Systematics of Liagora with diffuse gonimoblasts based on rbcL sequences and carposporophyte development, including the description of the new genera Neoizziella and Macrocarpus (Liagoraceae, Rhodophyta)

Figures & data

Table 1. List of species of the Liagoraceae used for rbcL analysis in this study and their GenBank accession numbers.

Species	Collection information	rbcL accession no.
Akalaphycus setchelliae (Yamada) Huisman, I.A. Abbott & A.R. Sherwood	Jian Shan, Penghu Archipelago, W Taiwan, coll. S.-M. Lin, 28 April 2003	^1GU357697
Akalaphycus setchelliae (Yamada) Huisman, I.A. Abbott & A.R. Sherwood	Bulusan, Luzon Island, The Philippines, S.-L. Liu, 12 February 2003	^1GU357698
Dermonema virens (J. Agardh) Pedroche & Ávila Ortíz	Lung Keng, Pingtung Co., S Taiwan, coll. S.-M. Lin, 9 March 2008	^2HQ901787
Ganonema farinosum (J.V. Lamouroux) K.C. Fan & Y.C. Wang	Horse Bridge, Green Island, Taitung Co., E Taiwan, coll. S.-L. Liu, 29 August 2002	^1GU357699
Ganonema farinosum (J.V. Lamouroux) K.C. Fan & Y.C. Wang	Bulusan, Luzon Island, The Philippines, S.-L. Liu, 12 February 2003	^1GU357700
Helminthocladia australis Harvey	Chuan Fan Shi, Kenting National Park, Pingtung Co., S Taiwan, coll. S.-L. Liu, 15 January 2006	^1GU357701
Izziella formosana (Yamada) Showe-Mei Lin, Shi-Yuan Yang & Huisman	(♀) Chuan Fan Shi, Kenting National Park, Pingtung Co., S Taiwan, coll. S.-L. Liu, 15 January 2006	^1GU357688
Izziella kuroshioensis Showe-Mei Lin, Shi-Yuan Yang & Huisman	(♀) ‘slender form’, Hai Kuo, Pingtung Co., S Taiwan, coll. S.-M. Lin, 12 February 2009	^1GU357685
Liagora albicans J.V. Lamouroux	Feng Chui Sha, Pingtung Co., S Taiwan, coll. S.-M. Lin, 27 April 2006	^2HQ901786
Liagora boergesenii Yamada	Wan Li Tong, Kenting National Park, Pingtung Co., S Taiwan, coll. S.-M. Lin, 1 November 2003	^1GU357679
Liagora ceranoides J.V. Lamouroux	Nan Wan, Kenting National Park, Pingtung Co., SE Taiwan, coll. S.-L. Liu, 17 April 2005	^1GU357681
Liagora ceranoides J.V. Lamouroux	Florida, USA, coll. M.H. Hommersand, 12 March 1997	^1GU357682
Liagora harveyana Zeh	New Zealand, coll. W. Freshwater, 12 November 2004 (NZ 4-432)	^1HM572263
Liagora harveyana Zeh	Port Fairy, Victoria, South Australia, coll. A.H.S. Lucas, January 1932 (NSW 401401)	^1HM572264
Liagora viscida (Forsskål) C. Agardh	El Golfet, Calella de Palafrugell, Spain, coll. C. Rodriguez-Prieto, 12 August 2005	^1GU357678
Macrocarpus perennis (I.A. Abbott) Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et comb. nov.	Wan Li Tong, Pingtung Co., S Taiwan, coll. S.-M. Lin, 23 December 2007	^2HQ901783
Macrocarpus perennis (I.A. Abbott) Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et comb. nov.	Hougtou, Orchid Island, SE Taiwan, coll. S.-M. Lin, 28 April 2002	^2HQ901784
Macrocarpus perennis (Abbott) Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et comb. nov.	Xian Gang Kou, Pingtung Co., S Taiwan, coll. S.-M. Lin, 12 February 2009	^2HQ901785
Neoizziella asiatica Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et sp. nov.	Won Li Tong, Pingtung Co., S Taiwan, coll. S.-M. Lin, 23 December 2007	^2HQ901778
Neoizziella asiatica Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et sp. nov.	Feng Chui Sha, Pingtung Co., S Taiwan, coll. S.-M. Lin, 18 March 2003	^2HQ901779
Neoizziella asiatica Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et sp. nov,	Shin Lan, Taitung Co., E Taiwan, coll. S.-M. Lin & X.-R. Hsueh, 8 March 2008	^2HQ901780
Neoizziella asiatica Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et sp. nov.	Pratas Island, Taiwan, coll. S.-M. Lin, 25 April 2004	^2HQ901788
Neoizziella asiatica Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et sp. nov.	Serangan I., Bali, Indonesia, coll. S.-M. Lin, 24 July 2008	^2HQ901789
Neoizziella asiatica Showe-Mei Lin, Shi-Yuan Yang & Huisman, gen et sp. nov.	Cebu, the Philippines, coll. S.-L. Liu, 07 November 2002	^2HQ901788
Neoizziella divaricata (C.K. Tseng) Showe-Mei Lin, Shi-Yuan Yang & Huisman, comb. nov.	Feng Gang, Pingtung Co., S Taiwan, coll. S.-L. Liu, 16 March 2006	^2HQ901781
Neoizziella divaricata (C.K. Tseng) Showe-Mei Lin, Shi-Yuan Yang & Huisman, comb. nov.	Hai Kou, Pingtung Co., S Taiwan, coll. S.-M. Lin, 12 February 2009	^2HQ901782
Stenopeltis gracilis (Yamada & Tanaka) Itono & Yoshizaki	Da Bai Sha, Green Island, E Taiwan, coll. S.-L. Liu, 28 August 2002	^1GU357695
Stenopeltis gracilis (Yamada & Tanaka) Itono & Yoshizaki	Lung-Meng Port, Orchid Island, Taitung Co., SE Taiwan, coll. S.-M. Lin, 26 April 2002	^1GU357696
Titanophycus setchellii (Yamada) Showe-Mei Lin, Shi-Yuan Yang & Huisman	(♂♀) Da Bai Sha, Green Island, E Taiwan, coll. S.-M. Lin, 26 April 2002	^1GU357694
Titanophycus validus (Harvey) Huisman, G.A. Saunders & A.R. Sherwood	(♀) As ‘Liagora’ #23, French Bay, San Salvador, Bahamas Islands, coll. Max H. Hommersand, 26 March 2006	^1GU357693
^1 Lin et al. (2011); ^2 this study.

