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A Guide to the Identification of ailedT Myxobolidae of the World: Dicauda, Hennegoides, Henneguya, Laterocaudata, Neohenneguya, Phlogospora, Tetrauromena, Trigonosporus and Unicauda

Eric J. Wagner

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Recommended Citation Wagner, Eric J., "A Guide to the Identification of ailedT Myxobolidae of the World: Dicauda, Hennegoides, Henneguya, Laterocaudata, Neohenneguya, Phlogospora, Tetrauromena, Trigonosporus and Unicauda" (2016). Textbooks. 1. https://digitalcommons.usu.edu/oer_textbooks/1

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ISBN Number 978-0-692-81303-4

Suggested citation format: Wagner, E.J. 2016. A guide to the identification of tailed Myxobolidae of the world: Dicauda, Hennegoides, Henneguya, Laterocapsulata, Neohenneguya, Phlogospora, Tetrauronema, Trigonosporus, and Unicauda. Fish Creek Records, Logan, Utah.

Cover art by Eric Wagner, inspired by M.C. Escher.

Preface

The impetus for this volume began with the Given the genetic data that has been finding of Henneguya species in the Logan River, generated in the last decade, it is apparent that Utah in the muscle of whitefish Prosopium the taxonomic relationships based on williamsoni. I quickly discovered there was no morphology may not fully characterize the true guide available to identify it. I resolved to evolutionary relatedness among myxozoan remedy that, and many years later, this species. So it is fully expected that the identification guide is a result. taxonomy of these species will change with The story of Henneguya and the other tailed time. Nevertheless, this summary should myxobolids began in Germany, with the first provide a benchmark and usuable summary of descriptions of spores by Müller (1841) and the known species and provide a framework for Creplin (1842). These later became Henneguya comparison with potential new species. In the psorospermica, the type species for the genus, past, several species have been described that and Henneguya (Myxobolus) creplini, through have already been published previously, in part, the works of Thélohan (1892) and Gurley due to a lack of this sort of guide. As part of this (1893), respectively. After Thélohan (1892) work, a few species have been synonomized created the genus Henneguya for tailed (Henneguya branchialis, H. alexandrinus, H. myxospores with two polar capsules, Davis nilotica, H. texta; See Henneguya-Taxonomic (1944) further subdivided the genus into Considerations section) or re-assigned to the Myxobilatus (striated spores with the suture proper genus, e.g. Hennegoides obpyriformis line between the polar capsules), Henneguya comb. nov., originally described as Henneguya (smooth spores with suture line around the obpyriformis (Ma et al. 1986). A parasite in spores, and Unicauda (spores with one tail elephantnose fish (Gnathonemus pertsii) made of different material than shell). There described by Caffara et al. (2007) was identified have been some efforts to summarize the as a unique species in this review and has been extant species of Henneguya, listing the metrics named Unicauda trentinii sp. nov. for each species in tabular form (Kudo 1920; I would like to thank several people who Eiras 2002; Eiras and Adriano 2012). Regional have assisted in making this document possible. summaries and, in some cases, keys also exist I am indebted to the staff at the Interlibrary for China (Chen and Ma 1998), Africa (Fomena Loan Office at Utah State University who helped and Bouix 1997) Brazil (Gioia and Cordeiro me obtain articles from obscure sources from 1996), and the U.S.S.R. (Bykhovskaya- around the world. I thank Patricia Lambert, Pavlovskaya et al. 1964). Cathryn Smith, Deborah Cartwright Iwanowicz, Given the trans-global shipment of fish, the and Maureen Judd for similar help in obtaining potential for movement of parasite species the original descriptions of species. I thank around the world is high. Hence, a broader Patricia Lambert for her support of my effort, as perspective is needed for an identification she was ‘widowed’ as I toiled away in my guide. This guide provides a summary of all the basement office month after month. tailed species of Myxobolidae described to date I would like to recognize the valuable from around the world. service provided by the Biodiversity Heritage Library for online access Cambridge University Press (Parasitology). (http://archive.org/details/biodiversity) to the Other publishers, requiring fees which were 19th century literature and to FishBase paid out of my pocket, are noted in Appendix B. (www.fishbase.org) for data on fish taxonomy. I thank Patricia Pilitt at the National For permission to redraw images from their Museum of Natural History for taking publications I thank the editors at Allen Press (J. photographs of Henneguya specimens in the Parasitol.), Current Science, Wiley and Sons (J. National Parasite Collection for me. I thank Fish Dis., J. Protozool., Trans. Am. Microsc. Michael Spooner for his guidance with the Assoc., J. Eukaryotic Microbiol.), American publication and insightful suggestions for Fisheries Society (J. Aquat. Animal Health), improving the guide. I thank the 3 anonymous University of Illinois (Illinois Biol. Monogr.), reviewers, Christine Swan, and Randy Oplinger Inter-Research (Dis. Aquat. Org.), Jagiellonian for taking the time to critically review the work. University (Acta Protozoologica), and

Dedication: To those who have the passion to those that have the fire to those who seek the truth with unwavering desire to those who thirst for knowledge to the curious at heart to those who find that answers are a brand new place to start. I salute your perserverence and the candles that you’ve lit while others curse the darkness or just don’t give a shit. When Saint Peter calls you and tallies up the list you’ll leave this world a better place —thank you for your gifts. Eric J. Wagner

Contents Introduction ...... 6 How to use this guide ...... 6 Key to Families within the phylum Myxozoa, Class Myxosporea...... 9 Key to Genera of Myxobolidae ...... 11 Dicauda Hoffman & Walker, 1978 ...... 13 Hennegoides Lom, Tonguthai, & Dykova 1991 ...... 14 Henneguya Thélohan 1892 ...... 16 Taxonomic considerations ...... 18 Freshwater Henneguya species ...... 22 Marine and Estuarine Henneguya species ...... 84 Key to the identification of Henneguya species with unequal polar capsules ...... 102 Henneguya species with unequal polar capsules ...... 105 Laterocaudata Chen et Hsieh 1984 ...... 112 Neohenneguya Tripathi 1952 ...... 113 Phlogospora Qadri 1962...... 113 Tetrauronema Wu, Wang et Jiang, 1988 ...... 114 Trigonosporus Hoshina 1952 ...... 115 Unicauda Davis 1944 ...... 116 Key to species of Unicauda ...... 118 References ...... 130 Appendix A (Henneguya species sorted by fish host taxon) ...... 151 Appendix B (Illustration Credits)……………….…………………………………………………………………………………… 159

Species Index ...... 164

Introduction The phylum Myxozoa contains over 2100 species of parasites that principally infect fish, although some species infect amphibian and reptilian hosts (Kudo 1920; Lom and Dykova 2006; Yokoyama et al. 2012). Myxozoans are multicellular, comprised of 1 to 7 spore shell valves, 1-2 infective germ sporoplasms, and 2 to 7 polar capsules (Lom and Dyková 1992). The polar capsules contain an extrudible filament used for attachment to a host, similar to the stinging cells of Cnidaria. After attachment, 1 to 2 amoeboid sporoplasms infect the target tissue and begin development. See Lom and Dyková (1992) for details on development of the trophozooite, plasmodia, and mature spores. Several myxozoan parasites studied to date have a life cycle with two hosts: fish and aquatic oligochaete or polychaete worms (Lom and Dyková 2006). The myxospore stage infects the worm host when ingested, which produces the actinospore stage that infects the fish host. Yokoyama et al. (2012) listed 33 myxozoan species for which the corresponding actinospore stage has been identified. These species include Myxobolus cerebralis (Wolf and Markiw 1984), the first species deduced to have a complex two- host life cycle. Other species within Myxobolidae include M. cotti (El-Matbouli and Hoffman 1989), M. carassii (El-Matbouli and Hoffmann 1993), M. arcticus (Kent et al. 1993), M. drjagini (El-Mansy and Molnar 1997a), M. portucalensis (El-Mansy et al. 1998), M. hungaricus (El-Mansy and Molnar 1997b), M. psuedodispar (Székely et al. 2001), M. bramae (Eszterbauer et al. 2000), M. macrocapsularis (Székely et al. 2002), and M. parviformis (Kallert et al. 2005). Among Henneguya species, only H. exilis, H. ictaluri, H. mississippiensis and H. neusslini have had the actinospore stage discovered (aurantiactinomyxon or triactinomyxon; Yokoyama et al. 2012; Rosser et al. 2015). So, many species have not had their complete life cycle determined yet. Likely, future taxonomic revisions of Myxozoa will consider the morphology of the actinospore stage, which varies among the known life cycles (e.g., hexactinomyxon, raabeia, triactinomyxon; Lom and Dyková 2006). See Yokoyama et al. (2012) for a key to these actinospore types.

The organisms in this guide are in the family Myxobilidae within suborder Platysporina, order Bivalvulida, class Myxosporea, phylum Myxozoa. The family Myxobolidae Thélohan 1892 is characterized by spores flattened parallel to the straight sutural line and two polar capsules (which may be unequal in length). The genera of Myxobolidae that lack tails, i.e., Myxobolus, Spirosutoria, Neothelohanellus, Thelohanellus, Neothelohanellus, are not covered in this guide. This summary covers the genera that have appendages that look like ‘tails’, i.e., Henneguya, Hennegoides, Neohenneguya, Unicauda, Dicauda, Laterocaudata, Phlogospora, and Tetrauronema, and Trigonosporus. The tailed spores of Myxobilatus (family Parvicapsulidae, suborder Variisporina), which look similar to Henneguya spores, but have striated shells and the suture line bisects the polar capsules, are also not included in this guide, other than in the family key. I have chosen to use the word ‘tail’ instead of ‘caudal appendage’ or ‘caudal projection’ to make the guide more user friendly and to use less space in the text boxes. For similar reasons I have chosen to use the word ‘cyst’ instead of ‘trophozooite’.

How to use this guide To identify an unknown species of Myxobolidae with a tail, first use the key to the families of Myxozoa on page 9 to identify the species to family. If the species is keyed to Myxobolidae, use the key to genera of this family on page 11. Note that some sections of the keys have three choices to pick from instead of two. Once you have keyed an organism to the genus level, go to the section of the guide for that genus. For identification within the Henneguya genus, it is further divided into 1) unequal polar capsules (both marine and freshwater, ≥ 0.5 µm mean length difference), 2) freshwater origin, and 3)

6 marine origin. Estuarine species are cross listed on both freshwater and marine sections. Within each of these sections you will find illustrations for each species, organized in order by total length, from smallest to largest. Because of incomplete descriptions (e.g., no data for some metrics or no range data), the amount of overlap among species, and the large number of species to consider, a dichotomous key was not considered practical for identification of Henneguya species. For Henneguya with unequal polar capsules and for Unicauda species, however, a key is provided in addition to the descriptions and figures.

