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Rise of the Planet of the Apes
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Rise of the Planet of the Apes
Theatrical release poster
Directed by
Rupert Wyatt
Produced by
Peter Chernin
Dylan Clark
Rick Jaffa
Amanda Silver
Written by
Rick Jaffa
Amanda Silver
Based on
Premise suggested by Planet of the Apes
by Pierre Boulle
Starring
James Franco
Freida Pinto
John Lithgow
Brian Cox
Tom Felton
David Oyelowo
Andy Serkis
Music by
Patrick Doyle
Cinematography
Andrew Lesnie
Edited by
Conrad Buff
Mark Goldblatt
Production
company
Chernin Entertainment
Dune Entertainment
Big Screen Productions
Ingenious Film Partners
Distributed by
20th Century Fox
Release date
August 5, 2011 (2011-08-05)
Running time
110 minutes[1]
Country
United States[2][3][4]
Language
English
Budget
$93 million[5]
Box office
$481.8 million[1]
Rise of the Planet of the Apes is a 2011 American science fiction film directed by Rupert Wyatt and starring James Franco, Freida Pinto, John Lithgow, Brian Cox, Tom Felton, David Oyelowo, and Andy Serkis. Written by Rick Jaffa and Amanda Silver, it is 20th Century Fox's reboot of the Planet of the Apes series, intended to act as an origin story for a new series of films.[6] Its premise is similar to the fourth film in the original series, Conquest of the Planet of the Apes (1972), but it is not a direct remake of that film.
Rise of the Planet of the Apes was released on August 5, 2011, to critical and commercial success. The film was nominated for an Academy Award for Best Visual Effects. It was also nominated for five Saturn Awards including Best Director for Wyatt and Best Writing for Jaffa and Silver, winning Best Science Fiction Film, Best Supporting Actor for Serkis and Best Special Effects. Serkis's performance as Caesar was widely acclaimed, earning him many nominations from associations which do not usually recognize performance capture as traditional acting. A sequel to the film, Dawn of the Planet of the Apes, was released on July 11, 2014, and a third film, War for the Planet of the Apes, was released on July 14, 2017.
Contents
[hide]
1 Plot
2 Cast
2.1 Humans
2.2 Apes
3 Production
3.1 Development and writing
3.2 Filming
3.3 Visual effects
3.4 Music
4 Reception
4.1 Critical response
4.2 Box office
4.3 Home media
4.4 Awards
5 Sequels
6 See also
7 References
8 External links
Plot[edit]
Will Rodman, a scientist at the San Francisco biotech company Gen-Sys, is testing the viral-based drug ALZ-112 on chimpanzees to find a cure for Alzheimer's disease. ALZ-112 is given to a chimp named Bright Eyes, greatly increasing her intelligence. But then, during Will's presentation for the drug, Bright Eyes is forced from her cage, goes on a rampage, and is shot to death. Will's boss Steven Jacobs terminates the project and has the chimps slaughtered. However, Will's assistant Robert Franklin discovers that the reason for Bright Eyes' rampage was that she had recently given birth to an infant chimp. Will reluctantly agrees to take in the chimp, who is named Caesar. Will learns that Caesar has inherited his mother's intelligence and decides to raise him. Three years later, Will introduces Caesar to the redwood forest at Muir Woods National Monument. Meanwhile, Will treats his dementia-suffering father Charles with ALZ-112, which seems to restore his cognitive ability.
Five years later, when Caesar reaches adolescence and sees a dog on a leash like his own, he questions his identity and learns of his origins from Will. Meanwhile, Charles's condition returns as he becomes resistant to ALZ-112. Caesar injures an aggressive neighbor, Douglas Hunsiker, while defending a confused Charles from him. As a result, he is placed in a primate shelter where he is tormented by a chimp named Rocket and the chief guard, Dodge Landon, but also where he befriends an orangutan named Maurice who came from a circus. Caesar learns how to unlock his cage, gaining free access to the common area. With the assistance of a gorilla named Buck, he confronts the sanctuary's alpha, Rocket and claims that position. Meanwhile, Jacobs clears development of a more powerful, gaseous version of the drug – ALZ-113 – when Will tells him it can also improve intelligence. Will takes the drug home to try to save his father, but Charles declines further treatment and dies overnight.
After attempting to test the drug on a scarred bonobo test subject named Koba, Franklin becomes exposed to ALZ-113 and becomes ill. Attempting to warn Will at his home, he sneezes blood onto Hunsiker and is later discovered dead. Will attempts to reclaim Caesar, but the chimp refuses to go home with him so he could look after the other apes. Instead, he escapes from the facility and returns to Will's house, where he takes canisters of the ALZ-113. Upon returning, Caesar releases the gas and allows it to enhance the intelligence of the other apes. When Dodge attempts to get him back into his cage, Caesar speaks for the first time, yelling "No!" and gets in a fight which inadvertently leads to Dodge's death. The apes flee the facility, release the remaining chimps from Gen-Sys, and free more apes from the San Francisco Zoo on the way.
A battle ensues as the ape army fights their way past a police blockade on the Golden Gate Bridge to escape into the redwood forest. Buck sacrifices himself to save Caesar by jumping into the helicopter in which Jacobs is riding. The helicopter crashes onto the bridge, trapping Jacobs in the wreckage. While Caesar ignores Jacobs' pledges for help, Koba seizes his chance for revenge and sends Jacobs falling to his death in the bay below.
As the apes find their way into the forest, Will catches up to them and warns Caesar that the humans will hunt them down, and begs him to return home. In response, Caesar hugs him and says that, "Caesar is home." Will, realizing that this is indeed their last goodbye, respects Caesar's wishes. The apes embrace their new lifestyle in the forest.
In a mid-credits scene, an infected Hunsiker leaves his house for work as an airline pilot, arriving at San Francisco International Airport for his flight to Paris. His nose begins to drip blood onto the floor. A graphic traces the spread of the humanity-ravaging virus around the globe via international flight routes.
Cast[edit]
Humans[edit]
James Franco as Dr. William "Will" Rodman, a biologist who is trying to discover a cure for his father's Alzheimer's disease by testing ALZ-112 on chimps. He is a father figure to Caesar. James Franco was cast after talks with Tobey Maguire broke down.[7][8]
Freida Pinto as Dr. Caroline Aranha, a primatologist who begins a relationship with Will and grows attached to Caesar.
