Necrophage

Necrophages can be classified according to their nutritional reliance on carrion and also their level of adaptation to carrion feeding. Animals are described as 'obligate necrophages' if they use carrion as their sole or main food source and depend on carrion for survival or reproduction. The term 'specialists' is also sometimes used in recognition that these animals have traits favoring necrophagy and making other feeding behaviors difficult.{{cite journal |last1=Carrasco-Garcia |first1=R |last2= Barroso|first2=P |last3= Perez-Olivares |first3=J |last4=Montoro |first4=V |last5=Vicente |first5= J|title= Consumption of big game remains by scavengers: a potential risk as regards disease transmission in central Spain |journal= Frontiers in Veterinary Science|date=2018 |volume=5 |issue= |pages=4 |doi=10.3389/fvets.2018.00004|doi-access=free | pmid=29552564|pmc=5840163 }} For example, large wingspans facilitate the energy-efficient gliding vultures need to cover long distances in search of carrion,{{cite journal |last1=DeVault |first1=TL |last2= Rhodes |first2=OE |last3= Shivik |first3=JA|title= Scavenging by vertebrates: behavioral, ecological, and evolutionary perspectives on an important energy transfer pathway in terrestrial ecosystems |journal= Oikos|date=2003 |volume=102 |issue= 2|pages=225–234 |doi= 10.1034/j.1600-0706.2003.12378.x|bibcode=2003Oikos.102..225D }} but reduce the agility needed to kill prey.{{cite journal |last1=Whelan |first1=CJ |last2= Wenny|first2=DG |last3= Marquis |first3=RJ | title= Ecosystem services provided by birds |journal= Annals of the New York Academy of Sciences|date=2008 |volume=1134 |issue=1 |pages=25–60 |doi= 10.1196/annals.1439.003| pmid= 18566089|bibcode=2008NYASA1134...25W }} Animals that eat carrion opportunistically and retain the traits needed to find and consume other food sources are described as 'facultative necrophages' and 'generalists'. Both obligate and facultative necrophages are sometimes sub-classified as 'wet' and 'dry' feeders.{{cite journal |last1=Villet |first1=MH|title= African carrion ecosystems and their insect communities in relation to forensic entomology|journal= Pest Technology|date=2011 |volume=5 |issue=1 |pages=1–15 |url=http://www.globalsciencebooks.info/Online/GSBOnline/images/2011/PT_5(1)/PT_5(1)1-15o.pdf}} These terms differentiate animals feeding on moist, putrefying tissue from animals feeding on desiccated and keratinized tissues.

File:Chrysomya megacephala on baby bird.jpg, a species of blow fly, laying eggs in a dead baby bird.]]