So, to identify your unknown tailed myxobolids, the myxospore to be identified must be measured to determine the range of values for the various metrics that are used for identifying one species from another. The measurements to be made are shown in Figure 1 for a representative Henneguya myxospore. For polar capsule length and width, measure both capsules. At least 30 mature myxospores should be measured, preferably from fresh cysts. Formalin-fixed samples can also be measured, but some shrinkage can be expected (Parker and Warner 1970). Save some cysts or myxospores in 95% ethanol (preferable, though frozen samples work too) for potential DNA analysis. If multiple cysts are available, measure myxospores from several to insure you measure the full range of size variation1. Some species do not form cysts, so this is not always possible. Measure only the mature spores in the cyst, since cysts will have a mix of myxospores in various stages of development. In histological sections of polysporous trophozoites, the mature spores will typically be in the center of the cyst. For describing actinosporean stages, see Lom et al. (1997). See Lom and Dyková (1992) for guidance describing other myxospore types. In addition to these measurements, it is helpful to know the Figure 1. Representative Henneguya myxospore showing approximate size of the cyst (if present), the principal measurements used to differentiate among species. number of coils in the polar filament within the polar capsule, the host species and tissue infected, and whether the host is primarily a marine or freshwater species.

Once you have the measurements for your unknown myxospore and have determined the genus, start by identifying the species in the guide that overlap in total length with yours. Shaded rectangles for each spore indicate at a glance the range of values for total length and for spore body width. The figures display the relevant measurements, if available (ND = no data), from the original paper for the species. The figures are drawn to scale (note the ruler at top of the figure pages), redrawn from the original figures. The ruler range varies as the total length increases to provide clearer comparisons at the smaller scales. Continue to narrow down the list of possible species by comparing each of the other dimensions (spore body length & width, polar capsule length & width, tail length, etc.) and discarding

1 Roubal (1994) did not note much variation among different sized cysts, but some cysts may not be fully mature (spores with polar filaments coiled inside) 7 options that do not overlap with the range you have measured for each metric. The figures give quick guidance to the general shape of the spore (e.g., elongated versus rounded; proportion of the polar capsule length to the spore length) so that obviously different forms can quickly be withdrawn from consideration.

A corresponding text box contains additional information about the species host(s), host tissue, type location (site where the species was first identified from), the authority for the original description, and other relevant identifying character measurements, references, alternate hosts, and GenBank accession numbers. The data in the text boxes is also there to guide in deciding what species you have, showing what species overlap with it and what characters can be used to differentiate among the overlapping species. In some cases only the host species/family or host tissue differences can be used to differentiate among the known species.

Key to Families within the phylum Myxozoa, Class Myxosporea 1. a. Bivalvular, i.e., with two valves (shells) (Fig. 2) ……..…….2 - b. Multivalvular, with ≥ 3 valves (shells), radial symmetry Fig. 3)…………………………………………………………………………….. 11 2. a. (1a) Two polar capsules at opposite ends of the myxospore (e.g, Fig. 2a)…………………………………..……3 - b. Two polar capsules set apart, with capsule openings pointing at broad angle from each other (e.g. Figure 3. Apical view of Fig. 2b) Hexacapsula neothunni (redrawn from Lom and Figure 2. Examples of different (Ortholineidae/Sinuolineidae Dykova 1992) polar capsule location and complex)……………………………. 5 orientation among myxosporean - c. One, two, or four polar capsules clustered together (Fig. 2 taxa. c, Fig. 3) ……………………….……..6 3. a. (2a) Polar filament is folded, not spirally wound, short, flattened, wide at the base, tapering to tip (fig 4a); leaf-like plasmodia form polysporic pansporoblasts that float free in gall bladder of marine fishes. Sphaeromyxidae - b. Polar filament is coiled within polar capsule, similar thickness throughout its length (Fig. 4b)…………4 4. a. (3b) Ellipsoidal spores with pointed or bluntly pointed ends in which the two polar capsule openings are oriented in the same plane as the suture line, but in the opposite direction (Fig.2a) …..Myxidiidae Figure 4. Comparison of folded polar filament of - b. valves project like wings perpendicular to suture line (Fig. 5); gall Sphaeromyxidae (a) and bladder parasite in marine fish…..Fabesporidae coiled filament in other 5. a. (2b) Sutural line straight; coelozoic in marine and freshwater myxosporeans (b). fishes….Ortholineidae, genera Davisia, Myxoproteus, Schulmania of Sinuolineidae - b. suture line sinuous; coelozoic in urinary system of marine fishes…………Sinuolineidae 6. a. (2c) 1 or 2 polar capsules……….7 - b. 4 polar capsules…………………....8 7. a. (6a) single polar capsule and curved suture line; in gall bladder…Auerbachiidae Figure 5. Fabespora - b. single polar capsule with straight suture line, or two polar capsules nana. redrawn from ……9 Lom & Dykova 1992. 8. a. (6b) straight suture line, spherical or elongated spores that may have appendages ….Chloromyxidae - b. suture line curved, valves asymmetrical……. Parvicapsulidae

9. a. (7b) two polar capsules lie in the same plane as suture line, spores flattened parallel to the suture line (Fig 6a); one of the two polar capsules may be shorter or absent (i.e., 1 polar capsule)…..Myxobolidae b. two polar capsules lie in plane perpendicular to suture line (Fig. 6b,c)…10 10. a. (9b) In sutural view, the spore is tightly triangular around the polar capsule, with the shell forming wide Figure 6. Comparison of suture line wing-like lobular projections that are twisted from the orientation: in same plane as polar triangular portion (Fig. 7); coelozoic in gall bladders of capsules (Myxobolidae, a) vs. perpen- marine fish …….Alatosporidae dicular to polar capsule plane (Sphaerosporidae, b; Ceratomyxidae, c) b. Spore not tightly triangular around polar capsules; Elongate (wider than tall), wing-like projection as in Fig. 6c, not Fig. 7 ……………………...Ceratomyxidae c. spore shell valves not elongated perpendicular to suture line; spores circular to ovoid, with or without projections in the sutural plane (Fig.6b)…….Sphaerosporidae Figure 7. Alatospora samaroidea (redrawn from Lom & Dykova 1992). 11. a. (1b) 3 polar capsules per spore …...Trilosporidae b. 4 or more polar capsules per spore2…...….Kudoidae

2 species within the genera Pentacapsula ( formerly Pentacapsulidae), Hexacapsula (Hexacapsulidae), and Septemcapsula (Septemcapsulidae) are now assigned to Kudoidae based on recent genetic evidence (Whipps et al. 2004; Lom and Dyková 2006; Kaur and Singh 2012). 10

Key to Genera of Myxobolidae 1. a. spores with tail-like projections ……………………………….……..2 b. spores without tail-like projections ………………………………...9

2. a. (1a) ovoid spores that have a single tail-like projection …..3

b. spores with two or more tail-like projections or bifurcated for more than half the tail length, tail composed of same materials as the shell valve…………………………………………………..……..…4

3. a. (2a) single tail-like projection that often stains differently (pink) than the spore body (blue) when using Giemsa or that dissolves in sulfuric acid; projection is symmetrically located…….Unicauda Davis 1944 (Fig. 8a)

b. single tail-like projection, which may be forked at the very end, is of the same material as spore and projects asymmetrically (Fig. 8b)…Laterocaudata Chen & Hsieh 1984 4. a. (2b) tear-drop shaped spores with a single polar capsule, tail is bifurcated ..…… ……Phlogospora Qadri 1962 (Fig. 8c)

b. spores with two polar capsules and 2 or more projections…………………………..……5

5. a. (4b) spores with two projections ……6

b. spores with four projections ...….…..7

6. a. (5a) spores with two projections that point in opposite directions at a broad angle (Fig. 8d) …….Dicauda Hoffman & Walker 1978 b. spores with two projections that are in the same direction…………………….8

7. a. (5b) ellipsoidal shaped spores with two filamentous projections at each end of the spore; polar capsules situated asymmetrically from axis of the spore (Fig. 8e)……Neohenneguya Tripathi 1953

b. ovoid spore with four projections, two from Figure 8. Representative species from some genera of each shell valve, at the end of the spore Myxobolidae. a. Unicauda clavicauda, b. Laterocapsula mastacembala, c. Phlogospora mysti, d. Dicauda opposite the pair of polar capsules (Fig. atherinoidi, e. Neohenneguya tetraradiata, f. 8f)…...... Tetrauronema Wu, Wang & Jiang Tetrauronema macropodes, g. Trigonosporus acanthogobii, 1988 h. Hennegoides berlandi, & i. Thelohanellus horvorkai.

c. rhomboidal spore with four projections pointing in the opposite direction from the polar capsules; the projections are connected by transverse filaments that are attached to the projections, but not the spore body (Fig. 8g)……………………………….Trigonosporus Hoshina 1952

8. a. (6b) asymmetrical spores with non-axially projecting caudal appendage; polar capsules at an oblique angle from longitudinal axis of the spore body (Fig. 8h)….Hennegoides Lom, Tonguthai & Dyková 1991

b. symmetrical spores with coiled polar filaments, two tails, smooth shell valves (Fig. 2c)…Henneguya Thélohan 1892

9. a. (1b) single polar capsule…………………………………………….……….10

b. two polar capsules; ovoid, ellipsoid, or round shape……………11

10. a. (9a) spores tear-shaped, pyriform to broadly ellipsoidal with 1 polar capsule (PC) with coiled filament; axis of PC is parallel to long axis of the spore; smooth valves (Fig. 8i)…………..Thelohanellus Kudo 1933

b. smooth oval or egg shaped spores with 1 PC with long axis situated asymmetrically from axis of spore. The anterior end of the spores (where PC is) is broadly rounded, polar filament is coiled (Fig. 9)……..Neothelohanellus Figure 9. Das & Haldar 1986 (synonym: Lomosporus Gupta & Khera 1988) Neothelohanellus catlae. redrawn from c. spherical to ovoid spore at the base and pointed at the tip, which is recurved. Lom and Dykova 2006. Polar filament is irregularly coiled and indiscernible. The suture is bent and rarely visible. Gall bladder parasites of marine fish…Thelohanelloides Sarkar 20093

11. a. (9b) suture line is straight (Fig. 10b)…..Myxobolus Bϋtschli 1892

b. suture line is sinuous (Fig. 10a), slightly diverging from plane connecting the two polar capsules…..Spirosutoria Chen & Hsieh 1984

Figure 10. Comparison of the suture line shape between Spirosutoria (a) and Myxobolus (b).

3 Sarkar (2009) classified this new genus under Thellohanellidae. However, Lom and Dyková (1992) put Thellohanellus in Myxobolidae. Recent genetic evidence groups Thellohanellus with Myxobolus and Henneguya (Fiala and Bartošová 2010), so I have kept Thelohanellus and Neothelohanellus in Myxobolidae. Thelohanelloides has characters (irregularly coiled polar filament, gall bladder parasite of marine fish) that suggest it should be classified in Sphaeromyxidae, but it is provisionally located here until further genetic work clarifies its taxonomic position. 12

Dicauda Hoffman & Walker, 1978 Ovoid, flattened spores with two polar capsules and two rigid tails that extend in opposite directions. The tails extend from sutural groove, but are not an extension of the spore wall. Histozoic in freshwater fish.

