The early life and military career of John Sidney McCain III spans the first forty-five years of his life (1936–1981). McCain's father and grandfather were admirals in the United States Navy. McCain was born on August 29, 1936, in the Panama Canal Zone, and attended many schools growing up as his family moved among naval facilities. McCain graduated from the United States Naval Academy in 1958. He married the former Carol Shepp in 1965; he adopted two children from her previous marriage and they had another child together.
As a naval aviator, McCain flew attack aircraft from carriers. During the Vietnam War, he narrowly escaped death in the 1967 Forrestal fire. On his twenty-third bombing mission in October 1967, he was shot down over Hanoi and badly injured. He subsequently endured five and a half years as a prisoner of war, including periods of torture. In 1968, he refused a North Vietnamese offer of early release, because it would have meant leaving before other prisoners who had been held longer. He was released in 1973 after the Paris Peace Accords.
Upon his return, McCain studied at the National War College, commanded a large training squadron in Florida, and was appointed the Navy liaison to the U.S. Senate. He divorced his wife Carol in 1980 and married the former Cindy Hensley shortly thereafter. He retired from the Navy in 1981 as a captain.
Early years and education
John Sidney McCain III was born on August 29, 1936, at a United States Navy hospital at Coco Solo Naval Air Station in the Panama Canal Zone, which at that time was considered to be among the unincorporated territories of the United States. His parents were Navy officer John S. "Jack" McCain, Jr. (1911–1981) and Roberta (Wright) McCain (born 1912). McCain is of Scots-Irish and English ancestry.
John McCain's grandparents were natives of Arkansas, Mississippi, and Texas, and much of his ancestry was Southern on both his mother's side and father's side. The McCain family's patrilineal ancestral home is in Mississippi's Carroll County; they owned and ran a 2,000-acre (8.1 km2) plantation in Teoc from 1848 until 1952. The plantation had slaves before the American Civil War – some of whose descendants share the surname and call themselves the "black McCains" – and sharecroppers afterward; influential blues guitarist Mississippi John Hurt was born on the plantation to one of the latter.
The McCain family tree has a long heritage of American military service, with ancestors fighting as soldiers in the Indian Wars,American Revolutionary War (due to which McCain maintains a membership with the Sons of the American Revolution),War of 1812, for the Confederate States of America in the American Civil War, and in World War I. The tree also includes roguish behavior and economic success. John McCain's maternal grandfather, Archibald Wright (1875–1971), was a Mississippi native who migrated to Muskogee, Oklahoma, in his twenties, ran afoul of the law with several gambling and bootlegging charges, then became a strong-willed wildcatter who prospered on land deals during the early statehood years and struck oil in the Southwest. Rich by age forty, he never worked again and became a stay-at-home father. Raising a family in Oklahoma and Southern California, he instilled in Roberta and her twin sister Rowena a lifelong habit of travel and adventure. There is also independent-minded behavior in the family tree: Jack McCain and Roberta Wright eloped and married in a bar in Tijuana, Mexico, when Archibald Wright's wife Myrtle objected to Roberta's association with a sailor.
McCain's father and paternal grandfather eventually became Navy admirals, and were the first father–son pair to achieve four-star admiral rank. His grandfather, Admiral John S. "Slew" McCain, Sr. (1884–1945), was a pioneer of aircraft carrier operations who in 1942 commanded all land-based air operations in support of the Guadalcanal Campaign, and who ultimately in 1944–1945 aggressively led the Fast Carrier Task Force in the Pacific Ocean theater of World War II. His operations off the Philippines and Okinawa, and air strikes against Formosa and the Japanese home islands, caused tremendous destruction of Japanese naval and air forces in the closing period of the war. His death four days after the Japanese surrender ceremony in Tokyo Bay was front page news. Jack McCain was a submarine commander in several theaters of operation in World War II and was decorated with both the Silver Star Medal and Bronze Star Medal.
For his first ten years, "Johnny" McCain (the nickname he was given as part of a family tradition of distinguishing the generations) was frequently uprooted as his family, including older sister Sandy (born 1934) and younger brother Joe (born 1942), followed his father to New London, Connecticut, Pearl Harbor, Hawaii, and other stations in the Pacific Ocean. Summer vacations were sometimes spent at the family's Teoc plantation, but McCain always felt his heritage was military, not Southern. McCain attended whatever naval base school was available, often to the detriment of his education, as schools were sometimes substandard and their curricula often erratic. After the 1941 attack on Pearl Harbor, his father was absent for long stretches. His formal education was supplemented by the efforts of his mother, who took advantage of the family's many long-distance travels to expose him to historical and cultural sites. He later wrote, "She taught me to find so much pleasure in life that misfortune could not rob me of the joy of living." A Republican, she also made sure that he followed current events, although his parents avoided outward partisan affiliations due to his father's military career.
After World War II ended, his father stayed in the Navy, sometimes working political liaison posts. The family settled in Northern Virginia, and McCain attended the educationally stronger St. Stephen's School in Alexandria from 1946 to 1949. To his family, McCain had long been quiet, dependable, and courteous, while at St. Stephen's he began to develop an unruly, defiant streak. Another two years were then spent following his father to naval stations; altogether he attended about twenty schools during his youth. He was frequently disciplined in school for fighting. He later wrote, "The repeated farewells to friends rank among the saddest regrets of a childhood constantly disrupted by the demands of my father's career... At each new school I arrived eager to make, by means of my insolent attitude, new friends to compensate for the loss of others. At each new school I grew more determined to assert my crude individualism. At each new school I became a more unrepentant pain in the neck."
