Silicascaled Chrysophytes from Hainan, Guangdong Provinces And

Silica-scaled chrysophytes from Hainan, Guangdong Provinces and Hong Kong Special Administrative Region, China

Yin-Xin Wei , Xiu-Ping Yuan and J ø rgen Kristiansen

Y.-X. Wei ([email protected]) and X.-P. Yuan, Inst. of Hydrobiology, Chinese Academy of Sciences, CN-430072 Wuhan, P. R. China. – J. Kristiansen, Biological Inst., Univ. of Copenhagen, DK-1165 Copenhagen, Denmark.

Twenty fi ve phytoplankton samples containing chrysophytes were collected in Hainan and Guangdong Provinces and in Hong Kong Special Administrative Region in January 2010 and October to November 2011. Forty nine taxa of silica-scaled chrysophytes were identifi ed by means of transmission electron microscopy, twelve of them belonging to Paraphysomonadaceae and thirty seven to Synurales. Spiniferomonas cornuta , Paraphysomonas sp., Mallomonas hexareticulata , M . ceylanica , M . plumosa , M . ocellata , M . elevata and M . spinosa are reported for the fi rst time from China. A new species, Paraphysomonas hainanensis Wei & Kristiansen, was found in a pond in Haikou City, Hainan Province. An amended description of Mallomonas spinosa Gusev is proposed. Th e sampled localities are situated in the tropical and subtropical region of southern China. Th ere is great similarity in the geographical distribution of the observed silica-scaled chrysophytes with that of east Asia, southeast Asia and south Asia, and their ecological characteristics are discussed.

Since Fott (1955) fi rst reported the fi ne structures of In order to compile a freshwater silica-scaled chrysophyte Mallomonas scales using transmission electron microscopy, fl ora of China, extensive investigations on geographical dis- many electron microscope-based silica-scaled chrysophyte tribution have been performed since 1988 (Kristiansen studies have been made in diff erent parts of the world, espe- 1989, 1990, Kristiansen and Tong 1988, 1989a, 1989b, cially in Europe and North America (Takahashi 1978, Siver 1991, Wei and Kristiansen 1994, 1998, Wei and Yuan 2001, 1991, Kristiansen 2002, 2005, Kristiansen and Preisig 2013, Ma and Wei 2013, Pang and Wang 2013). Th e pres- 2007, Š kaloud et al. 2013). Silica-scaled chrysophytes ent study focuses on the occurrence and distribution of these are often important components of the phytoplankton algal groups in the Hainan and Guangdong Provinces and in community of freshwater lakes and reservoirs. Th ey not lim- Hong Kong Special Administrative Region. Th ese regions ited to cold and oligotrophic areas, as rich silica-scaled belong to the tropical and subtropical region of southern chrysophyte fl oras have been found in diverse localities China which have not been studied in detail with regards to including temperate, tropical and subtropical regions and in chrysophyte diversity before. eutrophic water bodies (D ü rrschmidt and Cronberg 1989, Hansen 1996, Kristiansen 2001, Gusev 2012). Th e freshwater silica-scaled chrysophytes in localities of Material and methods 20 Chinese provinces have previously been investigated using transmission electron microscopy. In total, eighty Twenty fi ve phytoplankton samples were collected during two taxa have been reported of which forty eight taxa belong- 12 – 21 January 2010, 17 October and 7 – 17 November ing to the genus Mallomonas . Mallomonas asymmetrica and 2011 in Hainan and Guangdong Provinces and in Hong Synura morusimila are two endemic species which have only Kong Special Administrative Region (18° 17 ′ 34.95 ″ – been found in northeast China so far. Mallomonas koreana is 23 ° 11 ′ 19.52 ″ N, 109 ° 11 ′ 13.16 ″ – 114 ° 16 ′ 32.01 ″ E). Biologi- a rare species found in Korea (Kim et al. 2008) and France cally important ponds, lakes and reservoirs of diff erent (N ě mcov á et al. 2012), and afterwards recorded from habitat types were selected (Fig. 1, Table 1). Water bodies the Zhejiang Province, China (Wei and Yuan 2013). with diff erent ecological features have been collected in these Mallomonas grata is distributed in eastern Asia, North and regions, but in the twenty fi ve examined ponds, lakes Central America and has been recorded from the Heilongjiang, and reservoirs containing chrysophytes the water is mainly Zhejiang, Jiangsu, Hubei, Fujian, and Guangdong Prov- oligotrophic. All samples were collected using a plankton inces, China. net (10 μ m mesh size) and were immediately fi xed in the