The type species is Dicauda atherinoidi, was described by Hoffman and Walker (1978) from the subcutaneous tissue of the cyprinid Notropis atherinoides in western Lake Erie and New York, USA. No mucoid envelope was seen in India ink preparations. Thin-walled cysts of 0.1 to 1 mm occurred most often on the head and ventral body, occasionally on the fins or the lining of the mouth. No relationship was apparent between cyst size and spore maturity. Prevalence was highest in fish up to 55 mm total length during the first winter and during the next spring and early summer. Paired developmental stages were observed, suggesting disporous pansporoblasts, similar to some Unicauda. Measurements (µm) and other characteristics of fresh spores were {mean(range)}: spore body length… 10.3(8.5-12.5) spore body width… 9.3(8.5-11.0) spore body thickness… 7.3(7.0-9.0) polar capsule length… 5.2(4.0-7.0) polar capsule width… 3.3(2.5-4.0) polar filament coils… 4.3(3.0-6.0) Figure 11. Dicauda atherinoidi Hoffman et Walker 1978 tail length… 49(25-73) angle between tails (˚)… 70-170.

Formalin-fixed spores were slightly smaller, with a spore body of 7.9-11.1 x 7.1-10.3 µm, polar capsules of 3.2-5.5 x 2.4-4.0 µm, and tail length of 13-67 µm.

Hennegoides Lom, Tonguthai, & Dykova 1991 Asymmetric spore in valvular view, with two tails and two polar capsules. The principal distinguishing feature of the genus is that the two polar capsules are situated obliquely and asymmetrically from the longitudinal axis of the spore. Polysporous cysts found in gills, kidney, liver and intestinal mucosa. The type species is Hennegoides longitudinalis (Lom et al. 1991). The polar filament of Hennegoides longitudinalis is coiled along the long axis of the capsule, rather than perpendicularly as in Henneguya and Myxobolus, but other members of the genus have coils perpendicular to the long axis of the polar capsule. Hennegoides longitudinalis features an absence of pansporoblasts during development. The development of other Hennegoides species has yet to be studied.

Hennegoides longitudinalis Lom, Tonguthai, & Notes: Polar filament tapers gradually to the end. Dyková 1991 Type species for the genus. Shortest tail of all the Host: intestinal mucosa of Osphronemus gourami known Hennegoides species, as well as Cyst: 0.25 to 3 mm differences in body length and width, and the Type Location: Kanchnaburi Province, Thailand longitudinal coiling of the polar filament, easily separate this species from others. Hennegoides obpyriformis comb.nov. Notes: Differs in spore body length from other Ma, Wang, & Cai 1986 Hennegoides species. Formerly Henneguya Host: gills, spleen, liver of Noemachellus obpyriformis. yingjangensis, N. polytaenia Cyst: 100 to 153 µm Type Location: Taiping River, China Hennegoides berlandi Molnar, Székely, Mohamed, & Notes: The proportionality of the original spore Shaharom-Harrison 2006a drawing is not congruent with the reported width Host: gill of Pangasius hypophthalmus measurement. The true body width is likely Cyst: 2-4 mm on edge of gill filaments or 0.5-1.0 wider; indeed, micrograph bars in original mm round, bulging out from the side of the publication indicate spore is 6-10 µm wide. filaments Shorter body length than any other Hennegoides Type Location: Terengganu River, Malaysia species.

Hennegoides malayensis Molnar, Székely, Notes: The spore body is longer than that of Mohamed, & Shaharom-Harrison 2006a Hennegoides longitudinalis or H. berlandi, but Host: gill of Pangasius hypophthalmus shorter than that of H. pangasii. Polar filament is Cyst: inside cartilaginous gill rays; elliptical 40 x 60 17-20 µm long when extruded. or 40 x 300 µm Type Location: Terengganu River, Malaysia Polar capsule coils: 4 Hennegoides pangasii Molnar, Székely, Mohamed, Notes: Longest and widest spore body of the & Shaharom-Harrison 2006a known Hennegoides species. Host: gill of Pangasius hypophthalmus Polar filament length = 48-54 µm Cyst: 0.2-0.5 x 0.7-1.3 mm in central and apical regions of the gill filaments Type Location: Terengganu River, Malaysia Polar capsule coils: 9-10 (larger), 7-8 (smaller)

Henneguya Thélohan 1892 Symmetrical spore in valvular view, with two tails and two polar capsules. The filaments within the polar capsule are coiled perpendicular to the long axis of the spore and are uniform in diameter. Shell valves smooth. Spores oval, rounded, or fusiform in valvular view; biconvex in sutural view. The tail is a continuation of the shell, tapering to a fine point in the direction opposite the end of the spore with the polar capsules. Binucleate sporoplasm within the spore. The cysts, when present, are polysporic and typically milky or whitish-yellow in color. Most of the 219 described species are histozoic, particularly infecting gills (Appendix A). Coelozoic species infecting kidney, gut, gall bladder, ovary, and heart also exist.

The Henneguya species in this guide are split into three groups to aid in identification and discrimination among the species: 1) freshwater, p. 23, 2) marine, p. 91, and unequal polar capsules (both freshwater and marine), p. 111. Estuarine species are cross listed under both freshwater and marine. Anadromous fishes are in the freshwater group. A key is provided for the species with unequal polar capsules (difference of > 0.5 µm). For the freshwater and marine species, a key would be too onerous to use, so they are presented in order of total length, smallest to largest. In Appendix A, the host fishes and their corresponding Henneguya parasites are grouped by fish order, family, and species. The type species is Henneguya psorospermica described from gill cysts in yellow perch Perca fluviatilis by Thélohan (1892). H. tenuis is the smallest of the known Henneguya species, reaching a total length of only 4 μm (Vaney and Conte 1901). H. tenuis was found in mostly spherical cysts in the pyloric ceaca and conjunctive tissue of the alimentary canal of ruffe Gymnocephalus cernua L. (syn: Acerina cernua, A. vulgaris, G. cernuus) from France. In contrast, H. longicauda is the longest of the known Henneguya species, reaching a total length of 91-127 μm; it was found in gills of channel catfish Ictalurus punctatus from Mississippi, USA by Minchew (1977).

The economic impacts of Henneguya species can be significant. Henneguya Infections in muscle can reduce the quality and marketability of fish (Patashnik and Groninger 1964; Boyce et al. 1985). Some gonadal parasites may negatively affect reproduction, e.g., Henneguya testicularis, H. oviperda, but clinical cases and controlled studies have not been conducted yet to confirm this (Sitjá-Bobadilla 2009). Prevalence can be up to 75-100% (Nie 1996) and up to 8,000 cysts in a single fish have been reported (Fomena and Bouix 1996). As a result, gill infections can be severe, in some cases occluding the entire gill (Eiras et al. 2008). This can lead to suffocation, which can be significant in smaller fish (Duhamel et al. 1986; Whitaker et al. 2005). Mortality prevalences of up to 95-100% has been reported in farm-reared channel catfish infected with Henneguya (Bowser and Conroy 1965; Minchew 1972).

Clinical signs and lesions associated with Henneguya infection include lethargy, swollen gills, gill hyperplasia, and respiratory distress (Smith and Inslee 1980; Bowser and Conroy 1985; Eiras et al. 2008). In pond culture, fish parasitized with Henneguya sp. may congregate around high oxygen sites, such as the inflow or pond margins (Martins et al. 1997). Henneguya sebasta in heart muscle of the rockfish Sebastes paucispinis caused necrosis, hyperplasia, and hypertrophy of the connective tissue and smooth muscle (Heckman and Jensen 1978). A hematological study of striped bass Morone saxatilis infected with Henneguya noted that infected fish had significantly higher packed cell volume, resulting in higher blood viscosity (Lebelo et al. 2001). Caetano et al. (2012) observed Henneguya spores in the blood of Astyanax fasciatus, but it was not clear where the spores may have originated from in the fish.

Seasonal and spatial differences in infection prevalence have been noted by some authors. Cone (1994) documented an annual cycle in yellow perch Perca flavescens, in Nova Scotia, Canada, infected with Henneguya doori. The cycle was comprised of a summer infection, fall-to-winter development of plasmodia and sporogenesis, and spring release of spores. The settling rate of fresh H. doori myxospores was slow (8.34 x 10-6 m/sec), permitting broad dispersal (McConnell and Cone 1992). Seasonal changes similar to those observed by Cone (1994) have been reported by Shul’man (1989), Ashmawy et al. (1989), and Andrews (1979). In Hungary, Molnár (1998) observed young plasmodia of Henneguya creplini in the gills of Stizostedion lucioperca in April. These developed through the summer, reaching 800-900 µm by late autumn, and maturing by the end of winter. In central Finland, Haaparanta et al. (1994) noted mature cysts in Perca fluviatilis in spring and early summer, but early sporogonic stages of H. creplini were found throughout the year; no correlation was observed between prevalence and the pollution level of the four lakes studied.

Year to year differences in prevalence have also been noted (Molnár 1998; Urdes et al. 2010). For Henneguya salmonicola, which infects anadromous salmonids, the highest prevalence was observed among species spending more time in freshwater prior to seaward migration (Boyce et al. 1985). Differences among stocks of fish in parasite prevalence and diversity have been used as an indicator of stock origin (Quinn et al. 1987; Kent et al. 1994). Pronin and Batueva (2011) compared measurements of Henneguya zschokkei, H. cutanea, and H. cerebralis from different host species and locations; significant differences were noted for some metrics.

Chemical treatment of fish with Henneguya has been reported in only one case. The tapir fish Gnathonemus petersi infected with Henneguya was treated with the drug Toltrazuril; a 4-h bath in 10 µg/ml was effective against pansporoblasts, but did not control the mature spores (Schmahl et al. 1991). Some polymerase chain reaction (PCR) tests have been developed for diagnostic work, e.g., for H. ictaluri (and its corresponding actinospore and worm host), which causes proliferative gill disease in channel catfish (Whitaker et al. 2005).

There are a number of authors who have conducted ultrastructure examinations of Henneguya species with electron microscopy. Among these are studies of a gill parasite of the Brazilian fish Hoplosternum littorale (Azevedo and Matos 1989), the ovarian parasite Henneguya amazonica (Torres et al. 1994), H. pinnae (Schurbert 1968), H. paraensis (Velasco et al. 2016), H. melini (Mathews et al. 2016), H. jocu (Azevedo et al. 2014), H. aequidens (Videira et al. 2015) and other unknown species (Siau 1974). Ultrastructural aspects of sporogenesis were described for Henneguya exilis and H. adiposa in channel catfish (Current and Janovy 1977; Current 1979). The composition of the shell of Henneguya oviperda and Myxobolus pseudodispar was primarily Si (94%), with Ca, Mg, Fe, and Mn (Cu, Al, Ni, Cr in lesser amounts) comprising the remainder (Tyutyaev 2008). This shell is resistant to strong acids and bases (Kudo 1921), which is likely an adaptation to surviving digestion by the alternate host and by fish predators.

