|
[[File:Moofushi Kandu fish.jpg|thumb|300px|right|Un [[Banco (pesci)|banco]] di grossi pesci predatori pelagici ([[Caranx melampygus|caranghi a pinne blu]]) circonda un banco di [[Pesce azzurro|pesci azzurri]] ([[Engraulidae|acciughe]]).]]
{{Tassobox
Si definiscono '''pesci pelagici''' i pesci che vivono nella [[Dominio pelagico|zona pelagica]] degli oceani o dei laghi, ovvero in acque che non sono né vicine al fondo né prossime alla costa, a differenza dei [[Demersale|pesci demersali]], che abitano sul fondo o nelle sue vicinanze, e dei [[pesci di barriera]], associati alle [[Barriera corallina|barriere coralline]].<ref name="Lal8">{{cita libro | autore=Brij V. Lal e Kate Fortune | titolo=The Pacific Islands: An Encyclopedia | url=https://books.google.com/books?id=T5pPpJl8E5wC&pg=PA8 | anno=2000 | editore=University of Hawaii Press | isbn=978-0-8248-2265-1 | p=8}}</ref>
|nome=Giaguaro
|statocons=NT
L'ambiente pelagico marino rappresenta il più grande habitat acquatico sulla Terra, occupando 1370 milioni di chilometri cubi ed ospitando l'11% delle specie di [[Pesce|pesci]] conosciute. Gli [[Oceano|oceani]] hanno una profondità media di 4000 metri. Circa il 98% del volume totale dell'acqua si trova sotto i 100 metri di profondità, e il 75% è situato oltre i 1000 metri.<ref name="Moyle585">{{cid|Moyle e Cech, 2004|p. 585}}.</ref>
|statocons_versione=iucn3.1
|statocons_ref=<ref name=iucn>{{IUCN|summ=15953|autore=Quigley, H., Foster, R., Petracca, L., Payan, E., Salom, R. & Harmsen, B. 2017}}</ref>
I pesci pelagici marini si dividono in due categorie principali: pesci costieri e pesci oceanici. I [[Pesci costieri|pesci pelagici costieri]] vivono nelle acque relativamente poco profonde e illuminate dal sole al di sopra della [[piattaforma continentale]], mentre i [[#Pesci oceanici|pesci pelagici oceanici]] abitano le acque profonde e vaste al di là della piattaforma continentale, sebbene possano anche nuotare vicino alla costa.<ref>{{cita libro | curatore=A. H. McLintock | anno=1966 | url=http://www.TeAra.govt.nz/en/1966/fish-marine/page-2 | capitolo=Pelagic | titolo=Te Ara – The Encyclopaedia of New Zealand | accesso=29 settembre 2022}}</ref><ref>{{cita libro | autore=Carl Walrond | url=http://www.teara.govt.nz/EarthSeaAndSky/SeaLife/OceanicFish/1/en | capitolo=Oceanic fish | titolo=Encyclopedia of New Zealand | accesso=29 settembre 2022}}</ref>
|immagine=[[File:Standing jaguar.jpg|230px]]
|didascalia=
La dimensione dei pesci pelagici varia da piccoli [[Pesce azzurro|pesci azzurri]] costieri, come [[Clupeidae|aringhe]] e [[Engraulidae|sardine]], a grandi [[Superpredatore|predatori apicali]] oceanici, come il [[Thunnus thynnus|tonno rosso]] e gli [[Selachimorpha|squali]] oceanici.<ref name="Lal8"/> Solitamente, sono nuotatori agili con corpi idrodinamici, capaci di percorrere lunghe distanze durante le [[Migrazione ittica|migrazioni]]. Molti pesci pelagici nuotano in [[Banco (pesci)|banchi]] che possono pesare centinaia di tonnellate, mentre altri, come il grande [[Mola mola|pesce luna oceanico]], sono solitari.<ref name="Lal8"/> Esistono anche pesci pelagici d'acqua dolce in alcuni grandi laghi, come la [[Limnothrissa miodon|sardina del Lago Tanganica]].<ref>{{cita web | url=http://pcwww.liv.ac.uk/aquabiol/BIOL468/gtlakes/african_gt_lakes/tanganyika/tanganyika_main.htm | titolo=Lake Tanganyika | sito=pcwww.liv.ac.uk}}</ref>
|dominio=[[Eukaryota]]
|regno=[[Animalia]]
== Pesci epipelagici ==
|sottoregno=
{{Doppia immagine verticale|destra|Bluefin-big.jpg|Herring2.jpg|220|I grandi pesci predatori epipelagici, come questo [[Thunnus thynnus|tonno rosso]], hanno una coda profondamente biforcuta e un corpo liscio affusolato a entrambe le estremità, con una colorazione controombreggiata e tonalità argentate.|I piccoli [[Pesce azzurro|pesci azzurri]] epipelagici, come questa [[Clupea harengus|aringa]], condividono caratteristiche corporee simili a quelle dei pesci predatori descritti sopra.}}
|superdivisione=
I pesci epipelagici abitano la [[Zona eufotica|zona epipelagica]], lo strato più superficiale della [[colonna d'acqua]], che va dal [[livello del mare]] fino a 200 metri di profondità. Questa zona, conosciuta anche come «zona superficiale» o «zona illuminata», include la [[Zona eufotica|zona fotica]]. La zona fotica è definita come lo strato d'acqua in cui la luce solare [[Attenuazione|si riduce]] fino all'1% rispetto alla sua intensità superficiale. La profondità di questa zona varia a seconda della [[Turbidimetria|torbidità]] dell'acqua, ma in acque limpide può raggiungere i 200 metri, coincidenti con la zona epipelagica. La luce presente in questa zona permette al [[fitoplancton]] di svolgere la [[Fotosintesi clorofilliana|fotosintesi]].<ref name="Moyle571"/>
|divisione=
|sottodivisione=
La zona epipelagica costituisce un vasto habitat per la maggior parte dei pesci pelagici. È ben illuminata, favorendo i predatori che fanno affidamento sulla vista, ed è generalmente ben mescolata e ossigenata grazie al movimento delle onde, oltre a offrire un ambiente ideale per la crescita delle [[Alga|alghe]]. Tuttavia, questa zona presenta una scarsa varietà di habitat, il che si traduce in una limitata diversità di specie, supportando meno del 2% delle specie ittiche conosciute nel mondo. Inoltre, molte aree di questa zona sono povere di nutrienti necessari per sostenere la vita dei pesci. Per questo motivo, i pesci epipelagici tendono a concentrarsi nelle acque costiere sopra le [[Piattaforma continentale|piattaforme continentali]], dove i nutrienti arrivano tramite il [[ruscellamento]], o nelle zone oceaniche in cui i movimenti di [[Upwelling|risalita]] portano nutrienti in superficie.<ref name="Moyle571">{{cid|Moyle e Cech, 2004|p. 571}}.</ref>
|superphylum=
|phylum=[[Chordata]]
I pesci epipelagici si dividono principalmente in piccoli [[Pesce azzurro|pesci azzurri]] e grandi predatori che si nutrono di essi. I pesci azzurri si riuniscono in [[Banco (pesci)|banchi]] e filtrano il [[plancton]]. La maggior parte dei pesci epipelagici ha corpi idrodinamici, progettati per sostenere lunghe [[Migrazione ittica|migrazioni]]. Predatori e pesci azzurri condividono spesso caratteristiche [[Morfologia (biologia)|morfologiche]] simili. I predatori, di solito fusiformi, possiedono grandi bocche, corpi lisci e code profondamente biforcute. Molti usano la vista per cacciare zooplancton o pesci più piccoli, mentre altri filtrano il plancton dall'acqua.
|subphylum=
[[File:Herring Silvering.svg|thumb|160px|left|Le scaglie riflettenti dell'[[Clupeidae|aringa]] sono quasi verticali per garantire un mimetismo laterale.]]
|infraphylum=
La maggior parte dei pesci predatori epipelagici e delle loro prede più piccole presentano una colorazione controombreggiante, con toni argentei che riducono la visibilità [[Riflessione diffusa|disperdendo]] la luce incidente.<ref name="Moyle571"/> Questo effetto argentato è dovuto a delle [[Scaglia|scaglie]] riflettenti che agiscono come piccoli specchi, creando un effetto di trasparenza. A profondità intermedie, dove la luce proviene dall'alto, questi «specchi» orientati verticalmente rendono i pesci invisibili se visti di lato.<ref name=Herring2002>{{cita libro | autore=Peter Herring | anno=2002 | titolo=The Biology of the Deep Ocean | pp=192-95 | editore=[[Oxford University Press]] | isbn=978-0-19-854956-7}}</ref>
|microphylum=
|nanophylum=
Nelle acque più superficiali, le scaglie devono riflettere una gamma diversa di lunghezze d'onda, e per raggiungere questo scopo sono dotate di cristalli distanziati variabilmente. Nei pesci dal corpo arrotondato, le scaglie riflettenti sono costituite da molteplici piccoli riflettori orientati verticalmente per mantenere l'effetto specchio.<ref name=Herring2002/>
|superclasse=
|classe=[[Mammalia]]
Nonostante il numero limitato di specie, i pesci epipelagici sono estremamente abbondanti. Ciò che manca in diversità è compensato dalla quantità. I pesci azzurri formano banchi enormi, e i grandi predatori che se ne nutrono sono spesso molto ricercati nell'[[Pesce (alimento)|industria alimentare]]. Nell'insieme, i pesci epipelagici rappresentano la risorsa ittica più importante del mondo.<ref name="Moyle571"/>
|sottoclasse=
|infraclasse=
Molti pesci azzurri sono predatori facoltativi, in grado di catturare singoli [[Copepoda|copepodi]] o avannotti dalla colonna d'acqua, ma anche di passare a un'alimentazione per filtrazione di [[fitoplancton]] quando ciò risulta più vantaggioso in termini energetici. I pesci che si nutrono filtrando utilizzano spesso branchiospine lunghe e sottili per trattenere piccoli organismi dall'acqua. Alcuni dei più grandi pesci epipelagici, come lo [[Cetorhinus maximus|squalo elefante]] e lo [[Rhincodon typus|squalo balena]], sono filtratori, così come alcuni dei più piccoli, come gli [[Sprattus|spratti]] adulti e le [[Engraulidae|acciughe]].<ref name="Moyle572">{{cid|Moyle e Cech, 2004|p. 572}}.</ref>
|superordine=
|ordine=[[Carnivora]]
Le acque oceaniche particolarmente limpide contengono poco cibo. Le aree ad alta produttività tendono a essere leggermente torbide a causa delle [[Fioritura algale|fioriture di plancton]], che attirano i pesci filtratori, seguiti dai predatori più grandi. La pesca del tonno è spesso ottimale in acque con una torbidità tale che un [[disco di Secchi]] sia visibile tra 15 e 35 metri di profondità durante una giornata di sole.<ref>{{cita pubblicazione | autore=Blackburn | anno=1965 | titolo=Oceanography and the ecology of tunas | rivista=Oceanography and Marine Biology: An Annual Review | volume=3 | pp=299-322}}</ref>
|sottordine=[[Feliformia]]
|infraordine=
===Floating objects===
|superfamiglia=
{{multiple image
|famiglia=[[Felidae]]
| align = left
|sottofamiglia=[[Pantherinae]]
| direction = vertical
|tribù=
| width = 220
|sottotribù=
| image1 = Sargassum weeds closeup.jpg
|genere=[[Panthera]]
| alt1 =
|genereautore=
| caption1 = Drifting ''[[Sargassum]]'' seaweed provides food and shelter for small epipelagic fish. The small round spheres are floats filled with carbon dioxide which provide buoyancy to the algae.
|sottogenere=
| image2 = Great Barracuda off the Netherland Antilles.jpg
|specie='''P. onca'''
| alt2 =
|sottospecie=
| caption2 = [[Great barracuda]] accompanied by a school of [[Carangidae|jacks]]
|biautore=([[Linneo|Linnaeus]]
}}
|binome=Panthera onca
{{multiple image
|bidata=[[1758]])
| align = right
|triautore=
| direction = vertical
|trinome=
| width = 220
|tridata=
| image1 = Lines of sargassum Sargasso Sea.jpg
|sinonimi=''Felis onca''<br/>{{zoo|Linnaeus|1758}}
| alt1 =
|nomicomuni=
| caption1 = Lines of ''Sargassum'' can stretch for miles along the ocean surface.
|suddivisione=[[Areale]]
| image2 = Histrio histrio by A. H. Baldwin.jpg
|suddivisione_testo=[[File:Panthera onca distribution.svg|230px]]<br/>{{legenda|red|areale attuale}}{{legenda|pink|areale storico}}
| alt2 =
| caption2 = The camouflaged [[sargassum fish]] has evolved to live among drifting ''Sargassum'' seaweed.
}}
Il '''giaguaro''' ('''''Panthera onca''''') è una grossa [[specie]] di [[Felidae|felino]], unico rappresentante [[Neontologia|vivente]] del [[Genere (tassonomia)|genere]] ''[[Panthera]]'' originario delle [[America|Americhe]]. Con una lunghezza (coda esclusa) che può raggiungere i 185 cm e un peso che può raggiungere i 158 kg, è il più grande felino delle Americhe e il terzo più grande del mondo. Il suo manto caratteristico presenta una colorazione di fondo dal giallo pallido al marroncino ricoperta da macchie che si dispongono a formare rosette sui fianchi, sebbene alcuni esemplari [[Melanismo|melanici]] presentino un manto interamente nero. Il potente morso del giaguaro gli consente di perforare il [[Guscio di tartaruga|carapace]] di [[Testudines|tartarughe]] e [[Testudinidae|testuggini]] e di utilizzare un insolito modo per uccidere le [[Predazione|prede]], mordendo direttamente il cranio dei [[Mammalia|mammiferi]] tra le orecchie per sferrare un colpo fatale al cervello.
Epipelagic fish are fascinated by floating objects. They aggregate in considerable numbers around objects such as drifting flotsam, rafts, jellyfish, and floating seaweed. The objects appear to provide a "visual stimulus in an optical void".<ref>{{cite journal|author=Hunter, JR |author2=Mitchell CT |year=1966|title=Association of fishes with flotsam in the offshore waters of Central America|journal=Fishery Bulletin|volume=66|pages=13–29}}</ref> Floating objects may offer [[refuge (ecology)|refuge]] for [[juvenile fish]] from predators. An abundance of drifting seaweed or jellyfish can result in significant increases in the survival rates of some juvenile species.<ref>{{cite journal|author=Kingsford MJ |year=1993|title=Biotic and abiotic structure in the pelagic environment: Importance to small fishes |journal=Bulletin of Marine Science |url=https://www.researchgate.net/publication/233550840|volume=53|issue=2|pages=393–415}}</ref>
Gli antenati del moderno giaguaro probabilmente raggiunsero le Americhe dall'[[Eurasia]] durante il [[Pleistocene inferiore]] attraverso il [[Ponte continentale|ponte di terra]] che un tempo attraversava lo [[stretto di Bering]]. Attualmente, l'areale della specie si estende dagli [[Stati Uniti sud-occidentali|Stati Uniti sud-occidentali]] al [[Paraguay]] e all'[[Argentina]] settentrionale, attraverso il [[Messico]], gran parte dell'[[America centrale]] e la [[Amazzonia|foresta amazzonica]]. Abita una gran varietà di terreni forestali e aperti, ma predilige [[habitat]] costituiti da [[Foresta pluviale tropicale|foreste pluviali di latifoglie tropicali e subtropicali]], [[Zona umida|zone umide]] e aree boschive. È un eccellente nuotatore ed è per lo più un [[superpredatore]] solitario e opportunista che tende imboscate alle prede. Si tratta di una [[specie chiave]], che svolge un ruolo importante nella stabilizzazione degli [[Ecosistema|ecosistemi]] e nella regolazione delle popolazioni di prede.
Many coastal juveniles use seaweed for the shelter and the food that is available from invertebrates and other fish associated with it. Drifting seaweed, particularly the pelagic ''[[Sargassum]]'', provide a niche habitat with its own shelter and food, and even supports its own unique fauna, such as the [[sargassum fish]].<ref name="Moyle572"/> One study, off Florida, found 54 species from 23 families living in flotsam from ''Sargassum'' mats.<ref>{{cite journal|author=Dooley JK |year=1972|title=Fishes associated with the pelagic sargassum complex, with a discussion of the sargassum community|journal=Contributions in Marine Science|volume=16|pages=1–32}}</ref> Jellyfish also are used by juvenile fish for shelter and food, even though jellyfish can prey on small fish.<ref name="Moyle576">[[#Moyle|Moyle and Cech]], p. 576</ref>
Il giaguaro è minacciato dalla [[Distruzione dell'habitat|distruzione]] e dalla [[Frammentazione ambientale|frammentazione dell'habitat]], dal [[bracconaggio]] per il commercio illegale di alcune parti del corpo e dalle uccisioni in situazioni di conflitto tra uomini e fauna selvatica, soprattutto ad opera degli [[Ranch|allevatori]] dell'America centrale e [[America meridionale|meridionale]]. Figura tra le «[[Specie prossima alla minaccia|specie prossime alla minaccia]]» (''Near Threatened'') sulla [[Lista rossa IUCN|lista rossa della IUCN]] dal 2002. Si ritiene che la popolazione sia diminuita a partire dalla fine degli anni '90. Gli studiosi hanno individuato 51 aree prioritarie per la conservazione della specie (indicate come JCU, Jaguar Conservation Units): queste, dislocate in 36 distinte regioni geografiche che vanno dal Messico all'Argentina, sono vaste aree in cui vivono almeno 50 esemplari in età riproduttiva.
Mobile oceanic species such as [[tuna]] can be captured by travelling long distances in large [[fishing vessel]]s. A simpler alternative is to leverage off the fascination fish have with floating objects. When fishermen use such objects, they are called [[fish aggregating device]]s (FADs). FADs are anchored rafts or objects of any type, floating on the surface or just below it. Fishermen in the Pacific and Indian oceans set up floating FADs, assembled from all sorts of debris, around tropical islands, and then use [[purse seine]]s to capture the fish attracted to them.<ref name="Moyle574/5" />
Il giaguaro ebbe un posto di rilievo nella mitologia delle [[Nativi americani|popolazioni indigene delle Americhe]], comprese le civiltà [[Aztechi|azteche]] e [[maya]].
A study using [[Fisheries acoustics|sonar]] in French Polynesia, found large shoals of juvenile [[bigeye tuna]] and [[yellowfin tuna]] aggregated closest to the devices, 10 to 50 m. Farther out, 50 to 150 m, was a less dense group of larger yellowfin and [[albacore tuna]]. Yet farther out, to 500 m, was a dispersed group of various large adult tuna. The distribution and density of these groups was variable and overlapped. The FADs also were used by other fish, and the aggregations dispersed when it was dark.<ref>{{Cite journal | doi = 10.1016/S0990-7440(00)00051-6| title = Typologie et comportement des agrégations thonières autour de dispositifs de concentration de poissons à partir de prospections acoustiques en Polynésie française| journal = Aquatic Living Resources| volume = 13| issue = 4| pages = 183–192| year = 2000| last1 = Josse | first1 = E. }}</ref>
== Etimologia ==
Probabilmente la parola «giaguaro» deriva del termine [[Lingue tupi-guaraní|tupi-guaraní]] ''yaguara'', che significa «bestia selvatica che abbatte la preda con un balzo».<ref name=Guggisberg1975>{{cita libro | autore=C. A. W. Guggisberg | titolo=Wild Cats of the World | anno=1975 | editore=Taplinger Publishing | città=New York | isbn=978-0-8008-8324-9 | capitolo=Jaguar ''Panthera onca'' (Linnaeus, 1758) | pp=247-265}}</ref><ref name=Seymour/> I popoli indigeni della Guyana lo chiamano ''jaguareté''.<ref>{{cita libro | autore=[[Jean-Baptiste Labat|J. B. Labat]] | anno=1731 | capitolo=Once, espèce de Tigre | titolo=Voyage du chevalier Des Marchais en Guinée, isles voisines, et à Cayenne, fait en 1725, 1726 & 1727 | città=Amsterdam | editore=La Compagnie | volume=III | p=285 | url=https://books.google.com/books?id=dfdWAAAAcAAJ&pg=RA6-PA5 | accesso=13 agosto 2020 | urlarchivio=https://web.archive.org/web/20211129021340/https://books.google.com/books?id=dfdWAAAAcAAJ&pg=RA6-PA5 | urlmorto=no}}</ref> L'appellativo specifico ''onca'' deriva dalla parola portoghese ''onça'', che indicava un felino maculato del Brasile non meglio identificato di dimensioni superiori a quelle di una [[Lynx (zoologia)|lince]] (cfr. l'epiteto specifico ''uncia'' del [[Panthera uncia|leopardo delle nevi]].<ref>{{cita libro | autore=[[John Ray|J. Ray]] | anno=1693 | capitolo=Pardus an Lynx brasiliensis, ''Jaguara'' | titolo=Synopsis Methodica Animalium Quadrupedum et Serpentini Generis. Vulgarium Notas Characteristicas, Rariorum Descriptiones integras exhibens | editore=S. Smith & B. Walford | città=Londra | url=https://archive.org/details/synopsismethodic00rayj/page/168/mode/1up | p=168}}</ref> ''Panthera'', invece, è la forma [[Lingua latina|latinizzata]] del [[Lingua greca antica|greco antico]] ''pánthēr'' (πάνθηρ).<ref>{{cita libro | autore=H. G. Liddell e R. Scott | anno=1940 | capitolo=el:πάνθηρ | titolo=A Greek-English Lexicon | edizione=aggiornata | città=Oxford | editore=Clarendon Press | url=https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 | accesso=20 febbraio 2021 | urlarchivio =https://web.archive.org/web/20080411203109/http://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 | urlmorto=no}}</ref>
Larger fish, even predator fish such as the [[great barracuda]], often attract a retinue of small fish that accompany them in a strategically safe way. [[Scuba diving|Skindivers]] who remain for long periods in the water also often attract a retinue of fish, with smaller fishes coming in close and larger fishes observing from a greater distance. [[Marine turtle]]s, functioning as a mobile shelter for small fish, can be impaled accidentally by a swordfish trying to catch the fish.<ref>{{Cite journal | doi = 10.1007/BF00004759| title = Impalement of marine turtles (Reptitia, Chelonia: Cheloniidae and Dermochelyidae) by billfishes (Osteichthyes, Perciformes: Istiophoridae and Xiphiidae)| journal = Environmental Biology of Fishes| volume = 39| pages = 85–96| year = 1994| last1 = Frazier | first1 = J. G. | last2 = Fierstine | first2 = H. L. | last3 = Beavers | first3 = S. C. | last4 = Achaval | first4 = F. | last5 = Suganuma | first5 = H. | last6 = Pitman | first6 = R. L. | last7 = Yamaguchi | first7 = Y. | last8 = Prigioni | first8 = C. M. | s2cid = 23551149}}</ref>
Nel 1758 Carlo Linneo descrisse il giaguaro nella sua opera Systema Naturae e gli diede il nome scientifico Felis onca. Nel XIX e XX secolo, diversi esemplari di tipo giaguaro hanno costituito la base per le descrizioni delle sottospecie. Nel 1939, Reginald Innes Pocock riconobbe otto sottospecie in base alle origini geografiche e alla morfologia del cranio di questi esemplari. Pocock non aveva accesso a esemplari zoologici sufficienti per valutare criticamente il loro stato sottospecifico, ma espresse dubbi sullo stato di diversi. Un successivo esame del suo lavoro suggerì che si dovessero riconoscere solo tre sottospecie. La descrizione di P. o. palustris era basato su un teschio fossile.
