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Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia
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Original article
Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia§ Candela Blanco-Moreno a,*, Bernard Gomez b, A´ngela Buscalioni a a b
Universidad Auto´noma de Madrid, Departamento de Biologı´a, Unidad de Paleontologı´a, Calle Darwin, 2, Cantoblanco, 28049 Madrid, Spain Laboratoire de Ge´ologie de Lyon, UMR CNRS 5276 LGLTPE, Universite´ Claude-Bernard Lyon 1, 2, rue Raphae¨l-Dubois, 69622 Villeurbanne, France
A R T I C L E I N F O
A B S T R A C T
Article history: Received 31 May 2017 Accepted 29 May 2018 Available online xxx
Eighty-three publications have been examined to recover information on the palaeogeography, the chronostratigraphy and the presence of the fossil fern genus Weichselia Stiehler in different depositional environments. This fossil fern foliage was reported in 72 localities worldwide. Its fossil record ranges from the Bathonian to the Cenomanian, predominating from the Berriasian to the Barremian, but still common during the Aptian to Cenomanian. Weichselia mostly occurred in continental depositional environments in Europe, whereas it has been mostly related to marginal marine depositional environments in Africa, North America, and India. Three species have been described: W. reticulata, W. peruviana, and W. negevensis. We analyse the measurements on specimens collected from sixteen localities as well as published data from twelve others. Our results suggest that: (i) the climate changes during the Aptian might have affected the pinnule size of Weichselia, as the Aptian-Cenomanian pinnules are larger than those from the Berriasian-Barremian; (ii) some differences might be explained by the metric variation of the frond parts; (iii) there are no clear differences in size between remains collected from different depositional environments; (iv) there is no sufficient evidence to determine how many species of Weichselia have existed.
C 2018 Elsevier Masson SAS. All rights reserved.
Keywords: Pteridophyta Gleicheniales Paleobotany Paleobiogeography Paleoclimatology Taphonomy Depositional environment
1. Introduction Weichselia reticulata (C. Stokes et Webb) Fontaine was first described from the English Wealden as Pecopteris reticulata by Stokes and Webb (1824), the generic name being later changed to Weichselia reticulata by Fontaine (1899). This fern has drawn attention due to its broad distribution, architecture and histology. Weichselia reticulata has been included within the Matoniaceae (Sender et al., 2015) or placed close (Bommer, 1910), in its own family, the Weichseliaceae (Alvin, 1971). In addition to W. reticulata, two other species have been proposed so far: Weichselia peruviana Zeiller from the Berriasian of two Peruvian localities, which probably had tripinnate instead of bipinnate sterile fronds (Zeiller, 1914), and Weichselia negevensis Silant. et Krassilov from the Aptian–Albian of Makhtesh Ramon, Israel, distinguished by its soral clusters (Silantieva and Krassilov, 2006). Nonetheless, for many authors W. reticulata is the only accepted species (e.g., Alvin, 1971; Sender et al., 2015).
§
Corresponding editor Jose´ B. Diez. * Corresponding author. E-mail address: [email protected] (C. Blanco-Moreno).
Weichselia shows a broad chronostratigraphic and geographic distribution and has been reported from a wide range of environments. The chronostratigraphic occurrence of Weichselia ranges from the Bathonian (Middle Jurassic) of Tunisia (Boureau and Caillon, 1958) to the Cenomanian–Turonian (lower Upper Cretaceous) of Italy (Gomez et al., 2002). The genus was geographically distributed throughout Africa, America, Asia, Europe, and India (Barnard, 1973). Finally, Alvin (1974) associated the occurrence of Weichselia with two types of deposits: continental deposits, and marine sediments and conditions. In the present study, the references published for Weichselia are explored with the objective of recovering data on the fossil record of the genus, its paleobiogeography and chronostratigraphic distribution. The depositional environments related to the sediments where Weichselia was collected were also collected. Finally, the metric variability of Weichselia is examined, based on the literature and on direct measurements, some of which consist of nearly complete primary and secondary pinnae. All these data should provide preliminary insights into the variability of Weichselia related to its biogeographic distribution, chronostratigraphy, and depositional environments. The three putative species of Weichselia are also discussed accordingly.
https://doi.org/10.1016/j.geobios.2018.05.001 C 2018 Elsevier Masson SAS. All rights reserved. 0016-6995/
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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Fig. 1. Specimens of Weichselia reticulata with various preservations. A, B. Bernissart (Barremian–Aptian, Belgium). A: IRSNB b 6838; B: IRSNB b 6839. C. Weald Clay Formation (Barremian, England), NHMUK PB V 51405. D. Lower Greensand Formation (Aptian, England), NHMUK PB V 40938. E. El Montsec (Barremian, Spain), MNHN.F 17701. F. Las
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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Table 1 Material measured for the genus Weichselia Stiehler. Locality, Country
Species
Chronostratigraphy
Depositional environment
Reference
Collection
Bernissart, Belgium
W. reticulata
Barremian-Aptian
Continental
IRSNB b
Bahariya Formation, Egypt Weald Clay Formation, England Isle of Wight, Lower Greensand Formation, England Beauvais, France
W. reticulata W. reticulata W. reticulata
Cenomanian Barremian Aptian
Marginal marine Continental Marginal marine
Seward, 1900; Alvin, 1971; Bommer, 1910; Present study Lejal-Nicol and Dominik, 1990 Harris, 1981; Present study Present study
W. reticulata
Barremian
Marginal marine
Present study
Fe´ron-Glageon, France Quedlinburg, Germany Makhtesh Hathira, Israel Makhtesh Ramon, Israel Todai Formation, Japan Jarash Formation, Jordan Mahis, Jarash Formation, Jordan Nahr es Zerka, Kurnub group Jordan Wadi Jabbok, Kurnub grooup, Jordan Wadi Kenetri, Kurnub group, Jordan Kailta, Kenya Mudayrij, Lebanese Republic Caleta del Paraiso, Peru ˜ onate, Peru Pin Preznosza and Lipnik, Poland Wasia Formation, Saudi Arabia El Montsec, Spain Escucha Formation, Spain Las Hoyas, Spain Jebel Dirra, Sudan Bir el Karma and Foum el Hassen, Tunisia Sweetwater, Texas, USA various African localities
W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W.