In 1951, McCain enrolled at Episcopal High School in Alexandria, an academically superior, all-male private boarding school with a rigorous honor code, tradition of hazing, and spartan living environment. Most of the children there were sons of wealthy Southerners, from whom McCain got a glimpse of life and career aspirations outside the Navy culture. Nicknamed "Punk" and "McNasty" due to his combative, fiery disposition, McCain enjoyed and cultivated a tough guy image; he also made a few friends. McCain earned two varsity letters in wrestling, excelling in the lighter weight classes. He also played on the junior varsityfootball team and the tennis team, and participated in the student newspaper, yearbook, and drama club. English teacher William Bee Ravenel III, who was also his football coach, became a great influence towards his sense of learning, honor, and self-image. With what he later termed an "undistinguished, but acceptable" academic record, McCain graduated from high school in 1954.
Having done well on its entrance exams, McCain entered the United States Naval Academy in Annapolis, Maryland, in June 1954, following in the footsteps of his father and grandfather. He had neither been ordered to go there by his parents nor discussed alternatives; as he later wrote, "I remember simply recognizing my eventual enrollment at the Academy as an immutable fact of life, and accepting it without comment."
Ambivalent about his presence there, McCain chose not to conform to the Academy's rules and some of its traditions. Each year he was given over a hundred demerits – earning him membership in the "Century Club" – for offenses such as shoes not being shined, formation faults, room in disorder, and talking out of place. His father came to the Academy to reprimand him on his behavior a number of times. He hated "plebe year", the trial by ordeal and hazing of entering midshipmen that would eventually weed out one quarter of the class. He did not take well to those of higher rank arbitrarily wielding power over him – "It was bullshit, and I resented the hell out of it" – and occasionally intervened when he saw it being done to others. At 5-foot 7 inches and 127 pounds (1.70 m and 58 kg), he competed as a lightweight boxer for three years, where he lacked skills but was fearless and "didn't have a reverse gear". In his final year, he managed the battalion boxing team to a brigade championship.
Possessed of a strong intelligence, McCain did well in a few subjects that interested him, such as English literature, history, and government. There was a fixed Bachelor of Science curriculum taken by all midshipmen; McCain's classmates were impressed by his cramming abilities on mathematics, science, and engineering courses and thought his low grades were by inclination and not ability, while McCain would later acknowledge that those courses were a struggle for him. His class rank was further lowered by poor grades for conduct and leadership, which reflected his sloppy appearance, rebellious attitude, and poor relations with his company officer. Despite his low standing, he was popular and a leader among his fellow midshipmen, in what biographer Robert Timberg called a "manic, intuitive, highly idiosyncratic way". Good at attracting women, he was famed for organizing off-Yard activities with a group who called themselves "the Bad Bunch"; one classmate said that "being on liberty with John McCain was like being in a train wreck." Other midshipmen were annoyed by his behavior. A June 1957 training cruise aboard the destroyer USS Hunt found McCain showing good skills at the conn, and the destination stop in Rio de Janeiro led to a dream-like romance with Brazilian fashion model and ballerina Maria Gracinda that persisted through a Christmastime reunion.
McCain graduated from the Naval Academy in June 1958; he was fifth from the bottom in class rank, 894th out of 899. Despite his difficulties, McCain later wrote that he never defamed the more compelling traditions of the Academy – courage, resilience, honor, and sacrifice for one's country – and he never wavered in his desire to show his father and family that he was of the same mettle as his naval forebears. Indeed, Slew and Jack McCain had not had sterling records at the Academy themselves, finishing in the bottom third and bottom twentieth respectively. McCain realized later that the Academy had taught him that "to sustain my self-respect for a lifetime it would be necessary for me to have the honor of serving something greater than my self-interest", a lesson that he would need to carry him through a "desperate and uncertain" time a decade later.
Naval training, early assignments, first marriage, and children
McCain was commissioned an ensign. He spent two years as a naval aviator in training, first at Naval Air Station Pensacola in Florida through September 1959, and then at Naval Air Station Corpus Christi in Texas, during which time he was promoted to lieutenant, junior grade. He earned a reputation as a party man, as he drove a Corvette, dated an exotic dancer named "Marie the Flame of Florida", spent all his free time on the beach or in a Bachelor Officer Quarters room turned bar and friendly gambling den, and, as he later said, "generally misused my good health and youth". He began as a sub-par flier: he had limited patience for studying aviation manuals, and spent study time reading history books instead. He was not assigned to the elite units flying fighter aircraft, and instead became a pilot of attack aircraft. During a March 1960 practice run in Texas, he lost track of his altitude and speed, and his single-seat, single-engined AD-6 Skyraider crashed into Corpus Christi Bay and sank to the bottom. Although momentarily knocked unconscious by the impact, he squeezed out of the cockpit and swam ten feet to the surface, escaping without major injuries. He graduated from flight school at Corpus Christi in May 1960. He joined squadron VA-42 at Naval Air Station Oceana in Virginia for five months of further training on the Skyraider.
Starting in November 1960, McCain flew Skyraiders with the VA-65 "World Famous Fighting Tigers" squadron on the aircraft carriers USS Intrepid and USS Enterprise. The carriers were based at Naval Station Norfolk and cruised in the Caribbean and in several deployments to the Mediterranean. His aviation skills improved, but around December 1961 he collided with power lines while recklessly flying too low over southern Spain. The area suffered a power outage, but McCain was able to return his damaged Skyraider to Intrepid.