The European bone skipper, Thyreophora cynophila, is an obligately necrophagous fly. It relies on carrion bone marrow in the first stage of its life cycle.{{cite journal |last1=Gu|first1=X|last2=Haelewaters|first2=D |last3=Krawczynski |first3=R|last4=Vanpoucke|first4=S|last5= Wagner|first5=H|last6= Wiegleb|first6=G|title= Carcass ecology – more than just beetles|journal=Entomologische Berichten|date=2014 |volume=74 |issue=1-2|pages=68-74|issn=0013-8827}} Many other types of fly are facultatively necrophagous. Examples commonly found on land include blow flies, flesh flies, muscid flies, ensign flies and thread-horns. Other necrophagous flies, for example black flies and lake flies, are semi-aquatic.{{Cite journal|last1=Oses-Rivera|first1=CA|last2=Tosti-Croce Astesiano|first2=EC|date=2020|title=First report of Rhantus validus Sharp (Coleoptera: Dytiscidae) as necrophage and generator of postmortem artifacts in a human corpse found in an artificial freshwater pond from the Región de La Araucanía, Chile|journal=Revista Chilena de Entomología|volume=46|issue=1|pages=81–86|doi=10.35249/rche.46.1.20.11|s2cid=212861052 |doi-access=free}} Types of carrion fed upon include wildlife,{{Cite journal|last1=Okiwelu|first1=SN|last2=Ikpamii|first2=T|last3=Umeozor|first3=OC|date=2010|title=Arthropods associated with mammalian carcasses in Rivers State, Nigeria.|journal=African Journal of Biomedical Research|volume=11|issue=3|doi=10.4314/ajbr.v11i3.50754 |doi-access=free|hdl=1807/54150|hdl-access=free}}{{cite journal |last1=Al-Mesbah |first1=H |last2=Moffatt |first2=C |last3=El-Azazy |first3=OME |last4=Majeed |first4=QAH |title=The decomposition of rabbit carcasses and associated necrophagous Diptera in Kuwait |journal=Forensic Science International |date=2012 |volume=217 |issue=1–3 |pages=27–31 |doi=10.1016/j.forsciint.2011.09.021 |pmid=22018747 }}{{cite journal |last1=Amat |first1=E |title=Notes on necrophagous flies (Diptera: Calyptratae) associated to fish carrion in Colombian Amazon |journal=Acta Amazonica |date=2010 |volume=40 |issue=2 |pages=397–400 |doi=10.1590/s0044-59672010000200018 |doi-access=free }} livestock and poultry carcasses, slaughterhouse and fishing discards, and human bodies. Necrophagous flies detect these dead bodies and body parts via minute traces of decomposition odor in the air.{{cite journal |last1=Mondor |first1=EB |last2=Tremblay |first2=MN |last3=Tomberlin|first3=JK|last4=Benbow|first4=EM|last5=Tarone|first5=AM|last6=Crippen|first6=TL|date=2012 |title=The ecology of carrion decomposition|journal=Nature Education Knowledge|volume=3 |issue=10 |pages=21 |url=https://www.nature.com/scitable/knowledge/library/the-ecology-of-carrion-decomposition-84118259/}}{{cite journal |last1=Carmo |first1=RFR |last2=Vasconcelos |first2=SD |title=Assemblage of necrophagous Diptera in Atlantic insular environments and response to different levels of human presence |journal=Neotropical Entomology |date=2016 |volume=45 |issue=5 |pages=471–481 |doi=10.1007/s13744-016-0394-x |pmid=27040531 |bibcode=2016NeEnt..45..471C |s2cid=8820120 }}

The diversity and abundance of necrophagous fly species vary geographically and seasonally. For example, Chrysomya species are present in subtropical regions of the USA but are rare in most of Canada. This geographic variation is attributable to factors such as soil type and meteorological conditions, and the effects these have on carrion decomposition. Whether urbanization affects fly species richness is open to dispute.{{cite journal |last1=Hwang |first1=C. |last2=Turner |first2=BD |title=Spatial and temporal variability of necrophagous Diptera from urban to rural areas |journal=Medical and Veterinary Entomology |date=2005 |volume=19 |issue=4 |pages=379–391 |doi=10.1111/j.1365-2915.2005.00583.x |pmid=16336303 |s2cid=10442916 }} Seasonally, many necrophagous fly species are observed in higher abundance in summer, but Thyreophora cynophila is more active in winter.

Flies play a critical role in forensic science as they are often the first insects to discover and colonize human remains. Blow flies can arrive within minutes and begin laying eggs in the nose, mouth and other openings. Because adult flies very rarely deposit eggs in live hosts, the age of the developing fly larvae can be used to estimate time of death. Fly larvae can also provide information regarding cause of death because necrophagous flies deposit their eggs in any open wounds.

