Waterloo Farm lagerstätte

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Reconstruction of paleoenvironment of Waterloo Farm.
Waterloo Farm in 2016.

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

History and discovery

Waterloo Farm in 1988.
Waterloo Farm main fossil site in 1999 preceding roadworks.
Robert Gess, main researcher of Waterloo Farm Lagerstätte

The Waterloo Farm

Makhanda (former Grahamstown) within the Eastern Cape Province, South Africa.[1][2] Before its 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

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.

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.

Priscomyzon riniensis), that have overturned major, long held, perspectives on vertebrate origins.[7]

Site characteristics

One of the sheds of shale blocks rescued (with the help of SANRAL) from Waterloo Farm roadworks in 2008, for ongoing scientific excavation.
Waterloo Farm during the excavation in 1985.

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

Grahamstown
, during construction of a bypass road. 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] 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 that 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

Serenichthys kowiensis).[21]

Other species represented include several species of armour plated (

charophyte
waterweeds, and a diverse range of plants.

Paleobiota

Animalia

Vertebrata

Agnatha
Agnatha
Genus Species Notes Images
Priscomyzon[20] P. riniensis The oldest known lamprey, known from multiple growth series.[7]
Placodermi
Placodermi
Genus Species Notes Images
Bothriolepis[22] B. africana
paleolatitude
.
Groenlandaspis[22] G. riniensis A
arthrodiran
placoderm. Probably the most common vertebrate in environment.
Africanaspis[23][22] A. doryssa A groenlandaspidid placoderm.
A. edmountaini
Arthrodira indet. At least two taxa are known.[17]
Acanthodii
Acanthodii
Genus Species Notes Images
Diplacanthus[24] D. acus Exceptionally deep-bodied and long-spined
acanthodian. Redescription of genus did not considered it as a species of Diplacanthus.[25]
Gyracanthides[17][26] G. riniensis
Diplacanthidae indet.[17]
Acanthodiformes indet.[17]
Chondrichthyes
Chondrichthyes
Genus Species Notes Images
Plesioselachus[27][28] P. macracantha 0.5–1 m (1 ft 8 in – 3 ft 3 in) long
chondrichthyan
with long spine.
Antarctilamna[28] A. ultima Large-sized
elasmobranch
. Known from multiple skeletal materials and complete juvenile.
Actinopterygii
Actinopterygii
Genus Species Notes Images
Actinopterygii indet.[17]
Sarcopterygii (includes Tetrapoda)
Sarcopterygii
Genus Species Notes Images
Serenichthys[21] S. kowiensis Coelacanth known from complete juvenile specimens.
Hyneria[29] H. udlezinye Up to 2.7 m (8 ft 10 in) long
tristichopterid
.
Rhizodontida indet.[17]
Isityumzi[30] I. mlomomde Lungfish known from parasphenoid bones.
Tutusius[18] T. umlambo Early tetrapod. First known African Devonian tetrapods.
Umzantsia[18] U. amazana

Invertebrates

Mollusca
Mollusca
Genus Species Notes Images
Naiadites[31][32] N. devonicus
bivalve
. Only known species of Naiadites from Devonian.
Brachiopoda
Brachiopoda
Genus Species Notes Images
Dignomia[33] D. sp. Described from newly found site in Witpoort Formation, not exactly from Waterloo Farm
Arthropoda
Arthropoda
Genus Species Notes Images
Hibbertopteridae indet.[17] 1.5 m (4 ft 11 in) long sweep-feeding
stylonurine eurypterid
. Described as "Cyrtoctenid", which is later considered as a growth stage of hibbertopterid.
Gondwanascorpio[19] G. emzantsiensis Partially known scorpion.

Plantae

Algae

Rhodophyta or Phaeophyta
Rhodophyta or Phaeophyta
Genus Species Notes Images
Hungerfordia[34][35] H. fionae Algae with unknown affinity, possibly brown algae.
Yeaia[34][35] Y. africana Brown algae.
Charophyta
Charophyta
Genus Species Notes Images
Hexachara[36][35] H. setacea
Charophyte
algae.
H. riniensis
Octochara[36][35] O. crassa
O. gracilis

Tracheophyta

Uncertain
Uncertain
Genus Species Notes Images
Flabellopteris[37] F. lococannensis Euphyllophyte with uncertain classification.
Zosterophyllopsida
Zosterophyllopsida
Genus Species Notes Images
Zosterophyll indet.[38]
Lycopsida
Lycopsida
Genus Species Notes Images
Colpodexylon[39] C. mergae Lycopsid in the order Protolepidodendrales.
C. pullumpedes
Leptophloeum[40] L. rhombicum A lycophyte.
Kowieria[41] K. alveoformis A
lycopsid
lycophyte.
Iridopteridales
Iridopteridales
Genus Species Notes Images
Iridopteridales indet.[17]
Sphenopsida
Sphenopsida
Genus Species Notes Images
Rinistachya[42] R. hilleri
Sphenopsid
. Only sphenopsid known from the Devonian of Gondwana.
Progymnospermopsida
Sphenopsida
Genus Species Notes Images
Archaeopteris[43] A. notosaria A progymnosperm tree. The only high latitude species of Archaeopteris as yet described.


References

  1. ^ .
  2. .
  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. ^
    OCLC 811026675.{{cite book}}: CS1 maint: multiple names: authors list (link
    )
  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. S2CID 4302716
    .
  7. ^ .
  8. .
  9. ^ .
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  13. ^ Gess, Robert Wolfgang (September 2008). "6th Annual Meeting Society of Vertebrate Paleontology". Cleveland Museum of Natural History. 28: 2.
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  27. ^ 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.
  28. ^
    S2CID 140726995
    .
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  34. ^ 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".
  35. ^
    S2CID 211122587
    .
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