auctores canonici” and
intermingles scientific observations with encyclopedism and
pure erudition.
Although this approach may appear contradictory to the
modern-day scientist, it should be noted (Cassirer, 1967; Olmi,
1976) that during the 16th century the old and the new were
mutually interpenetrating and not fully independent from each
other. Thus, Olmi (1976) and Vai and Cavazza (2006) advise
A. BAUCON
the reader to shed any excessively rigid frames of interpretation
when approaching Aldrovandi.
As anticipated above, Aldrovandi’s works cover a wide
range of naturalistic subjects while providing ample evidence
of his profound humanistic culture. The scope and vivacity of
Aldrovandi’s interests are reflected in the episodes of his life,
and he acknowledges his own juvenile curiosity (Aldrovandi,
1565): “essendo io spinto dal dessiderio insin dalla mia prima
età di sapere [. . .]” (“being desirous of knowledge from early
childhood”). At university, Aldrovandi studied the humanities,
law, mathematics, medicine and philosophy, pursuing the ideal
of “universal knowledge” that was typical of the Renaissance.
Aldrovandi writes about his education in his Discorso Naturale
(Aldrovandi, 1565): “[. . .] havendo atteso alli principi necessarii
delle scientie et, con ogni diligenza, alle polite et belle lettere
et dopo il studio delle lettere humane, fondamenti certissimi
et solidissimi d’ogni disciplina, et per conseglio de gli miei
parenti [. . .] mi diedi alli faticosi studii delle leggi” (“having
acquired an education in science and literature, and a solid
fundament in every discipline, I followed my parents’ advice
and took up the study of law”). Aldrovandi exhibits his vast
knowledge of the humanities in his very first work, dedicated
to the statues of Rome. However, where he proved most
prolific was in the natural sciences. His writings on zoology
are especially bountiful, with a treatise on birds (Aldrovandi,
1599), snakes and other animals, on “bloodless” animals and on
insects (Aldrovandi, 1606). These works, as most of his writings,
were often accompanied by beautiful woodcut illustrations. To
Aldrovandi the art of illustration was more than mere ornament;
in the Discorso Naturale (Aldrovandi, 1565) he says that “to
understand plants and animals there is no better way than to
depict them from life” (“in verità non si puol’ fare più bella
impresa, per venire in cognitione di queste piante et animali
diversi, che depingerli vivamente”). Aldrovandi refers that he
practised drawing himself: (“non voglio già tacere me stesso,
che sopra modo di queste pitture varie mi sono dilettato,” “I
cannot conceal that I have dedicated much time to drawing”)
(Aldrovandi, 1565). However, he also admits to have often
commissioned the illustrations for his works to other painters
(“E’ ben vero che, per non dare impedimento alli miei studii,
ho avuto pittori appresso di me continuamente,” “in order not
to interrupt my studies, I have had many painters work for me”)
(from Aldrovandi, 1565).
Aldrovandi’s works were also illustrated with hand-colored
woodcuts, lately reprinted for their artistic value (Alessandrini
and Ceregato, 2007). Particularly noteworthy are the illustra-
tions of plants, recently reprinted in Antonino et al. (2003),
showing Aldrovandi’s deep interest in botany. The illustrations
made for Monstruorum Historia, picturing “monstrua,” or
wonders of nature, are justly renowned. It was Aldrovandi’s
desire to gather all he could find amazing or unusual in nature,
including fantastic creatures and freaks (Fig. 1).
 ALDROVANDI: TRACES IN THE RENAISSANCE
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FIG. 1. Aldrovandi was especially curious about nature’s more unusual
manifestations. A. Many of the “monstrous” images are based on the observation
of malformations and deformities. For example, Aldrovandi describes this
image as a “12-year-old hairy girl” (Aldrovandi, 1648); the subject is a girl
affected by hypertrichosis, a condition that is well known to modern medicine.
B. Other times the sources of Aldrovandi’s illustrations are more difficult to
identify, as for this “winged and horned monster, similar to a Cacodemon”
(Aldrovandi, 1648). C. In Serpentem et Draconem, Aldrovandi shows a marked
interest for herpetology, to the point of investigating the skeletal anatomy
of a number of snakes. D. Aldrovandi’s background in humanities emerges
frequently throughout his work. This figure is presented as a “seven-headed
Hydra,” explicitly referring to the Hydra of classical mythology and known to
Renaissance figurative imagery.
