{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T06:00:53Z","timestamp":1792044053932,"version":"4.4.0"},"reference-count":51,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2017,3,28]],"date-time":"2017-03-28T00:00:00Z","timestamp":1490659200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"funder":[{"DOI":"10.13039\/100005835","name":"AMNH","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100005835","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Journal of Morphology"],"published-print":{"date-parts":[[2017,6]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Miniaturization, the evolution of extremely small adult body size, is widespread amongst animals and commonly associated with novel ecological, physiological, and morphological attributes. The phenotypes of miniaturized taxa are noteworthy because they combine reductions and structural simplifications with novel traits not developed in their larger relatives. Previous research on miniature cyprinid fishes (focused predominantly on South and South East Asian taxa of a single subfamily) has identified two distinct classes of miniature taxa: proportioned dwarves and developmentally truncated miniatures. Miniaturization has also occurred independently in the subfamily Cyprininae, particularly in African lineages. We investigate the skeletal anatomy of <jats:italic>Barboides<\/jats:italic>, a genus of miniature African cyprinids that includes Africa's smallest known species of vertebrates, to assess whether miniaturization has resulted in similar organismal outcomes in different lineages of the Cyprinidae. The skeleton of <jats:italic>Barboides<\/jats:italic> is characterized by the complete absence of a number of dermal and endochondral ossifications, and marked reduction in size and\/or complexity of other skeletal elements, particularly those of the dermatocranium. Absent skeletal elements in <jats:italic>Barboides<\/jats:italic> include those which develop relatively late in the ossification sequence of the non\u2010miniature African relative \u2018<jats:italic>Barbus<\/jats:italic>\u2019 <jats:italic>holotaenia<\/jats:italic> suggesting that their absence in <jats:italic>Barboides<\/jats:italic> can be explained by a simple scenario of developmental truncation. In contrast to this theme of loss and reduction, the os suspensorium of <jats:italic>Barboides<\/jats:italic> is enlarged and the outer arm distally trifid and associated with a novel bulbous muscle in males. An evaluation of the skeleton of <jats:italic>Barboides<\/jats:italic> provides further evidence for a link between developmental truncation and evolutionary morphological novelty in Cyprinidae. In the spectrum of miniature cyprinids ranging from proportioned dwarves with few bones missing to highly progenetic taxa with dozens of missing bones, the two species of <jats:italic>Barboides<\/jats:italic> range roughly in the middle showing that the extremes are connected by intermediate levels of truncatedness.<\/jats:p>","DOI":"10.1002\/jmor.20670","type":"journal-article","created":{"date-parts":[[2017,3,29]],"date-time":"2017-03-29T03:22:47Z","timestamp":1490757767000},"page":"750-767","source":"Crossref","is-referenced-by-count":14,"title":["Morphological novelty and modest developmental truncation in <i>Barboides<\/i>, Africa's smallest vertebrates (Teleostei: Cyprinidae)"],"prefix":"10.1002","volume":"278","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0161-8385","authenticated-orcid":false,"given":"Kevin W.","family":"Conway","sequence":"first","affiliation":[{"name":"Department of Wildlife and Fisheries Sciences and Biodiversity Research and Teaching Collections Texas A&amp;M University College Station Texas 77843 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kole M.","family":"Kubicek","sequence":"additional","affiliation":[{"name":"Department of Wildlife and Fisheries Sciences and Biodiversity Research and Teaching Collections Texas A&amp;M University College Station Texas 77843 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ralf","family":"Britz","sequence":"additional","affiliation":[{"name":"Department of Life Sciences Natural History Museum Cromwell Road London SW7 5BD United Kingdom"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2017,3,28]]},"reference":[{"key":"e_1_2_7_2_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.part.18603"},{"key":"e_1_2_7_3_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.1052080102"},{"key":"e_1_2_7_4_1","first-page":"5","article-title":"Systematics