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Taxonomy, biogeography and evolution of plants
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RESEARCH ARTICLE
Contents Vol 18(1)

Generic concepts within hornworts: historical review, contemporary insights and future directions

D. Christine Cargill A D , Karen S. Renzaglia B , Juan Carlos Villarreal B and R. Joel Duff C
+ Author Affiliations
- Author Affiliations

A Centre for Plant Biodiversity Research, GPO Box 1600, Canberra, ACT 2601, Australia.

B Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901, USA.

C Department of Biology, University of Akron, Akron, Ohio 44325, USA.

D Corresponding author. Email: Christine.Cargill@deh.gov.au

Australian Systematic Botany 18(1) 7-16 https://doi.org/10.1071/SB04012
Submitted: 3 June 2004  Accepted: 1 November 2004   Published: 29 March 2005

Abstract

Although the hornworts (anthocerotes) are a relatively small assemblage of approximately 150 species, generic boundaries and relationships within the group are controversial. The four prevailing classification schemes are based mainly on morphology and show little congruency. Here we set the foundation for contemporary phylogenetic and taxonomic studies by presenting an historical overview of generic concepts within the anthocerotes. An overview of recent morphological and molecular studies that concentrate on hornworts points to intuitive, novel relationships and a degree of diversity hitherto unknown in the group. Phylogenetically informative characters at the morphological level are identified, with emphasis on newly acquired ultrastructural data. A recent molecular analysis based on rbcL sequences is presented and the levels of suitability of several molecular markers to answer phylogenetic questions within the group are explored. On-going intensive studies that sample a wider range of species and utilise multiple genes and comprehensive morphological data are likely to revolutionise interpretations of the taxonomic relationships and character evolution within hornworts.


Acknowledgments

We thank Dr Helen Hewson for her reading and discussion of drafts of the manuscript. We would also like to thank two anonymous referees for their comments. The senior author would also like to acknowledge an ABRS grant for the revision of the Australian Anthocerotales and the Centre for Plant Biodiversity Research for ongoing support and availability of facilities for research into Australian hornworts. Support is acknowledged from National Science Foundation grants DEB-0235919, DEB-0235985 and DEB-0228679 as part of the Systematic biology and assembling the Tree of life programs, the Green Tree of Life Project (see http://ucjeps.berkeley.edu/TreeofLife/ validated 28 October 2004)


References


Asthana, AK ,  and  Srivastava, SC (1991). ‘Indian hornworts: a taxonomic study.’ (J Cramer: Berlin)

Bagchee K (1924) The spermatogenesis of Anthoceros laevis. Annals of Botany 38, 105–112. open url image1

Bartlett EM (1928) A comparative study of the development of the sporophyte in the Anthocerotaceae, with special reference to the genus Anthoceros. Annals of Botany 42, 409–430. open url image1

Beckert S, Steinhauser S, Muhle H, Knoop V (2000) A molecular phylogeny of bryophytes based on nucleotide sequences of the mitochondrial nad5 gene. Plant Systematics and Evolution 218, 179–192.
| Crossref |
open url image1

Beckert S, Hermann M, Pruchner D, Knoop V (2001) The mitochondrial nad2 gene as a novel marker locus for phylogenetic analysis of early land plants: a comparative analysis of mosses. Molecular Phylogenetics and Evolution 18, 117–126.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Bharadwaj DC (1971) On Folioceros, a new genus of Anthocerotales. Geophytology 1, 6–15. open url image1

Bharadwaj DC (1972) On some Asian and African species of Folioceros Bharadwaj. Geophytology 2, 74–89. open url image1

Bharadwaj, DC (1981). Taxonomy of Anthocerotales. In ‘Recent advances in cryptogamic botany. Part I: living cryptogams’. pp. 132–151. (The Palaeobotanical Society: Lucknow)

Bowe LM, dePamphilis CW (1996) Effects of RNA editing and gene processing on phylogenetic reconstruction. Molecular Biology and Evolution 13, 1159–1166.
| PubMed |
open url image1

Burr FA (1969) Reduction in chloroplast number during gametophyte regeneration in Megaceros flagellaris. The Bryologist 72, 200–209. open url image1

Campbell DH (1898) On the structure and development of Dendroceros Nees. Journal of the Linnaean Society 33, 467–478. open url image1

Campbell DH (1907) Studies on some Javanese Anthocerotaceae. I. Annals of Botany 21, 467–486. open url image1

Campbell DH (1908) Studies on some Javanese Anthocerotaceae. II. Annals of Botany 22, 91–102. open url image1

Campbell DH (1924) A remarkable development of the sporophyte in Anthoceros fusiformis. Annals of Botany 38, 473–483. open url image1

