YEAR IN REVIEW 1996: LIFE-SCIENCES: ZOOLOGY

ZOOLOGY: Entomology. Most modern insects possess wings and can fly. The evolutionary development of wings and flight in insects, however, is obscure because of the lack of transitional forms between earlier wingless and later winged forms in the fossil record. To demonstrate a possible intermediate step in wing evolution, James H. Marden and Melissa G. Kramer of Pennsylvania State University investigated primitive aquatic insects called stoneflies that have wings but do not fly. Taeniopteryx burksi move across water by surface skimming, with the body supported by water but propelled forward by the wings. Allocapnia vivipara sail across water by raising their wings to catch the wind. In experiments comparing surface-skimming and sailing abilities in stoneflies having artificially shortened wings of various sizes, the researchers found that greater wing area generally resulted in greater speed. They proposed that ancestral stoneflies and other semiaquatic insects used gill structures to move across the water's surface and that over time the advantages of faster surface skimming or sailing favoured an increase in the size of those structures, ultimately leading to wings and wing-powered flight. The defensive behaviour with which some honeybees (genus Apis) respond to attacks by hornets may be an example of coevolution, according to a study carried out in Japan by Masato Ono and colleagues of Tamagawa University, Tokyo. Giant hornets (Vespa mandarinia japonica) make orchestrated attacks on other social hymenopterans such as honeybees, which they kill with their mandibles and feed to their larvae. The investigators confirmed that an individual giant hornet marks a bee colony with a pheromone (chemical attractant) from specialized glands. Additional giant hornets then congregate and initiate a slaughter attack. Introduced European honeybees (A. mellifera) appear defenseless against the hornets and are killed at rates as high as 40 per minute. Native Japanese honeybees (A. cerana japonica), however, can detect the hornet pheromone and change their behaviour by increasing the number of defending bees. More than 500 bees swarm around an attacking hornet, forming a ball whose internal temperature reaches 47 C (117 F), high enough to kill the hornet but not the bees. European bees seem unaware of the hornet pheromone and do not respond effectively to the hornet attacks. The differential responses of the two bee species suggest that the Japanese honeybees have coevolved with the predator and developed an effective defense. The evolutionary history of symbiotic relationships between fungus-growing ants (tribe Attini) and their fungi was investigated by Ulrich G. Mueller and colleagues of Cornell University, Ithaca, N.Y., and the U.S. Department of Agriculture. Using ribosomal DNA analysis and morphological characteristics to compare phylogenies, or evolutionary family trees, for the ants and their fungi, they concluded that whereas the ants originated from a single ancestral form, the cultivated fungi had more than one ancestral line, which indicated that ants developed symbiotic relationships with different fungal lineages. They also found that the less primitive, generally more specialized species, including the leaf-cutting ants, have been associated with the same fungal lineage for at least 23 million years. In a related study Mitchell Sogin of Woods Hole (Mass.) Marine Biological Laboratory and colleagues, using ribosomal RNA analysis, concluded that the less-primitive leaf-cutting ant species and their fungal symbionts have undergone long-term coevolution. A notable feature of the relationship that exists between ants and fungi is that one symbiont may be inconspicuous yet be essential to the survival of the other. (ANNE R. GIBBONS) This updates the article insect1. ZOOLOGY: Ornithology. Identifying the factors that regulate the number of birds of a particular species breeding in a particular area has been a difficult task but is one of fundamental importance in the study of the natural regulation of animal numbers. I. Newton of the Institute of Terrestrial Ecology, part of the U.K. Natural Environment Research Council, reviewed the results of experiments on the limitation of the densities of breeding birds. In general, densities can be limited by resources such as food or suitable nest sites or held at a lowered level by predators, parasites, or other natural enemies. Among a group of 18 experiments in which supplementary winter food was provided by the experimenter, 11 showed an increase in nesting-season densities compared with control areas. Four experiments in which the summer food of insect-eating forest birds was depleted by the use of insecticides resulted in no reduction in the density of nesting pairs. In a third group of experiments in which additional nest sites (boxes) were provided, density increased in 30 cases out of 32. Among experiments in which natural predators of the birds were removed, 14 out of 15 led to increased hatching success, 4 out of 8 to higher post-breeding numbers, and 6 out of 11 to increased breeding density. Taken together, the experiments confirmed that all major potential external limiting factors can affect breeding density of one bird species or another. They also confirmed that a particular species limited by one factor in certain years or areas may be limited