Barremian Barremian Aptian-Albian Aptian-Albian Aptian Aptian-Albian Albian Lower Cretaceous Lower Cretaceous Lower Cretaceous Lower Cretaceous Lower Cretaceous Berriasian Berriasian Hauterivian-Barremian Cenomanian Barremian Albian Barremian Albian Aptian-Albian Albian Unknown
Continental Continental Marginal marine Marginal marine Unknown Marginal marine Continental Unknown Unknown Unknown Unknown Unknown Unknown Unknown Marginal marine Marginal marine Continental Marginal marine Continental Unknown Marginal marine Marginal marine Unknown
Carpentier, 1927 Daber, 1968 Present study Silantieva and Krassilov, 2006 Kimura and Aiba, 1985 Abu Hamad et al., 2016 Hu and Taylor, 2014 Present study Present study Present study Present study Edwards, 1929; Present study Zeiller, 1914; Present study Zeiller, 1914; Present study Reymano´wna, 1965 El-Khayal, 1985; Present study Barale, 1979; Present study Sender, 2012 Present study Edwards, 1926; Present study Barale and Ouaja, 2001 Berry, 1928 Edwards, 1933
reticulata reticulata reticulata negevensis reticulata reticulata reticulata reticulata reticulata reticulata reticulata reticulata peruviana peruviana reticulata reticulata reticulata reticulata reticulata reticulata reticulata reticulata reticulata
NHMUK PB V NHM MNHN.F.; NHMUK PB V
NHMUK PB V
NHMUK PB NHMUK PB NHMUK PB NHMUK PB NHMUK PB UCBL-EM UCBL-EM
V V V V V
NHMUK PB V MNHN.F. MCCM-LH NHMUK PB V
Institutional abbreviations for collections: IRSNB b: Palaeontology collection housed in the Royal Belgian Institute of Natural Sciences, Brussels, Belgium; MCCM-LH: Las Hoyas collection in the Museo de Las Ciencias de Castilla-La Mancha, Cuenca, Spain; MNHN.F.: Palaeobotany collection in the Muse´um national d’Histoire naturelle, Paris, France; NHMUK PB V: Palaeobotany collection in the Natural History Museum, London, UK; UCBL-EM: palaeontological collections in the Universite´ de Lyon 1, Villeurbanne, France.
2. Material and methods 2.1. Material In this study, we combine metric information from the literature together with direct measurements taken on 459 specimens identified as Weichselia reticulata (Fig. 1), and collected from sixteen localities (Table 1; Appendix A). Weichselia has been the subject of many publications, some of which were published in local journals that are hard to find, and therefore were not included. Overall, 83 references were consulted (Appendix B). Those references with general or imprecise information (Berry, 1911; Koeniguer, 1966; Alvin, 1971; Barnard, 1973; Lejal-Nicol and Dominik, 1990; Vakhrameev, 1991; Schrank, 1992) were only considered when others backed the data. 2.2. Sorting categories
floral provinces (1991), including Africa, North America, South America, East Asia, Europe, and India. These provinces have been mapped onto their correspondent Cretaceous landmasses (Fig. 2(A)); Chronostratigraphy based on the Weichselia fossil record. It is organised in three time intervals: from Bathonian (168.3 Ma) to Tithonian (145 Ma), Berriasian to Barremian (125 Ma), and Aptian to Cenomanian (93.9 Ma); Depositional environments. These have been grouped in two kinds (Continental and Marginal marine) according to the descriptions of the sediments where Weichselia was unearthed (Appendix C). Finally, the relationship between burnt specimens and size might have been another interesting value to analyse, but direct preservational information associated with a significant number of specimens is available only in a few collections (e.g., collection of Las Hoyas).
The criteria used for collecting the data analysed on Weichselia encompass the following three categories:
2.3. Metrics
Biogeographic distribution of Weichselia. For this purpose, the distributions listed in previous works such as Barnard (1973), ElKhayal (1985), and Daber (1990), were incorporated. The geographic distribution is presented following Vakhrameev’s
The metric variables that have been considered on different parts of the frond of Weichselia are depicted in Fig. 3. Since the measurements in the literature are given either as mean or as minmax ranges, the data of the directly observed material follows the
Hoyas (Barremian, Spain), MCCM-LH 32687. G. Beauvais (Barremian, France), NHMUK PB V 8759. H. Wasia Formation (Cenomanian, Saudi Arabia), NHMUK PB V 68758. I. Kailta (Lower Cretaceous, Kenya), NHMUK PB V 32249. J. Mudayrij (Lower Cretaceous, Lebanese Republic), NHMUK PB V 20486. K. Caleta del Paraı´so (Berriasian, Peru´), UCBL-EM009. L. Wadi Kenetri (Lower Cretaceous, Jordan), NHMUK PB V 20500. M. Jebel Dirra (Albian, Sudan), NHMUK PB V 21712. N. Makhtesh Hathira (Aptian–Albian, Israel), NHMUK PB V 35146. All NHMUK specimen images are copyright of The Trustees of the Natural History Museum, London. Scale bars: 1 cm.
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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Fig. 2. A. Floral provinces used for the geographic distribution criterion, based on Vakhrameev (1991), and drawn onto an upper Lower Cretaceous map (120 Ma) (palaeomap based on Martı´n-Abad and Poyato-Ariza, 2013). B. Distribution of localities per geographic unit (left), chronostratigraphical unit (middle), and environment (right) for all Weichselia-bearing fossil localities. C. Distribution of Weichselia-bearing fossil localities with available metric information. EU: Europe; NA: North America; EA: East Asia; AF: Africa; SA: South America; IN: India.
same criterion. Specimens were either measured directly with a ruler or photographed with a Nikon Digital Camera D5100 and measured using ImageJ (Schneider et al., 2012). Measurements for each locality are recorded in Appendix D. All the information addressing the differences in size of the Weichselia species (W. reticulata, W. negevensis, and W. peruviana) have been combined with the sorting categories described above (Table 2).
but none of the Bathonian-Tithonian specimens, Indian localities, as well as Berriasian–Barremian Cretaceous African and East Asian localities provide measurements (Fig. 2(C)). Apart from these absences, the localities with metric information are more or less homogenously represented for the studied categories, although the African and the Aptian-Cenomanian localities are overrepresented (Fig. 2(B, C)).
3. Results
3.1. Geographic, chronostratigraphic, and environmental data
Our literature survey shows that, at the moment, Weichselia has been reported from 72 localities worldwide. Sixty-two publications give chronostratigraphic information: 3 Bathonian–Tithonian, 34 Berriasian–Barremian, and 25 Aptian–Cenomanian. The depositional environment is described in 44 articles: 20 continental and 24 marginal marine. Only 21 publications include metric data,
Weichselia appeared in the fossil record during the Middle Jurassic in North Africa (Boureau and Caillon, 1958) and Georgia (Shatilova et al., 2011). During the Berriasian–Barremian, Weichselia had a worldwide distribution including Africa, South America, East Asia, Europe, and India (Fig. 4). For the three time intervals, Weichselia has been mostly reported from Europe (42% of the
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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Fig. 3. Metric variables collected for this study, including the width of the primary and secondary rachis (RW1 and RW2, respectively), the length and width of the pinnules (PL and PW, respectively), the distance between secondary pinnae (DBP), and the insertion angle of the secondary pinnae (IA2).
