Cenozoic biogeography

The Cenozoic was an important time of global tectonic reorganisation; India was colliding with Asia, there was no land between North and South America creating a link between the Atlantic and Pacific oceans, the Andes mountain range was rising, Africa was colliding with Europe, and Australia with Southeast Asia. All these events created distinct global bioprovinces of flora and fauna, including larger benthic foraminifera.

There are three main global bioprovinces; the American, Tethyan and Indo-Pacific provinces. Each bioprovince has its own unique evolutionary history including speciation and extinction events, and migratory events from other provinces (BouDagher-Fadel, 2018). It is because each province is unique that global temporal range charts of, especially larger benthic, foraminifera are not reliable. Regional biostratigraphy and biogeography of particular provinces should be studied.

During the Neogene global atmospheric and ocean circulation began to settle in to their present day arrangement. However, at the start of the Aquitanian the Tethyan seaway remained open enabling the migration of foraminifera between the proto-Mediterranean Sea and proto-Indian Ocean. During this time lineages such as the miogypsinids and lepidocyclinids that originated in the Americas had migrated into the Tethyan bioprovince by reaching North Africa on ocean currents and debris. Cycloclypeus speciated in the Tethyan province and migrated quickly east towards Southeast Asia but not west towards the Americas. At the end of the Aquitanian, global extinctions were low and only 8% of global larger benthic foraminifera (LBF) became extinct. The Indo-Pacific bioprovince, however, was an exception to this rule and of the global 8%, 43% of the LBF in the region became extinct opening up new ecological niches to be exploited (BouDagher-Fadel, 2018).

The Burdigalian saw much diversification of species of LBF and new genera of Rotalioids within the Tethyan province, but few in the Indo-Pacific. Acervulinids and planorbulinids were common globally, soritids became diverse and nummulitids and lepidocyclinids thrived together in reef and fore-reef environments. Lepidocyclinids and miogypsinids continued their migration from the Americas to the Indo-Pacific via the Tethys by algal rafting and planktonic gametes through the Mediterranean and Arabian coast before spreading fast on ocean currents to Southeast Asia to fill the niches left behind at the end of the Aquitanian (BouDagher-Fadel, 2018).

The end of the Burdigalian saw marked extinction event in which ~30% of global LBFs became extinct. It was most pronounced in the Americas and Tethys where 50% and 40% became extinct respectively. Casualties of this event included the extinction of lepidocyclinids and miogypsinids in the American Province and Cycloclypeus in the Tethys. In the American province this extinction is attributed the eruption of the Colombia River Basalts which may have increased atmospheric, and as a result ocean, CO2 leading to ocean acidification. The extinction of the lepidocyclinids and miogypsinids here also coincides with the extinction of many hermatypic corals. The extinct corals and lepidocyclinids/miogypsinids of the Caribbean and West Atlantic hold on until later in the Indo-Pacific region (BouDagher-Fadel, 2018).

The collision of Africa with Europe in the Serravallian sealed off the eastern proto-Mediterranean Sea and proto-Indian Ocean leading to 60% of global extinctions of LBF at this time occurring in the Tethyan province due to increased salinities (BouDagher-Fadel, 2018). In the Indo-Pacific the lepidocyclinids and miogypsinids finally become extinct at the end of the Serravallian. Cyclocylpeus, which had disappeared from the Mediterranean in the Burdigalian, continued to thrive, and does so to the present day, branching off to form Katacycloclypeus although this form was short-lived with Katacycloclypeus becoming extinct in the middle of the Serravallian.

In the Mid-Late Miocene the partial emergence of the Panamanian Isthmus closed off equatorial Atlantic-Pacific throughflow. This led to a diversification of the Soritoidea in the American province, such as Archais in the Caribbean. Although in the Indo-Pacific, the Soritoidea were dominated by just two genera; Amphisorus and Marginopora.

By the Late Miocene approximately 75% of LBF in the Mediterranean had become extinct due to the Messinian Salinity Crisis (BouDagher-Fadel, 2018). LBF endemic to the Indo-Pacific were not particularly affected and although no new forms emerged, there was a faunal turnover following the extinction of lepidocyclinids and miogypsinids to being dominated by Cycloclypeus, Marginopora, Alveolinella and several other foraminifera.

During the Pliocene, the emergence of new LBFs was absent globally except in the Indo-Pacific which was the only bioprovince in which new species appeared (BouDagher-Fadel, 2018). These were restricted to the western and central tropical Pacific and included the calcarinids such as Calcarina spengleri and Quasirotalia sp., which have preferences towards shallow-water, high-energy environments.

Global climate cooling from the Pliocene to the present day suppressed speciation and diversification so that long-lived and already established foraminifera continued to thrive and adapt. Today, Indo-Pacific sandy shoals are dominated by Calcarina, Baculogypsina, Amphistegina and the foreslopes by Heterostegina, Operculina and Cycloclypeus.

