Passalia Et Al (2022) Bosque Petrificado Valcheta
Passalia Et Al (2022) Bosque Petrificado Valcheta
Passalia Et Al (2022) Bosque Petrificado Valcheta
Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes
a r t i c l e i n f o a b s t r a c t
Article history: The Valcheta Petrified Forest is located in the northeast of the Río Negro Province, Argentina. This work
Received 3 August 2022 provides a geological characterization of the site and a survey of the fossil logs exposed on the surface
Received in revised form and their systematic study. The fossiliferous levels bearing the silicified logs are here referred to the Allen
4 October 2022
Formation (middle-upper Campanian e lower Maastrichtian, Malargüe Group). These deposits crop out
Accepted in revised form 9 October 2022
in patches in the study area, and have been deposited on a paleorelief carved in Eopaleozoic rocks of the
Available online 14 October 2022
Nahuel Niyeu Formation being mostly covered by thin layers of Quaternary sediments. The Valcheta
Petrified Forest was developed on the margins of a braided fluvial system. Numerous silicified logs have
Keywords:
Upper Cretaceous
been located at the site. They consist of large axes, not found in life position, with monopodial growth,
Patagonia some of which retain the bases of lateral branches and flared-like base. The estimated height and ages of
Somún Cura Massif these trees suggests that it consisted of a mature forest. No other associated vegetative or reproductive
Allen Formation structures have been found. Anatomically, a single xylological type has been distinguished and assigned
Podocarpoxylon mazzonii to Podocarpoxylon mazzonii (Petriella) Müller-Stoll et Schultze-Motel, (Podocarpaceae). This taxon has
been previously identified from other localities of the Upper Cretaceous to the Danian of Patagonia
although never preserved in such complete tree grow architecture. This shows that, along others co-
nifers, cycads and palms, P. mazzonii would have constituted an important element of the arboreal strata
in the North Patagonian forest communities during the CampanianeDanian interval producing mono-
typic patches in some areas.
© 2022 Elsevier Ltd. All rights reserved.
https://doi.org/10.1016/j.cretres.2022.105395
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M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Somún Cur a Massif (Río Negro Province). The first mention of large also taken using a Brunton-type geological compass, based on
fossil woods in the site is present in a report of Willis (1911e1914). sedimentary indicators (primary tractive sedimentary structures)
Tortorelli (1941) described one incompletely preserved wood (Suppl. mat. 2).
collected from the Valcheta area, without stating a probable age,
and compared it with Ginkgo biloba L. wood. Later references about 3. Geological Setting
the existence of silicified logs in the Valcheta town area were made
by Nún ~ ez et al. (1975), Nún~ ez and Rossi de García (1981) and In a regional geological context, the Valcheta Petrified Forest
Caminos et al. (2001) which related the fossiliferous strata to Upper (Fig. 1) is located in the north-eastern region of the Somún Cura
Cretaceous deposits. Finally, Vejsbjerg et al. (2008) provided a first Massif (Stipanicic and Methol, 1972; Page et al., 1999), a mor-
preliminary and integrated report on the paleobotanical and phostructural unit that has also been referred by other authors as
geological aspects of the site. In spite of them, an exhaustive study the “Comarca Nordpatago nica” (Stipanicic and Methol, 1980) or
on the fossil trunks assemblage of the Valcheta site have not been “Macizo Nordpatago nico” (Bonaparte et al., 1988) among others
formally carried out until now. names. Located between 40 and 44 S and 65 e70 W, the Somún
The objective of this study has been to stratigraphically locate, Cura Massif is characterized by the expose the crystalline basement
measure and anatomically analyze the fossil trunks exposed in the (metamorphic and plutonic rocks) of the Lower Paleozoic Nahuel
entire area of the Valcheta Petrified Forest, reconstruct their ver- Niyeu Formation and Valcheta Plutonic Complex (although alter-
tical structure and characterize the paleoenvironmental context in native names have been used by other authors).
which the trees lived. At the beginning of the Mesozoic, the region was subjected to
intense tectonic stresses related to the fragmentation of Gondwana.
2. Materials and methods Consequently, during the Triassic a set of small cratonic basins
associated with rift systems were developed, which was followed
Wood remains of the Valcheta Petrified Forest are exposed on by significant magmatism and acidic to mesosilicic volcanism
the surface in an area of ~0.23 km2 (around 40 400 3300 S, 66 090 0000 during the Early to Middle Jurassic (Uliana et al., 1985; Kay et al.,
W) in the surround of the Valcheta town, in the Northeast Río 1989; Nullo, 1991). In the surrounding area of the Valcheta town,
Negro Province, Argentina (Fig. 1). The fossil-bearing levels of these thickness successions of volcanic beds are integrated into the
silicified logs are here referred to deposits of the Allen Formation so-called Marifil Volcanic Complex or Marifil Formation (Nún ~ ez
(middle-upper Campanian e lower Maastrichtian, Malargüe et al., 1975; Franchi et al., 1975; Page et al., 1999; Caminos et al.,
Group) as discussed below. The most complete fossil trees consist of 2001; Garrido, 2019). Even though, during the Jurassic and most
large trunks that retain basal parts of their branches and roots. Of of the Cretaceous periods, the Somún Cura Massif would have been
ca. 200 petrified logs surveyed in the area (Vejsbjerg et al., 2008), a positive relief, serving as detrital source area for the surrounding
only those specimens that made it possible to clearly identify their basins. During the Campanian (Late Cretaceous) there was a reor-
sense of growth (from base to top) were considered for the mea- ganization of the South American tectonic plate resulting in a
surement of their orientation. Likewise, all those specimens strong regional subsidence. Consequently, the first pulse of Atlantic
(generally fragmentary) with evidence of displacement from their Ocean ingression into the region took place from the East (Uliana
original position by anthropic action were also discarded from this and Biddle, 1988; Legarreta et al., 1989; Macdonald et al., 2003).
