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Waterloo Farm lagerstätte

January 07, 2023

The Waterloo Farm lagerstätte is a Famennian lagerstätte in South Africa that constitutes the only known record of a near-polar Devonian coastal ecosystem.

Waterloo Farm in 2016.

History and discovery

The Waterloo Farm Lagerstätte is an approximately 360 million year old Famennian (latest Devonian) fossil-rich locality of the Witpoort Formation (Witteberg Group, Cape Supergroup) in Makhanda (former Grahamstown) within the Eastern Cape Province, South Africa.[1][2] Before it’s discovery very little was known of life during the Famennian (see Late Devonian extinction) in what is now southern Africa . This is largely due to the fact that fossils in the Witpoort Formation generally occur in black anaerobically deposited metashale that rapidly degrades near surface and is therefore rarely seen in natural outcrop.[3][4] As is the case with many other scientific discoveries, the discovery of Waterloo Farm was accidental.[5]

Uprisings against the apartheid system in South Africa had by the mid-1980s escalated to violent protests, particularly in the ‘townships’ (designated black residential areas). Prior to that the N2, the main road linking the industrial ports of Port Elizabeth (now Gqeberha) and East London, ran through then Grahamstown, including its township.[5] By 1985, protests and stoning of cars had made it unsafe for traffic to drive through the townships.[5] As a result the local government proposed the construction of a new road that would serve as a bypass around the townships.[5] Due to limited options, it was decided that the bypass would be constructed through the valleyed countryside to the south of then Grahamstown.[5] This involved excavation of extensive road cuttings through hills and spurs of the Rietberg and utilisation of the resultant rubble to build viaducts across the intervening valleys.[5] Amongst the small farms from which land was appropriated for the road was Waterloo Farm, owned by Rhodes University Botany professor, Roy Lubke - with one cutting passing virtually through the farm yard.[5] At a braai (barbeque) held by Roy Lubke one afternoon, a colleague Dr Mark Aken wandered off to the top of the cutting and peered down the slope. He returned to his colleagues to inform them that there were plant fossils in the disturbed black shale. Roy Lubke mentioned this to Dr Fred Gess, an entomologist, who passed the information on to his son Robert Gess. As Robert Gess had a great interest in the fossils of the Eastern Cape he immediately made his way out to the roadworks where he was struck by the uniquely well-preserved nature of the fossils. With the encouragement of his supervisor and mentor Dr Norton Hiller of the Rhodes Geology Department, he started assembling a collection of the fossils.[5] In 1991 Fiona Taylor, an honours student of Dr Hiller, conducted a short study on the geological context of the deposit, and also published a preliminary note on the fossils, including some of those from Gess’ collection. During a short combined excavation by Gess and Taylor, more material was found including remains of large armour-plated (placoderm) fish, later identified as Africa’s only known species of Bothriolepis. Taylor’s sedimentological work was presented in an article titled “Late Devonian shoreline changes: an analysis of Witteberg Group stratigraphy in the Grahamstown area” published in 1992 (Hiller and Taylor).[3] In 1993, Hiller employed Robert Gess to conduct a thorough palaeontological study and his excavations continued until 1995, after the emigration of Dr Hiller. This phase of research resulted in a number of papers , which were the subject of his master’s degree.[5][4]

By the mid-90’s, the road cutting was frequently collapsing due to the dip direction of the strata being towards the road.[5] This led to a near permanent closure of one lane for safety reasons.[5] After there had been a failed attempt to mitigate the situation with steel barriers in the mid-nineties, the newly established South African National Road Agency Limited (SANRAL) published a call for tenders to upgrade the road.[5] This would prove a propitious opportunity for Robert Gess as, by this time, he was struggling to secure further funding for excavations and research.[5] In September 1999, Gess, who had taken up residence in Bathurst (45 km away from Makhanda), heard that a group of consulting engineers from Jeffares and Green (now known as JG Africa) had been awarded the tender to cut back and stabilise the cuttings.[5] He immediately contacted the company and SANRAL to inform them of the sensitivity of the site.[5] SANRAL made funds available for Gess to rescue some of the shale for scientific research.[5] This was hand mined in blocks by Gess and a team under his direction before being transported on a flat-bed truck to his property in Bathurst.[5] Gess built a shed roof over the sample to protect them from the rapid decay that characterises the shale when exposed to weather.[5] It is from meticulous excavation of these recued blocks that most subsequent discoveries have been and continue to be made.[5] A similar process during further roadworks in 2007 and 2008 yielded an even larger haul of fossiliferous blocks which are stored in sheds constructed by SANRAL.[5]

The fossils are housed and researched by the Devonian Ecosystems project, (funded by the Millennium Trust and South African Centre of Excellence in Palaeosciences) in the Albany Museum’s Devonian lab at 87 Beaufort Street, Makhanda (formerly Rhini or Grahamstown). They have provided the material for extensive research by Dr Rob Gess and his collaborators, and an ongoing stream of significant papers.[1][6][7][8]

Contributions to palaeontological material

Waterloo Farm is a globally significant site, providing the only record of a high latitude (near polar) coastal ecosystem, overturning numerous assumptions about high latitude conditions during the latest Devonian.[9][1][7] Previous sparse evidence from sub-Saharan Africa, South America and Antarctica had previously led researchers to believe that high latitude conditions precluded extensive vegetation of land and high vertebrate diversity, and that for example Devonian tetrapods only occurred within tropical settings.[10] Waterloo farm has totally disrupted such beliefs, providing evidence for a diversely vegetated adjacent terrestrial habitat with plants including Archaeopteris trees, a diverse estuarine vertebrate fauna[1] and the only known non-tropical Devonian tetrapods.[9] Exceptional soft tissue preservation at Waterloo Farm is unique for a Famennian estuary and allows for reconstruction of an entire estuarine ecosystem,[7] grading from brackish to more marine conditions and including delicate waterweeds,[11] invertebrates[12] and diverse vertebrates.[13][9]  As yet 25 species have been diagnosed from Waterloo Farm and many others provisionally identified. An excess of 50 organisms are however believed to be represented in the extensive collections of the Devonian Ecosystems Project. Together this represents the most holistically studied Devonian tetrapod-bearing locality. Insights provided by the unique soft tissue preservation at the site are also making major contributions to evolutionary biology, such as a growth series of ancient lampreys (Priscomyzon riniensis), that have overturned major, long held, perspectives on vertebrate origins.[7]

