Historical Biology

An International Journal of Paleobiology

ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20

Las Hoyas: a Cretaceous wetland: a

multidisciplinary synthesis after 25 years of
research on an exceptional fossil Lagerstätte from
Spain

James O. Farlow

To cite this article: James O. Farlow (2017): Las Hoyas: a Cretaceous wetland: a multidisciplinary
synthesis after 25 years of research on an exceptional fossil Lagerstätte from Spain, Historical
Biology, DOI: 10.1080/08912963.2017.1367081

To link to this article: http://dx.doi.org/10.1080/08912963.2017.1367081

Published online: 22 Aug 2017.

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 Historical Biology, 2017
https://doi.org/10.1080/08912963.2017.1367081

BOOK REVIEW

Las Hoyas: a Cretaceous wetland: a multidisciplinary The ‘wet facies’ assemblages accumulated during intervals of
synthesis after 25 years of research on an exceptional deeper water. Here fewer specimens occur, but species richness is
greater than in ‘dry facies’ assemblages. Taxa include a mixture of
fossil Lagerstätte from Spain, edited by F. J. Poyato-
autochthonous aquatic and parauthochthonous terrestrial organ-
Ariza and Á. D. Buscalioni, Munich, Verlag Dr. Friedrich isms. Specimens are generally not as well-preserved as those of
Pfeil, 2016, 262 pp., €75.00 (hardcover), ISBN the dry facies. Fossils include abundant plant fragments and a
10 3899371534; ISBN 13 978-3-89937-153-6 fauna dominated by insects, small teleosts, and pycnodont fishes.
The fishes are often larger than those preserved in the dry facies.
Spending much of my childhood on a lake in Indiana led me to Sediments of the dry facies accumulated on very shallow
a lifelong interest in freshwater ecosystems. One of my recent or even emergent mud flats, covered by substantial microbial
projects focused on a late Neogene sinkhole pond/wetland fossil mats. Many more fossils occur than in the wet facies, but with
assemblage (Farlow et al. 2010). I have also worked extensively relatively low species richness. Individual fossils are often beau-
on Mesozoic vertebrates. Consequently, I picked this book up tifully preserved. Animal fossils are generally smaller than those
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with considerable interest. of the wet assemblage, often occurring in monospecific mass
Poyato-Ariza and Buscalioni have assembled an interna- death associations of individuals of about the same size. Some
tional team of specialists to describe and interpret the fossils and species are represented by ontogenetic growth series. The fauna
sediments of Las Hoyas (‘the sinkholes’), an Early Cretaceous is dominated by small fishes and crustaceans, insects other than
Konservat-Lagerstätte in southeastern Spain. The preface and water bugs being less common.
two initial chapters summarise the history of study of the sites, As a Lagerstätte, Las Hoyas is characterised by many spec-
documenting the recognition of Las Hoyas as an unusually well imens showing exquisite preservation of skeletal material and
preserved paleobiota. Groups of organisms constituting the fauna even soft parts. Plants are generally represented as impressions
and flora are described in 24 chapters and 4 appendices/addenda; and carbonised compressions, the aquatic taxa being more often
5 chapters deal with taphonomy (including experimental studies of preserved intact than terrestrial plants. Charophyte gyrogonites
carcass decomposition, disarticulation, and diagenesis); and two are found attached to the algal thallus. Insects are generally rep-
chapters synthesise the paleoecology of Las Hoyas. In some chap- resented by external molds of the exoskeleton, but the skeleton
ters the English writing is a bit awkward, but never so much as to may be replaced by calcite, silica, or pyrolusite. Bivalves are most
limit comprehension. The book is well illustrated throughout with often preserved as steinkerns. The notochordal canal of teleost
clear line-drawings, crisp and often lovely photographs (many in fishes may be filled with calcite, and it is not unusual for fish and
color), reconstructions of the Cretaceous landscape, and restora- tetrapod skeletons to be preserved in articulation.
tions of individual organisms. I read the book with great pleasure. Soft parts of animal carcasses are preserved as siderite, apatite,
Throughout the systematic chapters, the authors give due or stable organic molecules. Eyeballs and muscles are the most
attention to the phylogeny and biogeography of the groups commonly preserved soft tissues. Several tetrapod specimens show
under discussion. Almost a third of the animal taxa identified are skin impressions, and feathers are preserved by carbonised residues
endemic to Las Hoyas. In this review, however, I will emphasise or limonite. Traces of the digestive tracts of both invertebrates and
the paleoecology of Las Hoyas. fishes are known. Color patterns, however, are not often preserved.
The fossil assemblage is preserved in sediments that accu- Experimental observations of decay and disarticulation of
mulated in a half-graben on a subtropical, low-relief karstic small fishes imply that the Las Hoyas fishes are unlikely to have
landscape. Finely laminated, freshwater, biogenic limestones undergone much postmortem transport; they probably died
and marls comprise the La Huérgina Limestone Formation, where they were preserved. Similar observations of modern frog
of Hauterivian-Barremian age. Initially regarded as infilling a carcasses show that they are highly susceptible to disarticulation
deep meromictic lake, this interpretation was cast in doubt by in fresh water. Microbial mats are thought to have counteracted
the discovery of a crocodylomorph trackway (more about this this tendency to keep Las Hoyas frog skeletons from falling apart.
below) and subsequent finds of other crocodile and dinosaur Experiments indicate that cyanobacterial mats (for which
tracks. Together with desiccation cracks and other sedimen- there is sedimentological evidence at Las Hoyas) form biofilms
tary features, this led to a revised interpretation. The habitats around small carcasses, and these act to slow decomposition
are now thought to have been a complex of lakes and swamps/ and hold skeletons together. Such mats create calcium-rich films
marshes, with pronounced seasonal fluctuations in water depth; around dead fishes, serving as molds that can be filled by auth-
the Florida Everglades are a possible modern analog. Overall, the igenic minerals, thereby potentially contributing to excellent
paleobiota is dominated by aquatic organisms. preservation.
 2   
BOOK REVIEW