by a different factor in other years or areas. It was well known that birds act as important dispersers of plant seeds by voiding not only the seeds of consumed fruit but also the remains of the fruit material, which has been converted into useful fertilizer. That a fruit has evolved to contain a laxative for speeding the seed through the bird's digestive system was revealed for the first time by Greg Murray of Hope College, Holland, Mich. He showed that the fruits of Witheringia solanacea, a Central American bush, pass quickly through the gut of the black-faced solitaire (Myadestes melanops) of Panama and Costa Rica and are thus more likely to germinate. Newly discovered bird species included the chestnut-bellied cotinga (Doliornis remseni), a thrush-sized fruit eater from the Andes of Ecuador, and the diademed tapaculo (Scytalopus schulenbergi), a small, secretive, fast-running bird of the cloud forest, which was discovered near La Paz, Bolivia, but later was shown to be common at 900 m (3,000 ft) altitude and above in Bolivia and neighbouring Peru. In a semideciduous Brazilian forest was found a previously unknown member of the Tyrannidae (the tyrant flycatcher family), which was named the Bahia tyrannulet (Phylloscartes beckeri). Brazil also yielded a new nighthawk, Chordeiles viellardi, a bird of the caatinga vegetation common in the state of Bahia. From Africa was reported a new nightjar (a close relative of the nighthawk, both groups being insect feeders active at dawn and dusk) dubbed the Nechisar nightjar (Caprimulgus solala). Nechisar is a plain in southern Ethiopia. The Indian Ocean revealed a new long-winged seabird, the Mascarene shearwater (Puffinus atrodorsalis). The monumental, nine-volume The Birds of the Western Palearctic (i.e., Europe, North Africa, and the Middle East), easily the most detailed reference to the birds of any major region of the Earth, was completed with publication of its last two volumes. The first volume had appeared in 1977. In total the series covered 770 species. (JEFFERY BOSWALL) This updates the article bird. Literature The 1995 Nobel Prize for Literature was awarded to the Irish poet Seamus Heaney. (See NOBEL PRIZES.) Heaney had moved his home from Northern Ireland, part of the United Kingdom, across the border to the Irish republic. Nevertheless, he remained perhaps the most respected and admired poet in the U.K., better known than any other poet of the flourishing Northern Irish school. His award was generally reckoned to be associated with the recent successes in furthering peace in the troubled island. Earlier in the year, John Redmond had written in the London Review of Books about the continuing achievements of Northern Irish poetry, observing that the poetry displayed "the kind of integrity and intertextuality which English poetry last had in the Thirties." He recognized that some attributed the phenomenon to "the concentrating pressure of 'The Troubles' "--that being the euphemistic term for the political violence that had, for so long, disfigured Ireland. Redmond held, however, that an equally important factor was "the symbolic coherence of Northern Irish poetry"--"poets as diverse as Heaney, Muldoon and Mahon are to a certain degree sustained by a single symbolic world." Asked by an interviewer about the Troubles, the prizewinning Heaney replied, "That's all over now." Some, however, remembered his poetic reference to an atrocity committed by the Irish rebels as the "tribal, intimate revenge" and held that Heaney was too tolerant of such activities. Wole Soyinka, the Nobel laureate of 1986, drew attention to the troubles of his own homeland, Nigeria, from which he was exiled. His play The Beatification of Area Boy had been banned by the Nigerian military government, and it received its world premiere in Britain at the West Yorkshire Playhouse. The production, a comedy about petty rogues and large-scale corruption in Nigeria, was broadcast by BBC radio. Soyinka, a strong opponent of several Nigerian governments since the nation gained independence in 1960, expressed his sympathy for another Nigerian playwright, his friend Ken Saro-Wiwa (see OBITUARIES), who with eight others was executed for alleged complicity in the murders of four chieftains in the oil-rich Ogoni territory. Saro-Wiwa had long been a leader of the Ogoni protest movement against the spoliation of the area by the oil company Shell and its friends in the government. Although many held him to be innocent of the murder charges, it was his status as a writer that seemed to stimulate international opinion against his execution. At a meeting of Commonwealth heads of government, Nigeria was formally suspended from membership. Saro-Wiwa's best-known novel was Sozaboy ("Soldier Boy"), a satire on the military government, and he also developed the highly popular television series Basi and Company, which was shown on British television. Doubts about Saro-Wiwa's innocence were expressed publicly by a few Britons, including Auberon Waugh, the editor of Literary Review. In France the Prix Goncourt was awarded to Andre Makine, a 38-year-old Russian novelist living in Paris. His latest novel, Le Testament franais, concerned a boy trapped between the cultures of France and Russia. It seemed ironic that Makine's application for French citizenship had recently been rejected. It was generally expected, however, that Makine's success, not