Site characteristics

Witpoort Formation black shales within the Eastern Cape often exhibit cyclical changes in composition, which likely reflect (potentially seasonal) fluctuations in water salinity. Water stratification within the estuarine lake frequently led to anoxic bottom waters, resulting in episodes of exceptional preservation.[14]

Witpoort Formation sediments were deeply buried due to continued basinal subsidence through the Carboniferous, and were subsequently metamorphosed during the massive Permian aged Cape Fold Belt orogenesis.[15] Hundreds of millions of years of erosion and uplift brought the Waterloo Farm shales back up to near surface, they were exposed in 1985, in new road cuttings south of Makhanda/Grahamstown, during construction of a bypass road.

 
Waterloo Farm during the excavation in 1985.

On-site excavations were conducted in the 1990s, but the instability of the road cutting led to it being cut back in 1999 and in 2008. On both occasions large quantities of shale were rescued which provides for ongoing excavation. Decades of research has revealed the most important Late Devonian fossil site from what was the southern portion of Gondwanan region incorporating present-day sub-saharan Africa, South America and western Antarctica.[16]

 
Waterloo Farm in 1988.
 
Waterloo Farm main fossil site in 1999 preceding roadworks.
 
One of the sheds of shale blocks rescued (with the help of SANRAL) from Waterloo Farm roadworks in 2008, for ongoing scientific excavation.

Because the original fine black mud was often very low in oxygen, plants and animals rapidly buried in it sometimes left behind impressions of their soft parts. This is extremely rare in the fossil record which normally only preserves bones, teeth and other hard bits. Exceptionally, what is recorded is the remains of an entire estuarine ecosystem, from delicate waterweeds and seaweeds to small clams, baby fish and the bones of larger fish. Land plants which grew nearby are also preserved, from the remains of small undergrowth species to fronds from the earliest types of trees.[17]

Fossils

More than 20 species new to science have been named from Waterloo Farm, which probably represent about a third of the total number of taxa indicated by remains preserved in the shale. Taxa include the tetrapods Tutusius umlambo and Umzantsia amazana,[18] which are Africa's earliest known tetrapods and the only non-tropical Devonian tetrapods known. The first described fossils from Waterloo Farm comprise remains of sub-Saharan Africa's earliest woody trees (Archaeopteris notosaria).

 
Archaeopteris notosaria fronds from the Waterloo Farm lagerstätte in the Eastern Cape, South Africa. These represent the only high latitude species of Archaeopteris as yet described.

[19] Other fossils from Waterloo Farm include the oldest known land-living animal from Gondwana (the scorpion Gondwanascorpio emzantsiensis),[20] the oldest fossil lamprey in the world (Priscomyzon riniensis)[21] and Africa's oldest coelacanth from the world's earliest known coelacanth nursery (Serenichthys kowiensis).[22]

Other species represented include several species of armour plated (placoderm) fish, spiny finned (acanthodian) fish, sharks, ray-finned (actinopterygian) fish, a range of lobe-finned fish, bivalves; seaweeds; charophyte waterweeds, and a diverse range of plants.

List of published taxa from Waterloo Farm

Animalia

Invertebrates

Mollusca, Bivalvia

Naiadites form Devonica[23]

Arthropoda
Eurypterida

Cyrtoctenid eurypterid indet.[17]

Arachnida, Scorpiones

Gondwanascorpio emzantsiensis[20]

Vertebrata

Agnatha

Priscomyzon riniensis[21]

Placodermi

Bothriolepis africana[24]

Groenlandaspis riniensis[24]

Africanaspis doryssa[25][24]

Africanaspis edmountaini[25]

Gen. et sp. indet.

Acanthodii

Diplacanthus acus[17]

Diplacanthus indet[17]

acanthodidid indet[17]

gyracanthid indet[17]

Chondrichthyes

Plesioselachus macracantha [26][22]

Antarctilamna ultima[22]

Actinopterygii

gen. et. sp. indet.

Sarcopterygii

Serenichthys kowiensis[22]

Hyneria sp.[27][28]

Rhizodont indet.[17]

Isityumzi mlomomde (gen. et sp. nov.) [29]

Tetrapoda

Tutusius umlambo[18]

Umzantsia amazana[18]

Plantae

Algae

Rhodophyta or Phaeophyta

Hungerfordia fionae (sp. nov.)[30][31]

Yeaia africana (sp. nov.)[30][31]

Charophyta

Hexachara setacea (gen. et sp. nov.)[32][31]

Hexachara riniensis (gen. et sp. nov.)[32][31]

Octochara crassa (gen. et sp. nov.)[32][31]

Octochara gracilis (gen. et sp. nov.)[32][31]

Tracheophyta

Zosterophyllopsida

Zosterophyll indet.[33]

Lycopsida

Leptophloeum rhombicum[34]

Kowieria alveoformis[35]

Iridopteridales

Iridopterlean indet[17]

Sphenopsida

Rinistachya hilleri[36]

Progymnospermopsida

Archaeopteris notosaria[19]