BODY AND TRACE FOSSILS IN THE MUSAEUM
METALLICUM
The work by Aldrovandi in which geology and paleontology
are discussed at the greatest length is the Musaeum Metallicum
(Aldrovandi, 1648). The subject matter of this work is evident
from the title, in which the Latin term metallicum bears reference
to mineral research. Aldrovandi himself gives the etymology
of the term (Aldrovandi, 1565, though not referring to the
Musaeum Metallicum): “[. . .] dal vocabolo greco µ
τ αλλ
υω,
che significa cercare et cavvare” (“from the Greek word
µ
τ αλλ
υω, which means to search and to quarry”).
It should be noted, however, that the title Musaeum
Metallicum was not chosen by Aldrovandi but by Bartolomeo
247
Ambrosini, who edited the book, which, like the majority of
Aldrovandi’s works, was published posthumously. Ambrosini
made a number of changes, including the title (see Marabini
et al., 2003; Alessandrini and Ceregato, 2007). The book’s
original title was De Fossilibus, clearly more relevant than
Ambrosini’s choice: the book describes and classifies “fossilia,”
or objects found underground, that is, minerals, rocks and
fossils. Marabini et al. (2003) point out that the title chosen
by Ambrosini recalls De Re Metallica, the well-known book on
mining by Agricola.
As previously mentioned, the Musaeum Metallicum is first
of all an immense collection of “fossilia,” but that is not its only
point of interest. Another notable feature is the author’s classi-
ficatory intention, which emerges from the organization of the
book’s content. The Musaeum Metallicum is divided into four
books, each corresponding to a specific group of “fossilia”: met-
als, soils, petrified fluids and rocks (Table 1). This division, prob-
ably inspired by Dioscorides and Agricola (see Marabini et al.,
2003), is also found in Aldrovandi’s manuscript Historia Fossil-
ium (unpublished, reproduced in part in Marabini et al., 2003).
Each book of the Musaeum Metallicum is, in turn, divided
into chapters, with each chapter dedicated to a particular
“family” of fossilia. For example, Book IV, “De Lapidibus” (On
Rocks), includes the following chapters: “De Lapide Arenario”
(On Sandstone), “De Lapide Calcario” (On Carbonate Rocks)
and “De Marmore” (On Marble).
Within the chapters the presentation often follows a descrip-
tive order that begins with “aequivoca, synonima, etymum,”
or improper use, synonyms and etymology of the term in
discussion. The exposition continues with a description of
the main features of the given “family” of fossils (“natura
ed affectiones”), followed by a list of its various “species”
(“differentiae”).
Hence, in the Musaeum Metallicum we can recognize both
a classificatory and a taxonomic intent; Sarti (2003) and
Vai (2003a, p. 80) rightly compare Aldrovandi’s systematic
approach to Linnaeus’. Yet while Aldrovandi’s classification
presents important elements of scientific renewal, the author
cannot be considered a progressive tout court: his body of
works displays the erudition and encyclopedism that were
characteristic of 16th century scholars.
The classification contained in the Musaeum Metallicum is
not the only one found in Aldrovandi’s works, nor is it the most
explicit. For example, a classification of the sciences is contained
in the unpublished manuscript Artium et Scientiarum Methodus
(reproduced and discussed in Pattaro, 1981), in addition to the
one in Historia Fossilium mentioned above. The latter deserves
particular attention because it contains an attempt to interpret
the “fossilia” that “generantur in animalibus, aquis, terrae” (“are
produced in animals, in water or in the ground”).
Aldrovandi does not limit himself to descriptions but tries
to offer interpretations (Fig. 2). This can also be noted in the
Musaeum Metallicum where, among other things, the origin
of fossils sensu stricto is discussed. Aldrovandi argues an
 248 A. BAUCON

TABLE 1
This plate shows the layout of the Musaeum Metallicum: four books divided in chapters, with each chapter presenting various
“differentiae” of fossilia. The layout is systematic and, for all practical purposes, equivalent to a classification: four orders
divided in families, and these divided in species. The plate shows a few examples of chapters and “differentiae”
Liber (Book) Capitulum (Chapter) Differentiae (“Species”)
I. De Metallis (metals) De Aere (copper) Aes nativum figurae pangoniae (faceted native copper)
Aes Corynthium figuratum (copper sculpture from Corynth)
De Argento (silver) Argentum Trichites sive capillare (“Trichites” silver, that is,
“hairy”)
Argentum purum nativum (pure native silver)
De Ferro (iron) Ferrum in Magnetem conversum (magnetized iron)
Siderammonites (“iron” ammonite)
II. De Terra (clays) De Terra figlina, et Argilla Argilla candida (white clay)
(terracotta and clay)
