taxonID	type	description	language	source
03E287B9D5567361FF12A423FA862ED7.taxon	materials_examined	Material studied Type host: Macruronus magellanicus Lönnberg, 1907 Site of infection: gall bladder	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D5567361FF12A423FA862ED7.taxon	description	Localities, dates and depths: (1) off southern Chile, 55 º 30´S, 71 º 30´W, October 2007, 350 m; (2) off Chiloe Island, Chile, 43 º 0 0 ˏS, 73 º 0 0 ˏW, June 2007, 300 m; (3) west of Falkland Islands, 51 º 00´to 52 º 30´S, 62 º 00´to 62 º 30´W, October 2007, 200 – 250 m; (4) west of Falkland Islands, 51 º 00´to 52 º 30´S, 62 º 00´to 62 º 30´W, October 2009, 200 – 250 m.	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D5567361FF12A423FA862ED7.taxon	materials_examined	Type locality: (1). Prevalence: (1) 40 % (4 of 9); (2) 64 % (16 of 23); (3) 23 % (7 of 30); (4) 57 % (24 of 42).	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D5567361FF12A423FA862ED7.taxon	description	Host length range: 25 – 42 cm. Collection numbers: NHMUK 2012.3.19.1, 2012.3.19.2, 2012.3.19.3. Morphological description. Sporoblast globular, disporic (Fig. 1). No pseudopodia observed. Ectoplasm dense and clearly demarcated from fine endoplasm. Dimensions, based on 10 fixed specimens: 28.4 – 38.4 x 28.0 – 36.0. Spore (Figs. 2 – 6) triangular with sharply pointed tips in side view, flat anteriorly and curved in apical view. Sutural line prominent and raised. Sporoplasm deeply staining and binucleate. Valves drawn out into two delicate broad alate processes joined together at their proximal extremities to form a parachute-like structure over the valves. Polar capsules spherical, close together. Polar filament thick, number of coils not visible. Dimensions, based on 30 fixed spores, as ranges with means ± SD in parentheses: spore length 8.0 – 10.5 (9.1 ± 0.68); spore thickness, including alate processes 30.0 – 35.5 (31.8 ± 1.52), excluding alate processes 8.8 – 11.6 (9.0 ± 1.2); spore width 14.0 – 20.0 (15.7 ± 1.57); polar capsule diameter 2.5 – 3.0 (2.7 ± 0.16); polar capsule length: spore length = 1: 2.8 – 3.8; spore length: spore width = 1: 3.0 – 4.9. Molecular results: Pseudalataspora kovalevae. PCR amplification of the myxosporean 18 S rRNA gene from four infected gall bladders from sample (4) above generated a product of approximately 1650 nucleotides in both of the samples processed from each fish. Forward and reverse 18 S rRNA gene sequences were obtained for PCR products from both samples from each individual host, and internal sequences from a single sample from each host. All sequences were identical. Two polymorphic positions were found. A final sequence of 1594 nucleotides was submitted to Genebank under Accession No. JX 467675. The sequence was novel with respect to previously generated data from myxosporean species. Pseudalataspora kovalevae 18 S rRNA gene sequence showed closest sequence identity to Ceratomyxa anko Freeman et al., 2008 (DQ 301510) and Ceratomyxa pantherini Gunter et al., 2010 (GU 136393) based on BLAST analysis with 92 % and 97 % identity over 92 % and 85 % of its sequence respectively. Following removal of gaps and ambiguous bases, 1034 nucleotides remained for phylogenetic analyses. Cystodiscus melleni (Jirku et al., 2006) was chosen as the outgroup based on its position in the distance tree generated following initial BLAST analyses of the P. kovalevae sequence. MP and ML trees showed similar topologies, P. kovalevae grouping with C. pantherini and C. anko in both cases, with strong bootstrap support (100) (Fig. 7).	