Vulture bees are a small group of obligately necrophagous bees in the Trigona genus.{{cite journal |last1=Mateus |first1=S |last2=Noll |first2=FB |title=Predatory behavior in a necrophagous bee Trigona hypogea (Hymenoptera; Apidae, Meliponini) |journal=Naturwissenschaften |date=2004 |volume=91 |issue=2 |pages=94–96 |doi=10.1007/s00114-003-0497-1 |pmid=14991148 |bibcode=2004NW.....91...94M |s2cid=26518321 }} Trigona worker bees play a similar role to worker bees in the Apis genus; however, along with collecting pollen, nectar, and plant resins, Trigona workers also collect carrion.{{cite journal |last1=Camargo |first1=JMF |last2=Roubik |first2=DW |title=Systematics and bionomics of the apoid obligate necrophages: the Trigona hypogea group (Hymenoptera: Apidae; Meliponinae) |journal=Biological Journal of the Linnean Society |date=1991 |volume=44 |issue=1 |pages=13–39 |doi=10.1111/j.1095-8312.1991.tb00604.x }} Although pollen is associated with higher energy value, carrion is preferred by Trigona bees because it is biochemically easier to extract energy from. This dead animal tissue is used as a source of amino acids too.{{cite journal|last1=Roubik |first1=DW|title=Stingless bee (Apidae: Apinae: Meliponini) ecology |journal=Annual Review of Entomology |date=2023 |volume=68 |issue= |pages=231-256|doi=10.1146/annurev-ento-120120-103938 |pmid=3619840|doi-access=free }}

Cerumen pots are utilized by some Trigona species, such as T. necrophaga, as vesicles to store foodstuff.{{cite journal |last1=Serrão |first1=JE |last2=da Cruz-Landim |first2=C |last3=Silva-de-Moraes |first3=RLM |title=Morphological and biochemical analysis of the stored and larval food of an obligate necrophagous bee, Trigona hypogea |journal=Insectes Sociaux |date=1997 |volume=44 |issue=4 |pages=337–345 |doi=10.1007/s000400050055 |s2cid=33884946 |url=http://www.locus.ufv.br/handle/123456789/22326 |url-access=subscription }} The foodstuff of T. necrophaga consists of both honey and carrion from vertebrate carcasses. Ultimately, the stored food is utilized by developing larvae and the worker bee itself as a source of nutrition and energy. Due to the rapid decomposition of carrion, especially in warm temperatures, the bees must efficiently metabolize the carrion to avoid rotten carrion in their cerumen pots.

Trigona hypogea communicate the presence of a valuable carcass through olfactory signals. The bees create an odour trail between their nest and the prospective animal carcass; thus, the bees recruit the other nest members to respond and exploit the corpse's resources rapidly. Additionally, interspecific competition is observed in Trigona hypogea bees. The bees are observed to defend their colonized food item, including but not limited to a monkey, lizard, fish, or snake carcass, from competing necrophages, such as flies.

File:Oiceoptoma thoracicum01.jpg, a species of silphine beetle, feeding on fish carrion.]]

