Forgotten women of Paleontology: Margaret Benson

Margaret Jane Benson. Portrait in the Archives of Royal Holloway, University of London (RHC PH/282/13) From Fraser & Cleal, 2007

It is a truth universally acknowledged, that women has always work harder than men to gain some recognition. It was true in the 16th, and it’s true now. In “A Room of One’s Own”, Virginia Woolf explores the conflicts that a gifted woman must have felt during the Renaissance through the fictional character of Judith Shakespeare, the sister of William Shakespeare, and cites as obstacles the indifference of most of the world, the profusion of distractions, and the heaping up of various forms of discouragement. But not only in the Elizabethan times. In the Victorian times there was the common assumption that the female brain was too fragile to cope with mathematics, or science in general. In a letter from March 1860, Thomas Henry Huxley wrote to great geologist Charles Lyell FRS: “Five-sixths of women will stop in the doll stage of evolution, to be the stronghold of parsonism, the drag on civilisation, the degradation of every important pursuit in which they mix themselves – intrigues in politics and friponnes in science.”

Margaret Crosfield on a Geologists’ Association fieldtrip to Leith Hill with Professor Lapworth (From Burek and Malpas, 2007).

Women have played  various and extensive roles in the history of geology. Unfortunately, their contribution has not been widely recognised by the public or academic researchers. In the 18th and 19th centuries women’s access to science was limited, and science was usually a ‘hobby’ for intelligent wealthy women. Early female scientists were often born into influential families, like Grace Milne, the eldest child of Louis Falconer and sister of the eminent botanist and palaeontologist, Hugh Falconer; or Mary Lyell, the daughter of the geologist Leonard Horner. They collected fossils and mineral specimens, and were allowed to attend scientific lectures, but they were barred from membership in scientific societies. But by the first half of the 20th century, a third of British palaeobotanists working on Carboniferous plants were women. The most notable were  Margaret Benson, Emily Dix, and Marie Stopes.

Newnham began as a house for five students in Regent Street in Cambridge in 1871

Margaret Benson was born on the 20th October 1859 in London. Between 1878 and 1879, she studied at Newnham College Cambridge. After obtaining her BSc at University College London (UCL) in 1891, she started research on plant embryology.  In 1893, Benson was appointed head of the new Department of Botany at Royal Holloway College, the first woman in the United Kingdom to hold such a senior position in the field of botany. Her palaeobotanical research centred on the anatomy of reproductive structures, especially of Carboniferous pteridosperms and lycophytes. In 1904, she was among the first group of women to be elected as Fellows of the Linnean Society, and in 1912 she was appointed Professor of Botany at the University of London. Her major study on lycophyte fructifications was on the cones of the Sigillaria plant. She also speculated on the relationship between the Palaeozoic arborescent lycophytes and the Recent Isoetes, with the Triassic Pleuromeia as a possible intermediate form. She worked with ferns and cordaites and described a new species, Cordaites felicis. Benson’s work is characterized by careful description. One of her most important theoretical works concerns the phylogenetic significance of the sporangiophore in lycophytes, sphenophytes and ferns. After her retirement in 1922, she was encouraged by D. H. Scott to write up some of her earlier unpublished work on the root anatomy of the early Carboniferous pteridosperm Heterangium. She even continued with fieldwork when she was in her 70s. There is an unpublished manuscript in which she described a new fertile Rhacopteris that she collected from Teilia Quarry in North Wales in 1933. She died on 20th June 1936 at Highgate, Middlesex.


H. E. Fraser and C. J. Cleal, The contribution of British women to Carboniferous palaeobotany during the first half of the 20th century, Geological Society, London, Special Publications, 281, 51-82, 1 January 2007,

C. V. Burek (2007). The role of women in geological higher education – Bedford College, London (Catherine Raisin) and Newnham College, Cambridge, UK, Geological Society, London, Special Publications, eds Burek C. V., Higgs B. 281, pp 9–38



Molecular signatures of fossil leaves

Leaves of Ptilophyllum mueller, from Emmaville, New South Wales. Scale bars=10 mm (From McLoughlin et al., 2011)

The first plants colonized land approximately 450 million years ago. The transition from an exclusively aquatic to a terrestrial life style implied the evolution of a new set of morphological and physiological features. The most critical adaptive trait for survival during terrestrialization was the ability to retain water in increasingly dehydrating habitats. Consequently, the capacity to maintain a hydrophobic surface layer, or cuticle, over the surfaces of aerial organs was arguably one of the most important innovations in the history of plant evolution.

Spores, pollen and leaf cuticles, are among the most resilient organic structures in the geological record. These components may retain some phylogenetically unique signals, not only in well-preserved fossils, but also in remains with a high level of diagenetic maturity.

