![Waterloo Farm in 2016](https://upload.wikimedia.org/wikipedia/commons/1/18/Waterloo_Farm_in_2016.jpg)
The Waterloo Farm lagerstätte is a
Famennian
The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used b ...
lagerstätte
A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
in South Africa that constitutes the only known record of a near-polar
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
coastal ecosystem.
History and discovery
The Waterloo Farm
Lagerstätte
A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
is an approximately 360 million year old
Famennian
The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used b ...
(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 provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha.
The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in 1994 ...
, South Africa.
Before it’s discovery very little was known of life during the Famennian (see
Late Devonian extinction
The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major exti ...
) 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: , ; , "aparthood") was a system of institutionalised racial segregation that existed in South Africa and South West Africa (now Namibia) from 1948 to the early 1990s. Apartheid was ...
system in South Africa had by the mid-1980s escalated to violent protests, particularly in the ‘
townships
A township is a kind of human settlement or administrative subdivision, with its meaning varying in different countries.
Although the term is occasionally associated with an urban area, that tends to be an exception to the rule. In Australia, Ca ...
’ (designated black residential areas). Prior to that the N2, the main road linking the industrial ports of
Port Elizabeth
Gqeberha (), formerly Port Elizabeth and colloquially often 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 Municipality, Sou ...
(now Gqeberha) and
East London
East or Orient is one of the four cardinal directions or points of the compass. It is the opposite direction from west and is the direction from which the Sun rises on the Earth.
Etymology
As in other languages, the word is formed from the f ...
, 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 university, public research university located in Makhanda, Eastern Cape, Makhanda (Grahamstown) in the Eastern Cape Province of South Africa. It is one of four universities in the province. Established in 1904, ...
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 () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
, 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) fish, later identified as Africa’s only known species of ''
Bothriolepis
''Bothriolepis'' (from el, βόθρος , 'trench' and el, λεπίς 'scale') was a widespread, abundant and diverse genus of antiarch placoderms that lived during the Middle Devonian, Middle to Late Devonian period of the Paleozoic Era. Hist ...
''. 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 palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (geology), epoch (roughly 11,700 years before present). It includes t ...
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 recued 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 is situated in Grahamstown in 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 ...
’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, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
, 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
Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
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
Witpoort is a suburb of Midrand in Gauteng, South Africa, just west of Blue Hills. It is located in Region A of the City of Johannesburg Metropolitan Municipality. It was the site of the Battle of Witpoort during the Second Boer War
The S ...
Formation black shales within the
Eastern Cape
The Eastern Cape is one of the provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha.
The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in ...
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 period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
, and were subsequently metamorphosed during the massive
Permian
The Permian ( ) is a geologic period and 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 251.9 Mya. It is the last period of the Paleoz ...
aged
Cape Fold Belt
The Cape Fold Belt is a fold and thrust belt of late Paleozoic age, which affected the sequence of sedimentary rock layers of the Cape Supergroup in the southwestern corner of South Africa. It was originally continuous with the Ventana Mount ...
orogenesis
Orogeny is a mountain building process. An orogeny is an event that takes place at a convergent plate margin when plate motion compresses the margin. An ''orogenic belt'' or ''orogen'' develops as the compressed plate crumples and is uplifted t ...
. 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, also known as Grahamstown, is a town of about 140,000 people in the Eastern Cape province of South Africa. It is situated about northeast of Port Elizabeth and southwest of East London, Eastern Cape, East London. Makhanda is the lar ...
, during construction of a bypass road.
![Waterloo during excavation in 1985](https://upload.wikimedia.org/wikipedia/commons/8/86/Waterloo_during_excavation_in_1985.jpg)
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, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
n region incorporating present-day sub-saharan Africa, South America and western Antarctica.
![Waterloo Farm in 1988](https://upload.wikimedia.org/wikipedia/commons/0/07/Waterloo_Farm_in_1988.jpg)
![Shed Salvaged Road cut](https://upload.wikimedia.org/wikipedia/commons/4/44/Shed_Salvaged_Road_cut.jpg)
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.
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''; plural 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 nam ...
indicated by remains preserved in the shale. Taxa include the tetrapods ''
Tutusius umlambo'' and ''
Umzantsia amazana'',
which are Africa's earliest known
tetrapods
Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsids ...
and the only non-tropical
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
tetrapods known. The first described fossils from Waterloo Farm comprise remains of sub-Saharan Africa's earliest woody trees (''
Archaeopteris notosaria'').
![Archaeopteris (Progymnosperm) twigs and leaves](https://upload.wikimedia.org/wikipedia/commons/4/4f/Archaeopteris_%28Progymnosperm%29_twigs_and_leaves.jpg)
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 ...
''),
the oldest fossil
lamprey
Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
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
The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
nursery ('' Serenichthys kowiensis'').
Other species represented include several species of armour plated ( placoderm) fish, spiny finned (acanthodian
Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, which ...
) fish, sharks
Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...
, ray-finned
Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or hor ...
(actinopterygian
Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or ho ...