analysis. For this reason, the complete data collection of each log This process meant a deep change in the morphostructural
(location, orientation and maximum diameter) was reduced to 59 configuration of the latest Cretaceous basins and their depocentres,
specimens (Table 1, Suppl. mat. 1). The orientation has been taken giving rise to a coastal to marine sedimentary succession, strati-
by using a Brunton-type geological compass considering the graphically identified in the north-Patagonian sector as the
growth direction of the tree (from base to top) whiles the location Malargüe Group. The Malargüe Group (middle Campanian e
was taken with GPS Garmin Etrex 20X. The maximum diameter was Selandian) is integrated in the Valcheta area, by the Allen Forma-
employed to estimate the original height of each specimen tion (middle-upper Campanian e lower Maastrichtian) and the
applying the methodology proposed by Niklas (1993, 1994). The age Arroyo Barbudo Formation (lower Maastrichtian e Danian) (Lizuaín
of the Valcheta trees was estimated by means of the equations of Fuentes and Sepúlveda, 1979; Manera de Bianco, 1996; Page et al.,
Veblen et al. (1981) as explained below. 1999; de la Fuente et al., 2001; Caminos et al., 2001; Garrido,
Logs are silicified and consist mainly of secondary xylem (tra- 2019); while toward the west in the central Neuque n basin, the
cheidoxyls). All specimens studied are housed in the Paleobotanical Group is composed from base to top by the Allen, Jagüel, Roca and
collection of the Museo Provincial María Ine s Kopp (Valcheta, Río El Carrizo Formations (Uliana and Dellape , 1981; Barrio, 1990a;
Negro, Argentina), under the initials #specimen/P/2022. Nine fossil Rodríguez et al., 2007; Rodríguez, 2011; among others).
woods were thin sectioned in transverse (TS), longitudinal tangen- Previous studies suggested the presence of continental deposits
tial (TLS) and longitudinal radial (RLS) sections mounted in micro- from the Neuque n Group (Cenomanian e middle Campanian) in
scope slides and studied using light Microscopy (Olympus CH30 and the study area, which in turn would be the carriers of the petrified
BX51). At least 25 measures were taken when possible. Measures are logs of the Valcheta town area (Nún ~ ez et al., 1975; Nún
~ ez and Rossi
presented as the mean value, along with maximum and minimum de García, 1981; Caminos et al., 2001). However, based on the
measures. For the generic classification of the studied woods we analysis of the regional distribution of the Cretaceous units, their
followed the key for fossil genera proposed by Philippe and Bamford tectosedimentary evolution and paleocurrent design, these de-
(2008). Descriptive terminology used here follows the IAWA list of posits and consequently the logs that it carries are here attributed
microscopic features for softwood identification (IAWA Committee, to the Allen Formation (middle-upper Campanian e lower Maas-
2004), with the addition of terms defined in Philippe and Bamford trichtian), the lower unit of the Malargüe Group.
(2008) and Boura et al. (2021). Seriation and contiguity indices The Allen Formation is regionally characterized by exhibiting a
proposed by Pujana et al. (2016) were estimated. remarkable vertical and lateral variation of facies. Its depositional
An idealized geological profile was performed on the basis of environments range from shallow and coastal marine environ-
multiple outcrops (Fig. 2). A lithofacial analysis of the deposits ments (tidal plains) to fully continental ones with development of
carried out the fossil logs are here provided for the first time for the fluvial systems and eolian deposits (Andreis et al., 1974; Barrio,
Valcheta Petrified Forest area. Paleocurrent measurements were 1990b, 1991; Hugo and Leanza, 2001; Armas and S anchez, 2015).
2
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 1. A. Map showing the approximate limits of the Macizo Somún Cura (dark gray area, based on Page et al., 1999) and the Valcheta area reproduced in B (square). Black triangles
show Allen Formation localities with fossil plants (Bajo de Santa Rosa, BSR and Salitral Ojo de Agua, SOA) and palynomorphs (Lomas Coloradas, LC and Lago Pellegrini, LP). Black
diamond shows the uppermost Cretaceous locality El Quiosco (EQ, Puntudo Chico Formation) which also includes records of Podocarpoxylon mazzonii. B. Geology of the Valcheta
area, partially reproduced from the geological map 1:25,000 “Area Valcheta, Monumento Natural Bosque Petrificado” elaborated by Alberto C. Garrido (in Vejsbjerg et al., 2019). C.
Detail of the Valcheta Petrified Forest area showing the outcrops of the Allen Formation.
Table 1
Valcheta Petrified Forest, summary of orientation, length and diameter measured on 59 logs and their original estimated height (Hest), critical height (Hcrit) and security factor
(SF) using the allometric equations of Niklas (1992, 1993, 1994) and Creber and Ash (2004). The age range of the trees was estimated using the growth rate of extant southern
South American podocarp species Podocarpus nubigenus and Saxegothaea conspicua as modern analogs (regression equations of Veblen et al., 1981). The complete data is
provided on Supplementary material 1.
