Palaeogeography, Palaeoclimatology, Palaeoecology 293 (2010) 265–270

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Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

Preface

An island of dwarfs — Reconstructing the Late Cretaceous Haţeg palaeoecosystem

Zoltan Csiki a,⁎, Michael J. Benton b
a
Department of Geology and Geophysics, University of Bucharest, Bd. N. Bălcescu 1, RO-010041 Bucharest, Romania
b
Department of Earth Sciences, University of Bristol, Bristol, BS8 1RJ, UK

a r t i c l e i n f o a b s t r a c t

Article history: The Cretaceous was a special time in the evolution of terrestrial ecosystems, and yet the record from Europe
Received 3 February 2010 in particular is patchy. This special issue brings together results of multidisciplinary investigations on the
Received in revised form 4 May 2010 Late Cretaceous Haţeg area in southwestern Romania, and its continental fossil assemblage, with the aim of
Accepted 25 May 2010
exploring an exceptional palaeoecosystem from the European Late Cretaceous. The Haţeg dinosaurs, which
Available online 1 June 2010
seem unusually small, have become especially well known as some of the few latest Cretaceous dinosaurs
Keywords:
from Europe, comparable with faunas from the south of France and Spain, and preserved at a time when
Cretaceous most of Europe was under the Chalk Seas. Eastern Europe then, at a time of exceptionally high sea level, was
Tetrapods an archipelago of islands, some of them inhabited, but none so extraordinary as Haţeg. If Haţeg truly was an
Dinosaurs island (and this is debated), the apparently small dinosaurs might well be dwarfs, as enunciated over
Island dwarfing 100 years ago by the colourful Baron Franz Nopcsa, discoverer of the faunas. The dwarfing of dinosaurs, and
Island rule other taxa, is explored in this volume. The Haţeg dinosaurs appear to be very latest Cretaceous
Maastrichtian (Maastrichtian) in age, and they provide unique evidence, at a time when there are few dinosaurs known
from Europe, about some of the last faunas before the KT mass extinction. Further, the flora and fauna
(ostracods, fishes, frogs, turtles, lizards, crocodilians, pterosaurs, dinosaurs, and mammals) have never been
reviewed comprehensively, and we provide here the current best evidence of what was there, and how the
taxa fit in a global context.
© 2010 Elsevier B.V. All rights reserved.

1. Islands and dwarfs as well as those of miniaturized mammals (Busk, 1867; Adams, 1874;
Bate, 1903). This pattern of certain (usually large-sized) animals
The unusual nature and composition of island faunas, as well as the becoming smaller, while others became larger after colonizing an
causes underlying these oddities, have been a matter of scrutiny since island habitat, was identified as one of Nature's recurrent phenomena
Darwin's (1859) and Wallace's (1860, 1876) revolutionary contribu- (“rules”) by Foster (1964), and named as such (“island rule”) by Van
tions to the biological sciences. Islands are remarkable in the ways Valen (1973). The processes of phylogenetic size changes (dwarfing/
they control (and, by consequence, allow for tracking and under- giantism) that occur on islands (the “island rule”) became one of the
standing) evolutionary phenomena such as speciation (e.g., Mayr, hallmark aspects of the theory of island biogeography (MacArthur and
1942), survival of dwindling evolutionary lineages within refugia Wilson, 1967; Whittaker, 1999; Cox and Moore, 2005), despite
(e.g., Vartanyan et al., 1993; Guthrie, 2004; Vargas, 2007) or, ongoing controversy on its generality, patterns and underlying causes
conversely, extinction of such lineages restricted to over-shrunken (for a review, see Benton et al., 2010-this issue).
habitats (e.g., O'Regan et al., 2002), development of significant within- Examples of insular dwarfism in the fossil record are not
clade evolutionary divergence (adaptive radiations sensu Simpson, uncommon; however, these have mainly been described from the
1953; e.g., Fritts, 1984; Schluter, 2000; Grant and Grant, 2002; Glor Pliocene–Pleistocene (see, e.g., de Vos et al., 2007 and references
et al., 2004; Baker et al., 2005; Parent et al., 2008) often occurring at a therein), while far fewer cases have been documented from earlier
fast pace (e.g., Kapralov and Filatov, 2006; Millien, 2006; Herrel et al., time periods (e.g., Dalla Vecchia, 2002; Sander et al., 2006; Benton
2008), as well as phyletic size changes (“dwarfing” and “giantism”; et al., 2006). This rarity is at odds with the fact that one of the earliest
e.g., Foster, 1964; Sondaar, 1977; Vartanyan et al., 1993). documented examples of a fossil assemblage with putative insular
Of these special evolutionary features of island biotas, size changes dwarfs was the latest Cretaceous reptilian fauna of the Haţeg Basin
by island colonists has received wide attention, beginning with early (southern Carpathians, western Romania). This fauna was suggested
reports of oversized birds (Owen, 1843) and tortoises (Harlan, 1827), to have inhabited an island at the eastern margin of a palaeo-
archipelago stretching across Tethyan Europe during most of the
Cretaceous, with the particularly small size of its dinosaurs a direct
⁎ Corresponding author. consequence of their restrictive habitat (Nopcsa, 1914). Although the
E-mail address: zoltan.csiki@g.unibuc.ro (Z. Csiki). Haţeg dinosaurian assemblage was subsequently often cited as a

