Living Fossils, Links for Palaeobotanists
Links for Palaeobotanists

Home / Evolution & Extinction / Living Fossils

Web Sites about Evolution
Focussed on the Fossil Record
The Molecular Clock and/or/versus the Fossil Record
Insect Evolution
Evolution Sciences versus Doctrines of Creationism and Intelligent Design
Web Sites about Mass Extinctions
The Mass Extinction at the End of the Permian
Biotic Recovery from the Permian-Triassic Mass Extinction
The Mass Extinction at the End of the Triassic
! Teaching Documents about Evolution@
! Focused on the Evolution of Plants@
! Articles in Palaeobotany@
! The Gaia Hypothesis@
Teaching Documents about Palaeontology and Palaeoecology@
Databases focused on Palaeobotany and Palaeontology@
Databases focused on Botany and Biology@
Glossaries, Dictionaries and Encyclopedias: Palaeontology@
Glossaries, Dictionaries and Encyclopedias: Biology@

Living Fossils

Y. Asar et al. (2022): Evaluating the accuracy of methods for detecting correlated rates of molecular and morphological evolution. In PDF, bioRxiv.
See also here.
! Note figure 1 (on PDF-page 9): A flowchart of simulation study. About molecular and morphological phylograms, morphological characters and sequence alignments.

D.J. Beerling et al. (1998): Stomatal responses of the "living fossil" Ginkgo biloba L. to changes in atmospheric CO2 concentrations. PDF file, Journal of Experimental Botany, 49: 1603-1607.

D.J. Bennett et al. (2018): Quantifying the living fossil concept. Palaeontologia Electronica.

D.J. Bennett et al. (2017): Evolutionarily distinct “living fossils” require both lower speciation and lower extinction rates. In PDF, Paleobiology, 43: 34-48. See also here.

!: D.J. Bennett (2017): An appraisal of the ‘Living Fossil’ Concept. In PDF, Thesis, 262 p. Imperial College London.
See also here.

M.J. Benton and P.N. Pearson (2001): Speciation in the fossil record. PDF file, Trends in Ecology and Evolution, 16.

Museum of Paleontology, University of California, Berkeley, CA: Why all the fuss? The fascination of coelacanths. This expired link is available through the Internet Archive´s Wayback Machine.

B. Bomfleur et al. (2014): Fossilized Nuclei and Chromosomes Reveal 180 Million Years of Genomic Stasis in Royal Ferns. In PDF, Science, 343. See also here.

D.R. Brooks and S.M. Bandoni (1988): Coevolution and Relicts. Abstract, Systematic Biology, 37: 19–33.
"... Phylogenetic relicts are “living fossils,” members of old groups that have persisted relatively unchanged for a long period of time, such as horse-shoe crabs. Coevolved phylogenetic relicts are either restricted to host groups that are themselves phylogenetic relicts, or inhabit members of a non-relictual host group that share a common plesiomorphic ecology. ..."

! Mark Carnall (2016): Let's make living fossils extinct. The Guardian.

! D. Casane and P. Laurenti (2013): Why coelacanths are not ‘living fossils’. A review of molecular and morphological data. In PDF, Bioessays, 35: 332–338.

L. Cavin and N. Alvarez (2022): Why Coelacanths Are Almost “Living Fossils”? Free access, Front. Ecol. Evol., 10:896111. doi: 10.3389/fevo.2022.896111.
Note fig. 1: Selected life history traits and stratigraphical range for Latimeria and two Mesozoic coelacanths.

S. Conway Morris (1993): The fossil record and the early evolution of the Metazoa. PDF file, Nature, 361: 219–225.
See also here.

! C.C. Davis and H. Schaefer (2011): Plant Evolution: Pulses of Extinction and Speciation in Gymnosperm Diversity. See also here (abstract).

W.A. DiMichele et al. (2004): Long-term stasis in ecological assemblages: evidence from the fossil record. PDF file, Annu. Rev. Ecol. Evol. Syst., 35: 285-322. This expired link is available through the Internet Archive´s Wayback Machine.

! M.J. Donoghue et al. (2015): Confluence, synnovation, and depauperons in plant diversification. Open access, New Phytologist, 207: 260–274.

George Dvorsky, 12 of the most astounding "living fossils" known to science.

K.C. Elliott et al. (2016): Conceptions of Good Science in Our Data-Rich World. In PDF, Bioscience, 66: 880–889. S ee also here.