Fixed spores may vary in size slightly from fresh spores. Kudo (1921) compared a few different fixatives and found that minimal shrinkage of the myxosporidian Leptotheca ohlmacheri occurred with either Schaudinn’s fluid (2:1 mix of saturated HgCl and ethanol) or 50% alcohol, but shrinkage was worse with Bouin’s fluid or absolute alcohol. Kudo (1921) also noted that fixation made the polar filament invisible. Romuk-Wodoracki (1990) measured spores of Henneguya wolinensis in either glycerin-gelatin (Donetz and Shul’man 1973) or fresh; values were similar, though slightly larger in the glycerin-gelatin. A comparison of a few reports comparing fixed versus fresh spores is given in Table 2. 17

Table 2. Comparison of the effect of different fixatives on myxospore measurements.

Metric Mean change in value relative to fresh spores (µm) 50% alcohol: 70% alcohol: Schaudinn’s 4% 10% Glycerin- Giemsa1 Delafield1 fluid1 formalin1 formalin2 gelatin3 Total length -2.0 1.7 Spore length -2.5 0.6 Tail length -1.0 1.3 Body width 1.2 1.4 1.2 1.7 -0.2 0.9 Body depth 2.75 3.35 2.35 3.25 -0.5 0.9 Polar capsule -0.3 -0.5 length Polar capsule 0.5 0.6 0.6 0.6 -0.1 0.3 width 1Kudo 1921, Leptotheca ohlmacheri; 2Obiekezie & Enyenihi 1988, Henneguya chrysichthyi; 3Romuk-Wodororacki 1990, Henneguya wolinensis Taxonomic considerations Davis (1944) transferred several species previously assigned to the genus Henneguya to the new genus Myxobilatus. This genus was for spores that had shells with longitudinal striations and the suture line was perpendicular to the plane connecting the polar capsules (i.e., suture line between the polar capsules when viewing both). These species include Henneguya wisconsinensis (Mavor and Strasser 1916), H. rupestris (Herrick 1941), H. ohioensis (Herrick 1941), H. mictospora (Kudo 1920), and H. gasterostei (Parisi 1912). H. media (Thélohan 1890) and H. gasterostei (Parisi 1912) are considered to be members of Myxobilatus as well.

A few Henneguya parasites have been described but not named or compared with species already published. The Henneguya parasite described by Caffara et al. (2003) is likely H. mauritaniensis, which has dimensions that overlap with it and also infects the same host family. The Henneguya parasite described by Bahri et al. (1996) is likely H. pagri, which has dimensions that overlap with it and also infects the same host family. Bahri and Marques (2008) published a description and the parasite was later given the name Henneguya tunisiensis (Bahri et al. 2010). Meyers et al. (1977) described a Henneguya parasite from heart muscle of bluefish Pomatomux saltatrix; the only marine species that overlaps with it is Henneguya sebasta, but H. sebasta is not as wide-bodied (5.6-11.0 µm) as the unknown form (11.5 µm). Unfortunately other data is lacking on polar capsule dimensions, coil numbers, and developmental stages to adequately describe a new species. Alas, several species were not sufficiently described to differentiate them from other species and are not considered in this guide; these include Henneguya kolesnikovi (Gurley 1893), H. linearis (Gurley 1893), H. diplurus (Gurley 1893), and H. jubili (Qadri 1969).

Henneguya texta is considered a junior synonym of H. psorospermica, as explained below. Henneguya psorospermica was the first species of Henneguya, so named by Thélohan (1892) for the ‘Psorospermies de J. Müller’, who described tailed myxospores from the cell tissue of the eye muscle of ‘Hechtes’ (Pike), presumably Esox lucius (Müller 1841). Müller’s (1841) myxosporidian dimensions were given as “Der Längsdurchmesser des ovalen Körpers beträgt 0,0054, der Breitendurchmesser 0,0026 Linie.” Linie, in that time period, was equivalent to either 1/12 or 1/10th of an inch, which was either

1/10, 1/11th, or 1/12th of a foot (fuß; Wikipedia 2013). The foot varied widely from place to place, so interpretation of Müller’s measurements is difficult. Fortunately, another author from Germany provided the information needed for conversion 30 years later (Weltner 1892). Using the conversion provided in Weltner (1892), Müller’s (1841) dimensions were 12.1 μm for spore body length and 5.9 μm for spore body width; the tail length was 3-4 times the body length. For the form in Perca gills, Thélohan (1890) provides figures of the whole spore, gives the polar capsule length as 7 to 8 μm, but does not give other dimensions. When naming the species, he gave dimensions of 35-40 μm for total spore length and tail length approximating the same length as spore body (Thélohan 1892), but did not note which of the two hosts (pike or perch) these data were from. Based on Thélohan’s 1890 article, which notes the species examined, perch were studied, pike were not. So, the measurements were likely from the form infecting perch. The spore width for H. psorospermica (maximum of 7 μm) was provided later (Thélohan 1895). Notably, between the two descriptions, the tail length is 3-4 times the body length in the form infecting pike, and only about 1x for the form infecting perch.

Additional measurements of H. psorospermica have been subsequently reported by other authors. Lom (1961) measured spores found in the gills of Perca fluviatilis in Czechoslovakia: total spore length was about 36 μm, spore body length about 16 μm, spore body width 5-6 μm, polar capsules 8-9 x 2.2 μm. In comparison to the original description, the total length is within the range reported, but the width is slightly smaller and polar capsules slightly longer. Andrews (1979) found cysts (1-3 mm) on the gills of Perca fluviatilis from Llyn Tegid, the largest freshwater lake in Wales. Cysts were not found in any other tissue examined. The cyst occurred seasonally, present from November to May, but absent from June-October. Spore dimensions from samples taken at various times of the year were: spore body length 10.0-15.5 μm, spore body width 7.5-9.5 μm, body thickness 5.5-7.5 μm, tail length 19.0-32.0 μm, polar capsules 5.0-7.5 x 2.0-2.5 μm.

Thélohan (1892) described H. psorospermica as infecting gills of both pike and perch. However, Müller’s form infected a different tissue entirely. Gurley (1894) also lists both Esox lucius and Perca fluviatilis as hosts, but there is doubt in his statement “I believe we are justified in referring all the forms figured to one species, although fig. 34(pl.4) differs somewhat from the rest.” There are also a few errors in the dimensions given by Gurley (1894); the polar capsule length in Thélohan’s earlier description of the species (1890, p.207) is 7 to 8 μm, not 6-8 μm; also, 30 μm is given for the total length of the spore found in Perca, but the dimension given by Thélohan (1890) is actually for carp (species not given, but presumably is Cyprinus carpio). Thélohan (1895) later concedes that perhaps the myxobolid in Esox and Perca are two different species: “Peut-étre s’agit-il en réalité d’un organism différent; dans les figures donnés par Müller, la partie antérieure de la spore est plus renflée, la queue est plus longue, etc. Il pourrait donc se faire que le nom de Psorospermies ait été appliqué á une autre espéce que je n’ai pas eu la chance de retrouver.” (Maybe it really is a different organism; in the figures given by Müller, the anterior part of the spore is swollen, the tail is longer, etc. It may therefore be that the name of Psorospermies was applied to another species that I have not had the chance to observe.)

Cohn (1896) examined spores from the two different hosts and concluded they are different species. He assigned H. psorospermicus to the form in Esox, and gave the name H. texta to the form in Perca. The separation was based principally upon the difference in the cyst dimensions between the two forms (1.25 x 1.00 mm in Esox lucius, 0.75 x 0.375 mm in Perca fluviatilis). Complicating the assignation, however, is the different host tissue between Muller’s myxobolid (eye muscle) and Cohn’s (gills). Cohn (1896) also describes another myxobolid, Myxobolus (=Henneguya) lobosa, from the gill of

19 pike. As a further complication, Gurley (1893), assigned the name Myxobolus schizurus (now recognized as Henneguya shizura4) to the same myxobolid Müller (1841) described. Cohn (1896) also describes a third species found in gills of P. fluviatilis, H. minuta, the dimensions of which differed substantially from H. texta. Despite Cohn’s paper, in the following years, the linking of both hosts to H. psorospermica continues in the literature, e.g., Kudo (1920), Lom (1961), Andrews (1979), and Eiras (2002). Based on the tail length differences, the host species differences, and the different host tissue affinity, I believe that the myxobolid infecting perch gill and the myxobolid infecting pike eye muscle are two distinct species: H. psorospermica (perch gill) and H. shizura (pike eye muscle).

To sort out the taxonomic implications, we must also revisit the Henneguya parasites of Esocidae. Gurley (1893, 1894) added another host, Esox lucius, to the species H. creplini, based on Weltner’s (1892) observations of spores in the ovary with similar dimensions (spore body length 18 μm, body width 6.8 μm, tail 11 μm). Using the same data of Weltner, Cohn (1896) names the ovarian parasite Myxobolus oviperdus (currently recognized as Henneguya oviperda). A photomicrograph of H. oviperda by Tyutyaev (2008) shows the tail length about equal to the spore length. So, H. oviperda and H. creplini have very similar dimensions, but differ principally in host tissue affinity, and host family (and order). The combination of these two factors suggests that the myxobolid in the gill of Ruffe and the ovarian myxobolid in Pike are two different species.

So, are H. oviperda and H. creplini the same species? Are H. psorospermica and H. lobosa the same species? If so, that would imply that a Henneguya species has the ability to infect fish from two different taxonomic orders. Recent genetic studies on Henneguya and other myxosporeans have indicated that Henneguya species that infect the same host fish family are more related than those that infect other families (Lu and Nie 2004; Yokoyama et al. 2012). While studies on host specificity are needed for nearly all known Henneguya species, evidence suggests that a species that infects one fish family is unlikely to be able to infect a different family (Martins et al. 1999). In other myxozoans, such as the genus Kudoa, the species are related more by host and geography than by spore morphology (Hervio et al. 1997; Burger and Adlard 2011). For the species Myxobolus cerebralis, even species within the same family may be refractory to infection (O’Grodnick 1979), supporting the concept of a narrow genetic range of hosts for any particular myxosporean species. Further evidence is given by Yokoyama and Urawa (1997) who observed that the actinospore stage of Thelohanellus horvorkai would infect common carp, but not goldfish, which is also a cyprinid. Griffin et al. (2010) noted that after long term exposure to Henneguya ictaluri, prevalence was 66.2% in channel catfish Ictalurus punctatus, yet only 3.7% in blue catfish Ictalurus furcatus. In the current discussion, it is possible that Esocidae (Esociformes) and Percidae (Perciformes) share a common Henneguya parasite, but all the evidence to date suggests that the opposite is true.

Based on the above discussion, it is proposed that H. psorospermica be the name assigned for the myxobolid in perch gills described by Thélohan (1890, 1892, 1895), H. shizura (Gurley 1893) be the name assigned to the myxobolid in the eye muscles of Esox, and H. lobosa (Cohn 1896) be the name assigned to the myxobolid found in the gill of Esox species. This means that Henneguya texta, named for the form in perch gill by Cohn (1896), is a junior synonym of H. psorospermica. Another taxonomic implication of host order difference is that H. creplini (Gurley 1893) is the name for the myxobolid in

4 Gurley (1893, 1894), did not recognize the genus Henneguya, keeping all the tailed myxobolids in the genus Myxobolus. Cohn (1896) also did the same, but Labbé (1899), in his text on Sporozoa, later recognized Henneguya, setting the new standard. 20 ruffe and H. oviperda (Cohn 1896) is the name for the myxobolid in pike ovary (rather than H. creplini, which would have priority).