References

  1. ^ a b c d Gess, Robert W.; Whitfield, Alan K. (August 2020). "Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent". Biological Reviews. 95 (4): 865–888. doi:10.1111/brv.12590. ISSN 1464-7931. PMID 32059074. S2CID 211122587.
  2. ^ Frey, Linda; Coates, Michael; Ginter, Michal; Klug, Christian; Ginter, Michał (2017-07-07). "Skeletal remains of Phoebodus politus Newberry 1889 (Chondrichthyes: Elasmobranchii) from a Famennian Konservatlagerstätte in the eastern Anti-Atlas (Morocco) and its ecology". Ichthyolith Issues. Chęciny, Poland: University of Warsaw: 36. doi:10.5167/uzh-138397. S2CID 197550063.
  3. ^ a b Hiller & Taylor, Norton & Fiona (1992). "Late Devonian shoreline changes: an analysis of Witteberg Group stratigraphy in the Grahamstown area". South African Journal of Geology. 5: 203–214 – via AJA.
  4. ^ a b Wolfgang, Gess, Robert (2012-05-14). High latitude Gondwanan famennian biodiversity patterns : evidence from the South African Witpoort Formation (Cape Supergroup, Witteberg Group). University of the Witwatersrand. OCLC 811026675.
  5. ^ a b c d e f g h i j k l m n o p q r s t Gess, Robert Wolfgang (2002). "The palaeoecology of a coastal lagoon of the Witpoort Formation (Upper Devonian, Famennian) in the Eastern Cape Province, South Africa". University of Fort Hare, South Africa: 6–11 – via Fort Hare University, Southa Africa.
  6. ^ Gess, Robert W.; Coates, Michael I.; Rubidge, Bruce S. (October 2006). "A lamprey from the Devonian period of South Africa". Nature. 443 (7114): 981–984. Bibcode:2006Natur.443..981G. doi:10.1038/nature05150. ISSN 0028-0836. PMID 17066033. S2CID 4302716.
  7. ^ a b c d Miyashita, Tetsuto; Gess, Robert W.; Tietjen, Kristen; Coates, Michael I. (2021-03-18). "Non-ammocoete larvae of Palaeozoic stem lampreys". Nature. 591 (7850): 408–412. Bibcode:2021Natur.591..408M. doi:10.1038/s41586-021-03305-9. ISSN 0028-0836. PMID 33692547. S2CID 232192889.
  8. ^ Harris, Christopher; Gess, Robert W.; Penn-Clarke, Cameron; Rubidge, Bruce S. (2021-03-29). "Coombs Hill: A Late Devonian fossil locality in the Witpoort Formation (Witteberg Group, South Africa)". South African Journal of Science. 117 (3/4). doi:10.17159/sajs.2021/9190. ISSN 1996-7489. S2CID 233689294.
  9. ^ a b c Gess, Robert; Ahlberg, Per Erik (2018-06-07). "A tetrapod fauna from within the Devonian Antarctic Circle". Science. 360 (6393): 1120–1124. Bibcode:2018Sci...360.1120G. doi:10.1126/science.aaq1645. ISSN 0036-8075. S2CID 46965541.
  10. ^ Montes, C.; Cardona, A.; Jaramillo, C.; Pardo, A.; Silva, J. C.; Valencia, V.; Ayala, C.; Perez-Angel, L. C.; Rodriguez-Parra, L. A.; Ramirez, V.; Nino, H. (2015-04-10). "Middle Miocene closure of the Central American Seaway". Science. 348 (6231): 226–229. Bibcode:2015Sci...348..226M. doi:10.1126/science.aaa2815. ISSN 0036-8075. PMID 25859042. S2CID 206633672.
  11. ^ Hiller, Norton; Gess, Robert W. (March 1996). "Marine algal remains from the Upper Devonian of South Africa". Review of Palaeobotany and Palynology. 91 (1–4): 143–149. doi:10.1016/0034-6667(95)00062-3. ISSN 0034-6667.
  12. ^ Gess, Robert W. (December 2013). "The Earliest Record of Terrestrial Animals in Gondwana: A Scorpion from the Famennian (Late Devonian) Witpoort Formation of South Africa". African Invertebrates. 54 (2): 373–379. doi:10.5733/afin.054.0206. ISSN 1681-5556. S2CID 6336474.
  13. ^ Gess, Robert Wolfgang (September 2008). "6th Annual Meeting Society of Vertebrate Paleontology". Cleveland Museum of Natural History. 28: 2.
  14. ^ Gess, Robert W. (2016). "Vertebrate Biostratigraphy of the Witteberg Group and the Devonian-Carboniferous Boundary in South Africa". Origin and Evolution of the Cape Mountains and Karoo Basin. Regional Geology Reviews. pp. 131–140. doi:10.1007/978-3-319-40859-0_13. ISBN 978-3-319-40858-3.
  15. ^ Blewett, Scarlett C. J.; Phillips, David (2016). "An Overview of Cape Fold Belt Geochronology: Implications for Sediment Provenance and the Timing of Orogenesis". Origin and Evolution of the Cape Mountains and Karoo Basin. Regional Geology Reviews. pp. 45–55. doi:10.1007/978-3-319-40859-0_5. ISBN 978-3-319-40858-3.
  16. ^ Coates, Michael I.; Gess, Robert W. (2007). "A New Reconstruction of Onychoselache Traquairi, Comments on Early Chondrichthyan Pectoral Girdles and Hybodontiform Phylogeny". Palaeontology. 50 (6): 1421–1446. doi:10.1111/j.1475-4983.2007.00719.x.
  17. ^ a b c d e f g h Gess, Robert W.; Whitfield, Alan K. (14 February 2020). "Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent". Biological Reviews. 95 (4): 865–888. doi:10.1111/brv.12590. PMID 32059074. S2CID 211122587.