Orientation ( ) Max. length Diameter Hest (m) Hcrit (m) SF Age range (years)
measured (m) measured (m)
This succession of facies is related to the first pulse of the Atlantic At the Valcheta Petrified Forest, the Allen Formation deposits
Ocean ingression in northern Patagonia during the Late Cretaceous are patchy exposed and consist of small and discontinuous outcrops
which flooded the areas with the greatest subsidence (Uliana and (Figs. 1, 3). They have been deposited on a paleorelief carved in eo-
Biddle, 1988; Legarreta et al., 1989). Paleozoic rocks of the Nahuel Niyeu Formation and are mostly
3
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 3. Outcrops and lithofacies of Allen Formation in the Valcheta Petrified Forest area. A. General aspect of outcrops. B. Fragment of silicified log (white arrows) included in
psammitic channel-fill deposits. C. Detail of massive gravelly-sandstone facies (SGm). D. Detail of massive sandstone facies (Sm) with variable granulometry. E. Gravelly sandstone
facies with horizontal and low-angle cross-bedding facies (SGh/l). F. gravelly sandstones with planar cross-bedding facies (SGp).
Petrified Forest area. Many of the logs, in fact, are largely contained trunks may have suffered some displacement with respect to their
in this detrital cover showing that they were exposed on the surface original deposit. The Quaternary sediment shows a domain of
before being buried again by Quaternary sediments. These cir- medium to fine sandstones, texturally immature, with a significant
cumstances arise reasonable doubt about the possibility that the percentage of silty-cineritic matrix. They are massive in appearance
5
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 4. AeB. Alluvial Quaternary deposits in the Valcheta Petrified Forest area containing small reworked wood fragments (white arrows).
and include isolated clasts (pebble size) of basic and acidic vol- diameter, often preserved as Steinkerne (Gothan, 1905; Boura et al.,
canites together with small reworked wood fragments (Fig. 4). 2021). Tracheid tangential diameter 85.1 (22.3e74.0) mm. Intercel-
These deposits, which rarely exceed 1.5 m in thickness, indicate an lular spaces rare to absent. Tangential pits and axial parenchyma
alluvial origin with flows of high sediment load. The energy of the not observed. Cross-field pitting podocarpoid, 1e2 circular to
flows that originated the Quaternary detrital cover would have pointed half-bordered (with wide borders) pits per cross-field, with
been insufficient to move large logs; fact that is also suggested by oblique apertures not exceeding the width of the border at its
the good state of conservation of the logs which preserve in some widest point, 13.8 (9.6e18.0) mm in vertical diameter. Rays homo-
cases small portions of branches and roots. Likewise, the orienta- geneous, parenchymatous, with cells 22.6 (16e29.2) mm in vertical
tion exhibited by the logs, and its correspondence with the sedi- diameter. Rays mostly biseriate, up to 15 cells high, but including
mentological structures, agrees in that logs still reflex the original some uniseriate very low (4 cells high) and others 1-2-seriate in
position in the Cretaceous deposits. part very high (up to 40 cells high) or, less frequently, triseriate and
multiseriate with up to four cells wide. Ray frequency of 6 (4e8)
4. Systematic paleontology rays per mm.
Comparisons. The anatomical features of the Valcheta fossil woods
Fossil Genus Podocarpoxylon Gothan 1905 (tracheid radial pits of abietinean type, cross-field pitting podo-
Type species Podocarpoxylon juniperoides Gothan, in Gagel 1906 carpoid with <4 pits per-cross field and absence of both helical
thickenings and resin channels) allow to include them in the genus
Podocarpoxylon mazzonii (Petriella) Müller-Stoll et Schultze-
Podocarpoxylon Gothan 1905, according to the distinctive features
Motel 1990
of this taxon reviewed by Philippe and Bamford (2008) and Pujana
(Figs. 6e8)
and Ruiz (2017). Additionally, the distinctively presence of biseriate
Messembrioxylon mazzonii Petriella 1972, p. 173, fig. 3, Pl. 3, AeE and very high rays, tracheids with biseriate opposite radial pitting,
Circoporoxylon gregussii Del Fueyo 1998, p. 45, Pl. 1 abundant septa like-structures and the 1e2 circular pits per cross
Studied specimens. 1750/P/2022 e 1758/P2022 field justify their assignment to the species P. mazzoni (Petriella)
Locality. Valcheta Petrified Forest, Valcheta town, Río Negro Prov- Müller-Stohl et Schultze-Motel 1990.
ince, Argentina. Podocarpoxylon mazzonii has been previously identified in central
Stratigraphic horizon. Allen Formation; Upper Cretaceous (middle- and northern Patagonia sequences (Puntudo Chico, Bororo , Sala-
upper Campanian e lower Maastrichtian). manca and Pen ~ as Coloradas Formations) ranging from the upper-
Description. Silicified trunks with a monopodial growth consisting most Cretaceous to the Danian/Selandian (Petriella, 1972;
of a main orthotropic axis with lateral branch bases helically ar- Raigemborn et al., 2009; Brea et al., 2011; Vera et al., 2019).
ranged. Maximum lengths measured from the most completely Recently, Ro mbola et al. (2021) informed the presence of P. mazzonii
preserved specimens ˷22e23 m. Logs with 21e62 cm in diameter in the Cardiel Formation at the northern area of the Cardiel Lake,
(without cortex), mostly without considerable lateral compression. southwestern Patagonia, considered as Cenomanian in age in this
Tracheidoxyl (picnoxylic) wood with growth ring boundaries area due to its apparent continuity with the underlying Albian
mostly distinct, difficult to observe in some specimens. Latewood Piedra Clavada (¼Kachaike) Formation (Ramos, 1982; Panza et al.,
consisting of few layers (1e3 cell in thickness) of tangentially- 2018). Nevertheless, the age of these deposits should be regarded
compressed tracheids. Transition from earlywood to latewood as tentative. In fact, some authors have postulated a wider tentative
abrupt. In longitudinal section, tracheids usually show transverse age range (Cenomanian to lower Coniacian, or even younger; Panza
interruptions (septa-like structures, Philippe et al., 2013). Tracheid et al., 2018, and cites therein). Thus, besides this possible older
radial pitting areolate, uni (68%) to biseriate opposite (32%), very record, P. mazzonii seem to be bracketed between the Campanian to
infrequently triseriate opposite, abietinean type with no Sanio rims the Danian.