John Lithgow as Charles Rodman, Will's Alzheimer's-afflicted father and a former music teacher who improves after Will gives him the ALZ-112 and forms a grandfatherly bond with Caesar.
Brian Cox as John Landon, manager of the San Bruno Primate Shelter where Caesar is confined for a time. His full name is a reference to one of the astronauts in the original Planet of the Apes.
Tom Felton as Dodge Landon, John's son and an animal caretaker at the shelter, who abuses the apes for sport. His first and last name are references to two of the astronauts in the original Planet of the Apes.
David Oyelowo as Steven Jacobs, Will's arrogant boss. His last name is a reference to Arthur P. Jacobs, the producer of the original Planet of the Apes series.[citation needed]
Tyler Labine as Robert Franklin, a chimp handler at Gen-Sys and one of Will's friends. His surname is a reference to Franklin Schaffner, the director of the original film.
Jamie Harris as Rodney, a caretaker and a nightwatchman who is much kinder to the apes at the sanctuary and is regularly mocked by Dodge for this.
David Hewlett as Hunsiker, Will's hot-headed neighbor.
Chelah Horsdal as Irena, a nurse who is looking after Charles.
Apes[edit]
Andy Serkis as Caesar, a common chimpanzee whose intelligence is increased from being exposed in the womb to ALZ-112 when the drug is administered to his pregnant mother, and who is raised by Will for eight years.
Karin Konoval as Maurice, a wise and benevolent Bornean orangutan who was retired from the circus and knows sign language; he becomes Caesar's closest ally. Konoval also cameos as the court clerk whom Will briefly argues with about his appeal. His name is a reference to Maurice Evans who portrays Dr. Zaius in the original Planet of the Apes.[9]
Terry Notary as Rocket, the dominant common chimpanzee at the ape sanctuary, until Caesar overthrows him. Notary also plays Bright Eyes, Caesar's mother, who was captured in Africa. "Bright Eyes" was the name given by chimpanzee scientist Zira to Taylor, the human astronaut in the Planet of the Apes movie.
Richard Ridings as Buck, a Western lowland gorilla who pledges his allegiance to Caesar after he is freed by him.
Devyn Dalton as Cornelia, a female common chimpanzee in the ape sanctuary.
Jay Caputo as Alpha, the dominant male common chimpanzee of Bright Eyes' troop and Caesar's father.
Christopher Gordon as Koba, a scarred and aggressive bonobo who has spent most of his life in laboratories and holds a grudge against humans.
Production[edit]
Development and writing[edit]
In 2006, screenwriter-producer Rick Jaffa was searching for a script idea. As Jaffa searched a newspaper articles clipping, one about pet chimpanzees that become troublesome to their owners and heartbroken for not adapting well to the human environment intrigued him. As Jaffa eventually realized it fit the Planet of the Apes series, he called his wife and screenwriting partner Amanda Silver to express his ideas of such a chimpanzee eventually starting the ape revolution, and then the couple started developing the character of Caesar. Jaffa indicated that "it's a reinvention" and if he had to pick between calling it a prequel or a reboot he would say it is a reboot: "It's a different story of who Caesar is, and how he came to be. So it's really kind of hard to put a label on it. We are hopefully rebooting it." He went on to say that "we tried really hard to create a story that would stand on its own and yet also pay homage and honor the movies that came before us."[10] Jaffa and Silver then wrote a script and sold it to Fox, producers of the Apes franchise. The script added other elements which the couple had researched, such as genetic engineering.[11] Several tributes to specific scenes, characters, and cast and crew from the previous Apes film series were added in the script. In particular, Caesar's treatment at the primate sanctuary parallels Taylor's treatment as a captive in the original film.[12][9]
In a segment of a video blog post, director Rupert Wyatt commented on the originality of the plot: "This is part of the mythology and it should be seen as that. It's not a continuation of the other films; it's an original story. It does satisfy the people who enjoy those films. The point of this film is to achieve that and to bring that fan base into this film exactly like Batman Begins."[6] In a 2009 interview, Wyatt said, "We've incorporated elements from Conquest of the Planet of the Apes, in terms of how the apes begin to revolt, but this is primarily a prequel to the 1968 film...Caesar is a revolutionary figure who will be talked about by his fellow apes for centuries...This is just the first step in the evolution of the apes, and there's a lot more stories to tell after this. I imagine the next film will be about the all-out war between the apes and humans."[13] Mark Bomback did an uncredited rewrite of the script.[14]
Filming[edit]
Filming began in July 2010 in Vancouver, British Columbia.[15] Filming also happened in San Francisco, California (the primary setting of the film),[15] and around Oahu, Hawaii, which doubled for the African jungle as the schedule and budget did not allow for location shooting in Africa.[16]
Visual effects[edit]
As the apes in Rise were meant to be actual apes, the producers decided not to use actors in ape suits. After considering real apes, instead Weta Digital created the apes digitally in almost every case through performance capture.[17] Advances in the technology allowed the use of performance capture in an exterior environment, affording the film-makers the freedom to shoot much of the film on location with other actors, as opposed to the confines of a soundstage.[18][19] The main breakthrough was a camera that enabled viewing the motion capture dots in daylight, employed mostly for the Golden Gate Bridge battle. A maximum of six actors could have their movements captured, with larger ape crowds using fully digital animals animated using Weta's move library. The Golden Gate Bridge set used both a physical set which was extended digitally, and a fully computer-generated model of the bridge that also included the ocean and nearby hills.[20]
After shooting the actors playing humans interacting with others wearing the motion capture suits, a clean plate was shot with actors for extra reference. Actor-stuntman Terry Notary guided the actors on realistic ape movement, while Weta studied the chimps in the Wellington Zoo for reference. The digital apes also received detailed models with skeletons, muscles and nerve tissue layers for accurate animation. Cast models of apes' heads and limbs helped the texture department replicate skin details such as wrinkles and pores. Given the difference between human and chimpanzee facial muscles, the animators tweaked the performance through a new facial muscle system adding dynamics, ballistics, and secondary motion. As the silent performance required expressive eyes, a new eye model was made to depict both greater accuracy in muscle movement in and around the eyes, and also tears, pupil dilation, and light refraction.[17][21] While Andy Serkis was the primary performer for Caesar, as the effects team considered that at times "Andy overcame the character," other motion capture team actors were also used, especially Devyn Dalton, whose height matched that of a chimpanzee. Along with that, they used Notary to play Caesar in stunt-filled scenes such as the Golden Gate Bridge scene.[22]