Fig. 1. RbcL phylogeny: ML tree (ln L = –10165.9564) of the proposed new genera Neoizziella and Macrocarpus from the northwestern Pacific Ocean, using a selection of the related genera within the families Liagoraceae, Scinaiaceae and Galaxauraceae. Numbers above branches are ML bootstrap values, MP bootstrap values and Bayesian posterior probabilities in %, respectively. A dash indicates support values <50%.

Figs 2–11. Neoizziella asiatica, sp. nov. 2. Holotype, a female plant. 3. Habit, a liquid preserved, female plant. 4. Close-up of a female plant showing branching pattern and cystocarps (arrows). 5. Young cortex showing spermatangial initials. 6. Differentiating spermatangia (arrowheads) and the discharged spermatangia (white arrows). 7. Mature spermatangia (arrowheads) in cluster. 8. Close-up of assimilatory filaments and carpogonial branches (arrows). 9. An early post-fertilization stage showing supporting cell (sc), carpogonial branch (cb), fertilized carpogonium (ca), gonimoblast initial (gi) and initials of involucral filaments (arrows). 10. Another post-fertilization stage showing supporting cell (sc), carpogonial branch (cb), fertilized carpogonium (ca), early gonimoblasts (g) and branched involucral filaments (arrows). 11. A later post-fertilization stage showing supporting cell (sc), elongated carpogonium (ca), developing gonimoblasts (arrowheads) and elongated involucral filaments (arrows).

