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The diatom genus Gomphonema Ehrenberg in India: Checklist and description of three new species
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B. Karthick 1,    J.P. Kociolek 2*,    M.K. Mahesh 3    and    T. V. Ramachandra 1
1 Energy & Wetlands Research Group, Centre for Ecological Sciences, Indian Institute of Science, Bangalore – 560 012, India
2 Museum of Natural History and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO, USA
3 Department of Botany, Yuvaraja’s College, University of Mysore, Mysore - 570 005, India
*Corresponding author, email: patrick.kociolek@colorado.edu

INTRODUCTION

The diatom genus Gomphonema Ehrenberg is large, including over 500 taxa worldwide (Fourtanier & Kociolek 2009). While its members are almost exclusively freshwater in terms of habitats, and many species are associated with impacted trophic levels (e.g. Patrick in Patrick & Reimer 1966), the genus is quite diverse morpho-logically. Variation in valve ultrastructure includes presence or absence of stigmata, areolar structure, structure and position of the apical pore fields, and presence/absence of spines to name a few (Kociolek & Stoermer 1993, but also see Metzeltin & Lange-Bertalot 1998, Reichardt 1999). While some species of Gomphonema appear to be cosmopolitan in their distribution (for example, it appears that G. parvulum (Kützing) Kützing has been reported from all continents on earth), there are many reports of endemic species of Gomphonema from South America (Fricke 1904; Metzeltin & Lange-Bertalot 2007), Africa (Compère 1995, Hustedt 1949, Kociolek & Stoermer 1991), Madagascar (Spaulding & Kociolek 1998), North America (Kociolek & Kingston 1999, Thomas et al. 2009), Europe (Hustedt 1945, Reichardt 1999, 2005), Asia (Lange-Bertalot & Genkal 1999, Li et al. 2006) and Australia and environs (Hustedt 1942, Kociolek et al. 2004).

India is a large, geographically complex country with ten different biogeographic zones covering over 3 million sq. km, about a third the size of all of Europe, but with nearly 50% greater human population (World Population Prospects, 2008). The country shares borders with Bangladesh, Bhutan, Myanmar, China, Nepal and Pakistan. The complexity of India’s freshwater environments includes three major, complex watersheds (Himalayas and the Karakoram ranges; Vindhya, Satpura ranges and the Chota Nagpur Plateau; and the Western Ghats) that cut across the country, related to their sources in the mountains. Freshwater environments range in elevation from sea level at the coast to over 8000 m above sea level.

The diatom genus Gomphonema in India has been documented for over 160 years. First report of the genus in India was by Ehrenberg (1845), who reported G. clavatum, G. gracile and G. turris from "Kolkatta" (then Calcutta) and River Ganges. Since then, 49 separate papers have identified 130 taxa of Gomphonema from India, mostly from lowland habitats. Of these, about 30% have been newly described taxa. In the region around India, gomphonemoid diatoms have been considered by Hustedt (1922), Jüttner et al. (2004) from Nepal, and Mereschkowsky (1906), Kociolek (1992), and Li et al. (2006, in press) from China. Though many species of Gomphonema have been described from India, there are extensive parts of the country that have not been investigated. The Western Ghats mountain range in Southern India, one of the Gondwanaland breakup landmasses, is amongst the most important hotspots of biodiversity. Western Ghats is a chain of mountains that runs parallel with the west coast of India for over 1600 km from 8°15'N to 21°00'N. Due to its distinct physiographic, edaphic and climatic gradients, this region harbors a wide array of habitats with unique geographic position and that support unique sets of flora and fauna. The Western Ghats harbors approximately 5000 species of vascular plants belonging to nearly 2200 genera; about 1700 species (34%) are endemic. There are also 58 endemic plant genera, while nearly three-quarters of the endemic genera have only a single species (Conservation International, 2008). The fauna and flora of this region has attracted attention of systematic and evolutionary biologists because of the mixture of high-level endemism (Inger, 1999; Myers et al. 2000) and various affinities with other biogeographic regions (Bossuyt & Milinkovitch, 2001).