  18. ^ a b c Gess, Robert; Ahlberg, Per Erik (8 June 2018). "A tetrapod fauna from within the Devonian Antarctic Circle". Science. 360 (6393): 1120–1124. Bibcode:2018Sci...360.1120G. doi:10.1126/science.aaq1645. PMID 29880689.
  19. ^ a b Anderson, Heidi M.; Hiller, Norton; Gess, Robert W. (1 April 1995). "Archaeopteris(Progymnospermopsida) from the Devonian of southern Africa". Botanical Journal of the Linnean Society. 117 (4): 305–320. doi:10.1006/bojl.1995.0021.
  20. ^ a b Gess, Robert W. (December 2013). "The Earliest Record of Terrestrial Animals in Gondwana: A Scorpion from the Famennian (Late Devonian) Witpoort Formation of South Africa". African Invertebrates. 54 (2): 373–379. doi:10.5733/afin.054.0206.
  21. ^ a b Gess, Robert W.; Coates, Michael I.; Rubidge, Bruce S. (October 2006). "A lamprey from the Devonian period of South Africa". Nature. 443 (7114): 981–984. Bibcode:2006Natur.443..981G. doi:10.1038/nature05150. PMID 17066033. S2CID 4302716.
  22. ^ a b c d Gess, Robert W.; Coates, Michael I. (1 October 2015). "Fossil juvenile coelacanths from the Devonian of South Africa shed light on the order of character acquisition in actinistians". Zoological Journal of the Linnean Society. 175 (2): 360–383. doi:10.1111/zoj.12276.
  23. ^ Scholze, Frank; Gess, Robert W. (1 April 2017). "Oldest known naiaditid bivalve from the high-latitude Late Devonian (Famennian) of South Africa offers clues to survival strategies following the Hangenberg mass extinction". Palaeogeography, Palaeoclimatology, Palaeoecology. 471: 31–39. Bibcode:2017PPP...471...31S. doi:10.1016/j.palaeo.2017.01.018.
  24. ^ a b c Long, J. A.; Anderson, M. E.; Gess, R.; Hiller, N. (19 June 1997). "New placoderm fishes from the Late Devonian of South Africa". Journal of Vertebrate Paleontology. 17 (2): 253–268. doi:10.1080/02724634.1997.10010973.
  25. ^ a b Gess, Robert W.; Trinajstic, Kate M.; Smith, Thierry (5 April 2017). "New morphological information on, and species of placoderm fish Africanaspis (Arthrodira, Placodermi) from the Late Devonian of South Africa". PLOS ONE. 12 (4): e0173169. Bibcode:2017PLoSO..1273169G. doi:10.1371/journal.pone.0173169. PMC 5381866. PMID 28379973.
  26. ^ Anderson, M. Eric; Long, John A.; Gess, Robert W.; Hiller, Norton (1999). "An unusual new fossil shark (Pisces: Chondrichthyes) from the Late Devonian of South Africa". Records of the Western Australian Museum. 57: 151–156.
  27. ^ Daeschler, Edward B.; Downs, Jason P. (May 2018). "New description and diagnosis of Hyneria lindae (Sarcopterygii, Tristichopteridae) from the Upper Devonian Catskill Formation in Pennsylvania, U.S.A.". Journal of Vertebrate Paleontology. 38 (3): e1448834. doi:10.1080/02724634.2018.1448834. S2CID 89661336.
  28. ^ Büttner, Steffen H.; Prevec, Stephen A.; Gess, Robert. (PDF). Rhodes University. S2CID 132638533. Archived from the original (PDF) on 2020-02-09.
  29. ^ Gess, Robert W.; Whitfield, Alan K. (August 22, 2020). "Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent". Biological Reviews. 95 (4): 865–888. doi:10.1111/brv.12590. PMID 32059074. S2CID 211122587 – via Wiley Online Library.
  30. ^ a b Gess, Robert W. (1995). "A Preliminary Catalogue of Fossil Algal, Plant, Arthropod, and Fish Remains from a Late Devonian Black Shale Near Grahamstown, South Africa".
  31. ^ a b c d e f Gess, Robert W.; Whitfield, Alan K. (2020). "Estuarine fish and tetrapod evolution: Insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent". Biological Reviews. 95 (4): 865–888. doi:10.1111/brv.12590. PMID 32059074. S2CID 211122587.
  32. ^ a b c d Gess, R.W.; Hiller, N. (1995). "Late Devonian charophytes from the Witteberg Group, South Africa". Review of Palaeobotany and Palynology. 89 (3–4): 417–428. doi:10.1016/0034-6667(95)00007-8.
  33. ^ Gess, R. W.; Hiller, N. (1 December 1995). "Late Devonian charophytes from the Witteberg Group, South Africa". Review of Palaeobotany and Palynology. 89 (3): 417–428. doi:10.1016/0034-6667(95)00007-8.
  34. ^ Prestianni, C.; Gess, R. W. (1 October 2014). "The rooting system of Leptophloeum Dawson: New material from the Upper Devonian, Famennian Witpoort Formation of South Africa". Review of Palaeobotany and Palynology. 209: 35–40. doi:10.1016/j.revpalbo.2014.05.007.
  35. ^ Gess, R. W.; Prestianni, C. (1 February 2018). "Kowieria alveoformis gen. nov. sp. nov., a new heterosporous lycophyte from the Latest Devonian of Southern Africa" (PDF). Review of Palaeobotany and Palynology. 249: 1–8. doi:10.1016/j.revpalbo.2017.10.002.
  36. ^ Prestianni, Cyrille; Gess, Robert W. (28 November 2018). "Rinistachya hilleri gen. et sp. nov. (Sphenophyllales), from the upper Devonian of South Africa" (PDF). Organisms Diversity & Evolution. 19 (1): 1–11. doi:10.1007/s13127-018-0385-3. S2CID 53811583.