Music[edit]
The score for the film was written by Patrick Doyle and performed by the Hollywood Studio Symphony conducted by James Shearman.[23] The main concern was to have the music help progress the plot in the scenes without dialogue, for instance, conveying the emotions of Caesar's relationships with Will and Charles. To turn the score into a "driving force that keeps audiences paying attention," Doyle employed an African-American chorus and focused on percussion and "low and deep" orchestra sounds. Doyle collaborated closely with the sound department to make the music complement the sound effects, including writing a recurring theme based on their recording of a chimpanzee.[24]
Reception[edit]
Critical response[edit]
On review aggregator Rotten Tomatoes, the film has an approval rating of 81% based on 258 reviews, with an average rating of 7.1/10. The site's critical consensus reads, "Led by Rupert Wyatt's stylish direction, some impressive special effects, and a mesmerizing performance by Andy Serkis, Rise of the Planet of the Apes breathes unlikely new life into a long-running franchise."[25] Another review aggregator, Metacritic, gave the film a score of 68 out of 100, based on 39 critics, indicating "generally favorable reviews".[26]
Roger Ebert gave the film 3 out of 4 stars and praised the role of Caesar and Andy Serkis by stating it was a "wonderfully executed character" and "one never knows exactly where the human ends and the effects begin, but Serkis and/or Caesar gives the best performance in the movie."[27] Giving the film 5 out of 5 stars, Joe Neumaier of Daily News labelled Rise of the Planet of the Apes as the summer's best popcorn flick.[28] Nick Pinkerton of The Village Voice wrote, "Caesar's prison conversion to charismatic pan-ape revolutionist is near-silent filmmaking, with simple and precise images illustrating Caesar's General-like divining of personalities and his organization of a group from chaos to order."[29] Roger Moore of Orlando Sentinel wrote, "Audacious, violent and disquieting, "Rise of the Planet of the Apes" is a summer sequel that's better than it has any right to be." He gave the film 3.5 out of 4 stars.[30] Manohla Dargis of The New York Times praised the film by saying, "Precisely the kind of summer diversion that the studios have such a hard time making now. It's good, canny-dumb fun." She also gave it 3.5 out of 4 stars.[31]
Peter Travers of Rolling Stone noted that the film has mixed "twists lifted from 1972's Conquest of the Planet of the Apes and 1999's Deep Blue Sea".[32]
Box office[edit]
Rise of the Planet of the Apes made its debut in the United States and Canada on roughly 5,400 screens within 3,648 theaters.[33] It grossed $19,534,699 on opening day and $54,806,191 in its entire opening weekend, making it #1 for that weekend as well as the fourth-highest-grossing August opening ever.[34] The film held on to the #1 spot in its second weekend, dropping 49.2%, and grossing $27,832,307.[35] Rise of the Planet of the Apes crossed the $150 million mark in the United States and Canada on its 26th day of release. Entertainment Weekly said that this was quite an accomplishment for the film since the month of August is a difficult time for films to make money.[36]
The film ended its run at the box office on December 15, 2011, with a gross of $176,760,185 in the U.S. and Canada as well as $305,040,864 internationally, for a total of $481,801,049 worldwide.[1]
Home media[edit]
Rise of the Planet of the Apes was released on Blu-ray Disc, DVD, and Digital Copy on December 13, 2011.[37]
Awards[edit]
Award
Category
Recipient
Result
84th Academy Awards Best Visual Effects Joe Letteri, Dan Lemmon, R. Christopher White and Daniel Barrett Nominated
Alliance of Women Film Journalists[38] Best Supporting Actor Andy Serkis Nominated
Annie Awards[39] Character Animation in a Live Action Production Eric Reynolds Won
Broadcast Film Critics Association[40] Best Supporting Actor Andy Serkis Nominated
Best Visual Effects Won
Best Action Film Nominated
Empire Awards[41] Best Film Nominated
Best Sci-Fi/Fantasy Nominated
Best Director Rupert Wyatt Nominated
Best Actor Andy Serkis Nominated
Genesis Awards Best Feature Film Rick Jaffa and Amanda Silver Won
Houston Film Critics Society[42] Best Supporting Actor Andy Serkis Nominated
Technical Achievement Won
IGN Best of 2011[43] Best Movie Nominated
Best Sci-Fi Movie Won
Best Movie Actor Andy Serkis (also for The Adventures of Tintin) Nominated
Best Movie Director Rupert Wyatt Nominated
IGN Summer Movie Awards[44]
Best Summer Movie Nominated
Funniest Line "Why cookie Rocket?" Nominated
Best All-Out Brawl Apes vs. Humans on the Golden Gate Bridge Nominated
Favorite Kill Helicopter Pushed Over the Golden Gate Bridge Nominated
Coolest Creature Caesar Won
Favorite Hero Caesar – Andy Serkis Won
Las Vegas Film Critics Society[45] Best Visual Effects Won
London Film Critics Circle Technical Achievement Joe Letteri Nominated
Phoenix Film Critics Society Best Visual Effects Nominated
San Diego Film Critics Society[46] Best Supporting Actor Andy Serkis Nominated
Satellite Awards[47] Best Supporting Actor – Motion Picture Andy Serkis Nominated
Best Visual Effects Jeff Capogreco, Joe Letteri, R. Christopher White Nominated
Saturn Awards[48] Best Science Fiction Film Won
Best Supporting Actor Andy Serkis Won
Best Director Rupert Wyatt Nominated
Best Writing Rick Jaffa and Amanda Silver Nominated
Best Special Effects Dan Lemmon, Joe Letteri, R. Christopher White, and Daniel Barrett Won
Visual Effects Society[49] Outstanding Visual Effects in a Visual Effects-Driven Feature Motion Picture Dan Lemmon, Joe Letteri, Cyndi Ochs, Kurt Williams Won
Outstanding Animated Character in a Live Action Feature Motion Picture Caesar – Daniel Barrett, Florian Fernandez, Matthew Muntean, Eric Reynolds Won
Outstanding Virtual Cinematography in a Live Action Feature Motion Picture Thelvin Cabezas, Mike Perry, R. Christopher White, Erik Winquist Nominated
Outstanding Compositing in a Feature Motion Picture Jean-Luc Azzis, Quentin Hema, Simon Jung, Christoph Salzmann Nominated
Washington D.C. Area Film Critics Association[50] Best Supporting Actor Andy Serkis Nominated
Sequels[edit]
Main articles: Dawn of the Planet of the Apes and War for the Planet of the Apes