Figs 12–15. Neoizziella asiatica, gen. et sp. nov. Figs 12, 13. Two views of a young carposporophyte. 12. Focal plane of the involucral filaments (arrows), which surround the gonimoblasts completely. 13. Focal plane of distinct carpogonial branch (cb), the gonimoblasts (arrowheads) and the rear involucral filaments (arrows). 14. Nearly mature carposporophyte showing the gonimoblasts (arrowheads) intermingling with involucral filaments (arrows). Note that the cells of carpogonial branch (cb) are not fused. 15. Fully developed gonimoblasts bearing terminal, mature carposporangia (arrowheads). Note that the cells of carpogonial branch (cb) and the supporting cell (sc) remain distinct.

Figs 16–21. Neoizziella divaricata, comb. nov. (Hico, Pintung County, Taiwan). 16. Habit, a female plant. 17. Close-up of assimilatory filaments and a carpogonial branch (arrow). 18. An early post-fertilization stage showing supporting cell (sc), carpogonial branch (cb), gonimoblast initial (gi) and young involucral filaments (arrows). 19. A later post-fertilization stage showing carpogonial branch (cb), young gonimoblasts (arrowheads) and elongated involucral filaments (arrows). 20. Fully developed gonimoblasts (arrowheads) intermingling with involucral filaments (arrows). 21. Close of another mature cystocarp showing undivided carposporangia (arrowheads) and involucral filaments (arrows).

Figs 22–25. Macrocarpus perennis, gen. et comb. nov. (Figs 22, 23: Won-Li-Dong, PinTung County, Taiwan; Figs 24, 25: Houtou, Orchid Island, Taiwan). 22. Habit of a female plant. 23. Close-up of assimilatory filaments and carpogonial branches (cb). 24. An early post-fertilization stage showing carpogonial branch (cb), gonimoblast initial (gi), primary gonimoblasts (arrowheads) and young involucral filaments (arrows). 25. A later post-fertilization stage showing carpogonial branch (cb), gonimoblast initial (gi), young gonimoblasts (arrowheads) and involucral filaments (arrows).

Figs 26–33. Liagora spp. Figs 26–29. Liagora viscida (Figs 26, 27: Cabrera, Illes Balears, Spain; Figs 28, 29: Capraia, Tuscan Islands, Italy). 26. Close-up of assimilatory filaments and a carpogonial branch (cb). 27. An early post-fertilization stage showing carpogonial branch (cb), gonimoblast initial (gi) and young involucral filaments (arrows). 28. A later post-fertilization stage showing young gonimoblasts (arrowheads) and elongated involucral filaments (arrows). 29. Fully developed gonimoblasts (arrowheads) showing mature carposporangia, fusion cell (fc) and surrounding involucral filaments (arrows). Note that some gonimoblasts are squeezed out (white arrows). Figs 30–33. Liagora harveyana, carposporophyte development (Figs 30, 31, 33: Port Fairy, Victoria, southern Australia; Fig. 32: Leigh, Auckland Island, New Zealand). 30. An early post-fertilization stage showing carpogonial branch (cb), gonimoblast initial (gi) and young involucral filaments (arrows). 31. A later post-fertilization stage showing young gonimoblasts (arrowheads) and elongated involucral filaments (arrows). 32. Fully developed gonimoblasts (arrowheads) showing undivided carposporangia (arrowheads), fusion cell (fc) and involucral filaments (arrows). Note that the involucral filaments loosely surround the gonimoblasts. 33. Mature cystocarp showing divided (white arrowheads) and undivided (black arrowheads) carposporangia, fusion cell (fc) and involucral filaments (arrows). Note that the involucral filaments intermingle with the gonimoblasts.

Lin , S-M , Yang , S-Y and Huisman , JM . 2011 . Systematic revision of the genera Liagora and Izziella (Liagoraceae, Rhodophyta) from Taiwan based on molecular analyses and carposporophyte development, with the description of two new species . J. Phycol. , 47 : 352 – 365 .

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