) fish, a range of lobe-finned fish, bivalves
Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
; 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 group of freshwater green algae, called charophytes (), sometimes treated as a phylum, division, yet also as a superdivision or an unranked clade. The terrestrial plants, the Embryophyta emerged within Charophyta, possibly fro ...
waterweeds, and a diverse range of plants.
List of published taxa from Waterloo Farm
Animalia
Invertebrates
= Mollusca, Bivalvia
=
'' Naiadites'' form Devonica
= Arthropoda
=
= Eurypterida
=
Cyrtoctenid eurypterid
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 Myr, million yea ...
indet.[
]
= Arachnida, Scorpiones
=
''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 ...
''[
]
Vertebrata
= Agnatha
=
''Priscomyzon
''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 ...
riniensis''
= Placodermi
=
''Bothriolepis
''Bothriolepis'' (from el, βόθρος , 'trench' and el, λεπίς 'scale') was a widespread, abundant and diverse genus of antiarch placoderms that lived during the Middle Devonian, Middle to Late Devonian period of the Paleozoic Era. Hist ...
africana''
''Groenlandaspis
''Groenlandaspis'' is an extinct genus of arthrodire from the Late Devonian. Fossils of the different species are found in late Devonian strata in all continents except eastern Asia. The generic name commemorates the fact that the first specimens ...
riniensis''
''Africanaspis
''Africanaspis'' is an extinct genus of Groenlandaspididae, groenlandaspidid placoderm known from two species, ''Africanaspis doryssa'', named in 1997 from fossils discovered in South Africa and ''Africanaspis edmountaini'', named from fossils de ...
doryssa''
''Africanaspis
''Africanaspis'' is an extinct genus of Groenlandaspididae, groenlandaspidid placoderm known from two species, ''Africanaspis doryssa'', named in 1997 from fossils discovered in South Africa and ''Africanaspis edmountaini'', named from fossils de ...
edmountaini''
Gen. et sp. indet.
= Acanthodii
=
'' Diplacanthus acus''[
'']Diplacanthus
''Diplacanthus'' is an extinct genus of Mid to Late Devonian fish in the class Acanthodii, known as spiny sharks.
Classification
The genus was named by Louis Agassiz in 1843. It was formerly regarded as belonging to the Climatiformes but r ...
indet''[
''acanthodidid indet''][
''gyracanthid indet''][
]
= Chondrichthyes
=
'' Plesioselachus macracantha'' [
'' Antarctilamna ultima''][
]
= Actinopterygii
=
gen. et. sp. indet.
= Sarcopterygii
=
'' Serenichthys kowiensis''[
'']Hyneria
''Hyneria'' is a genus of large prehistoric predatory lobe-finned fish which lived in freshwater during the Devonian period around 360 million years ago.
Etymology
The genus name ''Hyneria'' is a reference to the village of Hyner, Pennsylvan ...
'' sp.
Rhizodont indet.[
'' Isityumzi mlomomde'' (gen. et sp. nov.)
]
= Tetrapoda
=
'' Tutusius umlambo''[
'' Umzantsia amazana''][
]
Plantae
Algae
= Rhodophyta or Phaeophyta
=
'' Hungerfordia fionae'' (sp. nov.)
'' Yeaia africana'' (sp. nov.)
= Charophyta
=
'' Hexachara setacea'' (gen. et sp. nov.)
'' Hexachara riniensis'' (gen. et sp. nov.)
''Octochara
''Octochara'' is a genus of fossil charophyte (aquatic green alga) from the Famennian (Late Devonian). It is one of two genera of charophyte described from the Waterloo Farm lagerstätte in southern Africa. It and ''Hexachara'', from the same loca ...
crassa'' (gen. et sp. nov.)
''Octochara
''Octochara'' is a genus of fossil charophyte (aquatic green alga) from the Famennian (Late Devonian). It is one of two genera of charophyte described from the Waterloo Farm lagerstätte in southern Africa. It and ''Hexachara'', from the same loca ...
gracilis'' (gen. et sp. nov.)
Tracheophyta
= Zosterophyllopsida
=
Zosterophyll
The zosterophylls are a group of extinct land plants that first appeared in the Silurian period. The taxon was first established by Banks in 1968 as the subdivision Zosterophyllophytina; they have since also been treated as the division Zosteroph ...
indet.
= Lycopsida
=
''Leptophloeum rhombicum
''Leptophloeum'' is an extinct genus of vascular plants in the lycophyte clade. It is widely distributed being, known from the Laurasian and Gondwanan settings between the Devonian and Early Carboniferous periods.
Classification
The type speci ...
''
'' Kowieria alveoformis''
= Iridopteridales
=
Iridopterlean indet[
]
= Sphenopsida
=
''Rinistachya hilleri
''Rinistachya'' is a genus of Late Devonian Sphenopsida, sphenopsid, currently containing one species, ''Rinistachya hilleri''. It comprises the only sphenopsid as yet known from the Devonian of Gondwana, and is currently known from only a small n ...
''
= Progymnospermopsida
=
'' Archaeopteris notosaria''
References
{{Geology of South Africa, paleontology
Fossils of South Africa
Geography of Africa