The demise of Leptotheca Thelohan, 1895 (Myxozoa: Myxosporea: Ceratomyxidae) and assignment of its species to Ceratomyxa Thelohan, 1892 (Myxosporea: Ceratomyxidae), Ellipsomyxa Koie, 2003 (Myxosporea: Ceratomyxidae), Myxobolus Butschli, 1882 and Sphaerospora Thelohan, 1892 (Myxosporea: Sphaerospo...

Robert Adlard

The genus Ceratocystis was established in 1890 and accommodates many important fungi. These include serious plant pathogens, significant insect symbionts and agents of timber degradation that result in substantial economic losses. Virtually since its type was described from sweet potatoes, the taxonomy of Ceratocystis has been confused and vigorously debated. In recent years, particulary during the last two decades, it has become very obvious that this genus includes a wide diversity of very different fungi. These have been roughly lumped together due to their similar morphological structures that have clearly evolved through convergent evolution linked to an insect-associated ecology. As has been true for many other groups of fungi, the emergence of DNA-based sequence data and associated phylogenetic inferences, have made it possible to robustly support very distinct boundaries defined by morphological characters and ecological differences. In this study, DNA-sequence data for three carefully selected gene regions (60S, LSU, MCM7) were generated for 79 species residing in the aggregate genus Ceratocystis sensu lato and these data were subjected to rigorous phylogenetic analyses. The results made it possible to distinguish seven major groups for which generic names have been chosen and descriptions either provided or emended. The emended genera included Ceratocystis sensu stricto, Chalaropsis, Endoconidiophora, Thielaviopsis, and Ambrosiella, while two new genera, Davidsoniella and Huntiella, were described. In total, 30 new combinations have been made. This major revision of the generic boundaries in the Ceratocystidaceae will simplify future treatments and work with an important group of fungi including distantly related species illogically aggregated under a single name.

Ellipsomyxa arariensis n. sp. was found in the gallbladder of Pygocentrus nattereri Kner, 1858 and Pimelodus ornatus Kner, 1858 from the Arari River on Marajó Island in Pará, Brazil. The new species has disporous plasmodium that varies in size and shape, with ellipsoidal mature spores in the sutural view that have a curved suture line. The spores are 12.6 (12.0-13.4) μm in length and 7.3 (6.7-8.0) μm in width. The two polar capsules present in the spore are pyriform and of equal size, with subterminal openings that project in opposite directions. The polar capsules are 3.5 (3.4-4.0) μm long and 2.6 (2.5-3.2) μm wide. Based on the partial sequences of the SSU rRNA gene of the Ellipsomyxa arariensis n. sp. spores found in Pygocentrus nattereri Kner, 1858 (1325 bps) and Pimelodus ornatus Kner, 1858 (1240 bps), the new species is clearly distinct from all the other myxozoan sequences deposited in GenBank. Based on Bayesian inference and p distances, the new species belongs to the "...

In the present study, 2 species of myxozoan parasites, the novel Ceratomyxa scophthalmi sp. nov. and Myxidium finnmarchicum, were documented from the gallbladder of Scophthalmus maeoticus inhabiting the Turkish Black Sea coast at Sinop. C. scophthalmi sp. nov. had mostly spherical, clavate, and fine-grained polymorphic plasmodia that ranged in diameter from 16.0 to 29.0 μm. Spores were arcuate with a convex anterior margin and concave posterior. Valves of spores were highly elongated and unequal in thickness with one valve 1.1−1.3 times thicker than the other. Valves narrowed gradually toward slightly truncated ends. The mean (range) dimensions of spores were 9.4 (8.5−10.0) μm in length and 49.6 (44.5−55.0) μm in thickness, with polar capsules that were 3.1 (2.7−3.5) μm in length and 3.2 (2.7−3.5) μm in width. Polar filaments were coiled with 4−5 turns. Large numbers of immature forms of C. scophthalmi sp. nov., which were often ‘crumpled’, and mature forms were found together in the gallbladder bile. Along with morphological differences, phylogenetic analysis based on 18S rDNA, including pairwise nucleotide similarities with other related Ceratomyxa species, suggests C. scophthalmi as a novel species. Moreover, this report provides the first morphological and molecular descriptions of M. finnmarchicum outside of its original geographical location and type host species.

Syst Parasitol (2010) 75:81–104 DOI 10.1007/s11230-009-9227-1 The demise of Leptotheca Thélohan, 1895 (Myxozoa: Myxosporea: Ceratomyxidae) and assignment of its species to Ceratomyxa Thélohan, 1892 (Myxosporea: Ceratomyxidae), Ellipsomyxa Køie, 2003 (Myxosporea: Ceratomyxidae), Myxobolus Bütschli, 1882 and Sphaerospora Thélohan, 1892 (Myxosporea: Sphaerosporidae) Nicole Gunter • Robert Adlard Received: 16 April 2009 / Accepted: 1 August 2009 Ó Springer Science+Business Media B.V. 2009 Abstract A revision of Leptotheca Thélohan, 1895 is presented. The boundaries that separate Leptotheca from Ceratomyxa Thélohan, 1892 and Sphaerospora Thélohan, 1892 are vague and have been highlighted as an area of concern within myxosporean classification. A survey of the literature revealed 63 species that are currently assigned to Leptotheca and a further 11 species that have been relegated as synonyms in Ceratomyxa, Sphaerospora or Myxobolus Bütschli, 1882. The placement of some species in the genus is unclear and demonstrates the need for a revision. The type-species of Leptotheca (L. agilis Thélohan 1892) has many Ceratomyxa-like characters, such that a minor amendation of the diagnosis of Ceratomyxa will then accept the type-species of Leptotheca, rendering the latter genus its synonym. We propose the suppression of Leptotheca, with all species currently assigned to that genus reassigned to Ceratomyxa, Ellipsomyxa Køie, 2003, Myxobolus or Sphaerospora on the basis of appropriate morphological N. Gunter R. Adlard (&) Biodiversity Program, Queensland Museum, P.O. Box 3300, South Brisbane, QLD 4101, Australia e-mail: robert.adlard@qm.qld.gov.au N. Gunter R. Adlard School of Chemistry and Molecular Biosciences, The University of Queensland, Brisbane, QLD 4072, Australia and biological traits. The diagnoses of Ceratomyxa and Ellipsomyxa are amended appropriately. Molecular analysis may change the placement of some species in the future; however, the aim of this review was to eliminate the ambiguity of assignment of species in the genera Leptotheca, Ceratomyxa and Sphaerospora by suppressing Leptotheca. Introduction Members of the myxozoan genus Leptotheca Thélohan, 1895 are predominantly parasites of the gallbladder or urinary system of teleost fishes and amphibians (Lom & Dyková, 2006). They have ‘‘oval, ellipsoidal spores, sometimes arcuate where the length of the individual laterally prolonged shell valve does not exceed the axial diameter of the spore, but significantly exceeds one half of this diameter’’ (Lom & Dyková, 2006). However, this spore-based classification has proved problematic. The boundaries between Leptotheca Thélohan, 1895 and Ceratomyxa Thélohan, 1892 and between Leptotheca and Sphaerospora Thélohan, 1892 are indistinct and remain a significant concern in myxosporean classification (Meglitsch, 1960; Lom & Noble, 1984; Lom & Dyková, 1992; Lom & Dyková, 2006). The classification of Leptotheca has a long and confusing history. Thélohan (1895) proposed the 123 82 genus on the basis of spore morphology. Six species were attributed to the genus with the type-species by original designation, L. agilis (Thélohan, 1892), originally having being recognised and described as Ceratomyxa agilis Thélohan, 1892 (see Thélohan, 1895). Doflein (1899) erected the family Ceratomyxidae, which contained only Ceratomyxa and Leptotheca, when he proposed a classification of the Myxosporidia based on vegetative development. The family was maintained even though the division based on development was found to be unacceptable by later taxonomists (Davis, 1917; Kudo, 1919, 1933). The currently accepted classification system is based fundamentally on spore morphology, which is the guiding principle in the taxonomy of the Myxosporea. Jameson (1929) did not believe the separation of Leptotheca and Ceratomyxa was warranted based on the variability of spores and the presence of intermediate spore types within the same species. He proposed that Leptotheca spp. be allocated to Ceratomyxa and suggested