In the present report we have compiled a checklist of the Gomphonema taxa previously reported and described from India (Table 1). We also document and describe with light and scanning electron microscopy three new species of Gomphonema from Central Western Ghats, Karnataka State, India. We also present information about each species' autecology, with particular attention paid to water chemistry and seasonality.

Table 1. Listing of Gomphonema taxa for India and references that have reported each taxon.
(Note: * denotes the species described from Indian administrative boundary; † indicates the studies on fossil material)

  Species Name Reference
1 Gomphonema abbreviatum Kutz. Abdul-Majeed, 1935; Srinivasan, 1965; Suxena and Venkateshwarlu, 1970
2 Gomphonema abbreviatum Kutz. f. minor? Krishnamurthy* Krishnamurthy, 1954; Kumawat et al., 2008
3 Gomphonema abbreviatum Kutz. v. pulneyensis? Krishnamurthy* Krishnamurthy, 1954
4 Gomphonema acuminatum  Ehr. Carter, 1926 ; Biswas, 1936 ; Krishnamurthy, 1954
5 Gomphonema acuminatum Ehr. v. elongata (Rabh.) W.Sm. Biswas, 1936
6 Gomphonema acuminatum Ehr. v. directum A.Cl. Gandhi, 1959b
7 Gomphonema acuminatum Ehr. v. turris (Ehr.) Cl. Krishnamurthy, 1954; Gandhi, 1960a; Sarode and Kamat, 1980; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
8 Gomphonema aequatoriale Hust. Gandhi, 1960a; Gandhi, 1964; Sarode and Kamat, 1980; Sarode and Kamat., 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
9 Gomphonema angustatum (Kutz.) Rabh. Krishnamurthy, 1954; Srinivasan, 1965; Kumawat et al., 2008
10 Gomphonema angustatum (Kutz.) Rabh. v. productaGrun Krishnamurthy, 1954; Kumawat et al., 2008
11 Gomphonema angustatum (Kutz.) Rabh. v. producta Grun. f. indica Gandhi* Gandhi, 1960a; Sarode and Kamat, 1984
12 Gomphonema apicatum Ehr. Singh, 1963
13 Gomphonema augur Ehr. West and West, 1907; Gandhi, 1959a; Gandhi, 1960a; Singh, 1961; Gandhi, 1962a; Gandhi, 1966; Gandhi, 1983a†; Sarode and Kamat, 1984; Kumawat et al., 2008
14 Gomphonema augur Ehr. v. gautieri V.H Gandhi, 1983a†
15 Gomphonema augur Ehr. v. genuinum May. Gandhi, 1956; Singh, 1961; Singh, 1963; Kumawat et al., 2008
16 Gomphonema balatonis Pant. Gandhi, 1960a; Srinivasan, 1965; Sarode and Kamat, 1984; Kumawat et al., 2008
17 Gomphonema balatonis Pant. v. lanceolata Gandhi Gandhi, 1960a; Sarode and Kamat, 1984;
18 Gomphonema brasiliense Grun. deToni J.B., 1891-94; Srinivasan, 1965
19 Gomphonema bengalensis Grun* deToni J.B., 1891-94;
20 Gomphonema bohemicum Reichelt et Fricke Gandhi, 1983a†
21 Gomphonema capitatum Ehr. Abdul-Majeed, 1935
22 Gomphonema clavatoides Gandhi* Gandhi, 1960a; Gandhi, 1964; Srinivasan, 1965; Gandhi, 1966; Sarode and Kamat, 1984; Gandhi, 1998; Kumawat et al., 2008
23 Gomphonema clavatoides Gandhi v. valida Gandhi* Gandhi, 1960a; Sarode and Kamat, 1984