Campbell EO (1981a) Notes on some Anthocerotae of New Zealand. Tuatara 25, 7–13. open url image1

Campbell EO (1981b) Notes on some Anthocerotae of New Zealand (2). Tuatara 25, 65–70. open url image1

Campbell EO (1982a) Notes on some Anthocerotae of New Zealand (3). Tuatara 26, 20–26. open url image1

Campbell EO (1982b) Some Anthocerotae of New Zealand with particular reference to their geographical distribution. Journal of the Hattori Botanical Laboratory 52, 37–44. open url image1

Campbell EO (1984) Notes on some Anthocerotae of New Zealand (4). Tuatara 27, 105–120. open url image1

Campbell EO (1986) Notes on some Anthocerotae of New Zealand (5). Tuatara 26, 83–94. open url image1

Campbell EO (1993) Phaeoceros hirticalyx (Steph.) Haseg. (Anthocerotae) new to New Zealand. New Zealand Journal of Botany 31, 127–131. open url image1

Campbell EO (1995) Phaeoceros delicatus a new species of Anthocerotae from New Zealand. New Zealand Journal of Botany 33, 285–290. open url image1

Cargill DC, Scott GAM (1997) Taxonomic studies of the Australian Anthocerotales I. Journal of the Hattori Botanical Laboratory 82, 47–60. open url image1

Crum, H (2001). ‘Structural diversity of bryophytes.’ (The University of Michigan Herbarium: Ann Arbor)

Davis BM (1899) The spore mother cell of Anthoceros. Botanical Gazette 28, 89–109.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Dillenius, JJ (1741). ‘Historia muscorum.’ (Theatro Sheldoniano: Oxford)

Duff RJ, Nickrent DL (1999) Phylogenetic relationships of land plants using mitochondrial small-subunit rDNA sequences. American Journal of Botany 86, 372–386.
| PubMed |
open url image1

Duff RJ, Cargill DC, Villarreal JC, Renzaglia KS (2004) Phylogenetic relationships of the hornworts based on rbcL sequence data: novel relationships and new insights. Annals of the Missouri Botanical Gardens 98, 41–58. open url image1

Frey W, Hofman M, Hilger HH (2001) The gametophyte-sporophyte junction: unequivocal hints for two evolutionary lines of archegoniate land plants. Flora 196, 431–445. open url image1

Funk VA (1985) Cladistics and generic concepts in the Compositae. Taxon 34, 72–80. open url image1

Gray, SF (1821). ‘A natural arrangement of British plants.’ (Baldwin, Cradock and Joy: London)

Grolle R (1976) Verzeichnis der Lebermoose Europas und benachbarter Gebiete. Feddes Repertorium 87, 171–279. open url image1

Hasegawa J (1988) A proposal for a new system of the Anthocerotae, with a revision of the genera. Journal of the Hattori Botanical Laboratory 64, 87–95. open url image1

Hasegawa J (1994a) A remarkable new species of Phaeoceros (Anthocerotae) with canaliculated-striate spore surface. Journal of the Hattori Botanical Laboratory 75, 267–273. open url image1

Hasegawa J (1994b) New classification of Anthocerotae. Journal of the Hattori Botanical Laboratory 76, 21–34. open url image1

Hässel de Menéndez GG (1976) Taxonomic problems and progress in the study of the Hepaticae. Journal of the Hattori Botanical Laboratory 41, 19–36. open url image1

Hässel de Menéndez GG (1984) Anthoceros punctatus L., sus esporas. Cryptogamie. Bryologie, Lichenologie 5, 201–209. open url image1

Hässel de Menéndez GG (1986a) Leiosporoceros Hässel n. gen. and Leiosporocerotaceae Hässel n. fam. of Anthocerotopsida. Journal of Bryology 14, 255–259. open url image1

Hässel de Menéndez GG (1986b) Neue Lebermoosfunde aus dem Norden Argentiniens. Veröffentlichungen des Geobotanischen Institutes der ETH, Stiftung Rübel, in Zürich 91, 293–304. open url image1

Hässel de Menéndez GG (1987) Phaeoceros laevis (L.) Prosk. And P. carolinianus (Michx.) Prosk., their spores. Journal of the Hattori Botanical Laboratory 62, 281–288. open url image1

Hässel de Menéndez GG (1988) A proposal for a new classification of the genera within the Anthocerotophyta. Journal of the Hattori Botanical Laboratory 64, 71–86. open url image1

Hässel de Menéndez GG (1989) Las especies de Phaeoceros (Anthocerotophyta) de América del Norte, Sud y Central; la ornamentación de sus esporas y taxonomía. Candollea 44, 717–739. open url image1