Regarding the story setting up possible sequels, director Rupert Wyatt commented: "I think we're ending with certain questions, which is quite exciting. To me, I can think of all sorts of sequels to this film, but this is just the beginning."[51] Screenwriter and producer Rick Jaffa also stated that Rise of the Planet of the Apes would feature several clues as to future sequels: "I hope that we're building a platform for future films. We're trying to plant a lot of the seeds for a lot of the things you are talking about in terms of the different apes and so forth."[6]
On May 31, 2012, 20th Century Fox announced that the sequel would be named Dawn of the Planet of the Apes.[52] Reports said that Wyatt was leaving the sequel due to his concern that a May 2014 release date would not give him enough time to properly make the film;[53] he was replaced by Cloverfield director Matt Reeves.[54] Jaffa and Silver returned as producers and to pen the screenplay, with rewrites from Scott Z. Burns[55] and Mark Bomback.[14]
Taking place ten years after Rise, Dawn follows Caesar's growing nation of evolved apes threatened by human survivors that put both sides in the brink of war.[citation needed] Andy Serkis, Terry Notary and Karin Konoval reprised their roles as Caesar, Rocket and Maurice.[56] James Franco returned as Will Rodman in a "cameo via video".[57] Judy Greer and Toby Kebbell replace Devyn Dalton and Christopher Gordon as Cornelia and Koba, respectively.[citation needed] Filming of Dawn of the Planet of the Apes started on North Vancouver Island in April 2013.[58] The film was released July 11, 2014.
On January 6, 2014, 20th Century Fox announced a third installment with Reeves returning to direct and co-write along with Bomback, with a planned July 2016 release.[59][60] In January 2015, Fox delayed the release to July 14, 2017.[61][62] On May 14, 2015, the title was given as War of the Planet of the Apes,[63] later re-titled to War for the Planet of the Apes.
See also[edit]
Uplift (science fiction)
References[edit]
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External links[edit]
Official website
Rise of the Planet of the Apes on IMDb
[show]
v
t
e
Planet of the Apes
Films
Original series
Planet of the Apes (1968)
Beneath the Planet of the Apes (1970)
Escape from the Planet of the Apes (1971)
Conquest of the Planet of the Apes (1972)
Battle for the Planet of the Apes (1973)
Remake
Planet of the Apes (2001)
Reboot series
Rise of the Planet of the Apes (2011)
Dawn of the Planet of the Apes (2014)
War for the Planet of the Apes (2017)
Television
Planet of the Apes (1974)
Return to the Planet of the Apes (1975)
Behind the Planet of the Apes (1998)
Video games
Planet of the Apes (2001)
Revenge of the Apes (2003)
Planet of the Apes: Last Frontier (2017)
Other
La Planète des singes
Characters
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Sockeye salmon
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For other uses, see Sockeye (disambiguation).
Sockeye salmon
Male ocean-phase sockeye
Conservation status

Least Concern (IUCN 3.1)[1]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Salmoniformes
Family: Salmonidae
Genus: Oncorhynchus
Species: O. nerka
Binomial name
Oncorhynchus nerka
(Walbaum, 1792)
Sockeye salmon (Oncorhynchus nerka), also called red salmon, kokanee salmon, or blueback salmon, is an anadromous species of salmon found in the Northern Pacific Ocean and rivers discharging into it. This species is a Pacific salmon that is primarily red in hue during spawning. They can grow up to 84 cm (2 ft 9 in) in length and weigh 2.3 to 7 kg (5.1–15.4 lb). Juveniles remain in freshwater until they are ready to migrate to the ocean, over distances of up to 1,600 km (990 mi). Their diet consists primarily of zooplankton. Sockeye salmon are semelparous, dying after they spawn. Some populations, referred to as kokanee, do not migrate to the ocean and live their entire lives in freshwater.
Contents
[hide]
1 Classification and name origin
2 Description
3 Range and habitat
3.1 Landlocked populations
4 Diet
5 Life cycle
6 Reproduction
6.1 Sexual selection and natural selection
6.2 Energy cost
7 Competition
8 Fisheries and consumption
9 Conservation status
9.1 United States
9.2 Canada
10 References
11 Technical reports
12 External links
Classification and name origin[edit]
Sockeye salmon is the third-most common Pacific salmon species, after pink and chum salmon.[2] Oncorhynchus comes from the Greek ὄγκος (onkos) meaning "barb", and ῥύγχος (rhynchos) meaning "snout". Nerka is the Russian name for the anadromous form.[3] The name "sockeye" is an anglicization of suk-kegh (sθə́qəy̓), its name in Halkomelem, the language of the indigenous people along the lower reaches of the Fraser River (one of British Columbia's many native Coast Salish languages). Suk-kegh means "red fish".[4][5]
Description[edit]
The sockeye salmon is sometimes called red or blueback salmon, due to its color.[5] Sockeye are blue tinged with silver in color while living in the ocean.[3] When they return to spawning grounds, their bodies become red and their heads turn green. Sockeye can be anywhere from 60 to 84 cm (2 ft 0 in–2 ft 9 in) in length and weigh from 2.3 to 7 kg (5.1–15.4 lb).[5] Two distinguishing features are their long, serrated gill rakers that range from 30 to 40 in number, and their lack of a spot on their tail or back.[3]
Range and habitat[edit]
Sockeye salmon range as far south as the Columbia River in the eastern Pacific (although individuals have been spotted as far south as the 10 Mile River on the Mendocino Coast of California) and in northern Hokkaidō Island in Japan in the western Pacific. They range as far north as the Bathurst Inlet in the Canadian Arctic in the east and the Anadyr River in Siberia in the west. The farthest inland sockeye salmon travel is to Redfish Lake, Idaho, over 900 miles (1,400 km) from the ocean and 6,500 feet (2,000 m) in elevation.[6]
Landlocked populations[edit]
Male spawning-phase sockeye
Completely landlocked populations of the same species also are known. Some sockeye live and reproduce in lakes and are commonly called kokanee, which is red-fish name in the Sinixt Interior Salish language and silver trout in the Okanagan language.[7][8] They are much smaller than the anadromous variety and are rarely over 35 cm (14 in) long. In the Okanagan Lake and many others, there are two kinds of kokanee populations – one spawns in streams and the other near lake shores. Landlocked populations occur in the Yukon Territory and British Columbia in Canada, as well as, in Alaska, Washington, Oregon, California, New York, Utah, Idaho, Montana, Nevada, Colorado, New Mexico, and Wyoming in the United States. Nantahala Lake is the only place in North Carolina where kokanee salmon are found.[3] The fish, which is native to western North America, was stocked in Nantahala Lake in the mid-1960s by the NC Wildlife Resources Commission in an attempt to establish the species as a forage fish for other predator fishes in the lake. This stock has remained and become a favorite target for anglers.[9]