Plants are represented in the La Huérgina paleobiota at Las Most of these fishes are small teleosts (generally 3–4 cm long),
Hoyas by palynomorphs as well as megafossils. Both wetland although growth series of some species reach 10–15 cm in length.
palynomorphs and assemblages characteristic of drier habitats Bigger fishes are also present, including coelacanths (up to 30
are present. Fern spores predominate in terms both of species or even 50  cm long), pycnodonts (as much as 13.5  cm long),
richness and numbers of specimens, but freshwater algae, bryo- semionotiforms (up to 16 cm long), and amiiforms (up to 65 cm
phytes, gymnosperms, and angiosperms are also represented in long). The presence of these larger fishes presumably indicates
the microflora. Plant megafossils are abundant, comprising all that the Las Hoyas habitats at least occasionally included larger
of the large fossils in some beds. and deeper bodies of water, or they were connected to such larger
Three kinds of habitat are indicated by the plant assemblages. lentic environments. Even so, the biggest Las Hoyas fishes are
One consists of somewhat deeper-water lake, pond, or pool sit- smaller than their relatives known in other faunas; elsewhere,
uations, where charophytes are especially common. There is semionotiforms reach up to 2 m in length.
disagreement among the authors as to whether this indicates Most of the Las Hoyas fishes were nektonic; there are no
oligotrophic (p. 54) or eutrophic (p. 101) nutrient levels in the specialised demersal species in the fauna. The small teleosts
ecosystem. Surrounding these aquatic habitats were marshes and presumably occupied relatively low positions in aquatic food
swamps, where the conifer Frenelopsis formed dense, possibly webs, preying on zooplankton and pleuston. Some apparently
monospecific stands. The higher, drier surrounding landscape nibbled on charophyte gyrogonites. Bigger fishes (amiiforms,
hosted a savanna type of vegetation, dotted by Weichselia tree coelacanths) probably ate insects, molluscs, and of course smaller
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ferns. Here, the land plants show xeric adaptations (thick cuti- fishes; some pycnodonts may have cropped pondweed.
cles, scale leaves, low humidity configurations of stomata), and The relative abundance of fishes in the Las Hoyas fauna
the frequent occurrence of charcoal in the sediments indicates (assuming it reflects the relative abundance of fishes living in the
that fire likely played a significant role in ecosystem processes. paleocommunity) may account for something that surprised me
Molluscs are represented in the aquatic facies by unionid about this assemblage: the relative scarcity of amphibians com-
bivalves and freshwater snails. Bivalves are usually preserved with pared with that of fishes. In modern and at least some Neogene
their valves open (in ‘butterfly position’), indicating that they wetland assemblages, amphibians can be extremely abundant,
lived close to the sediment surface. Two spiders of one species even numerically dominating vertebrate faunas (cf. Farlow
belong to a family whose modern representatives construct webs et al. 2010). For example, in Ellenton Bay, an isolated, fish-free,
along the margins of bodies of water. Sixty-four millipedes are long-hydroperiod wetland in South Carolina, up to 10 ha in area,
also mostly members of a single species. transforming young-of-the-year amphibians leaving the pond
Insects are the most diverse animals in the Las Hoyas fauna. have been shown to include an astonishing 1036 salamanders/ha,