As previously laid out, the global history of foraminiferal migrations and extinctions varies between bioprovinces so that the ‘Indo-Pacific Letter Stages’ (Fig. 1) based purely on LBF were proposed as an alternative to European biostratigraphic stages that were based on mixed molluscan and foraminiferal assemblages. It was noted that molluscan faunas of the Indo-Pacific developed separately from those in Europe so that European stages could not be correlated with confidence to those in Indonesia. In Indonesia foraminifera are more abundant than molluscs and knowledge and recognition of only a few key taxa enabled quick dating of rocks in the field (BouDagher-Fadel, 2018). Therefore the Indo-Pacific Letter Stage (Fig. 1) subdivision of the Cenozoic was first proposed by van der Vlerk and Umbgrove in 1927 and was later refined by Leupold and Van der Vlerk (1931); Adams (1970); Chaproniere (1984); BouDagher-Fadel and Banner (1999). These stages are assemblage zones based on the first and last appearance of key taxa that form a sequence from ‘Ta’ (oldest) to ‘Th’ (youngest; Fig. 1).

The Indo-Pacific Letter Stages are now well-established and their use is widespread. However, they are problematic. Correlation with global chrono- and biostratigraphy is still uncertain, although attempts have been made (Fig. 1). Problems arise from LBF assemblages being strongly facies controlled determined by changes in relative sea-level affecting photic zone, wave base and sediment supply (BouDagher-Fadel, 2018).

Fig. 1. Cenozoic ‘letter stages’ of the Indo-Pacific based on assemblage zones of highlighted benthic foraminifera (Modified from BouDagher-Fadel, 2008)


Adams, C.G. 1970. A reconsideration of the East Indian letter classification of the Tertiary, Bulletin of the British Museum (Natural History). Geology, 19, 87-137

Boudaugher-Fadel, M.K., 2018. Evolution and geological significance of larger benthic foraminifera. UCL Press.

BouDagher-Fadel, M.K. and Banner, F.T. 1999. Revision of the stratigraphic significance of the Oligocene-Miocene ‘Letter Stages’. Revue de Micropaléontologie, 42, 93-97

Chaproniere, G.C.H. 1984.The Neogene larger foraminiferal sequence in the Australian and New Zealand regions, and its relevance to the East Indies letter stage classification. Palaeogeography, Palaeoclimatlogy, Palaeoecology, 46, 25-35

Leupold, W. and van der Vlerk, I.M. 1931. The Tertiary. Leidsche Geologische Mededelingen, 5, 611-648

van der Vlerk, I.M. and Umbgrove, J.H. 1927. Tertiary Gidsforaminifera van Nederlandisch Oost-Indie. Wetenschappelijke Mededelingen van de Dienst van de Mijnbouw in Nederlandsch-Oost Indie, 6, 3-35

Jungle creatures of New Guinea

This post contains pictures of some of the other animals I came across while doing fieldwork in New Guinea. These include amphibians, spiders, insects, lizards, snakes and some marine creatures. I have tried to identify most of what I saw, although many remain a mystery to me. If anyone as any suggestions to what some of my unidentified creatures are, I would be very interested to hear your thoughts.


Possible ground frog (Platymantis sp.)

This Australian green tree frog (Litoria caerulea) hopped on to our balcony one night for a visit.


Spiders are generally known as ‘laba-laba’ in Indonesian. This is a type of Huntsman spider (Heteropoda sp.) that was finding sanctuary on this island in the middle of a puddle.

Possible ?long jawed orb spider (?Leucauge sp.)

Tent web spider (Cyrtophora moluccensis)

Spiny orb-web spider (Gasteracantha fornicata)


Not sure but quite happy on this leaf


This Huntsman spider (Heteropoda venatoria) was looming over me in one bathroom (‘Mandi’) I visited.

These large Golden orb-web spiders (Nephilia pilipes) are very common in the jungle. They spin large webs particularly from one side of a road or river to another to catch any insects that use them as a highway.


Green jumping spider, a member of the Salticidae

Cross spiders (Argiope spp.) creating a stabilimentum

By far the worst animal in the jungle is the ant, known locally as ‘semoot’. These critters bite when threatened, particularly when they crawl into your shirt or trousers and find themselves pressed against your arms, legs or chest.  In some of these pictures an army of ants is carrying the head of a crab along a rock.




Beautiful red dragonfly

This rather curious praying mantis (Hierodula sp. or Rhombodera sp.) took a liking to my camera


A giant stick insect looking like a stick


Giant cricket, or ‘jengkerik’.

Not sure what these two species of butterflies are, any ideas welcomed!


Giant Hercules moth (Coscinocera hercules), see pencil for scale, the wingspan was easily 25cm across.



These playful spinner dolphins (Stenella longirostris) frequently followed our boat

Marine gastropod

Pacific nautilus

Carnivorous pitcher plants, animals that are attracted to their nectar crawl into their funnels and get trapped in a pool of powerful enzymes at the base before being digested by the plant.

