{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T09:13:11Z","timestamp":1775725991874,"version":"3.50.1"},"reference-count":45,"publisher":"American Association for the Advancement of Science (AAAS)","issue":"5376","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Science"],"published-print":{"date-parts":[[1998,7,24]]},"abstract":"<jats:p>\n            The phylogeny of the Phytophaga, the largest and oldest radiation of herbivorous beetles, was reconstructed from 115 complete DNA sequences for the 18\n            <jats:italic>S<\/jats:italic>\n            nuclear ribosomal subunit and from 212 morphological characters. The results of these analyses were used to interpret the role of angiosperms in beetle diversification. Jurassic fossils represent basal lineages that are still associated with conifers and cycads. Repeated origins of angiosperm-feeding beetle lineages are associated with enhanced rates of beetle diversification, indicating a series of adaptive radiations. Collectively, these radiations represent nearly half of the species in the order Coleoptera and a similar proportion of herbivorous insect species.\n          <\/jats:p>","DOI":"10.1126\/science.281.5376.555","type":"journal-article","created":{"date-parts":[[2002,7,27]],"date-time":"2002-07-27T09:43:20Z","timestamp":1027763000000},"page":"555-559","source":"Crossref","is-referenced-by-count":668,"title":["\"Inordinate Fondness\" Explained: Why Are There So Many Beetles?"],"prefix":"10.1126","volume":"281","author":[{"given":"Brian D.","family":"Farrell","sequence":"first","affiliation":[{"name":"Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA."}]}],"member":"221","reference":[{"key":"e_1_3_1_2_2","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1086\/282070","volume":"93","author":"Hutchinson G. E.","year":"1959","unstructured":"Hutchinson G. E., Am. Nat. 93, 145 (1959);","journal-title":"Am. Nat."},{"key":"e_1_3_1_2_3","unstructured":". Haldane himself often repeated this quip although the circumstances and precise wording of the original remark have been controversial [see the summary of recent exchanges by S. J. Gould Nat. Hist. 1 4 (1993)]."},{"key":"e_1_3_1_3_2","doi-asserted-by":"crossref","unstructured":"N. Stork Biol. J. Linn. Soc. 35 321 (1988).","DOI":"10.1111\/j.1095-8312.1988.tb00474.x"},{"key":"e_1_3_1_4_2","unstructured":"D. R. Strong J. H. Lawton T. R. E. Southwood Insects on Plants (Harvard Univ. Press Cambridge MA 1984)."},{"key":"e_1_3_1_5_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.11536548"},{"key":"e_1_3_1_6_2","unstructured":"F. M. Carpenter Treatise on Invertebrate Paleontology Part R of Arthropoda vol. 4 of Superclass Hexapoda (Geological Society of America Boulder CO 1992)."},{"key":"e_1_3_1_7_2","unstructured":"J. F. Lawrence in Synopsis and Classification of Living Organisms S. P. Parker Ed. (McGraw-Hill New York 1982) vol. 2 pp. 482\u2013553."},{"key":"e_1_3_1_8_2","doi-asserted-by":"crossref","unstructured":"R. T. Thompson J. Nat. Hist. 26 835 (1992).","DOI":"10.1080\/00222939200770511"},{"key":"e_1_3_1_9_2","unstructured":"P. Jolivet and T. J. Hawkeswood Host-Plants of Chrysomelidae of the World (Backhuys Leiden Netherlands 1995)."},{"key":"e_1_3_1_10_2","unstructured":"V. G. Gratshev and V. V. Zherikhin Paleontol. J. 29 112 (1995)."},{"key":"e_1_3_1_11_2","unstructured":"L. V. Arnoldi'i V. V. Zherikhin L. M. Nikritin A. G. Ponomorenko Mesozoic Coleoptera (Oxonian Press New Delhi India 1991)."},{"key":"e_1_3_1_12_2","unstructured":"Beetle groups that are restricted to particular higher plant taxa were scored for the least inclusive plant taxon that contained their hosts. The cerambycid subfamilies Prioninae and Parandrinae were left unscored for host as these do not feed on live plant tissues but on dead or decaying wood [E. G. Linsley Univ. Calif. Berkeley Publ. Entomol. 