Peter Forey, The Natural History Museum, London: Nature online > Life > Reptiles, amphibians and fishes > Living fossils: coelacanths (PDF file). Coelacanths and the ancestry debate. See also here., Vienna: On the phenomenon of living fossils. PDF file, in German.
This expired link is now available through the Internet Archive´s Wayback Machine.

S. Gilmore and K.D. Hill (1997): Relationships of the Wollemi Pine (Wollemia nobilis) and a molecular phylogeny of the Araucariaceae. PDF file, Telopea 7. See also here.

! P. Grandcolas et al. (2014): Relict species: a relict concept? In PDF, Trends in Ecology & Evolution, 29: 655–663.
Still available via Internet Archive Wayback Machine.
See also here.

K.-J. Gu et al. (2022): GinkgoDB: an ecological genome database for the living fossil, Ginkgo biloba. Free access, Database, 2022: article ID baac046; DOI:

P. Gueriau et al. (2016): A 365-Million-Year-Old Freshwater Community Reveals Morphological and Ecological Stasis in Branchiopod Crustaceans. Open access, Current Biology, 26: 383-390.

Nina Horvath, Linz, Austria: Living Fossils (in German).
Now recovered from the Internet Archive´s Wayback Machine.

R. Heady (2012): The Wollemi Pine—16 years on. In PDF, Chapter 15: Australia’s Ever-changing Forests VI: Proceedings of the Eighth National Conference on Australian Forest History. Brett J. Stubbs et al. (ed.).
Snapshot provided by the Internet Archive´s Wayback Machine.

Thomas M. Iliffe and Louis S. Kornicker (2009): Worldwide Diving Discoveries of Living Fossil Animals from the Depths of Anchialine and Marine Caves. PDF file, Smithsonian Contributions to the Marine Sciences.
Snapshot provided by the Internet Archive´s Wayback Machine.

Susan L. Jewett, Division of Fishes, Smithsonian Institution's National Museum of Natural History: On the Trail of the Coelacanth, a Living Fossil. November 11, 1998; The Washington Post.

N. Jürgens et al. (2021): Welwitschia: Phylogeography of a living fossil, diversified within a desert refuge. Open access, Scientific Reports, 11.
"... With a fossil record of 112 My and phylogenetically isolated within the order Gnetales, the monotypic genus Welwitschia has survived only in the northern Namib Desert in Angola and Namibia. Despite its iconic role, the biogeography, ecological niche, and evolutionary history of the species remain poorly understood. Here we present the first comprehensive map of the strongly disjunct species range, and we explore the genetic relationships among all range fragments based on six SSR markers. ..."

J.L. King and R. Hanner (1998): Cryptic species in a “living fossil” lineage: Taxonomic and phylogenetic relationships within the genus Lepidurus (Crustacea: Notostraca) in North America. Abstract, Molecular Phylogenetics and Evolution, 10: 23-36.

! A. Knowlton (2012): Quick guide Equisetum. In PDF, Current Biology, 10.

! N.M. Koch and L.A. Parry (2020): Death is on Our Side: Paleontological Data Drastically Modify Phylogenetic Hypotheses. Free access, Syst. Biol., 69: 1052–1067.
See also here and there.
"... Since the early years of phylogenetic systematics, different studies have dismissed the impact of fossils due to their incompleteness, championed their ability to overturn phylogenetic hypotheses or concluded that their behavior is indistinguishable from that of extant taxa. Based on taxon addition experiments on empirical data matrices, we show that the inclusion of paleontological data has a remarkable effect in phylogenetic inference. ..."

R. Leinfelder (2004): „Lebende Fossilien“ Eine Einführung zur Ausstellung (PDF-file, in German).
See also:
Von der Evolution vergessen? Lebende Fossilien (in German, Paläontologischen Museums München).

! S. Lidgard and E. Kitchen (2023): Revealing the rise of a living fossil menagerie. Front. Ecol. Evol. 11:1112764. doi: 10.3389/fevo.2023.1112764
"... In this paper we&xnbsp;attempt the first comprehensive analysis of the use of the term ‘living fossil’ from 1860 to the present.
[...] future utilization of living fossils reflects increasingly complex challenges to deep-seated expectations at the heart of a living fossil research agenda: omnipresent evolutionary change and inevitable disappearance or extinction.

! S. Lidgard and A.C. Love (2021): The living fossil concept: reply to Turner. Open access, Biology & Philosophy, 36.
"... The LF concept—both in the past and even more so in the present—is multifaceted and theoretically rich because it corresponds to an ongoing research program in evolutionary biology. It includes biodiversity considerations based on aesthetic values but is by no means exhausted by them. ..."