Hässel de Menéndez GG (1990) Las especies de Anthoceros y Folioceros (Anthocerotophyta) de América del Norte, Sud y Central; la ornamentación de sus esporas y taxonomía. Candollea 45, 201–220. open url image1

Hedderson TA, Chapman RL, Rootes WL (1996) Phylogenetic relationships of bryophytes inferred from nuclear-encoded rRNA gene sequences. Plant Systematics and Evolution 200, 213–224.
| Crossref |
open url image1

Hofmeister, W (1851). ‘Vergleichende Untersuchungen der Keimung, Entfaltung und Fruchtbildung höherer Kryptogamen.’ (F Hofmeister: Leipzig)

Hyvönen J, Piippo S (1993) Cladistic analysis of the hornworts (Anthocerotophyta). Journal of the Hattori Botanical Laboratory 74, 105–119. open url image1

Janczewski E (1872) Vergleichende Untersuchungen über die Entwicklungsgeschichte des Archegoniums. Botanische Zeitung 30, 377–443. open url image1

Jones EW (1958) An annotated list of British hepatics. Transactions of the British Bryological Society 3, 353–374. open url image1

Kenrick, P ,  and  Crane, PR (1997). ‘The origin and early diversification of land plants: a cladistic study.’ (Smithsonian Institute Press: Washington DC)

Kugita M, Kaneko A, Yamamoto Y, Takeya Y, Matsumoto T, Yoshinaga K (2003) The complete nucleotide sequence of the hornwort (Anthoceros formosae) chloroplast genome: insight into the earliest land plants. Nucleic Acids Research 31, 716–721.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Leitgeb, H (1879). ‘Untersuchungen über die Lebermoose. V. Die Anthoceroteen.’ (Leuschnert and Lubensky: Graz)

Ligrone R, Duckett JG, Renzaglia KS (1993) The gametophyte-sporophyte junction in land plants. Advances in Botanical Research 19, 231–317. open url image1

Linnaeus, C (1753). ‘Species plantarum II.’ (Salvius: Stockholm)

Malek OK, Lättig R, Hiesel A, Brennicke A, Knoop V (1996) RNA editing in bryophytes and a molecular phylogeny of land plants. EMBO Journal 15, 1403–1411.
| PubMed |
open url image1

Meijer W (1954) Notes on some Malaysian species of Anthoceros L. (Hepaticae) — I. Reinwardtia 2, 411–423. open url image1

Micheli, PA (1729). ‘Nova plantarum genera.’ (Bernardi Paperinii: Florence)

Milde J (1859) über das Genus Notothylas Sullivant. Botanische Zeitung 17, 33–55. open url image1

Mishler BD, Churchill SP (1984) A cladistic approach to the phylogeny of the bryophytes. Brittonia 36, 406–424. open url image1

Mishler BD, Lewis LA, Buchheim MA, Renzaglia KS, Garbary DJ, Delwiche CF, Zechman FW, Kantz TS, Chapman RL (1994) Phylogenetic relationships of the ‘green algae’ and ‘bryophytes’. Annals of the Missouri Botanical Gardens 81, 451–483. open url image1

Mottier DM (1894) Contributions to the life-history of Notothylas. Annals of Botany 8, 391–402. open url image1

Nees von Esenbeck, CG (1846). Dendroceros. In ‘Synopsis hepaticarum. Fasc.4’. pp. 579. (Meissner: Hamburg)

Nickrent DL, Parkinson CL, Palmer JD, Duff RJ (2000) Multigene phylogeny of plants: hornworts are basal and mosses are sister to liverworts. Molecular Biology and Evolution 17, 1885–1895.
| PubMed |
open url image1

Nishiyama T, Kato M (1999) Molecular phylogenetic analysis among bryophytes and tracheophytes based on combined data of plastid coded genes and the 18S rRNA gene. Molecular Biology and Evolution 16, 1027–1036.
| PubMed |
open url image1

Piippo S (1993) Bryophyte flora of the Huon Peninsula, Papua New Guinea. LIV. Anthocerotophyta. Acta Botanica Fennica 148, 27–51. open url image1

Proskauer J (1948a) Studies on the morphology of Anthoceros I. Annals of Botany 12, 237–267. open url image1

Proskauer J (1948b) Studies on the morphology of Anthoceros I. Annals of Botany 12, 427–439. open url image1

Proskauer J (1951) Studies on Anthocerotales. III. Bulletin of the Torrey Botanical Club 78, 331–349. open url image1

Proskauer J (1953) Studies on Anthocerotales. IV. Bulletin of the Torrey Botanical Club 80, 65–75. open url image1