881 Figure 1. Map (A) Th e map of China showing studied regions; (B) Locations of 25 sampling sites see Table 1. a: 1-2 (Haikou City); b: 3 (Wenchang City); c: 4-8 (Xinglong County); d: 9-11 (Sanya City); e: 12-15 (Guangzhou City); f: 16-21 (Shenzhen City); g: 22-25 (Hong Kong) .

fi eld using Lugol ’ s solution with acetic acid. Th e water tem- Institute of Hydrobiology, Chinese Academy of Sciences. perature and pH of each site were measured in the fi eld Environmental parameters and geographic coordinates are with a CT-6021A pH meter. Th e samples were examined summarized in Table 1. preliminary using an Olympus BH-2 light microscope in order to determine species composition, focusing on the chrysophytes. Th e fi xed material was dried onto Results formvar-coated copper grids (150 mesh) and rinsed with distilled water. After drying, the grids were examined in a In total, forty four species and fi ve varieties of silica- JEOL, JEM-1230 transmission electron microscope in the scaled chrysophytes were identifi ed from the twenty fi ve

Table 1. Surveyed water bodies, their water temperature ( o C), pH and geographic coordinates.

Locality Water body type Date Temp. (o C) pH Geographic coordinates Hainan 1. Haikou City Shabo Reservoir 12/01/2010 22 6.2 19 ° 57 ′ 36.03 ″ N, 110 ° 19 ′ 27.97 ″ E 2. Haikou City Pond 12/01/2010 22 7.2 20 ° 0 ′ 55.62 ″ N, 110 ° 19 ′ 2.22 ″ E 3.Wenchang City Pond 13/01/2010 16 7.5 19 ° 38 ′ 46.83 ″ N, 110 ° 54 ′ 5.46 ″ E 4. Xinglong County small lake 14/01/2010 21 6.9 18 ° 44 ′ 3.52 ″ N, 110 ° 11 ′ 45.84 ″ E 5. Xinglong County Shengtai Lake 14/01/2010 21 6.9 18 ° 44 ′ 3.5 ″ N, 110 ° 11 ′ 45.86 ″ E 6. Xinglong County Pond 15/01/2010 22 7.2 18 ° 44 ′ 3.55 ″ N, 110 ° 11 ′ 45.86 ″ E 7. Xinglong County Pond 15/01/2010 22 7.1 18 ° 44 ′ 12.04 ″ N, 110 ° 11 ′ 32.58 ″ E 8. Xinglong County Xiao Lake 15/01/2010 24 6.8 18 ° 44 ′ 17.36 ″ N, 110 ° 11 ′ 56.23 ″ E 9. Sanya City Pond 17/01/2010 24 6.9 18 ° 17 ′ 34.95 ″ N, 109 ° 11 ′ 17.86 ″ E 10. Sanya City Pond 17/01/2010 24 6.9 18 ° 17 ′ 37.06 ″ N, 109 ° 11 ′ 15.21 ″ E 11. Sanya City Pond 17/01/2010 24 7.2 18 ° 17 ′ 44.43 ″ N, 109 ° 11 ′ 13.16 ″ E Guangdong 12. Guangzhou City Pond 20/01/2010 18 6.3 23 ° 11 ′ 12.58 ″ N, 113 ° 21 ′ 49.21 ″ E 13. Guangzhou City Pond 20/01/2010 18 6.3 23 ° 11 ′ 19.52 ″ N, 113 ° 21 ′ 51.06 ″ E 14. Guangzhou City Pond 20/01/2010 18 7.1 23 ° 11 ′ 13.86 ″ N, 113 ° 21 ′ 45.52 ″ E 15. Guangzhou City Pond 21/01/2010 19 7.5 23 ° 11 ′ 4.96 ″ N, 113 ° 21 ′ 43.23 ″ E 16. Shenzhen City Tiegang Reservoir 17/10/2011 25 7.4 22 ° 37 ′ 16.02 ″ N, 113 ° 53 ′ 55.06 ″ E 17. Shenzhen City Bijiashan Reservoir 17/10/2011 25 6.9 22 ° 34 ′ 0.42 ″ N, 114 ° 4 ′ 38.36 ″ E 18. Shenzhen City Shatoujiao Reservoir 17/10/2011 26 7.1 22 ° 35 ′ 7.48 ″ N, 114 ° 16 ′ 27.89 ″ E 19. Shenzhen City Xili Reservoir 17/10/2011 25 7.4 22 ° 35 ′ 26.96 ″ N, 113 ° 57 ′ 35.41 ″ E 20. Shenzhen City Donghu Reservoir 7/11/2011 26 7.4 22 ° 34 ′ 2.11 ″ N, 114 ° 8 ′ 54.91 ″ E 21. Shenzhen City Meisha Reservoir 7/11/2011 25 6.8 22 ° 37 ′ 15.33 ″ N, 114 ° 16 ′ 32.01 ″ E Hong Kong 22. Hong Kong Datan Reservoir 14/11/2011 25 6.9 22 ° 15 ′ 33.01”N, 114 ° 12 ′ 36.21”E 23. Hong Kong Bofulincun Reservoir 16/11/2011 25 6.8 22 ° 15 ′ 36.53”N, 114 ° 8 ′ 12.69”E 24. Hong Kong Shilibei Reservoir 16/11/2011 25 6.5 22 ° 21 ′ 21.55”N, 114 ° 8 ′ 52.88”E 25. Hong Kong Shibi Reservoir 17/11/2011 24 7.5 22 ° 13 ′ 57.48”N, 113 ° 53 ′ 55.01”E