With crustaceans and fishes they numerically dominate the and 38,612 anurans/ha (Gibbons et al. 2006)! Granted that these
assemblage. Some 13 insect orders and 90 species and/or mor- are only temporary residents of the pond, there are numerous
photypes are represented, larval or pupal stages being known adult amphibians (especially frogs) in and around this body of
as well as adults of some species. Aquatic (especially faunivo- water that could potentially contribute carcasses year-round to
rous) forms comprise 61% of the insect taxa, and dryland forms bottom sediments for a future fossil fauna.
36%. Groups include water bugs, dragonflies, caddisflies, beetles, Fishes are important predators of amphibian eggs, larvae, and
grasshoppers, cockroaches, and wasps. One of the most common even adults (Wells 2007), so amphibians often breed in fish-free
forms is the heteropteran Iberonepa, which appears similar, at habitats (Colburn 2004). Perhaps the abundance of Las Hoyas
least superficially, to bugs I have seen living in today’s shallow fishes had a damping effect on amphibian abundance, in turn
ponds. suggesting that standing water was present long enough to allow
Like insects, crustaceans are especially common, represented fish populations to grow sufficiently large to restrict amphibian
by decapods, copepods, ostracodes, peracarids, isopods, possi- numbers. Be that as it may, amphibians were nonetheless pres-
bly mysids, and others. In contrast with other constituents of ent in the fauna (even if they didn’t numerically predominate),
the fauna, ostracodes in the Las Hoyas assemblage are poorly and in the abundance needed to feed those predaceous insects
preserved, usually with their carapaces closed. This is taken to (water bugs, beetles) likely to have dined (at least in part) on
indicate deposition in quiet-water situations. The taxa present are amphibian larvae.
typical of fresh to slightly saline water; some probably required Among the Las Hoyas amphibians are albanerpetontids (6–8
long hydroperiods, but others made desiccation-resistant eggs. specimens, up to about 6 cm in snout-vent length [SVL]), ani-
Twenty-five specimens of a possible cavernicolous crustacean mals with a salamander-like body form, but with scaly skins, toe
(spelaeogriphacean) are known; to my eye, the reconstruction claws, and non-pedicellate teeth. They probably preyed on ter-
of this animal looks rather like a stretched-out pillbug. Decapod restrial arthropods. Salamanders are more common, represented
crustaceans are among the most abundant and well-preserved by 50 specimens in two taxa, making them the most common
animal fossils, but unlike the insects, their species richness is tetrapods in the assemblage. One of these (Valdotriton) is a ter-
relatively low. One of the crayfishes is represented by individuals restrial form with a snout-pelvis (SPL) length (close to SVL) of
up to 7 cm long. 3–4 cm. The other salamander was slightly bigger (SPL 3–5 cm),
Fishes are the third abundant animal fossils in the Las Hoyas neotenic and aquatic as an adult. The Las Hoyas fauna includes
assemblage. Two small (probably juvenile) articulated hybodont at least three kinds of frog, represented by tadpoles and adults,
shark specimens are known, but these were probably not regular the latter attaining SVLs of 2–4 cm.
members of the fauna. Osteichthyans, in contrast, are both abun- Another surprising feature of the Las Hoyas fauna (at least to
dant (about 5000 specimens) and diverse (at least 16 genera). me) is the scarcity of turtles. Only seven specimens of a single
 HISTORICAL BIOLOGY   3

