The red algal genus Reticulocaulis from the Arabian

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54 Phycologia, Vol. 42 (1), 2003 Table 2. Comparison of morphological and anatomical characteristics among the genera of the Naccariaceae (Zerlang 1889; Kylin 1928; Kylin 1956; Fan 1961; Womersley & Abbott 1968; Dixon & Irvine 1977; Abbott 1985; Hommersand & Fredericq 1990; Millar 1990; Womersley 1996; Abbott 1999; this article). Feature Atractophora Naccaria Reticulocaulis Branching pattern of determinate branches whorls of four periaxial cells per axial cell two periaxial cells per axial cell; branching on an irregular spiral of 1/4 two periaxial cells per axial cell; branching on an irregular spiral of 1/4 Sheath of inflated cells around absent present present axial strand Secondary pit connections absent present between jacket cells present between jacket cells Adventitious and rhizoidal filaments present present present Hairs on terminal cortical filament present present or absent present or absent cells Gametophytes monoecious monoecious or dioecious monoecious or dioecious Carpogonial branch (including three- to four-celled two- to eight-celled seven- to thirteen-celled carpogonium) Laterals on carpogonial branch cells short short primary laterals lengthy, secondary sterile filaments present Nutritive cells present absent or restricted to the hypogynous cell Immediate post-fertilization events Gonimoblast composition Localized swelling of the branches by mature cystocarps Position of spermatangial mother cells fusion of carpogonium with supporting cell by a short process fusion of carpogonium with hypogynous cell by widening of pit connection diffuse gonimoblast intermixed with vegetative filaments always present on the hypogynous cell, small clusters on the carpogonial branch cells #3, #4 fusion of carpogonium with hypogynous cell by widening of pit connection diffuse gonimoblast intermixed compact gonimoblast lacking with vegetative filaments vegetative filaments present present absent ? dendroid distal clusters or in catenate series dendroid distal clusters or in catenate series surround their central-axial filaments with a jacket of uninflated cells, have three- to four-celled recurved carpogonial branches bearing more than two sterile groups and produce a carposporophyte that surrounds the axial strand and intermingles with vegetative filaments. Liagorothamnion may thus prove to have a closer alliance with the Naccariaceae than with the Ceramiaceae. Abbott (1985) has suggested that Reticulocaulis ‘might be looked upon as reduced from Acrosymphyton-like forms in terms of the carpogonial branch . . .’ and hence related to the Dumontiaceae, a family then placed in the order Cryptonemiales and now a member of the Gigartinales (Saunders & Kraft 1997). The proposed affinity between Reticulocaulis and Acrosymphyton is not supported by recent evidence. According to a phylogenetic analysis of the Dumontiaceae (Tai et al. 2001), Acrosymphyton (Acrosymphytaceae; Lindstrom 1987) is sister to the Gelidiales, whereas Bonnemaisonia is basal to the included Gigartinales and Gelidiales. Ongoing molecular research (G.W. Saunders & C.A. Maggs, personal communication) shows that the Naccariaceae is almost certainly not monophyletic (Naccaria groups weakly with the Bonnemaisoniaceae and may not belong in the Bonnemaisoniales, whereas Atractophora receives solid support as a member of the Bonnemaisoniaceae), and it clearly does not belong in either the Nemaliales or the Gigartinales. Saunders & Kraft (1997, p. 130) suggested that DNA studies of the Naccariaceae, to establish its ordinal affinities, should be a top priority for molecular systematists, and this recommendation still holds. ACKNOWLEDGEMENTS We greatly appreciate the useful comments of Gerry Kraft and Max Hommersand, which helped improve the manuscript. We also give sincere thanks to Isabella Abbott for her help and enthusiasm in providing Hawaiian specimens, and the Senckenberg Research Institute, Germany (Michael Apel and Friedhelm Krupp), the UNDP Socotra Marine Team, and the Ardoukoba Organization (France) for providing excellent organization and a pleasant ambience during both field trips. Special thanks are due to the fellow ‘missionaries’, whose cheerful and professional spirit kept our little commune going efficiently. T.S. is much indebted to the diving partners Mohammed Ismail, Ali Bin Naser Al Rasibi and André Germé, who all became intrigued with hunting static sea life. The Latin translation was kindly supplied by Paul Goetghebeur. T.S. and O.D.C. are indebted to the Fund for Scientific Research Flanders (Belgium) for research assistant and postdoctoral research grants, respectively. Financial support was provided by the FKFO project 3G002496. REFERENCES ABBOTT I.A. 1985. 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The occurrence of Jolyna laminarioides (Phaeophyta) in the Arabian Sea and the Indian Ocean and a new report of Melanothamnus somalensis (Rhodophyta). Botanica Marina 33: 213–218. WYNNE M.J. & JUPP B.P. 1998. The benthic marine algal flora of the Sultanate of Oman: new records. Botanica Marina 41: 7–14. ZERLANG O.E. 1889. Entwicklungsgeschichtliche Untersuchungen über die Florideengattungen Wrangelia und Naccaria. Flora 72: 371–407. Accepted 28 July 2002
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The red algal genus Reticulocaulis from the Arabian

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