882 Table 2. Occurrence of silica-scaled chrysophytes found in Hainan, Guangdong Provinces and Hong Kong Special Administrative Region, China. abundant ϭ a; common ϭ c; occasional ϭ o; single (one ∗ scales on three grids) ϭ s. Species marked with are new to China.

Locations Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 pH Temp. ( ° C) Chrysophyceae Paraphysomonadaceae Chrysosphaerella annulata Kristiansen & Tong a a a a o c c 6.2 – 7.2 21 – 26 coronacircumspina Wujek & Kristiansen c c c a a c c c c c a c c a a o c 6.2 – 7.5 21 – 26 longispina Lauterbornem. Nicholls s 7.2 22 septispina (Nicholls) Kristiansen & Tong c 7.4 25 Spiniferomonas ∗ cornuta Balonov o c s 6.3 – 7.1 18 trioralis Takahashi c c o c c c a o o a c c 6.3 – 7.5 18 – 25 Paraphysomonas gladiata Preisig & Hibberd c c 7.4 25 ∗ hainanensis Wei & Kristiansen sp. nov. c 7.2 22 imperforata Lucas o o c o o o o 6.2 – 7.4 18 – 25 runcinifera Preisig & Hibberd o 6.2 22 ∗ sp. c c o o c 6.5– 7.5 24 – 25 vestita (Stokes) de Saedeleer a a c c o a a c o o c o c o c c c c 6.2 – 7.5 18 – 26 Synurales Mallomonas acaroids Perty em. Ivanov c o 6.8– 7.4 25 akrokomos Ruttner in Pascher a a o o 6.3 – 7.5 18 – 25 alpina Pascher & Ruttner em. Asmund & Kristiansen o c o a c 6.3– 7.5 18 – 24 areolata Nygaard o s o 6.3 – 7.4 18 – 25 caudata Ivanov em. Krieger o s c o a a o c o o c c c c 6.3 – 7.5 16 – 25 ∗ ceylanica D ü rrschmidt & Cronberg o 7.2 22 corymbosa Asmund & Hilliard c a a a a o c c o a a s o o c c a 6.2 – 7.5 18 – 26 costata D ü rrschmidt s s s 6.3 – 7.4 18 – 25 crassisquama (Asmund) Fott s o o c a c o c o a c c c c c o 6.3 – 7.5 16 – 26 ∗ elevata Kim S. H., Kim H. J., Shin & Jo a o 6.8 – 7.4 25 – 26 elongata Reverdin o c c a o o o o o 6.5 – 7.4 24 – 26 eoa Takahashi in Asmund & Takahashi c 6.8 25 favosa Nicholls c c 7.1– 7.4 26 grata Takahashi in Asmund & Takahashi c c c a a c c 6.3– 7.4 18 – 26 guttata var. simplex Nicholls a 7.1 26 heterospina Lund o o o a a o 6.3 – 7.5 16 – 24 ∗ hexareticulata Jo, Shin, Kim, Siver & Andersen c 7.2 24 matvienkoae (Matvienko) Asmund & Kristiansen o o a o o o a s s o c c c 6.2– 7.4 18 – 26 mangofera Harris & Bradley o c o o 6.2– 7.4 22 – 26 mangofera var. foveata (D ü rrschmidt) Kristiansen o o o o 6.3 – 7.1 18 – 26 mangofera var. gracilis (D ü rrschmidt) Kristiansen o 6.8 25 mangofera var. reticulata (Cronberg) Kristiansen c 7.2 22