Another conflict in taxonomy involves the species H. sinensis (Chen and Hsieh 1960), H. branchialis (Ashmawy et al. 1989), and H. nilotica (Marwan 1998 as cited in Rabie et al. 2009), all of which infect fish within the genus Clarias. The measurements shown in the table below indicate broad overlap in measurements for parasites taken from the same host species. The parasite also has a broad list of tissues that it was found in. Since Henneguya sinensis has priority, H. nilotica and H. branchialis are considered junior synonyms. H. assiuti (Mandour et al. 1988) also infects gills of Clarias spp., but the spore body is shorter (9.8-13.5 µm) and polar capsules may be unequal or equal (Rabie et al. 2009).

Table 3. Comparison of measurements (all in µm, except for coil number; mean, with range in parentheses) and host data for Henneguya nilotica (Marwan 1998), H. branchialis (Ashmawy 1989), and H. sinensis (Chen and Hseih 1960). Data for H. nilotica was derived from Rabie et al. (2009).

H. nilotica H. branchialis H. sinensis

Host species Clarias lazera Clarias spp. Clarias batrachus, Channa argus, Misgurnus anguillicaudatus, Channa maculatus Host tissue Suprabranchial Suprabranchial organ, gill Intestine, ovary organ filaments, internal organs Total spore length (35.0-45.6) (27.9-41.2a) 42.7(31.8-52.2)

Spore body length (12.6-17.4) 14.5(13.3-15.4) 15.1(13.8-16.2)

Spore body width No data 5.6(4.7-5.6) 5.4(4.8-6.0)

Tail length 22-30 17.3(14.6-25.8) 27.6(18-36)

Polar capsule length 7.2-8.4 6.2(5.5-7.7) 7.9(7.2-8.4)

Polar capsule width 1.5-2.7 2.2(2.15-2.58) 1.9(1.4-2.4)

Polar capsule coils 7-9 No data 7-8

atotal length by addition of minima or maxima of spore length and tail length

Henneguya alexandrinus (Ashmawy et al. 1989) is considered a junior synonym of Henneguya laterocapsulata (Landsberg 1986). Both spores feature a unique one-behind-the-other arrangement of the polar capsules. Both species were found in Clarias spp. Both have similar dimensions (H. laterocapsulata presented first in the following paired comparisions): body length 13.8-16.0 versus 12.9- 34.4 µm, body width 3.7-5.3 versus 3.8-7.3 µm, polar capsules 4.1 -5.3 x 2.2-3.0 versus 4.9 x 2.9 µm, and tail length 15.2-20.2 versus 12.9-34.4 µm.

A number of Henneguya species have overlapping dimensions and can only be differentiated by fish host or fish tissue affinity. For example, H. yoffensis and H. mauritaniensis both infect fish of the family Sparidae, but the former infects gills and the latter heart tissue. H. salmonis and H. zschokkei both infect muscle tissue, but in different host species within Salmonidae. These species have been kept as separate distinct species until future genetic work can clarify their taxonomic status. 21

Freshwater Henneguya with equal polar capsules

Henneguya tenuis Vaney & Conte 1901 Notes: Distinguished from all other Henneguya by Host: connective tissue of gut, pyloric caeca of small size and short tail Acerina cernua Cyst: spherical, 30-150 μm Type location: France Henneguya renicola Schuurmans-Stekhoven Notes: Distinguished from other species of similar 1920 length by narrower body width and anterior spore shape which is narrower. Dimensions overlap with Host: kidney of Gasterosteus pungitius H. nandi, which infects a different family and host Cyst: none described tissue. Type location: Netherlands Tail length calculated by subtraction.

Henneguya cesarpintoi Guimarães 1931 Notes: Shorter body length than other species of similar length Host: branchial cavity of Astyanax fasciatus Cyst: none described Tail length calculated by subtraction. Type location: Agua Funda, São Paolo State, See also Gioia and Cordeiro 1996. Brazil

Henneguya tengchongensis Ma, Wang, & Cai Notes: Narrower body width than H. brevis, H. 1986 caudicula, & H. multimorpha

Host: gills of Pseudecheneis sulcata Cyst: oval, 167 x100 μm Type location: China

Henneguya brevis Thélohan 1892 Notes: No figure was present in the original article. Length was 15 µm, but it is not clear if this was total Host: kidney and ovaries of Gasterosteus length or spore length. Here I am assuming it is total aculeatus length. Cyst: none described Type location: not provided in description, but host Tail is longer than for H. caudicula, but shorter than species is found in Europe south to Iberian for H. multimorpha; body is wider than for H. Peninsula, and in circum-Arctic and temperate tengchongensis, but narrower than for H. ntemensis areas worldwide.

Henneguya caudicula Eiras, Takemoto, & Notes: Tail is shorter than other species of similar Pavenelli 2008 length, except H. ntemensis which has a wider spore body. Host: gills of Leporinus lacustris Cyst: 50 μm, white, polysporic Type location: Paraná River, Brazil Polar capsule coils: 3

Henneguya multimorpha Li & Wang 1993 Notes: Wider body width than H. renicola, H. tengchongensis, or H. bergamini, but narrower than H. Host: gills of Acanthogobius hasta ntemensis; shorter body length than for H. caudicula or H. Cyst: 115-163 x 115-136 µm, oval-elongate to ctenopomae, but longer than for H. rondoni; polar round capsules shorter than for H. camarounensis, H. Type Location: China periophthalmusi, or H. ovaliformis, and wider than for H. chydadea or H. hemiodopsis.

Henneguya ntemensis Fomena & Bouix 1996 Notes: Wider body width than other Henneguya species of similar total length, except H. ctenopomae which has Host: gall bladder wall, kidneys and spleen narrower polar capsules. of Brienomyrus brachyistus Cyst: none observed Type Location: Nyong & Ntem basins, Cameroon Polar capsule coils: 5-6 Polar filament length: 36.7(33-48) µm

Henneguya rondoni Azevedo, Casal, Matos & Notes: Shorter polar capsules than H. renicola or H. Matos 2008 cydadae; narrower body width than for H. multimorpha, Host: peripheral nervous system of H. ntemensis, H. camarounensis, Gymnorhamphichthys rondoni H. periophthalmusi, H. ctenopomae, H. ovaliformis, but Cyst: spherical-ellipsoid; up to 110 um long wider than H. bergamini. Type Location: Amazon River, Brazil Polar capsule coils: 6-7 Henneguya camerounensis Fomena & Bouix Notes: Longer spore body than for H. cesarpintoi, H. 1987 renicola, H. rondoni, but shorter than for H. legeri; longer polar capsule than for H. tengchongensis, H. caudicula, H. Host: gills of Synodontis batesii multimorpha, H. chydadea, H. santae or H. sichuanensis; Cyst: 135-490 x 75-300 um narrower body width than H. brevis, H. ntemensis, H. Type Location: Marigot Awout, Cameroon periophthalmusi, H. ctenopomae, but wider than H. Polar filament length: 29.0(20.0-38.0) µm bergamini or H. hemiodopsis narrower polar capsule width than H. ovaliformis. Henneguya chydadea Barassa, Cordeiro, & Notes: Distinguished from H. renicola by different host Arana 2003 tissue; longer tail than H. multimorpha; narrower spore body than H. ntemensis, H. periothalmusi, H. ctenopomae, Host: gills of Astyanax altiparanae but wider than H. bergamini; longer body length than H. Cyst: white, oval, 40-64 x 64-80 um rondoni, but shorter than H. legeri; shorter polar capsule Type Location: Campinas, São Paulo, Brazil length than for H. camerounensis, but longer than H. PC Coils: 9-10 sichaunensis; narrower PC width than for H. ovaliformis; more polar filament coils than for H. hemiodopsis (5-6); longer tail than H. santae, but shorter than H. yunannensis.

Henneguya periophthalmusi Wu, Notes: Wider spore body than H. with overlapping total length, Wang & Jiang 1993 except H. ntemensis, zikaweiensis, & renalis, which are wider, H. caudicula, ctenopomae, & legeri, which have longer spore body Host: gills of Periopthalmus length, and H. sichuanensis, which has shorter polar capsules; all cantonensis the spore dimensions overlap with H. brevis & H. multimorpha, but Cyst: oval-round, 0.2-0.3 mm H. brevis has a shorter tail on average and multimorpha polar Type Location: China; freshwater and capsules are shorter and narrower on average. estuarine Henneguya bergamini Guimarães 1931 Notes: Narrower spore body than other species of overlapping Host: peritoneal cavity of Astyanax total length fasciatus Cyst: none Found in Piroacicaba River, Brazil Type Location: Sao Paulo, Brazil See also Jakowska and Nigrelli 1953

Henneguya ctenopomae Fomena & Notes: Wider spore body than most H. with overlapping total Bouix 1997 lengths, except H. ntemensis, zikaweiensis, & miyairii, which have Host: gills of Ctenopoma nanum shorter body lengths, and H. chongqingensis, which has a longer Cyst: 100-140 x 70-100, spherical to tail. Spore dimensions overlap with H. renalis and percae, but the ovoid, between lamellae body form of H. percae differs notably and its polar filaments have 10-15 coils; H. renalis has wider polar capsules on average, 4-5 Type Location: Cameroon coils, and infects fish of a different family. PC coils: 3

Henneguya ovaliformis Ma, Wang & Notes: Narrower spore body width than H. periophthalmusi, Cai 1986 ntemensis, zikaweiensis, ctenopomae, legeri, & renalis, but wider than H. rondoni, bergamini, hemiodopsis, & zahoori. Longer spore Host: intestine of Channa striatus body length than H. renicola, sichuanensis, bleekeri, massii, Cyst: oval, 200-250 μm long leporinicola; shorter tail than for H. santae, pinnae, yunanensis, Type Location: China and pseudoplatystoma; longer polar capsule than H. multimorpha, but shorter than H. camerounensis; greater polar capsule width than H. chydadea. Henneguya hemiodopsis Azevedo, Notes: Narrower spore body width than most H. of overlapping Casal, Mendonça, & Matos 2009 total length, except H. bergamini which has an even narrower Host: gills of Hemiodopsis microlepis spore body, H. renicola & bleekeri which have shorter spore body Cyst: to 200 μm, asynchronous lengths, and H. zahoori, pinnae, macropodi, leporinicola, and Type Location: Poty River, Brazil psuedoplatystoma, which have longer tails. PC coils: 5-6

Henneguya santae Guimarães & Notes: Narrower spore body width than ntemensis, ctenopomae, Bergamin 1934 periophthalmusi, zikaweiensis, renalis, but greater width than Host: gills of Tetragnopterus santae renicola, hemiodopsis, zahoori, & pseudoplatystoma; shorter Cyst: oval, 1.0-1.2 x 0.70-0.75 mm spore body length than H. legeri, pinnae, visceralis, macropodi; Type Location: Pinheiros River, Sao shorter tail than H. chydadea, sichuanensis, massii, hainanensis, Paolo, Brazil gambusi, & leporinicola, but tail longer than for H. artigasi or PC Coils: ND ovaliformis; H. camerounensis has longer PC. Dimensions overlap with H. nandi & yunannensis, which have not been reported to infect fish of Characidae. See also Jakowska & Nigrelli 1953; Gioia

& Cordeiro 1996. 26

Henneguya nandi Gupta & Khera 1987 Notes: Longer tail and differs in spore body length and width and Host: gill and heart of Nandus polar capsule length from other H. of overlapping total length nandus except H. hemiodopsis, which has a narrower polar capsule. Cyst: rounded, opaque white, 399- Dimensions overlap with H. renicola, chydadea, santae, bleekeri, 450 x 342-402 µm yunnanensis, & garavelli, which can only be differentiated by the different host family or, in some cases (H. renicola, bleekeri & Type location: Ludhiana and Harike, yunannensis), different tissues. Punjab, India Polar filament length: 12-20 µm Henneguya legeri Cepede 1906 Notes: Longer spore body length and shorter tail than other H. of similar total length, except H. ntemensis, renalis & ctenopomae, Host: urinary bladder of Cobitis which have wider spore bodies, and H. camerounensis, chydadea, barbatula ovaliformis, hemiodopsis, artigasi, macropodi, & zahoori, which Cyst: none described have a narrower body width. No figure in original paper. Cepede named the species after his Type Location: France professor, Léger.