On board for Enterprise's maiden voyage in January 1962, McCain gained visibility with the captain and shipboard publicity that fellow sailors and aviators attributed to his famous last name. McCain was made a lieutenant in June 1962, and was on alert duty on Enterprise when it helped enforce the naval quarantine of Cuba during the October 1962 Cuban Missile Crisis. In November 1963, he was rotated back to shore duty, serving nine months on the staff of the Naval Air Basic Training Command at Pensacola. In September 1964, he became a flight instructor with the VT-7 training squadron at Naval Air Station Meridian in Mississippi, where McCain Field had been named for his grandfather.
During the 1964 stint at Pensacola, McCain began a relationship with Carol Shepp, a successful swimwear and runway model originally from Philadelphia, Pennsylvania. They had known each other at the Naval Academy and she had married and then divorced one of his classmates. McCain told her he wanted to do something important with his life, so he would be recorded in history. On July 3, 1965, McCain married Shepp in Philadelphia. She already had two children, Douglas and Andrew, born in 1959 and 1962 respectively; he adopted them in 1966. Carol and he then had a daughter named Sidney in September 1966.
In July 1965, McCain appeared as a contestant on the quiz showJeopardy! (during the Art Fleming era). McCain won the first day, but lost in Final Jeopardy on the second day. He recalls the final question being "Cathy loves him, but she married Edgar Linton instead". McCain wrote down "What is Wuthering Heights" although the question was looking for the specific character, "Who is Heathcliff". .
In November 1965, he had his third accident when apparent engine failure in his T-2 Buckeye trainer jet over the Eastern Shore of Virginia led to his ejecting safely before his plane crashed. While at Meridian, McCain requested a combat assignment. In October 1966, he was slated for upcoming Vietnam War duty, and so reported to the VA-44Replacement Air Group squadron at Naval Air Station Cecil Field in Florida for training on the A-4 Skyhawk, a single-seat jet attack aircraft. There McCain was seen as a good pilot, albeit one who tended to "push the envelope" in his flying. Promoted to lieutenant commander in January 1967, McCain joined the aircraft carrier USS Forrestal by May 1967, flying Skyhawks with the VA-46 "Clansmen" squadron.Forrestal conducted training exercises in the Atlantic early in the year, then set sail for the Pacific in June. By this time, Jack McCain had risen in the ranks, making rear admiral in 1958 and vice admiral in 1963; in May 1967, he was promoted to four-star admiral, and became Commander-in-Chief, U.S. Naval Forces, Europe, stationed in London.
On July 25, 1967, Forrestal reached Yankee Station in the Gulf of Tonkin and joined Operation Rolling Thunder, the 1965–1968 air interdiction and strategic bombing campaign against North Vietnam. The alpha strikes flown from Forrestal were against specific, pre-selected targets such as arms depots, factories, and bridges. They were quite dangerous, due to the strength of the North Vietnamese air defenses, which used Soviet-designed and -supplied surface-to-air missiles, anti-aircraft artillery, and MiG jet interceptors. McCain's first five attack missions over North Vietnam went without incident, and while still unconcerned with minor Navy regulations, McCain had garnered the reputation of a serious aviator. McCain and his fellow pilots were frustrated by the micromanagement of Rolling Thunder from Washington; he later wrote, "The target list was so restricted that we had to go back and hit the same targets over and over again... Most of our pilots flying the missions believed that our targets were virtually worthless. In all candor, we thought our civilian commanders were complete idiots who didn't have the least notion of what it took to win the war."
McCain was almost killed on board Forrestal on July 29, 1967. While the air wing was preparing to launch attacks, a Zuni rocket from an F-4 Phantom accidentally fired across the carrier's deck. The rocket struck either McCain's A-4E Skyhawk or one near it. The impact ruptured the Skyhawk's fuel tank, which ignited the fuel and knocked two bombs loose. McCain later said, "I thought my aircraft exploded. Flames were everywhere." McCain escaped from his jet by climbing out of the cockpit, working himself to the nose of the jet, and jumping off its refueling probe onto the burning deck. His flight suit caught on fire as he rolled through the flames, but he was able to put it out. He went to help another pilot trying to escape the fire when the first bomb exploded; McCain was thrown backwards ten feet (three meters) and suffered minor wounds when struck in the legs and chest by fragments. McCain helped crewmen throw unexploded bombs overboard off the hangar deck elevator, then went to Forrestal's ready room and with other pilots watched the ensuing fire and the fire-fighting efforts on the room's closed-circuit television. The fire killed 134 sailors, injured scores of others, destroyed at least 20 aircraft, and took 24 hours to control. In Saigon a day after the conflagration, McCain praised the heroism of enlisted men who gave their lives trying to save the pilots on deck, and told New York Times reporter R. W. Apple, Jr., "It's a difficult thing to say. But now that I've seen what the bombs and the napalm did to the people on our ship, I'm not so sure that I want to drop any more of that stuff on North Vietnam." But such a change of course was unlikely; as McCain added, "I always wanted to be in the Navy. I was born into it and I never really considered another profession. But I always had trouble with the regimentation."