evidences (11% uniseriate contiguous, 57% uniseriate distant, 32% According to Vera et al. (2019) the specimens originally assigned to
biseriate opposite; Si ¼ 1.32), with Cp ¼ 14%. When biseriate, Circoporoxylon gregussi by Del Fueyo (1998) are conspecific with
mostly opposite. Radial pits circular 21.3 (15.5e27.2) mm in vertical P. mazzonii, resulting a junior synonym of the latter. This implies a
6
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 5. A. location of exposed logs (circles) in the main area of Valcheta Petrified Forest. Red circles indicate those specimens sampled for anatomical study. Triangles indicate the
midpoint of the outcrops where the paleocurrent data were obtained. The Latitude and longitude geographic coordinates expressed as decimal degrees. B. Rose diagrams of
paleocurrent (left) and orientation of petrified logs (right). Class interval: 30 . Both diagrams are presented in different scales, adjusted for best comparison. C. Histogram showing
the frequency distribution of the estimated tree height (left) and basal diameter measured (right), based on 59 specimens. D. Southenorth transect (aea’ in figure A) of Valcheta
forest logs showing their estimated height (when possible) and latitudinal distribution in the area with the highest concentration of specimens. Red circles indicate those specimens
sampled for anatomical study. Dotted line indicates the average estimated height (21,59 m). Vertical bars on the left indicate (maximum height/maximum diameter in meters) of the
extant selected South American Podocarpaceae (Podocarpus parlatorei, P. lambertii, P. nubigenus, P. saligna, Prumnopitys andina and Saxegothaea conspicua, data from Lusk, 1996;
Donoso, 2013). (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article).
previous record for this species at the same Allen Formation, but the mid-Cretaceous Mata Amarilla Formation in southern Patagonia
from the Bajo de Santa Rosa, a locality some 100 km northwest of the (Varela et al., 2016), is clearly distinguished from P. mazzonii by
Valcheta Petrified Forest. Podocarpoxylon garciae Del Fueyo (1998) is having uniseriate radial pitting and uniseriate rays.
another species erected to encompass fossil logs from the Allen Other Podocarpoxylon species have been identified from Mesozoic
Formation at Bajo de Santa Rosa locality. P. garciae, also recognized in and lower Paleogene deposits of Patagonia, including: P. atuelensis
7
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 6. Logs exposed in the visitor area of the Valcheta Petrified Forest. A. Specimen with its root system partially preserved showing flared-like base. B. Detail of log with blocky
texture suggesting possible evidence of fungal decay (brown-rot type). C. View from the cup to the base of the specimen illustrated in figure A showing lateral branches partially
preserved (white arrows). D. Specimen with the largest measured basal diameter, also showing flared-like base. (For interpretation of the references to color in this figure legend,
the reader is referred to the Web version of this article.)
Gnaedinger et al. (2015), P. austroamericanum (Gnaedinger, 2007), 2001). Noteworthy, none of these species has the profusely biseri-
€usel, 1924; Novas et al., 2019), P. feruglioi (Gnaedinger,
P. dusenii (Kra ate and very tall rays characteristic of P. mazzonii.
2007), P. multiparenchymatosum (Pujana and Ruiz, 2017; Ruiz et al., Tortorelli (1941) described a single specimen of wood collected in
2017), P. paleoandinum (Nishida, 1984), P. paleosalignum (Nishida, the Valcheta area (most certainly representing part of the same
1984), and P. prumnopityoides (Gnaedinger et al., 2017). Addition- assemblage here studied), tentatively referring it to Ginkgoales.
ally, species of this genus reported from Antarctica include Preservation quality of this specimen is not good, and radial sec-
P. aparenchymatosum (Gothan, 1908; Pujana et al., 2014), tions show scarce details, in particular in reference to cross-fields
P. chapmanae (Poole and Cantrill, 2001), P. communis (Poole and pitting, which is not preserved. Most informative features of the
Cantrill, 2001) and P. fildense (Zhang and Wang, 1994; Poole et al., wood described by Tortorelli (1941) are the presence of growth
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M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 7. Podocarpoxylon mazzonii (Petriella) Müller-Stoll et Schultze-Motel. AeC. Transverse section. A. Growth rings with distinct boundaries (1754/P/2022). BeC. Detail of growth
rings showing abrupt transition from earlywood to latewood, the latter with little development (white arrowheads) (1754/P/2022). DeJ. Longitudinal tangential sections. D. General
view showing the variability in size and width of rays (1755/P/2022). E. Uniseriate (white arrowheads) and biseriate rays (black arrowhead) (1751/P/2022). F. Biseriate (white
arrowhead) and triseriate rays (black arrowhead) (1754/P/2022). G. Detail of uniseriate, very low rays (white arrowheads) and a biseriate ray of medium height (black arrowhead).