small, typically aquatic turtle (appropriately named Hoyasemys) crest and a throat pouch on Pelecanimimus. Toe pads are present
are known. on the foot of Concavenator, and scales are preserved along the
Squamates are represented by 17 specimens, 11 of which ankle and tail. Bumps on its ulna are reminiscent of avian quill
are assigned to Meyasaurus, a rather standard-model lizard. A knobs. The oddest feature of Concavenator is elongation of neural
more unusual form is Scandensia, a lizard with broad, stiff ribs, spines on vertebrae above the hip, suggesting development of a
elongate penultimate phalanges on both the hand and foot, and triangular dorsal sail. Given the undoubtedly predatory habits of
forelimbs that are roughly the same length as the hind limbs, this dinosaur, I cannot look at this putative sail without having
all suggesting that it was a climber. Hoyalacerta is a lizard with the phrase ‘land shark’ pop into my head. Tridactyl dinosaur
relatively short limbs. The Las Hoyas lizards were small tetrapods footprints provide unambiguous evidence that at times the site
(generally 3–4 cm SVL), probably mostly insect-eaters, although was exposed above water level, or only just below it.
a specimen of Meyasaurus from one locality had a fish in its gut The dinosaurs for which Las Hoyas is most famous are the
region. A bone assemblage consisting of a half-dozen, disarticu- enantiornithine birds Iberomesornis, Concornis, and Eoalvulavis.
lated small lizards (mostly Meyasaurus) is interpreted as the gut They are represented by several partial skeletons, four of them
contents of a larger predator. occurring together in what is interpreted as a regurgitated pellet,
Among the potential lizard-eaters at Las Hoyas are croco- perhaps upchucked by a non-avian theropod or pterosaur. Gut
dylomorphs, represented by ten mostly articulated skeletons contents that are preserved indicate that Eoalvulavis dined on
and additional fragments. These reptiles include a gobiosuchid crustaceans. Enantiornithines seem to have grown more slowly
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(a long-legged, possibly terrestrial form), the atoposaurid than modern birds of comparable size. None of these birds is
Montsecosuchus, the neosuchian Unasuchus, and an advanced very big. They all appear to have been capable of powered flight,
neosuchian. These are all small crocs, with body lengths less although their performance would have been less impressive
than 1 m; they include both juveniles and ‘subadult’ individuals. than those of modern birds.
However, the trackway mentioned above indicates the pres- More than a dozen feather specimens are known from Las
ence of a much bigger crocodylomorph than any represented Hoyas. Some were attached to specimens of Eoalvulavis and
in the skeletal fauna. This creature had a pes length of 20 cm, Concornis, but most (molts?) were found isolated, preserved
indicating an animal comparable in size to an adult American as carbonaceous residues. Some record traces of color pattern.
alligator. The trackmaker is inferred in this book to have been Although birds are obvious candidates for the erstwhile own-
walking in extremely shallow water (possibly as little as a few ers of isolated feathers, some may have belonged to non-avian
centimeters in depth). It was the discovery of this trackway that dinosaurs.
initially prompted the change in interpretation of the Las Hoyas An important recent addition to the Las Hoyas paleofauna
paleoenvironment from a deep lake to a shallow wetland (p. 16). is the nearly complete articulated skeleton of a gobiconodontid
Compared with other crocodylomorph trackways, the Las mammal, Spinolestes. Preserved soft tissues include hair, some of
Hoyas trackmaker’s stride length is unusually long relative to it spiny, and an ear pinna. Spinolestes is estimated to have been
its pes print length (Farlow et al. 2017). The prints might have 24 cm long, with a body mass of 52–72 g. Presumably it was a
been made by an unusually long-legged form – perhaps a bigger small terrestrial faunivore.
version of the gobiosuchid? – but Farlow et al. (2017) hypoth- Trace fossils are reasonably common at Las Hoyas. In addition
esised that the trackmaker might have been moving underwa- to the vertebrate trackways, several kinds of invertebrate and
ter, employing a combination of punting and bottom-walking fish traces occur. Most common is Treptichnus pollardi, a three-
(Grigg and Kirshner 2015). This style of locomotion can occur in dimensional feeding trace possibly made by larval flies. Simpler
water depths of several meters, implying that a crocodile track- horizontal traces (Helminthoides, Palaeophycus, Planolites) var-
way alone may not document extremely shallow water. However, iously interpreted as dwelling structures or feeding traces of
as already noted, there is ample additional support for at least worms or larval insects also occur. Tiny versions of Cruziana
occasional periods of very shallow water and aerial exposure at are presumed to have been made by crustaceans, perhaps bran-
Las Hoyas. chiopods. Lockeia is thought to have been made by ostracodes
Pterosaurs occur only as scrappy material at Las Hoyas. Five or conchostracans. Given the abundance of fishes in the skeletal
isolated teeth and a partial skull represent three taxa in as many fauna, it isn’t surprising that a number of traces thought to have
families: Istiodactylidae, Ornithocheiridae, and Tapejaridae. The been created by fishes also occur. Swish marks (Undichna) were
last is a toothless species of Europejara. The istiodactylid was of possibly made by pycnodonts.
modest size (teeth only about 6 mm long). The ornithocheirid Overall, the trace fossil assemblage (assigned to the Mermia
was much bigger, the largest tooth being 37 m long. All three ichnofacies) is thought to have developed during periods of low
pterosaurs are inferred to have fed on fishes of various sizes, the water (as little as a few centimeters in depth). Allowing dino-
istiodactylid possibly also eating crustaceans and insects. saur footprints to be impressed and preserved, shallow water
Non-avian dinosaurs have scant representation in this might also have forced fishes (and perhaps a crocodylomorph?)
fauna, but the material that does occur is quite interesting. A to interact with the bottom during swimming. Apart from traces
gorgeous, articulated lower leg and hindfoot of the iguanodont confined to bedding planes or that involved only limited vertical
Mantellisaurus hints at the kinds of large herbivores that were penetration, the sediments at Las Hoyas show little indication
cropping the vegetation. Two theropods are known, the basal, of bioturbation.
toothed ornithomimosaur Pelecanimimus and the even toothier Las Hoyas provides an unusually detailed record of a vanished
basal carcharodontosaur Concavenator. Skin impressions occur paleoecosystem, inviting reflection on how it was similar to, and
in both cases. These suggest the presence of a triangular head different from, wetland systems of other times and places. The
 4   
BOOK REVIEW