0031-0182/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2010.05.032
266 Z. Csiki, M.J. Benton / Palaeogeography, Palaeoclimatology, Palaeoecology 293 (2010) 265–270

classical example of an island dwarf palaeofauna, this claim has not 3. The Late Cretaceous Haţeg Island — current research and prospectus
been investigated in detail up to now; moreover, the dwarf status of
some of the dinosaurian components was questioned recently by Le The Haţeg area became a focus of geological and palaeontological
Loeuff (2005) and Pereda-Suberbiola and Galton (2009). interest after the discovery of remains of fossil vertebrates in
continental beds (Nopcsa, 1897) referred subsequently to the
uppermost Cretaceous as equivalents of the “Rognacian” or “Gar-
2. The European Late Cretaceous ecosystems: islands umnian” of Western Europe (Nopcsa, 1905, 1915, 1923). Subsequent-
of palaeodiversity ly, through the efforts of Nopcsa and, after him, of several other
researchers, a large amount of data has been amassed on the Upper
The Late Cretaceous continental assemblages of Europe are Cretaceous deposits and their biotas from the Haţeg Basin (see a
somewhat overshadowed by the much better studied ones from synthesis of this research in Grigorescu, 2010-this issue). While early
North America and Asia, as well as by those described recently from work was mainly concerned with establishing the age and geological
South America. This is largely because of the more extensive outcrops context of the deposits, as well as description of the most conspicuous
of continental units in all these areas compared to those in Europe, members of the macrofauna, more recent research activity has been
correlated with the larger number of fossiliferous localities (see, e.g., more multidisciplinary, bringing together specialists working in
the reviews of Weishampel et al., 2004; and Kielan-Jaworowska et al., different fields of geosciences, from palaeozoology and palynology
2004 for dinosaurs and mammals, respectively), as well as the higher to stable isotope geochemistry and palaeomagnetism. The main aim of
diversity of local faunal assemblages relative to those described from this activity (although not always stated) was to achieve as complete
Europe. Starting as early as 1915 (Nopcsa, 1915), the different as possible a reconstruction of this latest Cretaceous island ecosystem
European Late Cretaceous faunas were often interpreted as depau- and to better understand its relationships with other contemporane-
perate compared to those known from other continental landmasses ous ecosystems from Europe and abroad (e.g., Csiki and Grigorescu,
(low alpha diversity, i.e., raw local species richness; see Whittaker, 2007). Documenting research efforts on the Haţeg localities and
1972) both at the level of the major clades present, and of the number faunas over the past 111 years since they were discovered reveals an
of individual taxa represented; moreover, the overall composition of all-time peak in the number of scientific papers in the past 15 years
the European Late Cretaceous palaeobiocoenosis was considered to be (see Csiki, 2005; Csiki and Grigorescu, 2007; Grigorescu, 2010-this
comparatively less diverse (gamma diversity, i.e., total diversity issue).
across a larger geographical area; Whittaker, 1972). This low-level Several results of this increased recent research activity on Haţeg
diversity was considered to be complemented by several other have already been published, most notably on the vertebrate
unusual features, such as the survival of several basal tetrapod palaeontology, sedimentology, and distribution of the Maastrichtian
lineages up to the terminal Cretaceous, giving these faunas a vertebrate-bearing beds, but other topics are less well represented.
“primitive” aspect (e.g.; Nopcsa, 1915, 1923; Weishampel et al., This volume brings together 14 contributions that address previously
1991; Gaffney and Meylan, 1992), the relative uniformity of the less well represented research topics, from igneous petrography and
faunas across the different parts of Europe (see, e.g., Nopcsa, 1915; Le stable isotope analyses to ichnology (dinosaur eggs) and taphonomy;
Loeuff, 1991) supporting relatively low beta diversity (i.e., degree of the institutional affiliations and research interests of the 27