Note also the septa transversally disposed in the tracheids (black arrows) (1755/P/2022). H. Detail of a high ray (between white arrowheads) with biseriate ends and uniseriate
central portion (1755/P/2022). IeJ. Detail of a triseriate (1755/P/2022) and an infrecuent multiseriate ray respectivelly (1755/P/2022). Scale bars: A ¼ 1 mm; B, D, H ¼ 500 mm;
EeG ¼ 200 mm; C, IeJ ¼ 100 mm.
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M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
rings, uniseriate to biseriate spaced radial pitting, and uniseriate of the original trees from partial log. A comparison with the results
rays (although a single biseriate ray is present), and transverse obtained by these authors for two Jurassic-Cretaceous forests shows
interruptions in the tracheids. Although absence of cross-fields tree heights values up to c. 50% higher than those that would be
pitting precludes exhaustive comparisons and taxonomic assign- obtained by applying the equation of Niklas (1993, 1994) here used
ments, resulting in a tracheidoxyl with indeterminate generic for the Valcheta Forest. The last indicate that the heights predicted
placement, it is worth mentioning that most of the identified fea- for the Valcheta Petrified Forest may be underestimated, and the
tures are shared with Podocarpoxylon mazzonii (i.e. defined growth obtained values may be considered as minimum tree height esti-
rings, tracheids with uni to biseriate spaced radial pitting, and mations. Notwithstanding this, the Hest.av. obtained is consistent
transverse interruptions), but the presence of rays almost exclu- with maximum lengths measured from the most completely pre-
sively uniseriate (with only a biseriate ray recognized) seem to served fossil logs (20.08 me23.02 m) suggesting that the Niklas
separate it from all described specimens of this fossil species. Brea equation used for the tree height prediction gives plausible values.
et al. (2011) describe Paleocene Podocarpoxylon mazzonii specimens A histogram of frequencies show that the Hest values follow an
with rays dominantly uniseriate, occasionally biseriate, and very almost normal bell-shaped distribution, with the ~78% of the tree
rarely triseriate, indicating that uniseriate rays may be propor- with heights between ˷18e26 m and a younger and older cohorts
tionally dominant in the species. Nevertheless, biseriate rays are containing less than ~13% of the trees. This distribution is in accord
always regularly present (and indicated in the diagnosis provided with a mature forest. The security factor (SF) is a measure of the
by Petriella, 1972), thus the almost absence in the specimen studied capacity of a structural design (in this case the tree trunk) to
by Tortorelli (1941) seems to suggest a different taxonomic entity. withstand stressful situations (for example, the variable intensity of
Nevertheless, given the shared features between this specimen and the wind) without collapsing. Follow to Niklas (1992) the SF is here
the ones here studied, but also the absence of some elements in the obtained as the ratio of Hcrit/Hest. The average SF of ~3 for the Val-
sample studied by Tortorelli (1941), as well as the lack of infor- cheta Petrified Forest is between the expected value for a woody
mation regarding the proportion of uniseriate and biseriate rays in axis of perennial plant (Niklas, 1992).
the latter fossil, it is here preferred to avoid suggesting it may have The age of the Valcheta trees was estimated using the regression
been (or may have been not) the same taxon. equations of Veblen et al. (1981) obtained for the podocarp extant
species Podocarpus nubigenus Lindl. (Loge A ¼ 3.0844 þ 0.5717 Loge
D) and Saxegothaea conspicua Lindl. (Loge A ¼ 3.4062 þ 0.5576 Loge
5. Spatial distribution and estimated heights and ages of the D), where A is the predicted tree age (years) and D the log diameter
trees in the Valcheta forest measured (cm). Lusk (1996) point out that linear regression showed
diameter to be a significant predictor of age for P. nubigenus and
Most of the logs exposed on the surface are found within an area S. conspicua according to their own observations in tree populations
of ~0.23 km2, within the approximate limits of the protected natural from different sites. Both extant species are part of the temperate
area (Fig. 5A, Suppl. mat. 1). In this area, 59 logs were located, their rain forest of southwestern South America. The estimated age
orientation recorded (see above section) and their diameter and values for the Valcheta Petrified Forest specimens are expressed as
maximum length of each one was measured (Suppl. mat. 1, Table 1). a range whose limits are given by the differential annual growth
The diameter was recorded at a log height where this segment was rate in diameter of P. nubigenus and S. conspicua (being greater in
approximately constant. According to the proposal of Mosbrugger this last species, Enright and Jaffre, 2011). According to these pre-
(1990), logs with a critical diameter of less than 8 cm were not dictive equations, the mean estimated age for the trees of Valcheta
considered since they could be lateral branches. Based on the di- forest would be between 218 and 285 years with minimum and
ameters of trunks their critical buckling heights (Hcrit) were maximum ages of 125 and 514 years respectively range (Table 1 and
calculated using the formula proposed by Niklas (1993, p. 166; Suppl. mat. 1). Coomes and Bellingham (2011) pointed out that a
1994): Hcrit ¼ C (E/r)1/3D2/3, where C is a constant (0.792); E is great longevity is not a feature of podocarps, whereas it is for other
Young's modulus (kg m2); r is the wood density (kg m3) and D is conifers in the same forests. Southern Hemisphere extant podocarp
the log diameter (m). The values of E and r are species specific and species living in lowlands and lower montane tropical to temperate
cannot be obtained for fossil logs. Follow to Creber and Ash (2004) forests mostly exhibit longevities ranging 400-to 600-years
and Brea et al. (2011), the values used for E and r have been (Enright and Jaffre , 2011 and cites therein). The latter may sug-
958.1 106 kg m2 and 468.9 kg m3 respectively which consist of gest that the trees of the Valcheta would have been part of a mature
the averages for 18 extant conifers cited by Niklas (1993, table 3.4). forest with an older cohort close to its maximum life span.