Papuan mulch skink (Glaphyromorphus crassicaudus)


Emerald tree skink (Lamprolepis smaragdina)

New Guinea Blue-tailed Emo Skink (Emoia caeruleocauda)

Not sure what this is although it is known locally as a ‘Goural’, possibly a ?Littoral whiptail skink (?Emoia atrocostata)


Definitely a Littoral whiptail skink (Emoia atrocostata)

Crocodile or Papuan monitor (Varanus salvadorii) known locally as ‘Biawek’. Endemic to New Guinea and often found climbing trees or on rock faces.

Biawek tracks

Snakes are common in New Guinea although you have to be incredibly lucky to see one in the wild as they tend to avoid humans. Unfortunately most of the snakes I saw were in captivity like this Boelen’s python (Morelia boelen) which was shedding its skin, apparent from the appearance of a glazed eye.

New Guinea small-eyed snake (Micropechis ikaheka) or ‘Ular putih’ the ‘White snake’. This snake is very dangerous with powerful venom and is often found in Palm Oil plantations.  These images are taken from web (Credit: Paul Freed and Scott Frazier) as the only one I saw was roadkill and I don’t like taking pictures of dead animals.

Strikingly-coloured juvenile and adult Yapen Green Tree Boa (Morelia viridis) or ‘Ular ijo’ the ‘Green snake’

Birds of Jamaica

Jamaica is another place I’ve been fortunate to do fieldwork in. It is a beautiful place and part of the Greater Antilles series of islands in the Western Caribbean. It too has an extraordinary biodiversity of bird species, many endemic to Jamaica or the Caribbean. A good guide to bird identification is Haynes-Sutton et al. (2009) ‘A photographic guide to the birds of Jamaica’. As with my post on the Birds of New Guinea, all pictures are taken with my trusty 14 MP Fujifilm Finepix S4500 with 30x (Wide 24mm) zoom. However I have now invested in an 18MP Canon Eos 100D with 75-300mm zoom for future fieldwork. Let’s start with bitterns, herons and egrets…


Green Heron (Butoroides virescens)


Cattle Egret (Bubulcus ibis)


Great Egret (Ardea alba)


Tricoloured Heron (Egretta tricolor ruficollis), endemic to the Caribbean.


Red-tailed Hawk (Buteo jamaicensis), known locally as a ‘Chicken Hawk’.


Loggerhead Kingbird (Tyrannus caudifasciatus), known locally as a ‘Petchary’. Kingbirds are very aggressive and are known to mob John Crows (see later).


Magnolia Warbler (Dendroica magnolia), possibly a female.


Jamaican Crow (Corvus jamaicensis), known locally as the ‘Jabbering or Jamming Crow’ due to its very vocal nature and loud calls.

Greater Antillean Grackle (Quiscalus niger cassirostris), known locally as a ‘Cling Cling’. This bird is endemic to Jamaica and conspicuous by its bright yellow eye and keeled tail.


Magnificent Frigatebird (Fregata magnificens), known locally as a ‘Man O’War Bird’ or ‘Scissor Tail’ due to its forked tail.


Vervian Hummingbird (Mellisuga minima), known locally as the ‘Little Doctor Bird’. Doctor Birds are the national bird of Jamaica, although this really applies to the Red-billed Streamertail (Trochilus polytmus).

Turkey Vulture (Cathartes aura), known locally as a ‘John Crow’. The John Crow mountains in the east of the island are named after this bird. The term John Crow is also used as an insult in Jamaica, I’m not sure why because these are quite beautiful birds!


American Kestrel (Falco sparverius), known locally as a ‘Sparrow Hawk’ or a ‘Killy-Killy’.


Haynes-Sutton, A., Downer, A. and Sutton, R.L., 2009. A photographic guide to the birds of Jamaica. A&C Black.

The Nutrient Paradox

Why do coral reefs, which are home to a high diversity of life, thrive in such nutrient-deficient (oligotrophic) waters? This ‘nutrient paradox’ is a question that has puzzled scientists since Charles Darwin (1842) observed this phenomenon during his voyage on the HMS Beagle. This is also true of foraminifera where a large number of species are also found in oligotrophic settings. So how do nutrients effect the life cycle and distribution of foraminifera? To answer this, we must first define what we mean by nutrients and how these determine marine trophic zones.

Clear waters of oligotrophic environments (Biak, Indonesia)

Nutrients primarily refers to fixed nitrogen (NH4+, NO2, NO3), phosphate (PO43-), iron (Fe2+, Fe3+) and silicate (SiO32-, SiO44−) ions. These are essential nutrients needed in small amounts by photosynthetic organisms to metabolise their cells and are used for cell growth, maintenance and reproduction (Hallock and Schlager, 1986; Hallock, 2001).