Ginkgo biloba, Eocene fossil leaf from the Tranquille Shale of MacAbee, British Columbia, Canada (From Wikipedia Commons)

Generally, the cuticle is divided into two major structural constituents: cutin and cutan. The fatty acid polyesters which constitute cutin, gives the cuticle considerable resistance to biodegradation. Cutan is a non-ester and non-hydrolyzable matrix of aliphatic compounds linked by ether bonds, which remain after cutin hydrolysis. Additionally, the surface of the cuticle may be covered by various long-chain hydrophobic waxes. All these components  favours the survival of the cuticle in many fossil plants, and can be used to resolve the stratigraphic ranges and relationships of extinct plants.

Data from infrared spectroscopy of modern plant cuticles, have been used successfully to support and clarify the species-level taxonomy of extant plants, for example, in Camellia and angiosperm pollen. Using infrared spectroscopy and statistical analysis, researchers at Lund University, the Swedish Museum of Natural History in Stockholm, and Vilnius University, analysed a selection of fossil Cycadales, Ginkgoales and conifers. The team obtained two major groups in the dendrogram of infrared spectra. One branch unites podocarpacean and araucariacean conifers (excluding the Jurassic Allocladus). A relationship consistent with all modern phylogenetic analyses of gymnosperm. The second branch unites a broad range of gymnosperms. Within this branch, Bennettitales (Otozamites and Pterophyllum) form a well-defined group in association with Ptilozamites and Nilssoniales. This cluster is linked to a group incorporating Cycadales on one sub-branch, and Leptostrobales, Ginkgoales and the putative araucariacean Jurassic conifer Allocladus on a second sub-branch.


Dendrogram based on HCA of infrared absorption spectra of an expanded group of 13 fossil gymnosperm taxa (From Vajda et al., 2017)

Early palaeobotanical studies generally linked Bennettitales to Cycadales, but more recent anatomical studies and cladistic analyses have indicated that Bennettitales are not closely related to Cycadales. By contrast, Bennettitales, Nilssoniales and Ptilozamites are grouped closely. Additionally,  the systematic position of Allocladus within Araucariaceae should be reassessed based on its close association with Ginkgo in the cluster analysis of infrared spectra.


Vivi Vajda, Milda Pucetaite, Stephen McLoughlin, Anders Engdahl, Jimmy Heimdal, Per Uvdal. Molecular signatures of fossil leaves provide unexpected new evidence for extinct plant relationships. Nature Ecology & Evolution, 2017; DOI: 10.1038/s41559-017-0224-5

Stephen McLoughlinRaymond J. Carpenter, and Christian Pott, Ptilophyllum muelleri (Ettingsh.) comb. nov. from the Oligocene of Australia: Last of the Bennettitales?, International Journal of Plant Sciences 2011 172:4574-585, DOI: 10.1086/658920

Terrestrial floras at the Triassic-Jurassic Boundary in Europe.

Proportions of range-through diversities of higher taxonomic categories of microfloral elements over the Middle Triassic–Early Jurassic interval (From Barbacka et al., 2017)

Over the last 3 decades, mass extinction events  have become the subject of increasingly detailed and multidisciplinary investigations. Most of those events are associated with global warming and proximal killers such as marine anoxia. Volcanogenic-atmospheric kill mechanisms include ocean acidification, toxic metal poisoning, acid rain, increased UV-B radiation, volcanic darkness, cooling and photosynthetic shutdown. The mass extinction at the Triassic-Jurassic Boundary (TJB) has been linked to the eruption of the Central Atlantic Magmatic Province (CAMP), a large igneous province emplaced during the initial rifting of Pangea. Another theory is that a huge impact was the trigger of the extinction event. At least two craters impact were reported by the end of the Triassic. The Manicouagan Impact crater in the Côte-Nord region of Québec, Canada was caused by the impact of a 5km diameter asteroid, and it was suggested that could be part of a multiple impact event which also formed the Rochechouart crater in France, Saint Martin crater in Canada, Obolon crater in Ukraine, and the Red Wing crater in USA (Spray et al., 1998).

Photographs of some Rhaetian–Hettangian spores and pollen from the Danish Basin (From Lindström, 2015)

Most mammal-like reptiles and large amphibians disappeared, as well as early dinosaur groups. In the oceans, this event eliminated conodonts and nearly annihilated corals, ammonites, brachiopods and bivalves. In the Southern Hemisphere, the vegetation turnover consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one. But there was no mass extinction of European terrestrial plants during the TJB. The majority of genera and a high percentage of species still existed in its later stages, and replacement seems to have been local, explainable as a typical reaction to an environmental disturbance. In Greenland, for example, the replacement of Triassic wide-leaved forms with Jurassic narrow-leaved forms was linked to the reaction of plants to increased wildfire. In Sweden, wildfire in the late Rhaetian and early Hettangian caused large-scale burning of conifer forests and ferns, and the appearance of new swampy vegetation. In Austria and the United Kingdom, conifers and seed ferns were replaced by ferns, club mosses and liverworts. In Hungary, there was a high spike of ferns and conifers at the TJB, followed by a sudden decrease in the number of ferns along with an increasing share of swamp-inhabiting conifers.