Proskauer J (1954a) A study of the Phaeoceros laevis complex. Rapports et Communications VIII. Congrés International de Botanique, Paris 14–16, 68. open url image1

Proskauer J (1954b) The European Anthocerotae. Rapports et Communications VIII. Congrés International de Botanique, Paris 14–16, 68–69. open url image1

Proskauer J (1957) [1958] Studies on Anthocerotales V. Phytomorphology 7, 113–135. open url image1

Proskauer, J (1958). Nachtrag zur Familie Anthocerotaceae. In ‘Die Lebermoose Europas, Rabenhorst’s Kryptogamen-Flora’. 3rd edn pp. 1303–1319. (Geest and Portig: Leipzig)

Proskauer J (1960) Studies on Anthocerotales. VI. Phytomorphology 10, 1–19. open url image1

Proskauer J (1967) Studies on Anthocerotales. VII. Phytomorphology 17, 61–70. open url image1

Proskauer J (1969) Studies on Anthocerotales. VIII. Phytomorphology 19, 52–66. open url image1

Qiu Y-L, Cho Y, Cox JC, Palmer JD (1998) The gain of three mitochondrial introns identifies liverworts as the earliest land plants. Nature 394, 671–674.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Renzaglia KS (1978) A comparative morphology and developmental anatomy of the Anthocerotophyta. Journal of the Hattori Botanical Laboratory 44, 31–90. open url image1

Renzaglia, KS ,  and  Vaughn, KC (2000). Anatomy, development and classification of hornworts. In ‘Bryophyte biology’. pp. 1–20. (Cambridge University Press: Cambridge)

Schuster RM (1987) Preliminary studies on Anthocerotae. Phytologia 63, 193–201. open url image1

Schuster, RM (1992). ‘The Hepaticae and Anthocerotae of North America, East of the Hundredth Meridian. Vol. 6.’ (Field Museum of Natural History: Chicago)

Singh, DK (2002). ‘Notothylaceae of India and Nepal.’ (Bishen Singh Mahendra Pal Singh: Dehra Dun, India)

Smith, GM (1955). ‘Cryptogamic botany. II. Bryophytes and Pteridophytes.’ (2nd edn) (McGraw-Hill Book Company: New York)

Stech M, Quandt D, Frey W (2003) Molecular circumscription of the hornworts (Anthocerotophyta) based on the chloroplast DNA trnL-trnF region. Journal of Plant Research 116, 389–398.
| Crossref |
open url image1

Stephani, F (1900). ‘Species hepaticarum. Vol. 1.’ (George and Cie: Geneva)

Stephani, F (1906). ‘Species hepaticarum. Vol. 2.’ (George and Cie: Geneva)

Stephani, F (1909). ‘Species hepaticarum. Vol. 3.’ (George and Cie: Geneva)

Stephani, F (1912). ‘Species hepaticarum. Vol. 4.’ (George and Cie: Geneva)

Stephani, F (1917). ‘Species hepaticarum. Vol. 5.’ (George and Cie: Geneva)

Stephani, F (1924). ‘Species hepaticarum. Vol. 6.’ (George and Cie: Geneva)

Stotler RE, Crandall-Stotler BJ (1977) A checklist of the liverworts and hornworts of North America. The Bryologist 80, 405–428. open url image1

Stotler RE, Crandall-Stotler BJ (2003) Proposals to conserve the name Anthoceros with a conserved type and the name Anthoceros agrestis against A. nagasakiensis (Anthocerotophyta). Taxon 52, 628–629. open url image1

Szmidt AE, Meng-Zhue L, Xiao-Ru W (2001) Effects of RNA editing on the coxI evolution and phylogeny reconstruction. Euphytica 118, 9–18.
| Crossref |
open url image1

Sullivant, WS (1845). ‘Musci alleghanienses exsiccatae.’ (Sullivant: Columbus, OH)

Udar, R ,  and  Singh, DK (1981). Recent concepts in the taxonomy of the genus Notothylas. In ‘Contemporary trends in plant sciences’. pp. 162–174. (Kalyani: New Delhi)

Vaughn KC, Ligrone R, Owen HA, Hasegawa J, Campbell EO, Renzaglia KS, Monge-Najera J (1992) The anthocerote chloroplast: a review. New Phytologist 120, 169–190. open url image1

Villarreal, JC ,  and  Salazar Allen, N (2002). Estudio preliminar de la morfología y distribución geográfica del genero Megaceros en Panamá. In ‘Libro de Resúmenes 8 Congreso Latinoamericano y Segundo Colombiano de Botánica.’ pp. 70. (Instituto de Ciencias Naturales: Universidad Nacional de Colombia, Bogotá)