For practical purposes and using these average values for E and r, Vertical reconstruction of the P. mazzonii tree population in the
this equation can be solved as follows: Hcrit ¼ 95.75 D2/3 (Creber Paleocene Ameghino Petrified Forest (Brea et al., 2011 Table 1) gives
and Ash, 2004). The Hcrit is the hypothetical maximum height to similar values in both average height (21.59 m versus 22.38 m in
which a tree trunk could be elevated before it undergoes elastic Valcheta forest) and height range (~14e34 m versus ~18e32 m in
buckling and collapse (Niklas, 1993). However, since the critical Valcheta forest). Brea et al. (2011) also provided a complete quan-
height is never reached, the original height of the Valcheta forest titative growth-ring analysis of P. mazzonii as autoecological and
canopy was estimated by applying the regression equation ob- paleoclimatological proxy. Based on these observations, those au-
tained by Niklas (1993, p.167): Hest ¼ 27.8 D1/3. thors suggest that this species had an evergreen habit and a
The Hest values obtained for the logs of the Valcheta Forest are possible long leaf retention times. According to Brea et al. (2011)
provided in Supplementary material 1 and summarized in Table 1 P. mazzonii specimens from the Ameghino Petrified Forest show a
and Fig. 5CeD. They suggest that the arboreal stratum had an latewood composed of 6e11 cell rows (Brea et al., 2011). As
average height (Hest.av) of 21.59 m, with a range of tree heights mentioned above, at the Valcheta Petrified Forest, the latewood
between ~14e34 m. does not exceed 2e3 cells in thickness, similarly to the informed for
Alternatively, to the curves of Mosbrugger (1990) and the here P. mazzonii from others latest Cretaceous to Paleocene floras
employed formula from Niklas (1993, 1994), other equations have (1e4 cell rows, Del Fueyo 1998; 1e6 cell rows, Vera et al., 2019 and
been proposed to estimate tree height from the diameter of fossil 1e2 cell rows, Petriella, 1972).
trunks (e.g. Pole, 1999; Philippe et al., 2009). Philippe et al. (2009) Varela et al. (2016) gave a characterization of a mid-Cretaceous
apply a useful alternative equation to calculate minimum heights podocarp-dominated forest based on silicified logs of
10
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Fig. 8. Podocarpoxylon mazzonii (Petriella) Müller-Stoll et Schultze-Motel. A-I. longitudinal radial sections. A. General view showing the variability in size of rays (between white
arrowheads) and tracheids with transverse septa (1751/P/2022). B. Growth ring showing abrupt transition from earlywood to latewood (black arrowheads) (1751/P/2022). C-E.
Detail of tracheids with areolate radial pitting, uni- and opposite biseriate (1755/P/2022). F. Detail of tracheid with opposite triseriate radial pitting (1757/P/2022). G. Detail of cross-
fields with one (white arrowheads) or two (black arrowhead) pits per field (1754/P2022). H-I. Detail of cross-fields showing single oculipore pits with an elliptical aperture included
in the border boundaries (1754/P2022). Scale bars: A-B = 500 mm; C-E = 40 mm; F-I = 20 mm.
11
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
Podocarpoxylon garciae Del Fueyo from the Mata Amarilla Forma- Urreta et al., 2011; Malumia n and Na n
~ ez, 2011). This is reflected
tion (Austral Basin, southern Patagonia). Among the attributes in a coastal to marine continental sedimentary succession, strati-
described, the authors provided a reconstruction of the vertical graphically known in the north-Patagonian as the Malargüe Group
structure of the tree population recorded in different localities. The (Garrido, 2019). In particular, the Allen Formation deposits are
estimated heights (based on 35 logs) ranging ˷15e30 m with regionally interpreted as continental and shallow marine envi-
average heights ranging ~18e25 m (Varela et al., 2016), values also ronment developed close to a marginal-littoral areas (Andreis et al.,
comparable to those obtained for the Valcheta Petrified Forest. 1974; Uliana and Dellape , 1981; Barrio, 1990b; Gonza lez Riga and
Casadio, 2000; Armas and S anchez, 2013, 2015). Furthermore,
6. The Valcheta Petrified Forest in the context of the north- Armas and S anchez (2011) recognized estuarine deposits at the top
patagonian plant communities and environments during the of the Anacleto Formation (underlying the Allen Formation) that
latest Cretaceous would correspond to the first evidence of the Atlantic ingression
toward an eastern sector of the Neuque n basin. The vertebrate re-
As mentioned above, the fossiliferous levels of the Valcheta cord from the Allen Formation at Bajo de Santa Rosa and neigh-
Petrified Forest can be correlated with the deposits bearing fossil boring areas includes terrestrial, freshwater and marine coastal/
logs located in the Bajo de Santa Rosa. Both sites consist of fluvial estuarine taxa which is congruent with the paleoenvironmental
deposits of the basal section of the Allen Formation. The identifi- reconstructions for this unit (Martinelli and Forasiepi, 2004;
cation of a single taxon of conifer in the Valcheta area could indicate Salgado et al., 2007; O'gorman et al., 2011). In the same sense, the
the development of a mature monotypic forest dominated by the mostly continental invertebrate association with few records of
evergreen species P. mazzonii, although it certainly expresses a marine ostracods seems to indicate that the Allen Formation at
certain preservation bias at the site. The paleoenvironmental Loma Puntuda (near to Bajo de Santa Rosa) was deposited in a
concordance between the fossil log deposits of Bajo de Santa Rosa shallow, fresh to oligohaline water body, near the sea coast line
and Valcheta allows us to suppose a certain continuity between (Carignano and Varela, 2011). In this paleogeographical context, the
both plant communities. In addition to the shared presence of braided channels identified for the Allen Formation in the area of
P. mazzonii in both sites, other plant taxa recognized only in the the Valcheta Petrified Forest, can be interpreted as part of a fluvial
Bajo de Santa Rosa, such as Podocarpoxylon garciae (Del Fueyo, system that would discharge its waters in a northeast direction in
1998) and a diversity of cycads (Artabe et al., 2004, 2005, 2010; the epicontinental sea.