In Japan, a landlocked variety termed black kokanee, or "kunimasu" in Japanese, was deemed to be extinct after 1940, when a hydroelectric project made its native lake in northern Akita Prefecture more acidic. The species seems to have been saved by transferring eggs to Saiko Lake, 500 kilometers to the south, however.[10] This fish has been treated as a subspecies of sockeye Oncorhynchus nerka kawamurae, or even an independent species Oncorhynchus kawamurae.[11]
Diet[edit]
Sockeye salmon use patterns of limnetic feeding behavior, which encompasses vertical movement, schooling, diel feeding chronology, and zooplankton prey selectivity. They can change their position in the water column, timing and length of feeding, school formation, and choice of prey to minimize the likelihood of predation. This also ensures they still get the minimum amount of food necessary to survive. All of these behaviors contribute to the survivability, and therefore fitness of the salmon. Depending on location and threat of predation, the levels of aggressive feeding behavior can vary.[12][13]
Sockeye salmon, unlike other species of Pacific salmon, feed extensively on zooplankton during both freshwater and saltwater life stages.[14] They also tend to feed on small aquatic organisms such as shrimp. Insects are part of their diets at the juvenile stage.[3]
Life cycle[edit]
See also: Salmon run
Sockeye salmon exhibit many different life histories with the majority being anadromous where the juvenile salmon migrate from freshwater lakes and streams to the ocean before returning as adults to their natal freshwater to spawn.[15] Similar to most Pacific salmon, sockeye salmon are semelparous, meaning they die after spawning once. Some sockeye, called kokanee, do not migrate to the ocean and live their entire lives in freshwater lakes.[16] The majority of sockeye spawn in rivers near lakes and juveniles will spend one to two years in the lake before migrating to the ocean, although some populations will migrate to saltwater in their first year.[17] Adult sockeye will spend two to three years in the ocean before returning to freshwater. Females will spawn in 3-5 redds over a period of a several days. The eggs usually hatch within six to nine weeks and the fry typically rear in lakes before migrating to the ocean.[18]
Reproduction[edit]
Spawning sockeye salmon
Males partake in competitive and sneaking tactics, formation of hierarchies, and non-hierarchical groupings around females who are ready to mate.[19] Reproductive success varies more in males than females. The greater variability in male reproduction is associated with the greater average size and exaggerated shape of males. Reproductive success in females is determined by the number of eggs she lays, her body size, and the survival of the eggs, which is due in part to the quality of the nest environment.[20] Male spatial distribution depends on shifts in reproductive opportunities, physical traits of breeding sites, as well as the operational sex ratio (OSR) of the environment.[19]
Non-dominant males adopt a subordinate behavior, acting as a satellite to mated pairs. During spawning, a subordinate male will move quickly into the redd and release their sperm. Nearby dominant males from other redds will also do this.[21] Male social status is positively correlated to length and dorsal hump size. Larger females tend to spawn in shallower water, which is preferred over deeper water.[20]
A male (near) and female (far) sockeye salmon spawning in the Adams River of British Columbia, Canada.
There is a dramatic sexual dimorphism at maturity.[20] Males go through numerous morphological changes at maturation including, an increase in body depth, hump height, and snout length. Snout size also increases in females, but hump height and adipose fin length do not increase. This could mean that longer snout sizes are sexually selected, but hump height and adipose fin length are not. Females develop large gonads that are about 25% of the body mass.[22]
Females are responsible for parental care. They select, prepare, and defend a nest site until they die or are displaced. Males do not participate in parental care at all, and they move between females after egg deposition.[23]
Sexual selection and natural selection[edit]
Sexual selection favors large males and females.[24] Males choose females based on their readiness to spawn and their size in order to maximize their breeding opportunities. Larger bodies allow females to reproduce larger and more numerous eggs, better nest choice and ability to defend it, and the ability to bury eggs deeper and provide more protection.[25] Females vary their breeding rate depending on the size of the courting male, mating more quickly with larger males. This increases the likelihood that larger males will displace attending, smaller males. Male sockeye salmon social status and greater reproductivity are directly associated with larger body size and more extreme body shapes; Larger bodies provide males with advantages when it comes to intrasexual competition and being selected for by females during reproduction.[25] Males preferentially spawn with females who are red, which is the usual color of females. Even small changes in wavelength, saturation, and brightness can affect preference. Both sexually naïve males and experienced males showed the same preferences, which illustrates that preference for a red hue may be innate.[23]
Some traits that lead to reproductive success, such as body size and sexual dimorphism can affect one’s survival. This leads to opposing pressures of natural selection and sexual selection. Larger males are favored, unless the risk of predation is very high. Sockeye salmon that die prematurely from predation are typically the larger ones in a population.[26] This shows natural selection against large bodies. Populations with higher levels of predation tend to evolve smaller body size.[27] Without the threat of predation, salmon that breed early in the season live longer than those that breed late in the season.[24]
Other ecological factors like stranding effect select for smaller body size in sockeye salmon when present in a habitat. Stranding is when salmon swim into dry land or shallow water during their migration for spawning and die from suffocation.[27] In fact, studies show that the Sockeye salmon with the largest bodies are most susceptible to stranding mortality.[28]
Energy cost[edit]