Table 2 Minima and maxima metric values (in mm; *: mean value) gathered from the literature and our own measurements (see Table 1 for a complete list of localities and references). Category
Group
RW1
DBP
IA2
RW2
PL
PW
Geographic location
Africa North America South America East Asia Europe Berriasian-Barremian Aptian-Cenomanian Freshwater Brackish W. reticulata W. negevensis W. peruviana
3–30.5 2.5 2–15 – 0.5–33 0.5–33 3–20 0.5–33 3–20 0.5–33 3–20 2–15
8–16 – 5–15 – 1.2–16.2 1.2–16.2 8–12 1.2–16.2 8–12 1.2–16.2 10–12 5–15
70–90 45 50–85 – 50–90 50–90 45–90 50–90 45–90 50–90 75–90 50–85
0.2–2.4 – – 1–2 0.01–2.7 0.01–2.7 0.2–2.4 0.01–2.7 0.3–1.5 0.01–2.7 1.5* –
1–17 – 1.5–6 7–9 0.5–8 0.5–7 1–13 0.5–7 2–13 0.5–17 7* 1.5–6
0.8–4 – 1.5–3 2–4 0.4–4 0.4–4 1–4 0.4–3.5 1–4 0.4–4 3* 1.5–3
Chronostratigraphy Depositional environment Species
RW1: primary rachis width; DBP: distance between secondary pinnae; IA2: insertion angle of the secondary pinnae; RW2: secondary rachis width; PL: pinnule length; PW: pinnule width.
localities) and in marginal marine depositional environments (55% of the localities) (Fig. 2(B)). In Europe, in 60% of the localities the depositional environment was continental, whereas 71% of the African localities and 100% of those from North America, were marginal marine (Fig. 4). Weichselia was more common in Europe and in continental environments during the Berriasian-Barremian. However, during the Aptian–Cenomanian, its distribution was slightly reduced, and W. reticulata was notably related to marginal marine depositional environments in Africa (Fig. 4). Weichselia appeared in North America in the Aptian and apparently vanished in South America and India before the Aptian. The last records of Weichselia correspond to the Cenomanian in Europe (Gomez et al., 2002) and the north of Vakhrameev’s African Province (El-Khayal, 1985; Barale and Azar, 2004; El-Saadawi et al., 2016). 3.2. Metric data In Europe, Weichselia showed the greatest variability for all the variables except for the pinnule length. Pinnules are longer and wider in Africa and East Asia. In North America, both rachis width and the insertion angles are the smallest among all the geographic
areas, and the same happens to a lesser extent in South American specimens. In African localities, the minimum values for all variables are seemingly higher than those of Europe (Table 2). The European fossil record is very significant for the Berriasian– Barremian and mostly consists of continental environments. Thus, for these groups, the metric variables showed the greatest variability, except for the insertion angle and the pinnule length and width. Moreover, because Aptian-Cenomanian and marginal marine localities are mostly located in Africa, the minimum values for all variables are seemingly higher than the Berriasian– Barremian and continental values (Table 2). Harris (1981) related the pinnule size to preservation, as pinnules may have shrunk during burning. The measurements taken on specimens from Las Hoyas denote no difference relative to the various types of preservation, including impressions, compressions, and permineralizations with limonite. The variability observed in the three species of Weichselia shows that values of W. negevensis are within the range of those observed for W. reticulata (Table 2). However, the minimum values of W. negevensis for all variables are always higher than those of W. reticulata and W. peruviana, as it occurs throughout the African specimens. W. peruviana is also within the observed variability for W. reticulata (Table 2). Nonetheless, although the values of the
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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Fig. 4. Occurrences of the genus Weichselia Stiehler plotted on three Cretaceous palaeomaps at 170 Ma (A), 120 Ma (B), and 105 Ma (C). Star: continental locality; dot: marginal marine locality; square: unknown depositional environment. Palaeomaps based on Martı´n-Abad and Poyato-Ariza (2013).
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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insertion angles of the secondary pinnae are within the range of W. reticulata, they are more acute than those of other specimens from the Berriasian–Barremian. The primary rachis width is also particularly small in W. peruviana (Table 2).
4. Discussion A survey of the literature allows for a preliminary scenario on the palaeobiogeographic distribution, the fossil record and most common depositional environment of the fossil fern Weichselia. The gathered information includes studies since 1824. Some references obviously lack precise information on the chronostratigraphy, depositional environment, and metric descriptions, and not all geographic areas were studied with the same intensity (Dunhill et al., 2014). There is also a bias in the direct metric data since the size of the different variables varies along the primary and secondary pinnae, and not all parts of the frond could be measured for all localities. However, the whole data allow for a thorough exploration of the distribution and metric variability of the fossil record of Weichselia. Although more localities were included in our study, the distribution of Weichselia resemble those described by Barnard (1973), El-Khayal (1985), Daber (1990), and Vakhrameev (1991), which already comprised records of this taxon in Africa, North America, South America, Europe, and India. The unique exception is East Asia, whose fossil record was published from 1975 onwards and included in Daber (1990) and Vakhrameev (1991). When the geographic distribution and the chronostratigraphy are crossed, a pattern of Weichselia dispersion emerges. Most leptosporangiate fern families have similar distributions and dispersion patterns. Most of them appeared in subtemperate humid areas, at latitudes higher than 308 N and S, and then later spread towards lower latitudes (Skog, 2001). However, our results show that the leptosporangiate family Weichseliaceae appeared near the Equator at latitudes of 0–208 N during the Bathonian–Tithonian. Weichselia dispersed very quickly both northward and southward, and reached latitudes of 508 N and S by the Berriasian–Barremian. At the end of the Barremian, its fossil record is confined to latitudes of 408 N–208 S, and declines during the Aptian–Cenomanian interval. The distribution of Weichselia is more consistent with that of the fern savannah biome, which existed during the Jurassic and Cretaceous. This biome lost importance in the Aptian, probably due to the increase in humidity during this period (Coiffard et al., 2007). Global dispersion patterns have not yet been studied for many taxa. One of the few groups that have been thoroughly studied are charophytes, and although not closely related to Weichselia, the pattern of dispersal of this taxon in the Northern Hemisphere is parallel to that of the charophytes Atopocara trivolvis (Martı´nClosas and Wang, 2008) and Clavator harrisii (Martı´n-Closas, 2015), particularly for the late colonization of North America, which occurred during the Aptian. The fact that charophytes and Weichselia follow the same pattern would suggest that the barriers for colonization prior to the Aptian might have been common to these groups, even though their ecological preferences are very different, and the localities where they have been collected differ in most cases. Our analysis of metric data also offers a new perspective as the metric differences are only observed for the pinnule size, and the variability of this metric is apparently associated with the chronostratigraphy. The length of Weichselia pinnules is larger in the Aptian-Cenomanian localities of Africa, East Asia, and Europe. The specimens collected and measured from the Albian localities of the Escucha Formation in NE Spain (Sender, 2012) show larger pinnules (i.e., 6–8 mm long), and differ in size with
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those collected from other European localities (0.5–7 mm long), which are all Berriasian–Barremian in age. Nevertheless, it should be noted that all localities reported so far from Africa and Eastern Asia are Aptian–Cenomanian in age. Therefore, this assessment requires further measurements in Berriasian–Barremian localities around the world. The size differences found in Weichselia might be related to a hothouse event with a rise in humidity, as suggested by Kidder and Worsley (2012) for the Aptian. This climatic shift might have positively affected the pinnule growth of Weichselia, because drier, nutrient-poor or high-irradiance conditions produce smaller pinnules in living ferns (Arens, 1997; Creese et al., 2011). The depositional environment does not seem to play a role in the differences in Weichselia frond size. The pinnules described from the marginal marine depositional environments of Preznosza and Lipnik (Reymano´wna, 1965) and Beauvais, are similar in size (5 mm long and 2–5 mm long, respectively) to the pinnules described in other continental Berriasian–Barremian localities. In most localities, Weichselia might have been parautochthonous because only fragments of fragile primary pinnae and disarticulated secondary pinnae preserved as charcoals have been collected (Gomez et al., 2001). Thus, the expected bias due to transportation and posterior burial in an environment where the fern did not grow is not so extreme. However, a more in-depth study of the taphonomy of each site must be conducted in order to fully understand the implications of the depositional environment on frond size. Finally, the metric variability among the species of Weichselia appears within the ranges obtained for W. reticulata for all variables. The min-max ranges for W. reticulata have been extended when direct measurements on the specimens have been performed. In addition, the differences found between the three species could probably be explained by the heterogeneity of the parts collected in each locality (e.g., W. peruviana is based on data from the apical part of the primary pinna) and/or chronostratigraphy (W. negevensis is only Aptian–Cenomanian in age). Acknowledgements We thank three anonymous reviewers for their comments which helped us to improve the original manuscript. We thank Santiago Langreo and Mercedes Llandres from the Museo de Paleontologı´a de Castilla-La Mancha, Cuenca, Spain, for enabling the study of the specimens from Las Hoyas, and Emmanuel Robert for granting us access to the material from Peru´ housed in the palaeontological collections at the Universite´ Lyon 1, Villeurbanne, France. Candela Blanco-Moreno is supported by a FPI-UAM Ph.D. scholarship from the Universidad Auto´noma de Madrid. Collections in the Universite´ Lyon 1 were studied thanks to a FPI-UAM travel grant provided by the same university. Collections in the Muse´um national d’Histoire naturelle, Paris; Natural History Museum, London; and Royal Belgian Institute of Natural Sciences, Brussels, were studied with the support of three grants from the SYNTHESYS Project (http://www.synthesys.info/), financed by the European Community Research Infrastructure Action under the FP7 Integrating Activities Program. We thank the staff from these three institutions for their assistance when accessing the collections. This work is part of the project CGL-2013-42643 P of the Spanish Ministry of Economy and Competitiveness and other projects.
