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Review of the Bennettitalean Genus Weltrichia Mihai Emilian Popa

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Review of the Bennettitalean Genus Weltrichia Mihai Emilian Popa Popa Journal of Palaeogeography (2019) 8:12 https://doi.org/10.1186/s42501-019-0023-9 Journal of Palaeogeography REVIEW Open Access Review of the bennettitalean genus Weltrichia Mihai Emilian Popa Abstract The bennettitalean male reproductive structure Weltrichia Braun 1849 emend. Harris 1969 (Family Williamsoniaceae) is discussed from several points of view, including anatomy and morphology, species diversity and validity, phytogeographical and stratigraphical distribution, and reproduction strategy. A very rare fossil, genus Weltrichia includes 25 valid species distributed in both hemispheres during Triassic and Jurassic times. This contribution critically reviews the systematics of the entire genus, together with its stratigraphical and palaeophytogeographical significance. Keywords: Bennettitales, Weltrichia, Male reproductive structure, Triassic, Jurassic 1 Introduction Cretaceous representatives. The Williamsoniaceae in- The Order Bennettitales Engler 1892 represents a highly cludes plants with a slender stem and unisexual and interesting group of extinct gymnosperms in terms of bisexual reproductive structures, such as the unisex- anatomy, high biodiversity and widespread phytogeograph- ual, male Weltrichia Braun 1849 emend Harris 1969, ical distribution during the whole Mesozoic (Stewart 1983; the female Williamsonia Carruthers 1870 or the bi- Taylor et al. 2009; Friis et al. 2011). In the same time, their sexual Williamsoniella Thomas 1915. evolutionary significance is related to the origin of angio- Weltrichia consists basically of a central cup to which sperms, as the Bennettitales innovated radial, complex, radially equal, centrifugal rays are attached, bearing bi- unisexual or bisexual reproductive structures resembling valve pollen sacs (also called synangia) with elliptical, the angiosperm flowers. Such reproductive structures are monocolpate pollen, the whole structure having a radial true gymnospermous cones and they represent a conver- symmetry. The substance of the cup and rays is usually gent evolutionary trend with true flowers, an experiment thick, and the rays occur in a single whorl. Variations to which the Bennettitales undertook before the first flower- this basic morphology include equal centripetal rays over ing plants originated (Friis et al. 2011; Pott 2014). The the central cup, attractants or resin (resinous) bodies reproductive structures resembling flowers were not the within the cup, unipinnate (pedicellate) pollen sacs, only peculiar characters of Bennettitales, they also pos- trichomes, appendages, striae and ridges over the centri- sessed syndetocheilic stomata in their epidermal tissues. fugal rays. Weltrichia is associated with Williamsonia These types of stomata separate them from morphologic- female reproductive structures, with foliage belonging to ally similar vegetative organs of plants belonging to the the genera Zamites, Otozamites and Ptilophyllum and Order Cycadales, among other characters. Bennettitales with other vegetative parts such as Bucklandia cortical are related to Gnetales and angiosperms in cladistic stud- fragments (Harris 1969). ies (Doyle 2006; Hilton and Bateman 2006). They are Genus Weltrichia was defined by Braun (1849), with divided into two separate families, based on vegetative and the type species Weltrichia mirabilis from Lower reproductive anatomy: Family Williamsoniaceae, domin- Jurassic sediments of Franconia, Germany. He also antly with Late Triassic and Jurassic representatives, and defined two more species of Weltrichia in the same Family Cycadeoidaceae (Bennettitaceae), with mainly paper, synonyms of W. mirabilis.Thenewtaxonwas dedicated to Johann P. A. Weltrich, who collected the material,anditwasconsideredasbelongingtothe Correspondence: [email protected] Faculty of Geology and Geophysics, Department of Geology and Doctoral Family Rafflesiaceae (Braun 1849). Initially, the Weltri- School of Geology, Laboratory of Palaeontology, University of Bucharest, 1, N. chia denomination did not reach a wide use, as such Bălcescu Ave, 010041 Bucharest, Romania © The Author(s). 