Trophic zones may be defined by the amount of chlorophyll from photosynthetic organisms and nutrients within a given environment. In the following examples the amount of phosphate defines the zone:

Low nutrients – Oligotrophic – 0.0-2.6 μg/l chlorophyll, 0.0-12.0 μg/l phosphate

Moderate nutrients – Mesotrophic – 2.6-20.0 μg/l chlorphyll, 12.0-24.0 μg/l phosphate

High nutrients – Eutrophic – 20.0-56.0 μg/l chlorophyll, 24.0-96.0 μg/l phosphate

Phosphate and iron in large amounts is known to inhibit carbonate crystal growth. Phosphate inhibits the growth of both calcite and aragonite, whereas iron inhibits the growth of calcite but not aragonite (van Langerak et al., 1999). The inhibition of crystal growth by trace amounts of ions is caused by the adsorption of the ions onto crystal growth sites   (van Langerak et al., 1999). This partially explains why some organisms that precipitate a carbonate (calcite or aragonite) skeleton do not flourish in meso- or eutrophic environments where these nutrients are present.

Another reason why nutrient-rich environments do not favour the growth of benthonic photosynthetic organisms is that with increased nutrients there is an increased abundance of phytoplankton (a phytoplankton ‘bloom’) at the near surface. These phytoplankton occur in large numbers as they take up the nutrients causing them to reproduce and multiply rapidly. This causes the surface waters to become murky and the amount of light reaching the seafloor to reduce – bad news for bottom-dwelling photosynthetic organisms. This scenario also occurs where there is significant terrestrial run-off, such as a river delta, that dumps a large amount of fine-grained material into the shallow marine environment causing increased murkiness and reduced light intensity. However, recent discoveries of coral reefs close to the mouths of large rivers near the Amazon and in Indonesia are challenging this idea.

Murky waters of mesotrophic environments (Biak, Indonesia)

A further explanation for the nutrient paradox is role of gross primary productivity (GPP) vs net primary productivity (NPP). Primary productivity is the rate at which energy is fixed into organic carbon, such as simple sugars, during photosynthesis (Hallock, 2001). GPP is a measure of the total amount of energy fixed to organic carbon regardless of whether this is used for respiration or cell growth (Hallock, 2001). NPP is the amount of energy fixed during photosynthesis that is available for cell growth which excludes the amount of energy used in respiration (Hallock, 2001). In reefs, GPP can be very high as nutrient-depleted clear waters with increased light penetration result in increased rates of photosynthesis and respiration, whereas NPP is actually very low (Hallock and Schlager, 1986; Hallock, 2001). Therefore, observations of thriving coral reefs in nutrient deficient waters relate to interpretations of GPP rather than NPP (Hallock and Schlager, 1986; Hallock, 2001).

These rotaliids (Kathina jamaicensis) can tolerate mesotrophic conditions. The micrite in the background of this sample may have been deposited as mud particles suspended in murky waters settled on the sea floor (Maastrichtian, Jamaica)

Most calcareous benthic foraminifera are oligotrophic (favour low nutrients), these include nummulitids, lepidocyclinids and orthophragminids.  This is because they house photosynthetic algal symbionts within their tests which need clear water to photosynthesise. These foraminifera often reach large sizes, with larger size corresponding to lower amounts of nutrients. These foraminifera grow large as a combination of having to provide a home for a thriving community of photosynthetic symbionts and by having a long life cycle to allow them enough time to gather enough of the required amount of scarcely available nutrients to enable them to reproduce. Some rotaliids, however, can tolerate high nutrients and can be found in mesotrophic settings. In mixed carbonate-siliciclastic (MCS) settings of the Mahakam Delta in East Kalimantan, Indonesia, Novak and Renema (2018) found distinct assemblages containing the genera Lepidosemicyclina, Anomalinella and Pseudorotalia which are restricted to MCS environments with fine-grained, muddy substrates. Small miogypsinids are dominant in shallow water settings with high terrigenous input (Novak and Renema, 2018).

The high abundance of thick-walled miliolids in this sample indicates deposition in an oligotrophic setting. This depositional environment is supported by the lack of micrite mud in the background of this sample. Instead, the large amount of spar in this grainstone suggests a high energy setting which would act to wash away any suspended mud from the site of deposition (Eocene, Jamaica)

Most miliolids are also oligotrophic and have thick microgranular walls which protect them from harmful UV radiation in shallow, clear water. However, some miliolids occur in deep water where nutrients are also scarce.  Other agglutinated foraminifera are opportunistic and can tolerate a variety of stressed environments including increased nutrients and salinity. Therefore a fossil assemblage dominated by agglutinated foraminifera may suggest deposition within a mesotrophic setting.   Other organisms that can tolerate high nutrients include bryozoa, echinoids and sponges where silicate is a nutrient for the latter.