Numerous beetles in the Nicrophorus genus are obligately necrophagous, for example Nicrophorus americanus and N. vespilloides. Many other beetles are facultative necrophages including checkered beetles, dermestid beetles, diving beetles,{{Cite journal|last=Silva-Soares|first=T|date=2019|title=Necrophagy on Rhinella granulosa (Amphibia, Anura, Bufonidae) by the aquatic beetle families Hydrophilidae and Dytiscidae (Insecta, Coleoptera) in Caatinga environment, Northeastern Brazil|url=https://www.biotaxa.org/hn/article/view/43969|journal=Herpetology Notes|volume=12|pages=869–872}} scarab beetles,{{Cite journal|last1=Messas|first1=YF|last2=Souza|first2=HS|last3=Schiffler|first3=G|last4=Sobczak|first4=JF|date=2012|title=First record of necrophagy by Scybalocanthon nigriceps Harold (Coleoptera, Scarabaeidae, Scarabaeinae)|journal=Revista Brasileira de Entomologia|volume=56|issue=2|pages=257–258|doi=10.1590/s0085-56262012005000026 |doi-access=free}} silphine beetles and water scavenger beetles. Types of carrion fed upon include wildlife, livestock and poultry carcasses, livestock viscera and human bodies. Necrophagous beetles locate this carrion using antennal chemoreceptors{{cite journal |last1= Dekeirsschieter|first1=J|last2=Verheggen|first2=F |last3= Lognay|first3=G|last4=Haubruge |first4=E.|title= Large carrion beetles (Coleoptera, Silphidae) in western Europe: a review|journal= Biotechnology, Agronomy, Society and Environment|date=2011 |volume=15 |issue=3|pages=435-447}}{{cite journal |last1=Scott|first1=MP|title=The ecology and behavior of burying beetles |journal=Annual Review of Entomology|date=1998 |volume=43 |issue=1|pages=595-618|doi= 10.1146/annurev.ento.43.1.595}} sensitive to sulfur-containing compounds.{{cite journal |last1= Kalinová |first1=B|last2=Podskalská |first2=H|last3=Růzicka |first3=J|last4=Hoskovec|first4=M|title= Irresistible bouquet of death-how are burying beetles (Coleoptera: Silphidae: Nicrophorus) attracted by carcasses |journal=Naturwissenschaften|date=2009 |volume=96 |issue=8|pages=889-99|doi= 10.1007/s00114-009-0545-6|pmid=19404598}}

N. vespilloides and other burying beetles preferentially feed on small carcasses (e.g. rodents and small birds) as these are easier to transport, clean and conceal from competitors.{{cite journal |last1=Duarte |first1=A|last2=Welch|first2=M|last3=Swannack |first3=C|last4=Wagner|first4=J |last5=Kilner|first5= RM|title= Strategies for managing rival bacterial communities: Lessons from burying beetles |journal=Journal of Animal Ecology|date=2018 |volume=87 |issue=2|pages=414-427|doi=10.1111/1365-2656.12725|pmid= 28682460|pmc=5836980}} Diving beetles, scarab beetles and water scavenger beetles have all been observed feeding on amphibian carrion (e.g. granular toads and tree frogs). The scarab beetle Scybalocanthon nigriceps uses its front legs and clypeus to shape frog carrion into pellets for eventual consumption. Other scarab beetles, for example, Coprophanaeus ensifer, build their burrows near carcasses for easier transportation of carrion pieces to offspring.{{Cite journal |last=Otronen |first=M |date=1988 |title=Intra-and intersexual interactions at breeding burrows in the horned beetle, Coprophanaeus ensifer |url=https://linkinghub.elsevier.com/retrieve/pii/S000334728880157X |journal=Animal Behaviour |language=en |volume=36 |issue=3 |pages=741–748 |doi=10.1016/S0003-3472(88)80157-X|url-access=subscription }}

Beetles that feed on human remains are important in forensic science. Terrestrial beetles such as checkered beetles and dermestid beetles colonize bodies in a predictable sequence and have well-characterized life cycles, so they can sometimes be used to estimate time of death.{{cite journal |last1= Kulshrestha |first1=P|last2= Satpathy |first2=DK|title= Use of beetles in forensic entomology|journal= Forensic Science International|date=2001 |volume=120 |issue=1-2|pages=15-17|doi=10.1016/S0379-0738(01)00410-8|pmid=11457603}} Aquatic beetles are less useful for estimating time of death but can cause physical damage to submerged bodies that must be distinguished from inflicted injuries when determining cause of death. For example, the facultatively necrophagous diving beetle Meridiorhantus validus creates postmortem channels and chambers in human bodies that must be differentiated from antemortem piercing injuries.