===Coastal fish===
== Tassonomia ed evoluzione ==
{{main|Coastal fish}}
=== Tassonomia ===
[[File:Sixfinger threadfin school.jpg|thumb|left|Schooling [[threadfin]], a coastal species]]
Nel 1758 [[Linneo|Carlo Linneo]] descrisse il giaguaro nella sua opera ''[[Systema Naturae]]'' e gli diede il [[Nomenclatura binomiale|nome scientifico]] ''Felis onca''.<ref name=Linn1758>{{cita libro | autore=C. Linnaeus | anno=1758 | capitolo=''Felis onca'' | titolo=Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis | edizione=10 | lingua=la | città=Holmiae | editore=Laurentius Salvius | volume=1 | p=42 | url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}}</ref> Tra il XIX e il XX secolo, vari [[Tipo nomenclaturale|esemplari tipo]] sono serviti da base per la descrizione di un certo numero di [[sottospecie]].<ref name=MSW3>{{MSW3|id=14000240}}</ref> Nel 1939 [[Reginald Innes Pocock]] ne riconobbe otto in base all'origine geografica e alla [[Morfologia (biologia)|morfologia]] del cranio.<ref name=Pocock1939>{{cita pubblicazione | autore=R. I. Pocock | anno=1939 | titolo=The races of jaguar (''Panthera onca'') | rivista=Novitates Zoologicae | volume=41 | pp=406-422 | url=https://archive.org/details/cbarchive_123320_theracesofjaguarpantheraonca9999}}</ref> Pocock non aveva accesso a esemplari zoologici sufficienti per valutare seriamente la validità del loro stato di sottospecie, ma espresse ugualmente dubbi riguardo alcune di esse. Un successivo riesame del suo lavoro suggerì che si dovessero riconoscere come tali solo tre sottospecie: la descrizione di ''P. o. palustris'', ad esempio, si basava sull'esame di un cranio [[fossile]].<ref name=Seymour>{{cita pubblicazione | autore=K. L. Seymour | anno=1989 | titolo=''Panthera onca'' | rivista=Mammalian Species | numero=340 | pp=1-9 | url=http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-340-01-0001.pdf | urlmorto=sì | urlarchivio=https://web.archive.org/web/20100620231016/http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-340-01-0001.pdf | doi=10.2307/3504096 | jstor=3504096}}</ref>
[[Coastal fish]] (also called [[neritic]] or inshore fish) inhabit the waters near the [[coast]] and above the [[continental shelf]]. Since the continental shelf is usually less than 200 metres deep, it follows that coastal fish that are not demersal fish, are usually epipelagic fish, inhabiting the sunlit epipelagic zone.<ref name="Moyle585"/>
Nel 2005 venivano ancora considerati taxa validi nove sottospecie, elencate qui sotto con il corrispondente olotipo:<ref name=MSW3/>
Coastal epipelagic fish are among the most abundant in the world. They include forage fish as well as the predator fish that feed on them. Forage fish thrive in those inshore waters where high productivity results from the upwelling and shoreline run off of nutrients. Some are partial residents that spawn in streams, estuaries, and bays, but most complete their life cycle in the zone.<ref name="Moyle572"/>
* ''P. o. onca'' {{zoo|Linnaeus|1758}}, un esemplare proveniente dal Brasile;<ref name=Linn1758/>
* ''P. o. peruviana'' {{zoo|[[Henri Marie Ducrotay de Blainville|de Blainville]]|1843}}, un cranio proveniente dal Perù;<ref>{{cita libro | autore=H. M. D. de Blainville | anno=1843 | capitolo=''F. leo nubicus'' | titolo=Ostéographie ou description iconographique comparée du squelette et du système dentaire des mammifères récents et fossils pour servir de base à la zoologie et la géologie | lingua=fr | città=Parigi | editore=J. B. Baillière et Fils | volume=2 | url=https://gallica.bnf.fr/ark:/12148/bpt6k6538959f/f171.item | accesso=13 agosto 2020 | urlarchivio=https://web.archive.org/web/20211129021313/https://gallica.bnf.fr/ark:/12148/bpt6k6538959f/f171.item | urlmorto=no}}</ref>
* ''P. o. hernandesii'' {{zoo|[[John Edward Gray|J. E. Gray]]|1857}}, un esemplare proveniente da [[Mazatlán]], in Messico;<ref>{{cita pubblicazione | autore=J. E. Gray | anno=1857 | titolo=Notice of a new species of jaguar from Mazatlan, living in the gardens of the Zoological Society | rivista=Proceedings of the Zoological Society of London | volume=25 | p=278 | url=https://archive.org/details/lietuvostsrmoksl56liet/page/278/mode/2up}}</ref>
* ''P. o. palustris'' {{zoo|[[Florentino Ameghino|Ameghino]]|1888}}, una [[mandibola]] [[fossile]] rinvenuta nelle [[Sierras Pampeanas]] del [[distretto di Córdova]], in Argentina;<ref>{{cita libro | autore=F. Ameghino | anno=1888 | capitolo=Formación Pampeana | titolo=Los Mamíferos fósiles de la República Argentina | lingua=es | città=Buenos Aires | editore=Pablo E. Coni é hijos | url=https://archive.org/details/mamferosfsil00ameg/page/476/mode/1up | pp=473-493}}</ref>
* ''P. o. centralis'' {{zoo|[[Edgar Alexander Mearns|Mearns]]|1901}}, il cranio di un maschio proveniente da [[Cantone di Talamanca|Talamanca]], in Costa Rica;<ref name=Mearns>{{cita pubblicazione | autore=E. A. Mearns | anno=1901 | titolo=The American Jaguars | rivista=Proceedings of the Biological Society of Washington | volume=14 | pp=137-143 | url=https://archive.org/details/3908800952802714biolrich/page/138/mode/2up}}</ref>
* ''P. o. goldmani'' {{zoo|Mearns|1901}}, una pelle proveniente da Yohatlan nello stato di [[Campeche]], in Messico;<ref name=Mearns/>
* ''P. o. paraguensis'' {{zoo|[[Ned Hollister|Hollister]]|1914}}, il cranio di un maschio proveniente dal Paraguay;<ref>{{cita pubblicazione | autore=N. Hollister | anno=1915 | titolo=Two new South American jaguars | rivista=Proceedings of the United States National Museum | volume=48 | numero=2069 | pp=169-170 | url=https://archive.org/details/proceedingsofuni481915unit/page/168/mode/2up | doi=10.5479/si.00963801.48-2069.169}}</ref>
* ''P. o. arizonensis'' {{zoo|[[Edward Alphonso Goldman|Goldman]]|1932}}, la pelle e il cranio di un maschio proveniente dalle vicinanze di [[Cibecue]], in Arizona;<ref name=Goldman1932>{{cita pubblicazione | autore=E. A. Goldman | anno=1932 | titolo=The jaguars of North America | rivista=Proceedings of the Biological Society of Washington | volume=45 | pp=143-146 | url=https://archive.org/details/proceedingsofbi451932biol/page/142/mode/2up}}</ref>
* ''P. o. veraecruscis'' {{zoo|[[Edward William Nelson|Nelson]] e Goldman|1933}}, il cranio di un maschio proveniente da [[San Andrés Tuxtla]], in Messico.<ref>{{cita pubblicazione | autore=E. W. Nelson e E. A. Goldman | anno=1933 | titolo=Revision of the jaguars | rivista=Journal of Mammalogy | volume=14 | numero=3 | pp=221-240 | doi=10.2307/1373821 | jstor=1373821}}</ref>
{{clear}}
Reginald Innes Pocock inserì il giaguaro nel [[Genere (tassonomia)|genere]] ''Panthera'' e notò che esso condivideva diverse caratteristiche morfologiche con il leopardo (''[[Panthera pardus|P. pardus]]''). Lo zoologo, pertanto, giunse alla conclusione che le due specie dovevano essere strettamente imparentate.<ref name=Pocock1939/> Di recente, le ricerche sui dati [[Morfologia (biologia)|morfologici]] e [[Genetica|genetici]] hanno indicato che la specie presenta una [[variazione clinale]] nord-sud tra le popolazioni, ma non hanno riscontrato alcuna prova di differenziazione sottospecifica.<ref name=Eizirik/><ref name=Larson>{{cita pubblicazione | autore=S. E. Larson | anno=1997 | titolo=Taxonomic re-evaluation of the jaguar | rivista=Zoo Biology | volume=16 | numero=2 | pp=107-120 | doi=10.1002/(SICI)1098-2361(1997)16:2<107::AID-ZOO2>3.0.CO;2-E}}</ref> Le analisi del [[DNA]] di 84 esemplari di giaguaro provenienti dall'America meridionale ha rivelato che in passato il [[flusso genico]] tra le popolazioni della [[Colombia]] era molto elevato.<ref>{{cita pubblicazione | autore=M. Ruiz-Garcia, E. Payan, A. Murillo e D. Alvarez | anno=2006 | titolo=DNA microsatellite characterization of the jaguar (''Panthera onca'') in Colombia | rivista=Genes & Genetic Systems | volume=81 | numero=2 | pp=115-127 | url=https://www.jstage.jst.go.jp/article/ggs/81/2/81_2_115/_pdf | doi=10.1266/ggs.81.115 | pmid=16755135 | accesso=13 dicembre 2014 | urlarchivio=https://web.archive.org/web/20141216093206/https://www.jstage.jst.go.jp/article/ggs/81/2/81_2_115/_pdf | urlmorto=no}}</ref> Dal 2017, il giaguaro è considerato un [[Monospecifico|taxon monotipico]].<ref name="catsg">{{cita pubblicazione | autore=A. C. Kitchener, C. Breitenmoser-Würsten, E. Eizirik, A. Gentry, L. Werdelin, A. Wilting, N. Yamaguchi, A. V. Abramov, P. Christiansen, C. Driscoll, J. W. Duckworth, W. Johnson, S.-J. Luo, E. Meijaard, P. O'Donoghue, J. Sanderson, K. Seymour, M. Bruford, C. Groves, M. Hoffmann, K. Nowell, Z. Timmons e S. Tobe | anno=2017 | titolo=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group | rivista=Cat News | volume=Special Issue 11 | pp=70-71 | url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=70 | accesso=13 maggio 2018 | urlarchivio=https://web.archive.org/web/20180730142355/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=70 | urlmorto=no}}</ref>
===Oceanic Evoluzione =fish==
[[File:Oceanic divisions.svg|thumb|400px|right|Oceanic fish inhabit the [[oceanic zone]], which is the deep open water which lies beyond the continental shelves.]]
[[File:Panthera onca augusta.JPG|thumb|right|Cranio fossile di ''P. o. augusta''.]]
Gli studiosi ritengono che la linea evolutiva ''Panthera'' si sia separata geneticamente dal [[progenitore comune]] dei [[Felidae|Felidi]] in un periodo che varia, a seconda delle stime, tra 9,32 e 4,47 milioni di anni fa e 11,75 e 0,97 milioni di anni fa,<ref name="Johnson2006">{{cita pubblicazione | autore=W. E. Johnson, E. Eizirik, J. Pecon-Slattery, W. J. Murphy, A. Antunes, E. Teeling e S. J. O'Brien | titolo=The late miocene radiation of modern Felidae: A genetic assessment | rivista=[[Science]] | volume=311 | numero=5757 | pp=73-77 | anno=2006 | pmid=16400146 | doi=10.1126/science.1122277 | bibcode=2006Sci...311...73J | s2cid=41672825 | url=https://zenodo.org/record/1230866 | accesso=24 novembre 2021 | urlarchivio=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 | urlmorto=no}}</ref><ref name="Werdelin2010">{{cita libro | autore=L. Werdelin, N. Yamaguchi, W. E. Johnson e S. J. O'Brien | capitolo=Phylogeny and evolution of cats (Felidae) | anno=2010 | pp=59-82 | editore=Oxford University Press | città=Oxford, UK | isbn=978-0-19-923445-5 | url=https://www.researchgate.net/publication/266755142 | curatore=D. W. Macdonald e A. J. Loveridge | titolo=Biology and Conservation of Wild Felids | accesso=24 novembre 2021 | urlarchivio=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 | urlmorto=no}}</ref><ref name=Li_al2016>{{cita pubblicazione | autore=G. Li, B. W. Davis, E. Eizirik e W. J. Murphy | anno=2016 | titolo=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) | rivista=Genome Research | volume=26 | numero=1 | pp=1-11 | doi=10.1101/gr.186668.114 | pmid=26518481 | pmc=4691742}}</ref> e che il genere abbia avuto origine molto probabilmente nell'Asia centro-settentrionale.<ref name=Tseng2014>{{cita pubblicazione | autore=Z. J. Tseng, X. Wang, G. J. Slater, G. T. Takeuchi, Q. Li, J. Liu e G. Xie | anno=2014 | titolo=Himalayan fossils of the oldest known pantherine establish ancient origin of big cats | rivista=Proceedings of the Royal Society B: Biological Sciences | volume=281 | numero=1774 | p=20132686 | doi=10.1098/rspb.2013.2686 | pmid=24225466 | pmc=3843846}}</ref> Alcune analisi genetiche collocano il giaguaro come specie sorella del leone, dal quale si sarebbe separato tra 3,46 e 1,22 milioni di anni fa,<ref name=Johnson2006/><ref name="Werdelin2010"/> mentre altre ricerche collocano il leone più prossimo al leopardo.<ref name="davis2010">{{cita pubblicazione | autore=B. W. Davis, G. Li e W. J. Murphy | titolo=Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, ''Panthera'' (Carnivora: Felidae) | rivista=Molecular Phylogenetics and Evolution | anno=2010 | volume=56 | numero=1 | pp=64-76 | doi=10.1016/j.ympev.2010.01.036 | pmid=20138224 | url=https://www.academia.edu/12157986}}</ref><ref name="mazak2011">{{cita pubblicazione | autore=J. H. Mazák, P. Christiansen, A. C. Kitchener e A. Goswami | titolo=Oldest known pantherine skull and evolution of the tiger | rivista=PLOS ONE | anno=2011 | volume=6 | numero=10 | pp=e25483 | doi=10.1371/journal.pone.0025483 | pmid=22016768 | pmc=3189913 | bibcode=2011PLoSO...625483M}}</ref>
Oceanic fish (also called open ocean or offshore fish) live in the waters that are not above the continental shelf. Oceanic fish can be contrasted with [[coastal fish]], who do live above the continental shelf. However, the two types are not mutually exclusive, since there are no firm boundaries between coastal and ocean regions, and many epipelagic fish move between coastal and oceanic waters, particularly in different stages in their life cycle.<ref name="Moyle572"/>
Sembra che la linea evolutiva del giaguaro abbia avuto origine in Africa e che da qui si sia diffusa in Eurasia tra 1,95 e 1,77 milioni di anni fa. La specie attuale potrebbe discendere da ''[[Panthera gombaszoegensis]]'', che si pensa abbia raggiunto il continente americano attraverso la [[Beringia]], il ponte di terra che all'epoca collegava le due sponde dello [[stretto di Bering]].<ref name=Argant2011>{{cita pubblicazione | autore=A. Argant e J. Argant | titolo=The ''Panthera gombaszogensis'' story: the contribution of the Château Breccia (Saône-et-Loire, Burgundy, France) | rivista=Quaternaire | numero=Hors-serie 4 | anno=2011 | pp=247-269 | url=https://www.researchgate.net/publication/286036249 | accesso=24 novembre 2021 | urlarchivio=https://web.archive.org/web/20211129021307/https://www.researchgate.net/publication/286036249_The_Panthera_Gombaszogensis_story_The_contribution_of_the_chateau_breccia_Saone-Et-Loire_Burgundy_France | urlmorto=no}}</ref><ref>{{cita pubblicazione | autore=Q. Jiangzuo e J. Liu | anno=2020 | titolo=First record of the Eurasian jaguar in southern Asia and a review of dental differences between pantherine cats | rivista=Journal of Quaternary Science | volume=35 | numero=6 | pp=817-830 | doi=10.1002/jqs.3222 | bibcode=2020JQS....35..817J | s2cid=219914902}}</ref> In America settentrionale sono stati rinvenuti fossili di giaguaro moderno risalenti a oltre {{M|850000}} anni fa.<ref name=Seymour/> I risultati dell'analisi del [[DNA mitocondriale]] di 37 giaguari indicano che le popolazioni attuali si siano evolute tra {{M|510000}} e {{M|280000}} anni fa nel nord dell'America meridionale e successivamente abbiano ricolonizzato l'America centrale e settentrionale dopo che la specie era scomparsa da queste zone durante il [[Pleistocene superiore]].<ref name=Eizirik>{{cita pubblicazione | autore=E. Eizirik, J. H. Kim, M. Menotti-Raymond, P. G. Crawshaw Jr., S. J. O'Brien e W. E. Johnson | anno=2001 | titolo=Phylogeography, population history and conservation genetics of jaguars (''Panthera onca'', Mammalia, Felidae) | rivista=Molecular Ecology | volume=10 | numero=1 | pp=65-79 | url=https://semanticscholar.org/paper/5bd14c9ccc8cc43ed15352fc86075b8d6b8cccee | doi=10.1046/j.1365-294X.2001.01144.x | pmid=11251788 | s2cid=3916428 | accesso=29 novembre 2019 | urlarchivio=https://web.archive.org/web/20211129021257/https://www.semanticscholar.org/paper/Phylogeography%2C-population-history-and-conservation-Eizirik-Kim/5bd14c9ccc8cc43ed15352fc86075b8d6b8cccee | urlmorto=no}}</ref>
Oceanic epipelagic fish can be true residents, partial residents, or accidental residents. True residents live their entire life in the open ocean. Only a few species are true residents, such as [[tuna]], [[billfish]], [[flying fish]], [[sauries]], [[Pilot fish|pilotfish]], [[remora]]s, [[dolphinfish]], ocean sharks, and [[ocean sunfish]]. Most of these species migrate back and forth across open oceans, rarely venturing over continental shelves. Some true residents associate with drifting jellyfish or seaweeds.<ref name="Moyle572"/>
Grazie ai ritrovamenti fossili sono note due sottospecie estinte di giaguaro, una nordamericana, ''P. o. augusta'', e una sudamericana, ''P. o. mesembrina''.<ref>{{cita pubblicazione | autore=A. Chahud e M. Okumura | anno=2020 | titolo=The presence of ''Panthera onca'' Linnaeus 1758 (Felidae) in the Pleistocene of the region of Lagoa Santa, State of Minas Gerais, Brazil | rivista=Historical Biology | volume=33 | numero=10 | pp=2496-2503 | doi=10.1080/08912963.2020.1808975 | s2cid=225408043}}</ref>
Partial residents occur in three groups: species that live in the zone only when they are juveniles (drifting with jellyfish and seaweeds); species that live in the zone only when they are adults (salmon, flying fish, dolphin, and whale sharks); and deep water species that make nightly migrations up into the surface waters (such as the [[lanternfish]]).<ref name="Moyle572"/> Accidental residents occur occasionally when adults and juveniles of species from other environments are carried accidentally into the zone by currents.<ref name="Moyle572"/>
<div style="text-align:center">'''Relazioni [[Filogenesi|filogenetiche]] del giaguaro come risultano dalle analisi del'''</div>
{{colonne|autoclear}}
<div style="text-align:center">'''DNA nucleare:'''<ref name=Johnson2006/></div>
{{clade |label1=Felidae
|1={{clade |label2=Pantherinae
|1=Felinae [[File:Felis torquata - 1834 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|70 px]]
|2={{clade |label1=''Panthera''
|1={{clade
|1={{clade
|1={{clade
|1=Leone (''[[Panthera leo|P. leo]]'') [[File:Felis leo - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(White Background).jpg|70 px]]
|2='''Giaguaro''' [[File:Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]}}
|2=Leopardo (''[[Panthera pardus|P. pardus]]'') [[File:Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]}}
|2={{clade
|1=Tigre (''[[Panthera tigris|P. tigris]]'') [[File:Stamp-russia2014-save-russian-cats-(tiger).png|70 px]]
|2=Leopardo delle nevi (''[[Panthera uncia|P. uncia]]'') [[File:Stamp-russia2014-save-russian-cats-(snow leopard).png|70 px]]}} }}
|2=''[[Neofelis]]'' [[File:Studienblatt Felis macroscelis Nebelparder (white background).jpg|70 px]]}} }} }}
{{colonne spezza}}
<div style="text-align:center">'''DNA mitocondriale:'''<ref name=Li_al2016/></div>
{{clade |label1=Felidae
|1={{clade |label2=Pantherinae
|1=Felinae [[File:Felis torquata - 1834 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|70 px]]
|2={{clade |label1=''Panthera''
|1={{clade
|1={{clade
|1={{clade
|1={{clade|1=Leone [[File:Felis leo - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(White Background).jpg|70 px]]|2=Leopardo [[File:Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]}}
|2=Leopardo delle nevi [[File:Stamp-russia2014-save-russian-cats-(snow leopard).png|70 px]]}}
|2='''Giaguaro''' [[File:Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]}}
|2=Tigre [[File:Stamp-russia2014-save-russian-cats-(tiger).png|70 px]]}}
|2=''Neofelis'' [[File:Studienblatt Felis macroscelis Nebelparder (white background).jpg|70 px]]}} }} }}
{{colonne fine}}
<gallery class="left" widths="225" heights="187">
== Descrizione ==
File:Sunfish.jpg|The huge [[ocean sunfish]], a true resident of the ocean epipelagic zone, sometimes drifts with the current, eating [[jellyfish]].
{{Immagine multipla
File:Whale shark Georgia aquarium.jpg|The giant [[whale shark]], another resident of the ocean epipelagic zone, filter feeds on [[plankton]], and periodically dives deep into the mesopelagic zone.
| allinea = right
File:Protomyctophum subparallelum (no common name).gif|[[Lanternfish]] are partial residents of the ocean epipelagic zone During the day they hide in deep waters, but at night they migrate up to surface waters to feed.
| direzione = verticale
</gallery>
| larghezza = 200
| titolo =
| immagine1 = Description iconographique comparée du squelette et du système dentaire des mammifères récents et fossiles (Panthera onca skull).jpg
| didascalia1 = Il cranio di un giaguaro
| immagine2 = Cheetah, leopard & jaguar (ita).jpg
| didascalia2 = Confronto tra [[Acinonyx jubatus|ghepardo]], leopardo e giaguaro.
| immagine3 = Black Jaguar (Panthera_onca).JPG
| didascalia3 = Un giaguaro nero. Questi esemplari [[Melanismo|melanici]], come quelli di leopardo, sono noti come [[Pantera nera|pantere nere]].