Darwin, C.R., 1842. The Structure and Distribution of Coral Reefs: Being the First Part of the Geology of the Voyage of the Beagle, Under the Command of Capt. Fitzroy,… During the Years 1832 to 1836. By Charles Darwin,… Published with the Approval of the Lords Commissioners of Her Majesty’s Treasury. Smith, Elder and Company, 65, Cornhill.

Hallock, P., 2001. Coral reefs, carbonate sediments, nutrients and global change. In: Stanley Jr., G.D. The history and sedimentology of ancient reef systems. p. 388-428

Hallock, P. and Schlager, W., 1986. Nutrient excess and the demise of coral reefs and carbonate platforms. Palaios, pp.389-398.

Novak, V. and Renema, W., 2018. Ecological tolerances of Miocene larger benthic foraminifera from Indonesia. Journal of Asian Earth Sciences, 151, pp.301-323.

van Langerak, E., Beekmans, M.M.H., Beun, J.J., Hamelers, H.V.M. and Lettinga, G., 1999. Influence of phosphate and iron on the extent of calcium carbonate precipitation during anaerobic digestion. Journal of Chemical Technology and Biotechnology, 74(11), pp.1030-1036.


New Guinea: Island of Birds

The main reason I became interested geology and palaeontology was a fascination with life on Earth. I wanted to know how the Earth worked and how life evolved on this planet. As a geologist I am lucky enough to visit some beautiful places as part of my job, one of the benefits of this is being able to see some truly wonderful wildlife.

In 2011 and 2013 I visited New Guinea, the second largest island on Earth.  It is a remarkable place as there are virtually no endemic large land predators. The only large land predators are species of crocodiles that are found in rivers and shoreline (even open marine) environments while in the highlands lives the very rare New Guinea Singing Dog. This is in contrast to the rest of Southeast Asia where there are several species of big cat and in Australia where there are marsupial and placental mammalian predators.  In New Guinea, birds rule. Birds of prey are the largest and most common predator in New Guinea, and there is an extraordinary biodiversity of birds of all types including approximately 320 endemic species. A handy resource for bird identification is Pratt and Beehler’s ‘Birds of New Guinea: 2nd Edition’.


In this post I have collated some pictures of birds I encountered in New Guinea. I have taken the following pictures using a 14 MP Fujifilm Finepix S4500 with 30x (Wide 24mm) zoom. It’s not great quality but the zoom is pretty good and the camera is robust enough to handle the rigours of fieldwork and intense tropical rainfall.

Yellow-faced Myna (Mino dumontii) locally known as ‘Beo’ have bright coloured yellow eye rings and are very vocal birds. These examples were observed in the wild, the last photograph is a captive bird.

Striated heron (Butorides striata papuensis) are common along the coast.

Blyth’s Hornbill (Rhyticeros plicatus) are one of my favourite Papuan birds. They are known locally as ‘Burung Tahun’ translated as the ‘Knowing Bird’ because they are believed to be incredibly wise. These are huge birds and you can hear them approaching by the loud ‘whoompf-whoompf’ sound of their heavy wing beats as they fly between rainforest canopies.

The Southern Cassowary (Casuarius casuarius), known locally as ‘Kasuari’, is the largest bird in New Guinea. It is also extremely dangerous and can easily disembowel a human with its powerful legs and sharp claws. You can appreciate the Cassowary’s non-avian dinosaur ancestry by looking at their feet.


I’m not sure what these are but they have very long forked tails.


Pheasant pigeon (Otidiphaps nobilis)

Gurney’s eagle (Aquila gurneyi) is a medium sized, stocky raptor.

Olive-backed Sunbird (Cinnyris jugularis), female – left, male – right


White-breasted Woodswallow (Artamus leucorynchus)


Greater Black Coucal (Centropus menbeki)?


Large basket-like nests.


Egret (Ardea sp.)

The Brahminy Kite (Haliastur indus), known locally as ‘Elang Laut’ or the ‘Sea Eagle’, is a common raptor found along the coast.

Leucistic Brahminy Kite? Known locally as ‘Elang Putih’ or the ‘White Eagle’.


Frigatebird, probably a Lesser Frigatebird (Fregata ariel), are huge birds and long distance travellers. They are conspicuous by their powerful tapered wings and forked tail.


The New Guinea Harpy Eagle (Harpyopsis novaeguineae) is the largest raptor on the island, capable of preying on large tree kangaroos.

The Sulphur-crested Cockatoo (Cacatua galerita), known locally as ‘Kakatua’, are very noisy and gregarious birds. Often found in large groups and tend to follow you around while screeching loudly.


This is a captive Palm Cockatoo (Probosciger aterrimus), known locally as ‘Kakatua Raja’ the ‘King Cockatoo’.

Rufous-bellied Kookaburra (Dacelo gaudichaud) has a powerful bill for catching large insects and sit patiently on exposed branches waiting for insects to fly by.


Singing Starling (Aplonis cantoroides) with startling blood red iris.