Appendix A. Supplementary data Supplementary data (including the list of examined material [A], the list of localities used for the palaeobiogeographic analysis [B], depositional environment criteria [C], and details of the metric
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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analysis for each locality [D]) associated with this article can be found, in the online version, at https://doi.org/10.1016/j.geobios. 2018.05.001. References Abu Hamad, A.M.B., Amireh, B., El Atfy, H., Jasper, J., Uhl, D., 2016. Fire in a Weichselia-dominated coastal ecosystem from the Lower Cretaceous (Barremian) of the Kurnub Group in NW Jordan. Cretaceous Research 66, 82–93. Alvin, K., 1971. Weichselia reticulata (Stokes et Webb) Fontaine from the Wealden of Belgium. Me´moire de l’Institut Royal des Sciences Naturelles de Belgique 166, 1–33. Alvin, K., 1974. Leaf anatomy of Weichselia based on fusainized material. Palaeontology 17, 587–598. Arens, N.C., 1997. Responses of leaf anatomy to light environment in the tree fern Cyathea caracasana (Cyatheaceae) and its application to some ancient seed ferns. Palaios 12, 84–94. Barale, G., 1979. De´couverte de Weichselia reticulata (Stokes & Webb) Fontaine emend. Alvin, filicine´e leptosporangie´e, dans le Cre´tace´ Infe´rieur de la province de Le´rida (Espagne): implications stratigraphiques et pale´oe´cologiques. Geobios 12, 313–319. Barale, G., Ouaja, M., 2001. De´couverte de nouvelles flores avec des restes a` affinite´s angiospermiennes dans le Cre´tace´ infe´rieur du Sud tunisien. Cretaceous Research 22, 131–143. Barale, G., Azar, D., 2004. Cuticules ve´ge´tales dans le Cre´tace´ infe´rieur du Sud du Liban. Comptes Rendus Palevol 3, 119–124. Barnard, P.D.W., 1973. Mesozoic Floras. Palaeontology, Special Papers 12, 175–187. Berry, E.W., 1911. The Lower Cretaceous deposits of Maryland. Maryland Geological Survey, Maryland. Berry, E.W., 1928. Weichselia from the Lower Cretaceous of Texas. Journal of the Washington Academy of Sciences 18, 1–5. Bommer, C., 1910. Contribution a` l’e´tude du genre Weichselia. Bulletin de la Socie´te´ Royale de botanique de Belgique 3, 296–304. Boureau, E., Caillon, P., 1958. Sur la flore fossile et l’aˆge des couches de Taouratine, dans le bassin de Fort-Polignac (Sahara). Comptes Rendus Hebdomadaire des Se´ances de l’Acade´mie des Sciences 247, 2173–2175. Carpentier, A., 1927. La flore wealdienne de Fe´ron-Glageon (Nord). Me´moires de la Socie´te´ Ge´ologique du Nord 1, 1–151. Coiffard, C., Gomez, B., The´venard, F., 2007. Early Cretaceous angiosperm invasion of Western Europe and major environmental changes. Annals of Botany 100, 545– 553. Creese, L., Lee, A., Sack, L., 2011. Drivers of morphological diversity and distribution in the Hawaiian fern flora: trait associations with size, growth form, and environment. American Journal of Botany 98, 956–966. Daber, R., 1968. A Weichselia–Stiehleria–Matoniaceae community within the Quedlinburg Estuary of Lower Cretaceous age. Botanical Journal of the Linnean Society London 61, 75–85. Daber, R., 1990. Zur Pala¨ogeographie und Biologie von Weichselia reticulata im Hauterive-Barreˆme (Unterkreide). Feddes Repertorium 101, 319–331. Dunhill, A.M., Hannisdal, B., Benton, M.J., 2014. Disentangling rock record bias and common-cause from redundancy in the British fossil record. Nature Communications 5, 1–9. Edwards, W.N., 1926. Fossil plants from the Nubian Sandstone of eastern Darfur. Quarterly Journal of the Geological Society 82, 94–100. Edwards, W.N., 1929. Lower Cretaceous plants from Syria and Transjordania. Annals & magazine of natural history 4, 394–405. Edwards, W.N., 1933. On the Cretaceous fern Paradoxopteris and its connexion with Weichselia. Annals of Botany 47, 317–341. El-Khayal, A., 1985. Occurrence of a characteristic Wealden fern (Weichselia reticulata) in the Wasia Formation, central Saudi Arabia. Scripta Geologica 79, 75–88. El-Saadawi, W., Osman, R., El-Faramawi, M.W., Bkhat, H., Kamal-El-Din, M., 2016. On the Cretaceous mangroves of Bahariya Oasis, 36, Taeckholmia, Egypt, pp. 1–16.