it would be more helpful to emphasise similarities rather than differences. However, Jameson’s proposal was apparently ignored at the time and many species described as Ceratomyxa by him were later allocated to Leptotheca by Kudo (1933). Many species originally described within Leptotheca have since been reassigned to other genera. L. parva Thélohan, 1895, L. coris Stempell, 1918, L. inaequalis Doflein, 1898, L. obovalis Fantham, 1930, L. macrospora Auerbach, 1909, L. platichthys Fujita, 1923 and L. yoichiensis Fujita, 1923 have been transferred to Ceratomyxa (see Kudo, 1933; Dogiel, 1948). L. acheilognathi Fujita, 1927 is now recognised as Myxobolus acheilognathi (see Landsberg & Lom, 1991). L. brevis Polyanskii, 1955, L. ohlmacheri Whinery, 1893 and L. ranae Thélohan, 1895 have been placed within Sphaerospora (see Arthur & Lom, 1985; Desser et al., 1986; Jirků et al., 2007). Ceratomyxa abbreviata Davis, 1917, C. fisheri Jameson, 1929, C. galeata Jameson, 1929, C. inconstans Jameson, 1929 and C. faba Meglitsch, 1960 have been reassigned to Leptotheca (see Kudo, 1933; Zhao & Song, 2003). Although these recommendations appear to have been generally accepted, the literature is scattered with competing opinions regarding these changes, and the original names still appear in some subsequent publications. Laird (1953) considers it ‘‘inadvisable to follow Kudo (1933) in transferring L. obovalis to Ceratomyxa’’. Noble 123 Syst Parasitol (2010) 75:81–104 (1966) discussed L. yoichiensis, while ignoring its reassignment to Ceratomyxa by Kudo (1933). A recent synopsis of Ceratomyxa (see Eiras, 2006) includes C. abbreviata, C. fisheri, C. galeata and C. inconstans. Jameson (1931) discussed C. (Leptotheca) limandae (Fujita, 1923) and C. (Leptotheca) scissura (Fujita, 1923) based on his 1929 proposal, but Fujita (1923) also designated a different myxosporean as C. limandae Fujita, 1923; this homonymy was left unresolved. All things considered, the placement of many species in the genus is in question and the nomenclature lacks clarity. A thorough survey of the literature revealed over 70 species that at one stage belonged to Leptotheca, many of which have since been synonymised. Lom & Dyková (2006) note 56 nominal species in this genus. We have compiled a list of 63 species, which we believe to be currently assigned to Leptotheca. Over the years, many species have been proposed or synonymised, without strictly following the rules of the ICZN, and the validity of such species is therefore questionable. Compounding this problem, type-material is unavailable for many species. To our best knowledge, the list of known and valid species we present here is exhaustive. We agree with Jameson’s (1929) proposal that it would be more helpful to emphasise the similarities between existing genera rather than the differences, but believe it is not as simple as collapsing the whole genus into Ceratomyxa. With this in mind, we attempt to resolve the problematic position of Leptotheca. Based on the morphological and biological attributes shared between Leptotheca and other myxozoan genera, the current diagnosis is unnecessary and inappropriate. On investigation of the genus, two main patterns become apparent, species that infect the gall-bladder of teleosts and species that infect the urinary system of teleosts and amphibians. To eliminate the confusion between Leptotheca and Ceratomyxa, we propose all but one species primarily infecting the gall-bladder be placed in Ceratomyxa and the diagnosis for that genus be amended. While we are becoming convinced that Ceratomyxa should be restricted to species infecting the gall-bladder, we do not seek to redefine those currently valid species of Ceratomyxa that occur in either the urinary system or other organs. The purpose of this review is to eliminate the ambiguity of Leptotheca. Syst Parasitol (2010) 75:81–104 A minor modification of the generic diagnosis of Ceratomyxa will enable the inclusion of the typespecies of Leptotheca, L. agilis (Thélohan, 1892), rendering Leptotheca its synonym. The spores (SP) of L. agilis are oval in sutural view and slightly elongate, with the length of the shell valve only slightly less than the axial diameter of the spore and occasionally equalling it (SP: 6–7 9 11–12 lm). Species of Ceratomyxa that have Leptotheca-like characteristics (i.e. axial diameter to valve length ratios of 1:1 or less; which is equivalent to a spore length/thickness ratio of 1:2 or less) include C. acanthuri Kpatcha, Diebakate, Faye & Toguebaye, 1996 (10.5 9 16.5 lm), C. allantoidea Gayeskaya & Kovaljova, 1984 (13.3– 16 9 16–22 lm), C. costata Aseeva, 2001 (11–14 9 22–30 lm), C. curvata Cunha & Fonseca, 1918 (12 9 24 lm), C. hungarica Molnar, 1992 (10.1 9 18.8 lm), C. insolita Meglitsch, 1960 (11.5 9 16.2 lm), C. labracis Sitjá-Bobadilla & Alvarez-Pellitero, 1993 (6.2 9 12.4 lm) and C. recurvata Davis, 1917 (8–9 9 16 lm). We believe that this character (the length/thickness ratio) is an inappropriate measure for separating the two genera, particularly when the placement of the type-species of Leptotheca is equivocal, and due to the morphological plasticity of the spores and the presence of species with intermediate spore types. Although molecular evidence is limited, it provides some support for this position. A recent phylogeny of Ceratomyxa spp. mapped morphological characters, including length to thickness ratios, against a molecular phylogeny (Gunter et al., 2009). Two Ceratomyxa species, C. labracis and C. ernsti Gunter, Whipps & Adlard, 2009, have spores with length/ thickness ratios similar to what has been recognised as Leptotheca (0.5 and 0.48, respectively) (Gunter et al., 2009) and are more closely related by molecular criteria to other species of Ceratomyxa than they are to each other. Additionally, an undescribed species with typical ‘Leptotheca’ spores is closely related by molecular criteria to other Ceratomyxa spp. and Palliatus indecorus Schulman, Kovaljova & Dubina, 1979 (see Gunter et al., 2009). An additional character Thélohan (1895) used to separate Ceratomyxa and Leptotheca was whether the sporoplasm filled the spore. Generally, Leptotheca has sporoplasm that completely fills the spore, whereas the sporoplasm of Ceratomyxa does not; however, this character has been applied inconsistently when assigning species to each genus (see Jameson, 83 1929). C. truncata Thélohan, 1895 (5 9 25 lm) and C. strepospora Davis, 1917 (4 9 34–39 lm) have sporoplasms that entirely fill the valves, whereas L. fusiformis Davis, 1917 (9 9 16 lm) and L. scissura Davis, 1917 (11 9 22 lm) do not. Upon investigation of the gall-bladder infecting species, L. fusiformis was found to have polar capsules close to the suture but not at the anterior pole of the spore. In apical view the polar capsules are at opposite sides of the spore and discharges in opposite directions. On the basis of these characters, L. fusiformis can be accommodated, with a minor amendment of the diagnosis, in Ellipsomyxa Køie, 2003, which currently only contains one species, E. gobii Køie, 2003 (see Lom & Dyková, 2006). To eliminate the confusion between Leptotheca and Sphaerospora, we propose that all species found infecting the urinary system be placed within Sphaerospora. All species in this category fit the current morphological diagnosis of Sphaerospora in having spherical or sub-spherical spores, in which the valvular diameter equals or, only in a few species, exceeds the sutural diameter (Lom & Dyková, 2006). Some species have valves with protuberances, bumps or prominent sutural ridges, which also are diagnostic characters of Sphaerospora (see Lom & Dyková, 2006). No site of infection or spore dimensions have ever been recorded for L. perlata Gurley, 1893. The history of this parasite is confusing. It was first recorded from Gymnocephalus cernuus Linnaeus (= Acerina cernua) in Balbiani (1883), but only a line drawing was made and no species name proposed. It was described as Myxobolus perlatum by Gurley (1893). Earlier, Thélohan (1892) had proposed Sphaerospora and Gurley (1894) subsequently moved M. perlatum to Sphaerospora. Labbé (1899) moved Sphaeorospora (Myxobolus) perlatum to Leptotheca and corrected the name from ‘perlatum’ to ‘perlata’. Based on morphological features visible in Balbiani’s (1883) original line drawing (i.e. the presence of protuberances and a prominent sutural ridge), we believe that L. perlata is a species of Sphaerospora. Although Gurley’s (1893) description itself is