Vasa figlina Lusitanica adversus venena (Portuguese pottery
against poisons)
De Terra creta nuncupata Creta Mauritana (“creta” clay from Mauritania)
(“creta” clay)
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Creta Syriaca [. . .] (“creta” clay from Syria)

III. De Succis Concretis
(petrified fluids)
IV. De lapidibus (rocks)
De Ochra (ochre clay)
De Sale (salt)
De Bitumine (tar)
De Succino, seu Electro
(amber)
De Lapide Arenario
(sandstone)
De Glossopetra
(tongue-like-stones)
De Marmore (marble)
Genus crustosum (encrusting variety [of ochre clay])
Veronensis ([ochre clay] from Verona)
Stiriae salis (salt icicles)
Spumae salis (salt “foams”)
Durum (solid [tar])
Liquidum (liquid [tar])
Succinum colore vini falernii in forma annuli redactum (amber,
with the colour of wine and shaped as a ring)
Succino figurae cordis (hearth-shaped amber)
Durus (hard [sandstone])
Mollis (soft [sandstone])
Glossopetra denticulata (dentate tongue-like-stone)
Glossopetrae non denticulatae (non-dentate tongue-like-stones)
Marmor Dendrites (marble with dendrites)
Marmo Cynites (marble with dog-like stains)

articulated position on fossils and lists opposite views. In fact,
the Musaeum Metallicum shows frequent inconstancies between
fossils treated as ex vivi and others treated as inorganic:
1. Fossils as inorganic curiosities. Chapter 62 of the Musaeum
Metallicum supports an observation that fossils show no
trace of viscera and draws the inference that fossils are
of inorganic origin. According to this interpretation, fossils
are not ex vivi but merely natural curiosities imitating the
organic world: fossils are produced by fluids circulating
within rocks (Sarti, 2003). This view occurs frequently in
the Musaeum Metallicum: see the specimen in Figure 2A,
literally described as a “rock pregnant with a shell.”
2. Fossils as “ex-vivi.” Aldrovandi appears to put aside his in-
organic theory when discussing mammal teeth fossils: in this
case he speaks of “petrifaction” (“Tabella cum dente Belluae
petrificato”) and when describing the teeth of Hippopotamus
amphibus (Aldrovandi, 1648, p. 828: “Tabella represantat
dentes. . . lapideos Elephanti”). Aldrovandi seems to support
the organic origin of fossils also in his manuscript Historia
Fossilium (Vai and Cavazza, pers. com.).
As suggested by Vai and Cavazza (pers. comm.), the internal
inconstancies of the Musaeum Metallicum can be partly related
with the editor, Bartolomeo Ambrosini. In fact, the Museum
Metallicum was published posthumously by Ambrosini, who
 ALDROVANDI: TRACES IN THE RENAISSANCE 249
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FIG. 2. Body fossils pictured in the Musaeum Metallicum. A. Aldrovandi describes this specimen as a “rock pregnant with a shell.” Aldrovandi considers most
fossils to be of inorganic origin made in imitation of living beings. B. Although Aldrovandi believes that fossils are not of organic origin, he often compares
them to existing animals. He calls this structure “Rhombites,” meaning a “(stone) resembling a fish of the Rhombus kind.” C. Detail from the plate entitled
“belemnitarum septem differentiae” (seven varieties of “belemnites”). D. Aldrovandi calls shark teeth “glossopetrae,” or “tongue-like stones.” This specimen is
given the attribute “Gesneri,” or “Gesner’s”: naturalist Konrad Gesner (Gesner, 1565) had already described shark teeth as “Glossopetre” in 1565. E. Aldrovandi
often calls echinoderm fossils “astroitis,” a word that comes from the Latin “aster,” meaning star. The “star-echinoderm” comparison is most likely based on
echinoderms’ pentameral symmetry, resembling the stylized figure of a star. The comparison was already made by Gesner (1565) in his description of fossil crinoids
similar to those depicted by Aldrovandi and shown here. F. Aldrovandi frequently uses the term “ophiomorphites,” or “snake-like stones,” in his descriptions
of ammonites. G, H. Fossil sea urchins, presented as “astroitis” (see E). I. When describing this mammoth tooth, Aldrovandi speaks of “petrifaction.” See also
Vai and Cavazza (2006, fig. 14) J. Fossil coral, also presented as “astroitis” (star-stone), presumably because of the polyps’ stellate morphology. K. Aldrovandi
distinguishes two main types of “glossopetre”: dentate (as in D) and nondentate (as in K).

supported the inorganic origin of fossils and probably amended
some of Aldrovandi’s original descriptions.
SYSTEMATIC ICHNOLOGY OF THE MUSAEUM
METALLICUM
The Musaeum Metallicum is Ulisse Aldrovandi’s most
extensive and important work in geology and paleontology. It
contains hundreds of body fossils, frequently described in detail
and with magnificent illustrations. Together with body fossils,
Aldrovandi describes a good number of trace fossils, presenting
his own theories about their origin.