}}
Il giaguaro è un felino dalla struttura compatta e muscolosa. È il più grande felino delle Americhe, nonché il terzo in assoluto: solo la tigre e il leone presentano dimensioni maggiori.<ref name=Seymour/><ref name="sizes">{{cita pubblicazione | autore=M. W. Hayward, J. F. Kamler, R. A. Montgomery e A. Newlove | anno=2016 | titolo=Prey Preferences of the Jaguar ''Panthera onca'' Reflect the Post-Pleistocene Demise of Large Prey | rivista=Frontiers in Ecology and Evolution | volume=3 | p=148 | doi=10.3389/fevo.2015.00148}}</ref><ref name="sizes22">{{cita pubblicazione | autore=M. K. Hope e S. L. Deem | anno=2006 | titolo=Retrospective Study of Morbidity and Mortality of Captive Jaguars (''Panthera onca'') in North America: 1982–2002 | rivista=Zoo Biology | volume=25 | numero=6 | pp=501-512 | url=https://repository.si.edu/bitstream/handle/10088/11687/Zoo%20Biology%2C%20Vol.%2025%2C%20Issue%206%20Retrospective%20Study%20of%20Morbidity%20and%20Mortality%20of%20Captive%20Jaguars....pdf?sequence=1&isAllowed=y | doi=10.1002/zoo.20112 | accesso=12 settembre 2018 | urlarchivio=https://web.archive.org/web/20180909000241/https://repository.si.edu/bitstream/handle/10088/11687/Zoo%20Biology%2C%20Vol.%2025%2C%20Issue%206%20Retrospective%20Study%20of%20Morbidity%20and%20Mortality%20of%20Captive%20Jaguars....pdf?sequence=1&isAllowed=y | urlmorto=no}}</ref> Misura 68-75 cm di altezza al garrese,<ref>{{cita pubblicazione | autore=M. S. Rich | anno=1976 | titolo=The jaguar | rivista=Zoonoz | volume=49 | numero=9 | pp=14-17}}</ref> ma il peso e le dimensioni variano considerevolmente: anche se la maggior parte degli esemplari rientra nel range compreso tra 56 e 96 kg, sono stati documentati maschi di dimensioni eccezionali che raggiungevano i 158 kg.<ref name=r10/><ref name="Animal">{{cita libro | autore=D. Burnie e D. E. Wilson | anno=2001 | titolo=Animal: The Definitive Visual Guide to the World's Wildlife | città=New York City | editore=[[Dorling Kindersley]] | isbn=978-0-7894-7764-4}}</ref> D'altro canto, le femmine più piccole pesano circa 36 kg. Il [[dimorfismo sessuale]] è evidente: le femmine sono del 10-20% più piccole dei maschi. La lunghezza dal naso alla base della coda varia tra 112 e 185 cm. La coda misura 45-75 cm ed è la più corta tra quelle di tutti i [[grandi felini]].<ref name=r10>{{cita libro | autore=R. M. Nowak, | anno=1999 | titolo=Walker's Mammals of the World | editore=[[Johns Hopkins University Press]] | città=Baltimora | volume=2 | p=831 | url=https://books.google.it/books?id=T37sFCl43E8C&pg=PAPA831&redir_esc=y#v=onepage&q&f=false | isbn=978-0-8018-5789-8}}</ref> Le zampe, molto muscolose, sono più corte di quelle delle altre specie del genere ''Panthera'' di dimensioni corporee simili.<ref name=Gonyea1976/>
===Deep water fish===
Un'ulteriore variabilità delle dimensioni è stata riscontrata a seconda della regione e dell'habitat: esse, infatti, tendono ad aumentare procedendo da nord a sud. I giaguari della [[riserva della biosfera Chamela-Cuixmala]], sulla costa pacifica del Messico centrale, pesano circa 50 kg, più o meno quanto una femmina di puma (''[[Puma concolor]]'');<ref name=foodhabits>{{cita pubblicazione | autore=R. Nuanaez, B. Miller e F. Lindzey | anno=2000 | titolo=Food habits of jaguars and pumas in Jalisco, Mexico | rivista=Journal of Zoology | volume=252 | numero=3 | pp=373-379 | url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 | doi=10.1111/j.1469-7998.2000.tb00632.x | accesso=8 settembre 2006 | urlarchivio=https://web.archive.org/web/20160810211518/http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 | urlmorto=no}}</ref> quelli del Venezuela e del [[Brasile]] sono molto più grandi, con un peso medio che si aggira sui 95 kg nei maschi e sui 56-78 kg nelle femmine.<ref name=Seymour/>
[[File:Pelagiczone.svg|thumb|160px|right|Scale diagram of the layers of the pelagic zone]]
{{See also|Deep sea fish}}
In the deep ocean, the waters extend far below the epipelagic zone and support very different types of pelagic fishes adapted to living in these deeper zones.<ref name="Moyle585"/>
Il mantello varia dal giallo pallido al marroncino o al giallo-rossastro, con la parte inferiore biancastra e ricoperta da macchie nere. La forma delle macchie è variabile: sui fianchi sono raggruppate a formare rosette che recano all'interno all'interno uno o due punti. Le macchie sulla testa e sul collo sono generalmente semplici, così come quelle sulla coda, che possono fondersi a formare bande verso l'estremità e creare una punta nera. Quelle al centro del dorso sono allungate e spesso si uniscono a creare una striscia mediana, mentre quelle sull'addome sono piene e di forma irregolare.<ref name=Seymour/> Questa colorazione funge da [[camuffamento]] nelle aree caratterizzate da una fitta vegetazione e da un'ombreggiatura irregolare.<ref>{{cita pubblicazione | autore=W. L. Allen, I. C. Cuthill, N. E. Scott-Samuel e R. Baddeley | anno=2010 | titolo=Why the leopard got its spots: relating pattern development to ecology in felids | rivista=Proceedings of the Royal Society B | volume=278 | numero=1710 | pp=1373-1380 | doi=10.1098/rspb.2010.1734 | pmc=3061134 | pmid=20961899}}</ref> I giaguari che vivono nelle foreste sono spesso più scuri e notevolmente più piccoli di quelli che vivono negli ambienti aperti, probabilmente a causa del minor numero di grandi prede erbivore nelle aree forestali.<ref name=CAP>{{cita libro | autore=K. Nowell e P. Jackson | anno=1996 | capitolo=Jaguar, ''Panthera onca'' (Linnaeus, 1758) | titolo=Wild Cats. Status Survey and Conservation Action Plan | editore=IUCN/SSC Cat Specialist Group | ___location=Gland, Svizzera | url=http://carnivoractionplans1.free.fr/wildcats.pdf#page=143 | pp=118-122 | accesso=31 agosto 2006 | urlarchivio=https://web.archive.org/web/20070807215533/http://carnivoractionplans1.free.fr/wildcats.pdf#page=143 | urlmorto=no}}</ref>
In deep water, [[marine snow]] is a continuous shower of mostly organic [[detritus]] falling from the upper layers of the water column. Its origin lies in activities within the productive [[photic zone]]. Marine snow includes dead or dying [[plankton]], [[protist]]s ([[diatom]]s), fecal matter, sand, soot, and other inorganic dust. The "snowflakes" grow over time and may reach several centimetres in diameter, travelling for weeks before reaching the ocean floor. However, most organic components of marine snow are consumed by [[microbe]]s, [[zooplankton]], and other filter feeding animals within the first 1,000 metres of their journey, that is, within the epipelagic zone. In this way marine snow can be considered the foundation of deep-sea [[mesopelagic]] and [[benthic]] [[ecosystem]]s: As sunlight cannot reach them, deep-sea organisms rely heavily on marine snow as an energy source.
Il giaguaro somiglia molto al leopardo, ma ha generalmente una struttura più robusta, con zampe più tozze e una testa più squadrata. Inoltre, le rosette del mantello sono più grandi, più scure e meno numerose e dotate di linee più spesse con un unico piccolo punto nel mezzo.<ref name=Gonyea1976>{{cita pubblicazione | autore=W. J. Gonyea | anno=1976 | titolo=Adaptive differences in the body proportions of large felids | rivista=Acta Anatomica | volume=96 | numero=1 | pp=81-96 | doi=10.1159/000144663 | pmid=973541}}</ref> Possiede mascelle potenti, in grado di produrre il terzo morso più potente tra i felini, dopo la tigre e il leone.<ref name=Wroe2006>{{cita pubblicazione | autore=S. Wroe, C. McHenry e J. Thomason | anno=2006 | titolo=Bite club: comparative bite force in big biting mammals and the prediction of predatory behavior in fossil taxa | rivista=Proceedings of the Royal Society B | volume=272 | numero=1563 | pp=619-625 | doi=10.1098/rspb.2004.2986 | pmc=1564077 | pmid=15817436}}</ref> Sulla punta del [[canino]] si registra una forza media di 887,0 [[Newton (unità di misura)|newton]] e un quoziente di forza di 118,6.<ref>{{cita pubblicazione | autore=P. Christiansen | anno=2007 | titolo=Canine morphology in the larger Felidae: implications for feeding ecology | rivista=Biological Journal of the Linnean Society | volume=91 | numero=4 | pp=573-592 | doi=10.1111/j.1095-8312.2007.00819.x}}</ref> Un giaguaro di 100 kg può infliggere un morso dalla potenza di 4,939 kN in corrispondenza dei canini e di 6,922 kN dei [[carnassiali]].<ref>{{cita pubblicazione | autore=A. Hartstone-Rose, J. M. G. Perry e C. J. Morrow | anno=2012 | titolo=Bite Force Estimation and the Fiber Architecture of Felid Masticatory Muscles | rivista=The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology | volume=295 | numero=8 | pp=1336-1351 | doi=10.1002/ar.22518 | pmid=22707481 | s2cid=35304260}}</ref>
Some deep-sea pelagic groups, such as the [[lanternfish]], [[ridgehead]], [[marine hatchetfish]], and [[Phosichthyidae|lightfish]] families are sometimes termed ''pseudoceanic'' because, rather than having an even distribution in open water, they occur in significantly higher abundances around structural oases, notably [[seamount]]s, and over [[continental slope]]s. The phenomenon is explained by the likewise abundance of prey species that also are attracted to the structures.
=== Variazioni cromatiche ===
I giaguari [[Melanismo|melanici]] sono noti anche come [[Panthera nera|pantere nere]]. La [[Polimorfismo (biologia)|forma]] nera è meno comune di quella maculata.<ref>{{cita libro | autore=D. E. Brown e C. A. Lopez-Gonzalez | anno=2001 | titolo=Borderland jaguars: tigres de la frontera | editore=University of Utah Press | città=Salt Lake City, UT}}</ref> La loro presenza è stata documentata sia in America centrale che meridionale. Nel giaguaro il melanismo è dovuto a delezioni nel gene del recettore della melanocortina 1 e viene ereditato attraverso un [[Dominanza (genetica)|allele dominante]].<ref>{{cita pubblicazione | autore=E. Eizirik, N. Yuhki, W. E. Johnson, M. Menotti-Raymond, S. S. Hannah e S. J. O'Brien | anno=2003 | titolo=Molecular Genetics and Evolution of Melanism in the Cat Family | rivista=Current Biology | volume=13 | numero=5 | pp=448-453 | doi=10.1016/S0960-9822(03)00128-3 | pmid=12620197 | s2cid=19021807}}</ref>
The fish in the different pelagic and deep water benthic zones are physically structured, and behave, in ways that differ markedly from each other. Groups of coexisting species within each zone all seem to operate in similar ways, such as the small mesopelagic [[Diel vertical migration|vertically migrating]] plankton-feeders, the bathypelagic [[anglerfish]]es, and the deep water benthic [[rattail]]s.<ref name="Moyle591">[[#Moyle|Moyle and Cech]], p. 591</ref>
Nel 2004, una fototrappola sulle montagne della [[Sierra Madre Occidentale]] catturò le immagini del primo giaguaro nero documentato nel Messico settentrionale.<ref name=Dinets>{{cita pubblicazione | autore=V. Dinets e P. J. Polechla | anno=2005 | titolo=First documentation of melanism in the jaguar (''Panthera onca'') from northern Mexico | rivista=Cat News | volume=42 | p=18 | url=http://dinets.travel.ru/blackjaguar.htm | urlmorto=sì | urlarchivio=https://web.archive.org/web/20060926024755/http://dinets.travel.ru/blackjaguar.htm}}</ref> Altri giaguari neri sono stati segnalati nella riserva biologica Alberto Manuel Brenes in Costa Rica, sulle montagne della [[Cordigliera di Talamanca]], nel [[parco nazionale Barbilla]] e nella parte orientale di [[Panama]].<ref>{{cita pubblicazione | autore=M. C. Núñez e E. C. Jiménez | anno=2009 | titolo=A new record of a black jaguar, ''Panthera onca'' (Carnivora: Felidae) in Costa Rica | rivista=Brenesia | volume=71 | pp=67-68 | url=https://www.researchgate.net/publication/313473228 | accesso=12 aprile 2021 | urlarchivio=https://web.archive.org/web/20211129021258/https://www.researchgate.net/publication/313473228_New_record_of_a_black_jaguar_Panthera_onca_Carnivora_Felidae_in_Costa_Rica | urlmorto=no}}</ref><ref>{{cita pubblicazione | autore=M. S. Mooring, A. A. Eppert e R. T. Botts | anno=2020 | titolo=Natural Selection of Melanism in Costa Rican Jaguar and Oncilla: A Test of Gloger's Rule and the Temporal Segregation Hypothesis | rivista=Tropical Conservation Science | volume=13 | pp=1-15 | doi=10.1177/1940082920910364}}</ref><ref>{{cita pubblicazione | autore=C. Sáenz-Bolaños, V. Montalvo, T. K. Fuller e E. Carrillo | anno=2015 | titolo=Records of black jaguars at Parque Nacional Barbilla, Costa Rica | rivista=Cat News | numero=62 | pp=38-39}}</ref><ref>{{cita pubblicazione | autore=M. Yacelga e K. Craighead | anno=2019 | titolo=Melanistic jaguars in Panama | rivista=Cat News | numero=70 | pp=39-41 | url=https://www.academia.edu/41977728 | accesso=17 aprile 2021 | urlarchivio=https://web.archive.org/web/20211129021257/https://www.academia.edu/41977728 | urlmorto=no}}</ref>
[[Acanthopterygii|Ray finned]] species, with spiny fins, are rare among deep sea fishes, which suggests that deep sea fish are ancient and so well adapted to their environment that invasions by more modern fishes have been unsuccessful.<ref name="Haedrich1996"/> The few ray fins that do exist are mainly in the [[Beryciformes]] and [[Lampriformes]], which also are ancient forms. Most deep sea pelagic fishes belong to their own orders, suggesting a long evolution in deep sea environments. In contrast, deep water benthic species are in orders that include many related shallow water fishes.<ref name="Moyle586">[[#Moyle|Moyle and Cech]], p. 586</ref>
==Distribution and habitat==
{{multiple image |direction=vertical |align=right
|image1=Jaguar (Panthera onca palustris) female Piquiri River.JPG |caption1=A female jaguar at [[Piquirí River (São Lourenço)|Piquiri River]], [[Mato Grosso]] state, Brazil
|image2=Jaguar (Panthera onca) male back in the water (29173428825).jpg |caption2=A jaguar in [[São Lourenço River (Mato Grosso)|São Lourenço River]]
}}
{{See also|North American jaguar|South American jaguar}}
In the 19th century, the jaguar was still sighted at the [[North Platte River]] in [[Colorado]] and coastal [[Louisiana]].<ref name="Daggett">{{cite journal |author1=Daggett, P. M. |author2=Henning, D. R. |year=1974 |name-list-style=amp |title=The Jaguar in North America |journal=American Antiquity |volume=39 |issue=3 |pages=465–469 |doi=10.2307/279437 |jstor=279437|s2cid=160927286}}</ref>
In 1919, sightings of jaguars were reported in the [[Monterey, California]] region.<ref>{{Cite journal |author=Merriam, C.H. |author-link=Clinton Hart Merriam |year=1919 |title=Is the Jaguar entitled to a place in the Californian fauna? |journal=Journal of Mammalogy |volume=1 |issue=1 |pages=38–42 |doi=10.1093/jmammal/1.1.38 |url=https://academic.oup.com/jmammal/article-abstract/1/1/38/875846?redirectedFrom=fulltext |access-date=18 November 2018 |archive-date=18 November 2018 |archive-url=https://web.archive.org/web/20181118210048/https://academic.oup.com/jmammal/article-abstract/1/1/38/875846?redirectedFrom=fulltext |url-status=live}}</ref>
In 1999, its historic range at the turn of the 20th century was estimated at {{cvt|19000000|km2}}, stretching from the southern United States through Central America to southern Argentina. By the turn of the 21st century, its global range had decreased to about {{cvt|8750000|km2}}, with most declines in the southern United States, northern Mexico, northern Brazil, and southern Argentina.<ref name=Sanderson2002>{{cite journal |last1=Sanderson |first1=E. W. |last2=Redford |first2=K. H. |last3=Chetkiewicz |first3=C. L. B. |last4=Medellin |first4=R. A. |last5=Rabinowitz |first5=A. R. |author5-link=Alan Rabinowitz |last6=Robinson |first6=J. G. |last7=Taber |first7=A. B. |year=2002 |name-list-style=amp |title=Planning to save a species: the jaguar as a model |journal=Conservation Biology |volume=16 |issue=1 |pages=58–72 |url=https://semanticscholar.org/paper/75d8d4f608e8e10f86dd0b1f2b35dce207decd76 |doi=10.1046/j.1523-1739.2002.00352.x |pmid=35701976 |s2cid=3955250 |access-date=29 November 2019 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021309/https://www.semanticscholar.org/paper/Planning-to-Save-a-Species%3A-the-Jaguar-as-a-Model-Sanderson-Redford/75d8d4f608e8e10f86dd0b1f2b35dce207decd76 |url-status=live}}</ref>
Its present range extends from Mexico through Central America to South America comprising [[Belize]], [[Guatemala]], [[Honduras]], [[Nicaragua]], [[Costa Rica]], particularly on the [[Osa Peninsula]], [[Panama]], [[Colombia]], [[Venezuela]], [[Guyana]], [[Suriname]], [[French Guiana]], [[Ecuador]], [[Peru]], [[Bolivia]], Brazil, [[Paraguay]] and [[Argentina]]. It is considered to be [[locally extinct]] in [[El Salvador]] and [[Uruguay]].<ref name=iucn />
Many species move daily between zones in vertical migrations. In the following table, they are listed in the middle or deeper zone where they regularly are found.
Jaguars have been occasionally sighted in [[Arizona]], [[New Mexico]] and [[Texas]].<ref>{{cite journal |author1=Brown, D. E. |author2=González, C. A. L. |year=2000 |name-list-style=amp |title=Notes on the occurrences of jaguars in Arizona and New Mexico |journal=The Southwestern Naturalist |volume=45 |issue=4 |pages=537–542 |doi=10.2307/3672607 |jstor=3672607}}</ref><ref name=Pavlik>{{Cite journal |last=Pavlik |first=S. |year=2003 |title=Rohonas and spotted Lions: The historical and cultural occurrence of the Jaguar, ''Panthera onca'', among the native tribes of the American Southwest |journal=Wíčazo Ša Review |volume=18 |issue=1 |pages=157–175 |doi=10.1353/wic.2003.0006 |jstor=1409436 |s2cid=161236104}}</ref>
Between 2012 and 2015, a male [[Vagrancy (biology)|vagrant]] jaguar was recorded in 23 locations in the [[Santa Rita Mountains]].<ref>{{cite report |author=Culver, M. |title=Open-File Report |year=2016 |chapter=Jaguar surveying and monitoring in the United States |publisher=U.S. Geological Survey |___location=Reston, VA |series=2016-1095 |doi=10.3133/ofr20161095 |doi-access=free}}</ref>
{| class="wikitable"
The jaguar prefers dense forest and typically inhabits dry [[deciduous forest]]s, [[tropical and subtropical moist broadleaf forests]], [[rainforest]]s and [[cloud forest]]s in Central and South America; open, seasonally flooded [[wetland]]s, dry [[grassland]] and historically also [[oak forest]]s in the United States. It has been recorded at elevations up to {{cvt|3800|m}} but avoids [[montane forest]]s. It favors riverine habitat and [[swamp]]s with dense vegetation cover.<ref name=CAP /> In the Mayan forests of Mexico and Guatemala, 11 GPS-collared jaguars preferred undisturbed dense habitat away from roads; females avoided even areas with low levels of human activity, whereas males appeared less disturbed by human population density.<ref>{{cite journal |author1=Colchero, F. |author2=Conde, D. A. |author3=Manterola, C. |author4=Chávez, C. |author5=Rivera, A. |author6=Ceballos, G. |year=2011 |name-list-style=amp |title=Jaguars on the move: modeling movement to mitigate fragmentation from road expansion in the Mayan Forest |journal=Animal Conservation |volume=14 |issue=2 |pages=1–9 |doi=10.1111/J.1469-1795.2010.00406.X |s2cid=62820463 |url=https://www.demogr.mpg.de/publications/files/4097_1300970681_1_ArticlePdf.pdf |access-date=14 March 2021 |archive-date=26 January 2021 |archive-url=https://web.archive.org/web/20210126211109/https://www.demogr.mpg.de/publications/files/4097_1300970681_1_ArticlePdf.pdf |url-status=live}}</ref> A young male jaguar was also recorded in the [[Semi-arid climate|semi-arid]] [[Sierra de San Carlos]] at a waterhole.<ref>{{cite journal |author1=Caso, A. |name-list-style=amp |author2=Domínguez, E. F. |year=2018 |title=Confirmed presence of jaguar, ocelot and jaguarundi in the Sierra of San Carlos, Mexico |journal=Cat News |issue=68 |pages=31–32}}</ref>
|+ Species by pelagic zone
|-
! Zone
! Species and species groups include:
|-
| Epipelagic<ref name="Moyle571"/>
|
* [[mackerel shark|mackerel]], [[requiem shark|requiem]] and [[whale shark]]s
* [[clupeiform]]s – [[herring]], [[anchovy]]
* [[Salmonidae]] – [[salmon]]
* [[atheriniform]]s – [[flyingfish]]es, [[halfbeak]]s, [[sauries]]
* [[perciform]]s – [[Carangidae|jacks]], [[dolphinfish]], [[pomfret]]s, [[barracuda]]s, [[tuna]]s, [[billfish]].
|-
| Mesopelagic
| [[Lanternfish]], [[opah]], [[longnose lancetfish]], [[barreleye]], [[ridgehead]], [[sabertooth fish|sabretooth]], [[stoplight loosejaw]], [[marine hatchetfish]]<ref>{{fishbase species |genus=Argyropelecus|species=aculeatus |month=August |year=2009}}</ref>
|-
| Bathypelagic
| Principally [[bristlemouth]] and [[anglerfish]]. Also [[fangtooth]], [[viperfish]], [[black swallower]], [[telescopefish]], [[hammerjaw]], [[daggertooth]], [[barracudina]], [[black scabbardfish]], [[bobtail snipe eel]], [[unicorn crestfish]], [[Saccopharynx|gulper eel]], [[flabby whalefish]].
|-
| Benthopelagic<ref name="Moyle571"/>
| [[Rattail]] and [[Ophidiidae|brotula]] are particularly abundant.
|-
| Benthic
| [[Flatfish]], [[hagfish]], [[eelpout]], [[greeneye]] [[eel]], [[stingray]], [[lumpfish]], and [[Pancake batfish|batfish]]<ref name="Moyle571"/>
|}
{| class="wikitable"
==Behavior and ecology==
|+ '''Comparative structure of pelagic fishes'''
The jaguar is mostly active at night and during [[twilight]].<ref>{{cite journal |author1=Scognamillo, D. |author2=Maxit, I. E. |author3=Sunquist, M. |author4=Polisar, J. |year=2003 |name-list-style=amp |title=Coexistence of jaguar (''Panthera onca'') and puma (''Puma concolor'') in a mosaic landscape in the Venezuelan llanos |journal=Journal of Zoology |volume=259 |issue=3 |pages=269–279 |doi=10.1017/S0952836902003230}}</ref><ref>{{cite journal |author1=Harmsen, B. J. |author2=Foster, R. J. |author3=Silver, S. C. |author4=Ostro, L. E. T. |author5=Doncaster, C. P. |year=2009 |name-list-style=amp |title=Spatial and temporal interactions of sympatric jaguars (''Panthera onca'') and pumas (''Puma concolor'') in a neotropical forest |journal=Journal of Mammalogy |volume=90 |issue=3 |pages=612–620 |doi=10.1644/08-MAMM-A-140R.1 |doi-access=free}}</ref><ref>{{cite journal |author1=Foster, V.C. |author2=Sarmento, P. |author3=Sollmann, R. |author4=Tôrres, N. |author5=Jácomo, A. T. |author6=Negrões, N. |author7=Fonseca, C. |author8=Silveira, L. |year=2013 |display-authors=4 |name-list-style=amp |title=Jaguar and Puma Activity Patterns and Predator-Prey Interactions in Four Brazilian Biomes |journal=Biotropica |volume=45 |issue=3 |pages=373–379 |url=https://www.researchgate.net/publication/234153943 |doi=10.1111/btp.12021 |s2cid=86338173 |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021315/https://www.researchgate.net/publication/234153943_Jaguar_and_Puma_Activity_Patterns_and_Predator-Prey_Interactions_in_Four_Brazilian_Biomes |url-status=live }}</ref>
|-
However, jaguars living in densely forested regions of the [[Amazon Rainforest]] and the [[Pantanal]] are largely active by day, whereas jaguars in the [[Atlantic Forest]] are primarily active by night.<ref name="Astate et al. 2008">{{cite journal |last1=Astete |first1=S.R. |last2=Sollmann |first2=R. |last3=Silveira |first3=L. |year=2008 |name-list-style=amp |title=Comparative ecology of jaguars in Brazil |journal=Cat News |issue=Special Issue 4 |pages=9–14 |citeseerx=10.1.1.528.3603}}</ref>
!
