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European Journal of Phycology This article was downloaded by:[UNAM] On: 26 September 2007 Access Details: [subscription number 768418234] Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK European Journal of Phycology Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713725516 A re-classification of the Acrochaetiales based on molecular and morphological data, and establishment of the Colaconematales ord. nov. (Florideophyceae, Rhodophyta) James T. Harper; Gary W. Saunders Online Publication Date: 09 September 2002 To cite this Article: Harper, James T. and Saunders, Gary W. (2002) 'A re-classification of the Acrochaetiales based on molecular and morphological data, and establishment of the Colaconematales ord. nov. (Florideophyceae, Rhodophyta)', European Journal of Phycology, 37:3, 463 - 476 To link to this article: DOI: 10.1017/S0967026202003840 URL: http://dx.doi.org/10.1017/S0967026202003840 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article maybe used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Eur. J. Phycol. (2002), 37: 463–476. # 2002 British Phycological Society 463 DOI: 10.1017\S0967026202003840 Printed in the United Kingdom A re-classification of the Acrochaetiales based on molecular and morphological data, and establishment of the Colaconematales ord. nov. (Florideophyceae, Rhodophyta) JAMES T. HARPER* AND GARY W. SAUNDERS Centre for Environmental and Molecular Algal Research, Department of Biology, University of New Brunswick, Fredericton, N.B., Canada, E3B 6E1 (Received 1 May 2002; accepted 1 May 2002) Systematics of the red algal order Acrochaetiales and related taxa was investigated using combined small- and large-subunit nuclear ribosomal DNA (SSU and LSU rDNA, respectively) sequence data. These data were subjected to distance, parsimony and maximum likelihood analyses. The resulting phylogenies were congruent with previously published SSU results in that all included orders (Balbianiales, Batrachospermales, Nemaliales, Palmariales and Thoreales) were resolved as monophyletic except the Acrochaetiales, which consisted of two lineages (Acrochaetiales I and II). The Batrachospermales and Thoreales occupied equivocal positions as early diverging lineages, while the Balbianiales generally Downloaded By: [UNAM] At: 00:57 26 September 2007 resolved as sister to an Acrochaetiales–Nemaliales–Palmariales (ANP) complex. Relationships among the four lineages of the ANP complex were not completely resolved, but detailed analyses weakly positioned Acrochaetiales II as sister to the Nemaliales, whereas Acrochaetiales I displayed a moderate to strong affiliation with the Palmariales. Acrochaetiales I included representatives of the genera Acrochaetium, Audouinella and Rhodochorton, whereas Acrochaetiales II had a number of acrochaetioid species including a representative of the genus Colaconema. Compared with published SSU phylogenies, bootstrap values within the two Acrochaetiales lineages increased substantially in combined SSU\LSU analyses. Based on these results, emended generic descriptions are provided for Acrochaetium, Audouinella, Colaconema and Rhodochorton, and a new family of acrochaetioid algae is described, the Colaconemataceae J. T. Harper et G. W. Saunders. The Acrochaetiaceae now includes the genera Acrochaetium, Audouinella and Rhodochorton, while the Colaconemataceae is considered monogeneric at this time. It is quite likely that additional genera will be recognized within the Colaconemataceae pending further investigation. Acrochaetiaceae is retained as the sole family of the Acrochaetiales, although the tenuous recognition of the Palmariales as distinct from this order is discussed, whereas Colaconemataceae is transferred to the new order Colaconematales J. T. Harper et G. W. Saunders. Key words: Acrochaetiaceae, Acrochaetiales, Colaconemataceae fam. nov., Colaconematales ord. nov., Florideophyceae, large-subunit rDNA, molecular systematics, phylogeny, Rhodophyta, small-subunit rDNA Introduction spermales, Corallinales, Nemaliales, Palmariales, Rhodogorgonales and Thoreales nom. int. (cf. Recent molecular systematic investigations have Sheath et al., 2000). resolved four higher-level lineages within the red Until recently, there were few molecular sys- algal class Florideophyceae (Saunders & Bailey, tematic investigations