of Barbus: State of the art and heuristic comments","volume":"45","author":"Berrebi P.","year":"1996","journal-title":"Folia Zoologica"},{"key":"e_1_2_7_5_1","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.10387"},{"key":"e_1_2_7_6_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1095-8649.2012.03307.x"},{"key":"e_1_2_7_7_1","doi-asserted-by":"crossref","first-page":"53","DOI":"10.11646\/zootaxa.2277.1.4","article-title":"Danionella priapus, a new species of miniature cyprinid fish from West Bengal, India (Teleostei: Cypriniformes: Cyprinidae)","volume":"2277","author":"Britz R.","year":"2009","journal-title":"Zootaxa"},{"key":"e_1_2_7_8_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10698"},{"key":"e_1_2_7_9_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.20486"},{"key":"e_1_2_7_10_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2009.0141"},{"key":"e_1_2_7_11_1","doi-asserted-by":"publisher","DOI":"10.1111\/zoj.12184"},{"key":"e_1_2_7_12_1","first-page":"415","article-title":"Paedocypris carbunculus, a new species of miniature fish from Borneo (Teleostei: Cypriniformes: Cyprinidae)","volume":"56","author":"Britz R.","year":"2008","journal-title":"The Raffles Bulletin of Zoology"},{"key":"e_1_2_7_13_1","first-page":"327","article-title":"Fangfangia spinicleithralis, a new genus and species of miniature cyprinid fish from the peat swamp forests of Borneo (Teleostei: Cyprinidae)","volume":"22","author":"Britz R.","year":"2012","journal-title":"Ichthyological Exploration of Freshwaters"},{"key":"e_1_2_7_14_1","first-page":"249","article-title":"Monophyly of the genus Boraras (Teleostei: Cyprinidae)","volume":"16","author":"Conway K. W.","year":"2005","journal-title":"Ichthyological Exploration of Freshwaters"},{"key":"e_1_2_7_15_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.2011.00698.x"},{"key":"e_1_2_7_16_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1095-8649.2007.01646.x"},{"key":"e_1_2_7_17_1","first-page":"251","article-title":"Review of the South East Asian miniature cyprinid genus Sundadanio (Ostariophysi: Cyprinidae) with descriptions of seven new species from Indonesia and Malaysia","volume":"22","author":"Conway K. W.","year":"2011","journal-title":"Ichthyological Exploration of Freshwaters"},{"key":"e_1_2_7_18_1","first-page":"73","article-title":"Barboides britzi, a new species of miniature cyprinid from Benin (Ostariophysi: Cyprinidae), with a neotype designation for B. gracilis","volume":"17","author":"Conway K. W.","year":"2006","journal-title":"Ichthyological Exploration of Freshwaters"},{"key":"e_1_2_7_19_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10228-007-0030-1"},{"key":"e_1_2_7_20_1","first-page":"400","article-title":"Supplement on some new species of American and African fishes","volume":"13","author":"Cope E. D.","year":"1867","journal-title":"Transactions of the American Philosophical Society"},{"key":"e_1_2_7_21_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1097-4687(199608)229:2<121::AID-JMOR1>3.0.CO;2-4"},{"key":"e_1_2_7_22_1","doi-asserted-by":"crossref","first-page":"37","DOI":"10.11646\/zootaxa.2571.1.2","article-title":"New species of Microhyla from Sarawak: Old World's smallest frogs crawl out of miniature pitcher plants on Borneo (Amphibia: Anura: Microhylidae)","volume":"2571","author":"Das I.","year":"2010","journal-title":"Zootaxa"},{"key":"e_1_2_7_23_1","volume-title":"The development of the vertebrate skull","author":"Beer G. R. (","year":"1930"},{"key":"e_1_2_7_24_1","unstructured":"Eschmeyer W. N. &Fong J. D. (2017).Species by family\/subfamily in the Catalog of Fishes. Retrieved fromhttp:\/\/researcharchive.calacademy.org\/research\/ichthyology\/catalog\/SpeciesByFamily.asp"},{"key":"e_1_2_7_25_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1981.tb01575.x"},{"key":"e_1_2_7_26_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.es.24.110193.002441"},{"key":"e_1_2_7_27_1","doi-asserted-by":"publisher","DOI":"10.2307\/1447174"},{"key":"e_1_2_7_28_1","doi-asserted-by":"publisher","DOI":"10.2307\/1447427"},{"key":"e_1_2_7_29_1","first-page":"1","article-title":"Anatomy and phylogeny of the Chinese Major Carps Ctenopharyngodon Steind., 1866 and Hypophalmichthys Blkr., 1860","volume":"41","author":"Howes G. J.","year":"1981","journal-title":"Bulletin