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D5567361FF12A423FA862ED7.taxon	discussion	Discussion. Twelve species of the genus Pseudalataspora have been described from the gall bladders of marine fishes but none has been sequenced to date. In addition, we were made aware of an unpublished manuscript discovered in the late Professor Kovaleva’s laboratory in Kaliningrad, which includes the morphological description of Pseudalataspora pacifica Kovaleva and Grudnev, a new species found in the gall bladder of M. magellanicus caught off the coast of Chile in 1983. The authors described the new species as being similar to P. umbraculiformis, originally described by Gaevskaya and Kovaleva (1984) from the gadid fish Gaidropsarus mediterraneus (L., 1758) in the Northeast Atlantic. However, they considered the two species to differ sufficiently in certain features to be considered different species. The main differences they identified were in the shape and size of the spores, the diameter of the polar capsules, and the number of coils in the polar filament. Because the description of P. pacifica remains unpublished, it cannot be considered to be a valid species, but in recognition of Professor Kovaleva’s discovery we decided to name our new species after her. Table 1 compares P. kovalevae with both P. umbraculiformis and P. pacifica. Pseudalataspora kovalevae is from the same host and locality as P. pacifica and these two species are clearly more similar in morphology than either is to P. umbraculiformis. Gunter et al. (2009) suggested that the Ceratomyxa clade currently contains a number of as yet undifferentiated genera and that additional sampling and further morphological and genetic data are needed to resolve the divisions within the group. As previously mentioned, there are currently no sequence data deposited in GenBank for any species of Pseudalataspora, and the new P. kovalevae 18 S rRNA gene sequence grouped most closely with some Ceratomyxa species. Morphologically the spores of these two genera are very similar, with the presence or absence of delicate alate processes being the main distinguishing feature between them. The morphological and molecular similarities between these genera therefore suggest that they are closely related, but they are currently classified in different families. The alate processes are fragile and not always easily observed, which raises the possibility that some species currently assigned to the genus Ceratomyxa may turn out on closer inspection to belong in Pseudalataspora. Feature P. umbraculiformis P. “ pacifica ” (*) P. kovalevae Plasmodium shape Club-shaped or spherical, Spherical, disporous Spherical, disporous disporous Plasmodium dimensions 18.0 – 21.0 29.28 – 32.28 28.4 – 38.4 x 28.0 – 36.0 Spore shape Wedge-shaped, expanded Rounded tops, sharply Triangular with pointed anteriorly, narrowed curved anteriorly extremities in side view; curved posteriorly and flat anteriorly in apical view Spore length 8.0 – 9.3 7.98 – 12.63 8.0 – 10.5 Spore thickness with alate 14.6 – 17.3 27.93 – 35.91 30.0 – 35.5 processes Spore thickness without alate 6.7 – 8.0 Not given 8.8 – 11.6 processes Spore width 14.6 – 17.3 Not given 14.0 – 20.0 Sutural line Clear and straight Clear and straight Prominent, raised Polar capsule diameter 2.7 3.0 - 3.32 2.5 – 3.0 No. of polar filament coils 5 6 Number not observed	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D552736DFF12A4DDFECC2F78.taxon	materials_examined	Type hosts: Macrourus holotrachys Günther, 1878 and Merluccius hubbsi Marini, 1933 Site of infection: gall bladder	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D552736DFF12A4DDFECC2F78.taxon	description	Localities, dates and depths: (1) off Chiloe Island, Chile, 43 ºS, 73 ºW, June 2007, 300 m; (2) west of Falkland Islands, 51 º 00´to 52 º 30´S, 62 º 00´to 62 º 30´W, October 2007, 200 – 250 m; (3) west of Falkland Islands, 51 º 00´to 52 º 30´S, 62 º 00´to 62 º 30´W, October 2009, 200 – 250 m.	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D552736DFF12A4DDFECC2F78.taxon	materials_examined	Type locality: (1). Prevalence: (1) 4 % (1 of 23); (2) 13 % (4 of 30); (3) 7 % (3 of 42).	