Fontaine, W.M., 1899. Notes on Lower Cretaceous plants from the Hay Creek Coal Field, Crook County, Wyoming. U.S. Geological Survey Nineteenth Annual Report 2, 645–702. Gomez, B., Martı´n-Closas, C., Me´on, H., The´venard, F., Barale, G., 2001. Plant ˜ a delta (Late Barremian, taphonomy and palaeoecology in the lacustrine Un Iberian Ranges, Spain). Palaeogeography, Palaeoclimatology. Palaeoecology 170, 133–148. Gomez, B., The´venard, F., Fantin, M., Giusberti, L., 2002. Late Cretaceous plants from the Bonarelli level of the Venetian Alps, northeastern Italy. Cretaceous Research 23, 671–685. Harris, T.M., 1981. Burnt ferns from the English Wealden. Proceedings of the Geological Association 92, 47–58. Hu, S., Taylor, D.W., 2014. Floristics and paleoecology of an Early Cretaceous flora from Jordan. Bulletin of the Peabody Museum of Natural History 55, 153–170. Kidder, D.L., Worsley, T.R., 2012. A human-induced hothouse climate? GSA Today 22, 4–11. Kimura, T., Aiba, H., 1985. Discovery of Weichselia reticulata (Stokes et Webb) from the Lower Cretaceous Todai Formation, in the Outer Zone of Japan. Proceedings of the Japan Academy: Physical and Biological Sciences, Series B 61, 356–358. Koeniguer, J.C., 1966. E´tude pale´ophytoge´ographique du Continental Intercalaire de l’Afrique Nord-E´quatoriale. Sur de nouveaux e´chantillons du genre Paradoxopteris. Me´moires de la Socie´te´ ge´ologique de France 105, 100–112. Lejal-Nicol, A., Dominik, W., 1990. Sur la pale´oflore a` Weichseliaceae et a` Angiospermes du Ce´nomanien de la re´gion de Bahariya (Egypte du Sud-Ouest). Berliner Geowissenschftliche Abhandlunger 120, 922–957. Martı´n-Abad, H., Poyato-Ariza, F.J., 2013. Historical patterns of distribution in Pycnodontiform and Amiiform fishes in the context of moving plates. Geologia Belgica 16, 217–226. Martı´n-Closas, C., 2015. Cosmopolitanism in Northern Hemisphere Cretaceous Charophyta (Clavatoroidae). Palaeogeography, Palaeoclimatology, Palaeoecology 438, 9–23. Martı´n-Closas, C., Wang, Q., 2008. Historical biogeography of the lineage Atopochara trivolvis PECK 1941 (Cretaceous Charophyta). Palaeogeography, Palaeoclimatology, Palaeoecology 260, 435–451. Reymano´wna, M., 1965. On Weichselia reticulata and Frenelopsis hoheneggeri from the western Carpathians. Acta Palaeobotanica 6, 15–25. Schneider, C.A., Rasband, W.S., Eliceiri, K.W., 2012. NIH Image to ImageJ: 25 years of image analysis. Nature Methods 9, 671–675. Schrank, E., 1992. Nonmarine Cretaceous correlations in Egypt and northern Sudan: palynological and palaeobotanical evidence. Cretaceous Research 13, 351–368. Sender, L.M., 2012. Paleobota´nica (macrorrestos) y paleoambientes del Albiense ˜ a). (Creta´cico inferior terminal) del norte de la provincia de Teruel (Espan Universidad de Zaragoza, Spain (Ph.D. Thesis, unpubl.). Sender, L.M., Villanueva-Amadoz, U., Pons, D., Diez, J.B., Garcı´a-A´vila, M., Ferrer, J., 2015. New reconstruction of Weichselia reticulata (Stokes et Webb) Fontaine in Ward emend. Alvin, 1971 based on fertile remains from the middle Albian of Spain. Historical Biology 27, 460–468. Seward, A.C., 1900. La flore wealdienne de Bernissart. Me´moire du Muse´um d’Histoire Naturelle de Belgique 1, 1–37. Shatilova, I., Mchedlishvili, N., Rukhadze, L., Kvavadze, E., 2011. The history of the flora and vegetation of Georgia (South Caucasus). Georgian National Museum Institute of Paleobiology, Tbilisi. Silantieva, N.A., Krassilov, V., 2006. Weichselia Stiehler from Lower Cretaceous of Makhtesh Ramon, Israel: new morphological interpretation and taxonomical affinities. Acta Palaeobotanica 46, 119–135. Skog, J.E., 2001. Biogeography of Mesozoic leptosporangiate ferns related to extant ferns. Brittonia 53, 236–269. Stokes, C., Webb, P.B., 1824. Description of some fossil vegetables of the Tilgate Forest in Sussex. Transactions of the Geological Society of London 1, 421–424. Vakhrameev, V.A., 1991. Jurassic and Cretaceous floras and climates of the Earth. Cambridge University Press, Cambridge. Zeiller, R., 1914. Sur quelques plantes wealdiennes recueillies au Pe´rou par M. le Capitaine Berthon. Revue ge´ne´rale de Botanique 25, 647–674.
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Original article
Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia§ Candela Blanco-Moreno a,*, Bernard Gomez b, A´ngela Buscalioni a a b
Universidad Auto´noma de Madrid, Departamento de Biologı´a, Unidad de Paleontologı´a, Calle Darwin, 2, Cantoblanco, 28049 Madrid, Spain Laboratoire de Ge´ologie de Lyon, UMR CNRS 5276 LGLTPE, Universite´ Claude-Bernard Lyon 1, 2, rue Raphae¨l-Dubois, 69622 Villeurbanne, France
A R T I C L E I N F O
A B S T R A C T
Article history: Received 31 May 2017 Accepted 29 May 2018 Available online xxx
Eighty-three publications have been examined to recover information on the palaeogeography, the chronostratigraphy and the presence of the fossil fern genus Weichselia Stiehler in different depositional environments. This fossil fern foliage was reported in 72 localities worldwide. Its fossil record ranges from the Bathonian to the Cenomanian, predominating from the Berriasian to the Barremian, but still common during the Aptian to Cenomanian. Weichselia mostly occurred in continental depositional environments in Europe, whereas it has been mostly related to marginal marine depositional environments in Africa, North America, and India. Three species have been described: W. reticulata, W. peruviana, and W. negevensis. We analyse the measurements on specimens collected from sixteen localities as well as published data from twelve others. Our results suggest that: (i) the climate changes during the Aptian might have affected the pinnule size of Weichselia, as the Aptian-Cenomanian pinnules are larger than those from the Berriasian-Barremian; (ii) some differences might be explained by the metric variation of the frond parts; (iii) there are no clear differences in size between remains collected from different depositional environments; (iv) there is no sufficient evidence to determine how many species of Weichselia have existed.
C 2018 Elsevier Masson SAS. All rights reserved.
Keywords: Pteridophyta Gleicheniales Paleobotany Paleobiogeography Paleoclimatology Taphonomy Depositional environment
1. Introduction Weichselia reticulata (C. Stokes et Webb) Fontaine was first described from the English Wealden as Pecopteris reticulata by Stokes and Webb (1824), the generic name being later changed to Weichselia reticulata by Fontaine (1899). This fern has drawn attention due to its broad distribution, architecture and histology. Weichselia reticulata has been included within the Matoniaceae (Sender et al., 2015) or placed close (Bommer, 1910), in its own family, the Weichseliaceae (Alvin, 1971). In addition to W. reticulata, two other species have been proposed so far: Weichselia peruviana Zeiller from the Berriasian of two Peruvian localities, which probably had tripinnate instead of bipinnate sterile fronds (Zeiller, 1914), and Weichselia negevensis Silant. et Krassilov from the Aptian–Albian of Makhtesh Ramon, Israel, distinguished by its soral clusters (Silantieva and Krassilov, 2006). Nonetheless, for many authors W. reticulata is the only accepted species (e.g., Alvin, 1971; Sender et al., 2015).