The activity pattern of the jaguar coincides with the activity of its main prey species.<ref name="Harmsen et al. 2011">{{cite journal |last1=Harmsen |first1=B.J. |last2=Foster |first2=R.J. |last3=Silver |first3=S. C. |last4=Ostro |first4=L.E.T. |last5=Doncaster |first5=C.P. |year=2011 |name-list-style=amp |title=Jaguar and puma activity patterns in relation to their main prey |journal=Mammalian Biology – Zeitschrift für Säugetierkunde |volume=76 |issue=3 |pages=320–324 |doi=10.1016/j.mambio.2010.08.007}}</ref> Jaguars are good swimmers and play and hunt in the water, possibly more than tigers. They have been recorded moving between islands and the shore. Jaguars are also good at climbing trees but do so less often than cougars.<ref name=Seymour />
! Epipelagic
! Mesopelagic
! Bathypelagic
! Deep sea [[benthic]]
|-
| muscles
|
| muscular bodies
| poorly developed, flabby
|
|-
| skeleton
|
| strong, ossified bones
| weak, minimal ossification
|
|-
| scales
|
| yes
| none
|
|-
| nervous systems
|
| well developed
| lateral line and olfactory only
|
|-
| eyes
|
| large and sensitive
| small and may not function
| variable (well developed to absent)
|-
| photophores
| absent
| common
| common
| usually absent
|-
| gills
|
| well developed
|
|
|-
| kidneys
|
| large
| small
|
|-
| heart
|
| large
| small
|
|-
| swimbladder
|
| vertically migratory fish have swimbladders
| reduced or absent
| variable (well developed to absent)
|-
| size
|
|
| usually under 25 cm
| variable, species greater than one metre are not uncommon
|}
===EcologicalMesopelagic rolefish===
[[File:california headlightfish.png|thumb|Most mesopelagic fishes are small filter feeders that ascend at night to feed in the nutrient rich waters of the epipelagic zone. During the day, they return to the dark, cold, oxygen-deficient waters of the mesopelagic where they are relatively safe from predators. [[Lanternfish]] account for as much as 65% of all deep sea fish [[biomass]] and are largely responsible for the [[deep scattering layer]] of the world's oceans.]]
[[File:Jaguar (Panthera onca palustris) male Three Brothers River 2.jpg|thumb|Jaguar at [[Three Brothers River]], Pantanal, Brazil]]
[[File:Coccorella atrata.png|thumb|Most of the rest of the mesopelagic fishes are ambush predators, such as this [[sabertooth fish]]. The sabertooth uses its telescopic, upward-pointing eyes to pick out prey silhouetted against the gloom above. Their recurved teeth prevent a captured fish from backing out.]]
The adult jaguar is an [[apex predator]], meaning it is at the top of the [[food chain]] and is not preyed upon in the wild. The jaguar has also been termed a [[keystone species]], as it is assumed that it controls the population levels of prey such as [[Herbivore|herbivorous]] and [[Seed predation|seed-eating]] mammals and thus maintains the structural integrity of forest systems.<ref name=foodhabits /><ref>{{cite journal |author=Nijhawan, S. |year=2012 |title=Conservation units, priority areas and dispersal corridors for jaguars in Brazil |journal=Cat News |issue=Special Issue |pages=43–47 |url=http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.7._SI_7/Nijhawan_2012_Conservation_units_and_corrdors_for_jaguars_in_Brazil.pdf |access-date=17 August 2020 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021300/http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.7._SI_7/Nijhawan_2012_Conservation_units_and_corrdors_for_jaguars_in_Brazil.pdf |url-status=live }}</ref><ref>{{cite journal |author1=Petracca, L.S. |author2=Ramírez-Bravo, O.E. |author3=Hernández-Santín, L. |year=2014 |name-list-style=amp |title=Occupancy estimation of jaguar ''Panthera onca'' to assess the value of east-central Mexico as a jaguar corridor |journal=Oryx |volume=48 |issue=1 |pages=133–140 |doi=10.1017/S0030605313000069 |s2cid=86460403 |doi-access=free}}</ref>
However, field work has shown this may be natural variability, and the population increases may not be sustained. Thus, the [[keystone predator]] hypothesis is not accepted by all scientists.<ref name=r30>{{cite journal |author1=Wright, S. J. |author2=Gompper, M. E. |author3=DeLeon, B. |year=1994 |name-list-style=amp |title=Are large predators keystone species in Neotropical forests? The evidence from Barro Colorado Island |journal=Oikos |volume=71 |issue=2 |pages=279–294 |url=https://www.researchgate.net/publication/270356084 |doi=10.2307/3546277 |jstor=3546277 |access-date=18 June 2019 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021300/https://www.researchgate.net/publication/270356084_Are_Large_Predators_Keystone_Species_in_Neotropical_Forests_The_Evidence_from_Barro_Colorado_Island |url-status=live }}</ref>
Below the epipelagic zone, conditions change rapidly. Between 200 metres and approximately 1000 metres, light continues to fade until darkness is nearly complete. Temperatures fall through a [[thermocline]] to temperatures between {{convert|4|°C|°F}} and {{convert|8|°C|°F}}. This is the twilight or [[mesopelagic]] zone. Pressure continues to increase, at the rate of one atmosphere every 10 metres, while nutrient concentrations fall, along with dissolved oxygen and the rate at which the water circulates.<ref name="Moyle585"/><ref name="salvanes2001" />
The jaguar is [[sympatric]] with the [[cougar]] (''Puma concolor''). In central Mexico, both prey on [[white-tailed deer]] (''Odocoileus virginianus''), which makes up 54% and 66% of jaguar and cougar's prey, respectively.<ref name=foodhabits /> In northern Mexico, the jaguar and the cougar share the same habitat, and their diet overlaps dependent on prey availability. Jaguars seemed to prefer [[deer]] and calves. In Mexico and Central America, neither of the two cats are considered to be the dominant predator.<ref name="GGLG2017">{{cite journal |last1=Gutiérrez-González |first1=C. E. |last2=López-González |first2=C. A. |year=2017 |name-list-style=amp |title=Jaguar interactions with pumas and prey at the northern edge of jaguars' range |journal=[[PeerJ]] |volume=5 |issue=5 |page=e2886 |doi=10.7717/peerj.2886 |pmc=5248577 |pmid=28133569}}</ref>
In South America, the jaguar is larger than the cougar and tends to take larger prey, usually over {{cvt|22|kg}}. The cougar's prey usually weighs between {{cvt|2|and|22|kg|0}}, which is thought to be the reason for its smaller size.<ref name=r31>{{cite journal |author1=Iriarte, J. A. |author2=Franklin, W.L. |author3=Johnson, W.E. |author4=Redford, K.H. |year=1990 |name-list-style=amp |title=Biogeographic variation of food habits and body size of the America puma |journal=[[Oecologia]] |volume=85 |issue=2 |pages=185–190 |bibcode=1990Oecol..85..185I |doi=10.1007/BF00319400 |pmid=28312554 |s2cid=10134066}}</ref>
This situation may be advantageous to the cougar. Its broader prey niche, including its ability to take smaller prey, may give it an advantage over the jaguar in human-altered landscapes.<ref name=foodhabits />
Sonar operators, using the sonar technology developed during World War II, were puzzled by what appeared to be a false sea floor 300–500 metres deep at day, and less deep at night. This turned out to be due to millions of marine organisms, most particularly small mesopelagic fish, with swimbladders that reflected the sonar.
===Hunting and diet===
[[File:Panthera onca at the Toronto Zoo 2.jpg|thumb|The jaguar has a powerful bite that allows it to pierce the shells of armored prey.]]
[[File:3 Jaguars killing a Caiman, Parque Estadual Encontro das Águas Thomas-Fuhrmann.jpg|thumb|Jaguars killing and feeding on a [[yacare caiman]]]]
The jaguar is an [[obligate carnivore]] and depends solely on flesh for its nutrient requirements. An analysis of 53 studies documenting the diet of the jaguar revealed that its prey ranges in weight from {{cvt|1|to|130|kg}}; it prefers prey weighing {{cvt|45-85|kg}}, with [[capybara]] (''Hydrochoerus hydrochaeris'') and [[giant anteater]] (''Myrmecophaga tridactyla'') being the most selected. When available, it also preys on [[marsh deer]] (''Blastocerus dichotomus''), [[southern tamandua]] (''Tamandua tetradactyla''), [[collared peccary]] (''Dicotyles tajacu'') and [[black agouti]] (''Dasyprocta fuliginosa'').<ref name="sizes" /> In floodplains, jaguars opportunistically take reptiles such as [[turtle]]s and [[caiman]]s. Consumption of reptiles appears to be more frequent in jaguars than in other big cats.<ref name="reptile">{{cite journal |last1=Miranda |first1=E |last2=Menezes |first2=J |last3=Rheingantz |first3=M. L. |year=2016 |name-list-style=amp |title=Reptiles as principal prey? Adaptations for durophagy and prey selection by jaguar (''Panthera onca'') |journal=Journal of Natural History |volume=50 |issue=31–32 |pages=2021–2035 |url=https://zenodo.org/record/3993366 |doi=10.1080/00222933.2016.1180717 |s2cid=89150920 |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021309/https://zenodo.org/record/3993366 |url-status=live }}</ref> One remote population in the Brazilian Pantanal is recorded to primarily feed on aquatic reptiles and fish.<ref name="Eriksson2021">{{cite journal|author=Eriksson, C.|display-authors=etal|year=2021|title=Extensive aquatic subsidies lead to territorial breakdown and high density of an apex predator|journal=Ecology|volume=103|issue=1|page=e03543|doi=10.1002/ecy.3543|pmid=34841521|s2cid=242197640}}</ref>
The jaguar also preys on livestock in [[cattle]] [[ranch]]ing areas where wild prey is scarce.<ref>{{cite journal |author1=Amit, R. |author2=Gordillo-Chávez, E.J. |author3=Bone, R. |year=2013 |name-list-style=amp |title=Jaguar and puma attacks on livestock in Costa Rica |journal=Human-Wildlife Interactions |volume=7 |issue=1 |pages=77–84}}</ref><ref>{{cite journal |author1=Zarco-González, M.M. |author2=Monroy-Vilchis, O. |author3=Alaníz, J. |year=2013 |name-list-style=amp |title=Spatial model of livestock predation by jaguar and puma in Mexico: conservation planning |journal=Biological Conservation |volume=159 |pages=80–87 |doi=10.1016/j.biocon.2012.11.007}}</ref>
The daily food requirement of a captive jaguar weighing {{cvt|34|kg}} was estimated at {{cvt|1.4|kg}} of meat.<ref name=Emmons1987 />
Mesopelagic organisms migrate into shallower water at dusk to feed on plankton. The layer is deeper when the moon is out, and may move higher when the sky is dark. This phenomenon has come to be known as the [[deep scattering layer]].<ref name="TeAraMZ">Ryan P [http://www.teara.govt.nz/EarthSeaAndSky/SeaLife/DeepSeaCreatures/2/en "Deep-sea creatures: The mesopelagic zone"] ''Te Ara – the Encyclopedia of New Zealand''. Updated 21 September 2007.</ref>
The jaguar's bite force allows it to pierce the [[Turtle shell#Carapace|carapace]]s of the [[yellow-spotted Amazon river turtle]] (''Podocnemis unifilis'') and the [[yellow-footed tortoise]] (''Chelonoidis denticulatus'').<ref name=Emmons1987>{{cite journal |author=Emmons, L. H. |year=1987 |title=Comparative feeding ecology of fields in a neotropical rain forest |journal=Behavioral Ecology and Sociobiology |volume=20 |issue=4 |pages=271–283 |url=https://www.researchgate.net/publication/225982805 |doi=10.1007/BF00292180 |s2cid=24990860 |access-date=13 April 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021300/https://www.researchgate.net/publication/225982805_Comparative_Feeding_Ecology_of_Felids_in_a_Neotropical_Rain-Forest |url-status=live}}</ref><ref>{{cite journal |author=Emmons, L. H. |year=1989 |title=Jaguar predation on chelonians |journal=Journal of Herpetology |volume=23 |issue=3 |pages=311–314 |doi=10.2307/1564460 |jstor=1564460}}</ref> It employs an unusual killing method: it bites mammalian prey directly through the [[skull]] between the ears to deliver a fatal bite to the brain.<ref name=rosa>{{cite book |last1=Rosa |first1=C. L. de la |last2=Nocke |first2=C. C. |year=2000 |name-list-style=amp |chapter=Jaguar (''Panthera onca'') |title=A guide to the carnivores of Central America: natural history, ecology, and conservation |publisher=[[University of Texas Press]] |chapter-url=https://books.google.com/books?id=x5ihAwAAQBAJ&pg=PT39 |pages=39–? |isbn=978-0-292-71604-9 |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021333/https://books.google.com/books?id=x5ihAwAAQBAJ&pg=PT39 |url-status=live}}</ref> It kills capybara by piercing its [[canine tooth|canine teeth]] through the [[temporal bone]]s of its skull, breaking its [[zygomatic arch]] and [[mandible]] and penetrating its brain, often through the ears.<ref>{{cite journal |author1=Schaller, G.B. |author1-link=George Schaller |author2=Vasconselos, J.M.C. |year=1978 |name-list-style=amp |title=Jaguar predation on capybara |journal=Zeitschrift für Säugetierkunde |volume=43 |pages=296–301 |url=https://www.zobodat.at/pdf/Zeitschrift-Saeugetierkunde_43_0296-0301.pdf |access-date=16 August 2020 |archive-date=25 June 2021 |archive-url=https://web.archive.org/web/20210625121230/https://www.zobodat.at/pdf/Zeitschrift-Saeugetierkunde_43_0296-0301.pdf |url-status=live }}</ref>
It has been hypothesized to be an adaptation to "cracking open" turtle shells; armored reptiles may have formed an abundant prey base for the jaguar following the late [[Pleistocene extinction]]s.<ref name=Emmons1987 /> However, this is disputed, as even in areas where jaguars prey on reptiles, they are still taken relatively infrequently compared to mammals in spite of their greater abundance.<ref name="reptile" />
Most mesopelagic fish make [[Diel vertical migration|daily vertical migrations]], moving each night into the epipelagic zone, often following similar migrations of zooplankton, and returning to the depths for safety during the day.<ref name="salvanes2001" /><ref name="Moyle585" /><ref>[[#Bone|Bone and Moore]], p. 38.</ref> These vertical migrations occur over hundreds of meters.
Between October 2001 and April 2004, 10 jaguars were monitored in the southern Pantanal. In the dry season from April to September, they killed prey at intervals ranging from one to seven days; and ranging from one to 16 days in the wet season from October to March.<ref>{{cite journal |author1=Cavalcanti, S. M. C. |author2=Gese, E. M. |year=2010 |name-list-style=amp |title=Kill rates and predation patterns of jaguars (''Panthera onca'') in the southern Pantanal, Brazil |journal=Journal of Mammalogy |volume=91 |issue=3 |pages=722–736 |url=https://www.academia.edu/3122316 |doi=10.1644/09-MAMM-A-171.1 |doi-access=free |access-date=8 March 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021307/https://www.academia.edu/3122316 |url-status=live }}</ref>
These fish have muscular bodies, ossified bones, scales, well developed gills and central nervous systems, and large hearts and kidneys. Mesopelagic [[Filter feeder|plankton feeders]] have small mouths with fine [[gill raker]]s, while the [[piscivore]]s have larger mouths and coarser gill rakers.<ref name="salvanes2001">{{cite journal |title=Mesopelagic Fishes |journal=Encyclopedia of Ocean Sciences |year=2001 |last1=Salvanes |first1=A.G.V. |last2=Kristoffersen |first2=J.B. |volume=3 |url=http://earthguide.ucsd.edu/fishes/environment/0_images/Original/myctophids/salvanes_01.pdf |access-date=4 November 2020 }}</ref><ref name="Moyle585" />
The jaguar uses a stalk-and-ambush strategy when hunting rather than chasing prey. The cat will slowly walk down forest paths, listening for and stalking prey before rushing or ambushing. The jaguar attacks from cover and usually from a target's blind spot with a quick pounce; the species' ambushing abilities are considered nearly peerless in the animal kingdom by both indigenous people and field researchers and are probably a product of its role as an apex predator in several different environments. The ambush may include leaping into water after prey, as a jaguar is quite capable of carrying a large kill while swimming; its strength is such that carcasses as large as a heifer can be hauled up a tree to avoid flood levels. After killing prey, the jaguar will drag the carcass to a [[thicket]] or other secluded spot. It begins eating at the neck and chest. The heart and lungs are consumed, followed by the shoulders.<ref>{{cite book| title = Guidelines for Captive Management of Jaguars |author=Baker, W. K. Jr.|pages=8–16 |editor=Law, C. |publisher=Jaguar Species Survival Plan. [[Association of Zoos and Aquariums]] |url=http://www.jaguarssp.com/Animal%20Mgmt/JAGUAR%20GUIDELINES.pdf |url-status = dead |archive-url=https://web.archive.org/web/20120113131137/http://www.jaguarssp.com/Animal%20Mgmt/JAGUAR%20GUIDELINES.pdf
|archive-date=13 January 2012}}</ref>
Vertically migratory fish have [[swimbladder]]s.<ref name="Haedrich1996">{{Cite journal | doi = 10.1111/j.1095-8649.1996.tb06066.x| title = Deep-water fishes: Evolution and adaptation in the earth's largest living spaces| journal = Journal of Fish Biology| volume = 49| pages = 40–53| year = 1996| last1 = Haedrich | first1 = R. L.}}
===Social activity===
</ref> The fish inflates its swimbladder to move up. Given the high pressures in the mesopelagic zone, this requires significant energy. As the fish ascends, the air in the swimbladder must decrease to prevent the swimbladder from bursting. To return to the depths, the swimbladder is deflated.<ref>{{Cite journal |last1=Douglas |first1=E. |last2=Friedl |first2=W. |last3=Pickwell |first3=G. |year=1976 |title=Fishes in oxygen-minimum zones: Blood oxygenation characteristics |journal=[[Science (journal)|Science]] |volume=191 |issue=4230 |pages=957–9 |bibcode=1976Sci...191..957D |doi=10.1126/science.1251208 |pmid=1251208}}</ref> The migration takes them through the [[thermocline]], where the temperature changes between 10 and 20 °C, thus displaying considerable temperature tolerance.<ref name="Moyle590">[[Pelagic fish#Moyle|Moyle and Cech]], p. 590</ref>
[[File:Jaguars (Panthera onca) after fight ...(Female left) (29070165892).jpg|thumb|Female (left) and male jaguar (right) at São Lourenço River]]
The jaguar is generally [[Solitary animal|solitary]] except for females with cubs. In 1977, groups consisting of a male, female and cubs, and two females with two males were sighted several times in a study area in the [[Paraguay River]] valley. A radio-collared female moved in a [[home range]] of {{cvt|25-38|km2}}, which partly overlapped with another female. The home range of the male in this study area overlapped with several females.<ref>{{cite journal |author1=Schaller, G. B. |author2=Crawshaw, P. G. Jr. |year=1980 |name-list-style=amp |title=Movement Patterns of Jaguar |journal=Biotropica |volume=12 |issue=3 |pages=161–168 |doi=10.2307/2387967 |jstor=2387967}}</ref>
Mesopelagic fish are adapted for an active life under low light conditions. Most of them are visual predators with large eyes. Some of the deeper water fish such as the [[Telescopefish]] have tubular eyes with big lenses and only [[rod cell]]s that look upward. These give binocular vision and great sensitivity to small light signals.<ref name="Moyle585" /> This adaptation gives improved terminal vision at the expense of lateral vision, and allows the predator to pick out [[squid]], [[cuttlefish]], and smaller fish that are silhouetted above them.<ref name="salvanes2001" />
The jaguar uses scrape marks, urine, and feces to [[Territorial marking|mark its territory]].<ref name=Rabinowitz>{{cite journal |author1=Rabinowitz, A. R. |author1-link=Alan Rabinowitz |author2=Nottingham, B.G. Jr. |year=1986 |name-list-style=amp |title=Ecology and behaviour of the Jaguar (''Panthera onca'') in Belize, Central America |journal=Journal of Zoology |volume=210 |issue=1 |pages=149–159 |doi=10.1111/j.1469-7998.1986.tb03627.x}}</ref><ref>{{cite journal |last1=Harmsen |first1=B. J. |last2=Foster |first2=R.J. |last3=Gutierrez |first3=S.M. |last4=Marin |first4=S.Y. |last5=Doncaster |first5=C.P. |year=2007 |name-list-style=amp |title=Scrape-marking behavior of jaguars (''Panthera onca'') and pumas (''Puma concolor'') |journal=Journal of Mammalogy |volume=91 |issue=5 |pages=1225–1234 |doi=10.1644/09-mamm-a-416.1 |doi-access=free}}</ref>
The size of home ranges depends on the level of deforestation and human population density. The home ranges of females vary from {{cvt|15.3|km2}} in the [[Pantanal]] to {{cvt|53.6|km2}} in the Amazon to {{cvt|233.5|km2}} in the [[Atlantic Forest]]. Male jaguar home ranges vary from {{cvt|25|km2}} in the Pantanal to {{cvt|180.3|km2}} in the Amazon to {{cvt|591.4|km2}} in the Atlantic Forest and {{cvt|807.4|km2}} in the [[Cerrado]].<ref>{{cite journal |author1=Morato, R.G. |author2=Stabach, J.A. |author3=Fleming, C.H. |author4=Calabrese, J.M. |author5=De Paula, R.C. |author6=Ferraz, K.M. |author7=Kantek, D.L. |author8=Miyazaki, S.S. |author9=Pereira, T.D. |author10=Araujo, G.R. |author11=Paviolo, A. |year=2016 |display-authors=4 |name-list-style=amp |title=Space use and movement of a neotropical top predator: the endangered Jaguar |journal=PLOS ONE |volume=11 |issue=12 |page=e0168176 |doi=10.1371/journal.pone.0168176 |doi-access=free |pmc=5193337 |pmid=28030568 |bibcode=2016PLoSO..1168176M}}</ref>
Studies employing [[Global Positioning System|GPS]] [[telemetry]] in 2003 and 2004 found densities of only six to seven jaguars per 100 km<sup>2</sup> in the Pantanal region, compared with 10 to 11 using traditional methods; this suggests the widely used sampling methods may inflate the actual numbers of individuals in a sampling area.<ref name=soisalo /> Fights between males occur but are rare, and avoidance behavior has been observed in the wild.<ref name=Rabinowitz /> In one wetland population with degraded territorial boundaries and more social proximity, adults of the same sex are more tolerant of each other and engage in more friendly and co-operative interactions.<ref name="Eriksson2021"/>
[[File:Jaguar saw.flac|thumb|Captive jaguar vocalizing while playing]]
The jaguar [[Roar (vocalization)|roars]] or grunts for long-distance communication;<ref name=Seymour /><ref name=Emmons1987 /> intensive bouts of counter-calling between individuals have been observed in the wild.<ref name=Emmons1987 /> This vocalization is described as "hoarse" with five or six [[guttural]] notes.<ref name=Seymour /> [[prusten|Chuffing]] is produced by individuals when greeting, during [[Courtship#In animals|courting]], or by a mother comforting her cubs. This sound is described as low intensity snorts, possibly intended to signal tranquility and passivity.<ref name=":0">{{Cite journal |last1=Peters |first1=G. |last2=Tonin-Leyhausen |first2=B. |year=1999 |name-list-style=amp |title=Evolution of Acoustic Communication Signals of Mammals: Friendly Close-Range Vocalizations in Felidae (Carnivora) |journal=Journal of Mammalian Evolution |volume=6 |issue=2 |pages=129–159 |doi=10.1023/A:1020620121416 |s2cid=25252052}}</ref><ref>{{cite journal |last1=Leuchtenberger |first1=C. |last2=Crawshaw |first2=P. G. |last3=Mourão |first3=G. |last4=Lehn |first4=C. R. |year=2009 |name-list-style=amp |title=Courtship behavior by Jaguars in the Pantanal of Mato Grosso do Sul |journal=Natureza & Conservaç~ao Revista Brasileira de Conservaç~ao da Natureza |volume=7 |issue=1 |pages=218–222 |url=https://www.researchgate.net/publication/263424625 |access-date=13 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021418/https://www.researchgate.net/publication/263424625_Courtship_behavior_by_Jaguars_in_the_Pantanal_of_Mato_Grosso_do_Sul |url-status=live }}</ref> Cubs have been recorded bleating, gurgling and mewing.<ref name=Seymour />
Mesopelagic fish usually lack defensive spines, and use colour for [[camouflage]].<ref name="salvanes2001" /> [[Ambush predator]]s are dark, black or red. Since the longer, red, wavelengths of light do not reach the deep sea, red effectively functions the same as black. Migratory forms use [[Countershading|countershaded]] silvery colours. On their bellies, they often display [[photophore]]s producing low grade light. For a predator from below, looking upward, this [[bioluminescence]] camouflages the silhouette of the fish. However, some of these predators have yellow lenses that filter the (red deficient) ambient light, leaving the bioluminescence visible.<ref>{{Cite journal | doi = 10.1017/S0025315400021019| title = On yellow lenses in mesopelagic animals| journal = Journal of the Marine Biological Association of the United Kingdom| volume = 56| issue = 4| pages = 963–976| year = 2009| last1 = Muntz | first1 = W. R. A. | s2cid = 86353657}}</ref>
===Reproduction and life cycle===
[[File:Jaguarpickingupcub08.jpg|thumb|Female jaguar picking up her cub]]
In captivity, the female jaguar is recorded to reach [[sexual maturity]] at the age of about 2.5 years. [[Estrus]] lasts 7–15 days with an [[estrus cycle]] of 41.8 to 52.6 days. During estrus, she exhibits increased restlessness with rolling and prolonged vocalizations.<ref>{{cite journal |author1=Wildt, D.E. |author2=Platz, C.C. |author3=Chakraborty, P.K. |author4=Seager, S.W.J. |year=1979 |name-list-style=amp |title=Oestrous and ovarian activity in a female jaguar (''Panthera onca'') |journal=Reproduction |volume=56 |issue=2 |pages=555–558 |doi=10.1530/jrf.0.0560555 |doi-access=free |pmid=383976}}</ref>