It might seem improper, or even provocative, to talk
about “systematic ichnology” when presenting the trace fossils
described in the Musaeum Metallicum. An ichnotaxonomic
analysis almost always requires the specimen to be examined
directly—sometimes even photographs can be misleading. Then
how do we justify speaking of systematic ichnology in the works
of Aldrovandi?
The first reason is the superlative quality of the iconographic
documentation. Many of the specimens pictured in the Museum
Metallicum are still preserved at the Museo Capellini and at
Palazzo Poggi in Bologna (see also Sarti, 2003), which makes it
possible to compare the original items with their representations
in the Musaeum Metallicum and evaluate the quality of the
illustrations. The comparison confirms the scientific value of the
geo-palaeontological illustrations in the Musaeum Metallicum,
given by their accuracy, wealth of detail and remarkable
 250
level of objectivity. Most of the illustrations examined were
virtually identical to the original specimens: see, for example,
the Ceraunias described in the Musaeum Metallicum and the
corresponding specimen of Charcharodon megalodon in the
Aldrovandi collection (Sarti, 2003).
The second reason for talking of systematic ichnology is
the value of the morphological descriptions in the Musaeum
Metallicum. Their accuracy was verified using the same method
applied to the illustrations, that is, by comparing the “fossilia” of
the Aldrovandi collection with the corresponding descriptions in
the Musaeum Metallicum. The comparisons show Aldrovandi’s
descriptions to be substantially objective, although a critical
attitude is always advisable. Aldrovandi does not always
distinguish between description and interpretation. For example,
he describes an ammonite as “ophiomorphites” (“rock similar
to a snake”) because of its snake-like morphology.
In conclusion, considering the objectivity of the illustrations
and the validity of the descriptions in the Musaeum Metallicum,
we can attempt a cautious ichnotaxonomic analysis of the trace
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fossils presented by Aldrovandi.
Gastrochaenolites
Diagnosis: The specimen is a rock (Fig. 3), “pitted here
and there by hollows of varying size” (“Erat [. . .] passim
sinubus diversae magnitudinis excavatus”; Aldrovandi, 1648).
Aldrovandi named it “Silicem dactylitem” because, he explains,
the “hollows” resemble the borings made by “Dactyli,” a genus
of lithophagous bivalves (the present-day date mussels). The
corresponding illustration confirms Aldrovandi’s description,
and allows a few additional observations. The “hollows”
FIG. 3. “De Silice” and Gastrochaenolites A. Gastrochaenolites pictured in Aldrovandi’s (1648) Musaeum Metallicum. B. Caption referring to the specimens of
Gastrochaenolites and Cosmoraphe shown in the Musaeum Metallicum: “plate 5, showing two varieties of flint.” C. This specimen, probably a limestone pebble
with a chert horizon, appears in the chapter entitled “De Silice.” Gastrochaenolites appears in the same chapter; as explained in the text, the borings are probably
made in a carbonate or siliciclastic rock.
A. BAUCON
mentioned by Aldrovandi have a circular cross-section, and from
the shading it can be surmised that they extend in an oblique
direction, probably ending with a clavate morphology.
Discussion: Aldrovandi’s “hollows” show evident morpho-
logical similarities with bioerosional structures, and the author
rightly compares them to the “hollows [. . .] resembling the
cavities in which date pholads (Pholas dactylus) seek shelter”
(“qui imitabantur illas cavitates, in quibus Dactyli animantes
delitescere solent,” Aldrovandi 1648). There is no doubt that
the author is explicitly referring to borings.
The borings described by Aldrovandi show features that
associate them with the ichnogenus Gastrochaenolites. The
circular cross-section, the oblique extension and the possible
clavate shape are fully consistent with the diagnosis of
Gastrochaenolites (Häntzschel, 1975).
In addition, it is quite possible that the specimen described
by Aldrovandi is a trace fossil and not a recent example of
bioerosion, because Aldrovandi clearly indicates the specimen’s
place of provenance as near “a Valley called Valdense, in
the territory of Siena” (“[. . .] perhibetur in Valley nuncupata
Valdense Agri Senensis”).
It is noteworthy that during the Renaissance Aldrovandi was
not the only Naturalist studying bioerosional trace fossils. Even
Leonardo da Vinci described bored fossil shells (Baucon, 2008):
“[. . .] the trace of the course [of the moving animal] is still
preserved on the shell that has been consumed in the same
manner of woodboring beetles [. . .]” (“[. . .] perchè ancora resta
il vestigio del suo andamento sopra la scorza che lui già, a uso di
tarlo sopra il legname, andò consumando”) (Leonardo da Vinci,
Leicester Codex, folio 9v).