poor, it is still an available species under the provisions of the ICZN, and likely valid as well. The taxonomic position of L. fugu Tun, Yokoyama, Ogawa & Wakabayashi, 2000 is intriguing. This species primarily infects the intestinal epithelium and 123 84 rectum, occasionally the stomach and rarely the gallbladder (Tun et al., 2000). It was first recorded as Ceratomyxa sp. 1 by Ogawa & Yokoyama (1998); however, recent phylogenetic analysis suggested it is more closely related to Sphaerospora spp. than to Ceratomyxa spp. (see Jirků et al., 2007). Additionally, some Sphaerospora spp. are known to infect the intestinal epithelium, e.g. S. dicentrachi Sitjá-Bobadilla & Alvarez-Pellitero, 1992 and S. reichenowi Jacob, 1953 (see Fioravanti et al., 2004; Jacob, 1953), whereas L. sparidarum Sitja-Bobadilla & AlvarezPellitero, 2001 primarily infects the urinary system but also the intestinal epithelium. Although the spores of L. fugu look slightly arched, we believe this is due to the presence of enlarged protuberances, similar to those of S. truttae Fischer-Scherl, El-Matbouli & Hoffmann, 1986 and S. molnari Lom, Pavlaskova & Grupcheva, 1983. The original description does not describe them as protuberances, but they are clearly visible as such in the figures of fresh and stained spores and in the line drawing (Tun et al., 2000). Based on molecular (Jirků et al., 2007) and morphological evidence, we propose that L. fugu be allocated to Sphaerospora. Leptotheca asymmetrica Lalitha Kumari, 1969, was described from the gall-bladder, gills and intestine of Osteochilus hasselti Valenciennes (=O. neilli (Dav)). The spores of this species are transversely elliptical, with the breadth of the spore being more than the sutural diameter. This led the author to assign the species to Leptotheca, but Lalitha Kumari (1969) appeared not to consider the position of the suture. The spores are flattened parallel to the sutural plane, with both polar capsules lying solely in this plane. For this reason, L. asymmetrica is more appropriately placed in the Platysporina, and we propose that it be transferred to Myxobolus. L. asymmetrica has striated shell valves and unequal polar capsules, which are rare but not unknown in Myxobolus. Generally, the shell valves of Myxobolus spp. are smooth; however the spores of at least one other species have striated shell valves (M. potaili Lalitha Kumari, 1969 — see Basu & Haldar, 2003). The polar capsules are also asymmetrical. Myxobolus spp. that have polar capsules of differing sizes include M. absonus Cellere, Cordeiro & Adriano, 2002, M. bilongi Fomena, 1995, M. inaequalis Gurley, 1893 and M. inaequus Kent & Hoffman, 1948 (see Cellere et al., 2002). 123 Syst Parasitol (2010) 75:81–104 Ceratomyxa Thélohan, 1892 Forty-two species formally belonging to Leptotheca are here assigned to Ceratomyxa on the basis of morphometric similarities and locality in the host tissues. All species are from the gall-bladders of marine fishes and, with minor amendments (in bold below), fit the diagnosis of Ceratomyxa. A synopsis and notes on species is provided below and complemented with morphometric and host data (Table 1). Amended diagnosis Spores elongate, generally crescent-shaped or arcuate, occasionally sub-spherical or oval; shell valves significantly exceeding in length half of axial diameter of spore. Shell valves conical or subhemispherical, often pliable rather than rigid as in other genera. Sub-spherical polar capsules located and open close to suture line, in plane perpendicular to it, at anterior pole of spore, but exceptionally open laterally from central suture line. Binucleate sporoplasm may fill spore cavity completely; in some species, two uninucleate sporoplasms have been reported. Trophozoites usually disporic, sometimes mono- to polysporic. Coelozoic parasites of marine fishes, exceptionally in freshwater and rarely histozoic. (Modified after Lom & Dyková, 2006.) Ceratomyxa abbreviata Davis, 1917 Syn. Leptotheca abbreviata (Davis, 1917) Kudo 1933 (Fig. 1) Description: Anterior margin of spores extremely convex and posterior margin concave, roughly crescent-shaped, almost circular; valves equal, having exceptionally large diameter in comparison with length; ends of valves rounded. Polar capsules spherical (Davis, 1917; Eiras, 2006). Ceratomyxa acanthopagri (Zhao & Song, 2003) n. comb. Syn. Leptotheca acanthopagri Zhao & Song 2003 (Fig. 2a–b) Description: Spores elongate ovoid to subspherical; valves with rounded ends; sutural line distinct with raised sutural ridge. Polar capsules spherical. Species Host(s) (original name) Length Thickness PCL PCW References C. abbreviata Davis, 1917 Rhizoprionodon terraenovae GB (= Scoliodon terranovae) Off USA 14 17 4.5 4.5 Davis (1917) C. acanthopagri (Zhao & Song, 2003) n. comb. Acanthopagrus schlegelii (= A. schlegeli) GB Off China 9.4 (8.7–10.0) 17.9 (16.0–20.0) 1.9 (1.7–2.3) 1.9 (1.7–2.3) Zhao & Song (2003) C. adeli (Bakay & Grudnev, Sebastes marinus, S. 1998) n. comb. mentella, S. fasciatus GB North Atlantic 12–13 18.62–20 4 4 Bakay & Grudnev (1998) C. agilis Thélohan, 1892 GB Mediterranean and Tyrrhenian coasts 6–7 11–12 – – Thélohan (1895) GB Peter the Great Bay, Japan 9.2–10 13–15 5 2.5–2.6 Aseeva (2001) Lepidonotothen squamifrons GB C. ampla (Kovaljova, Rodjuk & Grudnev, 2002) n. comb. Antarctic Ocean 14.7–17.3 25.3–29.3 4.6–6 4.6–6 Kovaljova et al. (2002) C. annulata (Meglitsch, 1960) n. comb. Thyrsites atun, Rexea solandri (= Jordanidia solandri), Arripis trutta GB Off New Zealand 5.8 (4.5–6.7) 10.9 (7.9–12.3) 2.3 (1.7–2.8) 2.3 (1.7–2.8) Meglitsch (1960) C. apogoni (Narasimhamurti, Kalavati, Anuradha & Padma Dorothy, 1990) n. comb. Apogon aureus, Nemipterus japonicus GB Off India 17.56 (16.8–18.2) 25.40 (24.6–26.2) 3.8 (2.4–4.2) 3.8 (2.4–4.2) Narasimhamurti et al. (1990) C. beveridgei (Moser, Kent Scorpaenopsis vittapinna & Dennis, 1989) n. comb. (= S. gibbosus) GB Off Australia 9.1 (8.0–10.0) 16.5 (15.0–18.0) 2.5 (2.0–3.0) 2.5 (2.0–3.0) Moser et al. (1989) C. coelorhyncha (Yoshino & Coelorhinchus Noble, 1973) n. comb. coelorhinchus (= Coelorhynchus coelorhynchus) GB Off Ireland 6.7 (6.0–8.0) 11.34 (9.0–13.0) 2.01 (1.5–3.0) 2.01 (1.5–3.0) Yoshino & Noble (1973) C. constricta Fujita, 1923 n. comb. Sebastolobus macrochir GB Off Japan 10.0 19.0 – – Fujita (1923) C. elongata (Thélohan, 1895) n. comb. Merluccius merluccius (= M. vulgaris) GB Off France 12–15 18–20 – – Thélohan (1895) C. etrumuci (Dogiel, 1948) n. comb. Etrumeus micropus GB Off Russia 7 7 – – Dogiel (1948) Dasyatis pastinaca (= Trygon vulgaris) C. amatea (Aseeva, 2001) n. Pseudopleuronectes comb. yokohamae (= Limanda yokohamae) Tissue Locality Syst Parasitol (2010) 75:81–104 Table 1 Morphological features of the species of Ceratomyxa 85 123 86 123 Table 1 continued Host(s) (original name) Tissue Locality Length Thickness PCL PCW References C. faba Meglitsch, 1960 Arnoglossus scapha (= Caulopsetta scapha) GB Off New Zealand 6.2 (5.6–6.7) 12.7 (10.7–14.1) 2.4 (2.0–3.1) 2.4 (2.0–3.1) Meglitsch (1960) C. fisheri Jameson, 1929 Hydrolagus colliei GB Off USA 5.1–7.1 9.3–13.3 – – Jameson (1929) C. fujitai nom. nov. [Syn. Leptotheca limandae] Limanda punctatissimus (= L. iridorum), L. aspera GB Off Japan 14.0–16.0 24.0 – 26.0 6.0–8.0 – Fujita (1923) C. galeata Jameson, 1929 Eptatretus stoutii (= Polistotrema stouti) GB Off USA 6.6–8.7 11.6–14.7 – – Jameson (1929) C. hepseti (Thélohan, 1895) n. comb. Atherina hepsetus GB Off France 7–8 12–15 – – Thélohan (1895) C. inconstans Jameson, 1929 Scomber japonicus GB Off USA 5.4–7.6 11.2–13.3 – – Jameson (1929) C. informis (Auerbach, 1910) n. comb. Merlangius merlangus GB North Sea 9.8 (9.0–12.0) 19.1 (15.0–21.0) 3.6 (2.5–4.0) 3.6 (2.5–4.0) Kalavati & MacKenzie (1999) C. kovaljovae (Bakay & Grudnev, 1998) n. comb. Sebastes viviparus GB North Atlantic 13.3 21.28 4.65–5.32 4.65–5.32 Bakay & Grudnev (1998) C. latesi (Chakravarty, 1943) n. comb. Lates calcarifer GB Off Bengal 6.2 10.3–12.4 3.1 3.1 Chakravarty (1943) C. longipes (Auerbach, 1910) n. comb. Brosme brosme, Melanogrammus aeglefinus GB North Sea 8.3 (7.5–8.5) 14.4 (12.5–16.5) 3.2 (2.5–3.5) 3.2 (2.5–3.5) Kalavati & MacKenzie (1999) C. lovei nom. nov. [Syn. Leptotheca sebasta] Sebastes serranoides GB Off California 8.0 (7.5–8.5) 13.8 (13.0–15.0) 3.5 (3.0–4.0) 3.5 (3.0–4.0) Moser et al. (1976) C. lubati nom. nov. [Syn. Leptotheca chromis] Chromis chromis GB Off France 6 (5.5–7) 14 (12.5–15) 3 2.75 Lubat et al. (1989) C. macroformis (Gaevskaya & Kovaljova, 1984) n. comb. Sebastes viviparus