24 Gomphonema clavatoides v. rostrata Gandhi* Gandhi, 1998
25 Gomphonema clavatum Ehr. Ehrenberg, 1845
26 Gomphonema clevei f. acuta Gandhi* Gandhi, 1966
27 Gomphonema clevei Fricke Gandhi, 1959b; Srinivasan, 1965; Gandhi, 1966
28 Gomphonema clevei v. bipunctata Gandhi* Gandhi, 1959b; Gandhi, 1966
29 Gomphonema clevei v. javanica Hust. Gandhi, 1966
30 Gomphonema clevei v. undulata Gandhi* Gandhi, 1966
31 Gomphonema constrictum Ehr. Abdul-Majeed, 1935; Biswas, 1936; Srinivasan, 1965; Gandhi, 1983a†; Kumawat et al., 2008
32 Gomphonema constrictum Ehr. v capitata (Ehr.) Cl. Gandhi, 1960a; Gandhi, 1983a†; Sarode and Kamat, 1984; Kumawat et al., 2008
33 Gomphonema constrictum Ehr. v. indica Gandhi* Gandhi, 1960a; Sarode and Kamat, 1984; Kumawat et al., 2008
34 Gomphonema constrictum Ehr. v. capitatum Cleve. f. italica Kuetz. Kumawat et al., 2008
35 Gomphonema constrictum Ehr. v. capitatum Cleve. f. turgida Mayer Kumawat et al., 2008
36 Gomphonema dharwarensis Gandhi* Gandhi, 1956; Srinivasan, 1965
37 Gomphonema dichotomum Kutz. Ehrenberg, 1845; Grunow, 1865; Schmidt, 1935
38 Gomphonema dubia Meister Meister, 1932
39 Gomphonema dubravicense Pant. Gandhi, 1998
40 Gomphonema geminatum Ag. v. hybrida Grun. Gandhi, 1983a†
41 Gomphonema ghosea Abdul-Majeed* Abdul-Majeed, 1935
42 Gomphonema gracile Ehr. Ehrenberg, 1845; Carter, 1926; Schmidt, 1935;
Krishnamurthy, 1954; Gandhi, 1955; Gandhi, 1957a; Gandhi, 1959c; Gandhi, 1960a; Gandhi, 1962a; Singh, 1962; Singh, 1963; Gandhi, 1966;
Srinivasan, 1965; Sarode and Kamat, 1984; Gandhi, 1998; Kumawat et al., 2008
43 Gomphonema gracile Ehr. f. turris (Ehr.) Hust. Sarode and Kamat, 1984
44 Gomphonema gracile Ehr. v. auritum A. Br. Gandhi, 1960a; Thomas and Gonzalves, 1965; Gandhi, 1966; Sarode and Kamat, 1983; Sarode and Kamat, 1984
45 Gomphonema gracile Ehr. v. dichotomum (W . Smith)Cleve Thomas and Gonzalves, 1965
46 Gomphonema gracile Ehr. v. frickei Gandhi* Gandhi, 1960a; Sarode and Kamat, 1984
47 Gomphonema gracile Ehr. v. hybridum A.Cl. Sarode and Kamat, 1984
48 Gomphonema gracile Ehr. v. intricatiforme May. Sarode and Kamat, 1983; Sarode and Kamat, 1984
49 Gomphonema gracile Ehr. v. lanceolata (Kutz.) Cl. Krishnamurthy, 1954; Gandhi, 1960a; Sarode and Kamat, 1984; Kumawat et al., 2008
50 Gomphonema gracile Ehr. v. major Grun. Thomas and Gonzalves, 1965; Sarode and Kamat, 1984; Kumawat et al., 2008
51 Gomphonema gracile Ehr. v. naviculoides (W. Sm.) Grun. Gandhi, 1960a; Gandhi, 1962b; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
52 Gomphonema gracile Ehr. v. subcapitata Gandhi* Sarode and Kamat, 1984; Gandhi, 1960a; Kumawat et al., 2008
53 Gomphonema grovei M.S Gandhi, 1983a†; Gandhi, 1998†
54 Gomphonema grovei M.S v. conspicua Gandhi et al. * Gandhi, 1983a†; Gandhi, 1998†
55 Gomphonema grovei M.S v. lanceolata Gandhi et al. * Gandhi, 1983a†; Gandhi, 1998†