 ALDROVANDI: TRACES IN THE RENAISSANCE
The Silex Problem: A problematic question arises from the
specimen’s (supposed) lithology. Aldrovandi describes it in the
chapter called “De Silice,” or “On Flint.” It is known that
in ancient times the meaning of the word “flint” was rather
vague. Aldrovandi (1648) is aware of this, “that the word flint
is employed more vaguely than would be desirable is evident in
Pliny, who used the word to designate both a rock and a variety
of marble.”
However, Aldrovandi (apparently) rejects the “vague” mean-
ing of flint and informs the reader that he will only discuss “true”
flint (“nos autem in praesentia de vero, et genuino Silice verba
facimus”) (Aldrovandi, 1648). Aldrovandi correctly mentions
some of the properties that characterize flint; he describes its
extreme hardness and reports that it gives off sparks when struck.
The presence of bioerosion on what is apparently a flint
pebble poses something of a problem, but a solution can be
found by examining Aldrovandi’s collections. Three specimens
in the collections are described in the Musaeum Metallicum
as “silex” (Silex quodammodo fungiformis, p. 727; Silex ex
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alveo Rheni Bonionensi, p. 739; Tabella cum novem Silicum
differentijs, p. 729) but are actually limestone pebbles (Sarti,
2003). Similarly, silex qui expressam a Natura crucem fert
pulcherrimam (Aldrovandi, 1648, p. 735) is actually a polished
serpentine pebble (Sarti, 2003). Together with these lithologies
we also find “true” flint, as in a flint nodule described as Silex
referens Impilium, genus tegumenti pedis (Aldrovandi, 1648, p.
740; cherty limestone formations are common in the Mesozoic
of the Alps and of the Appennines; Vai, pers. comm.).
FIG. 4. Cosmorhaphe A. Overall view of the plate with Cosmorhaphe and Gastrochaenolites (described above). B. Cosmorhaphe. This illustration is remarkably
realistic, thanks to its extreme detail and to the use of hatching to restore three-dimensionality and volume to the subject.
251
As can be seen from these examples, Aldrovandi uses the
term “silex” rather loosely, even applying it to the description
of certain types of limestone. In conclusion, the bioeroded
specimen is not made of flint but, in all likelihood, limestone.
Cosmorhaphe
Diagnosis: The plate on page 730 of the Musaeum Metal-
licum shows an unbranched structure with two orders of
meanders (Fig. 4). Aldrovandi (1648) describes this morphology
saying that the specimen “presented varied and curving stripes”
(“[. . .] erat variijs tenijs obliquis insignitus”). Both the illus-
tration and the description seem to indicate that we are dealing
with a convex relief. The surface presents parallel lineations and
sole marks.
This specimen appears on page 730 of the Musaeum
Metallicum, together with the Gastrochaenolites described
above. Aldrovandi (1648) presents the two trace fossils with
these words: “quinta tabella monstrat duas pulcherrimas Silicis
differentias,” or “the fifth figure shows two beautiful varieties
of flint.” As said when discussing the previous specimen, we
know that Aldrovandi uses the term “silex” (literally, “flint”)
to indicate different rocks, including certain types of limestone.
The structure in question does not seem attributable to a siliceous
rock and is more likely a carbonate or siliciclastic rock.
Discussion: The structure clearly shows two regular orders
of meanders, with no branching. These features are diagnostic
of the ichnogenus Cosmorhaphe, which is characterized by
regular, unbranched traces with two orders of meanders (after
 252
Häntzschel, 1975, and Seilacher, 1977). Cosmorhaphe is
frequently found as a convex hyporelief; this seems to be the
case for the structure in the Musaeum Metallicum, which is
associated with sole marks.
Describing and interpreting this structure, Aldrovandi
mentions curving stripes “quae prorsus figuras Serpentum
aemulabantur” (“that resemble snake figures”). It would
appear that he considers Cosmorhaphe as having an inorganic
origin, similarly to his suggestions regarding the body fossils.
Aldrovandi uses the verb “aemulare,” meaning “to imitate”;
hence, Cosmorhaphe would be a natural curiosity imitating the
sinuous curves of a snake.
Popular culture frequently refers to snake figures to de-
scribe meandering or winding trace fossils. In Portugal the
ichnogenus Cruziana is commonly associated with snakes
(see the “cobras”—snakes—of Penha Garcia, and the “bicha
pintada”—painted snake—of Milreu, in Carvalho and Cachão,
2005; some inhabitants of Penha Garcia are afraid of the
“snakes” that are Cruziana. Eddy Chambino, pers. comm.).