18 1 (1961)]. The only higher beetle taxa for which DNA sequences were not obtainable were the Sagrinae and Anthribidae."},{"key":"e_1_3_1_13_2","unstructured":"G. Kuschel Mem. Entomol. Soc. Wash. 14 5 (1995)."},{"key":"e_1_3_1_14_2","unstructured":"C. A. M. Reid in Biology and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson J. Pakaluk and S. A. Slipinski Eds. (Muzeum I Instytut Zoologi PAN Warsaw 1995) pp. 559\u2013631."},{"key":"e_1_3_1_15_2","unstructured":"Sequences were obtained for the complete 18 S ribosomal subunit gene from 115 of these beetle taxa and from the outgroup species from the Tenebrionidae ( Tenebrio molitor GenBank accession number 70810) Melyridae ( Collops quadrimaculatus ) and Erotylidae ( Cypherotylus boisduvali ) with methods that were given by"},{"key":"e_1_3_1_15_3","first-page":"1","volume":"46","author":"Whiting M. F.","year":"1997","unstructured":"Whiting M. F., Carpenter J. C., Wheeler Q. D., Wheeler W. C., Syst. Biol. 46:, 1 (1997);","journal-title":"Syst. Biol."},{"key":"#cr-split#-e_1_3_1_15_4.1","unstructured":". These sequences were aligned using Sequencher 3.0 (Gene Codes Corporation Ann Arbor MI 1995) producing a matrix of 2117 positions. Three \u223c40-base pair"},{"key":"#cr-split#-e_1_3_1_15_4.2","unstructured":"(bp) hypervariable regions could not be unambiguously aligned and were excluded from the analyses as were the two 50-bp ends of the gene to avoid excessive missing data in parts of the matrix. The remaining 1874 positions yielded 355 potentially informative characters. These characters were analyzed separately and together with the morphological matrix compiled from Kuschel and Reid ( 12 13 ). Analyses using the program PAUP* 4.0 version d59 included 100 initial heuristic searches using random taxon addition sequences and tree bisection-reconnection (TBR) branch swapping setting MAXTREES (maximum number of trees held in memory) to 200 and keeping two trees per replicate search. This set of 200 trees was then subjected to TBR branch swapping with MAXTREES set to 10 000. Bootstrap analysis used 1000 random taxon addition sequences with branch swapping limited to 100 trees per replicate. Tests of incongruence (using simple addition sequences and limiting MAXTREES to 100) between morphological and molecular data sets were not significant (incongruence length difference P > 0.5)."},{"key":"e_1_3_1_16_2","unstructured":"The Rhinorhynchinae subfamily includes the most morphologically plesiomorphic nemonychids and they currently consist of 14 genera associated with strobili of Araucariaceae or Podocarpaceae in Chile Argentina and Australia plus a single species living on Pinaceae in Colorado [G. Kuschel Rev. Chil. Hist. Nat. 54 97 (1954)]. The closely related Holarctic Doydirhynchinae comprise 19 species living on Pinaceae. Crowson removed the nominate genus Nemonyx to the Anthribidae [R. A. Crowson Entomol. Mon. Mag. 121 144 (1985)]."},{"key":"e_1_3_1_17_2","unstructured":"The Palophaginae consist of three species in two genera which develop in the male strobili of Araucariaceae in Chile Argentina Australia and New Zealand [G. Kuschel and B. M. May N. Z. Entomol. 19 1 (1996)]."},{"key":"e_1_3_1_18_2","unstructured":"The most plesiomorphic oxycorynine belid genus Oxycraspedus attacks Araucaria strobili in Chile and Argentina [G. Kuschel Invest. Zool. Chil. 5 229 (1959)]. Crowson also suggested that Oxycraspedus and Rhopalotria are sister taxa but did not place the morphologically disparate oxycorynine genera reported from the Hydnoraceae and Balanophoraceae which are families of tree parasites [R. A. Crowson in Advances in Coleopterology M. Zunino X. Belles M. Blas Eds. (European Association of Coleopterology Barcelona 1991) pp. 13\u201328]. The belid tribe Pachyurini comprises 13 genera associated with Araucaria and Agathis in Australia and New Zealand and a single genus associated with Podocarpaceae and Cupressaceae in Brazil."