only in winning the Goncourt but also in sharing the Prix Mdicis, would induce the immigration authorities to review their unfavourable decision. United Kingdom. The 50th anniversary of the conclusion to World War II, though well marked by public ceremonies, attracted less attention in the world of literature and publishing than might have been expected. The Allied victory, suggested Hew Strachan in the Times Literary Supplement, had been overshadowed by the collapse of one of those victorious allies, the Soviet Union; "Thus the notion of the 'short' twentieth century, begun in 1914 and concluded in 1989, diminishes the importance of 1945." Strachan was reviewing The Oxford Companion to the Second World War, edited by I.C.B. Dear and M.R.D. Foot, which he described as "an outstanding guide, as sensible and cogent on the big questions as it is instructive and informed on the lesser ones." There were, of course, other compendious new accounts of the war. Martin Gilbert's The Day the War Ended was reviewed without enthusiasm by Richard Overy, who himself published a book called Why the Allies Won. Armageddon: The Second World War, by Clive Ponting was also received with disfavour in the Times Literary Supplement on the grounds that its studied objectivity looked too much like a discreditable neutrality. Such arguments seemed rather narrow and esoteric to the general public. It was interesting to note that Strachan was engaged in writing a history of World War I, for there were indications that 1914-18 had become as fascinating a period for the general reader as 1939-45. The U.K. publishing world was "convulsed," according to David Sexton in the Sunday Telegraph, by the collapse of the price-fixing mechanism known as the Net Book Agreement. Best-sellers were discounted in the shops, and there was a general fear that more ambitious books, with less commercial appeal, would become more difficult to publish. The apparently pleasurable prospect of cheaper books was seen to be accompanied by unexpected disadvantages. The rumours of a revival of interest in poetry proved to be greatly exaggerated, despite a notable increase in the sales of Heaney's work after he won the Nobel Prize. The bicentenary of the birth of John Keats was marked by several essays in journals and numerous radio broadcasts, but it drew little attention from book publishers. The poet laureate, Ted Hughes, wrote an admiring poem about the Queen Mother on her 95th birthday, comparing her to a six-rooted oak tree, but his verse was received in a mocking spirit. One poet who was accorded serious attention was Robert Graves (1895-1985), a veteran of World War I, a mythopoeic fantasist, and a historical novelist as well as a pure-voiced poet of erotic love. His long, strange life, with his wives and his lovers, was recorded once more in a new biography by Miranda Seymour, while his nephew, R.P. Graves, proffered the third volume of his own biography--Robert Graves and the White Goddess, 1940-85. An earlier biography by Martin Seymour-Smith also reappeared in a new edition. The Carcanet Press began a 21-volume reprint program of the poet's work, starting with the first volume of Collected Poems, Complete Short Stories, and Collected Writings on Poetry. The press also offered The Centenary Selected Poems, which was found too limited by Neil Powell in the Times Literary Supplement, who called it "rather perverse" in omitting many admired poems. Perplexed by Graves's life, Powell remained impressed by his verse. Mark Ford, on the other hand, writing in the London Review of Books, was inclined to dismiss Graves's claim to universal significance and to see his poems only as "symptoms of his personal problems." The death of Kingsley Amis (see OBITUARIES), shortly after the publication of his latest novel, The Biographer's Moustache, seemed to mark the end of a significant genre of British fiction, the novel of snobbery. Amis had been the most accomplished writer of these class-conscious comedies since Evelyn Waugh, as interested as Nancy Mitford in the ways a choice of words and their pronunciation could be used to distinguish between "common" people and those thought to be "posh." Such distinctions, made to seem very old-fashioned, lingered on, quite credibly, in The Biographer's Moustache, which told of a rather charming elderly novelist confronted by an ambitious young biographer, somewhat dubious about the novelist's worth. The biographer, clever and common, seemed to resemble Amis in his youth, while the novelist, posh and snobbish, reflected certain apparent characteristics of the older Amis. Observant, subtle, and comical, the novel concluded straightforwardly, with a modern girl saying, "Oh, they're really there, all those distinctions are, but . . . it isn't class differences that keep people apart, it's thinking they bloody matter." This might be read as Amis' apologia. Amis had been a previous winner of the Booker Prize, but his novel did not appear on the shortlist for 1995. Nor did new novels by seven other previous winners, including Penelope Fitzgerald, whose historical novel about the German poet Novalis, The Blue Flower, had been much admired. Also omitted, to the surprise of many, was the new novel by Amis' son, Martin Amis, a writer whose career, marital situation, and dealings with publishers attracted great interest among journalists. His book The Information, another study