focused on the members of 1997, 1999; Saunders & Kraft, 1997; Harper & this lineage. However, studies using small-subunit Saunders, 2001b). Of these, Lineage 2 includes nuclear ribosomal DNA (SSU rDNA) data have those members possessing two caps on the cyto- resolved many novel relationships among the in- plasmic faces of the pit plugs, with or without a cluded taxa. Saunders et al. (1995) illustrated the membrane. This lineage includes the orders close relationship between members of the Acro- Acrochaetiales, Balbianiales, Balliales, Batracho- chaetiales, Nemaliales and Palmariales, whereas Saunders & Bailey (1997) provided the first mol- Correspondence to: J. T. Harper. e-mail: jtharper!interchange. ecular evidence for an association between the ubc.ca Corallinales and Rhodogorgonales. Vis et al. (1998) * Present address: Canadian Institute for Advanced Research, examined the systematics of the Batrachospermales, Evolutionary Biology Program, Department of Botany, University of British Columbia, 3529-6270 University Blvd., Vancouver, B. C., and highlighted the unusual position of the Canada, V6T 1Z4. Thoreaceae with respect to other members of the J. T. Harper and G. W. Saunders 464 order. Harper & Saunders (1998) provided the first siella, Chromastrum, Colaconema, Conferva, Grania, detailed molecular study of the Acrochaetiales and Kylinia, Liagorophila, Pseudacrochaetium, Pseu- discovered two relatively divergent lineages (Acro- dochantransia, Rhodochorton, Rhodothamniella, chaetiales I and II) within the order which seemed, Thamnidium and Trentepohlia. Of these, only seven at least morphologically, homogeneous. Sheath & are currently available for use with species of this Mu$ ller (1999) reported molecular evidence for the order: Acrochaetium, Audouinella, Colaconema, recognition of the Balbianiales as a distinct clade Grania, Kylinia, Liagorophila and Rhodochorton (for within Lineage 2, while the Balliales was proposed more detailed accounts see Woelkerling, 1971, by Choi et al. (2000) to accommodate a few red algal 1983). At least 24 different classification schemes species previously classified in the Ceramiales. involving taxa of generic and higher rank have been Lastly, Sheath et al. (2000) have proposed rec- proposed and these concepts have ranged from ognition of the Thoreales for the batrachosperm- recognizing a single genus (Drew, 1928; Dixon & alean Thoreaceae, which had consistently occupied Irvine, 1977; Garbary, 1978) to as many as eight a relatively distinct position in rbcL- and SSU- genera (Feldmann, 1962). These discrepancies in the based phylogenies (Harper & Saunders, 1998; Vis literature have led to multiple concepts associated et al., 1998). with each genus. In spite of these taxonomic advances, a number of Such an abundance of generic concepts has led to, relationships within Lineage 2 remain unresolved. and then been fuelled by, a lack of consensus as to In particular: (1) the relative positions of the which characters are useful for taxonomy at this Balbianiales, Batrachospermales and Thoreales; rank. In 1928, Drew advocated a monogeneric and (2) the relationships between the Acro- Acrochaetiaceae and placed all marine and fresh- chaetiales, Nemaliales and Palmariales. water acrochaetioid taxa within Rhodochorton. The Acrochaetiales presents an interesting prob- Thus, this genus included plants in which the Downloaded By: [UNAM] At: 00:57 26 September 2007 lem for red algal systematists. Compared with the chloroplasts vary from one to many per cell, and are remainder of the Florideophyceae, members of the parietal, stellate, ribbon-shaped, discoid or have Acrochaetiales possess simple vegetative (generally radiating lobes. Papenfuss (1945) and Feldmann heterotrichous, simple or branched filaments) and (1962), however, recognized multiple genera within reproductive (absence of a carpogonial branch, the Acrochaetiales and had a restricted concept of reduced postfertilization development) attributes. It Rhodochorton: asexual marine plants with cells is this apparent simplicity that has led to arguably possessing few to many discoid chloroplasts. The one of the most chaotic taxonomic histories in red other included genera, notably Acrochaetium and algal systematics. Since first proposed by Feldmann Audouinella, have suffered similarly owing
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