January 07, 2023
waterloo, farm, lagerstätte, this, article, lead, section, short, adequately, summarize, points, please, consider, expanding, lead, provide, accessible, overview, important, aspects, article, 2021, famennian, lagerstätte, south, africa, that, constitutes, only. This article s lead section may be too short to adequately summarize the key points Please consider expanding the lead to provide an accessible overview of all important aspects of the article May 2021 The Waterloo Farm lagerstatte is a Famennian lagerstatte in South Africa that constitutes the only known record of a near polar Devonian coastal ecosystem Waterloo Farm in 2016 Contents 1 History and discovery 2 Contributions to palaeontological material 3 Site characteristics 4 Fossils 5 List of published taxa from Waterloo Farm 5 1 Animalia 5 1 1 Invertebrates 5 1 1 1 Mollusca Bivalvia 5 1 1 2 Arthropoda 5 1 1 3 Eurypterida 5 1 1 4 Arachnida Scorpiones 5 1 2 Vertebrata 5 1 2 1 Agnatha 5 1 2 2 Placodermi 5 1 2 3 Acanthodii 5 1 2 4 Chondrichthyes 5 1 2 5 Actinopterygii 5 1 2 6 Sarcopterygii 5 1 2 7 Tetrapoda 5 2 Plantae 5 2 1 Algae 5 2 1 1 Rhodophyta or Phaeophyta 5 2 1 2 Charophyta 5 2 2 Tracheophyta 5 2 2 1 Zosterophyllopsida 5 2 2 2 Lycopsida 5 2 2 3 Iridopteridales 5 2 2 4 Sphenopsida 5 2 2 5 Progymnospermopsida 6 ReferencesHistory and discovery EditThe Waterloo Farm Lagerstatte is an approximately 360 million year old Famennian latest Devonian fossil rich locality of the Witpoort Formation Witteberg Group Cape Supergroup in Makhanda former Grahamstown within the Eastern Cape Province South Africa 1 2 Before it s discovery very little was known of life during the Famennian see Late Devonian extinction in what is now southern Africa This is largely due to the fact that fossils in the Witpoort Formation generally occur in black anaerobically deposited metashale that rapidly degrades near surface and is therefore rarely seen in natural outcrop 3 4 As is the case with many other scientific discoveries the discovery of Waterloo Farm was accidental 5 Uprisings against the apartheid system in South Africa had by the mid 1980s escalated to violent protests particularly in the townships designated black residential areas Prior to that the N2 the main road linking the industrial ports of Port Elizabeth now Gqeberha and East London ran through then Grahamstown including its township 5 By 1985 protests and stoning of cars had made it unsafe for traffic to drive through the townships 5 As a result the local government proposed the construction of a new road that would serve as a bypass around the townships 5 Due to limited options it was decided that the bypass would be constructed through the valleyed countryside to the south of then Grahamstown 5 This involved excavation of extensive road cuttings through hills and spurs of the Rietberg and utilisation of the resultant rubble to build viaducts across the intervening valleys 5 Amongst the small farms from which land was appropriated for the road was Waterloo Farm owned by Rhodes University Botany professor Roy Lubke with one cutting passing virtually through the farm yard 5 At a braai barbeque held by Roy Lubke one afternoon a colleague Dr Mark Aken wandered off to the top of the cutting and peered down the slope He returned to his colleagues to inform them that there were plant fossils in the disturbed black shale Roy Lubke mentioned this to Dr Fred Gess an entomologist who passed the information on to his son Robert Gess As Robert Gess had a great interest in the fossils of the Eastern Cape he immediately made his way out to the roadworks where he was struck by the uniquely well preserved nature of the fossils With the encouragement of his supervisor and mentor Dr Norton Hiller of the Rhodes Geology Department he started assembling a collection of the fossils 5 In 1991 Fiona Taylor an honours student of Dr Hiller conducted a short study on the geological context of the deposit and also published a preliminary note on the fossils including some of those from Gess collection During a short combined excavation by Gess and Taylor more material was found including remains of large armour plated placoderm fish later identified as Africa s only known species of Bothriolepis Taylor s sedimentological work was presented in an article titled Late Devonian shoreline changes an analysis of Witteberg Group stratigraphy in the Grahamstown area published in 1992 Hiller and Taylor 3 In 1993 Hiller employed Robert Gess to conduct a thorough palaeontological study and his excavations continued until 1995 after the emigration of Dr Hiller This phase of research resulted in a number of papers which were the subject of his master s degree 5 4 By the mid 90 s the road cutting was frequently collapsing due to the dip direction of the strata being towards the road 5 This led to a near permanent closure of one lane for safety reasons 5 After there had been a failed attempt to mitigate the situation with steel barriers in the mid nineties the newly established South African National Road Agency Limited SANRAL published a call for tenders to upgrade the road 5 This would prove a propitious opportunity for Robert Gess as by this time he was struggling to secure further funding for excavations and research 5 In September 1999 Gess who had taken up residence in Bathurst 45 km away from Makhanda heard that a group of consulting engineers from Jeffares and Green now known as JG Africa had been awarded the tender to