Lesser Bird of Paradise (Paradisaea minor) male (top) and female (below). These are captive examples, however they have a beautiful and distinctive song that can be heard carried through the forest for many miles. Birds of Paradise are known locally as ‘Cenderawasih’ in West Papua and lend their name to the embayment, ‘Cenderawasih Bay’, surrounding the Bird’s Head and Bird’s Neck in western New Guinea.

Rainbow Lorikeet (Trichoglossus haematodus) – left, and Black-capped Lory (Lorius lory) – right


A captive Brown Lory (Calcopsitta duivenbodei), known locally as ‘Nuri coklat’.


Victoria Crowned Pigeon (Goura victoria), known locally as ‘Mambruk’.

Little Tern (Sternula albifrons)

I will upload more pictures of birds from Jamaica in a following post. I will also post some images of reptiles and spiders from New Guinea at a later date.


Pratt, T.K. and Beehler, B.M., 2014. Birds of New Guinea. Princeton University Press.

What are forams and why are they useful?

If you were to think about single-celled organisms, you might imagine that they could only been seen under the microscope. Foraminifera (colloquially called ‘forams’), however, are single-celled organisms that can grow from several millimetres up to the size of small frisbees. The one in the above picture, Assilina exponens (Eocene, Germany), is the same size and shape as a ready salted crisp. The largest recorded foram measured 19cm in diameter, nearly 4 times the size of the Assilina exponens pictured above.

Forams are major rock-forming organisms that contribute 43 million tons of carbonate in the oceans per year (Langer et al., 1997) and make up approximately 20% of the carbonate in the geological record. They are also major components of beach ‘sand’ especially in subtropical environments (you may have seen pictures of the beautiful variety of morphologies in foraminifera that make up beach sand that occasionally do the rounds on social media), even the pyramids of Egypt are built from Nummulitic limestone, like the one below, formed of millions of ‘tests’ of foraminifera from the genus Nummulites.


Nummulites gizehensis in a hand specimen of the Eocene Faumai Limestone from New Guinea
Nummulites gizehensis in a hand specimen of the Eocene Faumai Limestone from New Guinea
Nummulites gizehensis in thin section from the same sample of Eocene Faumai Limestone, New Guinea
Nummulites gizehensis in thin section from the same sample of Eocene Faumai Limestone, New Guinea

These organisms are important predators and prey in the ocean ecosystem, and make up much of the biomass of the seafloor and water column, it is even estimated that most of the seafloor is even covered by the reticulate pseudopodia of these organisms. Yet, these creatures remain almost unknown to most outside of the geological or biological communities and have yet to achieve their deserved star status having been mostly overlooked on prime-time nature documentaries. So what exactly are they? Why are they so large? And why do people like me spend all day looking at them?


The taxonomy of foraminifera is problematic and highly disputed. For now, foraminifera are placed within the Rhizaria a ‘supergroup’ of Protists, one of the ‘5 Kingdoms’, although this kingdom is currently being phased out. What is not disputed is that they are eukaryotic single-celled organisms that resemble amoebae with pseudopodia (stringy, gelatinous appendages that are used for locomotion and collecting food).

Foraminifera first appeared during the time of the ‘Cambrian Explosion’ ~542 Ma, with oldest known foraminifera being Platysolenites sp. To date there are approximately 275,000 recognised species, of these 10,000 are extant with remainder extinct and consigned to the fossil record.

There are many ways to divide forams based on their skeletal structure or mode of life. Foraminifera are often described as being ‘benthic’ – living close to the sediment-water interface, or ‘planktonic’ – living within the water column. Planktonic foraminifera account for only around 50 species of 10,000 species around today. Benthic foraminifera account for the remaining extant species, these are often further subdivided by their size into smaller and larger benthic forams, or according to their test structure.

Foraminifera secrete a shell (generally composed of calcite), which is technically called a ‘test’ because cytoplasm covers the exterior of the shell in a ‘sheath’ of ectoplasm. Foraminiferal tests may be ‘agglutinated’ where they are made from sand grains, shell fragments from other organisms, or generally whatever other detritus the foram can find within the substrate in which it lives that it can stick to itself. Foraminiferal tests may be ‘granular/porcelaneous’ such as those of miliolids that do not let light though, or ‘hyaline/glassy’ in which the orientation of crystals permits light to pass giving a transparent or translucent appearance to the test (see below). Again, foraminifera can be further subdivided and subdivided further still based on the numerous shapes, chambers, and apertural openings etc. of the test.