GB North-east Atlantic 10.7–12.2 20.0–21.0 4.0–4.2 4.0–4.2 Gaevskaya & Kovaljova (1984) C. macronesi (Chakravarty, 1943) n. comb. Mystus gulio (= Macrones gulio) GB Off Bengal 6.18–7.2 10–14.4 3.1 3.1 Chakravarty (1943) C. macrouridonum nom. nov. [Syn. Leptotheca minuta] Coelorinchus occa (= Coryphaenoides occa), Nezumia condylura GB Lesser Antilles and off Japan 4.8 (4.0–6.5) 6.7 (5.0–8.5) 1.2 (1.0–2.0) 1.2 (1.0–2.0) Moser & Noble (1976) C. minima (Meglitsch, 1960) n. comb. Arripis trutta GB Off New Zealand 5.2 (5.1–6.2) 9.3 (7.9–11.2) 1.6 (1.1–2.1) 1.6 (1.1–2.1) Meglitsch (1960) Syst Parasitol (2010) 75:81–104 Species Species Host(s) (original name) Tissue Locality Length Thickness PCL PCW References C. mylionis (Ishizaki, 1960) n. comb. Acanthopagrus schlegelii (= Mylio macrocephalus) GB Off Japan 6.2 13.3 2.3 2.3 Ishizaki (1960) C. nebulifera (Zhao & Song, Pholis nebulosa (= Enedrias GB 2003) n. comb. nebulosus) Off China 5.8 (5.6–6.3) 11.5 (10–13) 1.7 (1.5–1.9) 1.7 (1.5–1.9) Zhao & Song (2003) C. noblei nom. nov. [Syn Leptotheca elegans] Gibbonsia elegans GB Off California 9.0 17.0 2.2 3.0 Noble (1938) C. opisthocornata (Evdokimova, 1977) n. comb. Odontesthes incisa (= Austroatherina incisa) GB Off Argentina 6.4–8 9.6–14.4 3.2–4 3.2–4 Evdokimova (1977) C. ovalis (Kovaljova & Gaevskaya, 1983) n. comb. Trachurus murphyi GB Pacific Ocean 6.0–6.65 9.97–10.64 2.0–2.66 2.0–2.66 Kovaljova & Gaevskaya (1983) C. parva Thélohan, 1895 [Syn Leptotheca scombri] Scomber scombrus (= Scomber scomber) GB Black Sea 8–9 12–14 – – Pogoreltseva (1964) C. pegusae (Kpatcha, Diebakate & Toguebaye, 1996) n. comb. Pegusa lascaris GB Off Senegal 8.32 (6.7–9) 12.78 (11.25–13.5) 4.06 (3.6–4.5) 4.06 (3.6–4.5) Kpatcha et al. (1996) C. pinguis (Meglitsch, 1960) Peltorhamphus n. comb. novaezeelandiae, Arnoglossus scapha (= Caulopsetta scapha) GB Off New Zealand 9.7 (8.3–10.8) C. polymorpha (Labbé, 1899) n. comb. Phycis phycis (= P. mediterranea) GB C. quadritaenia (Kovaljova & Zubtschenko, 1984) n. comb. Epigonus telescopus C. scissura (Davis, 1917) n. comb. Dasyatis hastata (= D. hastatus), Gymnura altavela (= Pteroplatea maclura) 2.9 (2.4–3.9) 2.9 (2.4–3.9) 2.9 (2.8–3.4) 2.9 (2.8–3.4) Off France 10–12 18–20 3 2.5 Labbé (1899) GB Atlantic Ocean 8.0–9.3 12.0–13.3 4.0 2.6 Kovaljova & Zubchenko (1984) GB Off North Carolina 11 22 4 4 Davis (1917) C. sebastisci (Zhao & Song, Sebastiscus sp. 2003) n. comb. GB Off China 10.2 (10.0–10.5) 20.6 (19–23) 2.9 (2.5–3.1) 2.9 (2.5–3.1) Zhao & Song (2003) C. subelegans (Laird, 1953) Callogobius atratus, n. comb. Diplocrepis puniceus GB Off New Zealand 11.6 (8.6–12.5) 22.6 (19.6–26.2) 2.5 (2.1–2.9) 2.5 (2.1–2.9) Laird (1953) 17.3 (16.2–20.3) 2.5 (2.1–2.9) 2.5 (2.1–2.9) C. vikrami (Tripathi, 1948) n. comb. GB 10.3 (10.3–13.7) 18.3 (17.1–20.6) 3.4 (3.2–3.4) 3.4 (3.2–3.4) Zeus faber 9.0 (8.3–10.7) Off England Meglitsch (1960) Tripathi (1948) Mean and/or range are provided in micrometres, with the range in parenthesis if both were reported. Tissue location listed: GB gall-bladder. Spore length, or thickness are listed according to the guidelines for spore descriptions recommended by Lom & Arthur (1989) 87 123 16.4 (13.7–18.6) 10.9 (10.2–11.0) 17.4 (16.0–19.0) Syst Parasitol (2010) 75:81–104 Table 1 continued 88 Ceratomyxa adeli (Bakay & Grudnev, 1998) n. comb. Syn. Leptotheca adeli Bakay & Grudnev, 1998 (Fig. 3a–b) Description: Spores ovoid; anterior margin arched and posterior margin slightly concave or flattened with rounded ends; valves equal; sutural line not protruding. Polar capsules spherical. Ceratomyxa agilis Thélohan, 1892 Syn. Leptotheca agilis (Thélohan, 1892) (Fig. 4) Description: Spores ovoid in sutural view slightly elongated; valves equal; sutural ridge not protruding. Syst Parasitol (2010) 75:81–104 Figs. 1–20 Diagrammatic illustrations of Ceratomyxa spp. c spores. 1. C. abbreviata (after Davis, 1917); 2a–b. C. acanthopagri n. comb. (after Zhao & Song, 2003); 3a–b. C. adeli n. comb. (after Bakay & Grudnev, 1998); 4. C. agilis (after Thélohan, 1895); 5. C. amataea n. comb. (after Aseeva, 2001); 6. C. ampla n. comb. (after Kovaljova, Rodjuk & Grudnev, 2002); 7a–b. C. annulata n. comb. (after Meglitsch, 1960); 8. C. apogoni n. comb. (after Narasimhamurti et al., 1990); 9. C. beveridgei n. comb. (after Moser et al., 1989); 10. C. coelorhyncha n. comb. (after Yoshino & Noble, 1973); 11. C. constricta n. comb (after Fujita, 1923); 12a–b. C. elongata n. comb. (after Thélohan, 1895); 13. C. faba (after Meglitsch, 1960); 14a–b. C. fisheri (after Jameson, 1929); 15a–b. C. fujitai nom. nov. (after Fujita, 1923); 16. C. galeata (after Jameson, 1929); 17a–b. C. inconstans (after Jameson, 1929); 18a–b. C. informis n. comb. (after Kalavati & MacKenzie, 1999); 19. C. kovaljovae n. comb. (after Bakay & Grudnev, 1998); 20a–b. C. latesi n. comb. (after Chakravarty, 1943). Scale-bar: 10 lm Ceratomyxa amatea (Aseeva, 2001) n. comb. Syn. Leptotheca amatea Aseeva, 2001 (Fig. 5) Description: Spores ovoid; anterior margin arched and posterior margin flattened with rounded ends; valves equal with ribbon-like projection at end of valve; sutural ridge indistinct and not protruding. Polar capsules pyriform. Ceratomyxa ampla (Kovaljova, Rodjuk & Grudnev, 2002) n. comb. Syn. Leptotheca ampla Kovaljova, Rodjuk & Grudnev, 2002 (Fig. 6) Description: Spores ovoid in sutural view with rounded ends; valves unequal; sutural line not protruding. Polar capsules spherical. Ceratomyxa annulata (Meglitsch, 1960) n. comb. Syn. Leptotheca annulata Meglitsch, 1960 (Fig. 7a–b) Description: Spores roughly ovoid with broadly curved ends; anterior margin convex and posterior angle generally straight; valves cylindrical and unequal; sutural line heavily ridged. Polar capsules spherical. transversely striated; sutural ridge prominent. Polar capsules spherical; one on either side of sutural ridge. Ceratomyxa beveridgei (Moser, Kent & Dennis, 1989) n. comb. Syn. Leptotheca beveridgei Moser, Kent & Dennis, 1989 (Fig. 9) Description: Spores oblong in sutural view with unequal rounded ends; valves unequal in size and shape; sutural ridge distinct but not protruding. Polar capsules spherical. Ceratomyxa coelorhyncha (Yoshino & Noble, 1973) n. comb. Syn. Leptotheca coelorhyncha Yoshino & Noble, 1973 (Fig. 10) Description: Spores reniform in sutural view with rounded ends; valves equal, thin and smooth; sutural line straight, with distinct sutural ridge most evident at anterior and posterior ends of spore. Polar capsules spherical. Ceratomyxa constricta (Fujita, 1923) n. comb. Ceratomyxa apogoni (Narasimhamurti, Kalavati, Anuradha & Padma Dorothy, 1990) n. comb. Syn. Leptotheca apogoni Narasimhamurti, Kalavati, Anuradha & Padma Dorothy, 1990 (Fig. 8) Description: Spores hemispherical or occasionally trapezoidal; valves lightly stained, thick and 123 Syn. Leptotheca constricta Fujita, 1923 (Fig. 11) Description: Spores transversely reniform, with obtuse extremities in sutural view and rounded ends; valves equal with anterior notch at sutural ridge; sutural line distinct and not protruding. Polar capsules spherical. Syst Parasitol (2010) 75:81–104 89 1 2a 3a 2b 3b 4 5 6 8 10 9 7a 7b 15a 12a 14a 11 13 12b 17a 14b 18a 19 16 17b 18b 15b 20a 20b 123 90 Comment: The name Ceratomyxa constricta (Fujita, 1923), resulting from the transfer of this species from Leptotheca to Ceratomyxa, is a senior secondary homonym of a morphologically distinct species, Ceratomyxa constricta Meglitsch, 1960. The senior name is retained, but the new replacement name C. centriscopsi (Meglitsch, 1960) nom. nov. is proposed for the latter, junior species, which is named after the host genus. Ceratomyxa elongata (Thélohan, 1895) n. comb. Syn. Leptotheca elongata Thélohan, 1895 (Fig. 12a–b) Description: Spores ovoid in sutural view; valves equal; sutural ridge not protruding. Polar capsules spherical. Comment: The name Ceratomyxa elonga (Thélohan, 1895), resulting from the transfer of this species from Leptotheca to Ceratomyxa, is a senior secondary homonym of a morphologically distinct species, Ceratomyxa elongata Meglitsch, 1960. The senior name is retained, but the new replacement name C. lepidopusi (Meglitsch, 1960) nom. nov. is proposed for the latter, junior species, which is named after the host genus. Ceratomyxa etrumuci (Dogiel, 1948) n. comb. Syn. Leptotheca etrumuci Dogiel, 1948 (no illustration) Description: Spores nearly spherical with two polar capsules located at anterior part of spore. Ceratomyxa faba Meglitsch, 1960 Syn. Leptotheca faba (Meglitsch, 1960) Zhao & Song, 2003 (Fig. 13) Description: Anterior margin of spores arched; posterior margin usually almost straight with slight concavity, with rounded ends; valves equal; sutural ridge prominent. Polar capsules spherical. Syst Parasitol (2010) 75:81–104 Figs. 21–41 Diagrammatic illustrations of Ceratomyxa spp. c spores. 