2019 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Popa Journal of Palaeogeography (2019) 8:12 Page 2 of 21 structures were often referred as “male Williamsonias”, Since Braun (1849), Weltrichia species were described in relation with and resembling the female bennettita- from Franconia, Germany (Schuster 1911), Banat, lean reproductive structures Williamsonia Carruthers Romania (Krasser 1915, 1922; Humml 1969; Givulescu 1870. Krasser (1912, 1913, 1915, 1919, 1922), Nathorst 1990, 2001, 2014), Yorkshire, United Kingdom (Nathorst (1909, 1911a, 1912)andSharma(1969)usedtheterm 1909, 1911a, 1911b, 1912, 1913; Florin 1933; Harris “male Williamsonia” for various Weltrichia species, 1969), Tirol, Germany (Krasser 1919), Veneto and while Schuster (1911) and Nathorst (1911b)used Sardinia, Italy (Saporta 1891; Krasser 1912), France Weltrichia.Saporta(1891)usedWeltrichia in his contri- (Saporta 1891), Alborz Mountains, Iran (Schweitzer and bution on Jurassic plants of France, erecting two new spe- Kirchner 2003), Rajmahal, Malda and Katchchh, India cies, W. fabrei from France and W. oolithica from Italy. (Sharma 1969, 1971, 1977; Sitholey and Bose 1971; Bose Discussion over the relationship between Weltrichia and and Banerji 1984; Pal and Ghosh 1985; Banerji 1992), Williamsonia and on their nomenclature was continued Malaysia (Asama 1974), Nepal (Kimura et al. 1985), by Harris (1969), Sitholey and Bose (1971), Schweitzer China (Sun et al. 2001; Li et al. 2004), Japan (Kimura and Kirchner (2003) and Popa (2014), next to emenda- and Ohana 1989) and Oaxaca, Mexico (Delevoryas tions of previous species or with definitions of new and Gould 1973; Delevoryas 1991;SilvaPinedaetal. species. As Weltrichia has priority over Williamsonia,it 2011; Velasco de Leon et al. 2012). The distribution was kept by Harris (1969) as a legitimate denomination of Weltrichia species in the world is illustrated in for male, unisexual structures, keeping Williamsonia only Fig. 1. for female, unisexual structures. Harris (1969)emended The stratigraphic ranges of Weltrichia species vary the diagnosis of the genus, supplementing it with micro- from Late Triassic to Late Jurassic, from the earliest rep- scopical characters, noting the high variation in attaching resentative Weltrichia alpina, Late Triassic in age, to its the pollen sacs to the centrifugal rays, directly or on sup- latest representatives, W. maldaensis and W. huangban- plementary, sterile appendages. Harris refrained from jigouensis, Late Jurassic in age. splitting the genus and erecting new genera based on this variation of characters, for practical reasons. Popa (2014) briefly reviewed the valid species of Weltrichia and their 2 Material and methods ranges when introducing new species of this genus, The Romanian material belonging to genus Weltrichia stressing the most important characters in Weltrichia: has been surveyed and described in a previous work shape and number of centrifugal rays, ornamentation and (Popa 2014), curated at the University of Bucharest and at shape of centripetal rays, distribution and types of pollen University of Petroșani. Previous palaeobotanical literature sacs, ornamentation of the central cup, and occurrence of was carefully surveyed and large sized differential tables resin bodies. were synthesized. I have produced the reconstructions Fig. 1 Geographical distribution of genus Weltrichia. 1. Oaxaca State (Mexico); 2. Yorkshire (United Kingdom); 3. Tirol, Franconia (Germany), Veneto (Italy), Lozére (France); 4. Sardinia (Italy); 5. Anina (Romania); 6. Alborz (Iran); 7. Kachchh (India); 8. Rajmahal and Malda (India); 9. Palpa (Nepal); 10. Ulu Endau (Malaysia); 11. Liaoning (China); 12. Chifeng (Inner Mongolia Autonomous Region, China); 13. Samenoura (Japan) Popa Journal of Palaeogeography (2019) 8:12 Page 3 of 21 manually, using a protractor, a ruler, a pair of com- 3 Terminology passes and soft lead 0.5 mm mechanical pencils (2B), The terminology used in Weltrichia parts is very variable as well as normal and semi-transparent paper for and sometimes confusing. Harris (1969) used the terms continuously refining the drawings. The numbers of central cup, rays or microsporophylls and pollen sacs for centrifugal and centripetal rays were considered, in the main parts of Weltrichia, as well as resin or resinous order to divide angles and circles accordingly, while bodies (sometimes also called resinous sacs), sterile scales, ray lengths, widths and diameters of central cups and fertile appendages (Fig. 2). The preference for pollen were reflected in reconstructions, for correct dimen- sacs instead of synangia was explained for the gymno- sional ratios in drawings. All dimensions and pre- spermous nature of Weltrichia (Harris 1969,p.159),in served anatomical and morphological characters were contrast with the more pteridophytic synangia. Although considered in order to produce accurate reconstruc- the term synangia is also correctly used for these tions. Drawings with adaxial and abaxial perspectives structures (Pott et al. 2017), pollen sacs being basically were produced, depending
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