! S. Lidgard and A.C. Love (2018): Rethinking Living Fossils. Open access, BioScience, 68: 760–770.
Note figure 2: Modern and ancient representatives of living fossils Triops, Ginkgo, and xiphosurans.
"... work on character evolution illustrates how analyzing living fossils and stasis in terms of parts (characters) and wholes (e.g., organisms and lineages) advances our understanding of prolonged stasis at many hierarchical levels. Instead of viewing the concept’s task as categorizing living fossils, we show how its primary role is to mark out what is in need of explanation, accounting for the persistence of both molecular and morphological traits ..."

Links for Palaeobotanists: Introductions to both Fossil and Recent Plant Taxa, e.g. Sphenophyta, Cycads, and Ginkgoales.

L.H. Liow (2006): Oddities, Wonders, and Other Tall-tales of "living Fossils". In PDF. Thesis, University of Chicago.

J. Ma (2003): The chronology of the "living fossil" Metasequoia glyptostroboides (Taxodiaceae): a review (1943-2003). PDF file, Harvard Papers in Botany, 8: 9-18. See also here.
"... On the basis of primary documents including letters, manuscripts, and original publications, plus personal experience, the major events, important publications, and main scientists related to this story are recorded chronologically for the first time 60 years after the species’ discovery ..."

Steven R. Manchester (website hosted by International Organisation of Palaeobotany): Living Fossils, Davidia - the Dove Tree and its fossil record.
The link is to a version archived by the Internet Archive´s Wayback Machine.

B. Mantovani et al. (2004): Molecular taxonomy and phylogeny of the "living fossil" lineages Triops and Lepidurus (Branchiopoda: Notostraca). PDF file, Zoologica Scripta, 33: 367-374.
Website outdated, download a version archived by the Internet Archive´s Wayback Machine.

T.C. Mathers et al.(2013): Multiple global radiations in tadpole shrimps challenge the concept of "living fossils". PeerJ., 1.
See also here (Der Spiegel, in German).

Palaeontological Museum Munich, Germany: Online-Ausstellung "Lebende Fossilien". In German.

J. Murienne et al. (2013): A living fossil tale of Pangaean biogeography. Proc. R. Soc., B 281.

! N.S. Nagalingum et al. (2011): Recent Synchronous Radiation of a Living Fossil. Abstract.
"Using fossil-calibrated molecular phylogenies, we show that cycads underwent a near synchronous global rediversification beginning in the late Miocene, followed by a slowdown toward the Recent. Although the cycad lineage is ancient, our timetrees indicate that living cycad species are not much older than ~12 million years". See also here. In PDF, Science 334.

R. Nuwer (2013): There’s No Such Thing as a ‘Living Fossil’. Smithsonian Magazine.
"... The tadpole shrimp, for example, turns out to include 38 distinct species, descended from two subtypes that diverged a relatively recent 73 million years ago. The organism is constantly adapting to new conditions, even if its general body plan remains the same. ..."

C.A. Offord et al. (1999): Sexual Reproduction and Early Plant Growth of the Wollemi Pine (Wollemia nobilis), a Rare and Threatened Australian Conifer. PDF file, Annals of Botany 84.

! K.E. Omland et al. (2008): Tree thinking for all biology: the problem with reading phylogenies as ladders of progress. In PDF, BioEssays, 30: 854–867.
See also here.

David N. Reznick and Robert E. Ricklefs (2009): Darwin´s bridge between microevolution and macroevolution. PDF file, Nature, 257. Provided by the Internet Archive´s Wayback Machine.

Mark Ridley, Department of Zoology, Oxford University: Evolution, 3rd Edition. Book announcement. See also: Evolution (Blackwell). Go to:
! Living Fossils.

! D.L. Royer et al. (2003): Ecological conservatism in the "living fossil" Ginkgo. In PDF, Paleobiology, 29: 84–104.
See also here.
"... The living species Ginkgo biloba is phylogenetically isolated, has a relictual distribution, and is morphologically very similar to Mesozoic and Cenozoic congenerics. [...] Ginkgo is an extreme example of a geologically long-lived genus, with its one living species arguably having a temporal range of >100 Myr. ..."

G. Sanchez et al. (2022): Beyond “living fossils”: Can comparative genomics finally reveal novelty? Free access, Molecular Ecology Resources, 22: 9–11.