883 Continued water bodies by means of transmission electron microscopy C) ° ( 22 26 24 25 26 25 25 25 (Table 2). None of the taxa found in the Hong Kong Special – – – – – – – – Administrative Region have previously been recorded in the region. Forty of the taxa have previously been found in China, but Spiniferomonas cornuta , Paraphysomonas sp., 7.5 18 7.2 24 7.4 18 7.27.2 18 22 7.4 22 7.4 18 7.5 16 Mallomonas hexareticulata , M . ceylanica , M . plumosa , M . – – – – – – – – 6.9 6.8 24 ocellata , M . elevata and M . spinosa are reported for the fi rst time in China. A new species, Paraphysomonas hainanensis Wei & Kristiansen, was found in a pond in Haikou City, Hainan Province. c 6.5 In the Chrysophyceae, in family Paraphysomonadaceae, twelve species belonging to three genera ( Chrysosphaerella , Spiniferomonas and Paraphysomonas ) were found (Table 2). o 6.9 25 In Chrysosphaerella , C . coronacircumspina (Fig. 5) was found in seventeen localities with pH 6.2– 7.5 and water temperature 21– 26 ° C. Th is species was the dominant a 7.4 26 o 7.4species in 26 fi ve water bodies. Of Chrysosphaerella longispina , only a short spine scale and a large elliptical plate scale (Fig. 12) were collected in a pond of Xinglong County o 7.1 26 with pH ϭ 7.2 and a water temperature of 22 ° C. Th is species is especially abundant in clear water lakes and ponds in the northern temperature region and occurred often in warmer summer waters (Siver 1993). During the present study it was, however, found in a tropical region. Chrysosphaerella annulata (Fig. 6) was found in two ponds, two lakes and three reservoirs with pH 6.2– 7.2 and water temperatures of 21 – 26° C, and constituted the dominant species in four water bodies. Chrysosphaerella septispina (Fig. 7) was found only in the Xili Reservoir with pH ϭ 7 . 4 and a water temperature of 25° C. It was previously reported from Quanzhou City and Wanning City, China (Wei and Kristiansen 1998, Wei and Yuan 2001). Within the genus Spiniferomonas , S . cornuta (Fig. 16 – 17) was found in three ponds with pH 6.3 – 7.1 and a water temperature of 18° C. Within the genus Paraphysomonas , P . runcinifera (Fig. 8) was found in the Shabo Reservoir cwith pH ϭ o 6.2 and c a water temperature of 22° C. It was 7.1 previously reported from Wuhan City, China (Kristiansen and Tong 1989a). Paraphysomonas vestita (Fig.13) was found in eighteen localities and has very wide pH and water temperature tolerances. It was the dominant species in four water bodies. Paraphysomonas sp. (Fig. 9– 10) was found in fi ve reservoirs with pH 6.5 – 7.5 and water temperatures of 24 – 25 ° C. Th e scales of this taxon closely resembled scales of P . vestita , but diff ered by the presence of 6– 8 radial ridges on the base plate. Similar scales with 5– 7 radial ridges on á c c s c o o c o o a c o o c c c