56 Gomphonema grovei M.S v. rhomboidea Gandhi et al. * Gandhi, 1983a†; Gandhi, 1998†
57 Gomphonema hebridense Ehr. Gandhi, 1970; Sarode and Kamat, 1980 ;
Sarode and Kamat, 1983 ; Sarode and Kamat, 1984; Kumawat et al., 2008
58 Gomphonema intermedium Hust. Prasad and Srivastava, 1986
59 Gomphonema intricatum Kutz. West and West, 1907; Abdul-Majeed, 1935; Krishnamurthy, 1954; Gandhi, 1958a; Gandhi, 1960a; Gandhi, 1983a†; Gandhi, 1983b†; Sarode and Kamat, 1984; Gandhi, 1998†
60 Gomphonema intricatum Kutz. v. dichotoma (Kutz.) Grun Gandhi, 1983a†; Gandhi, 1983b†; Gandhi, 1998†
61 Gomphonema intricatum Kutz. v vibrio Cl. f. subventricosa Gandhi* Gandhi, 1962b
62 Gomphonema intricatum Kutz. v. bohemicum (Reich. et Fricke) A.Cl. Gandhi, 1958a; Sarode and Kamat, 1984
63 Gomphonema intricatum Kutz. v. fossile Pant. Sarode and Kamat, 1980; Sarode and Kamat, 1983; Sarode and Kamat, 1984
64 Gomphonema intricatum Kutz. v. pumila Grun. Singh, 1962; Sarode and Kamat, 1984
65 Gomphonema intricatum Kutz. v. pusillum May. Sarode and Kamat, 1983; Gandhi, 1970
66 Gomphonema intricatum Kutz. v. vibrio (Ehr.) Cl. Venkataraman, 1939; Gandhi, 1959b; Gandhi, 1960a; Sarode and Kamat, 1980; Sarode and Kamat, 1984; Kumawat et al., 2008
67 Gomphonema lacus-rankala Gandhi* Gandhi, 1958a; Gandhi, 1964; Srinivasan, 1965; Sarode and Kamat, 1984; Gandhi, 1998
68 Gomphonema lacus-rankala Gandhi v. gracilis Gandhi* Gandhi, 1962b; Gandhi, 1964; Gandhi, 1967; Gandhi, 1970; Sarode and Kamat, 1980; Sarode and Kamat, 1984; Gandhi, 1998; Kumawat et al., 2008
69 Gomphonema lacus-rankala v. chandolensis Gandhi* Gandhi, 1964; Gandhi, 1998; Kumawat et al., 2008
70 Gomphonema lacus-rankala Gandhi v. robusta Gandhi* Gandhi, 1958a; Sarode and Kamat, 1980; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Gandhi, 1998; Kumawat et al., 2008
71 Gomphonema lapponicum A.Cleve Kumawat et al., 2008
72 Gomphonema lanceolatum Ehr. Grunow, 1865; Carter, 1926; Venkataraman, 1939; Gandhi, 1958a; Gandhi, 1959c; Gandhi, 1960a; Gandhi, 1960b; Gandhi, 1962a; Gandhi, 1962b; Gandhi, 1964; Gandhi, 1966; Gandhi, 1967; Gandhi, 1983a†; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
73 Gomphonema lanceolatum Ehr.  v. insingis (Greg.) Cl. Venkataraman, 1939; Gandhi, 1955; Gandhi, 1960a; Gandhi, 1966; Sarode and Kamat, 1983 ;
Sarode and Kamat, 1984;
74 Gomphonema lanceolatum Ehr. f. turris (Ehr.) Hust. Gandhi, 1959b
75 Gomphonema lanceolatum Ehr. v. affine (Kutz.) A.Cl. Gandhi, 1957b; Gandhi, 1960b; Sarode and Kamat, 1984
76 Gomphonema lingulatum Hust. Gandhi, 1960a; Sarode and Kamat, 1984
77 Gomphonema longiceps Ehr. Krishnamurthy, 1954
78 Gomphonema longiceps Ehr. v. subclavata Grun. Krishnamurthy, 1954; Sarode and Kamat, 1983
79 Gomphonema longiceps Ehr. v. subclavata Grun. f. gracilis Venkataraman* Venkataraman, 1956