},{"key":"e_1_3_1_19_2","unstructured":"The Orsodacninae comprise the Australian genus Cucujopsis which is associated with the male strobili of the araucariaceous genus Agathis and the Holarctic genus Orsodacne [J. S. Mann and R. A. Crowson J. Nat. Hist. 15 727 (1981)]. Although the larval affiliations of Orsodacne are still unconfirmed these are probably in the male strobili of Pinaceae (with which all eight species co-occur) a resource available during the early spring flights of the pollen-feeding adults."},{"key":"e_1_3_1_20_2","unstructured":"The belid subfamily Allocoryninae comprises >20 species in the Neotropical genus Rhopalotria which attack the male strobili of Zamia and Dioon."},{"key":"e_1_3_1_21_2","unstructured":"The chrysomelid subfamily Aulacoscelidinae comprises 18 species in two Neotropical genera restricted to the Cycadaceae."},{"key":"e_1_3_1_22_2","first-page":"496","volume":"19","author":"Brundin L.","year":"1965","unstructured":"Brundin L., Evolution 19, 496 (1965).","journal-title":"Evolution"},{"key":"e_1_3_1_23_2","doi-asserted-by":"crossref","unstructured":"G. Kuschel in Australian Weevils E. Zimmerman Ed. [Commonwealth Scientific and Industrial Research Organization (CSIRO) Melbourne Australia 1994] p. 569. Other nemonychids in the Karatau Formation apparently belong to the now-extinct subfamily Brenthorrhininae ( 9 ). The Nemonychidae are also represented by Libanorhinus succinus in Lower Cretaceous amber derived from Araucariaceae resins [G. Kuschel and G. O. Poinar Entomol. Scand. (Group 2) 24 143 (1993)] and by the Lower Cretaceous Slonik in the central Asian trans-Baikal deposits [G. Kuschel GeoJournal 7 499 (1983)].","DOI":"10.1163\/187631293X00253"},{"key":"e_1_3_1_24_2","unstructured":"The oxycorynine Archeorrhynchus paradoxopus (Belidae) is found in the Karatau Formation [G. Kuschel in Australian Weevils E. Zimmerman Ed. (CSIRO Melbourne Australia 1994) p. 244]. Oxycoryninae are also represented in the Lower Cretaceous Santana Formation of Brazil [D. A. Grimaldi Ed. Bull. Am. Mus. Nat. Hist. 195 8 (1990)]. Additional Karatau belids include the extinct subfamily Eobelinae [V. V. Zherikhin and V. G. Gratshev in Biology and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson J. Pakaluk and S. A. Slipinski Eds. (Muzeum I Instytut Zoologi PAN Warsaw 1995) p. 646]."},{"key":"#cr-split#-e_1_3_1_25_2.1","unstructured":"The belid subfamily Carinae which attacks strobili of the coniferous Cupressaceae occurs in the Jurassic Karatau beds as represented by Eccoptarthrus and Emanrhynchus [V. V. Zherikhin and V. G. Gratshev in Biology and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson J. Pakaluk and S. A. Slipinski Eds. (Muzeum I Instytut Zoologi PAN Warsaw 1995) pp. 634-777]. The Carinae also appear in the Lower Cretaceous trans-Baikal beds ( Cretonanophyes and Baissorhynchus)"},{"key":"#cr-split#-e_1_3_1_25_2.2","unstructured":"the Carinae presently contains Car which is found in Australia and Tasmania and Chilecar and Caenominurus which are found in Chile and Argentina [E. Zimmerman Ed. Australian Weevils (CSIRO Melbourne Australia 1994) p. 504."},{"key":"e_1_3_1_26_2","unstructured":"The chrysomelid Cerambyomima longicornis attributed to the Aulacoscelinae [G. Kuschel and B. M. May Invertebr. Taxon. 3 697 (1993)] resembles the orsodacnine Cucujopsis in the grooved frons and may be an intermediate form."},{"key":"e_1_3_1_27_2","unstructured":"Jurassic fossil cones of Araucaria mirabilis from Argentina closely resemble A. bidwellii and show damage similar to that caused by weevil larvae [see R. A. Stockey Paleontographica 166 1 (1978)]. A. bidwellii is host to extant species in both the Nemonychidae and Palophaginae."