She is an [[induced ovulator]] but can also ovulate spontaneously.<ref>{{cite journal |author1=Barnes, S.A. |author2=Teare, J.A. |author3=Staaden, S. |author4=Metrione, L. |author5=Penfold, L.M. |year=2016 |name-list-style=amp |title=Characterization and manipulation of reproductive cycles in the jaguar (''Panthera onca'') |journal=General and Comparative Endocrinology |volume=225 |pages=95–103 |doi=10.1016/j.ygcen.2015.09.012 |doi-access=free |pmid=26399935}}</ref>
[[Gestation]] lasts 91 to 111 days.<ref>{{cite book |author=Hemmer, H. |year=1976 |chapter=Gestation period and postnatal development in felids |title=The world's cats |editor=Eaton, R.L. |publisher=Carnivore Research Institute, Univ. Washington |___location=Seattle |volume=3. Contributions to Biology, Ecology, Behavior and Evolution |pages=143–165}}</ref>
The male is sexually mature at the age of three to four years.<ref>{{cite book |author1=Mondolfi, E. |author2=Hoogesteijn, R. |year=1986 |name-list-style=amp |chapter=Notes on the biology and status of the jaguar (''Panthera onca'') in Venezuela |title=Cats of the world: biology, conservation and management |editor1=Miller, S.D. |editor2=Everett, D.D. |publisher=National Wildlife Federation |___location=Washington, DC |pages=85–123 |isbn=978-091218679-5}}</ref>
His mean [[ejaculate]] volume is 8.6±1.3 ml.<ref>{{cite journal |author1=Morato, R. G. |author2=Guimaraes, M.A.B. |author3=Ferriera, F. |author4=Verreschi, I.T.d.N. |author5=Barnabe, R.C. |year=1999 |name-list-style=amp |title=Reproductive characteristics of captive male jaguars |journal=Brazilian Journal of Veterinary Research and Animal Science |volume=36 |issue=5 |doi=10.1590/S1413-95961999000500008 |doi-access=free}}</ref>
[[Generation time|Generation length]] of the jaguar is 9.8 years.<ref>{{cite journal |author1=Pacifici, M. |author2=Santini, L. |author3=Di Marco, M. |author4=Baisero, D. |author5=Francucci, L. |author6=Grottolo Marasini, G. |author7=Visconti, P. |author8=Rondinini, C. |name-list-style=amp |year=2013 |title=Generation length for mammals |journal=Nature Conservation |volume=5 |issue=5 |pages=87–94 |doi=10.3897/natureconservation.5.5734 |doi-access=free}}</ref>
<gallery class="left" widths="200px" heights="175px">
In the Pantanal, [[breeding pair]]s were observed to stay together for up to five days. Females had one to two cubs.<ref>{{cite journal |author1=Cavalcanti, S. M. C. |author2=Gese, E. M. |year=2009 |name-list-style=amp |title=Spatial ecology and social interactions of jaguars (''Panthera onca'') in the southern Pantanal, Brazil |journal=Journal of Mammalogy |volume=90 |issue=4 |pages=935–945 |doi=10.1644/08-MAMM-A-188.1 |doi-access=free}}</ref>
File:Dmawsoni Head shot.jpg|The [[Antarctic toothfish]] have large, upward looking eyes, adapted to detecting the silhouettes of prey fish.<ref>{{FishBase species|genus=Dissostichus|species=mawsoni|year=2009|month=August}}</ref>
The young are born with closed eyes but open them after two weeks. Cubs are [[wean]]ed at the age of three months but remain in the birth den for six months before leaving to accompany their mother on hunts.<ref name=SWWL>{{cite magazine |author=Egerton, J. |date=2006 |title=Jaguars: Magnificence in the Southwest |magazine=Wild Tracks |url=http://www.southwestwildlife.org/pdf/Newsletter/Spring06.pdf |url-status=dead |archive-url=https://web.archive.org/web/20110721144027/http://www.southwestwildlife.org/pdf/Newsletter/Spring06.pdf |access-date=6 December 2009 |archive-date=21 July 2011}}</ref>
File:Opisthoproctus soleatus.png|The [[Barreleye]] has barrel-shaped, tubular [[eye]]s that generally are directed upward, but may be swivelled forward.<ref>[https://www.sciencedaily.com/releases/2009/02/090223150331.htm Mystery Of Deep-sea Fish With Tubular Eyes And Transparent Head Solved] ''ScienceDaily'', 24 February 2009.</ref>
Jaguars remain with their mothers for up to two years. They appear to rarely live beyond 11 years, but captive individuals may live 22 years.<ref name=Seymour />
Image:Malacosteus niger.jpg|The [[stoplight loosejaw]] has a [[lower jaw]] one-quarter as long as its body. The jaw has no floor and is attached only by a hinge and a modified tongue bone. Large fang-like teeth in the front are followed by many small barbed teeth.<ref name="kenaley">{{cite journal|author=Kenaley, C.P|title=Revision of the Stoplight Loosejaw Genus ''Malacosteus'' (Teleostei: Stomiidae: Malacosteinae), with Description of a New Species from the Temperate Southern Hemisphere and Indian Ocean|journal=Copeia|volume=2007|issue=4|pages=886–900|year=2007|doi=10.1643/0045-8511(2007)7[886:ROTSLG]2.0.CO;2|s2cid=1038874 }}</ref><ref name="sutton">{{cite journal|author=Sutton, T.T.|title=Trophic ecology of the deep-sea fish ''Malacosteus niger'' (Pisces: Stomiidae): An enigmatic feeding ecology to facilitate a unique visual system?|journal=Deep-Sea Research Part I: Oceanographic Research Papers|volume=52|issue=11|date=Nov 2005|pages=2065–2076|doi=10.1016/j.dsr.2005.06.011|bibcode=2005DSRI...52.2065S|url=https://nsuworks.nova.edu/occ_facpresentations/244}}</ref>
Image:Malacosteus.JPG|The [[stoplight loosejaw]] is also one of the few fishes that produce red [[bioluminescence]]. As most of their prey cannot perceive red light, this allows it to hunt with an essentially invisible beam of light.<ref name="kenaley"/>
</gallery>
The [[brownsnout spookfish]] is a species of [[barreleye]] and is the only vertebrate known to employ a mirror, as opposed to a lens, to focus an image in its eyes.<ref name="wagner et al">{{cite journal |author=Wagner, H.J., Douglas, R.H., Frank, T.M., Roberts, N.W., and Partridge, J.C. |title=A Novel Vertebrate Eye Using Both Refractive and Reflective Optics |journal=Current Biology |volume=19 |pages=108–114 |date=27 January 2009 | pmid = 19110427 | doi = 10.1016/j.cub.2008.11.061 |issue=2
In 2001, a male jaguar killed and partially consumed two cubs in [[Emas National Park]]. [[DNA paternity testing]] of blood samples revealed that the male was the father of the cubs.<ref>{{cite journal |author1=Soares, T. N. |author2=Telles, M. P. |author3=Resende, L.V. |author4=Silveira, L. |author5=Jácomo, A.T.A. |author6=Morato, R.G. |author7=Diniz-Filho, J.A.F. |author8=Eizirik, E. |author9=Brondani, R.P. |author10=Brondani, C. |year=2006 |display-authors=4 |name-list-style=amp |title=Paternity testing and behavioral ecology: A case study of jaguars (''Panthera onca'') in Emas National Park, Central Brazil |journal=Genetics and Molecular Biology |volume=29 |issue=4 |pages=735–740 |url=https://www.scielo.br/pdf/gmb/v29n4/32128.pdf |doi=10.1590/S1415-47572006000400025 |doi-access=free |access-date=27 February 2021 |archive-date=27 October 2020 |archive-url=https://web.archive.org/web/20201027093753/https://www.scielo.br/pdf/gmb/v29n4/32128.pdf |url-status=live }}</ref> Two more cases of [[Infanticide (zoology)|infanticide]] were documented in the northern Pantanal in 2013.<ref>{{cite journal |author1=Tortato, F.R. |author2=Devlin, A.L. |author3=Hoogesteijn, R. |author4=Júnior, J.A.M. |author5=Frair, J.L. |author6=Crawshaw, P.G. |author7=Izzo, T.J. |author8=Quigley, H.B. |year=2017 |name-list-style=amp |title=Infanticide in a jaguar (''Panthera onca'') population – does the provision of livestock carcasses increase the risk? |journal=Acta Ethologica |volume=20 |issue=1 |pages=69–73 |doi=10.1007/s10211-016-0241-4 |s2cid=34002056 |url=https://www.researchgate.net/publication/308940647 |access-date=27 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021302/https://www.researchgate.net/publication/308940647_Infanticide_in_a_jaguar_Panthera_onca_population-does_the_provision_of_livestock_carcasses_increase_the_risk |url-status=live }}</ref> Infanticide may be combated by the female hiding her cubs and distracting the male with courtship behavior.<ref>{{cite journal |author1=Stasiukynas, D. C. |author2=Boron, V. |author3=Hoogesteijn, R. |author4= Barragán2, J. |author5=Martin, A. |author6=Tortato, F. |author7= Rincón, S. |author8=Payán, E.|year=2021|title=Hide and flirt: observed behavior of female jaguars (''Panthera onca'') to protect their young cubs from adult males|journal=Acta Ethologica|volume=25 |issue=3 |pages=179–183 |doi=10.1007/s10211-021-00384-9|s2cid=239539707 }}</ref>
|s2cid=18680315 |doi-access=free }}</ref><ref name="smith">Smith, L. (8 January 2009). [https://web.archive.org/web/20110629123143/http://www.timesonline.co.uk/tol/news/environment/article5469077.ece "Fish with four eyes can see through the deep sea gloom"]. ''Times Online''. Times Newspapers Ltd. Retrieved 14 March 2009.</ref>
Sampling via deep [[trawling]] indicates that [[lanternfish]] account for as much as 65% of all deep sea fish [[biomass]].<ref name=EoF>{{cite book |editor=Paxton, J.R. |editor2=Eschmeyer, W.N.|author= Hulley, P. Alexander|year=1998|title=Encyclopedia of Fishes|publisher= Academic Press|___location=San Diego|pages= 127–128|isbn= 978-0-12-547665-2}}</ref> Indeed, lanternfish are among the most widely distributed, populous, and diverse of all [[vertebrate]]s, playing an important [[ecology|ecological]] role as prey for larger organisms. The estimated global biomass of lanternfish is 550–660 million [[tonne]]s, several times the entire world fisheries catch. Lanternfish also account for much of the biomass responsible for the [[deep scattering layer]] of the world's oceans. [[Sonar]] reflects off the millions of lanternfish [[swim bladder]]s, giving the appearance of a false bottom.<ref>{{cite web | title = Deep-sea fish diversity and ecology in the benthic boundary layer |author1=Cornejo, R. |author2=Koppelmann, R. |author3=Sutton, T. |name-list-style=amp |year=2006 | url = http://www.agu.org/meetings/os06/os06-sessions/os06_OS45Q.html}}</ref>
===Attacks on humans===
{{Further|Man-eater}}
The Spanish [[conquistador]]s feared the jaguar. According to [[Charles Darwin]], the [[indigenous peoples of South America]] stated that people did not need to fear the jaguar as long as capybaras were abundant.<ref name="Porter1894">{{cite book |author=Porter, J. H. |year=1894 |chapter=The Jaguar |title=Wild beasts; a study of the characters and habits of the elephant, lion, leopard, panther, jaguar, tiger, puma, wolf, and grizzly bear |publisher=C. Scribner's sons |___location=New York |chapter-url=https://archive.org/stream/wildbeastsstud00port#page/n197/mode/2up |pages=174–195}}</ref>
The first official record of a jaguar killing a human in Brazil dates to June 2008.<ref>{{cite journal |author1=de Paula, R. |author2=Campos Neto, M. F. |author3=Morato, R. G. |year=2008 |name-list-style=amp |title=First Official Record of Human Killed by Jaguar in Brazil |journal=Cat News |issue=49 |pages=31–32}}</ref>
Two children were attacked by jaguars in Guyana.<ref name=r22>{{cite journal |author1=Iserson, K. V. |author2=Francis, A. M. |year=2015 |name-list-style=amp |title=Jaguar Attack on a Child: Case Report and Literature Review |journal=Western Journal of Emergency Medicine |volume=16 |issue=2 |pages=303–309 |doi=10.5811/westjem.2015.1.24043 |pmc=4380383 |pmid=25834674}}</ref>
The jaguar is the least likely of all big cats to kill and eat humans, and the majority of attacks come when it has been cornered or wounded.<ref>{{Cite book |last1=Seidensticker |first1=J. |last2=Lumpkin |first2=S. |date=2016 |name-list-style=amp |title=Cats in Question: The Smithsonian Answer Book |publisher=Smithsonian Institution |___location=Washington, D.C. |url=https://books.google.com/books?id=09LwCwAAQBAJ&pg=PT303 |isbn=978-158834546-2}}</ref>
The 2010 Malaspina Circumnavigation Expedition traveled 60,000 km, undertaking acoustic observations. It reported that mesopelagic biomass was 10 billion tonnes or more (10x prior estimates), comprising about 90 percent of all ocean fish biomass.<ref>{{Cite journal |last=Duarte |first=Carlos M. |date=2015-01-28 |title=Seafaring in the 21St Century: The Malaspina 2010 Circumnavigation Expedition |url=http://dx.doi.org/10.1002/lob.10008 |journal=Limnology and Oceanography Bulletin |volume=24 |issue=1 |pages=11–14 |doi=10.1002/lob.10008 |issn=1539-607X|hdl=10754/347123 |hdl-access=free }}</ref> Estimates of how much carbon these fish sequester remained highly uncertain as of 2024.<ref name=":0">{{Cite web |last=Donovan |first=Moira |date=November 21, 2023 |title=All the Fish We Cannot See |url=https://hakaimagazine.com/features/all-the-fish-we-cannot-see/ |access-date=2024-02-19 |website=Hakai Magazine |language=en}}</ref>
==Threats==
[[File:Colonel Roosevelt's first South American jaguar.jpg|thumb|A South American jaguar killed by [[Theodore Roosevelt]]]]
The jaguar is threatened by [[habitat loss|loss]] and [[habitat fragmentation|fragmentation]] of habitat, illegal killing in retaliation for livestock depredation and for illegal trade in jaguar body parts. It is listed as [[Near Threatened]] on the [[IUCN Red List]] since 2002, as the jaguar population has probably declined by 20–25% since the mid-1990s. [[Deforestation]] is a major threat to the jaguar across its range. Habitat loss was most rapid in drier regions such as the Argentine [[pampas]], the arid grasslands of Mexico and the southwestern United States.<ref name=iucn />
Mesopelagic fish do not constitute a major fishery as of 2024. Initial efforts in Iceland, Norway, and the Soviet Union did not create a commercial industry. The European Union funded the MEESO project to study abundance and fishing technologies for key mesopelagic species. To date, fish that appeal to the human palate have not been identified, leading harvesters to focus on animal feed markets instead.<ref name=":0" />
In 2002, it was estimated that the range of the jaguar had declined to about 46% of its range in the early 20th century.<ref name=Sanderson2002 /> In 2018, it was estimated that its range had declined by 55% in the last century. The only remaining stronghold is the Amazon rainforest, a region that is rapidly being fragmented by deforestation.<ref>{{cite journal |author1=De La Torre, J.A. |author2=González-Maya, J.F. |author3=Zarza, H. |author4=Ceballos, G. |author5=Medellín, R.A. |year=2018 |name-list-style=amp |title=The jaguar's spots are darker than they appear: assessing the global conservation status of the jaguar ''Panthera onca'' |journal=Oryx |volume=52 |issue=2 |pages=300–315 |doi=10.1017/S0030605316001046 |doi-access=free}}</ref>
Between 2000 and 2012, forest loss in the jaguar range amounted to {{convert|83.759|km2|abbr=on}}, with fragmentation increasing in particular in corridors between Jaguar Conservation Units (JCUs).<ref>{{cite journal |author1=Olsoy, P.J. |author2=Zeller, K.A. |author3=Hicke, J.A. |author4=Quigley, H.B. |author5=Rabinowitz, A.R. |author6=Thornton, D.H. |year=2016 |name-list-style=amp |title=Quantifying the effects of deforestation and fragmentation on a range-wide conservation plan for jaguars |journal=Biological Conservation |volume=203 |pages=8–16 |url=https://www.researchgate.net/publication/307954092 |doi=10.1016/j.biocon.2016.08.037 |access-date=13 April 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021307/https://www.researchgate.net/publication/307954092_Quantifying_the_effects_of_deforestation_and_fragmentation_on_a_range-wide_conservation_plan_for_jaguars |url-status=live }}</ref>
By 2014, direct linkages between two JCUs in Bolivia were lost, and two JCUs in northern Argentina became completely isolated due to deforestation.<ref>{{cite journal |author1=Thompson, J.J. |author2=Velilla, M. |year=2017 |name-list-style=amp |title=Modeling the effects of deforestation on the connectivity of jaguar ''Panthera onca'' populations at the southern extent of the species' range |journal=Endangered Species Research |volume=34 |pages=109–121 |url=https://www.int-res.com/articles/esr2017/34/n034p109.pdf |doi=10.3354/esr00840 |doi-access=free |access-date=2 March 2021 |archive-date=8 March 2021 |archive-url=https://web.archive.org/web/20210308132647/https://www.int-res.com/articles/esr2017/34/n034p109.pdf |url-status=live }}</ref>
[[Bigeye tuna]] are an epipelagic/mesopelagic species that is carnivorous, eating other fish. Satellite tagging has shown that bigeye tuna often spend prolonged periods cruising deep below the surface during the daytime, sometimes making dives as deep as {{Convert|500|m|ft|abbr=off}}. These movements are thought to be in response to the vertical migrations of prey organisms in the [[deep scattering layer]].
In Mexico, the jaguar is primarily threatened by [[poaching]]. Its habitat is fragmented in northern Mexico, in the [[Gulf of Mexico]] and the [[Yucatán Peninsula]], caused by changes in land use, construction of roads and tourism infrastructure.<ref name=Ceballos2016>{{cite book |author1=Ceballos, G. |author2=Zarza, H. |author3=Chávez, C. |author4=González-Maya, J.F. |year=2016 |name-list-style=amp |chapter=Ecology and Conservation of Jaguars in Mexico |title=Tropical conservation: Perspectives on local and global priorities |editor1=Aguirre, A. |editor2=Sukumar, R. |publisher=Oxford University Press |chapter-url=https://www.researchgate.net/publication/307985045 |pages=273–289 |isbn=978-019976698-7 |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021317/https://www.researchgate.net/publication/307985045_Ecology_and_conservation_of_jaguars_in_Mexico_state_of_knowledge_and_future_challenges |url-status=live }}</ref>
In Panama, 220 of 230 jaguars were killed in retaliation for predation on livestock between 1998 and 2014.<ref>{{cite journal |author1=Moreno, R. |author2=Meyer, N. |author3=Olmos, M. |author4=Hoogesteijn, R. |author5=Hoogesteijn, A.L. |year=2015 |name-list-style=amp |title=Causes of jaguar killing in Panama – a long term survey using interviews |journal=Cat News |issue=62 |pages=40–42 |url=https://repository.si.edu/bitstream/handle/10088/29640/CN62_Moreno_et_al_2015.pdf |access-date=18 November 2021 |archive-date=18 November 2021 |archive-url=https://web.archive.org/web/20211118072049/https://repository.si.edu/bitstream/handle/10088/29640/CN62_Moreno_et_al_2015.pdf |url-status=live }}</ref>
In Venezuela, the jaguar was extirpated in about 26% of its range in the country since 1940, mostly in dry [[savanna]]s and unproductive scrubland in the northeastern region of [[Anzoátegui]].<ref>{{cite journal |last1=Jędrzejewski |first1=W. |last2=Boede |first2=E.O. |last3=Abarca |first3=M. |last4=Sánchez-Mercado |first4=A. |last5=Ferrer-Paris |first5=J.R. |last6=Lampo |first6=M. |last7=Velásquez |first7=G. |last8=Carreño |first8=R. |last9=Viloria |first9=Á.L. |last10=Hoogesteijn |first10=R. |last11=Robinson |first11=H.S. |last12=Stachowicz |first12=I. |last13=Cerda |first13=H. |last14=Weisz |first14=M. del Mar |last15=Barros |first15=T.R. |last16=Rivas |first16=Gilson A. |last17=Borges |first17=G. |last18=Molinari |first18=J. |last19=Lew |first19=D. |last20=Takiff |first20=H. |last21=Schmidt |first21=K. |year=2017 |name-list-style=amp |title=Predicting carnivore distribution and extirpation rate based on human impacts and productivity factors; assessment of the state of jaguar (''Panthera onca'') in Venezuela |journal=Biological Conservation |volume=206 |pages=132–142 |url=https://www.researchgate.net/publication/312059394 |doi=10.1016/j.biocon.2016.09.027 |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021303/https://www.researchgate.net/publication/312059394_Predicting_carnivore_distribution_and_extirpation_rate_based_on_human_impacts_and_productivity_factors_assessment_of_the_state_of_jaguar_Panthera_onca_in_Venezuela |url-status=live }}</ref>
In Ecuador, the jaguar is threatened by reduced prey availability in areas where the expansion of the road network facilitated access of human hunters to forests.<ref>{{cite journal |author1=Espinosa, S. |author2=Celis, G. |author3=Branch, L.C. |year=2018 |name-list-style=amp |title=When roads appear jaguars decline: Increased access to an Amazonian wilderness area reduces potential for jaguar conservation |journal=PLOS ONE |volume=13 |issue=1 |page=e0189740 |doi=10.1371/journal.pone.0189740 |doi-access=free |pmc=5751993 |pmid=29298311 |bibcode=2018PLoSO..1389740E}}</ref>
In the [[Alto Paraná Atlantic forests]], at least 117 jaguars were killed in [[Iguaçu National Park]] and the adjacent [[Misiones Province]] between 1995 and 2008.<ref>{{cite journal |author1=Paviolo, A. |author2=De Angelo, C.D. |author3=Di Blanco, Y.E. |author4=Di Bitetti, M.S. |year=2008 |name-list-style=amp |title=Jaguar ''Panthera onca'' population decline in the Upper Paraná Atlantic Forest of Argentina and Brazil |journal=Oryx |volume=42 |issue=4 |pages=554–561 |doi=10.1017/S0030605308000641 |doi-access=free}}</ref>
Some [[Afro-Colombians]] in the Colombian [[Chocó Department]] hunt jaguars for consumption and sale of meat.<ref>{{cite journal |author1=Balaguera-Reina, S. |author2=Gonzalez-Maya, J.F. |year=2008 |name-list-style=amp |title=Occasional jaguar hunting for subsistence in Colombian Chocó |journal=Cat News |issue=48 |page=5 |url=https://www.researchgate.net/publication/233399315 |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021347/https://www.researchgate.net/publication/233399315_Occasional_Jaguar_Hunting_for_Subsistence_in_Colombian_Choco |url-status=live }}</ref>
Between 2008 and 2012, at least 15 jaguars were killed by livestock farmers in central Belize.<ref>{{cite journal |author1=Foster, R.J. |author2=Harmsen, B.J. |author3=Urbina, Y. L. |author4=Wooldridge, R.L. |author5=Doncaster, C.P. |author6=Quigley, H. |author7=Figueroa, O.A. |year=2020 |name-list-style=amp |title=Jaguar (''Panthera onca'') density and tenure in a critical biological corridor |journal=Journal of Mammalogy |volume=101 |issue=6 |pages=1622–1637 |doi=10.1093/jmammal/gyaa134 |pmid=33505226 |pmc=7816682 |doi-access=free}}</ref>
<gallery class="left" widths="187px" heights="167px">
The international trade of jaguar skins boomed between the end of the [[Second World War]] and the early 1970s.<ref name="SkinTrade">{{cite book |last=Broad |first=S. |date=1987 |title=The harvest of and trade in Latin American spotted cats (Felidae) and otters (Lutrinae) |publisher=IUCN Conservation Monitoring Centre |___location=Cambridge |url=https://www.biodiversitylibrary.org/item/119261 |access-date=21 February 2018 |archive-date=13 January 2020 |archive-url=https://web.archive.org/web/20200113095346/https://www.biodiversitylibrary.org/item/119261 |url-status=live }}</ref>
File:Longnoselancetfish.jpg|[[Longnose lancetfish]]. Lancetfish are ambush predators that frequent the mesopelagic. They are among the largest mesopelagic fishes (up to 2 metres).<ref name="Moyle336">[[#Moyle|Moyle and Cech]], p. 336</ref>
Significant declines occurred in the 1960s, as more than 15,000 jaguars were yearly killed for their skins in the [[Brazilian Amazon]] alone; the trade in jaguar skins decreased since 1973 when the [[CITES|Convention on International Trade in Endangered Species]] was enacted.<ref name="r32">{{cite journal |last1=Weber |first1=W. |author2=Rabinowitz, A. |year=1996 |name-list-style=amp |title=A global perspective on large carnivore conservation |journal=Conservation Biology |volume=10 |issue=4 |pages=1046–1054 |doi=10.1046/j.1523-1739.1996.10041046.x |url=http://www.jaguarnetwork.org/pdf/71.pdf |archive-url=https://web.archive.org/web/20120426042409/http://www.jaguarnetwork.org/pdf/71.pdf |archive-date=26 April 2012}}</ref>
File:gigantura chuni.png|The [[telescopefish]] has large, forward-pointing telescoping eyes with large lenses.<ref>{{FishBase species|genus=Gigantura|species=chuni|year=2010|month=October}}</ref>
Interview surveys with 533 people in the northwestern Bolivian Amazon revealed that local people killed jaguars out of fear, in retaliation, and for trade.<ref>{{cite journal |author1=Knox, J. |author2=Negrões, N. |author3=Marchini, S. |author4=Barboza, K. |author5=Guanacoma, G. |author6=Balhau, P. |author7=Tobler, M.W. |author8=Glikman, J.A. |year=2019 |name-list-style=amp |title=Jaguar persecution without "cowflict": insights from protected territories in the Bolivian Amazon |journal=Frontiers in Ecology and Evolution |volume=7 |page=494 |doi=10.3389/fevo.2019.00494 |doi-access=free}}</ref>
File:Daggertooth.PNG|The [[daggertooth]] slashes other mesopelagic fish when it bites them with its dagger-like teeth.<ref>{{FishBase species|genus=Anotopterus|species=pharao|year=2010|month=April}}</ref>
Between August 2016 and August 2019, jaguar skins and body parts were seen for sale in tourist markets in the Peruvian cities of [[Lima]], [[Iquitos]] and [[Pucallpa]].<ref>{{cite journal |author1=Braczkowski, A. |author2=Ruzo, A. |author3=Sanchez, F. |author4=Castagnino, R. |author5=Brown, C. |author6=Guynup, S. |author7=Winter, S. |author8=Gandy, D. |author9=O'Bryan, C. |year=2019 |name-list-style=amp |title=The ayahuasca tourism boom: An undervalued demand driver for jaguar body parts? |journal=Conservation Science and Practice |volume=1 |issue=12 |page=e126 |url=https://www.researchgate.net/publication/336450042 |doi=10.1111/csp2.126 |doi-access=free |access-date=28 February 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021318/https://www.researchgate.net/publication/336450042_The_ayahuasca_tourism_boom_An_undervalued_demand_driver_for_jaguar_body_parts |url-status=live }}</ref>
File:Thobe u0.gif|[[Bigeye tuna]] cruise the epipelagic zone at night and the mesopelagic zone during the day.