Reproduction is marked by depletion in energy stores. Fat, protein, and somatic energy stores decrease from the final moments in marine migration through freshwater entry, spawning, and death.[22] Sockeye salmon do not feed during reproduction.[21] Feeding ends once they enter into freshwater, which can be several months before spawning.[22] Embryos are maintained with only endogenous food supplies for about 3–8 months.[29] Reproduction in the sockeye salmon has to be accomplished with the energy stores brought to the spawning grounds. How the salmon use their energy during migration and spawning affects how successful they will be reproductively; energy used for migration cannot also be used for courtship. If they waste too much energy, they might not be able to spawn. Males must also make the decision whether to invest energy in fighting for a female or for longevity on the spawning grounds.[21]
Competition[edit]
Aggressive behavior displayed by dominant males is predominantly directed towards intruding dominant males. Sometimes sockeye salmon males behave aggressively towards subordinate males. These encounters are short, with the intruding male leaving after one or two aggressive interactions.[21] Spawning females direct their aggression primarily towards intruding females or other spawning females that are close by. However, they may also direct aggression towards intruding or subordinate males.[21] Aggressive interactions between females only last one or two charges and/or chases. The intruder retreats and the spawning female settles back in her redd.[21] These acts of aggression are important in terms of reproductive success, because they determine the quality of the nest site the female obtains and access to males.[21]
Competition for food or space while the salmon are in their lake residence period can exist. This happens when there is a more populous class of young sockeye or when there are multiple classes present. It can also happen when resources are in short supply. Interspecific competition can also occur and can lead to interactive segregation, which is when species emphasize their differences in diet and habitat to avoid competition. Interspecific competition can affect the growth rates of the salmon if their access to resources is limited.[30]
Fisheries and consumption[edit]
See also: Salmon (food)
The total registered fisheries harvest of the sockeye in 2010 was some 170,000 tonnes, of which 115,000 tonnes were from the United States and the rest was equally divided between Canada and Russia. This corresponds to some 65 million fish in all, and to some 19% of the harvest of all Pacific salmon species by weight.[31]
Smoked sockeye salmon ready for consumption
Commercial fishermen in Alaska net this species using seines and gillnets for fresh or frozen fillet sales and canning. The annual catch can reach 30 million fish in Bristol Bay, Alaska, which is the site of the world's largest sockeye harvest.[32]
Sockeye salmon have long been important in the diet and culture of the Coast Salish people of British Columbia.
The largest spawning grounds in Asia are located on the Kamchatka Peninsula of the Russian Far East, especially on the Ozernaya River of the Kurile Lake, which accounts for nearly 90% of all Asian sockeye salmon production,[33] and is recognized as the largest spawning ground outside of Alaska.[34] Illegal fishing in Kamchatka is subject to environmental concern.[35]
Sockeye is almost never farmed. A facility in Langley, BC harvested its first salmon in March 2013, and continues to harvest farmed salmon from its inland facility.[36][37]
Conservation status[edit]
See also: Salmon conservation and Steelhead and salmon distinct population segments
United States[edit]
Spawning Kokanee salmon in the Sawtooth Range of Idaho, United States
United States sockeye salmon populations are currently listed under the US Endangered Species Act[38] by the National Marine Fisheries Service as an endangered species in the Snake River and as a threatened species in Lake Ozette, Washington. The Snake River sockeye salmon was listed as endangered in November 1991, after the Shoshone-Bannock Tribe at Fort Hall Indian Reservation petitioned the National Marine Fisheries Service. Other sockeye populations in the upper Columbia River and in Puget Sound are not listed under the Act.
Sockeye is an exception to 2010's forecast resurgence of Oregonian fish stocks. Spring Chinook, summer steelhead, and Coho are forecast to increase by up to 100% over 2008 populations. The sockeye population peaked at over 200,000 in 2008 and were forecast to decline to just over 100,000 in 2010. As an early indication of the unexpectedly high sockeye run in 2010, on July 2, 2010, the United States Army Corps of Engineers reported over 300,000 sockeye had passed over Bonneville Dam on the Columbia River. Lower temperatures in 2008 North Pacific waters brought in fatter plankton, which, along with greater outflows of Columbia River water, fed the resurgent populations.[39]
Proposed legislative efforts, such as the Northern Rockies Ecosystem Protection Act, are attempting to protect the headwaters of the sockeye salmon by preventing industrial development in roadless areas.
Record numbers of a once-waning population of sockeye salmon have been returning to the Northwest's Columbia Basin (as of June 2012), with thousands more crossing the river's dams in a single day than the total numbers seen in some previous years.[40]
Canada[edit]
See also: Conservation status of British Columbia salmonids
Sockeye salmon jumping over a beaver dam, Aleknagik Lake, Alaska, United States
The conservation status of sockeye populations in Canada is under review by Fisheries and Oceans Canada as part of its Wild Salmon Policy strategy to standardize monitoring of wild salmon status.[41] Salmon runs of particular note are the Skeena and Nass river runs, and the most famous is the Fraser River sockeye run.
The Fraser River salmon run has experienced declines in productivity since the 1990s, mirroring a similar decline in the 1960s.[42]
The return abundance (population) of Fraser River sockeye in 2009 was estimated at a very low 1,370,000,[43] 13% of the pre-season forecast of 10,488,000.[44] That represented a decline from the recent (1993) historical cycle peak of 23,631,000 [45] and the return abundance was the lowest in over 50 years. The reasons for this (former) decline remain speculative. According to a consortium of scientists assembled to review the problem, the decline highlights the uncertainty in forecasting salmon returns. [46] After the low returns, the Government of Canada launched a formal inquiry into the decline, the Commission of Inquiry into the Decline of Sockeye Salmon in the Fraser River.[47] [48]
The Commission has been tasked with investigating all the factors which may affect Fraser River sockeye salmon throughout their life cycle. According to the terms of reference,[47] [48] the subjects of investigation are "the impact of environmental changes along the Fraser River, marine environmental conditions, aquaculture, predators, diseases, water temperature and other factors that may have affected the ability of sockeye salmon to reach traditional spawning grounds or reach the ocean."