Hayakawa c the base plate have previously been reported from Japan by & lov lov

č Takahashi (1976, 1978) as a taxon with the provisional

s c a c c sdesignation ‘P . vestita form. no. 1’ , and 6.3 from Madagascar Kyn Kyn &

terﬁ by Hansen (1996) as ‘P . vestita ssp.’ . Ř ez á č ov á and Š kaloud é Takahashi Takahashi P (2004) found the taxon in Ireland, with 6– 7 radial ridges on & Kristiansen

kaloud the base plate. In addition to these, a new species was found & Š Croome

Asmundas described below. o c o & Hansen) Asmund Hansen) s o o a a o o c c 6.3 & Tyler

& According to new taxonomical studies, the Synurales is & Bradley Tyler terﬁ rrschmidt o & nested within the class Chrysophyceae ( Š kaloud et al. bangladeshica é

& ü P 2013). In the genus Synura only two species, S . curtispina rrschmidt rrschmidt D Powse c o 6.8 24 ü var. var. (Harris) Momeu Korshikov o a a c 6.3 (G. S. West) Playfair em. Croome, Playfair West) (G. S. (Petersen (Petersen Croome Korshikov em. Korshikov (Fig. 76– 77) and S . petersenii (Fig. 78– 79) were found. In Teiling em. KriegerTeiling c c o o c o c a c a c c 6.3 rrschmidt s s s s o o 6.3 D Harris Gusev em. Wei Wei Gusev em. ü the genus Chrysodidymus , C . synuroideus (Fig. 80) was found D rrschmidt rrschmidt

ü in Xiao Lake and Bofulincum Reservoir with pH 6.8 and a D spinosa plumosa ocellata ° ° punctifera rasilis portae-ferreae ∗ peronoides ∗ peronoides multisetigera ∗ papillosa Table 2. Continued. Table Locations Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 pH Temp. splendens tonsurata Synuraceae Synura curtispina petersenii Chrysodidymus synuroideus water temperature of 24 C and 26 C, respectively.

884

Figure 2 – 4. Paraphysomonas hainanensis, whole cell, plate scales and spine (2). Figure 5: Chrysosphaerella coronacircumspina , whole cell (22). Figure 6: C . annulata , plate scales (7). Figure 7: C . septispina , whole cell (19). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1.

Twenty nine species and ﬁ ve varieties were found in the (Fig. 19 – 20) was found in a pond with pH ϭ 7.2 and a water genus Mallomonas . Mallomonas caudata (Fig. 18) and temperature of 24° C, and M . elevata (Fig. 51 – 52) was found M . matvienkoae (Fig. 21) were found in fourteen and in the Donghu Reservoir and the Meisha Reservoir with thirteen localities, respectively, and were sometimes domi- pH ϭ 6.8 to 7.4 and water temperatures of 25– 26 ° C. Th e nant in these water bodies. Mallomonas hexareticulata two species have recently been described by Jo et al. (2013)

885 Figure 8. Paraphysomonas runcinifera , plate scales (1). Figure 9 – 10: P. sp., spines (16). Figure 11: Spiniferomonas trioralis , spine and plate scale (4). Figure 12: Chrysosphaerella longispina , spine and plate scales (6). Figure 13: Paraphysomonas vestita , spine (6). Figure 14: P . imperforata , spine (5). Figure 15: P. gladiata , spine (16). Figure 16 – 17: Spiniferomonas cornuta , whole cell, plate scales (13). Scale bar ϭ 1 μ m, Th e numbers in parenthesis refer to site locations listed in Table 1. and Kim et al. (2014) from Jeju Island, South Korea where was the dominant species in two water bodies. Mallomonas the climate is generally subtropical. Mallomonas peronoides peronoides var. bangladeshica (Fig. 24 – 25) was found in a (Fig. 22 – 23) was found in four ponds and two lakes with pond and in the Shilibei Reservoir with pH ϭ 6.5 – 7.2 pH ϭ 6.3 – 7.2 and water temperatures of 18 – 24° C, and it and water temperatures of 22 – 25° C. Mallomonas ceylanica

886 Figure 18. Mallomonas caudata , whole cell (13). Figure 19 – 20: M . hexareticulata , body scale (11). Figure 21: M. matvienkoae , body scale (20). Figure 22– 23: M . peronoides, apical scale, body scale (6). Figure 24– 25: M . peronoides var. bangladeshica, body scales (2). Figure 26: M . ceylanica , body scale (2). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1.