[[Human-wildlife conflict]], opportunistic hunting and hunting for trade in domestic markets are key drivers for killing jaguars in Belize and Guatemala.<ref>{{cite journal |author1=Arias, M. |author2=Hinsley, A. |author3=Milner-Gulland, E.J. |year=2020 |name-list-style=amp |title=Characteristics of, and uncertainties about, illegal jaguar trade in Belize and Guatemala |journal=Biological Conservation |volume=250 |page=108765 |doi=10.1016/j.biocon.2020.108765 |s2cid=224967913 |url=https://www.researchgate.net/publication/344172628 |access-date=18 November 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021310/https://www.researchgate.net/publication/344172628_Characteristics_of_and_uncertainties_about_illegal_jaguar_trade_in_Belize_and_Guatemala |url-status=live }}</ref>
File:Lestidiops affinis (1).jpg|A collection of mesopelagic forage fishes trawled from the Gulf of Mexico that includes [[Myctophids]], larval [[anglerfishes]], [[bristlemouth]]s, and a [[barracudina]]
Seizure reports indicate that at least 857 jaguars were involved in trade between 2012 and 2018, including 482 individuals in Bolivia alone; 31 jaguars were seized in [[China]].<ref>{{cite journal |author1=Morcatty, T.Q. |author2=Bausch Macedo, J.C. |author3=Nekaris, K.A.I. |author4=Ni, Q. |author5=Durigan, C.C. |author6=Svensson, M.S. |author7=Nijman, V. |year=2020 |name-list-style=amp |title=Illegal trade in wild cats and its link to Chinese‐led development in Central and South America |journal=Conservation Biology |volume=34 |issue=6 |pages=1525–1535 |doi=10.1111/cobi.13498 |pmid=32484587 |doi-access=free}}</ref>
</gallery>
Between 2014 and early 2019, 760 jaguar fangs were seized that originated in Bolivia and were destined for China. Undercover investigations revealed that the [[smuggling]] of jaguar body parts is run by Chinese residents in Bolivia.<ref>{{cite report |author=Earth League International |year=2020 |title=Unveiling the criminal networks behind jaguar trafficking in Bolivia |publisher=IUCN National Committee of the Netherlands |___location=Amsterdam |url=https://www.iucn.nl/app/uploads/2021/03/iucn_nl_report_jaguar_trafficking_bolivia_media-1.pdf |access-date=26 July 2021 |archive-date=26 July 2021 |archive-url=https://web.archive.org/web/20210726070809/https://www.iucn.nl/app/uploads/2021/03/iucn_nl_report_jaguar_trafficking_bolivia_media-1.pdf |url-status=live }}</ref>
===Bathypelagic fish===
==Conservation==
[[File:Humpback anglerfish.png|thumb|right|The [[humpback anglerfish]] is a bathypelagic ambush predator, which attracts prey with a bioluminescent lure. It can ingest prey larger than itself, which it swallows with an inrush of water when it opens its mouth.<ref name="TeAraBZ" />]]
The jaguar is listed on [[CITES Appendix I]], which means that all international commercial trade in jaguars or their body parts is prohibited. Hunting jaguars is prohibited in Argentina, Brazil, Colombia, French Guiana, Honduras, Nicaragua, Panama, Paraguay, Suriname, the United States, and Venezuela. Hunting jaguars is restricted in Guatemala and Peru.<ref name=iucn /> In Ecuador, hunting jaguars is prohibited, and it is classified as threatened with extinction.<ref>{{cite book |author1=Zapata Ríos, G. |author2=Araguillin, E. |author3=Cevallos, J. |author4=Moreno, F. |author5=Ortega, A. |author6=Rengel, J. |author7=Valarezo, N. |date=2014 |name-list-style=amp |title=Plan de Acción para la Conservación del Jaguar en el Ecuador |trans-title=Action Plan for the Conservation of the Jaguar in Ecuador |publisher=Ministerio del Ambiente y Wildlife Conservation Society Ecuador |___location=Quito |language=es |url=http://www.wild4ever.com/wp-content/uploads/2012/05/Ecuador-National-Jaguar-Plan.pdf |access-date=13 February 2021 |archive-date=23 September 2020 |archive-url=https://web.archive.org/web/20200923073232/http://www.wild4ever.com/wp-content/uploads/2012/05/Ecuador-National-Jaguar-Plan.pdf |url-status=live }}</ref>
[[File:Gonostoma bathyphilum.jpg|thumb|right|Many [[bristlemouth]] species, such as the "spark anglemouth" above,<ref>{{FishBase species | genus = Gonostoma | species = bathyphilum | month = January | year = 2006}}</ref> are also bathypelagic ambush predators that can swallow prey larger than themselves. They are among the most abundant of all vertebrate families.<ref>{{FishBase family | family = Gonostoma | month = August | year = 2009}}</ref>]]
In Guyana, it is protected as an endangered species, and hunting it is illegal.<ref>{{cite book |author=Kerman, I. |date=2010 |title=Exploitation of the jaguar, ''Panthera onca'' and other large forest cats in Suriname |editor=Felix, M.-L. |publisher=WWF Guianas |___location=Paramaribo |url=https://www.a2000greetings.com/downloads/exploitation_of_the_jaguar_and_other_large_forest_cats_in_suriname_irvin_kerman.pdf}}</ref>
[[File:Flabby whalefish.jpg|thumb|right|Young, red [[flabby whalefish]] make nightly vertical migrations into the lower mesopelagic zone to feed on [[copepods]]. When males mature into adults, they develop a massive liver and then their jaws fuse shut. They no longer eat, but continue to metabolise the energy stored in their liver.<ref name="aparticle">{{cite web |url=https://www.nbcnews.com/id/wbna28801960 |title=Scientists solve mystery: 3 fish are all the same |date=22 January 2009 |author=Schmid, Randolph E. |publisher=Associated Press}}</ref>]]
[[File:El-jefe-jaguar-fws1.jpg|thumb|[[El Jefe (jaguar)|El Jefe]], a jaguar in Arizona]]
In 1986, the [[Cockscomb Basin Wildlife Sanctuary]] was established in Belize as the world's first protected area for jaguar conservation.<ref name=Weckel>{{cite journal |author1=Weckel, M. |author2=Giuliano, W. |author3=Silver, S. |year=2006 |name-list-style=amp |title=Cockscomb revisited: jaguar diet in the Cockscomb Basin Wildlife Sanctuary, Belize |journal=Biotropica |volume=38 |issue=5 |pages=687–690 |url=https://semanticscholar.org/paper |doi=10.1111/j.1744-7429.2006.00190.x |s2cid=85151201 |access-date=29 November 2019 |archive-date=27 May 2020 |archive-url=https://web.archive.org/web/20200527070312/https://www.semanticscholar.org/paper |url-status=live }}</ref>
Below the mesopelagic zone it is pitch dark. This is the '''midnight''' or [[bathypelagic zone]], extending from 1000 m to the bottom deep water [[benthic zone]]. If the water is exceptionally deep, the pelagic zone below {{Convert|4000|m|mi|abbr=off}} sometimes is called the '''lower midnight''' or [[abyssopelagic zone]].
===Jaguar Conservation Units===
In 1999, field scientists from 18 jaguar range countries determined the most important areas for long-term jaguar conservation based on the status of jaguar population units, stability of prey base and quality of habitat. These areas, called "Jaguar Conservation Units" (JCUs), are large enough for at least 50 breeding individuals and range in size from {{cvt|566|to|67598|km2}}; 51 JCUs were designated in 36 geographic regions including:<ref name=Sanderson2002 />
* the [[Sierra Madre Occidental]] and [[Sierra de Tamaulipas]] in Mexico
* the [[Selva Maya]] tropical forests extending over Mexico, Belize and Guatemala
* the [[Chocó–Darién moist forests]] from Honduras and Panama to Colombia
* [[Venezuelan Llanos]]
* northern [[Cerrado]] and [[Amazon basin]] in Brazil
* [[Tropical Andes]] in Bolivia and Peru
* [[Misiones Province]] in Argentina
Conditions are somewhat uniform throughout these zones, the darkness is complete, the pressure is crushing, and temperatures, nutrients, and dissolved oxygen levels are all low.<ref name="Moyle585"/>
Optimal routes of travel between core jaguar population units were identified across its range in 2010 to implement [[wildlife corridor]]s that connect JCUs. These corridors represent areas with the shortest distance between jaguar breeding populations, require the least possible energy input of [[dispersal (ecology)|dispersing]] individuals and pose a low mortality risk. They cover an area of {{cvt|2600000|km2}} and range in length from {{cvt|3|to|1102|km}} in Mexico and Central America and from {{cvt|489.14|to|1607|km}} in South America.<ref name=RabinowitzZeller2010>{{Cite journal |author1=Rabinowitz, A. |author2=Zeller, K.A. |year=2010 |name-list-style=amp |title=A range-wide model of landscape connectivity and conservation for the jaguar, ''Panthera onca'' |journal=Biological Conservation |volume=143 |issue=4 |pages=939–945 |doi=10.1016/j.biocon.2010.01.002 |url=https://www.panthera.org/cms/sites/default/files/documents/Rabinowitz_Zeller_2010_Arangewidemodeloflandscapeconnectivityandconservationforjaguar_BioCon.pdf |access-date=14 March 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021304/https://www.panthera.org/cms/sites/default/files/documents/Rabinowitz_Zeller_2010_Arangewidemodeloflandscapeconnectivityandconservationforjaguar_BioCon.pdf |url-status=live }}</ref>
Cooperation with local landowners and municipal, state, or federal agencies is essential to maintain connected populations and prevent fragmentation in both JCUs and corridors.<ref>{{cite book |author1=Zeller, K.A. |author2=Rabinowitz, A. |author3=Salom-Perez, R. |author4=Quigley, H. |year=2013 |name-list-style=amp |chapter=The Jaguar Corridor Initiative: A range-wide conservation strategy |title=Molecular population genetics, evolutionary biology and biological conservation of Neotropical carnivores |editor1=Ruiz-Garcia, M. |editor2=Shostell, J.M. |publisher=Nova Science Publishers |___location=New York |chapter-url=https://conservationcorridor.org/cpb/Zeller_et_al_2013.pdf |pages=629–657 |isbn=978-1-62417-071-3 |access-date=14 March 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129021304/https://conservationcorridor.org/cpb/Zeller_et_al_2013.pdf |url-status=live }}</ref>
Seven of 13 corridors in Mexico are functioning with a width of at least {{cvt|14.25|km}} and a length of no more than {{cvt|320|km}}. The other corridors may hamper passage, as they are narrower and longer.<ref>{{cite journal |author1=Rodríguez-Soto, C. |author2=Monroy-Vilchis, O. |author3=Zarco-González, M.M. |year=2013 |name-list-style=amp |title=Corridors for jaguar (''Panthera onca'') in Mexico: Conservation strategies |journal=Journal for Nature Conservation |volume=21 |issue=6 |pages=438–443 |doi=10.1016/j.jnc.2013.07.002 |url=https://www.academia.edu/35225702 |access-date=18 November 2021 |archive-date=29 November 2021 |archive-url=https://web.archive.org/web/20211129044432/https://www.academia.edu/35225702 |url-status=live }}</ref>
Bathypelagic fish have special [[adaptation]]s to cope with these conditions – they have slow [[metabolism]]s and unspecialized diets, being willing to eat anything that comes along. They prefer to sit and wait for food rather than waste energy searching for it. The behaviour of bathypelagic fish can be contrasted with the behaviour of mesopelagic fish. Mesopelagic are often highly mobile, whereas bathypelagic fish are almost all lie-in-wait predators, normally expending little energy in movement.<ref name="Moyle594">[[#Moyle|Moyle and Cech]], p. 594</ref>
In August 2012, the [[United States Fish and Wildlife Service]] set aside {{cvt|838232|acres|km2|order=flip}} in Arizona and New Mexico for the protection of the jaguar.<ref>{{cite journal |author=Department of the Interior Fish and Wildlife Service |year=2012 |title=Designation of Critical Habitat for Jaguar; Proposed Rule |journal=Federal Register |volume=77 |issue=161 |pages=50214–50242 |url=https://www.fws.gov/southwest/es/arizona/Documents/SpeciesDocs/Jaguar/Jaguar_pCH_FR_8-20-2012.pdf |access-date=13 March 2021 |archive-date=14 February 2020 |archive-url=https://web.archive.org/web/20200214001827/https://www.fws.gov/southwest/es/arizona/Documents/SpeciesDocs/Jaguar/Jaguar_pCH_FR_8-20-2012.pdf |url-status=live }}</ref> The Jaguar Recovery Plan was published in April 2019, in which [[Interstate 10]] is considered to form the northern boundary of the Jaguar Recovery Unit in Arizona and New Mexico.<ref>{{cite journal |author1=Sanderson, E.W. |name-list-style=amp |author2=Fisher, K. |author3=Peters, R. |author4=Beckmann, J.P. |author5=Bird, B. |author6=Bradley, C.M. |author7=Bravo, J.C. |author8=Grigione, M.M. |author9=Hatten, J.R. |author10=González, C.A.L. |author11=Menke, K. |author12=Miller, J.R.B. |author13=Miller, P.S. |author14=Mormorunni, C. |author15=Robinson, M.J. |author16=Thomas, R.E. |author17=Wilcox, S. |year=2021 |title=A systematic review of potential habitat suitability for the jaguar ''Panthera onca'' in central Arizona and New Mexico, USA |journal=Oryx |volume= 56|issue= |pages=116–127 |doi=10.1017/S0030605320000459 |doi-access=free}}</ref>
The dominant bathypelagic fishes are small [[bristlemouth]] and [[anglerfish]]; [[fangtooth]], [[viperfish]], [[daggertooth]], and [[barracudina]] are also common. These fishes are small, many about {{Convert|10|cm|in|abbr=off}} long, and not many longer than {{Convert|25|cm|in|abbr=on}}. They spend most of their time waiting patiently in the water column for prey to appear or to be lured by their phosphors. What little energy is available in the bathypelagic zone filters from above in the form of detritus, faecal material, and the occasional invertebrate or mesopelagic fish.<ref name="Moyle594"/> About 20% of the food that has its origins in the epipelagic zone falls down to the mesopelagic zone,<ref name="TeAraMZ" /> but only about 5% filters down to the bathypelagic zone.<ref name="TeAraBZ">Ryan P [http://www.teara.govt.nz/EarthSeaAndSky/SeaLife/DeepSeaCreatures/3/en "Deep-sea creatures: The bathypelagic zone"] ''Te Ara – the Encyclopedia of New Zealand''. Updated 21 September 2007.</ref>
In Mexico, a national conservation strategy was developed from 2005 on and published in 2016.<ref name=Ceballos2016/> The Mexican jaguar population increased from an estimated 4,000 individuals in 2010 to about 4,800 individuals in 2018. This increase is seen as a positive effect of conservation measures that were implemented in cooperation with governmental and non-governmental institutions and landowners.<ref>{{cite journal |author1=Ceballos, G. |name-list-style=amp |author2=Zarza, H. |author3=González-Maya, J.F. |author4=de la Torre, J.A. |author5=Arias-Alzate, A. |author6=Alcerreca, C. |author7=Barcenas, H.V. |author8=Carreón-Arroyo, G. |author9=Chávez, C. |author10=Cruz, C. |author11=Medellín, D. |year=2021 |title=Beyond words: From jaguar population trends to conservation and public policy in Mexico |journal=PLOS ONE |volume=16 |issue=10 |pages=e0255555 |doi=10.1371/journal.pone.0255555 |pmid=34613994 |pmc=8494370 |bibcode=2021PLoSO..1655555C |doi-access=free}}</ref>
Bathypelagic fish are sedentary, adapted to outputting minimum energy in a habitat with very little food or available energy, not even sunlight, only bioluminescence. Their bodies are [[wiktionary:elongate|elongated]] with weak, watery muscles and [[skeleton|skeletal]] structures. Since so much of the fish is water, they are not compressed by the great pressures at these depths. They often have extensible, hinged [[Jaw#The jaw in fish|jaws]] with recurved teeth. They are slimy, without [[Fish scale|scale]]s. The central nervous system is confined to the lateral line and olfactory systems, the eyes are small and may not function, and [[gill]]s, kidneys and hearts, and [[swimbladder]]s are small or missing.<ref name="TeAraBZ" /><ref name="Moyle587">[[#Moyle|Moyle and Cech]], p. 587</ref>
An evaluation of JCUs from Mexico to Argentina revealed that they overlap with high-quality habitats of about 1,500 mammals to varying degrees. Since co-occurring mammals benefit from the JCU approach, the jaguar has been called an [[umbrella species]].<ref>{{cite journal |author1=Thornton, D. |name-list-style=amp |author2=Zeller, K. |author3=Rondinini, C. |author4=Boitani, L. |author5=Crooks, K. |author6=Burdett, C. |author7=Rabinowitz, A. |author8=Quigley, H. |year=2016 |title=Assessing the umbrella value of a range‐wide conservation network for jaguars (''Panthera onca'') |journal=Ecological Applications |volume=26 |issue=4 |pages=1112–1124 |doi=10.1890/15-0602 |jstor=24818150 |pmid=27509752 |hdl=11573/893793 |url=https://iris.uniroma1.it/retrieve/handle/11573/893793/279408/Thornton_Assessing_2016.pdf |access-date=16 November 2021 |archive-date=16 November 2021 |archive-url=https://web.archive.org/web/20211116083400/https://iris.uniroma1.it/retrieve/handle/11573/893793/279408/Thornton_Assessing_2016.pdf |url-status=live }}</ref>
Central American JCUs overlap with the habitat of 187 of 304 regional endemic amphibian and reptile species, of which 19 amphibians occur only in the jaguar range.<ref>{{cite journal |author1=Figel, J.J. |name-list-style=amp |author2=Castañeda, F. |author3=Calderón, A.P. |author4=Torre, J. |author5=García-Padilla, E. |author6=Noss, R.F. |year=2018 |title=Threatened amphibians sheltered under the big cat's umbrella: conservation of jaguars ''Panthera onca'' (Carnivora: Felidae) and endemic herpetofauna in Central America |journal=Revista de Biología Tropical |volume=66 |issue=4 |pages=1741–1753 |doi=10.15517/rbt.v66i4.32544 |doi-access=free}}</ref>
These are the same features found in fish [[larvae]], which suggests that during their evolution, bathypelagic fish have acquired these features through [[neoteny]]. As with larvae, these features allow the fish to remain suspended in the water with little expenditure of energy.<ref>Marshall (1984) "Progenetic tendencies in deep-sea fishes", pp. 91–101 in Potts GW and Wootton RJ (eds.) (1984) ''Fish reproduction: strategies and tactics'' Fisheries Society of the British Isles.</ref>
===Approaches===
[[File:Obscured jaguar.jpg|thumb|A jaguar in [[Belize]]]]
In setting up protected reserves, efforts generally also have to be focused on the surrounding areas, as jaguars are unlikely to confine themselves to the bounds of a reservation, especially if the population is increasing in size. Human attitudes in the areas surrounding reserves and laws and regulations to prevent poaching are essential to make conservation areas effective.<ref name=gutierrez>{{cite journal |last1=Gutierrez-Gonzalez |first1=C.E. |last2=Gomez-Ramirez |first2=M.A. |last3=Lopez-Gonzalez |first3=C.A. |last4=Doherty |first4=P.F. |year=2015 |name-list-style=amp |title=Are Private Reserves Effective for Jaguar Conservation? |journal=PLOS ONE |volume=10 |issue=9 |page=e0137541 |bibcode=2015PLoSO..1037541G |doi=10.1371/journal.pone.0137541 |pmc=4580466 |pmid=26398115 |doi-access=free}}</ref>
Despite their ferocious appearance, these beasts of the deep are mostly miniature fish with weak muscles, and are too small to represent any threat to humans.