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Bioturbational ichnofossils also attracted the interest of
Leonardo da Vinci (Baucon, 2008), who described and in-
terpreted correctly trace fossils: “[B]etween one and another
rock layer, there are still the traces of the worms that crawled
in them when they were not yet dry. All the marine muds
still contain shells, and the shells are petrified together with
the mud” (“Come nelle falde, infra l’una e l’altra si trovano
ancora gli andamenti delli lombrici, che caminavano infra
esse quando non erano ancora asciutte. Come tutti li fanghi
marini ritengano ancora de’ nicchi, ed è petrificato il nicchio
insieme col fango.”) (Leonardo da Vinci, Leicester Codex,
folio 10 v.).
FIG. 5. “Stelechites.” Aldrovandi uses this term to indicate subcylindrical, arborescent elements. Some specimens of “Stelechites” represent trace fossils, but
most are concretions. A. “Three varieties of Stelechites,” fragments of a burrow. The specimens shown are currently preserved at Palazzo Poggi (Bologna). B.
Another specimen of Stelechites, possibly a trace fossil.
A. BAUCON
“Fragmentary Burrows” (Thalassinoides?;
Planolites/Palaeophycus?)
Diagnosis: The “Stelechites” described by Aldrovandi
(1648) are straight, subcylindrical structures (Fig. 5).
Aldrovandi writes: “[I]n nature we also find a stone that
resembles the trunk of a tree and shows the hardness of iron.”
(“Observatur etiam in rerum natura quidam lapis trunci alicuius
arboris speciem exprimens, & duritia ferrum aemulans.”) The
etymology of “stelechites” refers to their trunk-like shape
(Aldrovandi, 1648).
The “Stelechites” described in the Musaeum Metallicum can
still be seen in the Aldrovandi collections preserved at Palazzo
Poggi (Bologna). When arranging the geological material in
the Aldrovandi Museum, Sarti (2003) described “Stelechites”
as mineralized burrows of blackish color, slightly lighter on the
inside.
Discussion: These structures are not easy to classify, in lack
of any certain indication that might help to identify them as
trace fossils. The structures are fragmented and removed from
their geological context, making it difficult to assign them a
biogenic nature. However, the most plausible hypothesis is that
“Stelechites” are indeed trace fossils, and no evidence to the
contrary has emerged. Their interpretation as trace fossils is
also supported by Sarti (2003).
Assuming they are trace fossils, the fragmented conditions
of “Stelechites” do not allow to identify their corresponding
ichnogenus. “Stelechites” could be an unbranched burrow (e.g.,
Planolites or Palaeophycus) or part of a branched structure (e.g.,
Thalassinoides). By comparing “Stelechites” to the trunk of a
tree, Aldrovandi might be implicitly referring to an originally
branched structure.
 ALDROVANDI: TRACES IN THE RENAISSANCE 253

Aldrovandi uses this term to signify a specific animal behavior

FIG. 6. Gastrochaenolites pictured in the Musaeum Metallicum. Aldrovandi
associates borings to bivalve activity.
De Lapide Pholadis
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common to pholads and many other animals. He explains that
this behavior is characteristic of “aves, reptilia, ursae, et aliae
animantes,” or “birds, reptiles, bears and other animals.” The
meaning of “latebra,” hiding place, includes burrows sensu
stricto but also natural shelters (i.e., not actively dug out by
animals, e.g., the cave where a bear might spend the winter).
Aldrovandi specifies the nature of the shelters, distinguishing
“burrows dug underground, borings made in rocks, and other
kinds of shelters” ([. . .] “Sub terra, vel in saxis excavatis, vel
al.s latibulis [. . .] occultantur”).
Aldrovandi’s approach is not merely descriptive: he asks
himself why certain animals hide inside “latebrae.” His answer
is that animals use “latebrae” chiefly to avoid hostile environ-
ments, for example, cold weather (“frigoris”).
MORPHOLOGIES DOUBTFULLY ATTRIBUTABLE TO
TRACE FOSSILS
In addition to the examples mentioned above, the Musaeum

Chapter L of the Musaeum Metallicum, “De Lapide Pho-
ladis,” is entirely dedicated to the study of Recent bioerosion
(Fig. 6). As suggested by the title, Aldrovandi dedicates this
chapter to the study of rocks with pholads, which he defines
“little animals of the bivalve genus” (“animacula de genus
bivalvium”). He begins by explaining the etymology of Pholad,
a name of Greek derivation associated with the idea of “hiding”
or “living inside a shelter” (nam f. Graecis latere, vel in
latebris degere significat” [. . .]). “Latere” means to hide oneself.
Metallicum presents other images that might be associated
with trace fossils (Fig. 7). Contrary to the examples previously
mentioned, there are conflicting indications as to whether the
illustrations discussed in this chapter can be said to represent
trace fossils. Accordingly, we shall discuss these structures with
caution, pending further studies.