Nassa mud snails such as Nassarius festivus and Nassarius clarus scavenge on dead and decaying animal matter in the intertidal zone of eulittoral soft shores.{{cite journal |last1=Morton |first1=B |title=The biology of Hipponix australis (Gastropoda: Hipponicidae) on Nassarius pauperatus (Nassariidae) in Princess Royal Harbour, Western Australia |journal=Journal of Molluscan Studies |date=2001 |volume=67 |issue=3 |pages=247–255 |doi=10.1093/mollus/67.3.247 }} At Shark Bay in Australia, Nassarius clarus feeds on the carrion of fishes and bivalves. In the presence of carrion, the animal's proboscis performs a search reaction followed by a quick onset of feeding. When faced with a competitor, such as a hermit crab, at the site of the carrion, the Nassarius clarus attack the competition to defend their meal. Nassarius clarus are attracted to fish and bivalve carrion to a distance of 26 miles and have a heightened interest in areas where the sand has been disturbed; thus, indicating the potential presence of organic detritus or damaged fauna.

File:20191213_Neophron_percnopterus,_Jor_Beed_Bird_Sanctuary,_Bikaner_0926_8272.jpg) surrounding a mammalian carcass.]]

Many vulture species are obligately necrophagous including the bearded vulture, black vulture, cinereous vulture, Eurasian griffon, Himalayan vulture, king vulture and turkey vulture. Types of carrion fed upon include dead wildlife, livestock, poultry and companion animals, human remains (sky burial), hunting discards, slaughterhouse offal and roadkill. Typically, muscle tissue is consumed,{{Cite journal|last1=Demo|first1=C|last2=Cansi|first2=ER|last3=Kosmann|first3=C|last4=Pujol-Luz|first4=JR|date=2013|title=Vultures and others scavenger vertebrates associated with man-sized pig carcasses: a perspective in forensic taphonomy|journal=Zoologia (Curitiba)|volume=30|issue=5|pages=574–576|doi=10.1590/s1984-46702013000500010 |doi-access=free}} but bearded vultures feed on bones and bone marrow.{{cite web | title=Bearded Vultures — Europe's rarest vulture| website=Vulture Conservation Foundation | date= | url=https://4vultures.org/vultures/bearded-vulture/| access-date=March 28, 2025}} In addition to eating carrion, Egyptian vultures feed on small live animals such as turtles, eggs and rotting fruit.{{Cite journal|last1=Di Vittorio|first1=M|last2=López-López|first2=P|last3=Cortone|first3=G|last4=Luiselli|first4=L|date=2017|title=The diet of the Egyptian vulture (Neophron percnopterus) in Sicily: temporal variation and conservation implications |url=https://wwwphp.obs-banyuls.fr/Viemilieu/index.php/volume-67-2017/67-issue-1/671-article-2.html |journal=Vie et Milieu|volume=67 |issue=1 |pages=7–14 }}{{Cite journal| last1 = Prakash| first1 = V| last2 = Nanjappa| first2 = C|date=1988| title = An instance of active predation by scavenger vulture (Neophron p. ginginianus) on checkered keelback watersnake (Xenochrophis piscator) in Keoladeo National Park, Bharatpur, Rajasthan| journal = The Journal of the Bombay Natural History Society| volume = 85| issue = 2| page = 419| url= https://www.biodiversitylibrary.org/page/48804980}}{{cite web | title=Egyptian vultures — Europe's only globally endangered vulture| website=Vulture Conservation Foundation | date= |url=https://4vultures.org/vultures/egyptian-vulture/| access-date=March 28, 2025}}

Some human activities have had an adverse impact on vultures in Sicily, the Azerbaijan Republic{{cite journal |last1=Karimov |first1=T |title=Main limiting factors affecting biological parameters of necrophage birds |journal=The Journal of V.N.Karazin Kharkiv National University. Series «Biology» |date=2015 |volume=24 |issue=1153 |pages=68–72 |url=https://periodicals.karazin.ua/biology/article/view/4656 }} and other countries.{{cite journal|last1=Oaks |first1=JL|first2=M |last2=Gilbert |first3=MZ | last3=Virani |first4=RT |last4=Watson |first5=CU |last5=Meteyer |first6=BA |last6=Rideout |first7=HL |last7=Shivaprasad |first8=S |last8=Ahmed |first9=MJI |last9=Chaudhry |first10=M |last10=Arshad |first11=S |last11=Mahmood |first12=A |last12=Ali |first13=AA |last13=Khan |name-list-style=amp |title=Diclofenac residues as the cause of vulture population decline in Pakistan |journal=Nature |issue=6975 |pages=630–633 |date=2004 |doi=10.1038/nature02317 |pmid=14745453 |volume=427 |bibcode=2004Natur.427..630O|s2cid=16146840 }} For example, changes in farming practices such as the indoor raising of cattle and incineration or burial of cattle carcasses have reduced food availability for Eurasian griffon vultures. Shootings of birds, removal of nestlings from nests, and drug pollution have also contributed to declines in vulture populations.