(Fig. 26) was found in a pond with pH ϭ 7.2 and water described in Ontario, Canada (Nicholls 1989), and later temperatures of 22° C. Mallomonas guttata var. simplex reported from China’ s Fujian Province (Wei and Kristiansen (Fig. 29) was discovered in the Shatoujiao Reservoir with 1998). Th e present ﬁ nding thus represents the third record. pH 7.1 and a water temperature of 26 °C. It was originally Mallomonas splendens (Fig. 35) was found in the Donghu

887 Reservoir with pH ϭ 7.4 and a water temperature of proximally developed as a v-rib. Shield covered by a dense 26 ° C. It was previously discovered using light microscopy reticulation of fused papillae, but in some cases the apical in Nanren Lake and in a pond near Nanren Lake (Taiwan part only with papillae. Th e most remarkable feature is the Province) (Yamagishi 1992), and reported from the development of a rigid spine from the central part of one Xinchang County, China (Wei and Yuan 2013). Only body branch of the submarginal rib, in some cases only rudimen- scales of Mallomonas plumosa (Fig. 34) were found in tary, but typically as a strong spine oriented perpendicularly Datan Reservoir with pH ϭ 6.9 and a water temperature of to the scale axis; up to about 1.5 μ m long and giving it a 25 ° C. Mallomonas ocellata (Fig. 73– 75) was found only in triangular shape. Th e spines are best developed in the ante- the Donghu Reservoir with pH ϭ 7.4 and a water tempera- rior part of the cell. Th e submarginal rib is developed as a ture of 26° C. Mallomonas mangofera var. reticulata v-rib with its angle almost in the centre of the scale, with a (Fig. 56 – 57) was found in a pond with pH ϭ 7.2 and a very broad posterior fl ange, with reticulation and proximally water temperature of 22° C. It has previously been reported with a patch of pores. Th e posterior border (rim) is well from Wuxi City, China (Wei and Yuan 2013). Mallomonas developed, with distinct struts. Posterior scales are not dif- grata (Fig. 60– 61) was found in a pond and six reservoirs ferentiated, perhaps slightly smaller than body scales. Bristles with pH ϭ 6.3 – 7.4 and water temperatures of 18 – 26 ° C, (Fig. 63, 68) curved, smooth, strongly tapering acute, shorter and was the dominant species in two water bodies. than the cell, 14 – 16 μ m long. Mallomonas spinosa was described from a pool, a pond and a reservoir in Vietnam (Gusev 2012) on the basis of Paraphysomonas hainanensis Wei & Kristiansen sp. samples observed using a scanning electron microscope nov. (Fig. 2 – 4) (SEM). Here we report a rich population of Mallomonas spinosa (Fig. 62 – 72) in a pond (pH ϭ 7.2, 22° C) in the Holotype: TEM grids deposited at the Herbarium of algae, Xinlong County, Hainan Province. Specimens of this spe- Hydrobiology Museum, Institute of Hydrobiology, Chinese cies were also found in the Tiegang Reservoir and the Academy of Sciences, China, HN0112 (Fig. 2, the holo- Shatoujiao Reservoir (pH ϭ 7.4, 25° C and pH ϭ 7.1, 26° C, type). Isotype: TEM grids deposited at the Herbarium of respectively). Th e water temperature and pH were algae, Hydrobiology Museum, Institute of Hydrobiology, comparable to three Vietnamese localities (the average Chinese Academy of Sciences, China, HN20100112. annual temperature 25– 28 ° C, pH range ϭ 6.1 – 7.4). Th e type locality is an unnamed pond, Haikou City, ° ′ ″ ° ′ ″ Mallomonas spinosa inhabits both eutrophic and olig- Hainan Province, China (20 0 55.62 N, 110 19 2.22 E , ϭ ° otrophic waters with wide range of pH and higher water pH 7.2, water temperature 22 C, electrical conductivity μ Ϫ1 temperature. In Gusev ’ s original description, the species 250.3 S cm ), 12 Jan 2010. has domeless apical scales without spines and is referred to section Papillosae. But in Fig. 62 and 63 micrographs Etymology of a whole cell are now provided, and the dome-bearing Th e specifi c epithet ‘hainanensis ’ refers to the fact that the apical collar scales and bristles are illustrated. Gusev ’ s name species was fi rst found in Hainan Province, China. for the species must have priority, but because the species Description was only incompletely described, our description adds ϫ important details, and the description is emended. In adi- Th e cell (Fig. 2) is spherical to ellipsoid, 4.2 – 4.5 4.3 – 4.7 μ m. Two unequal fl agella, the longer approximately tion, it is evident that M . spinosa cannot belong to the sec- μ μ tion Papillosae. Th e arrangement and structure of the apical 6 m long, the shorter 2.5 m long. Cell body covered collar scales are reminding of those found in sect. Torqua- with two types of scales. Elliptical plate scales (Fig. 3) are 1.2 – 1.4 μm long and 0.8– 1.1 μ m wide, with a wide mar- tae, ser. Mangoferae. But the body scales are unique and μ defi nitely quite diff erent from any found in sect. Torquatae. ginal rim (0.2 – 0.23 m) of radiating ribs and a large ellipsoid lacuna in the centre. Spine scales (Fig. 4) are A lateral spine is not known from any other Mallomonas μ species in any of the existing series. 4.2 – 5.4 m long, and consist of a saucer-shaped base that supports three long and narrow wings, and fused along a median thickened rib that extends from the base to the apex. Mallomonas spinosa Gusev (2012), emend. Wei & Kristiansen (Fig. 62 – 72) Similar species Paraphysomonas hainanensis is rather similar to P. acantholepis Cells (Fig. 62 – 63) are ovoid or ellipsoidal, 18 – 24 ϫ μ Preisig & Hibberd, but diff ers from it by the ornamental 10 – 12 m. Apical collar scales (Fig. 62– 64, 66) are six in structure of the plate scales and long spine scales. number, asymmetrically sausage shaped, dome bearing, 5.8 – 6.2 ϫ 3.4 – 3.5 μm. Dome rounded rectangular, smooth, but with a small area covered with reticulations. Scale with Biogeographical considerations interrupted submarginal rib; shield with reticulation, but distally with papillae and proximally with a group of pores. Th e localities in this study were situated in the southern Proximal border broad, with struts. tropical and subtropical China. Characterization of the Body scales (Fig. 65, 67, 69– 72) domeless, ovoid, southern Chinese silica-scaled chrysophyte fl ora is based 4.5 – 6.2 ϫ 2.4 – 3.4 μ m. Anterior fl ange with a row of papil- on relative occurrences of the species. Most of the species lae, with a strong submarginal rib totally surrounding the found in this study are cosmopolitan and/or widely distri- scale apex, apical part of it supported internally by struts, buted in the world. Chrysosphaerella coronacircumspina ,