Henneguya artigasi Gioia & Cordeiro Notes: Longer spore body length than other H. of overlapping total 1987 length, except H. renalis, ovaliformis, & ctenopomae, which have a Host: gills of Astynanax scabripinnis wider spore body. Polar capsule coils: ND Cyst: 400 μm; rounded Type Location: Sao Paulo, Brazil Polar filament length: 21 μm Henneguya sichuanensis Ma 1998 Notes: Longer tail than other species of overlapping length, except (Chen & Ma 1998) H. psuedoplatystoma, which has a longer tail; H. zikaweiensis, ctenopomae, renalis, & percae, which have a wider spore body; H. Host: urinary bladder of Rhinogobius bergamini & zahoori, which have a narrower spore body; H. pinnae, typus miyairii, fonsecai, macropodi, visceralis & creplini, which have a Cyst: none described longer spore body; H. bleekeri, yunannensis, hainanensis & Type Location: Jiangjin, Sichuan gambusi, which have longer polar capsules. Measurements overlap Province, China with H. massii & leporinicola, but tail length of latter ranges higher; massii fish host is in Latidae, whereas sichuanensis host is in Polar capsule coils: ND Gobiidae.

Henneguya bleekeri Haldar & Notes: Shorter spore body length than other Henneguya of Mukherjee 1985 overlapping length, but longer than for H. leporinicola; narrower spore body width than for H. ntemensis, zikaweiensis, ctenopomae, Host: kidney of Mystus bleekeri renalis, periophthalmusi, massii, visceralis, hoimba, & percae, but Cyst: oval to round, whitish, 1.5-x 1.0 wider body than for H. zahoori; longer polar capsule length than for mm H. sichuanensis; longer tail than for H. chydadea. Measurements Type Location: West Bengal, India overlap with H. renicola, santae, nandi, yunannensis, & garavelli, Polar capsule coils: 6-8 but these species infect other fish families than Bagridae; H. renicola is also found in kidneys, but its anterior spore shape is more sharply pointed.

Henneguya shariffi Molnar, Notes: narrower bodied, with longer polar capsules and shorter tails Székeley, Mohamed, Shaharom- than other H. of overlapping total length, except H. renicola, H. rondoni Harrison 2006 and H. bergamini, which have shorter spore bodies. Dimensions overlap Host: gills of Pangasius with H. macropodi, which has a longer polar filament, slighter larger hypophthalmus cysts, and infects fish of a different taxonomic order. Cyst: 0.2-0.5 mm Type Location: Terengganu River, Malaysia Polar capsule coils:7 Polar filament length: 15.3 ± 4.0 (13.2-21.0) Henneguya zikaweiensis Sikama Notes: Wider spore body than most other H. of overlapping total 1938 length, except H. percae, creplini & minuta, which are wider; H. ntemensis, pinnae, hoimba, daoudi & H. latesi, which differ in polar Host: eye cornea of Carassius capsule dimensions; H. ctenopomae, renalis, salvelini, miyarii, auratus nanhaiensis, & chongqingensis, which have a longer spore body; H. Cyst: 350-485 x 230-480 µm, tchangi & nüsslini, which are deeper bodied; and H. psorospermica & H. white, round or oval cerebralis, which have longer tails. Measurements overlap with H. guanduensis, but H. zikaweiensis is more broadly rounded anteriorly Type Location: creek in Zikawei, than guanduensis and does not get as long in body or total length. Total near Shanghai, China length by addition of maxima, minima. Polar capsule coils: 5 See also Chen and Ma 1998 Henneguya pinnae Schubert 1967 Notes: Shorter polar capsule length than other H. of overlapping length, except H. renicola, santae, sichuanensis, bleekeri, massii, & leporinicola, Host: fins of Ctenopoma which have shorter spore bodies, and H. artigasi, percae, manipurensis, kingsleyae & corruscans, which have longer spore bodies. Longer tail than for H. Cyst: spherical, up to 0.7 mm ntemensis, legeri, hemiodopsis, but shorter tail than H. tchangi, Type Location: West Africa pseudoplatystoma, or suprabranchiae. Thinner body than for H. Polar capsule coils: 5 gambusi. Measurements overlap with H. miyairii, which also infect skin, but infects a different fish family. See also Schubert 1968. Henneguya massii Kostoingue, Notes: Caudal processes well separated; the spore body width is less Diebakate, Faye & Toguebaye 2001 than other species of overlapping length, except H. renicola, zahoori, Host: gills of Lates niloticus travassosi, hainanensis & leporinicola, which are narrower; tail length is Cyst: spherical, 0.5-1.0 mm longer than for H. periophthalmusi, camerounensis, chydadea, legeri,& Type Location: Chari River, Chad ovaliformis; spore body length is shorter than for H. hemiodopsis, Type specimen: N˚ Myxo-096, Biol. artigasi, pinnae, visceralis, macropodi, fonsecai, manipurensis, gambusi, Dept., Univ. C.A. DIOP, Dakar miyairii, & pseudoplatystoma; polar capsule length is shorter than for H. yunannensis. Measurements overlap with H. santae & sichuanensis, but these infect different fish families. Henneguya visceralis Jakowska & Notes: The longer polar capsules that comprise more than half the Nigrelli 1953 length of the spore body differentiates this species from most others of Host: kidney, liver, heart, similar length; narrower spore body width than H. renalis, hainanensis, stomach, spleen, gall bladder, & miyairii, but wider spore width than H. zahoori; longer spore body mesentery of Electrophorus length than H. santae, sichuanensis, bleekeri, or massii, but shorter than electricus H. artigasi; Polar capsule width differentiates H. visceralis from H. Cyst: white, solid, flat or raised macropodi & percae; longer tail than H. legeri. Type Location: Brazil Polar filament length: to 44 µm 30

Henneguya iheringi Pinto 1928 Notes: Unique shape distinguishes this from other spores of similar total length. Shorter polar capsule than H. hoimba and Host: gills (?) of Serrasalmo spilopleura not as wide. Cyst: unknown Type location: Rio Turvo, Pirangy, Brazil Polar capsule coils: ND

Henneguya yunannensis Ma, Wang & Cai Notes: Spore body width differs from most H. of overlapping 1986 total length, except: shorter body length than for H. artigasi, leporini, & wenyoni, but longer than for H. leporinicola; thinner Host: abdomen and tail of Glyptothorax body than for H. gambusi; shorter polar capsule length than for trilineatus H. camerounensis, visceralis, macropodi, fonsecai, Cyst: oval, 3 x 2.4 mm; manipurensis, corruscans, schizodon, but longer than for H. 267 x 142 μm sichuanensis, pinnae, massii, shorter tail than H. dini, malabarica, & suprabranchiae, but longer than for H. chydadea Type Location: China or ovaliformis. Measurements overlap with H. garavelli, santae, nandi & bleekeri, but these infect different fish families and different tissues (gill or kidney).

Henneguya hoimba Cordeiro & Gioia 1987 Notes: The description lacks data on variance, but the stout club-shape of the spore is distinctive from other species. Host: gills of Astyanax fasciatus Differences in spore body width or polar capsule length Cyst: round, white differentiate this species from others of overlapping length, except H. tchangi, which has wider polar capsules.

Type Location: Atibaia River, São Paulo, Brazil

Henneguya macropodi Shiba 1934 Notes: Narrower body width, shorter tail, and longer polar capsules than other H. of overlapping total length, except H. Host: skin and fins of Macropodus renicola, santae, sichuanensis, bleekeri, & massii, which have a opercularis (syn: M. chinensis) shorter spore body length; H. creplini, artigasi, Cyst: 0.3-0.7x 0.4 mm, milky-yellow multiplasmodialis & pisciforme, which have longer spore body Type Location: Keijo, Japan lengths; H. zahoori, which has a narrower spore body; H. visceralis, percae, & corruscans, which have a narrower polar Polar filament length: 35-47 μm. capsule width. Dimensions overlap with H. camerounensis, H.

shariffi, & ntondei, which infect different fish families. Total length by addition.

Henneguya zahoori Bhatt & Siddiqui 1964 Notes: Narrower body width than other spores of overlapping total length, except H. renicola & hemiodopsis, which have Host: gills of Ophiocephalus punctatus shorter tails, and H. gracilis, which has a longer tail. Cyst: 0.5 mm; pyriform Museum specimen: Aligarh Muslim Univ., India Type Location: India

Henneguya fonsecai Guimaraes 1931 Notes: Medium-sized spore that differs from other H. of overlapping length in body width or tail length, except H. Host: fins and cutaneous tissue bleekeri, massii, & leporinicola, which have a shorter spore body, of Leporinus copelandi H. multiplasmodialis, corruscans, leporini, & pisciforme, which Cyst: spherical, white, 1.5-3 mm have a longer spore body; H. pinnae, visceralis, macropodi, Type Location: Paraiba River, São Paulo, manipurensis, & gambusi, which differ in polar capsule length. Brazil See also Jakowska & Nigrelli 1953

Henneguya manipurensis Hemanand Notes: Longer spore body length than other H. of overlapping Meitei, Bandyopadhyay, & Mitra 2008 length, except H. ctenopomae, creplini, hoimba, renalis, percae, zikaweiensis, salvelini, macropodi, bleekeri, multiplasmodialis, Host: muscle of Anabas testudineus, corruscans, malabarica, schizodon, leporini, minuta, pisciforme producing skin ulcers & nuesslini, which differ in spore body width; H. yunnanensis, Cyst: no cyst testicularis, garavelli, and wenyoni, which have shorter polar Type Location: Manipur, India capsules; and H. daoudi, suprabranchiae, nanhaiensis & Type specimens: HP1/2006, HP2/12, guanduensis, which have longer tails. Manipur Univ., Life Sci. Dept.