21. C. longipes n. comb. (after Kalavati & MacKenzie, 1999); 22. C. lovei nom. nov. (after Moser et al., 1976); 23. C. lubati nom. nov. (after Lubat et al., 1989); 24a–b. C. macroformis n. comb. (after Gaevskaya & Kovaljova, 1984); 25a–c. C. macronesi n. comb. (after Chakravarty, 1943); 26. C. macrouridonum n. comb. (after Moser & Noble, 1975); 27a–b. C. minima n. comb. (after Meglitsch, 1960); 28. C. mylionis n. comb. (after Ishizaki, 1960); 29. C. nebulifera n. comb. (after Zhao & Song, 2003); 30a–b. C. noblei nom. nov. (after Noble, 1938); 31. C. opisthocornata n. comb. (after Evdokimova, 1977); 32. C. ovalis n. comb. (after Kovaljova & Gaevskaya, 1983); 33. C. parva (after Schulman, 1966); 34. C. pegusae n. comb. (after Kpatcha et al., 1996); 35. C. pinguis n. comb. (after Meglitsch, 1960); 36. C. polymorpha n. comb. (after Labbé, 1899); 37. C. quadritaenia n. comb. (after Kovalova & Zubchenko, 1984); 38. C. scissura n. comb. (after Davis, 1917); 39. C. sebastisci n. comb. (after Zhao & Song, 2003); 40. C. subelegans n. comb. (after Laird, 1953); 41. C. vikrami n. comb. (after Tripathi, 1949). Scale-bar: 10 lm Ceratomyxa fujitai nom. nov. Syn. Leptotheca limandae Fujita, 1923 (Fig. 15a–b) Description: Spores broadly oval to egg-shaped in sutural view with rounded ends; valves unequal; sutural ridge indistinct and not protruding. Polar capsules pyriform. Etymology: Named for Tsunenboui Fujita, who originally described the parasite. Comment: The name Ceratomyxa limandae (Fujita, 1923), resulting from the transfer of this species from Leptotheca to Ceratomyxa, would be a secondary homonym of a morphologically distinct species, Ceratomyxa limandae Fujita, 1923, described in the same work as the present species by Fujita (1923). Acting as First Reviser, we choose the latter of these two simultaneously described species as having priority and propose the replacement name C. fujitai nom. nov. for the species hitherto classified in Leptotheca. Ceratomyxa galeata Jameson, 1929 Ceratomyxa fisheri Jameson, 1929 Syn. Leptotheca fisheri (Jameson, 1929) Kudo, 1933 (Fig. 14a–b) Description: Spores ovoid; anterior margin convex and posterior margin straight; valves equal with rounded ends. Polar capsules large and round. 123 Syn. Leptotheca galeata (Jameson, 1929) Kudo, 1933 (Fig. 16) Description: Spores hat- or helm-like; anterior margin very strongly convex and posterior straight to slightly concave; valves equal, with brim-like ends. Polar capsules small and spherical. Syst Parasitol (2010) 75:81–104 91 21 24a 22 23 24b 25a 27a 25b 26 28 29 27b 25c 30a 31 34 33 32 30b 38 35 36 39 37 40 41 123 92 Syst Parasitol (2010) 75:81–104 Ceratomyxa hepseti (Thélohan, 1895) n. comb. Ceratomyxa lovei nom. nov. Syn. Leptotheca hepseti Thélohan, 1895 (no illustration) Description: Spore contour triangular in sutural view, with rounded ends. Syn. Leptotheca sebasta Moser, Love & Jensen, 1976 (Fig. 22) Description: Spore anterior margin arched and posterior margin moderately to deeply concave, with rounded ends; valves equal; sutural ridge prominent. Polar capsules spherical. Etymology: Named for Milton Love, one of the authors who originally described the parasite. Comment: The name Ceratomyxa sebasta (Moser, Love & Jensen, 1976), resulting from the transfer of this species from Leptotheca to Ceratomyxa, would be a secondary homonym of a morphologically distinct species, Ceratomyxa sebasta Moser, Love & Jensen, 1976, described in the same work as the present species by Moser et al. (1976). Acting as First Reviser, we choose the latter of these two simultaneously described species as having priority and propose the replacement name C. lovei nom. nov. for the species hitherto classified in Leptotheca. Ceratomyxa inconstans Jameson, 1929 Syn. Leptotheca inconstans (Jameson, 1929) Kudo, 1933 (Fig. 17a–b) Description: Spores variable in shape, generally rather stumpy with straight posterior edge, but sometimes appearing spherical; anterior margin strongly arched and posterior margin varying from straight to strongly concave; valves equal, with ends varying from strongly pointed to very blunt. Polar capsules small and ovoid. Ceratomyxa informis (Auerbach, 1910) n. comb. Syn. Leptotheca informis Auerbach, 1910 (Fig. 18a–b) Description: Spores broadly ovoid with slight curvature in the mid-region; valves smooth and equal; sutural line straight and distinct, not protruding. Polar capsules spherical (Kalavati & MacKenzie, 1999). Ceratomyxa kovaljovae (Bakay & Grudnev, 1998) n. comb. Syn. Leptotheca kovaljovae Bakay & Grudnev, 1998 (Fig. 19) Description: Spores ovoid with rounded ends; anterior margin arched and posterior margin slightly concave to slightly convex; valves equal; sutural line not protruding. Polar capsules spherical. Ceratomyxa latesi (Chakravarty, 1943) n. comb. Syn. Leptotheca latesi Chakravarty, 1943 (Fig. 20a–b) Description: Spores bean-shaped in sutural view, with both extremities rounded; valves smooth and symmetrical; sutural line prominent but sutural ridge not seen. Polar capsules spherical. Ceratomyxa longipes (Auerbach, 1910) n. comb. Syn. Leptotheca longipes Auerbach, 1910 (Fig. 21) Description: Spores reniform with ventral side slightly concave; valves smooth and equal; sutural line straight, not protruding. Polar capsules spherical (Kalavati & MacKenzie, 1999). 123 Ceratomyxa lubati nom. nov. Syn. Leptotheca chromis Lubat, Radujkovic, Marques & Bouix, 1989 (Fig. 23) Description: Spores round; anterior margin arched and posterior margin concave with rounded ends; valves equal; sutural line not protruding. Polar capsules sub-spherical. Etymology: Named for Vincent Lubat, one of the authors who originally described the parasite. Comment: The name Ceratomyxa chromis (Lubat, Radujkovic, Marques & Bouix, 1989), resulting from the transfer of this species from Leptotheca to Ceratomyxa, would be a secondary homonym of a morphologically distinct species, Ceratomyxa chromis Lubat, Radujkovic, Marques & Bouix, 1989, described in the same work as the present species by Lubat et al. (1989). Acting as First Reviser, we choose the latter of these two simultaneously described species as having priority and propose the replacement name C. lubati nom. nov. for the species hitherto classified in Leptotheca. Ceratomyxa macroformis (Gaevskaya & Kovaljova, 1984) n. comb. Syn. Leptotheca macroformis Gaevskaya & Kovaljova, 1984 (Fig. 24a–b) Syst Parasitol (2010) 75:81–104 93 Description: Spores ovoid in sutural view, with rounded ends; valves unequal; sutural line not protruding. Polar capsules spherical. Description: Spores elongate ovoid to subspherical; valves with rounded ends; sutural line distinct, with raised sutural ridge. Polar capsules spherical. Ceratomyxa macronesi (Chakravarty, 1943) n. comb. Ceratomyxa noblei nom. nov. Syn. Leptotheca macronesi Chakravarty, 1943 (Fig. 25a–c) Description: Spores elliptical, more or less flattened in sutural view, with rounded ends and one smaller than other; valves smooth, unequal and thin; sutural line fine; sutural ridge not prominent. Polar capsules spherical. Ceratomyxa macrouridonum nom. nov. Syn. Leptotheca minuta Moser & Noble, 1976 (Fig. 26) Description: Spores with anterior margin convex and posterior margin slightly convex or flattened in sutural view; valves equal; straight, distinct sutural line not protruding. Polar capsules spherical. Etymology: Named after the host family. Comment: The name Ceratomyxa minuta (Moser & Noble, 1976), resulting from the transfer of this species from Leptotheca to Ceratomyxa, would be a junior secondary homonym of a morphologically distinct species, Ceratomyxa minuta Meglitsch, 1960, therefore, the replacement name C. macrouridonum nom. nov. is proposed for the present species. Ceratomyxa minima (Meglitsch, 1960) n. comb. Syn. Leptotheca minima Meglitsch, 1960 (Fig. 27a–b) Description: Spores stubby with broadly rounded ends; anterior margin usually convex and occasionally straight; valves unequal; sutural line not protruding. Polar capsules spherical. Ceratomyxa mylionis (Ishizaki, 1960) n. comb. Syn. Leptotheca mylionis Ishizaki, 1960 (Fig. 28) Description: Spores reniform, with smooth asymmetrical