{{Main|Forensic entomology}}

Necrophagous flies and beetles play an important role in forensic entomology due to their postmortem colonization of human remains. For example, in homicide cases, forensic medical examiners can sometimes determine the minimum post-mortem interval based on the fly and beetle species present in the body and their development stage.{{cite journal |last1=Oliveira |first1=TC |last2=Vasconcelos |first2=SD |title=Insects (Diptera) associated with cadavers at the Institute of Legal Medicine in Pernambuco, Brazil: Implications for forensic entomology |journal=Forensic Science International |date=2010 |volume=198 |issue=1–3 |pages=97–102 |doi=10.1016/j.forsciint.2010.01.011 |pmid=20181449 }} This is because these insects rarely deposit eggs in live hosts, they colonize bodies in a predictable sequence following death, and information is available on how long it takes different species to reach different stages of development. Because insect arrival and departure time and larval development time are affected by seasonal changes,{{cite journal |last1=Archer |first1=MS |title=Annual variation in arrival and departure times of carrion insects at carcasses: implications for succession studies in forensic entomology |journal=Australian Journal of Zoology |date=2003 |volume=51 |issue=6 |pages=569 |doi=10.1071/zo03053 }} temperature,{{cite journal |last1=Charabidze |first1=D |last2=Hedouin |first2=V |title=Temperature: the weak point of forensic entomology |journal=International Journal of Legal Medicine |date=2019 |volume=133 |issue=2 |pages=633–639 |doi=10.1007/s00414-018-1898-1 |pmid=30043225 |s2cid=51714494 }} moisture levels,{{cite journal | vauthors = Campobasso CP, Di Vella G, Introna F | title = Factors affecting decomposition and Diptera colonization | journal = Forensic Science International | volume = 120 | issue = 1–2 | pages = 18–27 | date = 2001 | pmid = 11457604 | doi = 10.1016/S0379-0738(01)00411-X }} air exposure,{{cite book | vauthors = Goff ML | title = A Fly for the Prosecution. | location = Cambridge, Massachusetts | publisher = Harvard University Press | date = 2000 }} geographical region,{{cite journal |last1=Kulshrestha |first1=P |last2=Satpathy |first2=DK|title=Use of beetles in forensic entomology |journal=Forensic Science International |date=2001 |volume=120 |issue=1–2 |pages=15–17 |doi=10.1016/S0379-0738(01)00410-8 |pmid=11457603 }} and other factors, these must all be carefully considered when estimating minimum post-mortem interval.{{cite journal | vauthors = Amendt J, Campobasso CP, Gaudry E, Reiter C, LeBlanc HN, Hall MJ | title = Best practice in forensic entomology--standards and guidelines | journal = International Journal of Legal Medicine | volume = 121 | issue = 2 | pages = 90–104 | date = 2007 | pmid = 16633812 | doi = 10.1007/s00414-006-0086-x }}

• Thanatophage
• Detritivore
• Decomposer
• Saprotrophic nutrition
• Consumer-resource systems

{{Reflist|2}}

• {{Wiktionary-inline}}

{{Feeding}}

{{modelling ecosystems}}

Category:Eating behaviors

Category:Mycology

Category:Soil biology