Paleoparasitology
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Paleoparasitology (or "palaeoparasitology") is the study of
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s from the past, and their interactions with hosts and vectors; it is a subfield of
Paleontology 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 (roughly 11,700 years before present). It includes the study of fossi ...
, the study of living organisms from the past. Some authors define this term more narrowly, as "Paleoparasitology is the study of parasites in archaeological material." (p. 103) K.J. Reinhard suggests that the term " archaeoparasitology" be applied to "... all parasitological remains excavated from archaeological contexts ... derived from human activity" and that "the term 'paleoparasitology' be applied to studies of nonhuman, paleontological material." (p. 233) This article follows Reinhard's suggestion and discusses the
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
n and animal parasites of non-human animals and plants from the past, while those from humans and our
hominid The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the east ...
ancestors are covered in archaeoparasitology.


Sources of material

The primary sources of paleoparasitological material include
mummified A mummy is a dead human or an animal whose soft tissues and organs have been preserved by either intentional or accidental exposure to chemicals, extreme cold, very low humidity, or lack of air, so that the recovered body does not decay furt ...
tissues,
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...
s (fossilised dung) from mammals or dinosaurs,
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s, and amber inclusions. Hair, skins, and feathers also yield ectoparasite remains. Some archaeological artifacts document the presence of animal parasites. One example is the depiction of what appear to be mites in the ear of a "hyaena-like" animal in a tomb painting from ancient Thebes. Some parasites leave marks or traces ( ichnofossils) on host remains, which persist in the fossil record in the absence of structural remains of the parasite. Parasitic ichnofossils include plant remains which exhibit characteristic signs of parasitic insect infestation, such as
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
s or leaf minesStone, G.N., R.W.J.M. van der Ham, and J.G. Brewer (2008
Fossil oak galls preserve ancient multitrophic interactions
''Proceedings of the Royal Society, Series B. Biological Sciences'' 275(1648):2213-2219.
and certain anomalies seen in invertebrate endoskeletal remains. Plant and animal parasites have been found in samples from a broad spectrum of geological periods, including the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
(samples over 10,000 years old),
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
(over 550,000 years old),
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
(over 44 million years old),
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
(over 100 million years) and even
Lower Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago (m ...
(over 500 million years).


Evidence of parasitism

One of the most daunting tasks involved in studying parasitic relationships from the past is supporting the assertion that the relationship between two organisms is indeed parasitic. Organisms living in "close association" with each other may exhibit one of several different types of
trophic Trophic, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to: * Trophic cascade * Trophic coherence * Trophic egg * Trophic function * Trophic hormone * Trophic level index * Trophic level * Trop ...
relationships, such as
parasitism Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
, mutualism, and
commensalism Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
. Demonstration of true parasitism between existing species typically involves observing the harmful effects of parasites on a presumed host. Experimental infection of the presumed host, followed by recovery of viable parasites from that host also supports any claim of true parasitism. Obviously such experiments are not possible with specimens of extinct organisms found in paleontological contexts. Assumptions of true parasitism in paleontological settings which are based on analogy to known present-day parasitic relationships may not be valid, due to host-specificity. For example, ''
Trypanosoma brucei gambiense ''Trypanosoma brucei'' is a species of parasitic kinetoplastid belonging to the genus ''Trypanosoma'' that is present in sub-Saharan Africa. Unlike other protozoan parasites that normally infect blood and tissue cells, it is exclusively extracellu ...
'' and ''
Trypanosoma brucei rhodesiense ''Trypanosoma brucei'' is a species of parasitic kinetoplastid belonging to the genus ''Trypanosoma'' that is present in sub-Saharan Africa. Unlike other protozoan parasites that normally infect blood and tissue cells, it is exclusively extrace ...
'' are both devastating human parasites, but the related subspecies ''
Trypanosoma brucei brucei ''Trypanosoma brucei'' is a species of parasitic kinetoplastid belonging to the genus ''Trypanosoma'' that is present in sub-Saharan Africa. Unlike other protozoan parasites that normally infect blood and tissue cells, it is exclusively extracellu ...
'' will infect a number of animal hosts, but cannot even survive in the human blood stream, much less reproduce and infect a human host. So a related (or unidentifiable) species of ''Trypanosoma'' found in a paleontological or archaeological context may not be a true human parasite, even though it appears identical (or very similar) to the modern parasitic forms. The most convincing evidence of paleoparasitism is obtained when a presumed parasite is found in direct association with its presumed host, in a context that is consistent with known host-parasite associations. Some examples include
helminth Parasitic worms, also known as helminths, are large macroparasites; adults can generally be seen with the naked eye. Many are intestinal worms that are soil-transmitted and infect the gastrointestinal tract. Other parasitic worms such as schi ...
s caught in amber in the process of escaping from the body of an insect, lice found in the fur of guinea pig mummies, protozoans in the alimentary canal of flies in amber, nematode larvae found embedded in animal coprolites, and a mite caught in amber in the process of apparently feeding on a spider. Fossil organisms which are related to present-day parasites often possess the morphological features associated with a parasitic lifestyle, such as blood-feeding mouthparts. So fossil ticks and
hematophagous Hematophagy (sometimes spelled haematophagy or hematophagia) is the practice by certain animals of feeding on blood (from the Greek words αἷμα ' "blood" and φαγεῖν ' "to eat"). Since blood is a fluid tissue rich in nutritious pro ...
insects are generally assumed to be ectoparasites, even when their remains are found in the absence of a host. The presence of structures resembling leaf miner trails in leaf fossils provide indirect evidence of parasitism, even if remains of the parasite are not recovered. The dramatic tissue aberrations seen in present-day plant
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
s and gall-like structures in some invertebrates are direct physiological reactions to the presence of either
metazoan Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motility, able to move, ca ...
parasites or
microbial A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
pathogens In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
. Similar structures seen in fossil plant and invertebrate remains are often interpreted as evidence of paleoparasitism. Host-parasite interactions today are often exploited by other species, and similar examples have been found in the fossil record of plant galls and leaf mines. For example, there are species of wasps, called
inquilines In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the h ...
, which are unable to induce their own plant galls, so they simply take up residence in the galls that are made by other wasps. Another example is the
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
of plant galls or leaf mines, to eat the trapped insect larva inside the gall or mine.


Knowledge gained from ancient animal and plant parasites

Studies of parasite remains and traces from the past have yielded a vast catalog of ancient host-parasite associations. Genetic sequence data obtained directly from ancient animal parasites, and inferences of past relationships based on genetic sequences of existing parasite groups are also being applied to paleoparasitological questions. Data obtained by all of these methods are constantly improving our understanding of the origin and evolution of the parasites themselves and their vectors, and of the host-parasite and vector-parasite associations. In some cases, presumed host-parasite relationships of the past seem quite different from those known in the present, such as a fly which appears to be a parasite of a
mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
Paleoparasitological studies have also provided insight into questions outside the realm of parasitology. Examples include the migration and phylogeography of marine mammal hosts, the identity of domestic animal bones based on the known hosts of parasite remains found at the site, and the possible role of climatic changes on animal host genetic diversity.


References

{{reflist, 2 Parasitology
Parasitology Parasitology is the study of parasites, their hosts, and the relationship between them. As a biological discipline, the scope of parasitology is not determined by the organism or environment in question but by their way of life. This means it fo ...