of the British Museum (Natural History). Zoology"},{"key":"e_1_2_7_30_1","doi-asserted-by":"publisher","DOI":"10.3998\/mpub.17658"},{"key":"e_1_2_7_31_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2005.3419"},{"key":"e_1_2_7_32_1","first-page":"161","article-title":"Boraras micros, a new genus and species of minute freshwater fish from Thailand (Teleostei: Cyprinidae)","volume":"4","author":"Kottelat M.","year":"1993","journal-title":"Ichthyological Exploration of Freshwaters"},{"key":"e_1_2_7_33_1","doi-asserted-by":"publisher","DOI":"10.3897\/zookeys.154.1963"},{"key":"e_1_2_7_34_1","doi-asserted-by":"publisher","DOI":"10.1111\/azo.12122"},{"key":"e_1_2_7_35_1","first-page":"1","article-title":"Two new species of Barbus (Pisces: Cyprinidae) from southeastern Dahomey","volume":"83","author":"Loiselle P. V.","year":"1971","journal-title":"Revue de Zoologie et Botanique Africaine"},{"key":"e_1_2_7_36_1","first-page":"103","article-title":"Skeletal development and ossification sequence of the characiform Salminus brasiliensis (Ostariophysi: Characidae)","volume":"25","author":"Mattox G. M.","year":"2014","journal-title":"Ichthyological Exploration of Freshwaters"},{"key":"e_1_2_7_37_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.20477"},{"key":"e_1_2_7_38_1","first-page":"27","article-title":"Trends in the evolution of teleostean fishes","volume":"7","author":"Myers G. S.","year":"1958","journal-title":"Stanford Ichthyological Bulletin"},{"key":"e_1_2_7_39_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.2694"},{"key":"e_1_2_7_40_1","first-page":"77","volume-title":"Problems in vertebrate evolution","author":"Patterson C. (","year":"1977"},{"key":"e_1_2_7_41_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0029797"},{"key":"e_1_2_7_42_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00842977"},{"key":"e_1_2_7_43_1","first-page":"131","article-title":"The \u201ccelestial pearl danio\u201d, a new genus and species of colourful minute cyprinid fish from Myanmar (Pisces: Cypriniformes)","volume":"55","author":"Roberts T. R.","year":"2007","journal-title":"Raffles Bulletin of Zoology"},{"key":"e_1_2_7_44_1","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2148-7-38"},{"key":"e_1_2_7_45_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.200900116"},{"key":"e_1_2_7_46_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2015.05.025"},{"key":"e_1_2_7_47_1","first-page":"1","article-title":"Study of the dorsal gill\u2010arch musculature of teleostome fishes with special reference to the Actinopterygii","author":"Springer V. G.","year":"2004","journal-title":"Bulletin of the Biological Society of Washington, 11"},{"key":"e_1_2_7_48_1","first-page":"107","article-title":"Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study","volume":"9","author":"Taylor W. R.","year":"1985","journal-title":"Cybium"},{"key":"e_1_2_7_49_1","first-page":"132","article-title":"Description of a new genus and species for a small cyprinid fish from southern Cameroon","volume":"83","author":"Thys van den Audenaerde D. F. E.","year":"1971","journal-title":"Revue de Zoologie et de Botanique Africaines"},{"key":"e_1_2_7_50_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1095-8649.1992.tb03169.x"},{"key":"e_1_2_7_51_1","first-page":"444","article-title":"Miniaturization in South American freshwater fishes; an overview and discussion","volume":"101","author":"Weitzman S. H.","year":"1988","journal-title":"Proceedings of the Biological Society of Washington"},{"key":"e_1_2_7_52_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2015.01.014"}],"container-title":["Journal of Morphology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Fjmor.20670","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.20670","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,10,3]],"date-time":"2023-10-03T19:17:39Z","timestamp":1696360659000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/jmor.20670"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,3,28]]},"references-count":51,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2017,6]]}},"alternative-id":["10.1002\/jmor.20670"],"URL":"https:\/\/doi.org\/10.1002\/jmor.20670","archive":["Portico"],"relation":{},"ISSN":["0362-2525","1097-4687"],"issn-type":[{"value":"0362-2525","type":"print"},{"value":"1097-4687","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,3,28]]}}}