of a conflict between two writers, did not appeal to the Booker judges, however. The Booker candidate most generally favoured was Salman Rushdie, with a new novel about the history of an Indian family in Bombay from the last days of the British Empire to the 1970s. The Moor's Last Sigh was an extravagant saga--"a triumph of un-naturalism and a feast for anyone with a strong literary digestion," according to Victoria Glendinning in the Daily Telegraph. Though it did not win, Rushdie's novel was nominated for the Whitbread Book of the Year Award, and it was the winner in the fiction section. Barry Unsworth--like Rushdie a previous Booker Prize winner--was another strong contender with a historical novel, Morality Play, about a priest in 14th-century England who joins a company of traveling actors; it develops into a sort of detective story, its sombre realism vitiated by rather heavy moralizing. Justin Cartwright, born in South Africa, was nominated for his sour novel set in London, In Every Face I Meet, concerning a failing businessman, who was bred in Africa, and his dealings with a young London prostitute. "Deeply depressing," commented Patrick Gale, "as though Kingsley Amis had turned his hand to tragedy." A fourth contender, from Australia, was Tim Winton (see BIOGRAPHIES), who published The Riders, an eerie but perhaps sentimental tale of an Australian and his small daughter searching throughout Europe for a missing wife and mother. In the London Review of Books, Jonathan Coe called it a "bruising, exultant novel." The winner of the Booker Prize, however, was a woman writing about men at war. The Ghost Road was the third volume of Pat Barker's trilogy of World War I. In the book she dealt with the psychological traumas of the ex-combatants and the attempts to heal them. Barker's grasp of military systems and her understanding of the period were much admired. The chairman of the Booker Prize judges, however, after reading 141 new novels, remarked despondently, "Our art hankers after the past. Very few people write with any conviction about the present." This impression of a fin de sicle world yearning for the past did not receive much support from the year's crop of biographies. There were several lives of writers who had clearly lost their old appeal. Three modern playwrights--Terence Rattigan, William Douglas Home, and Dennis Potter--were rather defensively appreciated. Before Amis' novel The Biographer's Moustache was published, a biography of the novelist appeared, written by his genial friend Eric Jacobs. D.J. Taylor, writing in the Times Literary Supplement, took strong objection to the book and to its subject: "It would be a brave man who suggested that the life outlined here was particularly edifying or attractive." Margaret Drabble attempted to revive the reputation of a suddenly neglected novelist in her biography of Angus Wilson. "The conflict between Wilson's generous humanity and his apparently selfish delight in extravagant behaviour, in the crazy crowd, is a persistent theme of this large and satisfying biography," wrote Frank Kermode in the London Review of Books. He admired Drabble's handling of "the theme of male homosexual social relations" and noted that "there was a freakishness, a habit of clowning, an ebullience that was to become an ingredient of Wilson's huge but, as it turned out, transient popular success." It seemed that Wilson's reputation would not be revived. The detective story writers Arthur Conan Doyle and Dorothy L. Sayers each received another appraisal, as did that biographers' favourite Lewis Carroll (Charles Lutwidge Dodgson). More contentious perhaps was Ian MacKillop's biography F.R. Leavis: A Life in Criticism. The once-authoritative scholar and critic, born a century earlier, was gravely appreciated, his pugnacity and his sense of persecution comprehended. "Notwithstanding MacKillop's avowedly personal attachment to his subject," wrote Dan Jacobson in the Times Literary Supplement, "he does as much as anyone could to be fair-minded alike to friends, enemies and friends-late-revealed-as-enemies." Biographies of an encomiastic sort were published by two senior politicians with once-powerful reputations in the Labour Party. One was Roy Jenkins, who had broken with that party to become cofounder, in 1981, of the Social Democrats; he offered a new life of the scholarly Victorian statesman W.E. Gladstone, who broke with the Conservatives to become a Liberal prime minister. Jenkins was already recognized as an accomplished political biographer, and it was evident that his own ministerial experience had been of value to him in writing the book. The work was much admired, and it was the winner of the biographical section of the Whitbread Award. The other senior statesman-biographer was Michael Foot, former leader of the Labour Party, who further strengthened his literary credentials with H.G.: The History of Mr. Wells. Foot had known H.G. Wells and supported him in many of his political and social campaigns, and the biographer appeared as an advocate for Wells's utopian objectives and achievements. For those readers who had come to think of Wells primarily as a science-fiction romancer and a brilliant comic novelist, Foot's use of long quotations from his half-forgotten pamphlets and fiction supported the appreciation of the literary merits and modern relevance of this prolific author. (D.A.N. JONES)