Henneguya renalis Sarkar, Mazumder, & Notes: Wider spore than other H. of overlapping total length, Pramanik 1985 except H. ntemensis, zikaweiensis, latesi, miyairii, & cerebralis, which have shorter spore body lengths; H. creplini, pinnae, & Host: kidney of Channa marulius percae, which differ in polar capsule length; and H. salvelini, Cyst: not observed psorospermica & occulta, which have longer tails. Dimensions Type Location: West Bengal, India overlap with H. ctenopomae and doori, but H. ctenopomae has Polar capsule coils: 4-5 only 3 polar capsule coils, and H. doori has 9-14.

Henneguya hainanensis Chen & Ma 1998 Notes: Differs in spore body width from most other H. of overlapping total length except H. renicola, bleekeri, massii, Host: intestine of Misgurnus leporinicola, & mormyri, which have a shorter spore body anguillicaudatus length, H. creplini, artigasi, wenyoni, minuta, pisciforme, laterocapsulata & odzai, which have a longer spore body; H. Cyst: none described camerounensis & suprabranchiae, which have longer tails; polar capsule length differs from H. sichuanensis, pinnae, macropodi. Type Location: Hainan, China Dimensions overlap with H. fonsecai, travassosi, dini, maculosus

& garavelli, but these infect other fish families.

Henneguya schizodon Eiras et al. Notes: Narrower spore body than most other H. of overlapping 2004 length, except H. macropodi, which has a shorter tail; H. gracilis, Host: kidney of Schizodon fasciatus which has a longer tail; H. travassosi & leporinicola, which have a Cyst: whitish, 0.25-0.5 mm shorter spore body length; H. garavelli, psuedoplatystoma, & odzai, diameter, superficial or embedded which have shorter polar capsules; and H. laterocapsulata, which has in the tissue a wider polar capsule. Measurements overlap with H. dini, which infects gills (not kidney) of fish from a different family. Type Location: Amazon River at Holotype: INPA-001, Inst. Nacional de Pesquisas da Amazonia, Manaus, Brazil Manaus Polar capsule coils: 8-10 Henneguya travassosi Guimaraes & Notes: Narrower spore body and shorter polar capsule than most H. Bergamin 1933 of overlapping length, except H. zahoori, which is even narrower; H. yunnanensis, fonsecai, suprabranchiae, & pseudoplatystoma, which Host: muscle, skin of Astynanax differ in tail length; and H. dini, malabarica, wenyoni, leporinicola & fasciatus & Leporinus copelandi pisciforme, which differ in spore body length. Measurements Cyst: to 5 mm, white overlap with H. hainanensis, ntondei & garavelli, but these infect Type Location: Paraiba River, Brazil different tissues (intestine or gill) of different fish families. Henneguya gambusi Parker, Spall & Notes: Spore body is shorter & wider than most H. of overlapping Warner 1971 length, except H. hoimba, gambusi, tchangi, psorospermica, Host: skin of Gambusia affinis adherens, vovki, friderici, nyongensis, tegidiensis & shackletoni, Cyst: flattened, fresh cysts nearly which differ in polar capsule dimensions; & H. cartilaginis, which has colorless a longer tail. Thinner body than H. nuesslini. Overlaps with H. Type Location: USA, Oklahoma cerebralis & garavelli, but H. cerebralis target different tissues (cranial cartilage) in salmonids & H. garavelli infects a different fish Polar capsule coils: 3-5 family. Total length by addition. Polar filament length: 10 μm Henneguya percae Fantham, Porter Notes: Unusual polygonal shape of the spore body with a tail & Richardson 1939 undivided for more than half its length, as well as having a wider Host: gills of Perca flavescens spore body than most H. of similar length, distinguishes this species. Cyst: 0.25 mm, white Total length by addition. Divided part of the tail: 5.0-9.1 µm long. Type Location: Lake Memphremagog, Quebec, Canada Polar capsule coils: 10-15

Henneguya latesi Tripathi 1952 Notes: Shorter spore body and shorter polar capsule length than other H. of overlapping length, except H. bleekeri & massii, which Host: gills, buccal mucosa of Lates have even shorter bodies; H. zikaweiensis, ctenopomae, creplini, calcarifer sichuanensis, renalis, percae, tchangi, nuesslini, psorospermica, Cyst: small (size not given), travassosi, hainanensis, pseudoplatystoma, which differ in body containing only mature spores width; and H. santae & mormyri, which differ in tail length. Thicker body than for H. pinnae or H. yunnanensis. Dimensions overlap with Type Location: Chilca Lake, India H. dini & adherens, but these infect other fish families. Total length by addition. H. latesa (Wu et al. 1994) is found in the same fish host in South China Sea, but dimensions differ.

Henneguya tchangi Ma 1998 (Chen Notes: Differences in spore width or the wider polar capsule & Ma 1998) differentiate this species from H. with overlapping total length, except Host: urinary bladder of H. cerebralis, psorospermica, & tegidiensis, which have longer tails; and Schizothorax (Racona) davidi H. nanhaiensis, which has a longer spore body. Thicker bodied than H. Cyst: none described zikawiensis & H. daoudi. Dimensions overlap with H. guanduensis, but Type Location: Chongqin, China this infects fish of a different family and a different tissue. H. cuniculator Naldoni, Maia, da Notes: Polar capsule dimensions and differences in body length and Silva, & Adriano 2014 width differentiate this from other H. of overlapping total length Host: gills of Pseudoplatystoma except H. lobosa, which has a longer tail, & H. corruscans, maculosus, corruscans dini, fridirici, nagelii, guanduensis & multiplasmodialis, which have Cyst: elongated, to 1 cm long fewer polar filament coils. Dimensions overlap with H. macropodi, Type Location: Sao Francisco River, ntondei, leporini, & australis, which infect fish of different families. Minas Gerais, Brazil Polar Capsule coils: 10-11. Henneguya dini Kabre, Sakiti, Notes: Narrower spore body width than other H. overlapping length, Marques, & Sawadogo 1997 except H. travassosi, multiplasmodialis, corruscans, leporini, Host: gills of Heterotis niloticus leporinicola, pisciforme, laterocapsulata, odzai, pungitii, guanduensis, Cyst: 0.12-0.35 mm; oval & sinensis, which differ in spore body length; H. macropodi & fonsecai, Type Location: Burkino-Faso which have a shorter tail; H. mormyri, pseudoplatystoma, pellucida, Polar capsule coils: 5-8 nyongensis, shaharini, gracilis, suprabranchiae, & lobosa, which have longer tails. Fewer PC coils than H. schizodon or H. ntondei. Length/width: 3.01 Dimensions overlap with H. zahoori, hainanensis, gambusi, malabarica, garavelli, mystusia, australis & friderici, but these have not been reported in Arapaimidae. Henneguya multiplasmodialis Notes: Longer and narrower spore body and longer polar capsule than Adriano et al. 2012 H. of overlapping total length, except H. australis, nyongensis, & Host: gills of Pseudoplatystoma gracilis which have a longer tail, & H. doori & sinensis, which have a spp. longer polar capsule. H. corruscans also infects the same genus of fish Cyst: white, to 2.5 cm host, but is shorter in total length and has a slightly wider polar Type Location: Pantanal wetland, capsule. Dimensions overlap with H. maculosus & leporini; the latter Brazil has not been reported to infect species in Pimelodidae, but H. maculosus has; this species can only be differentiated by rDNA Polar capsule coils: 6-7 analysis. GenBank No.: KF296354, JQ654969 Henneguya miyairii Kudo 1920 Notes: Wider spore body than other H. of overlapping total length, (Miyairi 1909) except H. zikaweiensis, percae, latesi, cerebralis, minuta, creplini, doori, Host: subcutaneous tissue of varichorhinusi, salvelini, occulta, malapteruri & similis, which differ in head of Carassius auratus spore body length; H. ctenopomae & salmonis, which differ in tail Cyst: membrane thin, fibrous; length; H. tegidiensis, which has unequal lengths of the two tails. polysporous, 7-12 nuclei per Dimensions overlap with H. psorospermica, electrica, sarotherodoni, pansporoblast cartilaginis, shackletoni, pinnae & nuesslini; the latter 2 spp. also infect skin/fins, but not in Cyprinidae. Polar filament 23-40 µm. Type Location: Fukuoka, Japan Henneguya cerebralis Pronin 1972 Notes: Shorter and wider spore body than H. of overlapping length, Host: cranial cartilage, except H. renalis which is even wider; H. zikaweiensis, tchangi, & connective tissue of Thymallus tegidiensis, which differ in tail length; & H. gambusi, adherens & vovki, arcticus which differ in polar capsule length. Thinner body than H. neusslini, Cyst: 0.5-3, usually 0.8-1.0 mm; but thicker than H. nyongensis. Dimensions overlap with H. garavelli, spherical psorospermica, friderici, & cartilaginis; these species infect fish of different families, except H. cartilaginis, which is slightly longer in total Type Location: Mongolia length and deeper bodied. See also Batueva et al. (2013). GenBank#: JX1311380

Henneguya testicularis Azevedo, Notes: Longer polar capsules than H. of overlapping length, except H. Corral, & Matos 1997 percae, dini, miyairii, minuta, & creplini, which differ in spore body Host: testes of Moenkhausia width, and H. suprabranchiae, which has a longer tail. oligolepis Cyst: yes, no size given Type Location: Amazon estuary Brazil Polar capsule coils: 12-13 Henneguya corruscans Eiras, Notes: Longer and narrower spore body than other H. of overlapping Takemoto & Pavanelli 2009 total length, except H. schizodon, malabarica, & odzai, which are Host: gills of Pseudoplatystoma narrower in body width; H. yunnanensis, macropodi, garavelli, & corruscans latercapsulata, which differ in polar capsule dimensions; and H. Cyst: 100-300 μm guanduensis & maculosus, which has a longer tail. Dimensions overlap with H. ntondei, which infects a different fish family. Type Location: Parana River, Brazil Cyst is whitish-yellow, round-oval, elongate or irregular. Polar capsule coils: 5-6

Henneguya ntondei Fomena Notes: Differs in spore width and body length, and has a longer polar Folefack & Boiux 2008 capsule than other H. of overlapping total length, except H. lobosa & Host: gills of Schilbe mystus manipurensis, which have wider polar capsules. More polar filament Cyst: ovoid or subspherical, 170- coils than H. corruscans, maculosus, daoudi, guanduensis, nyongensis, 335 x 70-285 μm; up to 150/host friderici & sinensis. Dimensions overlap with H. macropodi, travassosi, dini, leporini, psorospermica, vovki, australis, pungitii, suprabranchiae Type Location: Sangé River, & electrica, but these infect other fish families. Cameroon