authors and editors of this beautiful book are to be congratulated Gibbons JW, Winne CT, Scott DE, Willson JD, Glaudas X, Andrews KM,
for giving this fossil assemblage so thorough and interesting a Todd BD, Fedewa LA, Wilkinson L, Tsaliagos RN, et al. 2006. Remarkable
amphibian biomass and abundance in an isolated wetland: implications
treatment. for wetland conservation. Conserv Biol. 20:1457–1465.
Grigg G, Kirshner D. 2015. Biology and evolution of crocodylians. Ithaca
(NY): Cornell University Press (Comstock); 649 p.
References Wells KD. 2007. The ecology and behavior of amphibians. Chicago (IL):
Colburn EA. 2004. Vernal pools: natural history and conservation. University of Chicago Press; 1148 p.
Blacksburg (VA): McDonald & Woodward; 426 p.
Farlow JO, Richards RL, Garniewicz RC, Greenan M, Wepler WR, Shunk AJ, James O. Farlow
Ludvigson GA, Shaffer NR, Wenning AL. 2010. Occurrence and features Department of Biology, Indiana-Purdue University, Fort
of fossiliferous sediments of the Pipe Creek Sinkhole (Late Neogene,
Wayne, IN, USA
Grant County, Indiana). In: Farlow JO, Steinmetz JC, DeChurch DA,
editors. Geology of the Late Neogene Pipe Creek Sinkhole (Grant County, [email protected]
Indiana). Indiana Geological Survey Special Report 69. Bloomington © 2017 James O. Farlow
(IN): Indiana University, Indiana Geological Survey; p. 1–60. https://doi.org/10.1080/08912963.2017.1367081
Farlow JO, Robinson NJ, Kumagai CJ, Paladino FV, Falkingham PL, Elsey
RM, Martin AJ. 2017. Trackways of the American crocodile (Crocodylus
acutus) in northwestern Costa Rica: implications for crocodylian
ichnology. Ichnos. DOI:10.1080/10420940.2017.1850856.
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