As Forrestal headed to port for repairs, McCain volunteered to join the undermanned VA-163 "Saints" squadron on board the USS Oriskany. This carrier had earlier endured its own deck fire disaster and its squadrons had suffered some of the heaviest losses during Rolling Thunder. The Saints had a reputation for aggressive, daring attacks, but paid the price: in 1967, one-third of their pilots were killed or captured, and all of their original fifteen A-4s had been destroyed. After taking some leave in Europe and back home in Orange Park, Florida, McCain joined Oriskany on September 30, 1967, for a tour he expected would finish early the next summer. He volunteered to fly the squadron's most dangerous missions right away, rather than work his way up to them. During October 1967, the pilots operated in constant twelve-hour on, twelve-hour off shifts. McCain would be awarded a Navy Commendation Medal for leading his air section through heavy enemy fire during an October 18 raid on the Lac Trai shipyard in Haiphong. On October 25, McCain successfully attacked the Phúc Yên Air Base north of Hanoi through a barrage of anti-aircraft artillery and surface-to-air missile fire; credited with destroying one aircraft on the ground and damaging two, the raid would garner him the Air Medal. Air defenses around Hanoi were at this point the strongest they would be during the entire war.
Prisoner of war
On October 26, 1967, McCain was flying his twenty-third mission, part of a twenty-plane strike force against the Yen Phu thermal power plant in central Hanoi that previously had almost always been off-limits to U.S. raids due to the possibility of collateral damage. Arriving just before noon, McCain dove from 9,000 to 4,000 feet on his approach; as he neared the target, warning systems in McCain's A-4E Skyhawk alerted him that he was being tracked by enemy fire-control radar. Like other U.S. pilots in similar situations, he did not break off the bombing run, and he held his dive until he released his bombs at about 3,500 feet (1,000 m). As he started to pull up, the Skyhawk's wing was blown off by a Soviet-made SA-2 anti-aircraft missile fired by the North Vietnamese Air Defense Command's 61st Battalion, commanded by Captain Nguyen Lan and with fire control officer Lieutenant Nguyen Xuan Dai. (McCain was later awarded the Distinguished Flying Cross for this day, while Nguyen Xuan Dai was awarded the title Hero of the People's Armed Forces. Decades later, Soviet Army Lieutenant Yuri Trushechkin claimed that he had been the missile guidance officer who had shot McCain down. In any case, the raid was a failure, as the power plant was not damaged and three of the attacking planes were shot down.)
McCain's plane went into a vertical inverted spin. McCain bailed out upside down at high speed; the force of the ejection fractured his right arm in three places, his left arm, and his right leg at the knee, and knocked him unconscious. McCain nearly drowned after parachuting into Trúc Bạch Lake in Hanoi; the weight of his equipment was pulling him down, and as he regained consciousness, he could not use his arms. Eventually, he was able to inflate his life vest using his teeth. Several Vietnamese, possibly led by Department of Industry clerk Mai Van On, pulled him ashore. A mob gathered around, spat on him, kicked him, and stripped him of his clothes; his left shoulder was crushed with the butt of a rifle and he was bayoneted in his left foot and abdominal area. He was then transported to Hanoi's main Hỏa Lò Prison, nicknamed the "Hanoi Hilton" by American POWs.
McCain reached Hỏa Lò in as bad a physical condition as any prisoner during the war. His captors refused to give him medical care unless he gave them military information; they beat and interrogated him, but McCain only offered his name, rank, serial number, and date of birth (the only information he was required to provide under the Geneva Conventions and permitted to give under the U.S. Code of Conduct). Soon thinking he was near death, McCain said he would give them more information if taken to the hospital, hoping he could then put his interrogators off once he was treated. A prison doctor came and said it was too late, as McCain was about to die anyway. Only when the North Vietnamese discovered that his father was a high-ranking admiral did they give him medical care, calling him "the crown prince". Two days after McCain's plane went down, that event and his status as a POW made the front pages of The New York Times and The Washington Post. Interrogation and beatings resumed in the hospital; McCain gave the North Vietnamese his ship's name, squadron's name, and the attack's intended target. This information, along with personal details of McCain's life and purported statements by McCain about the war's progress, would appear over the next two weeks in the North Vietnamese official newspaper Nhân Dân as well as in dispatches from outlets such as the Cuban news agency Prensa Latina. Disclosing the military information was in violation of the Code of Conduct, which McCain later wrote he regretted, although he saw the information as being of no practical use to the North Vietnamese. Further coerced to give future targets, he named cities that had already been bombed, and responding to demands for the names of his squadron's members, he supplied instead the names of the Green Bay Packers' offensive line.
McCain spent six weeks in the hospital, receiving marginal care in a dirty, wet environment. A prolonged attempt to set the fractures on his right arm, done without anesthetic, was unsuccessful; he received an operation on his broken leg but no treatment for his broken left arm. He was temporarily taken to a clean room and interviewed by a French journalist, François Chalais, whose report was carried on the French television program Panorama in January 1968 and later in the U.S. on the CBS Evening News. The film footage of McCain lying in the bed, in a cast, smoking cigarettes and speaking haltingly, would become one of the most widely distributed images of McCain's imprisonment. McCain was observed by a variety of North Vietnamese, including renowned Vietnamese writer Nguyễn Tuân and Defense Minister and Army commander-in-chief General Võ Nguyên Giáp. Many of the North Vietnamese observers assumed that McCain must be part of America's political-military-economic elite. Now having lost fifty pounds (twenty-three kilograms), in a chest cast, covered in grime and eyes full of fever, and with his hair turned white, in early December 1967 McCain was sent to a prisoner-of-war camp on the outskirts of Hanoi nicknamed "the Plantation". He was placed in a cell with George "Bud" Day, a badly injured and tortured Air Force pilot (later awarded the Medal of Honor) and Norris Overly, another Air Force pilot; they did not expect McCain to live another week. Overly, and subsequently Day, nursed McCain and kept him alive; Day later recalled that McCain had "a fantastic will to live".