Although certain taxa/families indeed became extinct by the end of the Triassic (e.g. Peltaspermales), the floral changes across Europe were rather a consequence of local changes in topography.


Maria Barbacka, Grzegorz Pacyna, Ádam T. Kocsis, Agata Jarzynka, Jadwiga Ziaja, Emese Bodor , Changes in terrestrial floras at the TriassicJurassic Boundary in Europe, Palaeogeography, Palaeoclimatology, Palaeoecology (2017), doi: 10.1016/j.palaeo.2017.05.024

S. Lindström, Palynofloral patterns of terrestrial ecosystem change during the end-Triassic event — a review, Geol. Mag., 1–23 (2015)

Van de Schootbrugge, B., Quan, T.M., Lindström, S., Püttmann, W., Heunisch, C., Pross, J., Fiebig, J., Petschick, R., Röhling, H.-G., Richoz, S., Rosenthal, Y., Falkowski, P. G., 2009. Floral changes across the Triassic/Jurassic boundary linked to flood basalt volcanism. Nat. Geosci. 2, 589–594. doi: 10.1038/NGEO577.

N.R. Bonis, W.M. Kürschner, Vegetation history, diversity patterns, and climate change across the Triassic/Jurassic boundary, Paleobiology, 8 (2) (2012), pp. 240–264

Forgotten women of Paleontology: Emily Dix


Dr Emily Dix and her assistant Miss Elsie White.

Dr Emily Dix and her assistant Miss Elsie White.

In the 18th and 19th centuries women’s access to science was limited, and science was usually a ‘hobby’ for intelligent wealthy women. It was common for male scientists to have women assistants, often their own wives and daughters. But by the first half of the 20th century, a third of British palaeobotanists working on Carboniferous plants were women. The most notable were  Margaret Benson, Emily Dix, and Marie Stopes.

Emily Dix was born on 21 May 1904 in Penclawdd, in the beautiful area of the Gower Peninsula. At age 18, she gained the Central Welsh Board Higher Certificate in history, botany and geography, with distinctions in both history and botany. In 1925, she graduated with first class honours in Geology at the University College Swansea. After graduation, Emily continued at Swansea to research the geology of the western part of the South Wales Coalfield. Her work was supervised by Arthur E. Trueman, Professor of Geology at Swansea, and a pioneer in developing stratigraphical theory. Trueman realized that the only accurate way to use fossils for correlation was to divide the stratigraphical succession into biozones defined exclusively by the assemblages of species present, independently of the lithology in which they were found. Trueman’s main interest were  non-marine bivalves, so Emily’s early work was on the non-marine bivalves of the South Wales Coalfield.

Emily Dix during the 2nd International Carboniferous Congress in 1935 (From Burek and Cleal, 2005)

Emily Dix during the 2nd International Carboniferous Congress in 1935 (From Burek and Cleal, 2005)

Emily initially studied all aspects of the Late Carboniferous biotas in South Wales, but soon, she realized that plant fossils also had considerable biostratigraphical potential. Although, Paul Bertrand developed macrofloral biozones for the French coalfields in 1914, Emily used stratigraphical range charts for the first time in paleobotany recognising the need to separate biostratigraphy from lithostratigraphy. In 1926 Emily was awarded an MSc based on her Gwendraeth Valley work: ‘The Palaeontology of the Lower Coal Series of Carmarthen and the Correlation of the Coal Measures in the Western Portion of the South Wales Coalfield’.  In 1929 she was elected a fellow of the Geological Society, and a year later she was appointed Lecturer in Palaeontology at Bedford College in London, a position that she held for the rest of her working life. The same year, she attended the International Botanical Congress in Cambridge where she met W. Gothan, P. Bertrand, W. J. Jongmans and A. Renier, leaders in Upper Carboniferous palaeobotany at the time. Five years later, she attended the Second International Carboniferous Congress in Heerlen (The Netherlands) and she delivered papers on Carboniferous biostratigraphy. In 1936, Emily was invited to become the only female on an 11-man discussion group of the British Association for the Advancement of Science on Coal Measures correlation. She was clearly at the international forefront of the field.

Emily Dix in the Auvergne 1936 (seated fourth from right, see white arrow). From Burek and Cleal, 2005.

Emily Dix in the Auvergne 1936 (seated fourth from right, see white arrow). From Burek and Cleal, 2005.

At the start of the World War II, she was evacuated to Cambridge, along with the rest of Bedford College Geology Department. She lost a lot of valuable literature and other records in a London Blitz in May 1941. Fortunately, much of her collection of fossils survived.