The Musaeum Metallicum frequently shows rocks bearing
figurative elements (e.g., animals, monsters). The color of these
elements is different from the color of the enclosing rock, which

FIG. 7. Morphologies doubtfully attributable to trace fossils. A. The specimen presents branched shapes whose color is different from that of the enclosing rock.
Bauhin (1600) used the same style to illustrate a specimen of Phymatoderma (Seilacher, 2007). B. Detail of the specimen shown in A. The figure shows the profile
of an open-mouthed animal morphologically resembling a branched structure. Another branched structure appears in front of the open mouth, probably indicating
the forked tongue of a snake. C. Detail of the specimen shown in A. This bird’s profile might be the interpretation of a branched trace fossil. D, E, F. More rocks
with figurative elements. G. This specimen presents a number of elongated elements whose color is different from that of the matrix. They could be bioclasts (e.g.,
bivalve shells), but the branches might indicate trace fossils (e.g., Chondrites). H. Detail of the previous specimen. The branches are clearly visible. I. Branched
structure; bioclasts (bivalve shells?) and concentric elements (coated grains?—Foraminifera?) can be seen. The structure might be Thalassinoides. J. The caption
says “petrified sea worm.” Both the morphology and the description might indicate a trace fossil, but the specimen is probably a sabellariid tube.
 254 A. BAUCON
is usually darker. Bauhin (1600) uses an almost identical style
to represent a bioturbational trace fossil, Phymatoderma. In
Bauhin’s illustration, the burrows are represented by intricate
angelic figures that contrast with the color of the matrix (see
Seilacher, 2007). It is possible that even the rocks depicted
by Aldrovandi represent branched trace fossils whose fill is
different from the matrix (e.g., Chondrites). As in Bauhin
(1600), the appendages of the creatures depicted may represent
the branching of biogenic structures. However, the illustrator
also may have had other structures in mind, such as bioclasts
or post-diagenetical alterations. There are other specimens in
the Musaeum Metallicum that exhibit branching, for example,
the one shown in Figures 7F and 7G. This specimen might be
Chondrites, but bioclasts cannot be excluded.
The Musaeum Metallicum includes a branched structure
with bioclasts (bivalve shells?) and concentric elements (coated
grains?—Foraminifera?). Based on its morphology, it could
be Thalassinoides, but given the lack of certain evidence, this
attribution cannot be confirmed.
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DISCUSSION AND CONCLUSION
The studies of Ulisse Aldrovandi discussed in this paper help
bring to light a little-known but fundamental chapter in the
history of ichnology. In particular, our analysis gives rise to the
following points for discussion.
The Roots of Ichnology: Aldrovandi, Bauhin and Leonardo
Aldrovandi’s studies in ichnology are historically important
because they come at a critical time in the history of scientific
thought. When Aldrovandi was working on his studies, in the
years following Leonardo Da Vinci and before Galileo, the mod-
ern scientific method was in its early stages of development (Vai
and Cavazza, 2006). Consequently, the Musaeum Metallicum
stands out as an important step in the history of ichnology,
containing some of the first examples of a scientific approach to
the study of traces.
Seilacher (2007) mentions Bauhin (1598, 1600) as one of the
first artists to illustrate and comment on trace fossils. Unfortu-
nately, the Musaeum Metallicum was published posthumously,
making it difficult to establish a link between Aldrovandi and
Bauhin.
Trace fossils are also mentioned in Leonardo’s notebooks. In
fact Da Vinci interpreted correctly bioerosional and bioturba-
tional trace fossils, but his ichnological observations have been
neglected until recently (Baucon, 2008). The reason may have
been the language of the notebooks—vernacular Italian—while
during Leonardo’s times Latin was the language of erudition. In
addition, Leonardo wrote in mirror-image, making his writings
extremely difficult to decipher.
Roots of Ichnology during the Flourishing of Geology
and Paleontology
During the Renaissance, there was a growing interest in earth
sciences. Many naturalists dedicated ample work to the study of
fossils (e.g., Leonardo, Gesner, Cesalpino, Agricola, Fracastoro,
Cardano, Falloppio, Encelius; see Morello, 2003), and even
the word geology dates back to the Renaissance, coined by
Aldrovandi himself (Vai, 2003a).
In this scenario, Aldrovandi’s studies in ichnology occupy a
place of distinction and corroborate an important historical fact:
the scientific roots of ichnology were developed during the same
period that saw the development of geology and paleontology.
Age of Fucoids, Age of Reaction, Modern Age:
Age of Naturalists?