To estimate population sizes within specific areas and to keep track of individual jaguars, [[camera trap]]ping and [[Telemetry#Fishery and wildlife research and management|wildlife tracking telemetry]] are widely used, and feces are sought out with the help of [[detection dog]]s to study jaguar health and diet.<ref name=soisalo>{{cite journal |last1=Soisalo |first1=M.K. |last2=Cavalcanti |first2=S.M.C. |year=2006 |name-list-style=amp |title=Estimating the density of a Jaguar population in the Brazilian Pantanal using camera-traps and capture-recapture sampling in combination with GPS radio-telemetry |journal=Biological Conservation |volume=129 |issue=4 |pages=487–496 |doi=10.1016/j.biocon.2005.11.023 |url=http://www.ekonoiz.com/Eco_Projects/Jaguar_Conservation/estimatingthedensityofjaguarsinthepantanal.pdf}}</ref><ref name=furtado>{{cite journal |last1=Furtado |first1=M.M. |last2=Carrillo-Percastegui |first2=S.E. |last3=Jácomo |first3=A.T.A. |last4=Powell |first4=G. |last5=Silveira |first5=L. |last6=Vynne |first6=C. |last7=Sollmann |first7=R. |year=2008 |name-list-style=amp |title=Studying jaguars in the wild: past experiences and future perspectives |journal=Cat News |issue=Special Issue 4 |pages=41–47 |url=http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.4._SI_4/Furtado_et_al_2008_Jaguar_field_methods_s.pdf |access-date=2 February 2019 |archive-date=2 February 2019 |archive-url=https://web.archive.org/web/20190202212055/http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.4._SI_4/Furtado_et_al_2008_Jaguar_field_methods_s.pdf |url-status=live}}</ref>
The swimbladders of deep sea fish are either absent or scarcely operational, and bathypelagic fish do not normally undertake vertical migrations. Filling bladders at such great pressures incurs huge energy costs. Some deep sea fishes have swimbladders that function while they are young and inhabit the upper epipelagic zone, but they wither or fill with fat when the fish move down to their adult habitat.<ref>{{cite journal|author=Horn MH |year=1970|url=http://biostor.org/reference/4182 |title=The swimbladder as a juvenile organ in stromateoid fishes|journal=Breviora|volume=359|pages=1–9}}</ref>
Current conservation efforts often focus on educating ranch owners and promoting [[ecotourism]].<ref name=WWF>{{cite web |author=Estévez, E. |date=2009 |title=Jaguar Refuge in the Llanos Ecoregion |publisher=[[World Wildlife Fund]] |url=http://wwf.panda.org/es/nuestro_trabajo/latinoamerica/venezuela/index.cfm?uProjectID=VE0854 |url-status=dead |archive-url=https://web.archive.org/web/20141217001905/http://wwf.panda.org/es/nuestro_trabajo/latinoamerica/venezuela/index.cfm?uProjectID=VE0854 |access-date= 1 September 2006 |archive-date=17 December 2014}}</ref> Ecotourism setups are being used to generate public interest in charismatic animals such as the jaguar while at the same time generating revenue that can be used in conservation efforts. A key concern in jaguar ecotourism is the considerable habitat space the species requires. If ecotourism is used to aid in jaguar conservation, some considerations need to be made as to how existing ecosystems will be kept intact, or how new ecosystems will be put into place that are large enough to support a growing jaguar population.<ref name=mossaz>{{cite journal |last1=Mossaz |first1=A. |last2=Buckley |first2=R.C. |last3=Castley |first3=J.G. |year=2015 |name-list-style=amp |title=Ecotourism contributions to conservation of African big cats |journal=Journal for Nature Conservation |volume=28 |pages=112–118 |doi=10.1016/j.jnc.2015.09.009 |hdl=10072/125191 |hdl-access=free}}</ref>
The most important sensory systems are usually the [[inner ear]], which responds to sound, and the [[lateral line]], which responds to changes in water pressure. The [[olfactory]] system also can be important for males who find females by smell.<ref>{{Cite journal | doi = 10.1086/285295|jstor=2462555| title = Location by Olfaction: A Model and Application to the Mating Problem in the Deep-Sea Hatchetfish ''Argyropelecus hemigymnus''| journal = The American Naturalist| volume = 138| issue = 6| pages = 1431| year = 1991| last1 = Jumper | first1 = J. | last2 = Baird | first2 = R. C. |s2cid=84386858 }}</ref>
==In culture and mythology==
Bathypelagic fish are black, or sometimes red, with few [[photophore]]s. When photophores are used, it is usually to entice prey or attract a mate. Because food is so scarce, bathypelagic predators are not selective in their feeding habits, but grab whatever comes close enough. They accomplish this by having a large mouth with sharp teeth for grabbing large prey and overlapping [[gill raker]]s that prevent small prey that have been swallowed from escaping.<ref name="Moyle587"/>
{{multiple image |align=right |direction=vertical
|image1=Jaguar warrior.jpg |caption1=Jaguar warrior in the Aztec culture
|image2=MocheJaguarLarcoMuseum.jpg |caption2=Moche jaguar figurine dating to 300 [[Common Era|CE]], at the [[Larco Museum]] in [[Lima]], Peru}}
{{Further|Jaguars in Mesoamerican cultures}}
In the [[pre-Columbian era|pre-Columbian]] Americas, the jaguar was a symbol of power and strength. In the Andes, a jaguar cult disseminated by the early [[Chavín culture]] became accepted over most of today's Peru by 900 [[Before Christ|BC]].<ref name=b1>{{cite book |title=The Earth and Its Peoples: A Global History |edition=Fifth |editor1=Bulliet, R.W. |editor2=Crossley, P. |editor3=Headrick, D. |editor4=Hirsch, S. |editor5=Johnson, L. |name-list-style=amp |year=2000 |publisher=Houghton Mifflin |volume=A 1: To 1200 |pages=75–76|url={{Google books|aujp0cT_TiEC|page=PA75|plainurl=yes}} |isbn=978-1-4390-8476-2}}</ref> The later [[Moche (culture)|Moche culture]] in northern Peru used the jaguar as a symbol of power in many of their ceramics.<ref name=r41>{{cite book|author=Park, Yumi |year=2012 |title=Mirrors of Clay: Reflections of Ancient Andean Life in Ceramics from the Sam Olden Collection |publisher=University Press of Mississippi|page=49|isbn=9781617037955}}</ref> In the [[Muisca religion]] in [[Altiplano Cundiboyacense]], the jaguar was considered a sacred animal, and people dressed in jaguar skins during religious rituals.<ref name=Ocampo_p231>{{cite book |last=Ocampo López |first=J. |author-link=Javier Ocampo López |year=2007 |title=Grandes culturas indígenas de América – Great indigenous cultures of the Americas |___location=Bogotá, Colombia |publisher=Plaza & Janes Editores Colombia S.A. |language=es |page=231 |isbn=978-958-14-0368-4}}</ref>
The skins were traded with peoples in the nearby [[Orinoquía Region]].<ref>{{cite thesis |author=Kruschek, M.H. |year=2003 |type=PhD thesis |title=The evolution of the Bogotá chiefdom: A household view |publisher=[[University of Pittsburgh]] |___location=Pittsburgh |url=http://d-scholarship.pitt.edu/7735/1/kruschek2003.pdf |access-date=12 January 2017 |archive-date=15 August 2016 |archive-url=https://web.archive.org/web/20160815211943/http://d-scholarship.pitt.edu/7735/1/kruschek2003.pdf |url-status=live }}</ref>
The name of the [[Muisca rulers|Muisca ruler]] [[Nemequene]] was derived from the [[Chibcha language|Chibcha]] words ''nymy'' and ''quyne'', meaning "force of the jaguar".<ref name=nymy>{{cite web |title=''nymy'' |publisher=Muysc cubun Dictionary Online |language=es |url=http://muysca.cubun.org/nymy |access-date=11 January 2017 |archive-date=13 October 2020 |archive-url=https://web.archive.org/web/20201013111835/http://muysca.cubun.org/nymy |url-status=live }}</ref><ref name=quyne>{{cite web |title=''quyne'' |publisher=Muysc cubun Dictionary Online |language=es |url=http://muysca.cubun.org/quyne |access-date=11 January 2017 |archive-date=10 January 2017 |archive-url=https://web.archive.org/web/20170110082526/http://muysca.cubun.org/quyne |url-status=live }}</ref>
It is not easy finding a mate in this zone. Some species depend on [[bioluminescence]]. Others are [[hermaphrodite]]s, which doubles their chances of producing both eggs and sperm when an encounter occurs.<ref name="TeAraBZ" /> The female anglerfish releases [[pheromone]]s to attract tiny males. When a male finds her, he bites onto her and never lets go. When a male of the anglerfish species ''[[Haplophryne mollis]]'' bites into the skin of a female, he release an [[enzyme]] that digests the skin of his mouth and her body, fusing the pair to the point where the two circulatory systems join up. The male then atrophies into nothing more than a pair of [[gonads]]. This extreme [[sexual dimorphism]] ensures that, when the female is ready to spawn, she has a mate immediately available.<ref name="doran">{{Cite journal | doi = 10.1038/256038a0| title = Precocious sexual parasitism in the deep sea ceratioid anglerfish, ''Cryptopsaras couesi'' Gill| journal = Nature| volume = 256| issue = 5512| pages = 38–40| year = 1975| last1 = Pietsch | first1 = T. W. | bibcode = 1975Natur.256...38P| s2cid = 4226567}}</ref>
Sculptures with "[[Olmec were-jaguar]]" motifs were found on the Yucatán Peninsula in [[Veracruz]] and [[Tabasco]]; they show stylized jaguars with half-human faces.<ref>{{cite journal |author1=Metcalf, G. |name-list-style=amp |author2=Flannery, K.V. |year=1967 |title=An Olmec "were-jaguar" from the Yucatan Peninsula |journal=American Antiquity |volume=32 |issue=1 |pages=109–111 |doi=10.2307/278787 |jstor=278787|s2cid=164201137 }}</ref> In the later [[Maya civilization]], the jaguar was believed to facilitate communication between the living and the dead and to protect the royal household. The Maya saw these powerful felines as their companions in the spiritual world, and several Maya rulers bore names that incorporated the Mayan word for jaguar ''b'alam'' in many of the [[Mayan languages]]. ''Balam'' remains a common Maya surname, and it is also the name of [[Chilam Balam]], a legendary author to whom are attributed 17th and 18th-centuries Maya [[miscellanies]] preserving much important knowledge. Remains of jaguar bones were discovered in a burial site in [[Guatemala]], which indicates that [[Mayans]] may have kept jaguars as [[pets]].<ref>{{cite web |last=Dapcevich |first=M. |date=2018 |title=Ancient Mayans Probably Kept Jaguars As Pets And Raised Dogs For Food |publisher=[[IFLScience]] |url=https://www.iflscience.com/plants-and-animals/ancient-mayans-probably-kept-jaguars-as-pets-and-raised-dogs-for-food/ |access-date=26 July 2017 |archive-date=27 March 2020 |archive-url=https://web.archive.org/web/20200327152614/https://www.iflscience.com/plants-and-animals/ancient-mayans-probably-kept-jaguars-as-pets-and-raised-dogs-for-food/ |url-status=live }}</ref>
Many animal forms other than fish live in the bathypelagic zone, such as squid, large whales, octopuses, sponges, [[brachiopod]]s, sea stars, and [[echinoid]]s, but this zone is difficult for fish to live in.
The [[Aztec]] civilization shared this image of the jaguar as the representative of the ruler and as a warrior. The Aztecs formed an elite warrior class known as the [[Jaguar warrior]]. In [[Aztec mythology]], the jaguar was considered to be the [[totem]] animal of the powerful deity [[Tezcatlipoca]].<ref>{{cite journal |author=Saunders, N.J. |year=1994 |title=Predators of culture: Jaguar symbolism and Mesoamerican elites |journal=World Archaeology |volume=26 |issue=1 |pages=104–117 |doi=10.1080/00438243.1994.9980264 |jstor=124867}}</ref><ref>{{cite book |last=Christenson |first=A.J. |year=2007 |title=Popol Vuh: The Sacred Book of the Maya |publisher=[[University of Oklahoma Press]] |___location=Oklahoma |chapter=The first four men |chapter-url={{Google books|hJx47n_sm-YC|page=PA196|plainurl=yes}} |pages=196–199 |isbn=978-0-8061-3839-8}}</ref>
<gallery class="left" widths="187px" heights="167px">
A [[conch shell]] gorget depicting a jaguar was found in a [[burial mound]] in [[Benton County, Missouri]]. The gorget shows evenly-engraved lines and measures {{cvt|104x98|mm}}.<ref name="Daggett"/>
File:Eurypharynx pelecanoides.jpg|The [[Saccopharynx|gulper eel]] uses its mouth like a net by opening its large mouth and swimming at its prey. It has a luminescent organ at the tip of its tail to attract prey.
Rock drawings made by the [[Hopi]], [[Anasazi]] and [[Pueblo]] all over the desert and [[chaparral]] regions of the American Southwest show an explicitly spotted cat, presumably a jaguar, as it is drawn much larger than an [[ocelot]].<ref name=Pavlik /><!--commented out, as this ref is without date + publisher, hence gray literature: Jaguar skins were sold for $18 apiece in the mid 19th century in the vicinity of San Antonio, Texas.<ref>{{Cite report |author=Robinson, M. J. |title=Suitable Habitat for jaguars in New Mexico |publisher=___location= |url=https://www.biologicaldiversity.org/species/mammals/jaguar/pdfs/NM_jaguar_habitat_report.pdf}}</ref>
Image:Chiasmodon niger.jpg|The [[black swallower]], with its distensible stomach, is notable for its ability to swallow whole [[bony fish]]es ten times its mass.<ref name="jordan">{{cite book|title=A Guide to the Study of Fishes|url=https://archive.org/details/aguidetostudyfi02jordgoog|author=Jordan, D.S.|publisher=H. Holt and Company|year=1905}}</ref><ref>{{fishbase species|genus=Chiasmodon|species=niger|year=2009|month=August}}</ref>
-->
Image:Hamol u0.gif|Female ''[[Haplophryne mollis]]'' anglerfish trailing attached males that have atrophied into a pair of [[gonads]], for use when the female is ready to spawn.
File:Anoplogaster cornuta 2.jpg|The widespread [[fangtooth]] has the largest teeth of any fish, proportionate to body size.<ref>{{FishBase species|genus = Anoplogaster|species = cornuta|month = August|year = 2009}}</ref> Despite their ferocious appearance, bathypelagic fish are usually weakly muscled and too small to represent any threat to humans.
File:Messina Straits Chauliodus sloani.jpg|The [[Sloane's viperfish]] can make nightly migrations from bathypelagic depths to near surface waters.<ref>{{FishBase species|genus= Chauliodus|species= sloani|year= 2010|month= April}}</ref>
</gallery>
==Demersal fish==
The jaguar is also used as a symbol in contemporary culture. It is the [[List of national animals|national animal]] of Guyana and is featured in its [[Coat of arms of Guyana|coat of arms]].<ref>{{cite news |author=Khan, A. |date=2021 |title=National symbols: The Coat-Of-Arms |website=Guyana News and Information |url=http://www.guyana.org/Handbook/symbols.html |access-date=10 February 2021 |archive-date=12 May 2012 |archive-url=https://web.archive.org/web/20120512212328/http://guyana.org/Handbook/symbols.html |url-status=live }}</ref>
[[File:Giant grenadier.jpg|thumb|right|[[Giant grenadier]], an [[wiktionary:elongate|elongate]] benthic fish with large eyes and well-developed [[lateral line]]s]]
The [[flag of the Department of Amazonas]] features a black jaguar silhouette leaping towards a hunter.<ref>{{cite web |author=Gutterman, D. |date=2008 |title=Amazonas Department (Colombia) |publisher=Flag of the World |url=https://www.fotw.info/flags/co-ama.html |access-date=2 April 2010 |archive-date=9 June 2020 |archive-url=https://web.archive.org/web/20200609071213/https://www.fotw.info/flags/co-ama.html |url-status=live }}</ref>
{{See also|Demersal fish}}
The crest of the [[Argentine Rugby Union]] features a jaguar.<ref>{{cite news |last=Davies |first=S. |date=2007 |title=Puma power: Argentinian rugby |publisher=BBC News |url=http://news.bbc.co.uk/sport2/hi/rugby_union/5389512.stm |access-date=8 October 2007 |archive-date=19 April 2020 |archive-url=https://web.archive.org/web/20200419112328/http://news.bbc.co.uk/sport2/hi/rugby_union/5389512.stm |url-status=live }}</ref>
[[Demersal fish]] live on or near the bottom of the sea.<ref name="seafloor">Walrond C [http://www.teara.govt.nz/en/coastal-fish/5 Carl . "Coastal fish – Fish of the open sea floor"] Te Ara – the Encyclopedia of New Zealand. Updated 2 March 2009</ref> Demersal fish are found by the [[seafloor]] in coastal areas on the [[continental shelf]], and in the open ocean they are found along the outer [[continental margin]] on the continental slope and the continental rise. They are not generally found at [[abyssopelagic]] or [[hadopelagic]] depths or on the [[abyssal plain]]. They occupy a range of seafloors consisting of mud, sand, gravel, or rocks.<ref name="seafloor" />
== References ==
{{Reflist|30em|refs=
<ref name=fossil>{{Cite web |url=http://www.fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&taxon_no=36760 |title=Fossilworks: Megaptera |website=[[Fossilworks]] |access-date=17 December 2021 }}</ref>
In deep waters, the fishes of the demersal zone are active and relatively abundant, compared to fishes of the [[bathypelagic zone]].<ref name="Moyle594"/>
<ref name=MSW3>{{MSW3 Cetacea|id=14300027}}</ref>
[[Rattail]]s and [[Ophidiidae|brotula]]s are common, and other well-established families are [[eel]]s, [[eelpout]]s, [[hagfish]]es, [[greeneye]]s, [[Pancake batfish|batfish]]es, and [[lumpfish]]es.<ref name="Moyle587"/>
<ref name=CITES>{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/appendices.php|access-date=2022-01-14|website=cites.org}}</ref>
The bodies of deep water [[benthic]] fishes are muscular with well developed organs. In this way they are closer to mesopelagic fishes than bathopelagic fishes. In other ways, they are more variable. [[Photophore]]s are usually absent, eyes and [[swimbladder]]s range from absent to well developed. They vary in size, with larger species greater than one metre not uncommon.
}}
Deep sea benthic fish are usually long and narrow. Many are eels or shaped like eels. This may be because long bodies have long [[lateral line]]s. Lateral lines detect low-frequency sounds, and some benthic fishes appear to have muscles that drum such sounds to attract mates.<ref name="Haedrich1996"/> Smell is also important, as indicated by the rapidity with which benthic fish find traps baited with [[bait fish]].
The main diet of deep sea benthic fish is invertebrates of the deep sea [[benthos]] and [[carrion]]. Smell, touch, and lateral line sensitivities seem to be the main sensory devices for locating these.<ref name="Moyle588">[[#Moyle|Moyle and Cech]], p. 588</ref>
Deep sea benthic fish can be divided into strictly benthic fish and benthopelagic fish. Usually, strictly benthic fish are negatively buoyant, while benthopelagic fish are neutrally buoyant. Strictly benthic fish stay in constant contact with the bottom. They either lie in wait as [[ambush predator]]s or move actively over the bottom in search for food.<ref name="Moyle588"/>
[[File:Orange roughy.png|thumb|right|[[Orange roughy]]]]
[[File:Toothfish.jpg|thumb|right|[[Patagonian toothfish]]]]
===Benthopelagic fish===
{{See also|Benthopelagic fish}}
Benthopelagic fish inhabit the water just above the bottom, feeding on [[benthos]] and benthopelagic [[zooplankton]].<ref>{{cite journal|author=Mauchline J |author2= Gordon JDM |year=1986|title=Foraging strategies of deep-sea fish|journal=Mar. Ecol. Prog. Ser.|volume=27|pages=227–238|doi=10.3354/meps027227|bibcode=1986MEPS...27..227M|doi-access=free}}</ref> Most dermersal fish are benthopelagic.<ref name="seafloor" />
They can be divided into flabby or robust body types. Flabby benthopelagic fishes are like bathopelagic fishes, they have a reduced body mass, and low metabolic rates, expending minimal energy as they lie and wait to [[Ambush predator|ambush]] prey.<ref>{{Cite journal | doi = 10.1111/j.1095-8649.1996.tb06067.x| title = Energetic and life-history patterns of deep-sea benthic, benthopelagic and seamount-associated fish| journal = Journal of Fish Biology| volume = 49| pages = 54–74| year = 1996| last1 = Koslow | first1 = J. A.}}</ref> An example of a flabby fish is the cusk-eel ''[[Acanthonus armatus]]'',<ref name="fishbase">{{fishbase species |genus=Acanthonus |species=armatus |year=2009 |month=August}}</ref> a predator with a huge head and a body that is 90% water. This fish has the largest ears ([[otolith]]s) and the smallest brain in relation to its body size of all known vertebrates.<ref name="Fine">{{Cite journal | doi = 10.1098/rspb.1987.0018|pmid=2884671|jstor=36061|url=https://www.researchgate.net/publication/20270929| title = ''Acanthonus armatus'', a Deep-Sea Teleost Fish with a Minute Brain and Large Ears| journal = Proceedings of the Royal Society B: Biological Sciences| volume = 230| issue = 1259| pages = 257–65| year = 1987| last1 = Fine | first1 = M. L.| last2 = Horn | first2 = M. H.| last3 = Cox | first3 = B.|bibcode=1987RSPSB.230..257F|s2cid=19183523}}</ref>
Robust benthopelagic fish are muscular swimmers that actively cruise the bottom searching for prey. They may live around features, such as [[seamount]]s, which have strong currents.<ref name="Fine"/> Examples are the [[orange roughy]] and [[Patagonian toothfish]]. Because these fish were once abundant, and because their robust bodies are good to eat, these fish have been harvested commercially.<ref>{{FishBase species|genus=Hoplostethus|species=atlanticus|year=2009|month=August}}</ref><ref>{{FishBase species|genus=Dissostichus|species=eleginoides|year=2009|month=August}}</ref>
===Benthic fish===
{{See also|Benthic fish}}
Benthic fish are not pelagic fish, but they are discussed here briefly, by way of completeness and contrast.
Some fishes do not fit into the above classification. For example, the family of nearly blind [[spiderfish]]es, common and widely distributed, feed on benthopelagic zooplankton. Yet they are strictly benthic fish, since they stay in contact with the bottom. Their fins have long rays they use to "stand" on the bottom while they face the current and grab zooplankton as it passes by.<ref>{{cite journal|author= Sulak KJ |title=The systematics and biology of ''Bathypterois'' (Pisces, Chlorophthalmidae) with a revised classification of benthic myctophiform fishes|journal= Galathea Rep.|volume= 14|url=https://www.researchgate.net/publication/237671743}}</ref>
The deepest-living fish known, the strictly benthic ''[[Abyssobrotula galatheae]]'', eel-like and blind, feeds on benthic invertebrates.<ref>{{cite journal|author=Nielsen JG |year=1977|title=The deepest living fish ''Abyssobrotula galatheae'': a new genus and species of oviparous ophidioids (Pisces, Brotulidae)|journal=Galathea Report|volume=14|pages=41–48|url=http://bionames.org/references/7b623b8692cc9de5ca095628affbc735}}</ref><ref>{{fishbase species |genus=Abyssobrotula |species=galatheae |month=August |year=2009}}</ref>
<gallery class="left" widths="200" heights="175">
File:Pacific hagfish Myxine.jpg|[[Pacific hagfish]] resting on bottom. Hagfish coat themselves and any dead fish they find with noxious slime, making them inedible to other species.