differentiation of local assemblages along habitat gradients or simply researchers co-authoring these contributions reflect well the breadth
between different part of a larger area; Whittaker, 1972), and the of international cooperation involved in this research project.
ambiguous palaeobiogeographic affinities of this Late Cretaceous The introductory chapter of the issue (Grigorescu, 2010-this issue)
ecosystem, in which the mixture of taxa of either southern offers an overview of the previous research done on the Upper
(Gondwanan) or northern (Laurasian) origin blurred its biogeograph- Cretaceous deposits of the Haţeg area and their palaeontological
ic individuality (Le Loeuff, 1991; Le Loeuff and Buffetaut, 1995; Le content, focusing mainly on the early years and the seminal and
Loeuff, 1997). These special palaeobiological features of the European prodigious activity of Baron Franz Nopcsa. The data synthesized in this
Late Cretaceous assemblages were paralleled by their particular overview are supplemented by the historical information offered by
palaeogeographic–tectonic setting, inhabiting an archipelago with a the other contributions to this issue.
geography undergoing large-scale temporal and spatial fluctuations Although not directly concerned with the Late Cretaceous Haţeg
(e.g., Tyson and Funnell, 1987; Smith et al., 1994; Dercourt et al., 2000; Island and its continental ecosystem, the next two contributions to
Csontos and Vörös, 2004), quite unlike the larger, spatially continuous this volume, focusing on the marine deposits directly beneath the
continental landmasses of North America and Asia (e.g., Smith et al., vertebrate-bearing beds, are nevertheless integral parts of the
1994). ongoing multidisciplinary research in the area. They are critical in
Recent advances in the study of Late Cretaceous European establishing the age of the Haţeg continental deposits, which do not
ecosystems appear to support the emergence of a significantly contain such age-diagnostic fossils as the better-dated underlying
different picture for this part of the world, as foreshadowed by Rage marine deposits. This information is here updated and supplemented
(2002). Discovery of rich and diverse fossil assemblages ranging from by Melinte-Dobrinescu's (2010-this issue) contribution on calcareous
the Cenomanian of France (e.g., Vullo and Néraudeau, 2008) to the nannoplankton biostratigraphy, which suggests that, despite earlier
Santonian of Hungary (Makádi et al., 2006; Ősi and Rabi, 2006) and to reports (e.g., Stilla, 1985; Pop, 1990), marine deposition in the area
the Campanian–Maastrichtian of the Ibero–Armorican landmass (e.g., continued only as late as the latest Campanian, thus allowing
Barroso-Barcenilla et al., 2009; Company Rodriguez et al., 2009) have continental deposition to start in the earliest Maastrichtian. These
revealed much greater alpha and gamma diversity within Late age constraints are confirmed by stable isotope studies reported by
Cretaceous European ecosystems than had been thought before, Melinte-Dobrinescu and Bojar (2010-this issue) through the identi-
while also pointing to a greater beta diversity than previously fication of several time-significant isotope excursions in these marine
acknowledged. A recent review of the palaeobiogeographical affinities deposits. Moreover, climatic trends identified in the late Late
of these faunas (Pereda-Suberbiola, 2009) has presented a rather Cretaceous (as recorded by the marine deposits) are hypothesized
complex picture of their inter- and intra-province relationships. In to continue into the Maastrichtian as well, thus contributing to our
view of these recent additions to our knowledge of Late Cretaceous understanding of the dominant palaeoclimate in the Haţeg area after
European ecosystems, an update on the Haţeg palaeobiota and its its emergence.
environment appears necessary in order to better understand its The age constraints offered by the study of the underlying marine
position and relationships within a European palaeobioprovince. deposits are supported by palaeomagnetic studies on the continental
 Z. Csiki, M.J. Benton / Palaeogeography, Palaeoclimatology, Palaeoecology 293 (2010) 265–270 267