! T.J.M. Schopf (1984): Rates of Evolution and the Notion of "Living Fossils". In PDF, Annual Review of Earth and Planetary Sciences, 12. See also here.

Andrew Simpson, Science Museums, Division of Environmental and Life Sciences, Macquarie University NSW: Andrew's Web Diary. A personal report from the Latin American Palaeontology Conference (Sociedade Brasileira de Paleontologia in Aracaju, in the north east of Brazil). Go to: DAY 5 - The plants of Gondwana and the Wollemi Pine.

P. Smarda et al. (2016): Polyploidy in a ‘living fossil’ Ginkgo biloba. Open access, New Phytologist, 212: 11–14.

Pamela S. Soltis et al. (2002): Rate heterogeneity among lineages of tracheophytes: Integration of molecular and fossil data and evidence for molecular living fossils. PDF file, PNAS, 99: 4430-4435. See also here (abstract).

B. Sterner (2023): Norms of evidence in the classification of living fossils. Frontiers in Ecology and Evolution, 11: 1198224. doi: 10.3389/fevo.2023.1198224.
See likewise here.

T.H. Struck and J. Cerca (2022): Extant Cryptic Species as Systems to Understand Macro-Evolutionary Stasis. Google books. In: Simon J. Mayo, Alexandre K. Monro (eds): Cryptic Species, Morphological Stasis, Circumscription, and Hidden Diversity. Cambridge University Press.

E. Thenius (2003): „Lebende Fossilien“ im Tier- und Pflanzenreich. Fiktion oder Realität?. PDF file, in German. Schriften Verein zur Verbreitung naturwissenschaftlicher Kenntnisse, 141: 99-123.

E. Thenius (1972): Versteinerte Urkunden: die Paläontologie als Wissenschaft vom Leben in der Vorzeit. Google books.

! D. Turner (2019): In Defense of Living Fossils. Biology and Philosophy. See also here.

G.J. Vermeij (2015): Paleophysiology: From Fossils to the Future. Trends in ecology & evolution.

A. Watkins (2021): The epistemic value of the living fossils concept. In PDF, Philosophy of Science, 88: 1221-1233.
See also here.
"... This paper's primary contribution has been to disambiguate the inferences that we can justifiably make on the basis of classifying a taxon as a living fossil. In doing so, I have specified some of the ways in which the living fossil concept may be epistemically useful.
Note table 1: Summary.

! A.J. Werth and W.A. Shear (2014): The evolutionary truth about living fossils. Abstract, American Scientist, 102. See also here (in PDF).

Wikipedia, the free encyclopedia:
Living fossil.
Lebendes Fossil (in German).
Triops cancriformis.
! Pages in category "Living fossils".

N. Wikström et al. (2022): No phylogenomic support for a Cenozoic origin of the “living fossil” Isoetes. OPen access, American Journal of Botany.

D. Wood et al. (2020): Phylogenomics indicates the “living fossil” Isoetes diversified in the Cenozoic. Open access, Plos One, 15.
"... genomic analyses coupled with a careful evaluation of the fossil record indicate that despite resembling forms from the Triassic, extant Isoetes species do not represent the remnants of an ancient and widespread group, but instead have spread around the globe in the relatively recent past. ..."

! G. Wörheide et al. (2016): Molecular paleobiology — Progress and perspectives. Abstract, Palaeoworld, 25: 138–148. See also here (in PDF).

Yale Peabody Museum of Natural History, New Haven, Connecticut:
The Dawn Redwood - A Living Fossil.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.
See also here.

! E. Yong (2013): The Falsity of Living Fossils. The Scientist Magazine. See also here (Richard Dawkins Foundation).

K. Yoshida (2002): Long survival of “living fossils” with low taxonomic diversities in an evolving food web. Abstract, Paleobiology, 28.

! Y.P. Zhao et al. (2019): Resequencing 545 ginkgo genomes across the world reveals the evolutionary history of the living fossil. Open access, Nature Communications, 10.
"... investigations provide insights into the evolutionary history of ginkgo trees and valuable genomic resources for further addressing various questions involving living fossil species.
[...] It is likely that the morphological stasis of living fossils is an effective adaptation strategy in response to environmental change, although the underlying mechanisms are unclear ..."

! Zhiyan Zhou (website hosted by International Organisation of Palaeobotany): Gingko biloba: its ancestors and allies.
Now recovered from the Internet Archive´s Wayback Machine.

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