cut back and stabilise the cuttings 5 He immediately contacted the company and SANRAL to inform them of the sensitivity of the site 5 SANRAL made funds available for Gess to rescue some of the shale for scientific research 5 This was hand mined in blocks by Gess and a team under his direction before being transported on a flat bed truck to his property in Bathurst 5 Gess built a shed roof over the sample to protect them from the rapid decay that characterises the shale when exposed to weather 5 It is from meticulous excavation of these recued blocks that most subsequent discoveries have been and continue to be made 5 A similar process during further roadworks in 2007 and 2008 yielded an even larger haul of fossiliferous blocks which are stored in sheds constructed by SANRAL 5 The fossils are housed and researched by the Devonian Ecosystems project funded by the Millennium Trust and South African Centre of Excellence in Palaeosciences in the Albany Museum s Devonian lab at 87 Beaufort Street Makhanda formerly Rhini or Grahamstown They have provided the material for extensive research by Dr Rob Gess and his collaborators and an ongoing stream of significant papers 1 6 7 8 Contributions to palaeontological material EditWaterloo Farm is a globally significant site providing the only record of a high latitude near polar coastal ecosystem overturning numerous assumptions about high latitude conditions during the latest Devonian 9 1 7 Previous sparse evidence from sub Saharan Africa South America and Antarctica had previously led researchers to believe that high latitude conditions precluded extensive vegetation of land and high vertebrate diversity and that for example Devonian tetrapods only occurred within tropical settings 10 Waterloo farm has totally disrupted such beliefs providing evidence for a diversely vegetated adjacent terrestrial habitat with plants including Archaeopteris trees a diverse estuarine vertebrate fauna 1 and the only known non tropical Devonian tetrapods 9 Exceptional soft tissue preservation at Waterloo Farm is unique for a Famennian estuary and allows for reconstruction of an entire estuarine ecosystem 7 grading from brackish to more marine conditions and including delicate waterweeds 11 invertebrates 12 and diverse vertebrates 13 9 As yet 25 species have been diagnosed from Waterloo Farm and many others provisionally identified An excess of 50 organisms are however believed to be represented in the extensive collections of the Devonian Ecosystems Project Together this represents the most holistically studied Devonian tetrapod bearing locality Insights provided by the unique soft tissue preservation at the site are also making major contributions to evolutionary biology such as a growth series of ancient lampreys Priscomyzon riniensis that have overturned major long held perspectives on vertebrate origins 7 Site characteristics EditWitpoort Formation black shales within the Eastern Cape often exhibit cyclical changes in composition which likely reflect potentially seasonal fluctuations in water salinity Water stratification within the estuarine lake frequently led to anoxic bottom waters resulting in episodes of exceptional preservation 14 Witpoort Formation sediments were deeply buried due to continued basinal subsidence through the Carboniferous and were subsequently metamorphosed during the massive Permian aged Cape Fold Belt orogenesis 15 Hundreds of millions of years of erosion and uplift brought the Waterloo Farm shales back up to near surface they were exposed in 1985 in new road cuttings south of Makhanda Grahamstown during construction of a bypass road Waterloo Farm during the excavation in 1985 On site excavations were conducted in the 1990s but the instability of the road cutting led to it being cut back in 1999 and in 2008 On both occasions large quantities of shale were rescued which provides for ongoing excavation Decades of research has revealed the most important Late Devonian fossil site from what was the southern portion of Gondwanan region incorporating present day sub saharan Africa South America and western Antarctica 16 Waterloo Farm in 1988 Waterloo Farm main fossil site in 1999 preceding roadworks One of the sheds of shale blocks rescued with the help of SANRAL from Waterloo Farm roadworks in 2008 for ongoing scientific excavation Because the original fine black mud was often very low in oxygen plants and animals rapidly buried in it sometimes left behind impressions of their soft parts This is extremely rare in the fossil record which normally only preserves bones teeth and other hard bits Exceptionally what is recorded is the remains of an entire estuarine ecosystem from delicate waterweeds and seaweeds to small clams baby fish and the bones of larger fish Land plants which grew nearby are also preserved from the remains of small undergrowth species to fronds from the earliest types of trees 17 Fossils EditMore than 20 species new to science have been named from Waterloo Farm which probably represent about a third of the total number of taxa indicated by remains preserved in the shale Taxa include the tetrapods Tutusius umlambo and Umzantsia amazana 18 which are Africa s earliest known tetrapods and the only non tropical Devonian tetrapods known The first described fossils from Waterloo Farm comprise remains of sub Saharan Africa s earliest woody trees Archaeopteris notosaria Archaeopteris notosaria fronds from the Waterloo Farm lagerstatte in the Eastern