At the end of the war, Emily suffered a mental breakdown. She was moved to a mental hospital run by the Quakers in the City of York. That was the end of her scientific career. She died in Swansea on 31 December 1972.

It was not until the late 1970s that her techniques were used again in Britain. Elsewhere in Europe, her approaches were adopted and can be seen in many of the papers presented at the International Carboniferous Congresses held at Heerlen during the 1950s and early 1960s.


Burek, C. V. & Cleal, C. J. (2005) The life and work of Emily Dix (1904-1972). In: Bowden, A. J., Burek, C. V. & Wilding, R (ed.) History of palaeobotany: selected essays. Geological Society of London, Special Publication, 241, 181-196

Burek, C. V. (2005). Emily Dix, palaeobotanist – a promising career cut short. Geology Today, 21(4), 144-145

A palaeobotanical perspective on the Permian extinction.


Leaf bank of Glossopteris leaves (Adapted from Mcloughlin, 2012)

Leaf bank of Glossopteris leaves (Adapted from Mcloughlin, 2012)

The fossil record indicates that more than 95% of all species that ever lived are now extinct. Occasionally, extinction events reach a global scale with many species of all ecological types dying out in a near geological instant. These are mass extinctions. They were originally identified in the marine fossil record and have been interpreted as a result of catastrophic events or major environmental changes that occurred too rapidly for organisms to adapt. Mass extinctions are probably due to a set of different possible causes like basaltic super-eruptions, impacts of asteroids, global climate changes, or continental drift. A central question in the understanding of mass extinctions is whether the extinction was a sudden or gradual event. This question may be addressed by examining the pattern of last occurrences of fossil species in a stratigraphic section.

Jack Sepkoski and David M. Raup identified five major extinction events in Earth’s history: at the end of the Ordovician period, Late Devonian, End Permian, End Triassic and the End Cretaceous. The most recently identified mass extinction occurred during the Middle Permian, about  262 million years ago, and it was first recognised in the marine realm as a turnover among foraminifera, with fusulinaceans among the principal casualties.

Sin título

Total diversity patterns of continental diversity (solid line) and marine diversity (dotted line) at the family level. Arrows indicate the mass extinction events. (From Cascales-Miñana and Cleal 2015)

Extinction dynamics in the marine and terrestrial biotas followed different trajectories, and only the Permo-Triassic event coincided with a clear and abrupt diminution of both realms. Moreover, analysis of the paleobotanical record has suggested that plants may have suffered an additional extinction event, that is not reflected significantly in the marine realm, at the Carboniferous–Permian boundary. Evidence also suggests that  terrestrial environments suffered a single global pulse of extinction in the latest Permian, affecting both the fauna and flora (Cascales-Miñana and Cleal 2015).

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. The palynological record suggests that wooded terrestrial ecosystems took four to five million years to reform stable ecosystems, while spore-producing lycopsids had an important ecological role in the post-extinction interval. A key factor for plant resilience is the time-scale: if the duration of the ecological disruption did not exceed that of the viability of seeds and spores, those plant taxa have the potential to recover (Traverse, 1988).



Borja Cascales-Miñana, José B. Diez, Philippe Gerrienne & Christopher J.Cleal (2015): A palaeobotanical perspective on the great end-Permian biotic crisis, HistoricalBiology, DOI: 10.1080/08912963.2015.1103237

Aberhan M. 2014. Mass extinctions: ecological diversity maintained. NatGeosci. 7:171–172.

Cascales-Miñana B, Cleal CJ. 2014. The plant fossil record reflects just two great extinction events. Terra Nova. 26(3):195–200.

Darwin and the flowering plant evolution in South America.


Retimonocolpites sp. (Adapted from Llorens and Loinaze, 2015)

Charles Darwin’s fascination and frustration with the evolutionary events associated with the origin and early radiation of flowering plants are legendary. In a letter to Oswald Heer, a famous Swiss botanist, and paleontologist, Darwin wrote: “the sudden appearance of so many Dicotyledons in the Upper Chalk appears to me a most perplexing phenomenon to all who believe in any form of evolution, especially to those who believe in extremely gradual evolution, to which view I know that you are strongly opposed”. Heer discussed about the early angiosperm fossil record with Darwin, in a letter dated 1 March 1875: “if we say that the Dicotyledons begin with the Upper Cretaceous, we must still concede that this section of the vegetable kingdom, which forms the bulk of modern vegetation, appears relatively late and that, in geological terms, it underwent a substantial transformation within a brief period of time.” 