},{"key":"e_1_3_1_28_2","doi-asserted-by":"crossref","unstructured":"R. A. Stockey J. Plant Res. 107 493 (1994).","DOI":"10.1007\/BF02344070"},{"key":"e_1_3_1_29_2","doi-asserted-by":"crossref","first-page":"881","DOI":"10.1086\/285258","volume":"138","author":"Farrell B. D.","year":"1991","unstructured":"Farrell B. D., Dussourd D., Mitter C., Am. Nat. 138, 881 (1991).","journal-title":"Am. Nat."},{"key":"e_1_3_1_30_2","unstructured":"The allocorynine Scelocamptus curvipes is found in the Karatau beds ( 10 )."},{"key":"e_1_3_1_31_2","unstructured":"The aulacosceline genera Protoscelis Protosceloides and Pseudomegamerus are found in the Karatau beds ( 5 )."},{"key":"e_1_3_1_32_2","unstructured":"T. N. Taylor and E. L. Taylor The Biology and Evolution of Fossil Plants (Prentice-Hall Englewood Cliffs NJ 1993)."},{"key":"e_1_3_1_33_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1558-5646.1964.tb01674.x"},{"key":"e_1_3_1_33_3","unstructured":"; B. D. Farrell and C. Mitter Biol. J. Linn. Soc. 68 533 (1998)."},{"key":"e_1_3_1_34_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.274.5292.1489"},{"key":"e_1_3_1_35_2","unstructured":"Assignments of feeding habits and numbers of recent genera are from Lawrence ( 6 )."},{"key":"e_1_3_1_36_2","unstructured":"The number of genera was extracted from the totals per beetle family in Lawrence ( 6 )."},{"key":"e_1_3_1_37_2","unstructured":"Estimates of diversity are from the following sources: Curculionoidea ( 7 ); Chrysomelidae [P. Jolivet E. Petitpierre T. H. Hsiao Eds. Biology of Chrysomelidae (Kluwer Academic Dordrecht Netherlands 1988)]; Cerambycidae [S. Bily and O. Mehl Longhorn Beetles (Coleoptera Cerambycidae) of Fennoscandia and Denmark vol. 22 of Fauna Entomologica of Scandinavica (Brill Leiden Netherlands 1989)]."},{"key":"e_1_3_1_38_2","unstructured":"For a discussion of the use of fossils to assign character optimizations see"},{"key":"e_1_3_1_38_3","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/0034-6667(87)90040-6","volume":"50","author":"Doyle J. M.","year":"1987","unstructured":"Doyle J. M., Donoghue M. J., Rev. Palaeobot. Palynol. 50, 63 (1987).","journal-title":"Rev. Palaeobot. Palynol."},{"key":"e_1_3_1_39_2","unstructured":"For supplying specimens or identifications of key or austral taxa I especially thank F. Andrews J. Chemsak L. Diego-Gomez J. Donaldson C. Duckett T. Erwin W. Flowers D. Furth C. D. Johnson J. Kingsolver G. Kuschel J. Lawrence A. Newton K. Norstog R. Oberprieler C. O'Brien and E. G. Riley among many others. I also thank A. Salmore M. Blair and L. Morrissey for technical lab support; A. Berry M. Donoghue D. Futuyma A. Knoll D. Lewontin E. Mayr C. Mitter N. Moran B. Normark S. Palumbi N. Pierce and E. O. Wilson for helpful discussions; and A. Knoll C. Labandeira and D. Maddison for detailed comments on a late draft. This research was supported by NSF USDA and the Putnam Expedition Fund of the Museum of Comparative Zoology."}],"container-title":["Science"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.science.org\/doi\/pdf\/10.1126\/science.281.5376.555","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,1,13]],"date-time":"2024-01-13T03:20:28Z","timestamp":1705116028000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.science.org\/doi\/10.1126\/science.281.5376.555"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[1998,7,24]]},"references-count":45,"journal-issue":{"issue":"5376","published-print":{"date-parts":[[1998,7,24]]}},"alternative-id":["10.1126\/science.281.5376.555"],"URL":"https:\/\/doi.org\/10.1126\/science.281.5376.555","relation":{},"ISSN":["0036-8075","1095-9203"],"issn-type":[{"value":"0036-8075","type":"print"},{"value":"1095-9203","type":"electronic"}],"subject":[],"published":{"date-parts":[[1998,7,24]]}}}