File:Bathypterois grallator.jpg|The tripodfish (''[[Bathypterois grallator]]''), a species of spiderfish, uses its fin extensions to "stand" on the bottom.<ref>{{FishBase species|genus=Bathypterois|species=grallator|year=2009|month=August}}</ref>
File:Taeniura meyeni reef.jpg|The [[blotched fantail ray]] feeds on bottom-dwelling fish, bivalves, crabs, and shrimps.<ref>{{fishbase species|genus=Taeniura|species=meyeni|month=August|year=2009}}</ref>
</gallery>
[[File:Oceanic basin.svg|thumb|Cross-section of an ocean basin, note significant [[vertical exaggeration]]]]
At great depths, food scarcity and extreme pressure works to limit the survivability of fish. The deepest point of the ocean is about {{Convert|11000|m|mi|abbr=off}}. Bathypelagic fishes are not normally found below {{Convert|3000|m|mi|abbr=off}}. The greatest depth recorded for a benthic fish is {{Convert|8370|m|mi|abbr=on}}.<ref name="nielsen">{{cite journal |title=The deepest living fish ''Abyssobrotula galatheae'': a new genus and species of oviparous ophidioids (Pisces, Brotulidae) |author=Nielsen, J.G. |journal=Galathea Report |year=1977 |volume=14 |pages=41–48}}</ref> It may be that extreme pressures interfere with essential enzyme functions.<ref name="TeAraBZ" />
Benthic fishes are more diverse and are likely to be found on the [[continental slope]], where there is habitat diversity and often, food supplies. Approximately 40% of the ocean floor consists of [[abyssal plain]]s, but these flat, featureless regions are covered with [[Pelagic sediments|sediment]] and largely devoid of benthic life ([[benthos]]). Deep sea benthic fishes are more likely to associate with canyons or rock outcroppings among the plains, where invertebrate communities are established. Undersea mountains ([[seamount]]s) can intercept deep sea currents and cause productive upwellings that support benthic fish. Undersea mountain ranges may separate underwater regions into different ecosystems.<ref name="Moyle591" />
{{clear}}
==Pelagic fisheries==
===Forage fish===
{{marine wild fish taxa}}
{{See also|Forage fish}}
Small pelagic fish are usually [[forage fish]] that are hunted by larger pelagic fish and other predators. Forage fish [[Filter feeder|filter feed]] on [[plankton]] and are usually less than {{Convert|10|cm|in|abbr=off}} long. They often stay together in [[Shoaling and schooling|schools]] and may [[Fish migration|migrate]] large distances between spawning grounds and feeding grounds. They are found particularly in [[upwelling]] regions around the northeast Atlantic, off the coast of Japan, and off the west coasts of Africa and the Americas. Forage fish are generally short-lived, and their [[Fish stocks|stocks]] fluctuate markedly over the years.<ref name="Checkley">Checkley D, Alheit J and Oozeki Y (2009) ''Climate Change and Small Pelagic Fish'', Cambridge University Press. {{ISBN|0-521-88482-9}}.</ref>
[[Herring]] are found in the [[North Sea]] and the [[North Atlantic]] at depths to {{Convert|200|m|ft|abbr=off}}. Important herring fisheries have existed in these areas for centuries. Herring of different sizes and growth rates belong to different populations, each of which have their own migration routes. When spawning, a female produces from 20,000 to 50,000 eggs. After spawning, the herrings are depleted of fat, and migrate back to feeding grounds rich in plankton.<ref name="pfa">[http://www.pfa-frozenfish.com/pfa2/fish1.html Pelagic species] {{webarchive|url=https://web.archive.org/web/20120211174604/http://www.pfa-frozenfish.com/pfa2/fish1.html |date=2012-02-11 }} Pelagic Freezer-trawler Association. Retrieved 22 July 2009.</ref> Around Iceland, three separate populations of herring were fished traditionally. These stocks collapsed in the late 1960s, although two have since recovered. After the collapse, Iceland turned to [[capelin]], which now account for about half of Iceland's total catch.<ref>[http://www.fisheries.is/main-species/pelagic-fishes/ Pelagic fishes] Icelandic fisheries. Retrieved 24 July 2009.</ref>
[[Blue whiting]] are found in the open ocean and above the [[continental slope]] at depths between 100 and 1000 meters . They follow vertical migrations of the [[zooplankton]] they feed on to the bottom during daytime and to the surface at night time.<ref name="pfa"/><ref>[http://www.imr.no/temasider/fisk/kolmule/kolmule/en Blue whiting] [[Institute of Marine Research]]. Retrieved 23 July 2009.</ref>
Traditional fisheries for [[Anchovy|anchovies]] and [[sardine]]s also have operated in the Pacific, the Mediterranean, and the southeast Atlantic.<ref name="Bone443">[[#Bone|Bone and Moore]], p. 443</ref> The world annual catch of forage fish in recent years has been approximately 22 million tonnes, or one quarter of the world's total catch.
<gallery class="left" widths="187" heights="167">
File:Pacific sardine002.jpg|These [[Shoaling and schooling|schooling]] [[Pacific sardine]]s are [[forage fish]].
File:Herringramkils.jpg|[[Herring]]s [[Forage fish#Hunting copepods|ram feeding]] on [[copepod]]s
File:Mallotus villosus.gif|[[Capelin]]
File:Anchovy closeup.jpg|[[Anchovy|Anchovies]]
File:Enrin u0.png|Peruvian anchoveta
</gallery>
===Predator fish===
{{See also|Predator fish}}
Medium size pelagic fishes include [[trevally]], [[barracuda]], [[flying fish]], [[bonito]], [[mahi mahi]], and coastal mackerel.<ref name="Lal8"/> Many of these fish hunt forage fish, but are in turn, hunted by yet larger pelagic fish. Nearly all fish are predator fish to some measure, and apart from the top predators, the distinction between predator fish and prey or forage fish, is somewhat artificial.<ref>[[FAO]]: LAPE project [http://www.fao.org/fishery/topic/4290/en Forage species] Rome. Updated 28 November 2008.</ref>
Around Europe there are three populations of coastal [[mackerel]]. One population migrates to the North Sea, another stays in the waters of the [[Irish Sea]], and the third population migrates southward along the west coast of Scotland and Ireland. The cruise speed of the mackerel is an impressive 10 kilometres per hour.<ref name="pfa"/><ref name="Mackerel">[http://www.imr.no/temasider/fisk/makrell/makrell/en Mackerel] [[Institute of Marine Research]]. Retrieved 23 July 2009.</ref>
Many large pelagic fish are oceanic nomadic species that undertake long offshore migrations. They feed on small pelagic forage fish, as well as medium-sized pelagic fish. At times, they follow their schooling prey, and many species form schools themselves.
Examples of larger pelagic fish are [[tuna]], [[billfish]], [[king mackerel]], sharks, and large [[Ray-finned fish|ray]]s.
Tuna in particular are of major importance to commercial fisheries. Although tuna migrate across oceans, trying to find them there is not the usual approach. Tuna tend to congregate in areas where food is abundant, along the boundaries of currents, around islands, near seamounts, and in some areas of upwelling along continental slopes. Tuna are captured by several methods: [[Seiner|purse seine vessels]] enclose an entire surface school with special nets, [[Fishing vessel#Line vessels|pole and line vessels]] that use poles baited with other smaller pelagic fish as [[baitfish]], and rafts called [[fish aggregating device]]s are set up, because tuna, as well as some other pelagic fish, tend to congregate under floating objects.<ref name="Lal8"/>
Other large pelagic fish are premier [[game fish]], particularly [[marlin]] and [[swordfish]].
<gallery class="left" widths="187" heights="167">
File:Yellowfin tuna nurp.jpg|[[Yellowfin tuna]] are being fished as a replacement for the now largely depleted [[Southern bluefin tuna]].
File:Brama brama.jpg|[[Atlantic pomfret]]
File:Xiphias gladius1.jpg|[[Swordfish]]
File:Sccav u0.gif|alt=King mackerels cruise on long migrations at 10 kilometres per hour|[[King mackerel]]s cruise on long migrations at 10 kilometres per hour.<ref name="pfa"/><ref name="Mackerel"/>
</gallery>
[[File:Ocean surface currents.jpg|thumb|right|450px|Major ocean surface currents]]
[[File:upwelling image1.jpg|thumb|right|300px|Areas of upwelling in red]]
[[File:La Nina and Pacific Decadal Anomalies - April 2008.png|thumb|right|300px|Pacific decadal anomalies – April 2008]]
===Productivity===
[[Upwelling]] occurs both along coastlines and in midocean when a collision of deep [[ocean current]]s brings cold water that is rich in nutrients to the surface. These upwellings support blooms of phytoplankton, which in turn, produce zooplankton and support many of the world's main fisheries. If the upwelling fails, then fisheries in the area fail.<ref name="Moyle574/5">[[#Moyle|Moyle and Cech]], pp. 574–575</ref>
In the 1960s the [[Peruvian anchoveta]] fishery was the world's largest fishery. The anchoveta population was greatly reduced during the 1972 [[El Niño]] event, when warm water drifted over the cold [[Humboldt Current]], as part of a 50-year cycle, lowering the depth of the [[thermocline]]. The upwelling stopped and [[phytoplankton]] production plummeted, as did the anchoveta population, and millions of [[seabird]]s, dependent on the anchoveta, died.<ref>{{Cite journal | doi = 10.1126/science.1075880| pmid = 12522241| title = From Anchovies to Sardines and Back: Multidecadal Change in the Pacific Ocean| journal = Science| volume = 299| issue = 5604| pages = 217–21| year = 2003| last1 = Chavez | first1 = F. P.| last2 = Ryan| first2 = John| last3 = Lluch-Cota| first3 = Salvador E.| last4 = Ñiquen c.| first4 = Miguel| bibcode = 2003Sci...299..217C| s2cid = 37990897}}</ref> Since the mid-1980s, the upwelling has resumed, and the Peruvian anchoveta catch levels have returned to the 1960s levels.
Off Japan, the collision of the [[Oyashio Current]] with the [[Kuroshio Current]] produces nutrient-rich upwellings. Cyclic changes in these currents resulted in a decline in the [[sardine]] ''sardinops melanosticta'' populations. Fisheries catches fell from 5 million tonnes in 1988 to 280 thousand tonnes in 1998. As a further consequence, [[Pacific bluefin tuna]] stopped moving into the region to feed.<ref>{{Cite journal | doi = 10.1111/j.1365-2419.1996.tb00110.x| title = Decadal variation in the trans-Pacific migration of northern bluefin tuna (''Thunnus thynnus'') coherent with climate-induced change in prey abundance| journal = Fisheries Oceanography| volume = 5| issue = 2| pages = 114–119| year = 1996| last1 = Polovina | first1 = J. J. | url = https://zenodo.org/record/1230607}}</ref><ref>[[FAO]]: Species Fact Sheets: [http://www.fao.org/fishery/species/2893/en ''Sardinops melanostictus'' (Schlegel, 1846)] Rome. Retrieved 18 August 2009.</ref>
Ocean currents can shape how fish are distributed, both concentrating and dispersing them. Adjacent ocean currents can define distinct, if shifting, boundaries. These boundaries can even be visible, but usually their presence is marked by rapid changes in salinity, temperature, and turbidity.<ref name="Moyle574/5" />
For example, in the Asian northern Pacific, [[albacore]] are confined between two current systems. The northern boundary is determined by the cold [[North Pacific Current]] and the southern boundary is determined by the [[North Equatorial Current]]. To complicate things, their distribution is further modified within the area defined by the two current systems by another current, the [[Kuroshio Current]], whose flows fluctuate seasonally.<ref>{{cite book|author=Nakamura, Hiroshi |title=Tuna distribution and migration|url=https://books.google.com/books?id=SygRAAAAYAAJ|year=1969|publisher=Fishing News|isbn=9780852380024}}</ref>
Epipelagic fish often [[spawn (biology)|spawn]] in an area where the eggs and larvae drift downstream into suitable feeding areas, and eventually, drift into adult feeding areas.<ref name="Moyle574/5" />
Islands and [[Fishing bank|banks]] can interact with currents and upwellings in a manner that results in areas of high ocean productivity. Large eddies can form downcurrent or downwind from islands, concentrating plankton.<ref>{{cite journal|author=Blackburn M |year=1965|title=Oceanography and the ecology of tunas|journal=Oceanography and Marine Biology: An Annual Review|volume=3|pages=299–322|url=http://fsf.fra.affrc.go.jp/bulletin/kenpoupdf/kenpou2-257.pdf}}</ref> Banks and reefs can intercept deep currents that upwell.<ref name="Moyle574/5" />
* [[Scombrid]]s
===Highly migratory species===
{{See also|Highly migratory species}}
[[File:Isurus-oxyrinchus.jpg|thumb|left|[[Shortfin mako shark]] make long seasonal migrations. They appear to follow temperature gradients, and have been recorded travelling more than 4,500 km in one year.<ref>{{Cite journal | doi = 10.1071/MF9920045| title = Tagging studies on the Shortfin Mako Shark (''Isurus oxyrinchus'') in the Western North Atlantic| journal = Marine and Freshwater Research| volume = 43| pages = 45| year = 1992| last1 = Casey | first1 = J. G. | last2 = Kohler | first2 = N. E. }}</ref>]]
Epipelagic fish generally move long distances between feeding and spawning areas, or as a response to changes in the ocean. Large ocean predators, such as salmon and tuna, can migrate thousands of kilometres, crossing oceans.<ref name="Moyle578">[[#Moyle|Moyle and Cech]], p. 578</ref>
In a 2001 study, the movements of [[Atlantic bluefin tuna]] from an area off North Carolina were studied with the help of special popup tags. When attached to a tuna, these tags monitored the movements of the tuna for about a year, then detached and floated to the surface where they transmitted their information to a satellite. The study found that the tuna had four different migration patterns. One group confined itself to the western Atlantic for a year. Another group also stayed mainly in the western Atlantic, but migrated to the Gulf of Mexico for spawning. A third group moved across the Atlantic Ocean and back again. The fourth group crossed to the eastern Atlantic and then moved into the Mediterranean Sea for spawning. The study indicates that, while there is some differentiation by spawning areas, there is essentially only one population of Atlantic bluefin tuna, intermixing groups that between them, use all of the north Atlantic Ocean, the Gulf of Mexico, and the Mediterranean Sea.<ref>{{Cite journal | doi = 10.1126/science.1061197|pmid=11509729|url=http://www.tunaresearch.org/reprints/migratory2001.pdf| title = Migratory Movements, Depth Preferences, and Thermal Biology of Atlantic Bluefin Tuna| journal = Science| volume = 293| issue = 5533| pages = 1310–4| year = 2001| last1 = Block | first1 = B. A.|last2=Dewar|first2=H|last3=Blackwell|first3=S. B.|last4=Williams|first4=T. D.|last5=Prince|first5=E. D.|last6=Farwell|first6=C. J.|last7=Boustany|first7=A|last8=Teo|first8=S. L.|last9=Seitz|first9=A|last10=Walli|first10=A|last11=Fudge|first11=D|bibcode=2001Sci...293.1310B|s2cid=32126319}}</ref>
The term [[highly migratory species]] (HMS) is a legal term that has its origins in Article 64 of the [[United Nations Convention on the Law of the Sea]] (UNCLOS).<ref>[[United Nations]] Convention on the [[Law of the Sea]]: [https://www.un.org/Depts/los/convention_agreements/texts/unclos/closindx.htm Text]</ref>
The highly migratory species include: [[tuna]] and tuna-like species ([[Albacore tuna|albacore]], Atlantic bluefin, [[bigeye tuna]], [[Skipjack tuna|skipjack]], [[Yellowfin tuna|yellowfin]], [[Blackfin tuna|blackfin]], [[Euthynnus alletteratus|little tunny]], [[Pacific bluefin tuna|Pacific bluefin]], [[Southern bluefin tuna|southern bluefin]] and [[Auxis rochei|bullet]]), [[Bramidae|pomfret]], [[marlin]], [[sailfish]], [[swordfish]], [[saury]] and oceangoing [[shark]]s, as well as mammals such as [[dolphin]]s, and other [[cetacean]]s.
Essentially, highly migratory species coincide with the larger of the "large pelagic fish", discussed in the previous section, if cetaceans are added and some commercially unimportant fish, such as the [[Molidae|sunfish]], are excluded. These are high [[trophic level]] species that undertake migrations of significant, but variable distances across oceans for feeding, often on forage fish, or reproduction, and also have wide geographic distributions. Thus, these species are found both inside the {{convert|200|nmi|km|adj=on}} [[exclusive economic zone]]s and in the [[high seas]] outside these zones. They are [[pelagic]] species, which means they mostly live in the open ocean and do not live near the sea floor, although they may spend part of their life cycle in [[nearshore waters]].<ref>[[Pacific Fishery Management Council]]: [http://www.pcouncil.org/hms/hmsback.html Background: Highly Migratory Species] {{Webarchive|url=https://web.archive.org/web/20090712150423/http://www.pcouncil.org/hms/hmsback.html |date=2009-07-12 }}</ref>
===Capture production===
According to the [[Food and Agriculture Organization]] (FAO), the [[Fishing by country|world harvest]] in 2005 consisted of 93.2 million [[tonne]]s captured by [[commercial fishing]] in [[wild fisheries]].<ref>[http://www.fao.org/fishery/ Fisheries and Aquaculture]. FAO. Retrieved on 2015-05-01.</ref> Of this total, about 45% were pelagic fish. The following table shows the world capture production in [[tonne]]s.<ref>[[FAO]] (2007) [http://www.fao.org/docrep/009/A0699e/A0699e00.htm ''State of the World Fisheries and Aquaculture 2006.''] Fisheries and Aquaculture Department. {{ISBN|978-92-5-105568-7}}</ref>
{| class="sortable wikitable"
|-
! colspan="9" style="text-align:center; width:640px;"| '''Capture production by groups of species in tonnes'''
|-
! style="width:45px;"| Type
! style="width:100px;"| Group
! 1999
! 2000
! 2001
! 2002
! 2003
! 2004
! 2005
|-
| Small pelagic fish
| [[Herring]]s, [[sardine]]s, [[Anchovy|anchovies]]
| 22 671 427
| 24 919 239
| 20 640 734
| 22 289 332
| 18 840 389
| 23 047 541
| 22 404 769
|-
| Large pelagic fish
| [[Tuna]]s, [[bonito]]s, [[billfish]]es
| 5 943 593
| 5 816 647
| 5 782 841
| 6 138 999
| 6 197 087
| 6 160 868
| 6 243 122
|-
| Other pelagic fish
|
| 10 712 994
| 10 654 041
| 12 332 170
| 11 772 320
| 11 525 390
| 11 181 871
| 11 179 641
|-
| [[Cartilaginous fish]]
| [[Shark]]s, [[Batoidea|rays]], [[chimaera]]s
| 858 007
| 870 455
| 845 854
| 845 820
| 880 785
| 819 012
| 771 105
|}
==Threatened species==
In 2009, the [[International Union for Conservation of Nature]] (IUCN) produced the first [[IUCN Red List|red list]] for threatened oceanic sharks and rays. They claim that approximately one third of open ocean sharks and rays are under [[Threatened species|threat of extinction]].<ref>[http://www.iucn.org/?3362/Third-of-open-ocean-sharks-threatened-with-extinction Third of open ocean sharks threatened with extinction] [[IUCN]]. 25 June 2009.</ref> There are 64 species of oceanic sharks and rays on the list, including [[Hammerhead shark|hammerhead]]s, giant [[Mobula|devil ray]]s, and [[porbeagle]].<ref name="guardian">[https://www.theguardian.com/science/2009/jun/25/sharks-extinction-iucn-red-list Fishing puts a third of all oceanic shark species at risk of extinction] ''[[guardian.co.uk]]'', 26 June 2009.</ref>
Oceanic sharks are [[incidental catch|captured incidentally]] by swordfish and tuna [[high seas]] fisheries. In the past there were few markets for sharks, which were regarded as worthless [[bycatch]]. Now sharks are being increasingly targeted to supply emerging Asian markets, particularly for [[Shark finning|shark fins]], which are used in [[shark fin soup]].<ref name="guardian"/>
The northwest Atlantic Ocean shark populations are estimated to have declined by 50% since the early 1970s. Oceanic sharks are vulnerable because they do not produce many young, and the young can take decades to mature.<ref name="guardian"/>
<gallery class="left" widths="200" heights="175">
File:Sphyrnalewini.jpg|The [[scalloped hammerhead]] is classified as endangered.
File:Oceanic Whitetip Shark.png|alt=The oceanic whitetip shark has declined by 99% in the Gulf of Mexico|The [[oceanic whitetip shark]] has declined by 99% in the [[Gulf of Mexico]].<ref name="guardian"/>
File:Atlantic mobula lisbon.jpg|The [[devil fish]], a large ray, is threatened.
File:Lamna nasus.jpg|The [[Porbeagle|porbeagle shark]] is threatened.
</gallery>
In parts of the world the [[scalloped hammerhead]] shark has declined by 99% since the late 1970s. Its status on the red list is that it is globally endangered, meaning it is near extinction.<ref name="guardian"/>
==See also==
{{div col|colwidth=22em}}
* [[Deep sea]]
* [[Deep sea fish]]
* [[Demersal fish]]
* [[Freshwater fish]]
* [[Nekton]]
* [[Ocean Tracking Network]]
* [[Oily fish]]
* [[Pacific Ocean Shelf Tracking Project]]
* [[Tagging of Pacific Predators]]
{{div col end}}
==References==
'''Notes'''
{{Reflist|30em}}
'''Bibliography'''
* {{cite book|ref=Bone|author1=Bone, Quentin |author2=Moore, Richard |title=Biology of Fishes|url=https://books.google.com/books?id=sLoqT_xWaqoC|date=2008|publisher=Garland Science|isbn=978-0-203-88522-2}}
*{{cite book|ref=Moyle|author1=Moyle, PB |author2=Cech, JJ |name-list-style=amp |year=2004|title=Fishes, An Introduction to Ichthyology|edition=5th|publisher=Benjamin Cummings|isbn=978-0-13-100847-2}}
==Further reading==
* Collette, BB (2010) [https://books.google.com/books?hl=en&lr=&id=_4xZ_QoYNSwC&oi=fnd&pg=PA21 "Reproduction and development in epipelagic fishes"] In: Kathleen S Cole, ''Reproduction and Sexuality in Marine Fishes: Patterns and Processes'', pp. 21–64, University of California Press. {{ISBN|978-0-520-26433-5}}.
* Freon, Pierre (1998) ''Dynamics of Pelagic Fish Distribution and Behaviour: Effects on Fisheries and Stock Assessment'', Wiley-Blackwell. {{ISBN|978-0-85238-241-7}}.
* {{cite journal | last1 = Johnsen | first1 = S | year = 2003 | title = Lifting the Cloak of Invisibility: The Effects of Changing Optical Conditions on Pelagic Crypsis1 | journal = Integrative and Comparative Biology | volume = 43 | issue = 4| pages = 580–590 | doi=10.1093/icb/43.4.580 | pmid=21680466| doi-access = free }}
* {{cite journal | last1 = Makris | first1 = N | last2 = Ratilal | first2 = P | last3 = Jagannathan | first3 = S | last4 = Gong | first4 = Z | last5 = Andrews | first5 = M | last6 = Bertsatos | first6 = I | last7 = Godo | first7 = OR | last8 = Nero | first8 = RW | last9 = Jech | first9 = JM | year = 2009 | title = Critical Population Density Triggers Rapid Formation of Vast Oceanic Fish Shoals | url = http://www.sciencemag.org/cgi/content/abstract/323/5922/1734 | journal = Science | volume = 323 | issue = 5922| pages = 1734–1737 | doi=10.1126/science.1169441 | pmid=19325116| bibcode = 2009Sci...323.1734M | s2cid = 6478019 }}
* Pepperell J (2011) [https://books.google.com/books?id=ZaSuYgEACAAJ ''Fishes of the Open Ocean: A Natural History and Illustrated Guide''] University of New South Wales Press, {{ISBN|978-1-74223-267-6}}.
* Salvanesa AGV and Kristoffersen JB [https://web.archive.org/web/20090521003502/http://www.bio.uib.no/inc/pdffiles/Pub/1325.pdf "Mesopelagic Fishes"] In: ''Encyclopedia of Ocean Sciences'', pp. 1711–1717. {{doi|10.1006/rwos.2001.0012}}
* [https://phys.org/news/2009-03-scientists-ids-genesis-animal-behavior.html Scientists IDs genesis of animal behavior patterns] ''[[PhysOrg.com]]'', 26 March 2009.
* [https://phys.org/news/2006-02-fish-mit-sensor.html One fish, two fish: New MIT sensor improves fish counts] ''[[PhysOrg.com]]'', 2 February 2006.
==External links==
{{Commons category|Deep sea fish}}
* [https://www.ted.com/talks/edith_widder_glowing_life_in_an_underwater_world Glowing life in an underwater world] TED video from [[Edith Widder]]
* [https://web.archive.org/web/20140625125029/http://marinebio.org/oceans/open-ocean.asp The Open Ocean] ''MarineBio.org''. MarineBio.org. Updated 28 August 2011. TED video from [[Edith Widder]]
* [https://web.archive.org/web/20140625125029/http://marinebio.org/oceans/open-ocean.asp The Open Ocean] ''MarineBio.org''. MarineBio.org. Updated 28 August 2011.
* [https://www.pelagic-ac.org/ Pelagic Advisory Council] of the [[European Commission]]
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