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D552736DFF12A4DDFECC2F78.taxon	description	Host length range: 25 – 42 cm Collection numbers: NHMUK 2012.3.19.3, 2012.3.19.4. Morphological description. Vegetative stages not found. Spore elongate with rounded ends in valvular view (Figs. 8, 10), flat elongate with attenuated ends in sutural view (Fig. 9). Sutural line straight and fine. Valve shells each with 4 – 5 surface striations. Sporoplasm located between the polar capsules, binucleate and deeply staining (Fig. 11). Polar capsules long, pyriform, rounded posteriorly and of equal size, at opposite ends of the spore and opening at the tips of the spores. Polar filament thick and with 4 – 6 coils. Dimensions, based on 25 fixed spores, as ranges with means ± SD in parentheses: spore length 14.4 – 20.8 (17.0 ± 2.0); spore width 5.4 – 9.6 (6.7 ± 1.2); spore thickness 3.2 – 5.2 (4.6 ± 0.6); polar capsule length 3.2 – 4.0 (3.6 ± 0.2); polar capsule width 1.6 – 3.2 (2.3 ± 0.5); polar capsule length: spore length = 1: 4.5 – 5.2; spore length: spore width = 1: 2.2 – 2.7. Molecular result. PCR amplification of myxosporean 18 S rRNA gene from one infected gall bladder from sample (3) above generated a product of approximately 1700 nucleotides in both of the samples processed from the individual fish in question. Forward and reverse sequences were obtained for each purified product. All sequences were identical. Seven polymorphic positions were observed. A final sequence of 1632 nucleotides was submitted to Genebank under Accession No. JX 467674. BLAST analysis revealed that the sequence was novel when compared to data currently deposited in Genbank. Based on the Max scores obtained following BLAST analysis, the sequence showed closest identity to Myxidium coryphaenoideum Noble, 1983 (DQ 377697), Cystodiscus melleni (DQ 003031) and Sphaeromyxa zaharoni Diamant et al., 2004 (AY 538662) with 89 %, 87 % and 86 % identity over 100 %, 100 % and 98 % of nucleotide coverage respectively. A sequence of 874 nucleotides remained for phylogenetic analysis following removal of gaps and ambiguous bases. Tetracapsuloides bryosalmonae Canning et al., 1999 was used as the outgroup. MP and ML trees showed similar topologies, with M. baueri grouping with M. coryphaenoideum in both cases, though bootstrap support was relatively poor at 58 and 62 for MP and ML respectively (Fig. 12).	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D552736DFF12A4DDFECC2F78.taxon	discussion	Discussion. The genus Myxidium includes more than 230 named species (Eiras et al., 2011), with more than 70 of these reported from marine fishes. The spores found in M. magellanicus most closely resemble those of Myxidium baueri Kovaleva & Gaevskaya 1982, described from two deepwater fishes in the Southwest Atlantic, one of which, Merluccius hubbsi, is closely related to M. magellanicus, both species being merluccids. Kalavati et al. (1995) also reported M. baueri from Merluccius australis (Hutton, 1872) and M. hubbsi caught in the Southwest Atlantic. These authors noted certain differences between their own observations and the original description, most notably the difference between the numbers of spores in the trophozoite and the presence of distinctive knob-like extremities on the spores in their material. We did not find any vegetative stages and we did not observe the knoblike extremities on the spores of our material from M. magellanicus. Despite these differences and because of the otherwise similar morphology, the common locality and the relatedness of the hosts (Table 2), we tentatively identify our material as M. baueri, for which M. magellanicus is a new host record. Unfortunately no sequence data are currently available for M. baueri in the public databases. Of the species of Myxidium for which molecular sequences are available, our material most closely resembles M. coryphaenoidium, collected from Coryphaenoides rupestris Gunnerus, 1765 caught in the North Atlantic (Fiala, 2006), though there is still a sufficient level of nucleotide differences to define it as a new species. Myxidium coryphaenoidium was originally described by Noble (1966) from “ Coryphaenoides sp. ” caught in the Pacific off Mexico, and has since been reported from 21 species of deepwater marine fish worldwide, most of them belonging to the family Macrouridae, but also including two morid species and one member of the family Synaphobranchidae (Yoshino & Noble, 1973; Yoshino & Moser, 1974; Moser et al., 1976; Kovaleva & Gaevskaya, 1982; Threlfall & Khan, 1990). Such large host and geographical ranges raise the possibility that M. coryphaenoidium may not be a single cosmopolitan species but a complex of sibling species. Moser et