deposits themselves, reported here by Panaiotu and Panaiotu (2010- agreement with those based on palaeomagnetism, paleosol sedimen-
this issue). Based on their magnetostratigraphic data, the authors tology and palynology (see above).
suggest that continental deposition started at the beginning of the The Haţeg Basin is also well known as the source of the only
Maastrichtian at the latest, but possibly even as early as the end of the documented field association between dinosaur eggs and hatchling
Campanian. Moreover, the recorded palaeomagnetic data also remains from the Upper Cretaceous of Europe (e.g., Grigorescu, 1993).
constrain the palaeogeographic position of the emergent Haţeg Island Somewhat ironically, this co-occurrence, usually held up as a good
and thus offer independent control for the previous palaeoclimatic opportunity to identify the egg-laying animals, has involved contro-
reconstructions based on palynology (e.g., Van Itterbeeck et al., 2005) versy (see, e.g., Grigorescu, 2003) over whether the eggs were laid by
and paleosol sedimentology (e.g., Therrien, 2005). titanosaur sauropods or by hadrosaurids. Grigorescu et al. (2010-this
Volcaniclastic deposits associated with the vertebrate-bearing issue) review the main stratigraphic, sedimentological, and tapho-
continental deposits of the Haţeg Basin (e.g., Grigorescu, 1992) were nomic features of the different nesting sites from the Maastrichtian of
often overlooked in previous palaeoenvironmental reconstructions, the Haţeg Basin, and their work further supports the originally
despite the major impact the explosive volcanic events might have suggested association between megaloolithid eggs and the hadro-
had upon the environments and biotas (e.g., Goldberg and Garcia, saurid Telmatosaurus.
2000; Guo et al., 2003; Fürsich et al., 2007; Zhao et al., 2007). The Csiki et al. (2010-this issue) present a thorough overview of
study of Bârzoi and Şeclăman (2010-this issue) offers new petro- taphonomic modes of the Haţeg vertebrates, a theme that has been
graphical and geochemical data for the interpretation of the palaeo- touched on by many authors since Nopcsa's (1902, 1914) original
geotectonic context of the volcaniclastics. According to these authors, suggestion that most of the vertebrate accumulations had been
volcanic activity responsible for generating these deposits must have produced by crocodilians gathering together carcasses of dead and
taken place in an island arc setting, relatively far from the geotectonic dying animals. In fact, as recent work has shown, the Haţeg deposits
settings within which the dinosaur-bearing beds were deposited; represent a wide range of taphonomic modes within the prevailing
accordingly, it can be hypothesized that these eruptions might have fluvial-dominated upland setting that experienced seasonal semi-arid
had a reduced impact upon the development of the terrestrial biotas. conditions. The major sedimentary settings are channel lags and fills,
Closing in on the Maastrichtian continental deposits, the contri- crevasse splays and channels, well-drained floodplains, mature non-
bution of Bojar et al. (2010a — this issue) analyses their heavy mineral calcareous paleosols, hydromorphic paleosols, abandoned channels
spectra in order to reconstruct possible source areas. The data derived and ponds, and marshy ponds. Most of the Haţeg vertebrates occur in
from their study allow a better understanding of the tectonic context fine-grained floodplain deposits, rather than in channels, the opposite
within which the Haţeg area evolved; according to the identification of the situation in many other dinosaur-bearing formations. Skeletons
of distinct northern and southern source areas that operated during show evidence of disturbance, breakage, and weathering, and bone-
the Maastrichtian, it can be reconstructed reliably as an intramontane beds, whether of larger or smaller elements, show remarkable mixing
basin fed by the uplifted marginal areas. Moreover, the tectonic of materials from many sources, some relatively fresh and others
evolution of the surrounding mountainous areas appears to have had much abraded. Further, although the smaller vertebrates show
significant impact upon the palaeoenvironmental evolution of the evidence of habitat preferences (amphibians and lizards in ponds;