The differences of the major types of foramiferal test structure in cross section
The differences of the major types of foramiferal test structure in cross section

Many forams are symbiotic and grow chambered tests in which to house their symbionts, it is these that make up the larger benthic foraminifera. As their name suggests it is this group that attain the largest sizes. The first formed chamber of a larger benthic foram is called the ‘proloculus’, it is here that the foram keeps its important cellular material with cytoplasm filling the rest of the test. From the proloculus the foram produces a series of large chambers (sometimes divided into smaller ‘chamberlets’). The cytoplasm in the chambers is linked to the proloculus by a canal-like structure called a ‘foramen’ (from which the foraminifera derive their name). It takes forams such as Heterostegina, Elphidium, and Rotalia 24 hours to grow a new chamber. It then takes them a further 5-7 days to recuperate and muster the strength to add a new chamber (Myers, 1941). Within these chambers the larger benthic foraminifera ‘farm’ their symbionts, often photosynthetic cyanobacteria. Rather counter-intuitively the largest larger benthic foraminifera are found within the most oligotrophic (low nutrient) environments. This is because they must reach a large size in order to house the necessary number of cyanobacteria to keep them alive.

Larger benthic foraminifera live typically from several weeks to a couple of years. Their life span is in part dependent on their reproductive cycle. Foraminifera have a complex life cycle in which they can reproduce sexually or asexually. Asexual reproduction produces a haploid ‘Megalospheric’ morphology via meiosis that contains a single set of chromosomes. Sexual reproduction produces a diploid ‘Microspheric’ morphology that contains a pair of chromosomes (see reproductive cycle figure below). Again rather confusingly, the Megalospheric form is often smaller than the Microspheric form. The terms ‘mega’ and ‘micro’ in this case does not refer to the size of the test but of the proloculus. The Megalospheric morphology has a larger proloculus than the Microspheric morphology.

The simplified life cycle of foraminifera. Foraminifera may reproduce sexually by two haploid cells coming together to form a diploid, or asexually by dividing by either mitosis or meiosis to form two new diploid or haploid cells, respectively
The simplified life cycle of foraminifera. Foraminifera may reproduce sexually by two haploid cells coming together to form a diploid, or asexually by dividing by either mitosis or meiosis to form two new diploid or haploid cells, respectively


Foraminifera are useful for palaeontologists and geologists like me in that they provide environmental information about the rocks in which they are contained were deposited. They are also key index fossils in helping to determine at what time these rocks were laid down. As mentioned previously, many larger benthic forams contain symbiotic cyanobacteria that photosynthesise to survive. Photosynthesis requires sunlight and because of this larger benthic forams are often found within the ‘photic zone’, the zone at which sunlight can still penetrate the water column.

Some smaller benthic foraminifera such as Amphistegina house photosynthetic symbionts adapted to blue light that can penetrate greater water depths of up to 150m, although healthy populations of Amphistegina thrive in water depths between 20-30m. Amphistegina is also commonly associated with sandy sediments in warm, tropical environments with their robust tests able to withstand high energy. Other smaller benthic foraminifera can be found in bathyal environments especially where they are agglutinating and not made of calcite (see CCD below).

Planktonic foraminifera are usually only found in the inner to outer neritic environment, but never within bathyal environments due to the ‘Carbonate Compensation Depth’ (or CCD), a depth below which carbonate is dissolved, typically this boundary is found between 4-5km. Occasionally planktonic foraminifera can be washed in to much shallower environments but these environments should be interpreted based on the greater number of benthic foraminifera than planktonics within the fossil assemblage.

The environmental preferences of some Neogene foraminifera through a transect of a typical carbonate rimmed shelf within 20° latitude either side of the equator.
The environmental preferences of some Neogene foraminifera through a transect of a typical carbonate rimmed shelf within 20° latitude either side of the equator.

Species of larger benthic and planktonic foraminifera have short temporal ranges which makes them perfect index fossils in determining the age of rocks through biostratigraphy. Good index fossils must fulfill several criteria 1) they must be common and easy to find, 2) they must be cosmopolitan (although migrations must be accounted for) so they can be correlated across the globe, 3) they must be easy to identify so they can be spotted in samples with varying degrees of preservation, 4) they must have short evolutionary histories so that one (preferably several) species can be used to define a particular period of time. Many smaller benthic foraminifera are hardy organisms that have long temporal ranges which make them not particularly useful for biostratigraphic purposes but, as previously illustrated, may have palaeoenvrionmental significance.

I will give more details on the principles of biostratigraphy in a later post.


Langer, M. R., Silk, M. T. B., Lipps, J. H. 1997. Global ocean carbonate and carbon dioxide production: The role of reef Foraminifera, Journal of Foraminiferal Research, 27(4), 271–277

Myers, E.H. 1941. Ecologic relationships of larger foraminifera: Report of the Committee on Marine Ecology as related to Paleontology, Division of Geology and Geography, National Research Council Annual Report, Appendix H, 1945.

About Me

My name is Dr. David Gold and I am a geologist specialising in basin analysis, carbonate sedimentology, petrography, and biostratigraphy of larger benthic foraminifera. I have been a full-time geologist since 2008 having pursued a life-long interest in rocks and fossils since I was a child. I have always had a great love for the planet we inhabit, both in terms of earth system processes, extant and extinct life, and how these two interact. I felt the best way to develop my love for Planet Earth was to go to university and find out more about how it worked.