al. (1976) described two forms of spore in M. coryphaenoidium, which they called “ long ” and “ stubby ”, and the occurrence of which they interpreted as polymorphism within the one species rather than mixed infections of two species. Kovaleva & Gaevskaya (1982) considered their new species M. baueri to be identical to the “ stubby ” form of M. coryphaenoideum. They also described and figured shorter forms of spore in M. baueri, which they described as “ anomalous ”. We occasionally observed longer spores similar to those of the “ long ” form of M. coryphaenoideum in our material. This suggests that polymorphism may be a common feature of those species of Myxidium infecting deepwater marine fishes. It is clear that the taxonomy of the species identified as M. coryphaenoidium and M. baueri requires further investigation, using both morphological and molecular methods. Table 2 compares our material from M. magellanicus with the morphological descriptions of both M. baueri and M. coryphaenoidium.	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D55E736BFF12A308FE172D38.taxon	materials_examined	Material studied Host: Macruronus magellanicus Lönnberg, 1907	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D55E736BFF12A308FE172D38.taxon	description	Site of infection: gall bladder Locality, date and depth: (1) Off Chiloe Island, Chile, 43 ºS, 73 ºW, June 2007, 300 m.	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D55E736BFF12A308FE172D38.taxon	materials_examined	Type locality: (1) Prevalence: 4 % (1 of 25).	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D55E736BFF12A308FE172D38.taxon	description	Host length range: 25 – 42 cm. Collection numbers: NHMUK 2012.3.19.2. Morphological description. Sporoblast oval or irregularly shaped, disporic (Fig. 13). Dimensions, based on 4 fixed specimens: 28.4 – 38.4 x 28.0 – 36.0. Developing sporoblasts show deeply staining cords (the origins of the membranous veil) twisted around the body (Fig. 14). These are clearly visible only when stained with Indian ink, but are indistinct and lightly stained in Giemsa preparations. Spore subspherical or broadly oval (Fig. 15). A smooth membranous veil originating anteriorly extends beyond the spore body posteriorly, enveloping the entire spore. Sutural ridge prominent anteriorly but thinner posteriorly. Keel-like appendages situated along the sutural ridge join together at the posterior extremity. Sporoplasm deeply staining and binucleate. Spore valves thin and smooth. Polar capsules pyriform, subterminal, one on either side of sutural line. Polar filament with 3 – 4 coils, not clearly visible. Dimensions, based on 15 fixed spores, as ranges with means and ± SD in parentheses: spore length 9.6 – 19.2 (13.64 ± 3.67); spore width 10.2 – 22.4 (19.03 ± 3.64); spore thickness 14.0 – 20.0 (16.09 ± 1.92); spore veil 32.0 – 48.0 (40.43 ± 6.47); polar capsule length 6.4 – 8.0 (6.93 ± 0.64); polar capsule width 3.2 – 5.4 (4.0 ± 0.80); spore length: spore width = 1: 0.8 – 0.96; polar capsule length: spore length = 1: 1.5 – 2.4.	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
03E287B9D55E736BFF12A308FE172D38.taxon	discussion	Discussion. This species was not observed during the initial examinations of gall bladders, but was discovered later as a double infection with Pseudalataspora kovalevae in a formalin-preserved sample from a fish originally identified as infected only with the latter species. We therefore have no molecular sequence and no photographs of fresh material. Only five species of Palliatus have been previously described from marine fishes, four from the gall bladder and one from the pancreas (Shulman et al., 1979; Padma Dorothy & Kalavati, 1998; Aseeva, 2003). The new species differs considerably in morphology from all of these and the host and locality are both new for the genus Palliatus (Table 3).	en	Kalavati, Chaganti, Mackenzie, Ken, Collins, Catherine, Hemmingsen, Willy, Brickle, Paul (2013): Two new species of myxosporean parasites (Myxosporea: Bivalvulida) from gall bladders of Macruronus magellanicus Lönnberg, 1907 (Teleostei: Merlucciidae). Zootaxa 3647 (4): 541-554, DOI: 10.11646/zootaxa.3647.4.4