Darwin’s defense of a gradualist perspective led him to suggest that prior to the Cretaceous record of flowering plants, angiosperms had slowly evolved and diversified on a remote landmass. On 22 July 1879, in a letter to Joseph Dalton Hooker, Darwin refers to the early evolution of flowering plants as an “abominable mystery”. Nearing the end of his life, he wrote to Hooker another letter about a lost fossil record in the earliest phases of angiosperm diversification:  “Nothing is more extraordinary in the history of the Vegetable Kingdom, as it seems to me, than the apparently very sudden or abrupt development of the higher plants. I have sometimes speculated whether there did not exist somewhere during long ages an extremely isolated continent, perhaps near the South Pole.”

Letter from Charles Darwin to Joseph Dalton Hooker, written 22 July 1879 (provenance: Cambridge University Library DAR 95: 485–488)

Letter from Charles Darwin to Joseph Dalton Hooker, written 22 July 1879 (provenance: Cambridge University Library DAR 95: 485–488)

While the oldest records of the different groups of angiosperms are still in discussion, the outcrops of the Baquero Group, located in Argentinean Patagonia, contain one of the richest and most diverse Early Cretaceous floras in the Southern Hemisphere. The unit comprises three formations: Anfiteatro de Tico, Bajo Tigre and Punta del Barco. The first reports of angiosperm remains for the Anfiteatro de Tico Formation were made in 1967. The dominant types are Clavatipollenites, and Retimonocolpites.

Pollen grains  could enter into the fossil record by falling directly into swamps or lakes, or being washed into them or into the rivers and seas. The ones which are not buried in reducing sediments will tend to become oxidized and be destroyed. They reflects the ecology of their parent plants and their habitats and provide a continuous record of their evolutionary history. Gymnosperms pollen often is saccate (grains with two or three air sacs attached to the central body), while Angiosperm pollen shows more variation and covers a multitude of combinations of features: they could be  in groups of four (tetrads),  in pairs (dyads),  or single (monads). The individual grains can be inaperturate, or have one or more pores, or slit-like apertures or colpi (monocolpate, tricolpate).


Clavatipollenites sp. SEM (Adapted from Archangelsky 2013)

Clavatipollenites pollen grains are interpreted as related to the modern family Chloranthaceae. The genus was established by Couper for dispersed monosulcate pollen grains recovered from the Early Cretaceous of Britain. Currently, the genus has a very broad definition. The genus Retimonocolpites include elongated to subcircular semitectate, columellate and microreticulate pollen grains with well defined monocolpate aperture (Llorens and Loinaze, 2015). The new species Jusinghipollisticoensis sp. nov. represents one of the oldest records of trichotomosulcate, and extends the geographical distribution of Early Cretaceous trichotomosulcate pollen grains to southern South America.

The data also indicates strong similarities between the Baquero Group assemblages and other coeval units from Argentina, Australia and United States.


M. Llorens, V.S. Perez Loinaze, Late Aptian angiosperm pollen grains from Patagonia: Earliest steps in flowering plant evolution at middle latitudes in southern South America, Cretaceous Research 57 (2016) 66-78

Archangelsky, S.,et al. (2009). Early angiosperm diversification: evidence from southern South America. Cretaceous Research, 30, 1073-1082.

Doyle, J. A., & Endress, P. K. (2014). Integrating Early Cretaceous fossils into the phylogeny of living Angiosperms: ANITA. Lines and relatives of Chloranthaceae. International Journal of Plant Sciences, 175, 555-560.

Marie Stopes and her legacy as paleobotanist.

Marie Stopes (1880-1958)  by George Bernard Shaw. (LSE Archives Image Record, 1921)

Marie Stopes (1880-1958) photographed by George Bernard Shaw. (LSE Archives Image Record, 1921).

George Bernard Shaw once wrote: “All progress is initiated by challenging current conceptions”. Those words describe exactly what Marie Stopes did during her entire life. Her book,  Married Love (1918), is considered one of the most influential of the 20th century. A dedicated feminist, her views on birth control and contraception drew to her the hostile attentions of conservative forces in British Society, and Stope’s ideas were attacked through an assault of her own character. But in addition to her well-known work on birth control and women’s rights, she was a prolific poet, playwright and a paleobotanist.

Marie Charlotte Carmichael Stopes was born in Edinburgh, Scotland, on October 15, 1880. Her father, Henry Stopes, a brewer, architect and amateur paleontologist and archeologist, amassed the largest private collection of fossils and ancient stone tools in Britain. Her mother, Charlotte Carmichael, wrote British Freewomen: Their Historical Privilege. The book, published in 1894, was a great influence in the early twentieth century British women’s suffrage movement. They were both members of the British Association for the Advancement of Science.