888 Figure 27. Mallomonas papillosa, body scales (18). Figure 28: M . rasilis , body scale (11). Figure 29: M . guttata var. simplex , body scales (18). Figure 30 – 31: M . heterospina , apical scale with bristle, body scale with bristle (13). Figure 32– 33: M . punctifera, apical scales, body scale (12). Figure 34: M . plumosa , body scale (22). Figure 35: M . splendens , body scale (20). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1.

889 Figure 36 – 37. Mallomonas akrokomos , apical scale, body scale (12). Figure 38: M . tonsurata , body scale (24). Figure 39: M . areolata , body scale (13). Figure 40: M . elongata , body scale (10). Figure 41: M . alpina , body scale with bristle (14). Figure 42– 43: M . corymbosa , dome-bearing body scale, domeless body scale (5). Figure 44: M . multisetigera , body scale (8). Scale bar ϭ 1 μm. Th e numbers in parenthesis refer to site locations listed in Table 1.

890 Figure 45 – 46. Mallomonas portae-ferreae, body scales (10, 11). Figure 47: M . mangofera var. foveata , body scales (12). Figure 48: M . crassisquama , body scale (12). Figure 49: M . costata , apical scale (2). Figure 50: M . eoa , collar and body scale (23). Figure 51 – 52: M . elevata , collar scale, rear scale (20, 21). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1.

891 Figure 53. Mallomonas acaroides , apical scale (16). Figure 54: M . mangofera , body scale (20). Figure 55: M . mangofera var. gracilis , body scale (23). Figure 56– 57: M . mangofera var. reticulata , collar scales, body scale (6). Figure 58 – 59: M . favosa , collar scale, body scale (20). Figure 60 – 61: M . grata , collar scale, body scales (17, 4). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations list in Table 1.