During the commission, hundreds of thousands of documents and scientific research papers were reviewed. Twelve technical reports were published using that information, looking at the possible impacts of diseases and parasites, hatchery diseases, contaminants, marine ecology, salmon farms, fisheries, predators, climate change and government management on the productivity of Fraser River sockeye runs.[49]
The commission submitted its final report on October 29, 2012.[50][51]
While the commission was holding public hearings, in the late summer of 2010, the largest run of sockeye since 1913 returned to the Fraser River system.[52] Final counts show that approximately 30 million salmon returned to the Fraser River and its tributaries in 2010. In total, approximately 11,591,000 Fraser sockeye were caught by Canadian fishers and 1,974,000 Fraser sockeye were caught by American fishers. The final projected escapement (fish which were not caught) was 15,852,990 fish.[53]
Recent unpredictable fluctuations in runs are speculated to be due to changing water temperatures.[54]
References[edit]
Jump up ^ "iucnredlist.org". iucnredlist.org. Retrieved 2013-01-17.
Jump up ^ "Sockeye Salmon". NOAA Fisheries Office of Protected Resources. Retrieved 2006-11-19.
^ Jump up to: a b c d e "Oncorhynchus nerka". Fishbase. Retrieved 15 November 2013.
Jump up ^ Bright, William (2004). Native American placenames of the United States. University of Oklahoma Press. p. 455. ISBN 978-0-8061-3598-4. Retrieved 11 April 2011.
^ Jump up to: a b c "nationalgeographic.com". Animals.nationalgeographic.com. Retrieved 2013-01-17.
Jump up ^ "Sockeye Salmon". Alaska Department of Fish and Game. Retrieved 2006-11-17.
Jump up ^ "Kokanee Lake". BC Geographical Names.
Jump up ^ "Fish Resources - Salmon/Steelhead". USDA Forest Service. Retrieved 9 September 2011.
Jump up ^ "Fish for Kokanee Salmon at Lake Nantahala". Reflection Lake Nantahala. Retrieved 21 November 2013.
Jump up ^ Scientist says he found Japan fish thought extinct Bay Ledger, December 14, 2010
Jump up ^ "Oncorhynchus kawamurae". Fishbase. Retrieved 21 November 2013.
Jump up ^ Eggers, Douglas M. (3 April 1978). "Limnetic feeding behavior of juvenile sockeye salmon in Lake Washington and predator avoidance". ASLO. 23 (6): 1114–1125. doi:10.4319/lo.1978.23.6.1114.
Jump up ^ Clark, Colin W.; David A. Levy (February 1988). "Diel Vertical Migrations by Juvenile Sockeye Salmon and the Antipredation Window". The American Naturalist. 131 (2): 271–290. doi:10.1086/284789.
Jump up ^ Alaska Department of Fish and Game. "Sockeye salmon". adfg.state.ak.us. Retrieved 2013-01-17.
Jump up ^ Burgner, RL (1991). Life history of sockeye salmon (Oncorhynchus nerka). Pacific salmon life histories: Vancouver, British Columbia: University of British Columbia Press. pp. 3–117.
Jump up ^ Nelson, Joseph S. (1968-02-01). "Distribution and Nomenclature of North American Kokanee, Oncorhynchus nerka". Journal of the Fisheries Research Board of Canada. 25 (2): 409–414. doi:10.1139/f68-032. ISSN 0015-296X.
Jump up ^ "Life history variation and population structure in sockeye salmon - ScienceBase-Catalog". www.sciencebase.gov. Retrieved 2017-11-11.
Jump up ^ "Archived copy". Archived from the original on 2014-12-30. Retrieved 2015-12-30.
^ Jump up to: a b Quinn, Thomas P.; Adkison, Milo D.; Ward, Michael B. (26 April 2010). "Behavioral Tactics of Male Sockeye Salmon (Oncorhynchus nerka) under Varying Operational Sex Ratios". Ethology. 102 (2): 304–322. doi:10.1111/j.1439-0310.1996.tb01127.x.
^ Jump up to: a b c Quinn, Thomas P.; Chris J. Foote (1 October 1994). "The effects of body size and sexual dimorphism on the reproductive behaviour of sockeye salmon, Oncorhynchus nerka". Animal Behaviour. 48 (4): 751–761. doi:10.1006/anbe.1994.1300.
^ Jump up to: a b c d e f g Healey, M.C.; R. Lake; S.G. Hinch (1 February 2003). "Energy expenditures during reproduction by sockeye salmon (Oncorhynchus nerka)". Behaviour. 140 (2): 161–182. doi:10.1163/156853903321671488.
^ Jump up to: a b c Hendry, Andrew P.; Ole K. Berg (1999). "Secondary sexual characters, energy use, senescence, and the cost of reproduction in sockeye salmon". Canadian Journal of Zoology. 77: 1663–1675. doi:10.1139/cjz-77-11-1663.
^ Jump up to: a b Foote, Chris J; Brown, Gayle S; Hawryshyn, Craig W (1 January 2004). "Female colour and male choice in sockeye salmon: implications for the phenotypic convergence of anadromous and nonanadromous morphs". Animal Behaviour. 67 (1): 69–83. doi:10.1016/j.anbehav.2003.02.004.
^ Jump up to: a b Quinn, Thomas P.; Andrew P. Hendry; Gregory B. Buck (2001). "Balancing natural and sexual selection in sockeye salmon: interactions between body size, reproductive opportunity and vulnerability to predation by bears". Evolutionary Ecology Research. 3: 917–937.
^ Jump up to: a b Quinn, Thomas P.; Foote, Chris J. "The effects of body size and sexual dimorphism on the reproductive behaviour of sockeye salmon, Oncorhynchus nerka". Animal Behaviour. 48 (4): 751–761. doi:10.1006/anbe.1994.1300.
Jump up ^ Carlson, Stephanie M.; Hilborn, Ray; Hendry, Andrew P.; Quinn, Thomas P. (2007-12-12). "Predation by Bears Drives Senescence in Natural Populations of Salmon". PLOS ONE. 2 (12): e1286. doi:10.1371/journal.pone.0001286. ISSN 1932-6203.
^ Jump up to: a b Quinn, Thomas P.; Buck, Gregory B. (2001-11-01). "Size- and Sex-Selective Mortality of Adult Sockeye Salmon: Bears, Gulls, and Fish Out of Water". Transactions of the American Fisheries Society. 130 (6): 995–1005. doi:10.1577/1548-8659(2001)1302.0.CO;2. ISSN 0002-8487.