Marie Stopes (From Wikimedia Commons)

Marie Stopes (From Wikimedia Commons)

Just before her twentieth birthday, Marie enrolled at University College London where she studied botany and geology. She graduated with honours after only two years and received the Gold Medal in Botany. At UCL she was employed by Francis Wall Oliver as a postgraduate research assistant on his pteridosperm project with Dukinfield Henry Scott. Shortly after, she went to study at the University of Munich, and received a Ph.D. in palaeobotany in 1904. She was the only female student among 5.000 men. During that time, she worked on the internal anatomy of cycad seed. In August 1904, Marie got her first academic job at the Victoria University of Manchester. She became more interested in Carboniferous coal balls. These concretions of calcite, dolomite, siderite, and pyrite, occur at many localities in northern England and preserved in beautiful anatomical detail the structure of the plants that formed the coal.

In 1907, she convinced the Royal Society to fund an excursion to Japan. During her work, she found what were then the earliest known flowers and fossil insects from the Cretaceous period. In 1910, she was commissioned by the Geological Survey of Canada to determine the age of the Fern Ledges, a geological structure at Saint John, New Brunswick. She proved that the rocks were Carboniferous and not Devonian or Silurian as others had earlier argued (Falcon-Lang, 2008).

In 1957 Marie Stopes was diagnosed with cancer. She died on October 2, 1958.


FALCON-LANG, H.J. & MILLER, R.F. 2007. Marie Stopes and the Fern Ledges of Saint John, New Brunswick. In Burek, C.V. (ed.) The Role of Women in the History of Geology. Special

Falcon-Lang, H.J., 2008. Marie Stopes: Passionate about Palaeobotany. Geology Today, 24: 132-136.

William Garrett, 2007,  Marie Stopes: Feminist, Eroticist, Eugenicist,

Stephanie Green (2013). The Public Lives of Charlotte and Marie Stopes. London: Pickering & Chatto.


A Permian lagerstätte from Antarctica.


Vertebraria solid-stele and polyarch roots colonised by fungal spores (From Slater et al., 2014)

Vertebraria solid-stele and polyarch roots colonised by fungal spores (From Slater et al., 2014)

A lagerstätte (German for ‘storage place’) is a site exhibiting an extraordinary preservation of life forms from a particular era. The term was originally coined by Adolf Seilacher in 1970. One of the most notable  is Burgess Shale in the Canadian Rockies of British Columbia. The site, discovered by Charles Walcott in 1909, highlight one of the most critical events in evolution: the Cambrian Explosion (540 million to 525 million years ago). The factors that can create such fossil bonanzas are: rapid burial (obrution), stagnation (eutrophic anoxia), fecal pollution (septic anoxia), bacterial sealing (microbial death masks), brine pickling (salinization), mineral infiltration (permineralization and nodule formation by authigenic cementation), incomplete combustion (charcoalification), desiccation (mummification) and freezing. The preservation of decay-resistant lignin of wood and cuticle of plant leaves  is widespread, but exceptional preservation also extends to tissues.

The Toploje Member chert of the Prince Charles Mountains preserves the permineralised remains of a terrestrial ecosystem before the biotic decline that began in the Capitanian and continued through the Lopingian until the Permo-Triassic transition (Slater et al., 2014). During the late Palaeozoic and early Mesozoic, Antarctica occupied a central position within Gondwana and played a key role in floristic interchange between the various peripheral regions of the supercontinent.


Singhisporites hystrix, a megaspore with ornamented surface.

The fossil micro-organism assemblage includes a broad range of fungal hyphae and reproductive structures. The macrofloral diversity in the silicified peats is relatively low and dominated by the constituent dispersed organs of arborescent glossopterid and cordaitalean gymnosperms.  The fossil palynological assemblage includes a broad range of dispersed bisaccate, monosaccate, monosulcate and polyplicate pollen. The roots (Vertebraria), stems (Australoxylon) and leaves (Glossopteris) of the arborescent glossopterid exhibited feeding traces caused by arthropods, but the identification is  difficult since plant and arthropod cuticles look similar in thin section. Tetrapods are currently unknown from Permian strata of the Prince Charles Mountains as either body fossils or ichnofossils (McLoughlin et al., 1997, Slater et al., 2014).

Times of exceptional fossil preservation are coincident with mass extinctions, oceanic anoxic events, carbon isotope anomalies, spikes of high atmospheric CO2, and transient warm-wet paleoclimates in arid lands (Retallack 2011). The current greenhouse crisis delivers several factors that can promote exceptional fossil preservation, such as eutrophic and septic anoxia, microbial sealing, and permineralization.


Benton, M.J., Newell, A.J., (2013), Impacts of global warming on Permo-Triassic terrestrial ecosystems. Gondwana Research.

Rees, P.M., (2002). Land plant diversity and the end-Permian mass extinction. Geology 30, 827–830.

Retallack, G., (2011), Exceptional fossil preservation during CO2 greenhouse crises?, Palaeogeography, Palaeoclimatology, Palaeoecology 307: 59–74.