In March 1968, McCain was put into solitary confinement, where he remained for two years. Unknown to the POWs, in April 1968, Jack McCain was named Commander-in-Chief, Pacific Command (CINCPAC) effective in July, stationed in Honolulu and commander of all U.S. forces in the Vietnam theater. In mid-June, Major Bai, commander of the North Vietnamese prison camp system, offered McCain a chance to return home early. The North Vietnamese wanted to score a worldwide propaganda coup by appearing merciful, and also wanted to show other POWs that members of the elite like McCain were willing to be treated preferentially. McCain turned down the offer of release, due to the POWs' "first in, first out" interpretation of the U.S. Code of Conduct: he would only accept the offer if every man captured before him was released as well. McCain's refusal to be released was remarked upon by North Vietnamese senior negotiator Lê Đức Thọ to U.S. envoy Averell Harriman, during the ongoing Paris Peace Talks. Enraged by his declining of the offer, Bai and his assistant told McCain that things would get very bad for him.
In late August 1968, a program of vigorous torture methods began on McCain. The North Vietnamese used rope bindings to put him into prolonged, painful positions and severely beat him every two hours, all while he was suffering from dysentery. His right leg was reinjured, his ribs were cracked, some teeth were broken at the gumline, and his left arm was re-fractured. Lying in his own waste, his spirit was broken; the beginnings of a suicide attempt were stopped by guards. After four days of this, McCain signed and taped an anti-American propaganda "confession" that said, in part, "I am a black criminal and I have performed the deeds of an air pirate. I almost died, and the Vietnamese people saved my life, thanks to the doctors." He used stilted Communist jargon and ungrammatical language to signal that the statement was forced. McCain was haunted then and since with the belief that he had dishonored his country, his family, his comrades and himself by his statement, but as he later wrote, "I had learned what we all learned over there: Every man has his breaking point. I had reached mine." Two weeks later his captors tried to force him to sign a second statement; his will to resist restored, he refused. He sometimes received two to three beatings per week because of his continued resistance; the sustained mistreatment went on for over a year. His refusals to cooperate, laced with loud obscenities directed towards his guards, were often heard by other POWs. His boxing experience from his Naval Academy days helped him withstand the battering, and the North Vietnamese did not break him again.
Other American POWs were similarly tortured and maltreated in order to extract "confessions" and propaganda statements. Many, especially among those who had been captured earlier and imprisoned longer – such as those in the "Alcatraz Gang" – endured even worse treatment than McCain. Under extreme duress, virtually all the POWs eventually yielded something to their captors. There were momentary exceptions: on one occasion, a guard surreptitiously loosened McCain's painful rope bindings for a night; when, months later, the guard later saw McCain on Christmas Day, he stood next to McCain and silently drew a cross in the dirt with his foot. In October 1968, McCain's isolation was partly relieved when Ernest C. Brace was placed in the cell next to him; he taught Brace the tap code the prisoners used to communicate.
A new species of Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) from the Quaternary of Chile
Juan Francisco Araya and David G. Reid
Article number: 19.1.8A
Copyright Paleontological Society, February 2016
Plain-language and multi-lingual abstracts
Submission: 19 September 2015. Acceptance: 29 January 2016
We describe a new fossil littorinid species, Echinolittorina nielseni sp. nov., from the Quaternary Caldera Strata, Región de Atacama, northern Chile. Fossils of littorinids are globally rare because of their high-intertidal habitat on rocky shores. The new species has a large, broad shell with strong spiral ribs and an angled periphery, differing from the two living littorinids currently found along the coasts of mainland Chile and from all the extant species distributed in the southeastern Pacific. In comparison with the living Chilean Echinolittorina peruviana, the new species shows stronger ribs and more inflated whorls, but they share an unusual detail in the irregular arrangement of spiral sculpture. We hypothesize that the new species may be ancestral or sister to E.peruviana and discuss the adaptive significance of shell sculpture.
Keywords: Quaternary; Pleistocene; SE Pacific Ocean; Littoraria; new species
Final citation: Araya, Juan Francisco and Reid, David G. 2016. A new species of Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) from the Quaternary of Chile. Palaeontologia Electronica 19.1.8A: 1-8. https://doi.org/10.26879/600
The shallow-water marine molluscs living in northern Chile are relatively well known, having been the subject of regional faunistic studies (e.g., Marincovich, 1973; Guzman et al., 1998; Ashton, 2007; Araya and Araya, 2015), while some members of widespread genera have been included in monographic revisions (e.g., McLean, 1984; Reid, 2002; Geiger, 2012). Nevertheless, new living species continue to be discovered in the area, particularly in the Región de Atacama (Osorio, 2012; Araya, 2013). In comparison, there have been fewer studies of fossil molluscs from coastal northern Chile. Many are old works dealing with the marine species from the Miocene to Pliocene Bahia Inglesa Formation, with a characteristic temperate to tropical fauna (Philippi, 1887; Möricke, 1896; Herm, 1969). Quaternary species from around Caldera were first mentioned by Herm (1969) and have recently been the subject of renewed study (Guzmán et al., 2000; Guicharrousse et al., 2015).