§
Corresponding editor Jose´ B. Diez. * Corresponding author. E-mail address: [email protected] (C. Blanco-Moreno).
Weichselia shows a broad chronostratigraphic and geographic distribution and has been reported from a wide range of environments. The chronostratigraphic occurrence of Weichselia ranges from the Bathonian (Middle Jurassic) of Tunisia (Boureau and Caillon, 1958) to the Cenomanian–Turonian (lower Upper Cretaceous) of Italy (Gomez et al., 2002). The genus was geographically distributed throughout Africa, America, Asia, Europe, and India (Barnard, 1973). Finally, Alvin (1974) associated the occurrence of Weichselia with two types of deposits: continental deposits, and marine sediments and conditions. In the present study, the references published for Weichselia are explored with the objective of recovering data on the fossil record of the genus, its paleobiogeography and chronostratigraphic distribution. The depositional environments related to the sediments where Weichselia was collected were also collected. Finally, the metric variability of Weichselia is examined, based on the literature and on direct measurements, some of which consist of nearly complete primary and secondary pinnae. All these data should provide preliminary insights into the variability of Weichselia related to its biogeographic distribution, chronostratigraphy, and depositional environments. The three putative species of Weichselia are also discussed accordingly.
https://doi.org/10.1016/j.geobios.2018.05.001 C 2018 Elsevier Masson SAS. All rights reserved. 0016-6995/
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Fig. 1. Specimens of Weichselia reticulata with various preservations. A, B. Bernissart (Barremian–Aptian, Belgium). A: IRSNB b 6838; B: IRSNB b 6839. C. Weald Clay Formation (Barremian, England), NHMUK PB V 51405. D. Lower Greensand Formation (Aptian, England), NHMUK PB V 40938. E. El Montsec (Barremian, Spain), MNHN.F 17701. F. Las
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Table 1 Material measured for the genus Weichselia Stiehler. Locality, Country
Species
Chronostratigraphy
Depositional environment
Reference
Collection
Bernissart, Belgium
W. reticulata
Barremian-Aptian
Continental
IRSNB b
Bahariya Formation, Egypt Weald Clay Formation, England Isle of Wight, Lower Greensand Formation, England Beauvais, France
W. reticulata W. reticulata W. reticulata
Cenomanian Barremian Aptian
Marginal marine Continental Marginal marine
Seward, 1900; Alvin, 1971; Bommer, 1910; Present study Lejal-Nicol and Dominik, 1990 Harris, 1981; Present study Present study
W. reticulata
Barremian
Marginal marine
Present study
Fe´ron-Glageon, France Quedlinburg, Germany Makhtesh Hathira, Israel Makhtesh Ramon, Israel Todai Formation, Japan Jarash Formation, Jordan Mahis, Jarash Formation, Jordan Nahr es Zerka, Kurnub group Jordan Wadi Jabbok, Kurnub grooup, Jordan Wadi Kenetri, Kurnub group, Jordan Kailta, Kenya Mudayrij, Lebanese Republic Caleta del Paraiso, Peru ˜ onate, Peru Pin Preznosza and Lipnik, Poland Wasia Formation, Saudi Arabia El Montsec, Spain Escucha Formation, Spain Las Hoyas, Spain Jebel Dirra, Sudan Bir el Karma and Foum el Hassen, Tunisia Sweetwater, Texas, USA various African localities
W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W. W.
Barremian Barremian Aptian-Albian Aptian-Albian Aptian Aptian-Albian Albian Lower Cretaceous Lower Cretaceous Lower Cretaceous Lower Cretaceous Lower Cretaceous Berriasian Berriasian Hauterivian-Barremian Cenomanian Barremian Albian Barremian Albian Aptian-Albian Albian Unknown
Continental Continental Marginal marine Marginal marine Unknown Marginal marine Continental Unknown Unknown Unknown Unknown Unknown Unknown Unknown Marginal marine Marginal marine Continental Marginal marine Continental Unknown Marginal marine Marginal marine Unknown
Carpentier, 1927 Daber, 1968 Present study Silantieva and Krassilov, 2006 Kimura and Aiba, 1985 Abu Hamad et al., 2016 Hu and Taylor, 2014 Present study Present study Present study Present study Edwards, 1929; Present study Zeiller, 1914; Present study Zeiller, 1914; Present study Reymano´wna, 1965 El-Khayal, 1985; Present study Barale, 1979; Present study Sender, 2012 Present study Edwards, 1926; Present study Barale and Ouaja, 2001 Berry, 1928 Edwards, 1933
reticulata reticulata reticulata negevensis reticulata reticulata reticulata reticulata reticulata reticulata reticulata reticulata peruviana peruviana reticulata reticulata reticulata reticulata reticulata reticulata reticulata reticulata reticulata
NHMUK PB V NHM MNHN.F.; NHMUK PB V
NHMUK PB V
NHMUK PB NHMUK PB NHMUK PB NHMUK PB NHMUK PB UCBL-EM UCBL-EM
V V V V V
NHMUK PB V MNHN.F. MCCM-LH NHMUK PB V
Institutional abbreviations for collections: IRSNB b: Palaeontology collection housed in the Royal Belgian Institute of Natural Sciences, Brussels, Belgium; MCCM-LH: Las Hoyas collection in the Museo de Las Ciencias de Castilla-La Mancha, Cuenca, Spain; MNHN.F.: Palaeobotany collection in the Muse´um national d’Histoire naturelle, Paris, France; NHMUK PB V: Palaeobotany collection in the Natural History Museum, London, UK; UCBL-EM: palaeontological collections in the Universite´ de Lyon 1, Villeurbanne, France.
2. Material and methods 2.1. Material In this study, we combine metric information from the literature together with direct measurements taken on 459 specimens identified as Weichselia reticulata (Fig. 1), and collected from sixteen localities (Table 1; Appendix A). Weichselia has been the subject of many publications, some of which were published in local journals that are hard to find, and therefore were not included. Overall, 83 references were consulted (Appendix B). Those references with general or imprecise information (Berry, 1911; Koeniguer, 1966; Alvin, 1971; Barnard, 1973; Lejal-Nicol and Dominik, 1990; Vakhrameev, 1991; Schrank, 1992) were only considered when others backed the data. 2.2. Sorting categories
floral provinces (1991), including Africa, North America, South America, East Asia, Europe, and India. These provinces have been mapped onto their correspondent Cretaceous landmasses (Fig. 2(A)); Chronostratigraphy based on the Weichselia fossil record. It is organised in three time intervals: from Bathonian (168.3 Ma) to Tithonian (145 Ma), Berriasian to Barremian (125 Ma), and Aptian to Cenomanian (93.9 Ma); Depositional environments. These have been grouped in two kinds (Continental and Marginal marine) according to the descriptions of the sediments where Weichselia was unearthed (Appendix C). Finally, the relationship between burnt specimens and size might have been another interesting value to analyse, but direct preservational information associated with a significant number of specimens is available only in a few collections (e.g., collection of Las Hoyas).