valves; sutural line forms faint ridge. Polar capsules spherical, with short necks and equal in size. Ceratomyxa nebulifera (Zhao & Song, 2003) n. comb. Syn. Leptotheca nebulifera Zhao & Song, 2003 (Fig. 29) Syn. Leptotheca elegans Noble, 1938 (Fig. 30a–b) Description: Spores egg-shaped in sutural view, with rounded ends; valves slightly unequal; sutural line indistinct, not protruding. Polar capsules pyriform. Etymology: Named for Elmer Noble, who originally described the parasite. Comment: The name Ceratomyxa elegans is preoccupied by a morphologically distinct species described by Jameson (1929). Ceratomyxa opisthocornata (Evdokimova, 1977) n. comb. Syn. Leptotheca opisthocornata Evdokimova, 1977 (Fig. 31) Description: Spores variable; anterior margin generally arched and posterior margin strongly concave, with rounded ends, rarely spherical; valves unequal; sutural ridge not protruding. Polar capsules spherical. Ceratomyxa ovalis (Kovaljova & Gaevskaya, 1983) n. comb. Syn. Leptotheca ovale Kovaljova & Gaevskaya, 1983 (Fig. 32) Description: Spores ovoid in sutural view, with rounded ends; valves equal to unequal; sutural line not protruding. Polar capsules spherical. Ceratomyxa parva (Thélohan, 1985) Kudo, 1933 Syn. Leptotheca scombri Pogoreltseva, 1964 (Fig. 33) Description: Spores arcuate, anterior margin convex and posterior margin concave, valves equal with rounded ends. Polar capsules spherical. Comments: Schulman (1966) suggested L. scombri Pogoreltseva, 1964 be revised because a previously described species, C. parva (Thélohan, 1985), from the same host (Scomber scomber) had spores and polar capsules that were identical in shape but slightly larger (8–9 9 12–14 vs. 3–4 9 8–10 lm). Rather than proposing a new combination, we take the conservative approach and place L. scombri as a synonym of C. parva based on similarities in 123 94 Syst Parasitol (2010) 75:81–104 Table 2 Morphological features of the species of Ellipsomyxa Species Host Tissue Locality Length Thickness PCL PCW Reference E. fusiformis (Davis, 1917) n. comb. Sphyrna zygaena (= Cestracion zygaena) GB Off USA 9 16 4.5 4.5 Davis (1917) Mean and/or range are provided in micrometres. Tissue location listed: GB gall-bladder morphology and type-locality (Mediterranean and Black Seas). valves equal with 2 ribbon-like appendages protruding from each; sutural ridge indistinct and not protruding. Polar capsules spherical. Ceratomyxa pegusae (Kpatcha, Diebakate & Toguebaye, 1996) n. comb. Ceratomyxa scissura (Davis, 1917) n. comb. Syn. Leptotheca pegusae Kpatcha, Diebakate & Toguebaye, 1996 (Fig. 34) Description: Spores ovoid to egg-shaped in sutural view, with rounded ends; valves unequal; sutural line not protruding. Polar capsules spherical. Syn. Leptotheca scissura Davis, 1917 (Fig. 38) Description: Spores ovoid with flattened posterior margin and rounded ends in sutural view, valves equal, sutural line distinct but not protruding. Polar capsules spherical. Ceratomyxa pinguis (Meglitsch, 1960) n. comb. Ceratomyxa sebastisci (Zhao & Song, 2003) n. comb. Syn. Leptotheca pinguis Meglitsch, 1960 (Fig. 35) Description: Spores stubby, with strongly arched anterior margin and broadly rounded ends; valves equal in Peltorhamphus novaezelandiae and unequal in Arnoglossus scapha (=Caulopsetta scapha); suture distinct, narrow and elevated. Polar capsules spherical. Syn. Leptotheca sebastisci Zhao & Song, 2003 (Fig. 39) Description: Anterior margin extremely convex and posterior margin concave; spores roughly crescentshaped; ends of valves rounded. Polar capsules spherical. Ceratomyxa polymorpha (Labbé, 1899) n. comb. Syn. Leptotheca polymorpha Labbé, 1899 (Fig. 36) Description: Spores round; anterior margin arched and posterior concave, with rounded ends; valves equal; sutural line not protruding. Polar capsules pyriform. Comment: The name Ceratomyxa polymorpha (Labbé, 1899), resulting from the transfer of this species from Leptotheca to Ceratomyxa, is a senior secondary homonym of a morphologically distinct species, Ceratomyxa polymorpha Meglitsch, 1960. The senior name is retained, but the new replacement name C. physiculusi (Meglitsch, 1960) nom. nov. is proposed for the latter, junior species, which is named after the host genus. Ceratomyxa quadritaenia btschenko, 1984) n. comb. (Kovaljova & Syn. Leptotheca subelegans Laird, 1953 (Fig. 40) Description: Spores elliptical to egg-shaped in sutural view, with rounded ends; valves generally equal in size and shape, with anterior notch at sutural ridge; sutural ridge slightly protruding. Polar capsules spherical. Ceratomyxa vikrami (Tripathi, 1948) n. comb. Syn. Leptotheca vikrami Tripathi, 1948 (Fig. 41) Description: Spores broadly ovoid in sutural view, with rounded ends; valves unequal; sutural line not protruding. Polar capsules spherical. Zu- Syn. Leptotheca quadritaenia Kovaljova & Zubtschenko, 1984 (Fig. 37) Description: Spore anterior margin arched and posterior margin slightly concave, with rounded ends; 123 Ceratomyxa subelegans (Laird, 1953) n. comb. Ellipsomyxa Køie, 2003 One species formerly belonging to Leptotheca is assigned to Ellipsomyxa on the basis of morphological similarities. This species is from the gall-bladders Syst Parasitol (2010) 75:81–104 95 Fig. 43 Diagrammatic illustrations of Myxobolus lalithakumarii nom. nov. (after Lilitha Kumari, 1969): a. side view, b. apical view. Scale-bar: 10 lm 42a 43a 43b 42b Fig. 42 Diagrammatic illustrations of Ellipsomyxa fusiformis n. comb. (after Davis, 1917): a. side view, b. apical view. Scale-bar: 10 lm of marine sharks and, with minor amendments (given in bold below), fits the generic diagnosis of the latter genus. A synopsis and notes on the species is provided below and complemented with morphometric and host data (Table 2). appears distinctly fusiform. Sutural plane slightly oblique to longitudinal axis; sutural line forms distinct ridge. Polar capsules spherical and distinct, located at some distance from capsular side and opening at opposite sides of spore. Myxobolus Bütschli, 1882 Amended diagnosis Spores elongate in direction perpendicular to straight, central, transverse suture. Thin-walled valves hemispherical to elongate. Two equal spherical polar capsules either close to or at some distance from sutural plane on opposite sides of spore close to surface, discharging in opposite directions. Disporic plasmodia coelozoic in gall-bladders of marine fishes including elasmobranchs. (Modified after Lom & Dyková, 2006.) One species formerly belonging to Leptotheca is assigned to Myxobolus on the basis of morphological similarities and tissue locality. This species is from the gills, intestine and gall-bladder of a freshwater fish and fits the diagnostic characters of Myxobolus. A synopsis and notes on the species is provided below and complemented with morphometric and host data (Table 3). Ellipsomyxa fusiformis (Davis, 1917) n. comb. Syn. Leptotheca asymmetrica Lalitha Kumari, 1969 (Fig. 43a–b) Description: Spores variable, generally transversely elliptical or oval, rarely sub-spherical, with breadth being more than sutural diameter. Valves thick, uniform and ornamented; each has three longitudinal curved striae which run lengthwise, converging Syn. Leptotheca fusiformis Davis, 1917 (Fig. 42a–b) Description: Spores elliptical in apical view; base of valves circular in cross-section but are somewhat compressed distally parallel to—longitudinal plane, so that when viewed from—capsular side—spore Myxobolus lalithakumarii nom. nov. Table 3 Morphological features of the species of Myxobolus Species Host Tissue Locality Length Width PCL PCW Reference M. lalithakumarii nov. nov. [Syn. Leptotheca assymetrica] Osteochilus hasseltii (= O. neilli) Gi, In, GB India 8.6–10.0 5.7–7.9 3.6–4.3 2.1–2.9 Lalitha-Kumari (1969) Mean and/or range are provided in micrometres. Tissue location listed: Gi gills, In intestine, GB gall-bladder. Spore length and width are listed according to the guidelines for spore descriptions recommended by Lom & Arthur (1989) 123 96 towards thick ridge and prominent horizontal or central transverse striation. Polar capsules pyriform and unequal. Etymology: Named for Dr P. S. Lalitha Kumari, who originally described the parasite. Comment: The name Myxobolus asymmetricus (Lalitha Kumari, 1969), resulting from the transfer of this species from Leptotheca to Myxobolus, would be a junior secondary homonym of a morphologically distinct species, Myxobolus asymmetricus Parasi, 1912, therefore, the replacement name C. lalithakumarii