In contrast to the older fossils from northern Chile, those of the Quaternary Caldera Strata include mainly warm-temperate species, of which a high proportion is extant on the nearby coasts. For example, the molluscan assemblage at the locality described here consists of 40 species, all but two of which are still living, including the bivalve Mulinia edulis (King and Broderip, 1832), the gastropod Aeneator aff. fontainei (d’Orbigny, 1839) and the chiton Acanthopleura echinata (Barnes, 1824). This assemblage indicates a coastal, shallow-water environment with sandy pockets and sparse rocky outcrops (cf. Araya, 2013; Araya and Araya, 2015). These species have been extensively reported from Quaternary deposits, while a few of them also have Miocene records (Nielsen, 2013); however, the extant muricid gastropod Concholepas concholepas (Bruguière, 1789) and the scallop Argopecten purpuratus (Lamarck, 1819) indicate that the assemblage is no older than the Pleistocene (Guzmán et al., 2000; Marquardt et al., 2000).
The systematics and phylogeny of the family Littorinidae have been as thoroughly investigated as those of any marine gastropod group (Reid et al., 2012, and references therein) and they have become a model system for the study of diversification in the marine realm. In a dated molecular phylogeny, most of the living littorinids are estimated to have originated in the Oligocene to Miocene, although 46% of the 60 extant Echinolittorina species diverged more recently (Reid et al., 2012). Littorinidae are represented in the living fauna of northern Chile by two common species, Echinolittorina peruviana (Lamarck, 1822) and Austrolittorina araucana (d’Orbigny, 1840) (Reid, 2002; but see Reid et al., 2012 for current generic assignments). The majority of littorinids, including these two, are typically found in abundance at high intertidal levels on wave-exposed rocky shores, where the potential for fossil preservation is generally low. Therefore, fossil records of the family are globally extremely scarce (review by Reid, 1989). As expected, the fossil record of these two species in Chile is very limited, with a single Pleistocene record (not figured) for Echinolittorina peruviana at Caleta Patillos (20°45’S; 70°12’W), in northern Chile (Rivadeneira and Carmona, 2008). For all these reasons, the discovery of a distinctive new species of Littorinidae in the Quaternary Caldera Strata is of interest and is reported here.
MATERIALS AND METHODS
The type locality of the new littorinid species is a small, low cliff to the east of El Morro Hill, one of a series of low terraces 10 km south of Caldera, Región de Atacama, northern Chile (Figure 1). It consists of a semi-consolidated and fossiliferous sedimentary formation composed of littoral marine coquinas, sandstone, cherts, phosphorites and diatomites (Marquardt et al., 2000; Godoy et al., 2003). The upper layer, of variable thickness and which contains the new species, is part of the Caldera Strata and the presence of Concholepas concholepas and Argopecten purpuratus indicates no more than a Pleistocene age (Guzmán et al., 2000; Marquardt et al., 2000). As described above, the molluscan assemblage is consistent with a shallow coastal environment of sand and sparse rocky outcrops (cf. Araya, 2013; Araya and Araya, 2015). No other littorinid taxa were present in the assemblage. The specific location where the new species was found was protected from the open ocean by the El Morro Hill, a massive 2 x 4 km outcrop aligned almost N-S, parallel to the coast, and up to 323 m high (Herm, 1969; Godoy et al., 2003).
Sampling was carried out in January 2012 at an eroded cliff. Bulk samples were obtained from the upper 2 cm of the sediment and further sieved, with some specimens picked out by hand. The specimens described here have been deposited in the collections of the Museo Nacional de Historia Natural, Santiago, Chile (SGO.PI), in the Natural History Museum, London, U.K. (NHMUK), and in the Museo Paleontológico de Caldera, Caldera, Chile (MPCCL). For the morphological descriptions, the shell height parallel to the axis of coiling (H), breadth (B) perpendicular to H and maximum length of aperture (LA) were measured with Vernier callipers (± 0.1 mm).
Phylum MOLLUSCA Linnaeus, 1758
Class GASTROPODA Cuvier, 1795
Subclass CAENOGASTROPODA Cox, 1960
Family LITTORINIDAE Anonymous, 1834
Genus ECHINOLITTORINAHabe, 1956
Type species. Litorina tuberculata Menke, 1828, by original designation.
Distribution. Worldwide tropical and warm temperate seas (Reid et al., 2012). Eocene to Recent.
Echinolittorina nielseni sp. nov.
v. 2015 Echinolittorina n. sp.; Araya and Reid, p. 85, figs 1.A-H.
Diagnosis. Large, broad Echinolittorina with rounded whorls, concave spire profile, 7-9 strong spiral ribs at and above angled periphery.
Description. Shell large (H = 15.7-22.2 mm). Shape high turbinate (H/B = 1.29-1.44; H/LA = 1.54-1.83); spire whorls well rounded, suture distinct; spire profile concave near apex; periphery of last whorl slightly but distinctly angled; rarely a slight angulation at shoulder (Figure 2.6-7). Columella straight, moderately broad, slightly hollowed and pinched at base; no eroded parietal area. Sculpture of 7-9 low, broad, rounded, spiral ribs above periphery (including the strongest rib at periphery), becoming less distinct near suture; ribs of unequal width and prominence, separated by narrow grooves or incised lines. Base with 6-8 finer ribs. No spiral microstriae visible. A trace of colour is preserved on one specimen, showing a pale spiral band on the base (Figure 2.4). None of the specimens showed fluorescence under ultraviolet light, which might have revealed original colour patterns.