The criteria used for collecting the data analysed on Weichselia encompass the following three categories:
2.3. Metrics
Biogeographic distribution of Weichselia. For this purpose, the distributions listed in previous works such as Barnard (1973), ElKhayal (1985), and Daber (1990), were incorporated. The geographic distribution is presented following Vakhrameev’s
The metric variables that have been considered on different parts of the frond of Weichselia are depicted in Fig. 3. Since the measurements in the literature are given either as mean or as minmax ranges, the data of the directly observed material follows the
Hoyas (Barremian, Spain), MCCM-LH 32687. G. Beauvais (Barremian, France), NHMUK PB V 8759. H. Wasia Formation (Cenomanian, Saudi Arabia), NHMUK PB V 68758. I. Kailta (Lower Cretaceous, Kenya), NHMUK PB V 32249. J. Mudayrij (Lower Cretaceous, Lebanese Republic), NHMUK PB V 20486. K. Caleta del Paraı´so (Berriasian, Peru´), UCBL-EM009. L. Wadi Kenetri (Lower Cretaceous, Jordan), NHMUK PB V 20500. M. Jebel Dirra (Albian, Sudan), NHMUK PB V 21712. N. Makhtesh Hathira (Aptian–Albian, Israel), NHMUK PB V 35146. All NHMUK specimen images are copyright of The Trustees of the Natural History Museum, London. Scale bars: 1 cm.
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Fig. 2. A. Floral provinces used for the geographic distribution criterion, based on Vakhrameev (1991), and drawn onto an upper Lower Cretaceous map (120 Ma) (palaeomap based on Martı´n-Abad and Poyato-Ariza, 2013). B. Distribution of localities per geographic unit (left), chronostratigraphical unit (middle), and environment (right) for all Weichselia-bearing fossil localities. C. Distribution of Weichselia-bearing fossil localities with available metric information. EU: Europe; NA: North America; EA: East Asia; AF: Africa; SA: South America; IN: India.
same criterion. Specimens were either measured directly with a ruler or photographed with a Nikon Digital Camera D5100 and measured using ImageJ (Schneider et al., 2012). Measurements for each locality are recorded in Appendix D. All the information addressing the differences in size of the Weichselia species (W. reticulata, W. negevensis, and W. peruviana) have been combined with the sorting categories described above (Table 2).
but none of the Bathonian-Tithonian specimens, Indian localities, as well as Berriasian–Barremian Cretaceous African and East Asian localities provide measurements (Fig. 2(C)). Apart from these absences, the localities with metric information are more or less homogenously represented for the studied categories, although the African and the Aptian-Cenomanian localities are overrepresented (Fig. 2(B, C)).
3. Results
3.1. Geographic, chronostratigraphic, and environmental data
Our literature survey shows that, at the moment, Weichselia has been reported from 72 localities worldwide. Sixty-two publications give chronostratigraphic information: 3 Bathonian–Tithonian, 34 Berriasian–Barremian, and 25 Aptian–Cenomanian. The depositional environment is described in 44 articles: 20 continental and 24 marginal marine. Only 21 publications include metric data,
Weichselia appeared in the fossil record during the Middle Jurassic in North Africa (Boureau and Caillon, 1958) and Georgia (Shatilova et al., 2011). During the Berriasian–Barremian, Weichselia had a worldwide distribution including Africa, South America, East Asia, Europe, and India (Fig. 4). For the three time intervals, Weichselia has been mostly reported from Europe (42% of the
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Fig. 3. Metric variables collected for this study, including the width of the primary and secondary rachis (RW1 and RW2, respectively), the length and width of the pinnules (PL and PW, respectively), the distance between secondary pinnae (DBP), and the insertion angle of the secondary pinnae (IA2).
Table 2 Minima and maxima metric values (in mm; *: mean value) gathered from the literature and our own measurements (see Table 1 for a complete list of localities and references). Category
Group
RW1
DBP
IA2
RW2
PL
PW
Geographic location
Africa North America South America East Asia Europe Berriasian-Barremian Aptian-Cenomanian Freshwater Brackish W. reticulata W. negevensis W. peruviana
3–30.5 2.5 2–15 – 0.5–33 0.5–33 3–20 0.5–33 3–20 0.5–33 3–20 2–15
8–16 – 5–15 – 1.2–16.2 1.2–16.2 8–12 1.2–16.2 8–12 1.2–16.2 10–12 5–15
70–90 45 50–85 – 50–90 50–90 45–90 50–90 45–90 50–90 75–90 50–85
0.2–2.4 – – 1–2 0.01–2.7 0.01–2.7 0.2–2.4 0.01–2.7 0.3–1.5 0.01–2.7 1.5* –
1–17 – 1.5–6 7–9 0.5–8 0.5–7 1–13 0.5–7 2–13 0.5–17 7* 1.5–6
0.8–4 – 1.5–3 2–4 0.4–4 0.4–4 1–4 0.4–3.5 1–4 0.4–4 3* 1.5–3
Chronostratigraphy Depositional environment Species
RW1: primary rachis width; DBP: distance between secondary pinnae; IA2: insertion angle of the secondary pinnae; RW2: secondary rachis width; PL: pinnule length; PW: pinnule width.
localities) and in marginal marine depositional environments (55% of the localities) (Fig. 2(B)). In Europe, in 60% of the localities the depositional environment was continental, whereas 71% of the African localities and 100% of those from North America, were marginal marine (Fig. 4). Weichselia was more common in Europe and in continental environments during the Berriasian-Barremian. However, during the Aptian–Cenomanian, its distribution was slightly reduced, and W. reticulata was notably related to marginal marine depositional environments in Africa (Fig. 4). Weichselia appeared in North America in the Aptian and apparently vanished in South America and India before the Aptian. The last records of Weichselia correspond to the Cenomanian in Europe (Gomez et al., 2002) and the north of Vakhrameev’s African Province (El-Khayal, 1985; Barale and Azar, 2004; El-Saadawi et al., 2016). 3.2. Metric data In Europe, Weichselia showed the greatest variability for all the variables except for the pinnule length. Pinnules are longer and wider in Africa and East Asia. In North America, both rachis width and the insertion angles are the smallest among all the geographic
areas, and the same happens to a lesser extent in South American specimens. In African localities, the minimum values for all variables are seemingly higher than those of Europe (Table 2). The European fossil record is very significant for the Berriasian– Barremian and mostly consists of continental environments. Thus, for these groups, the metric variables showed the greatest variability, except for the insertion angle and the pinnule length and width. Moreover, because Aptian-Cenomanian and marginal marine localities are mostly located in Africa, the minimum values for all variables are seemingly higher than the Berriasian– Barremian and continental values (Table 2). Harris (1981) related the pinnule size to preservation, as pinnules may have shrunk during burning. The measurements taken on specimens from Las Hoyas denote no difference relative to the various types of preservation, including impressions, compressions, and permineralizations with limonite. The variability observed in the three species of Weichselia shows that values of W. negevensis are within the range of those observed for W. reticulata (Table 2). However, the minimum values of W. negevensis for all variables are always higher than those of W. reticulata and W. peruviana, as it occurs throughout the African specimens. W. peruviana is also within the observed variability for W. reticulata (Table 2). Nonetheless, although the values of the
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Fig. 4. Occurrences of the genus Weichselia Stiehler plotted on three Cretaceous palaeomaps at 170 Ma (A), 120 Ma (B), and 105 Ma (C). Star: continental locality; dot: marginal marine locality; square: unknown depositional environment. Palaeomaps based on Martı´n-Abad and Poyato-Ariza (2013).