80 Gomphonema macropunctatum Krishnamurthy* Krishnamurthy, 1954; Srinivasan, 1965
81 Gomphonema magnifica Gandhi* Gandhi, 1960a; Srinivasan, 1965; Sarode and Kamat, 1984; Kumawat et al., 2008
82 Gomphonema magnifica Gandhi v. rhomboida Gandhi* Gandhi, 1960a; Sarode and Kamat, 1984
83 Gomphonema major A. Cl. F. unipuncta A. Cl. Thomas and Gonzalves, 1965
84 Gomphonema martini Fricke Sarode and Kamat, 1984; Gandhi, 1960a
85 Gomphonema moniliforme Gandhi* Gandhi, 1960a; Srinivasan K.S., 1965; Sarode and Kamat, 1984
86 Gomphonema montanum Schum Gandhi, 1960a; Gandhi, 1964; Sarode and Kamat, 1984
87 Gomphonema montanum Schum v. acuminatum May. Gandhi, 1956; Gandhi, 1960a; Gandhi, 1960b; Gandhi, 1964; Gandhi, 1967; Sarode and Kamat, 1980; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Gandhi, 1998; Kumawat et al., 2008
88 Gomphonema montanum Schum v. acuminatum May. f. indicum Sarode et Kamat* Sarode and Kamat, 1984; Kumawat et al., 2008
89 Gomphonema montanum Schum v. acuminatum May. f. maharashtrensis Sarode et Kamat* Sarode and Kamat, 1980; Sarode and Kamat, 1984
90 Gomphonema nagpurense Sarode et Kamat* Sarode and Kamat, 1984
91 Gomphonema olivaceoides Hust. Sarode and Kamat, 1984
92 Gomphonema olivaceum (Lung.) Kutz. Biswas, 1936; Krishnamurthy, 1954; Gandhi, 1958a; Gandhi, 1960a; Singh, 1962; Singh, 1963
Sarode and Kamat, 1980; Gandhi, 1983b†; Sarode and Kamat, 1984;
93 Gomphonema olivaceum (Lyng.) Kutz. v. calcarea Krishnamurthy, 1954;
94 Gomphonema olivaceum (Lyng.) Kutz. v. balticum Cl. Krishnamurthy, 1954; Gandhi, 1956
95 Gomphonema olivaceum (Lyng.) Kutz. v. genuinum Mayer. f. minutula Mayer Kumawat et al., 2008
96 Gomphonema oregonicum Ehr. Grunow, 1865
97 Gomphonema parvulum (Kutz.) Grun. Grunow, 1865; Schmidt, 1935; Skvortzow, 1935;
Venkataraman, 1939; Gandhi, 1955; Gandhi, 1957a; Gandhi, 1958b; Gandhi, 1959a; Gandhi, 1959c; Gandhi, 1960a; Gandhi, 1960b; Singh, 1961; Gandhi, 1962a; Gandhi, 1966; Gandhi, 1967; Sarode and Kamat, 1980; Sarode and Kamat, 1983; Sarode and Kamat, 1984;
Gandhi, 1998; Kumawat et al., 2008
98 Gomphonema parvulum (Kutz.) Grun. v. lagenula (Grun.) Hust. Gandhi, 1960a; Gandhi, 1962b; Gandhi, 1966;
Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
99 Gomphonema parvulum (Kutz.) Grun. v. micropus (Kutz.) Cl. Gandhi, 1960a; Gandhi, 1960b; Gandhi, 1962b
Gandhi, 1966; Sarode and Kamat, 1983; Sarode and Kamat, 1984
100 Gomphonema parvulum (Kutz.) Grun. v. subellipticum Cl. Gandhi, 1956; Gandhi, 1957a; Gandhi, 1958b; Gandhi, 1960a; Gandhi, 1960b; Gandhi, 1966; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
101 Gomphonema parvulum (Kutz.) Grun. v. subcapitata V. H. Venkataraman, 1957
102 Gomphonema parvulum (Kutz.) v. exlissma Grun. Gandhi, 1959c; Gandhi, 1962b; Gandhi, 1964; Thomas and Gonzalves, 1965; Gandhi, 1967; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008; Gandhi, 1998