Type material. Holotype: SGO.PI 23.100 (Figure 2.1-2); Paratypes NHMUKPAL PI TG 26769-26775 (Figure 2.3-8, seven specimens), MPCCL 14012016 (fifteen specimens). All the material collected at type locality by J. F. Araya, January 15, 2012.
Type locality. Eroded cliff east of El Morro Hill, about 10 km south of Caldera, Región de Atacama, northern Chile (27°09’13”S; 70°55’33”W, 123 m); Pleistocene.
Etymology. The name of the species honours our friend Sven Nielsen (Universidad Austral de Chile, Valdivia, Chile) for his extensive research on Chilean fossil molluscs.
Remarks. These relatively large, thick, spirally-sculptured shells with an entire aperture containing a basal white band, and lacking an umbilicus, unquestionably belong to the subfamily Littorininae (reviewed by Reid, 1989) and can be compared with members of its living genera. Superficially, the shells resemble members of the littorinid genus Littoraria; this assignment is suggested by the strong spiral sculpture, the enlarged peripheral rib that gives an angled profile to the final whorl, and the somewhat short and wide columella. In the recent fauna six Littoraria species occur on the Pacific mainland coast of Central and South America, but all have a tropical distribution and five are almost entirely restricted to mangrove and halophytic-grass habitats. The one rocky-shore species ( Littoraria pintado [Wood, 1828]; see Reid, 1999a; Reid et al., 2010) can be discounted because it is smooth-shelled and relatively narrow with a straight spire. The most southerly records of living Littoraria species are between 3° and 5°S in northernmost Peru (Reid, 1999a). Since the present fossil sample is from 27°S in northern Chile and accompanied by a fauna of temperate, rocky-shore gastropods, their identification as a Littoraria species associated with mangrove trees would be extremely surprising. Of the living Littoraria species, the new species most closely resembles Littoraria varia (Sowerby, 1832) (see Reid, 1999a) (Figure 2.14-15). However, close comparison shows that the sculpture of L. varia is stronger, the primary spiral ribs more equal in width, and that they are separated by wide grooves with secondary ribs and fine spiral microstriae, quite unlike the rounded ribs separated by grooves that are little more than incised lines in the new species. The peripheral angle is marked by a strong rib in L. varia, and there is sometimes a slight angle at the shoulder as well (Figure 2.14), which are points of similarity with the fossils (compare with Figure 2.6-7). One of the fossil shells bears a spiral white band on the base (Figure 2.4); this band is weakly present (and then only within the aperture) in only three of the 39 living Littoraria species (Reid, 1999b), but is present in a variety of other littorinid genera (Reid, 1989), including all members of the genus Echinolittorina (Reid, 2002, 2007, 2009, 2011). The sculptural details and presence of the white basal band exclude the present shells from Littoraria.
Turning to the modern rocky-shore fauna of the eastern Pacific Ocean, there are 18 littorinid species in the genera Echinolittorina and Austrolittorina (see Reid, 2002 for detailed descriptions and Reid et al., 2012 for current generic assignments based on molecular phylogeny). All have a basal white band visible within the aperture and often externally as well. None of the members of Echinolittorina in the tropical eastern Pacific is similar to the new species, being smaller, narrower, usually with regular spiral grooves, and sometimes nodulose or umbilicate. Two of the three temperate littorinid species can also be discounted: Austrolittorina fernandezensis (Rosewater, 1970) is large (to 21 mm), but macroscopically smooth, and is endemic to the Juan Fernández Archipelago, off central Chile; Austrolittorina araucana (d’Orbigny, 1840) is common in Chile, again macroscopically smooth, but does not exceed 13.8 mm in height. The only other living littorinid in the southeastern Pacific is Echinolittorina peruviana (Lamarck, 1822). This is a well-known species, common on temperate shores from northern Peru to southern Chile (5°05’S to 41°49’S; Reid, 2002; Castillo and Brown, 2010), where it is found in local assemblages that are much the same as that in which the fossil shells occurred. It too can reach a large size, up to 23.8 mm high. However, shells of E. peruviana are usually smooth, relatively elongate (H/B = 1.31-1.88; H/LA = 1.46-1.93), the whorls are not swollen, the columella is long and the anterior apertural margin protrudes slightly (Figure 2.9-12) (Reid, 2002)—characters which at first appear to exclude close relationship with the new species. However, a very few shells of E. peruviana (13/884 = 1.4% of shells examined from throughout geographical range) show “1-4 (rarely 6-8) faint incised lines above periphery” (Reid, 2002: 147) (Figure 2.9,12-13). Sometimes, even in smooth shells, irregularities in the axial colour stripes show a periodicity similar to that of the incised spiral lines (compare Figure 2.11 and 2.12). The incised lines are shown more clearly in an SEM of a well-preserved juvenile shell (Figure 2.13). The spacing of the incised spiral lines in these rare shells of E . peruviana is not as regular as the ‘primary spiral grooves’ of typical tropical Echinolittorina species (e.g., Reid, 2002), instead recalling the somewhat unequal ribs of the fossils. It is, therefore, proposed that E. nielseni sp. nov. is a member of Echinolittorina and is most closely related to E. peruviana.