nom. nov. is proposed for the present species. Sphaerospora Thélohan, 1892 Eighteen species formerly belonging to Leptotheca are assigned to Sphaerospora on the basis of morphological similarities or molecular evidence. Most of these species are from the urinary system of their hosts and all fit the morphological diagnosis of Sphaerospora. Lom & Dyková (2006) stated the diagnostic characters of Sphaerospora as ‘‘Spherical or subspherical spores, valvular diameter equals and only in a few species significantly exceeds the sutural diameter. Valves smooth or ridged, often with posterior or lateral protuberances or bumps, like in S. plagiognathopsis Chen & Hsieh, 1984. The sutural ridge is often prominent, polar capsules subspherical or pyriform. Two uninucleate sporoplasms. Mono- or disporic trophozoites are coelozoic, mainly in the urinary system of freshwater and marine fishes, some are histozoic’’. Sphaerospora spp. are also known from anuran amphibians (Lom & Dyková, 2006). All Leptotheca spp. we propose as belonging to Sphaerospora fit these criteria. However, molecular analysis has revealed that the genus is polyphyletic (Jirků et al., 2007). Jirků et al. (2007) have proposed that Sphaerospora spp. without DNA sequence data should be referred to as Sphaerospora (incertae sedis) to avoid the introduction of unnecessary taxonomic discrepancies. The only character that unites the ‘true’ Sphaerospora of Jirků et al. (2007), i.e. Sphaerospora (sensu stricto), is the presence of two areas within the V4 region of SSU rRNA with extensive nucleotide insertions, and this is the only case in myxozoan taxonomy that uses a molecular characteristic for the definition of a particular taxon (Jirků et al., 2007). Of the species we are proposing belong to Sphaerospora, 123 Syst Parasitol (2010) 75:81–104 only Leptotheca fugu has corresponding sequence data. Although, L. fugu forms a strongly supported clade with Sphaerospora (senso stricto) of Jirků et al. (2007), it does not have extensive nucleotide insertions in the V4 region and, therefore, does not fulfil the current criteria for Sphaerospora (senso stricto). Amendation of the diagnosis of Sphaerospora (senso stricto) to fit L. fugu would result in a diagnosis almost identical to that of Sphaerospora (sensu lato) given by Lom & Dyková (2006). It is not the purpose of this article to solve this problem, instead we propose to transfer L. fugu to Sphaerospora (sensu lato), until a better definition of the genus can be established, along with the other Leptotheca spp. reassigned here. A synopsis and notes on the species are provided below and complemented with morphometric and host data (Table 4). Sphaerospora armatura (Yoshino & Moser, 1974) n. comb. Syn. Leptotheca armatura Yoshino & Moser, 1974 (Fig. 44) Description: Spores semicircular, with anterior margin slightly flattened or rounded and concave posterior surface in sutural view; valves equal; distinct sutural ridge. Polar capsules spherical. Sphaerospora brevoides (Zhao & Song, 2009) n. comb. Syn. Leptotheca brevoides Zhao & Song, 2009 (Fig. 45a–b) Description: Spores ovoid to subspherical in sutural view; poster margin convex to straight; valves with rounded ends; sutural line not protruding. Polar capsules sub-spherical (Zhao & Song, 2003). Sphaerospora chagasi (Nemeczek, 1926) n. comb. Syn. Leptotheca chagasi Nemeczek, 1926 (Fig. 46) Description: Spores ovoid, almost spherical, with rounded ends; valves equal; suture distinct. Polar capsules spherical. Sphaerospora compressa (Noble, 1939) n. comb. Syn. Leptotheca compressa Noble, 1939 (Fig. 47) Description: Spores roughly pyramidal in sutural view, with rounded ends; valves equal; sutural ridge not protruding. Polar capsules sub-spherical. Species Host(s) (original name) Tissue Locality Length Thickness PCL PCW References S. armatura (Yoshino & Moser, 1974) n. comb Albatrossia pectoralis (Coryphaenoides pectoralis), C. leptolepis UB, RT Off California 12.9 (10–16) 20.9 (17–26) 5.7 (4–7) 5.7 (4–7) Yoshino & Moser (1974) S. brevoides (Zhao & Song, Chirolophis japonicus 2009) n. comb. (Azuma emmnion) UB China 6–8 7–10 2.1 (1.9–2.5) 1.7 (1.3–1.9) Zhao & Song (2009) S. chagasi (Nemeczek, 1926) n. comb. UD Brazil 10–11 15 8–8.5 8–8.5 Nemeczek (1926) S. compressa (Noble, 1939) Rimicola eigenmanni n. comb. UB Off California 10.5 12.2 3.3 4.0 Noble (1939) S. fugu (Tun, Yokoyama, Ogawa & Wakabayashi, 2000) n. comb. IE, R occ. S, GB Pacific Ocean Japan 9.0 (8.3–9.5) 14.0 (13.0–15.0) 2.8 (2.5–3.0) 2.8 (2.5–3.0) Tun et al. (2000) S. glomerosa (Davis, 1917) Paralichthys albigutta (P. n. comb. albiguttus) UB Off North Carolina 4.5 9 2 2 Davis (1917) S. koreana (Cho & Kim, 2001) n. comb. RT Off South Korea 8.59 13.42 3.86 3.86 Cho & Kim (2001) S. krogiusi (Konovalov & Oncorhynchus keta and Schulman 1965) n. comb. seven other salmonids UB Kamchatka 5.8–7.5 7.5–11 2.7–3.6 2.7–3.3 Schulman (1966) S. lobosa (Davis, 1917) n. comb. Paralichthys dentatus UB Off North Carolina 9–10 16–18 3 3 Davis (1917) S. lutjani (Kpatcha, Diebakate & Toguebaye, 1996) n. comb. Lutjanus fulgens Ki Off Senegal 6.06 (4.5–7.6) 8.95 (6.75–10.1) 2.12 (2–2.25) 2.12 (2–2.25) Kpatcha et al. (1996) S. mackenzii (Kalavati & Narayana Rao, 2005) n. comb. Fejervarya limnocharis (Lemnectus limnocharis) UB India 10.2 (8.8–11.6) 15.6 (14.4–18.0) 6.2 (4.8–6.8) 4.8 (4.6–5.0) Kalavati & Narayana Rao (2005) S. perlata (Gurley, 1893) n. comb. Gymnocephalus cernuus (Acerina cernua) – Off France (?) – – Kudo (1919) S. renicola (Thélohan, 1895) n comb. Scomber scombrus RT Off France 8 10 UB, Ki Onezhskaye Lake, Russia 6.5–7.5 7.5–8.5 3.5 3.5 Rumyantsev (1997) Atlantic Ocean 10.6–12.5 13.3–15.8 4.0 4.0 Kovaljova & Zubchenko (1984) Leptodactylus ocellatus Takifugu rubripes Sebastes schlegelii S. schulmani (Rumyantsev, Coregonus lavaretus, C. 1997) n. comb. albula Hippoglossus hippoglossus UB – Thélohan (1895) 97 123 S. simplex (Kovaljova & Zubtschenko, 1984) n. comb. – Syst Parasitol (2010) 75:81–104 Table 4 Morphological features of the species of Sphaerospora Mean and/or range are provided in micrometres, with the range in parenthesis if both were reported. Tissue location listed: UB urinary bladder, UD urinary ducts, R rectum, RT renal tubules, IE intestinal epithelium, S stomach, In intestine, Ki kidney, Ur ureters, Gl glomeruli, EG gut epithelium. Spore length or thickness given according to the guidelines for spore descriptions recommended by Lom & Arthur (1989) Zaika (1965) 3.5–4.2 9.8–10.5 7.0–8.4 S. subsphaerica (Zaika, 1965) n. comb Coregonus autumnalis migratorius, Thymallus arcticus 13 S. sphaerula (Noble, 1939) Gibbonsia metzi n. comb. 13 3.5–4.2 Lake Baikal RT, Ut Noble (1939) 5.0 Off California UB 123 4.6 6.02 (5–7.1) 10.65 (8.8–12.3) 2.91 (2.6–3.5) 2.75 (2.4– 3.2) Western Mediterranean coast, Spain Ki, RT, Ur, Gl, In, Ep Gu Dentex dentex, Sparus aurata S. sparidarum (SitjáBobadilla & AlvarezPellitero, 2001) n. comb. Thickness PCL PCW Length Locality Tissue Host(s) (original name) Species Table 4 continued Sitjá-Bobadilla & AlvarezPellitero (2001) Syst Parasitol (2010) 75:81–104 References 98 Sphaerospora fugu (Tun, Yokoyama, Ogawa & Wakabayashi, 2000) n. comb. Syn. Leptotheca fugu Tun, Yokoyama, Ogawa & Wakabayashi, 2000 (Fig. 48) Description: Spores trapezium-shaped, with flat or slightly curved anterior margin and slightly convex posterior margin; valves almost equal; sutural line indistinct and not protruding. Polar capsules spherical. Sphaerospora glomerosa (Davis, 1917) n. comb. Syn. Leptotheca glomerosa Davis, 1917 (Fig. 49) Description: Spores cylindrical, with rounded ends in sutural view; valves equal; sutural line not protruding. Sphaerospora koreana (Cho & Kim, 2001) n. comb. Syn. Leptotheca koreana Cho & Kim, 2001 (Fig. 50) Description: Spores broadly oval, spherical in sutural view, with rounded ends; valves equal with anterior notch at sutural ridge; sutural ridge not protruding. Polar capsules spherical. Sphaerospora krogiusi (Konovalov & Schulman, 1965) n. comb. Syn. Leptotheca krogiusi Konovalov & Schulman, 1965 (Fig. 51a–b) Description: Spores ovoid in sutural view, often with two or more growths on valves; valves equal; sutural line distinct, with keel-shaped projection. Polar capsules spherical (Schulman, 1966). Sphaerospora lobosa (Davis, 1917) n. comb. Syn. Leptotheca lobosa Davis, 1917 (Fig. 52) Description: Spores ovoid, with tapering ends in sutural view; valves unequal; sutural line prominent and forms sinuous ridge. Polar capsules spherical. Sphaerospora lutjani (Kpatcha, Diebakate & Toguebaye, 1996) n. comb. Syn. Leptotheca lutjani Kpatcha, Diebakate & Toguebaye, 1996 (Fig. 53) Description: Spores ovoid to spherical; valves equal; sutural line not protruding. Polar capsules spherical. Sphaerospora mackenzii (Kalavati & Narayana Rao, 2005) n. comb. Syst Parasitol (2010) 75:81–104 99 45a 44 46 47 45b 51a 48 50 49 51b 52 56 54 53 57 58 55 59 60 Figs. 44–60 Diagrammatic illustrations of Sphaerospora spp. spores. 