103 Gomphonema parvulum (Kutz.) V.H. v. genuinium May. Gandhi, 1956
104 Gomphonema punctatum Krasske Kumawat et al., 2008
105 Gomphonema saravanthense Gandhi* Gandhi, 1970
106 Gomphonema sphaerophorum Ehr. Krishnamurthy, 1954; Gandhi, 1957a; Gandhi, 1958b; Gandhi, 1959c; Gandhi, 1960a; Gandhi, 1964; Srinivasan, 1965; Gandhi, 1966; Sarode and Kamat, 1980; Sarode and Kamat, 1983; Sarode and Kamat, 1984; Kumawat et al., 2008
107 Gomphonema sphaerophorum Ehr. v. kolhapurense Sarode et Kamat* Sarode and Kamat, 1984; Kumawat et al., 2008
108 Gomphonema sphaerophorum Ehr. v. subcapitata Venkatraman* Venkataraman, 1939; Thomas and Gonzalves, 1965
109 Gomphonema sphaerophorum f. jogensis Gandhi* Gandhi, 1966
110 Gomphonema spiculoides Gandhi* Srinivasan, 1965; Gandhi, 1960a;  Sarode and Kamat, 1984
111 Gomphonema spiculoides Gandhi v. major Gandhi* Gandhi, 1960a; Sarode and Kamat, 1984
112 Gomphonema subapicatum Fiitsch et Rich v. okamurae (Skv) Gandhi Gandhi, 1960a; Sarode and Kamat, 1984
113 Gomphonema subapicatum Fritsch et Rich Abdul-Majeed, 1935; Gandhi, 1956; Gandhi, 1958a; Gandhi, 1960a; Gandhi, 1960b; Gandhi, 1962b; Gandhi, 1964; Gandhi, 1966; Gandhi, 1967; Sarode and Kamat, 1984 ; Gandhi, 1998
114 Gomphonema subcapitatum  v. curta Fritsch et Rich Abdul-Majeed, 1935
115 Gomphonema subclavatum Grun. Carter, 1926; Abdul-Majeed, 1935
116 Gomphonema submalayense Gandhi* Gandhi, 1970
117 Gomphonema substicature Fritsch v. stipitata Abdul-Majeed, 1935
118 Gomphonema subtile Ehr. Gandhi, 1958b; Gandhi, 1962a; Gandhi, 1966; Sarode and Kamat, 1984
119 Gomphonema subtile Ehr. v. malayensis Hust. Gandhi, 1960a; Gandhi, 1966; Gandhi, 1970; Sarode and Kamat, 1984
120 Gomphonema subventricosum Hust. Gandhi, 1962a; Gandhi, 1966; Sarode and Kamat, 1984
121 Gomphonema sumatrense Fricke Gandhi, 1960a; Sarode and Kamat, 1984
122 Gomphonema tenellum W.Sm. Dickie G, 1882; West and West, 1907; Gandhi, 1998
123 Gomphonema tenuis Gandhi* Gandhi, 1960a; Srinivasan, 1965; Sarode and Kamat, 1984
124 Gomphonema tergestinum (Grun.) Frickie Krishnamurthy, 1954
125 Gomphonema tropicale Brun Gandhi, 1959b
126 Gomphonema turris Ehr. Ehrenberg, 1845; Grunow, 1865
127 Gomphonema undulatum Hust. Gandhi, 1960a; Sarode and Kamat, 1984
128 Gomphonema varanasis Singh* Singh, 1961
129 Gomphonema vastum Hust v elongata Skv. Gandhi, 1958b
130 Gomphonema vidarbhense Sarode et Kamat* Sarode and Kamat, 1984

Citation: Karthick, B., J.P. Kociolek, M.K. Mahesh & T.V. Ramachandra (2011): The diatom genus Gomphonema Ehrenberg in India: Checklist and description of three new species. – Nova Hedwigia 93: 211–236.
Contact Address :
  Dr. T.V. Ramachandra
Energy & Wetlands Research Group,
Centre for Ecological Sciences, Indian Institute of Science, Bangalore – 560 012, INDIA.
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