Waterloo Farm Lagerstätte
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The Waterloo Farm lagerstätte is a
Famennian The Famennian is the later of two faunal stages in the Late Devonian epoch. The most recent estimate for its duration is that it lasted from around 371.1 to 359.3 million years ago. An earlier 2012 estimate, still used by the International Commis ...
lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
in South Africa that constitutes the only known record of a near-polar
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
coastal ecosystem.


History and discovery

The Waterloo Farm
Lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
is an approximately 360 million year old
Famennian The Famennian is the later of two faunal stages in the Late Devonian epoch. The most recent estimate for its duration is that it lasted from around 371.1 to 359.3 million years ago. An earlier 2012 estimate, still used by the International Commis ...
(latest Devonian) fossil-rich locality of the Witpoort Formation (Witteberg Group, Cape Supergroup) in Makhanda (former Grahamstown) within the
Eastern Cape Province The Eastern Cape ( ; ) is one of the nine provinces of South Africa. Its capital is Bhisho, and its largest city is Gqeberha (Port Elizabeth). Due to its climate and nineteenth-century towns, it is a common location for tourists. It is also kno ...
, South Africa. Before its discovery very little was known of life during the Famennian (see
Late Devonian extinction The Late Devonian mass extinction, also known as the Kellwasser event, was a mass extinction event which occurred around 372 million years ago, at the boundary between the Frasnian and Famennian ages of the Late Devonian period.Racki, 2005McGh ...
) 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. As is the case with many other scientific discoveries, the discovery of Waterloo Farm was accidental. Uprisings against the
apartheid Apartheid ( , especially South African English:  , ; , ) was a system of institutionalised racial segregation that existed in South Africa and South West Africa (now Namibia) from 1948 to the early 1990s. It was characterised by an ...
system in South Africa had by the mid-1980s escalated to violent protests, particularly in the ‘
townships A township is a form of human settlement or administrative subdivision. Its exact definition varies among countries. Although the term is occasionally associated with an urban area, this tends to be an exception to the rule. In Australia, Canad ...
’ (designated black residential areas). Prior to that the N2, the main road linking the industrial ports of
Port Elizabeth Gqeberha ( , ), formerly named Port Elizabeth, and colloquially referred to as P.E., is a major seaport and the most populous city in the Eastern Cape province of South Africa. It is the seat of the Nelson Mandela Bay Metropolitan Municipal ...
(now Gqeberha) and
East London East London is the part of London, England, east of the ancient City of London and north of the River Thames as it begins to widen. East London developed as London Docklands, London's docklands and the primary industrial centre. The expansion of ...
, ran through then Grahamstown, including its township. By 1985, protests and stoning of cars had made it unsafe for traffic to drive through the townships. As a result the local government proposed the construction of a new road that would serve as a bypass around the townships. Due to limited options, it was decided that the bypass would be constructed through the valleyed countryside to the south of then Grahamstown. 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. Amongst the small farms from which land was appropriated for the road was Waterloo Farm, owned by
Rhodes University Rhodes University () is a public research university located in Makhanda (formerly Grahamstown) in the Eastern Cape Province of South Africa. It is one of four universities in the province. Established in 1904, Rhodes University is the prov ...
Botany professor, Roy Lubke - with one cutting passing virtually through the farm yard. 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 Entomology (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
, 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. 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 Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...
) fish, later identified as Africa’s only known species of ''
Bothriolepis ''Bothriolepis'' (from , 'trench' and 'scale') was a widespread, abundant and diverse genus of antiarch placoderms that lived during the Middle to Late Devonian period of the Paleozoic Era. Historically, ''Bothriolepis'' resided in an array ...
''. 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). In 1993, Hiller employed Robert Gess to conduct a thorough
palaeontological Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
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. By the mid-90’s, the road cutting was frequently collapsing due to the dip direction of the strata being towards the road. This led to a near permanent closure of one lane for safety reasons. 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. This would prove a propitious opportunity for Robert Gess as, by this time, he was struggling to secure further funding for excavations and research. 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. He immediately contacted the company and SANRAL to inform them of the sensitivity of the site. SANRAL made funds available for Gess to rescue some of the shale for scientific research. 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. Gess built a shed roof over the sample to protect them from the rapid decay that characterises the shale when exposed to weather. It is from meticulous excavation of these rescued blocks that most subsequent discoveries have been and continue to be made. 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. 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 The Albany Museum, South Africa, situated in Makhanda, South Africa, is affiliated to Rhodes University and dates back to 1855,Chinsamy, Anusuya. (1997). "Albany Museum, Grahamstown, South Africa." ''Encyclopedia of Dinosaurs''. Edited by Philli ...
’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.


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. Previous sparse evidence from
sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
, 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
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s only occurred within tropical settings. 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 and the only known non-tropical Devonian tetrapods. Exceptional soft tissue preservation at Waterloo Farm is unique for a Famennian estuary and allows for reconstruction of an entire estuarine ecosystem, grading from brackish to more marine conditions and including delicate waterweeds, invertebrates and diverse vertebrates.  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 ''Priscomyzon riniensis'' is an extinct lamprey that lived some 360 million years ago during the Famennian (Late Devonian) in a marine or estuarine environment in South Africa. This small agnathan is anatomically similar to the Mazon Creek l ...
''), that have overturned major, long held, perspectives on vertebrate origins.