Martínez et al., 2012) and palm trees (Ancibor, 1995) may charac- Drought stress is thought to be a major driver of extant podo-
terize these communities. To date, no other megafloristic elements carp forest structure and composition leading to increased
have been described for the Allen deposits. vulnerability during dry periods (Enright and Jaffre, 2011; Kitayama
However, other sources of information can provide elements et al., 2011). Absence of marked seasonality as suggested by the
that allow a plausible a more comprehensive characterization of narrow latewood ring (Fig. 7A) and the fact that it consisted of a
the different types of vegetation and environments in northern mature well-developed Podocarpaceae forest may be suggesting a
Patagonia at the time the Valcheta forest was developed. Palyno- relatively constant humidity and favorable growth conditions for
logical assemblages have been recovered from the Allen Formation the Valcheta Petrified Forest.
deposits although they were obtained from localities at a consid- After the warm global conditions estimated for the mid-
erable distance northwest from the studied site (Palamarczuk and Cretaceous, the beginning of a significant cooling process is recor-
Gamerro, 1988; Vallati, 2010, 2013; Pe rez Pincheira and Di ded from the Campanian, a trend that although occurring at a
Pasquo, 2018a, 2018b). The palynological assemblages described slower rate, continued through the late Campanian and Maas-
by Vallati (2010) from the Lomas Coloradas section (Neuque n trichtian (Clarke and Jenkyns 1999; Huber et al., 2002; Friedrich
Province) include two associations: a lower with predominance of et al., 2012; Linnert et al., 2014). However, Huber et al. (2002)
algae phytoplankton and the presence of heterosporous aquatic observed that the latitudinal gradients of the sea surface temper-
ferns as well as spores of terrestrial ferns, lycophytes and bryo- ature during the middle CampanianeMaastrichtian (ranging
phytes; and an upper association characterized by a minor presence ~0.13e0.22 C per degree of latitude) were considerably lower than
of aquatic palynomorphs but with abundant Podocarpaceae the mean modern-day SST gradient of ~0.40 C per degree of lati-
(Podocarpidites spp., Phyllocladidites mawsonii and Dacrycarpites tude. This low latitudinal thermal gradient could explain the fact
australiensis), and a dominance of putative Proteaceae (triporate that the floras of the Allen Formation (e.g. Valcheta Forest, Bajo de
and tricolpoidate pollen grains) among the angiosperms. According Santa Rosa, Lomas Coloradas) and others north Patagonian as-
to Vallati (2013) the palynoflora of the Allen Formation represents semblages developed under the relatively cool conditions that
an association of marked affinity with the AntarcticeAustralian prevailed worldwide during the Late Cretaceous, even when they
region, with dominance of Podocarpaceae and Proteaceae but represent plant communities with an austral affinity with domi-
with South America endemic pollen grain Tricesticillus tetris along nance of Podocarpaceae and Proteaceae, also include elements of
with few other paleo-equatorial elements as the miospore Gabo- warmer paleoequatorial climates such as cycads and palm trees.
nisporis and the palm pollen grain Spinizonocolpites (palm). In a
recent communication Pe rez Pincheira and Di Pasquo (2018a,
2018b) report a palynological association presumably from the 7. Conclusions
middle section of the Allen Formation in the area of the Pellegrini
Lake (Neuque n Province). It is characterized by a predominance of Based on the analysis of the regional distribution of the Creta-
spores of bryophytes, lycophytes, ferns and pollen of conifers ceous units, their tectosedimentary evolution and paleocurrent
(Podocarpaceae and Cheirolepidiaceae), monocots (Liliacidites), design, it is concluded that the logs that constitute the Valcheta
putative Proteaceae (Peninsulapollis gilli) and a diversity of chlor- Petrified Forest are preserved in deposits of the Allen Formation
ophyceaen algae and dinoflagellates. (middle-upper Campanian e lower Maastrichtian) ruling out the
During the latest Cretaceous north Patagonia experimented a presence of continental deposits of the Neuquen Group in the study
strong regional subsidence related to the break-up of Gondwana, area as was previously proposed.
resulting in the first Atlantic transgression that flooded in part the The fossil tree population consisted of a mature forest, with the
Colorado and Neuque n Basins (Uliana and Biddle, 1988; Aguirre- older cohort possibly close to its maximum life span, developed on
12
M.G. Passalia, A. Garrido, A. Iglesias et al. Cretaceous Research 142 (2023) 105395
the banks of a braided fluvial system that in the area had a pref- Ballent, S.C., Carignano, A.P., 2008. Morphological abnormalities in Late Cretaceous
and early Paleocene foraminifer tests (Northern Patagonia, Argentina). Marine
erential flow in a NE direction.
Micropaleontology 67, 288e296.
A single xylological type was recognized and assigned to Podo- Barreda, V.D., Cúneo, N.R., Wilf, P., Currano, E., Scasso, R., Brinhuis, H., 2012.
carpoxylon mazzonii (Podocarpaceae). Cretaceous/Paleogene floral turnover in Patagonia: drop in diversity, low
This taxon has a previous record to the Allen Formation, but extinction, and a Classopollis spike. PLoS One 7, e52455.