44. Sphaerospora armatura n. comb. (after Yoshino & Moser, 1974); 45a–b. Sphaerospora brevoides n. comb. (after Zhao & Song, 2009); 46. Sphaerospora chagasi n. comb. (after Nemeczek, 1926); 47. Sphaerospora compressa n. comb. (after Noble, 1939); 48. Sphaerospora fugu n. comb. (after Tun, et al., 2000); 49. Sphaerospora glomerosa n. comb. (after Davis, 1917); 50. Sphaerospora koreana n. comb. (after Cho & Kim, 2001); 51a–b. Sphaerospora krogiusi n. comb. (after Konovalov & Schulman, 1965); 52. Sphaerospora lobosa n. comb. (after Davis, 1917); 53. Sphaerospora lutjani n. comb. (after Kpatcha, et al., 1996); 54. Sphaerospora mackenzii n. comb. (after Kalavati & Narayana Rao, 2005); 55. Sphaerospora perlata n. comb. (after Balbiani, 1883); 56. Sphaerospora schulmani n. comb. (after Rumyantsev, 1997); 57. Sphaerospora simplex n. comb. (after Kovalova & Zubchenko, 1984); 58. Sphaerospora sparidarum n. comb. (after Sitjá-Bobadilla & Alvarez-Pellitero, 2001); 59. Sphaerospora sphaerula n. comb. (after Noble, 1939); 60. Sphaerospora subsphaerica n. comb (after Zaika, 1963). Scale-bar: 10 lm 123 100 Syst Parasitol (2010) 75:81–104 Table 5 Replacement names for junior species with secondary homonyms created by transfer of Leptotheca species Replacement name Synonym Host (original name) Ceratomyxa centriscopsi (Meglitsch, 1960) nom. nov. Ceratomyxa constricta Meglitsch, 1960 Centriscops humerosus Ceratomyxa lepidopusi (Meglitsch, 1960) nom. nov. Ceratomyxa elongata Meglitsch, 1960 Lepidopus caudatus Ceratomyxa physiculus (Meglitsch, 1960) nom. nov. Ceratomyxa polymorpha Meglitsch, 1960 Pseudophycis bachus (Physciculus bachus) Sphaerospora dykovae (Dyková & Lom, 1982) nom. nov. Sphaerospora renicola Dyková & Lom, 1982 Cyprinus carpio Syn. Leptotheca mackenzii Kalavati & Narayana Rao, 2005 (Fig. 54) Description: Spores broadly oval and transversely elongated with round cornes; valves refringent, thick with 16–18 oblique striations; sutural line prominent and slightly curved, dumbbell shaped with broad ends. Polar capsules bulb shaped. Sphaerospora perlata (Gurley, 1893) n. comb. Syn. Leptotheca perlata (Gurley, 1893) (Fig. 55) Description: Spores elliptical; two small polar capsules convergent. Description: Spores ovoid; posterior margin slightly convex in sutural view, with rounded ends; valves equal; sutural ridge indistinct. Polar capsules spherical. Sphaerospora sparidarum (Sitjá-Bobadilla Alvarez-Pellitero, 2001) n. comb. & Syn. Leptotheca sparidarum Sitjá-Bobadilla & Alvarez-Pellitero, 2001 (Fig. 58) Description: Spores spherical in sutural view, with rounded ends; valves equal, each with posterior bulge in sutural view; sutural line conspicuous, straight with slightly protruding anterior and posterior notches. Polar capsules sub-spherical. Sphaerospora renicola (Thélohan, 1895) n. comb. Syn. Leptotheca renicola Thélohan, 1895 (no figure) Description: Spores globose, almost spherical. Polar capsules spherical. Comment: The name Sphaerospora renicola (Thélohan, 1895), resulting from the transfer of this species from Leptotheca to Sphaerospora, is a senior secondary homonym of a morphologically distinct species, Sphaerospora renicola Dyková & Lom, 1982. The senior name is retained, but the new replacement name S. dykovae (Dyková & Lom, 1982) nom. nov. is proposed for the latter, junior species, which is named for Iva Dyková, who originally described the parasite. Sphaerospora schulmani (Rumyantsev, 1997) n. comb. Syn. Leptotheca schulmani Rumyantsev, 1997 (Fig. 56) Description: Spores spherical in sutural view; valves equal, with an anterior and posterior notches at sutural ridge. Polar capsules spherical. Sphaerospora simplex (Kovaljova & Zubtschenko, 1984) n. comb. Syn. Leptotheca simplex Kovaljova & Zubtschenko, 1984 (Fig. 57) 123 Sphaerospora sphaerula (Noble, 1939) n. comb. Syn. Leptotheca sphaerula Noble, 1939 (Fig. 59) Description: Spores spherical in sutural view, with rounded ends; valves equal; sutural ridge not protruding. Polar capsules sub-spherical. Sphaerospora subsphaerica (Zaika, 1965) n. comb. Syn. Leptotheca subsphaerica Zaika, 1965 (Fig. 60) Description: Spores pyramidal in sutural view, with rounded ends; valves equal; sutural line distinct, appearing pointed at anterior pole and protruding. Polar capsules spherical. Conclusions The classification of myxosporean genera based on spore morphology has always been problematical, particularly when the boundaries between genera are based on small differences and the spores are known to be variable. The differences between Ceratomyxa and Leptotheca were based on whether the length of the spore exceeds the axial diameter of the spore or not, yet the spores of Ceratomyxa and Leptotheca spp. are known to be plastic (Jameson, 1929; Syst Parasitol (2010) 75:81–104 Sitjà-Bobadilla & Alvarez-Pellitero, 1993; Heiniger et al., 2008; Gunter et al., 2009). With the advances in molecular technology, the relationships between some of these biological characters are becoming apparent. Similarities in basic spore morphology do not tend to be good indicators of relatedness (Smothers et al., 1994; Andree et al., 1999; Kent et al., 2001; Fiala & Dyková, 2004; Whipps et al., 2004; Fiala, 2006; Jirků et al., 2007). However some biological correlates, such as tissue trophism, map well to the current phylogenies (Fiala, 2006; Burger et al., 2007). With this in mind we assign the majority of Leptotheca species to either Ceratomyxa or Sphaerospora based on the site of infection. Morphological traits of the species assigned to these genera also agree with the diagnosis of these genera for all species with the exception of two. L. fusiformis was assigned to Ellipsomyxa based on the unusual arrangement of polar capsules and L. asymmetrica was assigned to Myxobolus based on the possession of a longitudinal, rather than central, transverse suture. We have chosen to be conservative in our placement of species into only four genera, although for a few species their placement is uncertain. L. schulmani fits the diagnosis of Sphaerospora, but its spores are morphologically similar to those of Paramyxoproteus spp., which also infect the urinary system (Lom & Dyková, 2006). However, the original description is unclear as to whether the keel-like projections on the valves are actually keels. L. apogoni is another such species whose placement is questionable, and it may represent a new genus. This is the only species with trapezium-shaped spores with transversely striated valves from the gall-bladder of a marine fish and does not fit the diagnostic characters of any known genus. We take the conservative approach and place it within Ceratomyxa, although this species is the first representative of the genus with valves that are not smooth. Molecular analysis may change the placement of some species in the future; however, the aim of this review was to eliminate the ambiguity of the differences between Leptotheca, Ceratomyxa and Sphaerospora by suppressing Leptotheca. In summary we propose the following: (1) (2) Removal of the genus Leptotheca. Transfer of all Leptotheca spp. infecting the gall-bladder with polar capsules located at the anterior pole of the spore to Ceratomyxa. 101 (3) (4) (5) (6) (7) (8) (9) Amendment of the diagnosis of Ceratomyxa to ‘‘Spores elongated, generally crescent-shaped or arcuate, occasionally sub-spherical or ovoid, shell valves significantly exceed in length one half of the axial diameter of the spore. Shell valves conical or sub-hemispherical’’. Transfer of Leptotheca fusiformis to Ellipsomyxa. Amendment of the diagnosis of Ellipsomyxa to ‘‘Two equal spherical polar capsules either close to or at some distance from the sutural plane on opposite sides of the spore close to the surface, discharging in opposite directions. Disporic plasmodia coelozoic in gall-bladders of marine fishes and elasmobranchs’’. Transfer of Leptotheca asymmetrica to Myxobolus. Transfer of all Leptotheca spp. infecting the urinary system to Sphaerospora (sensu lato). 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