basin itself, promoting a shift towards wetter, more mesic environ- crocodiles, turtles, and mammals in less marshy areas), the dinosaurs
ments during the later part of the Maastrichtian. are found randomly across all sedimentary settings.
The palaeoclimatic conditions of Haţeg Island were also investi- Equating the Haţeg Basin with the Haţeg Island might seem
gated using stable isotopes derived from sediments and organic legitimate, and certainly has been done (see, e.g., Weishampel et al.,
remains (Bojar et al., 2010b — this issue). According to this 1991). However, this could be misleading, since island area is one of
investigation, reconstructed palaeotemperature values suggest a the main controlling factors of island ecosystems (e.g., MacArthur and
rather low-latitude palaeogeographic setting, in accordance with the Wilson, 1967; Whittaker, 1999; see Lomolino, 2001) and thus
conclusions of Panaiotu and Panaiotu (2010-this issue) discussed inferences concerning expected alpha diversity or distance from
above. The most important results of this stable isotope investigation surrounding landmasses could be seriously flawed. Consequently,
appear to concern, however, the ethology and physiology of several establishing the extent of the emergent Haţeg area is key for any
archosaurian taxa (Zalmoxes, Telmatosaurus, Allodaposuchus), part of a attempt to understand this island palaeoecosystem. Haţeg Island must
wider debate about dinosaurian metabolism and the use of isotopic have been significantly larger than the Haţeg Basin itself, as already
measurements to investigate environmental water composition. suggested by Nopcsa (1905). The contribution of Codrea et al. (2010-
These results open up the possibility of wider comparisons with this issue) attempts to estimate the area of emergent land by mapping
Late Cretaceous vertebrates from other areas (e.g., Amiot et al., 2006; the wider outcrop of continental deposits that preserve vertebrate
Fricke and Pearson, 2008; Fricke et al., 2008). remains reminiscent of those from the Haţeg Basin itself. According to
Plant remains are rare in the continental deposits of the Haţeg their data, Haţeg-type assemblages can be recognized all over the
Basin, being represented mainly by spores and pollen (e.g., Van southwestern, western and northwestern margin of the Transylva-
Itterbeeck et al., 2005; Csiki et al., 2008), while rare leaf impressions nian Basin, thus significantly increasing the mappable area of this
(Mărgărit and Mărgărit, 1967) make up the only known macrofloral former island. Preliminary work suggests some taxonomic and
remains. Similarly, plant remains, except for palynomorphs, are palaeoecological differences between these local assemblages (e.g.,
exceedingly rare in other Upper Cretaceous vertebrate-bearing between the Haţeg local fauna and those from outside the Haţeg
continental units of Western Europe (see, e.g., Nichols and Johnson, Basin), pointing to the need for further study to determine whether
2008), unlike the contemporaneous deposits of North America (e.g., these outlying assemblages really pertain to the same island as
Johnson, 2002). Consequently, the mesofloristic remains (seeds and represented by the Haţeg Basin itself.
fructifications) reported by May Lindfors et al. (2010-this issue) The larger palaeobiogeographic significance and relationships of
represent an important addition to our knowledge of the Maastrich- the Haţeg ecosystem are addressed by two contributions. Martin and
tian floras of Haţeg Island. These are even more significant in that they Delfino (2010-this issue) survey the stratigraphic and geographic
are associated with palynomorphs and microvertebrate remains at distribution of Cretaceous European eusuchians (represented on
the same site, thus allowing a detailed reconstruction of the local Haţeg Island by the apparently endemic Allodaposuchus precedens;
palaeoenvironment. The morphotypes and size compositions of the e.g., Delfino et al., 2008 and references therein) in order to identify
preserved Haţeg mesoflora suggests a semi-arid, seasonally variable patterns of their evolution. Their analysis allows the reconstruction of
climate, so providing a further independent palaeoclimatic signal in a Late Cretaceous European archipelago supporting widely divergent
268 Z. Csiki, M.J. Benton / Palaeogeography, Palaeoclimatology, Palaeoecology 293 (2010) 265–270

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