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insertion angles of the secondary pinnae are within the range of W. reticulata, they are more acute than those of other specimens from the Berriasian–Barremian. The primary rachis width is also particularly small in W. peruviana (Table 2).
4. Discussion A survey of the literature allows for a preliminary scenario on the palaeobiogeographic distribution, the fossil record and most common depositional environment of the fossil fern Weichselia. The gathered information includes studies since 1824. Some references obviously lack precise information on the chronostratigraphy, depositional environment, and metric descriptions, and not all geographic areas were studied with the same intensity (Dunhill et al., 2014). There is also a bias in the direct metric data since the size of the different variables varies along the primary and secondary pinnae, and not all parts of the frond could be measured for all localities. However, the whole data allow for a thorough exploration of the distribution and metric variability of the fossil record of Weichselia. Although more localities were included in our study, the distribution of Weichselia resemble those described by Barnard (1973), El-Khayal (1985), Daber (1990), and Vakhrameev (1991), which already comprised records of this taxon in Africa, North America, South America, Europe, and India. The unique exception is East Asia, whose fossil record was published from 1975 onwards and included in Daber (1990) and Vakhrameev (1991). When the geographic distribution and the chronostratigraphy are crossed, a pattern of Weichselia dispersion emerges. Most leptosporangiate fern families have similar distributions and dispersion patterns. Most of them appeared in subtemperate humid areas, at latitudes higher than 308 N and S, and then later spread towards lower latitudes (Skog, 2001). However, our results show that the leptosporangiate family Weichseliaceae appeared near the Equator at latitudes of 0–208 N during the Bathonian–Tithonian. Weichselia dispersed very quickly both northward and southward, and reached latitudes of 508 N and S by the Berriasian–Barremian. At the end of the Barremian, its fossil record is confined to latitudes of 408 N–208 S, and declines during the Aptian–Cenomanian interval. The distribution of Weichselia is more consistent with that of the fern savannah biome, which existed during the Jurassic and Cretaceous. This biome lost importance in the Aptian, probably due to the increase in humidity during this period (Coiffard et al., 2007). Global dispersion patterns have not yet been studied for many taxa. One of the few groups that have been thoroughly studied are charophytes, and although not closely related to Weichselia, the pattern of dispersal of this taxon in the Northern Hemisphere is parallel to that of the charophytes Atopocara trivolvis (Martı´nClosas and Wang, 2008) and Clavator harrisii (Martı´n-Closas, 2015), particularly for the late colonization of North America, which occurred during the Aptian. The fact that charophytes and Weichselia follow the same pattern would suggest that the barriers for colonization prior to the Aptian might have been common to these groups, even though their ecological preferences are very different, and the localities where they have been collected differ in most cases. Our analysis of metric data also offers a new perspective as the metric differences are only observed for the pinnule size, and the variability of this metric is apparently associated with the chronostratigraphy. The length of Weichselia pinnules is larger in the Aptian-Cenomanian localities of Africa, East Asia, and Europe. The specimens collected and measured from the Albian localities of the Escucha Formation in NE Spain (Sender, 2012) show larger pinnules (i.e., 6–8 mm long), and differ in size with
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those collected from other European localities (0.5–7 mm long), which are all Berriasian–Barremian in age. Nevertheless, it should be noted that all localities reported so far from Africa and Eastern Asia are Aptian–Cenomanian in age. Therefore, this assessment requires further measurements in Berriasian–Barremian localities around the world. The size differences found in Weichselia might be related to a hothouse event with a rise in humidity, as suggested by Kidder and Worsley (2012) for the Aptian. This climatic shift might have positively affected the pinnule growth of Weichselia, because drier, nutrient-poor or high-irradiance conditions produce smaller pinnules in living ferns (Arens, 1997; Creese et al., 2011). The depositional environment does not seem to play a role in the differences in Weichselia frond size. The pinnules described from the marginal marine depositional environments of Preznosza and Lipnik (Reymano´wna, 1965) and Beauvais, are similar in size (5 mm long and 2–5 mm long, respectively) to the pinnules described in other continental Berriasian–Barremian localities. In most localities, Weichselia might have been parautochthonous because only fragments of fragile primary pinnae and disarticulated secondary pinnae preserved as charcoals have been collected (Gomez et al., 2001). Thus, the expected bias due to transportation and posterior burial in an environment where the fern did not grow is not so extreme. However, a more in-depth study of the taphonomy of each site must be conducted in order to fully understand the implications of the depositional environment on frond size. Finally, the metric variability among the species of Weichselia appears within the ranges obtained for W. reticulata for all variables. The min-max ranges for W. reticulata have been extended when direct measurements on the specimens have been performed. In addition, the differences found between the three species could probably be explained by the heterogeneity of the parts collected in each locality (e.g., W. peruviana is based on data from the apical part of the primary pinna) and/or chronostratigraphy (W. negevensis is only Aptian–Cenomanian in age). Acknowledgements We thank three anonymous reviewers for their comments which helped us to improve the original manuscript. We thank Santiago Langreo and Mercedes Llandres from the Museo de Paleontologı´a de Castilla-La Mancha, Cuenca, Spain, for enabling the study of the specimens from Las Hoyas, and Emmanuel Robert for granting us access to the material from Peru´ housed in the palaeontological collections at the Universite´ Lyon 1, Villeurbanne, France. Candela Blanco-Moreno is supported by a FPI-UAM Ph.D. scholarship from the Universidad Auto´noma de Madrid. Collections in the Universite´ Lyon 1 were studied thanks to a FPI-UAM travel grant provided by the same university. Collections in the Muse´um national d’Histoire naturelle, Paris; Natural History Museum, London; and Royal Belgian Institute of Natural Sciences, Brussels, were studied with the support of three grants from the SYNTHESYS Project (http://www.synthesys.info/), financed by the European Community Research Infrastructure Action under the FP7 Integrating Activities Program. We thank the staff from these three institutions for their assistance when accessing the collections. This work is part of the project CGL-2013-42643 P of the Spanish Ministry of Economy and Competitiveness and other projects.
Appendix A. Supplementary data Supplementary data (including the list of examined material [A], the list of localities used for the palaeobiogeographic analysis [B], depositional environment criteria [C], and details of the metric
Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001
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Please cite this article in press as: Blanco-Moreno, C., et al., Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia. Geobios (2018), https://doi.org/10.1016/j.geobios.2018.05.001