Spiniferomonas trioralis , Paraphysomonas vestita , Mallomonas in more than nine localities) in habitats including ponds, caudata , M . matvienkoae , M . elongata , M . tonsurata , lakes and reservoirs covering very wide pH and water M . corymbosa , M . crassisquama , Synura curtispina and temperature ranges. Th ese species sometimes constituted the S . petersenii have highest appearance frequency (each found dominant species of these water bodies. Th is result is in

892

Figure 62 – 65. Mallomonas spinosa , whole cell, collar scales, body scales, rear scales (6). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1.

agreement with previous investigations (Takahashi 1978, Mallomonas spinosa which was discovered for the second Siver 1991, Kristiansen 2001, 2002, Kristiansen and Preisig time in southern China after it was ﬁ rst reported from 2007). Th e species composition in this region also includes Vietnam in southeast Asia (Gusev 2012). Mallomonas species with more or less scattered occurrences. Mallomonas ceylanica was previously discovered only in India and Sri splendens and Chrysodidymus synuroideus have scattered Lanka of south Asia, M . peronoides var. bangladeshica is global occurrence, and M . grata is scattered in east Asia, mainly distributed in the tropical and subtropical regions North and Central America (Kristiansen and Preisig 2007). (North and South America, Asia, Africa and Europe), Species considered to be restricted to temperate regions also M . ocellata was previously reported from east and southeast occurred, e.g. Mallomonas punctifera is distributed in Asia, M . plumosa was previously found scattered in the northern temperate regions, and M . corymbosa is south Asia, Australia and New Zealand (Dü rrschmidt and distributed in the temperate regions of the Northern and Cronberg 1989, Kristiansen 2001, Kristiansen and Preisig Southern Hemispheres (Kristiansen 2002, Kristiansen and 2007). Mallomonas mangofera var. reticulata has previously Preisig 2007). been found in tropical regions, including South America, Species restricted to tropical and subtropical regions Asia-India and Africa (Hansen 1996, Kristiansen and were also found in the present study. An example is Preisig 2007), but the variety has also been recorded in

893 Figure 66 – 72. Mallomonas spinosa , collar scale, body scales, bristle, body scales (6, 18). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1. the temperate Aquitaine region, France (Ně mcov á et al. pical or subtropical regions or during warmer seasons in 2012). Chrysosphaerella annulata , Mallomonas peronoides , temperate regions. Mallomonas hexareticulata and M . elevata M . portae-ferreae and M . mangofera var. foveata are cosmo- was recorded for the second time in southern China after it politan or widely globally distributed, but mainly in tro- was ﬁ rst reported from South Korea (Jo et al. 2013, Kim

894 Figure 73 – 75. Mallomonas ocellata , whole cell, body scale, rear scale (20). Figure 76 – 77: Synura curtispina , whole cell, body scale (13). Figure 78 – 79: S . petersenii , body scale, rear scale (9, 15). Figure 80: Chrysodidymus synuroideus, body scales with short spine (23). Scale bar ϭ 1 μ m. Th e numbers in parenthesis refer to site locations listed in Table 1.

895 et al. 2014). It is evident that the fl ora and geographical Kristiansen, J. and Tong, D. 1988. Silica-scaled chrysophytes distribution of the silica-scaled chrysophytes from this part of Wuhan, a preliminary note. – J. Wuhan Bot. Res. 6: of China is similar to that of east Asia (Japan, South Korea), 97 – 100. southeast Asia and south Asia (India, Sri Lanka, Vietnam Kristiansen, J. and Tong, D. 1989a. Studies on silica-scaled chrysophytes from Wuhan, Hanzhou and Beijing, P. R. China. and Malaysia). – Nova Hedw. 49: 183– 202. Kristiansen, J. and Tong, D. 1989b. Chrysosphaerella annulata n. sp., a new scale-bearing chrysophyte. – Nord. J. Bot. 9: Acknowledgements – Th is research was funded by A Major Direction 329 – 332. Project of the Knowledge Innovation Program of the Chinese Kristiansen, J. and Tong, D. 1991. Investigations on silica- Academy of Sciences (to Wei Yin-Xin, No. KSCX2-EW-Z-9). We scaled chrysophytes in China. – Verh. Int. Verein. 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