Many naturalists studied “fossilia” during the Renaissance,
but their relationship with ichnology remains poorly studied.
For this reason Aldrovandi, Leonardo and Bauhin could not
represent unique cases, even if, until now, they appear as
isolated. Hence, an “Age of Naturalists” could be erected,
complementing the three traditional stages of the history of
ichnology (established by Osgood, 1970). Possibly the naturalist
Conrad Gesner fits in this scenario; possibly some specimens
described in his “De rerum fossilium” (Gesner, 1565) represent
trace fossils. In fact, Gesner (1565) reports “Stelechites”
(“tree-stones,” p. 351; probably fragmentary burrows), “silex
pertusus foraminibus” (“Flint” with hollows, p. 31; doubtfully
bioerosional structures), “de lapidibus qui serpentes referut”
(“snake-like stones,” pp. 167–169; some snake-like stones could
be trace fossils, even though the most of them are representing
body fossils).
In conclusion, it emerged the important part of Aldrovandi
and his contemporaries in the study of trace fossils. Never-
theless, considerable caution must be exercised in using the
expression “Age of Naturalists” (here proposed) as the role of
naturalists in the history of ichnology deserves further attention.
Aesthetic Aspects of Trace Fossils
The illustrations in the Musaeum Metallicum are among the
oldest representations of trace fossils. This calls attention to two
interdependent aspects: the aesthetic value of traces as natural
objects, and the value of their representations.
The morphologies of trace fossils make them objects of
attraction. After all, that is what first sparked Aldrovandi’s
scientific interest: witness his appreciation for Cosmorhaphe
and Gastrochaenolites, which he called “pulcherrimas,” or
“beautiful.” The aesthetic appreciation of trace fossils is still
strong in our own time, as demonstrated by the ichnological-
artistic exhibition “Fossil Art” (Seilacher, 1997).
Aldrovandi gives much attention to illustration, as he
himself says in his Discorso Naturale (Aldrovandi, 1565).
This can be noticed in the illustration of the ichnogen-
era Cosmorhaphe and Gastrochaenolites, with the accu-
rate reproduction of their morphology, the care for details,
the use of hatching to enhance three-dimensionality. Else-
where the illustrations are more interpretative and “fanci-
ful”: certain branched structures in the Musaeum Metallicum
 ALDROVANDI: TRACES IN THE RENAISSANCE
(dubitatively, Chondrites, but perhaps not trace fossils at all)
become animals or monstrous creatures, all splendidly drawn.
The illustrations in the Musaeum Metallicum underline once
again Aldrovandi’s “double outlook”—his facing forward and
backwards at the same time.
A Renaissance Look at Traces
Aldrovandi’s “double outlook” can also be noted in his
interpretation of structures. His interpretation of traces is
bidirectional: the interpretation of bioerosive structures is
extraordinarily modern, but his ideas on Cosmorhaphe are
decidedly antiquated (Cosmorhaphe is interpreted as a rock
simulating the form of a snake). Conversely, his interpretation
of bioerosional structures is extraordinarily modern: Aldrovandi
associates borings to bivalve activity. In addition, stimulated by
this example, Aldrovandi lists a number of animals that make
burrows or borings and tries to interpret their behavior. His
opinion is that burrows and borings are used by the tracemakers
to protect themselves from adverse environmental conditions.
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Ichnology—A Young Science?
Ichnology is generally considered a young science, dating
back no further than the 19th century. In the light of the consider-
ations made above, one might be prompted to ask: Is ichnology
really a young science? At the current state of studies, this
question remains difficult to answer. In fact some of the major
figures of the Renaissance—Aldrovandi, Leonardo, Bauhin—
dedicated their interest to trace fossils and gave fundamental
contributions to the development of ichnology. However, it
seems that there was no continuity in ichnological studies
between the Renaissance and the 19th century. In conclusion,
ichnology can be considered a young science, but its roots are
much older, firmly anchored in the Renaissance.
ACKNOWLEDGMENTS
I am indebted to Serena Privitera and Michele Cammarosano
(Trieste) for valuable support regarding Latin translation. I am
very grateful to Giorgio Baucon and Chiara Santesso (Udine)
for language revision. I would like to thank the referees,
Gerhard Cadée (Netherlands), Gian Battista Vai and William
Cavazza (Bologna) for constructing criticism and valuable
comments on the manuscript. I thank Alessandro Ceregato
(Bologna) for his precious help about Aldrovandi’s work.
The paper greatly benefited from stimulating comments from
Adolf Seilacher (Tübingen) and Stephania Ern (Milan). I thank
Claudio Veronesi (Milan) for the useful observations. Finally,
I thank Francesca Pozzoli (Milan) for the help concerning
bibliographical research.
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