Site characteristics

Witpoort Formation black shales within the
Eastern Cape The Eastern Cape ( ; ) is one of the nine provinces of South Africa. Its capital is Bhisho, and its largest city is Gqeberha (Port Elizabeth). Due to its climate and nineteenth-century towns, it is a common location for tourists. It is also kno ...
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. Witpoort Formation sediments were deeply buried due to continued basinal subsidence through the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
, and were subsequently metamorphosed during the massive
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
aged
Cape Fold Belt The Cape Fold Belt (CFB) is a long fold-and-thrust mountain belt along the western and southern coastlines of Western Cape, South Africa. The Cape Fold Belt formed during the Permian period (300 to 250million years ago) in the late Paleozoic ...
orogenesis Orogeny () is a mountain-building process that takes place at a convergent plate margin when plate motion compresses the margin. An or develops as the compressed plate crumples and is uplifted to form one or more mountain ranges. This involv ...
. 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 Makhanda, formerly known as Grahamstown, is a town of about 75,000 people in the Eastern Cape province of South Africa. It is situated about northeast of Gqeberha and southwest of East London. It is the largest town in the Makana Local Mun ...
, 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
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n region incorporating present-day sub-Saharan Africa, South America and western Antarctica. 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.


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 In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
indicated by remains preserved in the shale. Taxa include the tetrapods ''
Tutusius ''Tutusius'' is a genus of extinct tetrapod from the Devonian of South Africa, containing a single species, ''Tutusius umlambo''. It was described from the +/- 360 myo Gondwana locality of Waterloo Farm lagerstätte on the south-eastern coast of ...
umlambo'' and ''
Umzantsia ''Umzantsia'' is an extinct genus of limbed stegocephalians ('tetrapods', in the broad sense) from the Devonian of South Africa. It contains a single species, ''Umzantsia amazana''. The genus is based on ~360 million year old skull and shoulder b ...
amazana'', which are Africa's earliest known
tetrapods A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
and the only non-tropical
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
tetrapods known. The first described fossils from Waterloo Farm comprise remains of sub-Saharan Africa's earliest woody trees ('' Archaeopteris notosaria''). Other fossils from Waterloo Farm include the oldest known land-living animal from Gondwana (the scorpion ''
Gondwanascorpio emzantsiensis ''Gondwanascorpio emzantsiensis'' is an extinct Gondwanan scorpion that lived 360 million years ago in the Devonian. Its fossil remains, clearly showing pincer and sting, were discovered in rocks of the Witteberg Group near Grahamstown in South A ...
''), the oldest fossil
lamprey Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
in the world (''
Priscomyzon riniensis ''Priscomyzon riniensis'' is an extinct lamprey that lived some 360 million years ago during the Famennian (Late Devonian) in a marine or estuarine environment in South Africa. This small agnathan is anatomically similar to the Mazon Creek l ...
'') and Africa's oldest coelacanth from the world's earliest known
coelacanth Coelacanths ( ) are an ancient group of lobe-finned fish (Sarcopterygii) in the class Actinistia. As sarcopterygians, they are more closely related to lungfish and tetrapods (the terrestrial vertebrates including living amphibians, reptiles, bi ...
nursery (''
Serenichthys kowiensis ''Serenichthys kowiensis'' is a fossil species of coelacanth described in 2015 from near Grahamstown in South Africa. Some 30 complete specimens of this new species were found in the black shale lagerstätte on Waterloo Farm, preserved by th ...
''). Other species represented include several species of armour plated (
placoderm Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...
) fish, spiny finned (
acanthodian Acanthodii or acanthodians is an extinct class of Gnathostomata, gnathostomes (jawed fishes). They are currently considered to represent a paraphyletic Evolutionary grade, grade of various fish lineages Basal (phylogenetics), basal to extant tax ...
) fish,
sharks Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
,
ray-finned Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin ...
(
actinopterygian Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin ...
) fish, a range of lobe-finned fish,
bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
;
seaweeds Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of ''Rhodophyta'' (red), '' Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
;
charophyte Charophyta () is a paraphyletic group of freshwater green algae, called charophytes (), sometimes treated as a division, yet also as a superdivision. The terrestrial plants, the Embryophyta emerged deep within Charophyta, possibly from terres ...
waterweeds, and a diverse range of plants.


Paleobiota


Animalia


Vertebrata


= Agnatha

=


= Placodermi

=


= Acanthodii

=


= Chondrichthyes

=


= Actinopterygii

=


= Sarcopterygii (includes Tetrapoda)

=


Invertebrates


= Mollusca

=


= Brachiopoda

=


= Arthropoda

=


Plantae


Algae


= Rhodophyta or Phaeophyta

=


= Charophyta

=


Tracheophyta


= Uncertain

=


= Zosterophyllopsida

=


= Lycopsida

=


= Iridopteridales

=


= Sphenopsida

=


= Progymnospermopsida

=


References

{{Geology of South Africa, paleontology Fossils of South Africa Geography of Africa Devonian south paleopolar deposits