Cape South Africa These represent the only high latitude species of Archaeopteris as yet described 19 Other fossils from Waterloo Farm include the oldest known land living animal from Gondwana the scorpion Gondwanascorpio emzantsiensis 20 the oldest fossil lamprey in the world Priscomyzon riniensis 21 and Africa s oldest coelacanth from the world s earliest known coelacanth nursery Serenichthys kowiensis 22 Other species represented include several species of armour plated placoderm fish spiny finned acanthodian fish sharks ray finned actinopterygian fish a range of lobe finned fish bivalves seaweeds charophyte waterweeds and a diverse range of plants List of published taxa from Waterloo Farm EditAnimalia Edit Invertebrates Edit Mollusca Bivalvia Edit Naiadites form Devonica 23 Arthropoda Edit Eurypterida Edit Cyrtoctenid eurypterid indet 17 Arachnida Scorpiones Edit Gondwanascorpio emzantsiensis 20 Vertebrata Edit Agnatha Edit Priscomyzon riniensis 21 Placodermi Edit Bothriolepis africana 24 Groenlandaspis riniensis 24 Africanaspis doryssa 25 24 Africanaspis edmountaini 25 Gen et sp indet Acanthodii Edit Diplacanthus acus 17 Diplacanthus indet 17 acanthodidid indet 17 gyracanthid indet 17 Chondrichthyes Edit Plesioselachus macracantha 26 22 Antarctilamna ultima 22 Actinopterygii Edit gen et sp indet Sarcopterygii Edit Serenichthys kowiensis 22 Hyneria sp 27 28 Rhizodont indet 17 Isityumzi mlomomde gen et sp nov 29 Tetrapoda Edit Tutusius umlambo 18 Umzantsia amazana 18 Plantae Edit Algae Edit Rhodophyta or Phaeophyta Edit Hungerfordia fionae sp nov 30 31 Yeaia africana sp nov 30 31 Charophyta Edit Hexachara setacea gen et sp nov 32 31 Hexachara riniensis gen et sp nov 32 31 Octochara crassa gen et sp nov 32 31 Octochara gracilis gen et sp nov 32 31 Tracheophyta Edit Zosterophyllopsida Edit Zosterophyll indet 33 Lycopsida Edit Leptophloeum rhombicum 34 Kowieria alveoformis 35 Iridopteridales Edit Iridopterlean indet 17 Sphenopsida Edit Rinistachya hilleri 36 Progymnospermopsida Edit Archaeopteris notosaria 19 References Edit a b c d Gess Robert W Whitfield Alan K August 2020 Estuarine fish and tetrapod evolution insights from a Late Devonian Famennian Gondwanan estuarine lake and a southern African Holocene equivalent Biological Reviews 95 4 865 888 doi 10 1111 brv 12590 ISSN 1464 7931 PMID 32059074 S2CID 211122587 Frey Linda Coates Michael Ginter Michal Klug Christian Ginter Michal 2017 07 07 Skeletal remains of Phoebodus politus Newberry 1889 Chondrichthyes Elasmobranchii from a Famennian Konservatlagerstatte in the eastern Anti Atlas Morocco and its ecology Ichthyolith Issues Checiny Poland University of Warsaw 36 doi 10 5167 uzh 138397 S2CID 197550063 a b Hiller amp Taylor Norton amp Fiona 1992 Late Devonian shoreline changes an analysis of Witteberg Group stratigraphy in the Grahamstown area South African Journal of Geology 5 203 214 via AJA a b Wolfgang Gess Robert 2012 05 14 High latitude Gondwanan famennian biodiversity patterns evidence from the South African Witpoort Formation Cape Supergroup Witteberg Group University of the Witwatersrand OCLC 811026675 a b c d e f g h i j k l m n o p q r s t Gess Robert Wolfgang 2002 The palaeoecology of a coastal lagoon of the Witpoort Formation Upper Devonian Famennian in the Eastern Cape Province South Africa University of Fort Hare South Africa 6 11 via Fort Hare University Southa Africa Gess Robert W Coates Michael I Rubidge Bruce S October 2006 A lamprey from the Devonian period of South Africa Nature 443 7114 981 984 Bibcode 2006Natur 443 981G doi 10 1038 nature05150 ISSN 0028 0836 PMID 17066033 S2CID 4302716 a b c d Miyashita Tetsuto Gess Robert W Tietjen Kristen Coates Michael I 2021 03 18 Non ammocoete larvae of Palaeozoic stem lampreys Nature 591 7850 408 412 Bibcode 2021Natur 591 408M doi 10 1038 s41586 021 03305 9 ISSN 0028 0836 PMID 33692547 S2CID 232192889 Harris Christopher Gess Robert W Penn Clarke Cameron Rubidge Bruce S 2021 03 29 Coombs Hill A Late Devonian fossil locality in the Witpoort Formation Witteberg Group South Africa South African Journal of Science 117 3 4 doi 10 17159 sajs 2021 9190 ISSN 1996 7489 S2CID 233689294 a b c Gess Robert Ahlberg Per Erik 2018 06 07 A tetrapod fauna from within the Devonian Antarctic Circle Science 360 6393 1120 1124 Bibcode 2018Sci 360 1120G doi 10 1126 science aaq1645 ISSN 0036 8075 S2CID 46965541 Montes C Cardona A Jaramillo C Pardo A Silva J C Valencia V Ayala C Perez Angel L C Rodriguez Parra L A Ramirez V Nino H 2015 04 10 Middle Miocene closure of the Central American Seaway Science 348 6231 226 229 Bibcode 2015Sci 348 226M doi 10 1126 science aaa2815 ISSN 0036 8075 PMID 25859042 S2CID 206633672 Hiller Norton Gess Robert W March 1996 Marine algal remains from the Upper Devonian of South Africa Review of Palaeobotany and Palynology 91 1 4 143 149 doi 10 1016 0034 6667 95 00062 3 ISSN 0034 6667 Gess Robert W December 2013 The Earliest Record of Terrestrial Animals in Gondwana A Scorpion from the Famennian Late Devonian Witpoort Formation of South Africa African Invertebrates 54 2 373 379 doi 10 5733 afin 054 0206 ISSN 1681 5556 S2CID 6336474 Gess Robert Wolfgang September 2008 6th Annual Meeting Society of Vertebrate Paleontology Cleveland Museum of Natural History 28 2 Gess Robert W 2016 Vertebrate Biostratigraphy of the Witteberg Group and the Devonian Carboniferous Boundary in South Africa Origin and Evolution of the Cape Mountains and Karoo Basin Regional Geology Reviews pp 131 140 doi 10 1007 