University of Portsmouth 2008-2011

I began studying ‘Palaeobiology & Evolution’ at the University of Portsmouth in 2008. I originally decided to study Palaeontology as when I was a child this was always something that I had wanted to do. I enjoyed learning about the taxonomy and evolutionary histories of many extant and extinct groups of vertebrate and invertebrate organisms. This also led to learning about some of the remarkable adaptations these organisms have through studying their anatomy and the comparative anatomy between organisms is something that still holds my interest to this day.

I soon learned that for fossils to be truly useful a good knowledge of the sediments that house them is required, and thus began my love for sedimentology. In some (most?!) cases the sedimentology of a sample is much more interesting than the fossils it contains. I believe that palaeontology and sedimentology are not mutually exclusive but compliment each other. A good grasp of the environmental preferences of the organisms and the depositional textures/fabrics within the rocks can tell us much about when and where these rocks were deposited, with implications for what the adjacent depositional environments are. Further study of the subsequent diagenetic textures/fabrics of the rock formed after it has been deposited gives insights into what processes the rock has been subjected to as it has been buried, and in some cases brought back to the surface.

I left the University of Portsmouth in 2011 with a First-Class Honours degree, having been awarded the ‘Palaeobiology & Evolution Project Prize’ for my final year dissertation on the ‘Taphonomy and Diagenesis of ammonites from the Whitby Mudstone Formation of the Yorkshire Lias’ which was also nominated for the 2011 BSRG prize in undergraduate sedimentology. I thoroughly enjoyed my time at the University of Portsmouth and would recommend it to any potential student thinking about a career in geoscience. I found the lecturers both knowledgable and inspiring, some of whom continue to support and inspire me to this day.

Southeast Asia Research Group (SEARG) 2011-2014

Having left Portsmouth I joined the Southeast Asia Research Group (SEARG) at Royal Holloway, University of London to work on a PhD project based in the little-explored island wilderness of New Guinea. SEARG have a long history of working in many regions of Southeast Asia (including Indonesia, Malaysia, Myanmar, and the Philippines to name a few) and has contributed significantly to the present day understanding of the history of this tectonically complex region. This is some feat as the research group is small and projects extend across a wide area, and I feel incredibly privileged to have been a part of it. The research group is funded by a consortium of oil & gas companies and much of the work is conducted closely with industry research interests.

The start of my PhD project was a steep learning curve as I was working on basin analysis that required me to have a good understanding of structural geology, something I had neglected at Portsmouth. But I soon found myself enjoying learning new concepts under the expert tutelage of my supervisor and staff in the Earth Sciences Department of Royal Holloway. My project also enabled me to spend a combined total of 6 months in the dense jungles of New Guinea.

Fieldwork in New Guinea is as challenging as it is rewarding. I feel extremely lucky to have been given the opportunity to spend such a long time in such a beautiful place and meeting some incredible animals and people along the way, as well as being given the responsibility to help with the logistics of conducting fieldwork in the area to industrial clients. I will be posting a lot more about fieldwork in New Guinea on this site in the near future.

By the time I had finished at Royal Holloway I had learned a great deal about structural geology, plate tectonics, and basin analysis, as well as furthering my skills in palaeontology and carbonate sedimentology. This contributed to me becoming the ‘well-rounded’ geologist that (I hope!) I am today.  During my time at SEARG I helped established working relationships with the University of Papua  (Universitas Negeri Papua or UNIPA) from academic collaboration with the Royal Botanical Gardens at Kew, and it is rewarding to know that I have left a legacy at SEARG for future and continuing work with UNIPA contributing to the better understanding of the region of West Papua in New Guinea.

I received my PhD from Royal Holloway, University of London in 2014 although I am continuing to publish the results of my research (which I will talk about in future) as part of SEARG and hope to conduct further research in the area in the future. Again, I thoroughly enjoyed my time at Royal Holloway and learned much from people who are some of the best experts in their field and who continue to support my work. I would recommend studying at Royal Holloway to any aspiring future geoscientists as a great place to start their academic career.

2014 to Present

I now work as a biostratigrapher and carbonate sedimentologist at a geoservice company in North Wales. We work with a number of industrial clients to produce reports on samples provided to us from exploration and appraisal wells across the globe. The biostratigraphy of samples we are given can be used to date various formations encountered during the drilling process. The sedimentology tells us further information about the depositional and burial history of the samples and their potential usefulness as source or reservoir rocks. My current work has enabled me to expand my knowledge of different places on the planet at different points in time and I enjoy continuing to learn new things on a weekly basis.



Hello and welcome to my personal blog site http://www.Gold-Geo.com. My name is Dr. David Gold and this site is designed to give readers an insight into life as a professional geoscientist, some thoughts into new and emerging (as well as historical) geological research and techniques, the status of my current research and stories about some of the beautiful places, people and creatures I have been lucky enough to visit.  I hope you find reading this site entertaining, educational and enjoyable!