Jump up ^ Carlson, Stephanie M.; Quinn, Thomas P. (2007-10-01). "Ten Years of Varying Lake Level and Selection on Size-at-Maturity in Sockeye Salmon". Ecology. 88 (10): 2620–2629. doi:10.1890/06-1171.1. ISSN 1939-9170.
Jump up ^ Quinn, Thomas P.; Andrew P. Hendry; Lisa A. Wetzel (December 1995). "The Influence of Life History Trade-Offs and the Size of Incubation Gravels on Egg Size Variation in Sockeye Salmon (Oncorhynchus nerka)". Oikos. 74: 425–438. doi:10.2307/3545987.
Jump up ^ Groot, C.; Margolis, L. (1991). Pacific salmon life histories. Vancouver: UBC Press. ISBN 0-7748-0359-2.
Jump up ^ Annual Statistics 2010: Commercial salmon catch by species and country North Pacific Anadromous Fish Commission Statistical Yearbook. Retrieved 2015 March 16. The statistics do not include fish taken in Russian waters by non-Russian fleet.
Jump up ^ "Species profile: Sockeye Salmon (Oncorhynchus nerka)". ADF&G Home. Alaska Department of Fish and Game. Retrieved 14 August 2012.
Jump up ^ Gustafson, R.G., T.C. Wainwright, G.A. Winans, F.W. Waknitz, L.T. Parker, and R.S. Waples. 1997. Status review of sockeye salmon from Washington and Oregon. U.S. Dept. Commer., NOAA Tech. Memo. NMFS-NWFSC-33, 282 p. Life History of Oncorhynchus nerka
Jump up ^ "Discovering Kamchatka: Terrestrial and aquatic fauna". The Royal Geographical Society. 2008. Retrieved 2012-08-14.
Jump up ^ Dronova, Natalia; Spiridonov, Vassily (2008). "Illegal, Unreported, and Unregulated Pacific Salmon Fishing in Kamchatka" (PDF). WWF Russia, IUCN. Archived from the original (PDF) on 24 April 2013. Retrieved 14 August 2012.
Jump up ^ Bland, Alastair (May 2, 2013). "Can Salmon Farming Be Sustainable? Maybe, If You Head Inland". NPR. Retrieved March 21, 2015.
Jump up ^ Shore, Randy (March 27, 2013). "World's first land-based-farm sockeye salmon ready for harvest in B.C." Vancouver Sun. Retrieved March 21, 2015.
Jump up ^ nmfs.noaa.gov, U.S. Endangered Species Act
Jump up ^ "Fish Boom Makes Splash in Oregon". Wall Street Journal. January 21, 2010. Archived from the original on February 9, 2013. Retrieved January 21, 2010.
Jump up ^ "Sockeye Salmon rebounding in Oregon's Columbia Basin". Retrieved 2012-11-21.
Jump up ^ "Wild Salmon Policy 1".
Jump up ^ "Speaking for the Salmon proceedings: Summit of Fraser River Sockeye Salmon: Understanding Stock Declines and Prospects for the Future March 30-31, 2010" (PDF).
Jump up ^ "NewsRelease10" (PDF). Pacific Salmon Commission News Releases. Sep 11, 2009. Archived from the original (PDF) on 2016-03-05. Retrieved Sep 2009. Check date values in: |access-date= (help)
Jump up ^ "NewsRelease01" (PDF). Pacific Salmon Commission News Releases. July 10, 2009. Archived from the original (PDF) on March 5, 2016. Retrieved Jul 2009. Check date values in: |access-date= (help)
Jump up ^ "Fraser River Annual Report to the Pacific Salmon Commission on the 2005 Fraser River sockeye and pink salmon fishing season" (PDF). Fraser River Panel reports to the Pacific Salmon Commission: 26. October 2009. Retrieved Oct 2009. Check date values in: |access-date= (help)[permanent dead link]
Jump up ^ "Managing Fraser sockeye in the face of declining productivity and increasing uncertainty: Statement from Think Tank of Scientists" (PDF). December 2009. Retrieved Dec 2009. Check date values in: |access-date= (help)
^ Jump up to: a b "Terms of Reference for the Commission of Inquiry into Decline of Sockeye Salmon in the Fraser River". November 2009. Archived from the original on 2010-01-13. Retrieved Nov 2009. Check date values in: |access-date= (help) Cite error: Invalid <ref> tag; name "Terms of Reference" defined multiple times with different content (see the help page).
^ Jump up to: a b "Commission of Inquiry into the Decline of Sockeye Salmon in the Fraser River". Archived from the original on June 22, 2012. Retrieved July 22, 2012.
Jump up ^ "Cohen Commission technical reports".
Jump up ^ I., Cohen, Bruce (2012). The uncertain future of Fraser River sockeye. Volume 1, The sockeye fishery. Commission of Inquiry into the Decline of Sockeye Salmon in the Fraser River. ISBN 9780660202143. OCLC 798416074.
Jump up ^ "Cohen Commission to submit final report by October 29, 2012". Archived from the original on May 18, 2013.
Jump up ^ "Consumers catch a deal in record sockeye run". August 27, 2010. Retrieved Sep 7, 2010.
Jump up ^ "Surveillance Report British Columbia Commercial Sockeye Salmon Fisheries. November 2011". Archived from the original on 2013-05-18.
Jump up ^ "Record number of sockeye salmon return in B.C." August 25, 2010. Retrieved Sep 7, 2010.
"Oncorhynchus nerka". Integrated Taxonomic Information System. Retrieved 30 January 2006.
Technical reports[edit]
Bristol Bay sockeye salmon inriver test fishing, 2004 / by Frederick W. West. Hosted by the Alaska State Publications Program.
Hetta and Eek Lakes subsistence sockeye salmon project : 2004 annual report / by Jan M. Conitz ... et al.. Hosted by Alaska State Publications Program.
External links[edit]
Wikimedia Commons has media related to Oncorhynchus nerka.
NOAA Fisheries sockeye salmon web page
FishBase entry for Oncorhynchus nerka
Animal Diversity Web entry for Oncorhynchus nerka
Species Profile
National Geographic Sockeye salmon
Watershed Watch Salmon Society A British Columbia advocacy group for wild salmon
Salmon/Steelhead page of the USDA Forest Service, Pacific-Northwest Fisheries Program.
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