Barrio, C.A., 1990a. Late Cretaceous e Early Tertiary seeimentation in a semi-arid
from outcrops of the Bajo de Santa Rosa, where it was part of a plant foreland basin (Neuque n Basin, western Argentina). Sedimentary Geology 66,
community also composed by another podocarp species (P. garciae) 255e275.
together with cycads and palm trees. Barrio, C.A., 1990b. Paleogeographic control of Upper Cretaceous tidal deposits,
Neuque n Basin, Argentina. Journal of South American Earth Sciences 3, 31e49.
The estimated average height of the canopy of P. mazzonii Barrio, C.A., 1991. Lateral variations in tidal deposits of the Upper Cretaceous
population in the Valcheta Petrified Forest is similar to that esti- Neuque n Basin, western Argentina. In: Smith, D.G., Reinson, G.E., Zaitlin, B.A.,
mated for this taxon from a younger (Danian) forest in Patagonia. Rahmani, R.A. (Eds.), Clastic Tidal Sedimentology. Canadian Society of Petro-
leoum Geologist Memoir, Vol. 16, pp. 321e334.
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the canopy in the northern Patagonian floristic communities Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de
developed in environments relatively close to shallow seas during Tucuma n. In: Serie Correlacio n Geolo gica Nº 2, Tomo I. San Miguel de Tucuma n,
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Acknowledgments Brea, M., Matheos, S.D., Raigemborn, M.S., Iglesias, A., Zucol, A.F., Pr amparo, M.,
2011. Paleoecology and paleoenvironments of Podocarp trees in the Ameghino
Petrified forest (Golfo San Jorge Basin, Patagonia, Argentina): Constraints for
Our acknowledgments to Laila Vejsbjerg who has been the early Paleogene paleoclimate. Geolo gica Acta 9, 13e28. https://doi.org/10.1344/
promoter of this study and the Secretaría de Cultura de Río Negro 105.000001647.
Caminos, Chernicoff, C.J., Fauque , L., Franchi, M., Espejo, P., 2001. Hoja Geolo gica
by the paleontological permission. We thank to Romina Rial (Museo
4166-I, Valcheta, Provincia de Río Negro. In: Programa Nacional de Cartas
Provincial María Ines Kopp) and to the visitor service staff at the Geolo gicas de la República Argentina 1:250.000. Instituto de Geología y
Petrified Forest and the Tourism Office of Valcheta (Dora Saco, Recursos Minerales, Servicio Geolo gico Minero Argentino, Boletín Nº 310.
Romina Martínez, Luja n Lucero, Marta Curuhuinca, Elías San Martín Buenos Aires.
Carignano, A., Varela, J., 2011. Ostr acodos (Crustacea) de la Formacio n Allen
and Mariela Guzma n). We also thank the Laboratorio Petrogra fico (Cretacico Tardío), Cuenca Neuquina, Argentina. Revista Brasileira de Paleon-
CICTERRA (CONICET-Universidad de Co rdoba) for the service of tologia 14, 169e178.
wood thin sections. This work is dedicated to the memory of María Clarke, L.J., Jenkyns, H.C., 1999. New oxygen isotope evidence for long-term Creta-
ceous climatic change in the Southern hemisphere. Geology 27, 699e702.
Victoria Martínez Direne (Area de Discapacidad y Adultos Mayores, Collinson, J.D., 1986. Alluvial Sediments. In: Reading, H.G. (Ed.), Sedimentary En-
Valcheta). Thanks are extended to Eduardo Koutsoukos (Journal vironments: Processes, Facies and Stratigraphy, third ed. Blackwell Science,
Editor) and the anonymous reviewers for their suitable comments Oxford, pp. 37e81.
Coomes, D.A., Bellingham, P.J., 2011. Temperate and tropical podocarps: How
and suggestions that improved the final version of the manuscript. ecologically alike are they? (Chapter 7). In: Turner, B.L., Cernusak, L.A. (Eds.),
Study supported by a grant of the Consejo Federal de Inversiones, Ecology of the Podocarpaceae in Tropical Forests. Smithsonian Contributions to
Argentina (CFI Exp. N 17851 11 01). Botany no. 95. Smithsonian Institution Scholarly Press, Washington,
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15
Cretaceous Research
THE VALCHETA PETRIFIED FOREST (UPPER CRETACEOUS), NORTHERN PATAGONIA, ARGENTINA: A GEOLOGICAL AND PALEOBOTANICAL SURVEY
M.G. PASSALIA, A. GARRIDO, A. IGLESIAS, E.I. VERA
Supplementary Material 1. Valcheta Forest, complete data of log location, orientation, length and diameter measured, the estimated original height (Hest), critical
height (Hcrit) and security factor (SF). Age estimation using the growth rate of extant southern South American podocarp species Podocarpus nubigenus (1) and
Saxegothaea conspicua (2) as modern analogues
n/d = no data
Cretaceous Research
THE VALCHETA PETRIFIED FOREST (UPPER CRETACEOUS), NORTHERN PATAGONIA, ARGENTINA: A GEOLOGICAL AND PALEOBOTANICAL SURVEY
M.G. PASSALIA, A. GARRIDO, A. IGLESIAS, E.I. VERA
45°
36°
40°
Outcrop #1 45°
40° 40' 24,3" S 68°
66° 08' 54,7" W 32°
34°
80°
64°
48°
Outcrop #2 63°
40° 40' 29,2" S 54°
66° 08' 51,4" 19°
36°