Generic assignment of littorinid shells in the subfamily Littorininae, even of living species, can be difficult because shell colour pattern and shape are quite uniform across the subfamily, whereas sculpture is variable, but prone to homoplasy (Reid, 1989). Shells are, therefore, often an unreliable guide to affinity (even at generic level), so that the advent of systematic anatomical studies (Reid, 1989) and of molecular phylogenetics (e.g., Reid et al., 2012) have had profound effects on littorinid classification. Littorinids mainly inhabit wave-swept intertidal shores, and their fossils are consequently extremely rare (Reid, 1989). These rare fossils are seldom well preserved and their classification presents considerable difficulty. The 23 shells in this sample are not significantly damaged by erosion or abrasion, so their surface sculpture is clear, and there is even a trace of colour pattern on one specimen. While their generic placement in Echinolittorina is not immediately obvious (as discussed in detail above), our observations of sculptural details and the basal white band make a strong case.
To assess whether the fossil shells differ sufficiently from the living form to justify diagnosis as a separate species, it is necessary to consider their respective ranges of morphological variation. Only about 1.4% of living specimens of E. peruviana bear faint incised spiral lines and none show the strong, rounded spiral ribs of the fossil shells. The living shells have less rounded whorls and are usually more elongate than the fossils, although there is overlap in the range of the ratio H/B. There are also subtle differences in the shape of the aperture. It could be argued that, since the fossil shells originate from a single locality, they represent an isolated population of extreme variants of E. peruviana. However, all Echinolittorina species have planktotrophic development and their larvae are likely able to disperse for more than 1400 km in ocean currents (Williams and Reid, 2004). Isolation of a small local population, followed by genetic drift or selection, is not likely and there is no evidence for such effects in this genus. The type locality of the new species (27°S) lies near the midpoint of the modern range of E. peruviana (5-42°S), so at least under current conditions the fossil locality is not near the edge of the range of the living species. Striking variation in shell sculpture on a small local scale does occur in some species of Echinolittorina, but is the result of ecophenotypic plasticity (Yeap et al., 2001; Reid, 2007). This usually takes the form of nodules and spiral striae in small individuals and spiral striae alone in large ones; the development of nodules is apparently connected with slow growth in unfavourable hot, dry microhabitats on horizontal rock surfaces. Reid (2002) examined 63 samples and over 800 shells of E. peruviana from throughout its range and reported no cases of localized variation consistent with ecophenotypic effects on shell sculpture. We, therefore, conclude that description of the new species is justified.
The great majority of Echinolittorina species bear strong spiral striae, spiral ribs or rows of nodules (Reid, 2002, 2007, 2009, 2011), as do most other members of the Littorininae (Reid, 1989). Outgroup comparison suggests, therefore, that the sculpture of E. nielseni is plesiomorphic. The only fossils of E. peruviana recorded from Chile were not figured (Rivadeneira and Carmona, 2008). With additional sampling of fossil shells it might be possible to address hypotheses such as whether E. nielseni is the sculptured ancestor of the smooth species E. peruviana, or whether the two are sister species. If the latter, then geographical separation can be predicted, because sister species of Echinolittorina are almost always allopatric, implying that geographical isolation is required for speciation (Williams and Reid, 2004). Strongly sculptured (both ribbed and nodulose) littorinid species tend to predominate in the tropics and at higher tidal levels on the shore (Vermeij, 1973). Furthermore, the one well-studied case of ecophenotypic plasticity in Echinolittorina (Yeap et al., 2001) suggested that strong sculpture might be functional in relation to convective heat loss. At present, smooth-shelled E. peruviana extends to higher latitudes than all but two of the 60 worldwide living Echinolittorina species (Williams and Reid, 2004; Reid et al., 2012). We, therefore, hypothesize that if E . peruviana and E. nielseni were once allopatric sister species, the latter may have had the more northerly (more tropical) distribution. Alternatively, anagenetic change from a sculptured ancestor to a smooth descendant species could suggest adaptation to a cooling climate. In the molecular phylogeny of Echinolittorina, no living species was identified as sister to E. peruviana; it was a basal member of an Eocene clade (Reid et al., 2012), indicating a long period of independent evolution or extinction of related species.
The assemblage of fossil molluscs at the present site indicates a coastal shallow-water environment. The abundant shell remains of the bivalve Mulinia edulis, which lives exclusively on open sandy beaches, and the presence of species of the gastropod genus Chlorostoma ( C. atrum [Lesson, 1830], C. luctuosa [d´Orbigny, 1841] and C . tridentata [Potiez and Michaud, 1838]), and of some polyplacophorans and limpets—known to live exclusively on rocky shores—points to a mixed area with rocky and sandy habitats, which at the time was protected from the open Pacific Ocean by the El Morro outcrop. With the exception of the new Echinolittorina described herein and one Cyclocardia species, all the molluscan taxa found at the site are currently represented in the living fauna of the area (Guicharrousse et al., 2015).
The authors are grateful to S. Nielsen (Universidad Austral de Chile, Valdivia, Chile) for his help with literature and information on related species; to the staff of the library of the Facultad de Ciencias and of the Departamento de Geología of the Universidad de Chile, in Santiago, Chile for their help with geological literature for the site; to M.E. Araya (Caldera, Chile) for her essential help in the field; to H. Taylor (NHMUK) for taking the photographs; to M. Guicharrouse and G. Roa (MPCCL) for their help in preparing the map of Figure 1; and to A. Beu (GNS Science, Lower Hutt, New Zealand) and an anonymous reviewer for corrections and suggestions that improved this work.
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