Slater, B.J., et al., (2014), A high-latitude Gondwanan lagerstätte: The Permian permineralised peat biota of the Prince Charles Mountains, Antarctica, Gondwana Research.

Seilacher, A., (1970) “Begriff und Bedeutung der Fossil-Lagerstätten: Neues Jahrbuch fur Geologie und Paläontologie“. Monatshefte (in German) 1970: 34–39.

The plant fossil record and the extinction events.

Odontopteris lingulata, seed fern from the Late  Late Pennsylvanian to Early Permian. (Image Credit: Taylor et al, 2009)

Odontopteris lingulata, seed fern from the Late Late Pennsylvanian to Early Permian. (Image Credit: Taylor et al, 2009)

Mass extinctions has shaped the global diversity of our planet several times during the geological ages. They were originally identified in the marine fossil record and have been interpreted as a result of catastrophic events or major environmental changes that occurred too rapidly for organisms to adapt.

In 1982, Jack Sepkoski and David M. Raup used a simple form of time series analysis at the rank of family to distinguish between background extinction levels and mass extinctions in marine faunas, and identified five major extinction events in Earth’s history: at the end of the Ordovician period, Frasnian (Late Devonian), Permian, Triassic and Cretaceous. But the plant fossil record reveals a different pattern of major taxonomic extinctions compared with marine organisms. The first of them took place at the Carboniferous-Permian transition, which is interpreted as result of the collapse of the tropical wetlands in Euramerica. The second mass extinction corresponds to the end-Permian event.

Glossopteris sp., seed ferns, Permian - Triassic - Houston Museum of Natural Science (From Wikimedia Commons)

Glossopteris sp., seed ferns, Permian – Triassic – Houston Museum of Natural Science (From Wikimedia Commons)

At the late Carboniferous the characteristic wetland families disappeared (e.g. Flemingitaceae, Diaphorodendraceae, Tedeleaceae, Urnatopteridaceae, Alethopteridaceae, Cyclopteridaceae, Neurodontopteridaceae). The downfall of rainforests probably reflects the complexity of the environmental changes that were taking place during the late Moscovian-early Sakmarian time interval (DiMichele et al., 2006, Sahney et al., 2010). This collapse probably drove the rapid diversification of Carboniferous tetrapods (amphibians and reptiles) in Euramerica (Sahney et al., 2010).

During the end-Permian Event, the woody gymnosperm vegetation (cordaitaleans and glossopterids) were replaced by spore-producing plants (mainly lycophytes) before the typical Mesozoic woody vegetation evolved. The palynological record suggests that wooded terrestrial ecosystems took four to five million years to reform stable ecosystems, while spore-producing lycopsids had an important ecological role in the post-extinction interval.

Schematic illustration comparing the three extinction events analized (From Vajda and Bercovici, 2014)

Schematic illustration comparing the three extinction events analized (From Vajda and Bercovici, 2014)

At the end-Triassic event,  the vegetation turnover in the Southern Hemisphere  consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one.

The end-Cretaceous biotic crisis had a significant effect on marine and terrestrial faunas, and caused localized loss of species diversity in vegetation. Patagonia shows a reduction in diversity and relative abundance in almost all plant groups from the latest Maastrichtian to the Danian, although only a few true extinctions occurred (Barreda et al, 2013). The nature of vegetational change in the south polar region suggests that terrestrial ecosystems were already responding to relatively rapid climate change prior to the K–Pg catastrophe.

Two examples of grains pollen from the Lopez de Bertodano Formation: Podocarpidites sp. (left) and Nothofagidites asperus (right)

Two examples of grains pollen: Podocarpidites sp. (left) and Nothofagidites asperus (right)

A key factor for plant resilience is the time-scale: if the duration of the ecological disruption did not exceed that of the viability of seeds and spores, those plant taxa have the potential to recover (Traverse, 1988).


Cascales-Miñana, B., and C. J. Cleal, 2014, The plant fossil record reflects just two great extinction events. Terra Nova. vol. 26, no. 3, pp. 195–200. DOI: 10.1111/ter.12086

Bambach, R.K., Knoll, A.H. and Wang, S.C., 2004. Origination, extinction, and mass depletions of marine diversity. Paleobiology, 30, 522–542.

Mayhew, Peter J.; Gareth B. Jenkins, Timothy G. Benton (January 7, 2008). “A long-term association between global temperature and biodiversity, origination and extinction in the fossil record”. Proceedings of the Royal Society B: Biological Sciences 275 (1630): 47–53.

Sahney, S., Benton, M.J. & Falcon-Lang, H.J. (2010). “Rainforest collapse triggered Pennsylvanian tetrapod diversification in Euramerica” (PDF). Geology 38 (12): 1079–1082. doi:10.1130/G31182.1.


Halloween special II: Lovecraft, Paleobotany and The Shadow Out of Time.