978 3 319 40859 0 13 ISBN 978 3 319 40858 3 Blewett Scarlett C J Phillips David 2016 An Overview of Cape Fold Belt Geochronology Implications for Sediment Provenance and the Timing of Orogenesis Origin and Evolution of the Cape Mountains and Karoo Basin Regional Geology Reviews pp 45 55 doi 10 1007 978 3 319 40859 0 5 ISBN 978 3 319 40858 3 Coates Michael I Gess Robert W 2007 A New Reconstruction of Onychoselache Traquairi Comments on Early Chondrichthyan Pectoral Girdles and Hybodontiform Phylogeny Palaeontology 50 6 1421 1446 doi 10 1111 j 1475 4983 2007 00719 x a b c d e f g h Gess Robert W Whitfield Alan K 14 February 2020 Estuarine fish and tetrapod evolution insights from a Late Devonian Famennian Gondwanan estuarine lake and a southern African Holocene equivalent Biological Reviews 95 4 865 888 doi 10 1111 brv 12590 PMID 32059074 S2CID 211122587 a b c Gess Robert Ahlberg Per Erik 8 June 2018 A tetrapod fauna from within the Devonian Antarctic Circle Science 360 6393 1120 1124 Bibcode 2018Sci 360 1120G doi 10 1126 science aaq1645 PMID 29880689 a b Anderson Heidi M Hiller Norton Gess Robert W 1 April 1995 Archaeopteris Progymnospermopsida from the Devonian of southern Africa Botanical Journal of the Linnean Society 117 4 305 320 doi 10 1006 bojl 1995 0021 a b Gess Robert W December 2013 The Earliest Record of Terrestrial Animals in Gondwana A Scorpion from the Famennian Late Devonian Witpoort Formation of South Africa African Invertebrates 54 2 373 379 doi 10 5733 afin 054 0206 a b Gess Robert W Coates Michael I Rubidge Bruce S October 2006 A lamprey from the Devonian period of South Africa Nature 443 7114 981 984 Bibcode 2006Natur 443 981G doi 10 1038 nature05150 PMID 17066033 S2CID 4302716 a b c d Gess Robert W Coates Michael I 1 October 2015 Fossil juvenile coelacanths from the Devonian of South Africa shed light on the order of character acquisition in actinistians Zoological Journal of the Linnean Society 175 2 360 383 doi 10 1111 zoj 12276 Scholze Frank Gess Robert W 1 April 2017 Oldest known naiaditid bivalve from the high latitude Late Devonian Famennian of South Africa offers clues to survival strategies following the Hangenberg mass extinction Palaeogeography Palaeoclimatology Palaeoecology 471 31 39 Bibcode 2017PPP 471 31S doi 10 1016 j palaeo 2017 01 018 a b c Long J A Anderson M E Gess R Hiller N 19 June 1997 New placoderm fishes from the Late Devonian of South Africa Journal of Vertebrate Paleontology 17 2 253 268 doi 10 1080 02724634 1997 10010973 a b Gess Robert W Trinajstic Kate M Smith Thierry 5 April 2017 New morphological information on and species of placoderm fish Africanaspis Arthrodira Placodermi from the Late Devonian of South Africa PLOS ONE 12 4 e0173169 Bibcode 2017PLoSO 1273169G doi 10 1371 journal pone 0173169 PMC 5381866 PMID 28379973 Anderson M Eric Long John A Gess Robert W Hiller Norton 1999 An unusual new fossil shark Pisces Chondrichthyes from the Late Devonian of South Africa Records of the Western Australian Museum 57 151 156 Daeschler Edward B Downs Jason P May 2018 New description and diagnosis of Hyneria lindae Sarcopterygii Tristichopteridae from the Upper Devonian Catskill Formation in Pennsylvania U S A Journal of Vertebrate Paleontology 38 3 e1448834 doi 10 1080 02724634 2018 1448834 S2CID 89661336 Buttner Steffen H Prevec Stephen A Gess Robert Field guide to geological sights in the Grahamstown area PDF Rhodes University S2CID 132638533 Archived from the original PDF on 2020 02 09 Gess Robert W Whitfield Alan K August 22 2020 Estuarine fish and tetrapod evolution insights from a Late Devonian Famennian Gondwanan estuarine lake and a southern African Holocene equivalent Biological Reviews 95 4 865 888 doi 10 1111 brv 12590 PMID 32059074 S2CID 211122587 via Wiley Online Library a b Gess Robert W 1995 A Preliminary Catalogue of Fossil Algal Plant Arthropod and Fish Remains from a Late Devonian Black Shale Near Grahamstown South Africa a b c d e f Gess Robert W Whitfield Alan K 2020 Estuarine fish and tetrapod evolution Insights from a Late Devonian Famennian Gondwanan estuarine lake and a southern African Holocene equivalent Biological Reviews 95 4 865 888 doi 10 1111 brv 12590 PMID 32059074 S2CID 211122587 a b c d Gess R W Hiller N 1995 Late Devonian charophytes from the Witteberg Group South Africa Review of Palaeobotany and Palynology 89 3 4 417 428 doi 10 1016 0034 6667 95 00007 8 Gess R W Hiller N 1 December 1995 Late Devonian charophytes from the Witteberg Group South Africa Review of Palaeobotany and Palynology 89 3 417 428 doi 10 1016 0034 6667 95 00007 8 Prestianni C Gess R W 1 October 2014 The rooting system of Leptophloeum Dawson New material from the Upper Devonian Famennian Witpoort Formation of South Africa Review of Palaeobotany and Palynology 209 35 40 doi 10 1016 j revpalbo 2014 05 007 Gess R W Prestianni C 1 February 2018 Kowieria alveoformis gen nov sp nov a new heterosporous lycophyte from the Latest Devonian of Southern Africa PDF Review of Palaeobotany and Palynology 249 1 8 doi 10 1016 j revpalbo 2017 10 002 Prestianni Cyrille Gess Robert W 28 November 2018 Rinistachya hilleri gen et sp nov Sphenophyllales from the upper Devonian of South Africa PDF Organisms Diversity amp Evolution 19 1 1 11 doi 10 1007 s13127 018 0385 3 S2CID 53811583 Retrieved from https en wikipedia org w index php title Waterloo Farm lagerstatte amp oldid 1131280736, wikipedia, wiki, book, books, library,

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