Howard Phillips Lovecraft_in_1915_(2)

Howard Phillips Lovecraft in 1915.

Howard Phillips Lovecraft was born on August 20, 1890 in Providence, Rhode Island. He was one of the most influential writers of the twentieth century.  Despite leaving school without graduating, in his writings, evidences an extensive knowledge of archaeology, astronomy, geology, and paleontology. As an amateur astronomer, Lovecraft attended several lectures from leading astronomers and physicists of his time. He  explicitly stated in a letter to a friend that Yuggoth is in fact the then recently discovered Pluto. This was one of the key aspects in Lovecraft’s literature: to reject the old spiritual world and use the advance of science as a source of inspiration.

“The Shadow Out of Time” (1935) was H. P. Lovecraft’s last major story. It’s told from the perspective of Nathaniel Wingate Peaslee, a professor of political economy at Miskatonic University. During five years, this man suffers a bizarre form of amnesia  followed by vivid dreams of aliens cities in ancient landscapes.  Later, Peaslee discovered that a small number of people throughout history suffered the same type of amnesia. They were possessed by the Great Race, a group of cone shaped creatures who developed the technique of swapping minds with creatures of another era with the purpose of learn the secrets of the Universe.

Lepidodendrom leaf cushions preserved in a Mazon Creek nodule. (Taylor et al., 2009)

Lepidodendrom leaf cushions preserved in a Mazon Creek nodule. (Image Credit: Taylor et al, 2009)

Peaslee describes the gardens that surround the cities of his visions with detail. There was calamites, cycads, trees of coniferous aspect, and small, colourless flowers.

Calamites was a genus of tree-sized, spore-bearing plants that lived during the Carboniferous and Permian periods (about 360 to 250 million years ago), closely related to modern horsetails. They reached their peak diversity in the Pennsylvanian and were major constituents of the lowland equatorial swamp forest ecosystems. The Cycadales are an ancient group of seed plants that can be traced back to the Pennsylvanian. Cycads have a stem or trunk that commonly looks like a large pineapple and composed of the coalesced bases of large leaves.  Today’s cycads are found in the tropical, subtropical and warm temperate regions of both the north and south hemispheres.

While angiosperm fossil pollen first appears in the Early Cretaceous, molecular data suggest that the first occurrence was in the early Permian (~275 Ma) to late Triassic (228-217 Ma). Recently, a new study describe six distinct pollen types that have angiosperm-like features from the Triassic of Switzerland.


Transverse section of Sigillaria approximata stem (Image Credit: Taylor et al, 2009)

Peaslee’s visions become more and more vivid:

The far horizon was always steamy and indistinct, but I could see that great jungles of unknown tree-ferns, calamites, lepidodendra, and sigillaria lay outside the city, their fantastic frondage waving mockingly in the shifting vapours.”

Lepidodendron was a tree-like (‘arborescent’) tropical plant, related to the lycopsids. The name of the genus comes from the Greek lepido, scale, and dendron, tree, because of the distinctive diamond shaped pattern of the bark. The name Lepidodendron is a generic name given to several fossil that clearly come from arborescent lycophytes but are difficult to assign to one species. Fossil remains indicate that some trees attained heights in excess of 40 m and were at least 2m in diameter at the base. They were dominant components of swamp ecosystems in the Carboniferous and frequently associated with Sigillaria, another extinct genus of tree-sized lycopsids from the Carboniferous Period. The absence of extensive branching and the structure of the leaf bases are the principal feature that distinguish Sigillaria from other lycopsids (Taylor et al, 2009). Sigillariostrobus is the name assigned to the reproductive organs or cones of Sigillaria. Unlike Lepidodendron cones, which were attached attached individually near the tip of the branches, Sigillaria cones occurred in clusters attached in certain places along the upper stem.

Later, on an expedition to Australia, Peaslee – accompanied by Professor William Dyer, leader of the Miskatonic Antarctic expedition of 1930-1931- discovered a manuscript written by himself eons ago when he was a captive mind of the Great Race.


H. P. Lovecraft, The Dreams in the Witch House and Other Weird Stories, Penguin Books, 2004.

Joshi, S. T. (2001). A dreamer and a visionary: H.P. Lovecraft in his time. Liverpool University Press, 302.

N. Taylor, Edith L. Taylor, Michael Krings: “Paleobotany: The Biology and Evolution of Fossil Plants”. 2nd ed., Academic Press 2009.

Kathy Willis, Jennifer McElwain, The Evolution of Plants, Oxford University Press, 2013

Hochuli, P. A., and Feist-Burkhardt, S.. (2013). Angiosperm-like pollen and Afropollis from the Middle Triassic (Anisian) of the Germanic Basin (Northern Switzerland). Frontiers in Plant Science. 4. doi: 10.3389/fpls.2013.00344