Polar capsule coils: 10 Henneguya garavelli Martins & Notes: Differs in polar capsule dimensions from other H. with Onaka 2006 overlapping total length, except H. zikaweiensis, ctenopomae, zahoori, Host: gills of Cyphocharax nagelli renalis, percae, miyairii, psorospermica, nuesslini, pseudoplatystoma, Cyst: dark, circular-ellipsoid, shaharini, gracilis, & creplini, which differ in body width; H. 60.9(42.8-63.2) x 34.7(24.5- manipurensis, mormyri, & pellucida, which differ in tail length; and H. 42.8)μm multiplasmodialis, wenyoni, leporinicola, pisciforme, laterocapsulata & minuta, which differ in spore body length. More polar filament coils Type Location: Rio do Peixe than for H. guanduensis. Dimensions overlap with H. bleekeri, nandi, Reservoir, Sao Paulo, Brazil yunnanensis, hainanensis, fonsecai, travassosi, dini, cerebralis, odzai & Polar capsule coils: 8-9; p.f.:24-33 µ friderici, which have not been reported to infect fish of Curimatidae. Henneguya malabarica Azevedo & Notes: Narrower spore body and shorter polar capsules than other H. Matos 1996 of overlapping total length, except H. zahoori & pseudoplatystoma, Host: gills of Hoplias malabaricus which are narrower-bodied; H. yunnanensis, macropodi, fonsecai, Cyst: polysporic, irregular pellucida, friderici, & suprabranchiae, which differ in tail length; H. Type Location: Amazon River, travassosi, gambusi, leporinicola, pisciforme, mormyri, laterocapsulata, & odzai, which differ in body length. Fewer PC coils than H. garavelli. near Belem, Brazil Dimensions overlap with H. hainanensis & dini, but these infect fish Polar capsule coils: 6-7 from different families. Henneguya leporini Nemeczek Notes: Narrower and longer spore body than other H. of overlapping 1926 total length, except H. zahoori, mystusia & schizodon, odzai, shaharini, Host: urinary ducts of Leporinus & gracilis, which are narrower; H. pisciforme, which has a longer spore mormyrops body; H. malabarica, garavelli, wenyoni, & adherens, which have Cyst: 30-40 x 60-80 μm shorter polar capsules; and H. lobosa, australis, nyongensis, which have Type Location: Sao Goncalo das longer tails. Dimensions overlap with H. macropodi, multiplasmodialis, Tabocas R., Minas Geraos, Brazil corruscans, ntondei, laterocapsulata, pungitii, guanduensis, sinensis; the latter species also infects internally, but all the overlapping spp.

infect different families.

Henneguya bicaudi Kaur and Attri Notes: Longer and narrower spore body than other H. of overlapping 2015 total length, except H. zahoori, H. macropodi & H hainanensis, which Host:gills of Cirrhinus mrigala differ in body width, and H. fonsecai, H. doori, & H. wenyoni, which Cyst: 0.5-0.7mm differ in polar capsule dimensions. Measurements overlap with H. Type location: Harike Wetland, India iheringi, which has a different shape and infects a different host on a different continent. Spore body/tail ratio=2.37, body length/width Polar capsule coils: 7-8; ratio=3.38 GenBank: KP099967 Henneguya wenyoni Pinto 1928 Notes: Longer spore body than other H. of overlapping total length, Host: gills of Tetragonopterus sp. except H. minuta, pisciforme, doori, & creplini, which have longer Cyst: 2 x 1 mm polar capsules. Type Location: Sao Paulo, Brazil Henneguya daoudi Szekely, Notes: Greater spore body width than other H. of overlapping length Shaharom-Harrison, Cech, Mohamed, and longer polar capsules than H. of similar width, except H. minuta, & Molnar 2009 miyairii, nuesslini & creplini, which have wider spore bodies; H. Host: gills of Trichogaster manipurensis, lobosa, & salvelini, which differ in tail length; H. trichopterus gambusi, cerebralis, testicularis, pisciforme, vovki, doori & Cyst: yes, no dimensions given nanhaiensis, which differ in spore body length. Dimensions overlap Type Location: Kelantan State, with H. psorospermica, ntondei, guanduensis, & tchangi, which infect Malasia fish of other families. Genbank: EU643625 Polar capsule coils: 6-7

Polar filament length: 29.4(27-31) µm Henneguya leporinicola Martins, Notes: Shorter spore body than other H. of overlapping total length, Souza, Moraes, & Moraes 1999 except H. renicola, bergamini, & santae, which have shorter tails; and H. massii & cerebralis, which have wider spore bodies. Host: gills of Leporinus Dimensions overlap with H. sichuanensis & mormyri, which have not macrocephalus been reported to infect fish of Anostomidae. Cyst: 23.4 to 70.3 μm No line drawing of spore in original paper; this drawing by the author to scale based on dimensions and photomicrograph. Type Location: Capivari, SP, Brazil

Henneguya pisciforme Cordeiro, Notes: Longer spore body than other H. of overlapping total length, Artigas, Gioia, & Lima 1983 except H. esocis, amiae, & umbri, which have longer tails; H. Host: gills of Hyphessobrycon wenyoni, minuta, curimata, creplini, doori, occulta, & rhinogobii, anisitsi which differ in polar capsule length; and H. renalis, which has a wider Cyst: 100.9(92-112) μm spore body. Type Location: Campinas, Sao Paulo, Brazil Henneguya nüsslini Schuberg & Notes: Wider spore body than other H. of overlapping total length, Schroeder 1905 (dimensions:Kallert et except H. electrica, creplini, salvelini, cartilaginis, salmonis, & al. 2005) shackletoni, which have longer tails; H. doori, occulta, psorospermica Host: subdermis of Salmo trutta & & similis, which have longer polar capsules; and H. minuta, which has Salvelinus fontinalis a longer spore body. Dimensions overlap with H. miyairii, cerebralis, Cyst: 1.5-2 mm & tegidiensis; The latter has unequal tails. H. miyairii has not been Type Location: Germany reported to infect any fish of Salmonidae. H. cerebralis has, but has a tropism for cranial tissue, as well as connective tissue. These two Polar capsule coils: 6-7 species may be the same, but mean spore width and length tend to Polar filament length: 48-60 μm differ. Actinospore stage is a triactinomxyon; does not infect GenBank #: AY669810 common carp; actinospore matures at 102 d (Kallert et al. 2005).

Henneguya oviperda Cohn Notes: Longer spore body than other H. of overlapping total length, 1896 except H. zikaweiensis, zahoori, renalis, wenyoni, creplini, amiae & Host: ovary of Esox lucius tegidiensis, which differ in spore body width. Dimensions overlap with H. Cyst: no real cyst, parasite pisciforme and H. doori, but these infect a different tissue and fish of occupies the ovum different orders. Dimensions (µm) in Tiutiaev 2008: spore length 17-22, total length up to Type Location: Germany 36, spore width 9-10, polar capsule up to 11 long x 2.5 wide. Dimensions Polar filament L: 47.9 μm in figure are from Weltner 1892. Auerbach (1911) notes cysts (1-6 mm) in See also Weltner 1892, Kennedy follicle epithelium of the ovary. 1974 Henneguya lobosa Cohn 1896 Notes: Differs in spore body width from other H. of overlapping total Host: gills of Esox lucius length and has a longer tail, except H. gambusi, cerebralis, pisciforme, Cyst: oval, 2.2-2.8 x 1-1.1 mm mormyri, vovki, curimata, umbri, amiae, rhinogobii , similis, which differ in Type Location: Germany spore body length; and H. cartilaginis, friderici & pseudorhinogobii which Polar filament length: 48-54 μm differ in polar capsule size. Dimensions overlap with H. guanduensis, See also Wegener 1910; nyongensis, suprabranchiae, electrica, doori, curvata, striolata, sinensis, Auerbach 1911; Kudo 1920 limatula, gigas, diversis, acuta, but these infect fish of different families. Mean values can also help differentiate, as many overlap only at the extremes of the size range.

Henneguya psorospermica Notes: Figure and dimensions shown are from Thélohan (1890, 1892, Thélohan 1892 1895), but body length estimated from proportion in figure relative to known polar capsule length. Tail length ≈ spore body length. Longer PC Host: gills of Perca fluviatilis length and differences in spore body width differentiate this species from Cyst: 1-3 mm (Andrews 1979) other H. of overlapping total length, except H. vovki which has a longer Type Location: Eurasia tail. Dimensions overlap with H. miyairii, electrica, bopleti, ntondei & See also Bütschli 1882, fig.16; umbri, which infect other fish families. H. similis also infects Perca, but is Doflein 1901; Dykova & Lom longer bodied. First species of H. described (Thélohan 1892). See 1978; Andrews 1979 Taxonomic Considerations section for additional comments.

Henneguya mormyri Notes: Shorter body length and narrower width than other H. of Kostoingue, Diebakate, Faye, & overlapping total length, except H. zahoori which has longer polar Toguebaye 2001 capsules. Dimensions overlap with H. leporinicola which infects fish of Host: gills of Mormyrus cashive Anostomidae in Brazil, not Mormyridae. Cyst: spherical, 0.4-0.8 mm Holotype: N˚ Myxo-097, Univ. C.A. DIOP, Dakar Type Location: Chari River, Chad Polar capsule coils: 3-4 Henneguya mystusia Sarkar Notes: Narrower width than other H. of overlapping total length, except 1985 H.leporinicola, psuedoplatystoma, & pellucida, which have a shorter spore body length, H. shaharini, gracilis, & eirasi, which differ in tail length, H. Host: gills of Mystus sp. odzai, which has a shorter polar capsule length, and H. laterocapsulata, Cyst: up to 0.1 x 0.2 mm, ovoid which has polar capsules that are not side-by-side. Dimensions overlap Type Location: West Bengal, with H. dini, australis, & acuta, but these infect fish of other families. India

Henneguya adherens Azevedo & Matos Notes: Differs in spore body width and has shorter polar 1995 capsules than other H. of overlapping total length, except H. Host: gills of Acestrorhynchus falcatus testicularis & corruscans, which have shorter tails; H. lobosa, Cyst: yes, no dimensions vovki & cartilaginis, which have longer tails; and H. wenyoni, Type Location: Amazon River estuary at leporinicola, pisciforme, doori, amiae & rhinogobii, which differ Belem, Brazil in spore body length. Dimensions overlap with H. gambusi, cerebralis & guanduensis, but these infect fish of different Polar capsule coils: 3-4 families. H. adherens is the sole Henneguya species reported to

date infecting the fish family Acestrorhynchidae. Henneguya laterocapsulata Landsberg Notes: The tandem polar capsule alignment in this species 1986 separates this from all the species that have polar capsules Host: skin of Clarias lazera arranged side-by-side. The only other species that share this Cyst: pale yellow, 500 x 750 μm trait is H. fusiformis, which is nearly twice as long as H. Type Location: Israel latercapsulata. The dimensions of H. alexandrinus (Ashmawy et Polar capsule coils: 5-6 al. 1989) overlap with H. laterocapsulata and are described from the same host, so are herein considered the same species. H. Polar filament length: 25.6(16.5-39.0) μm laterocapsulata, described first, takes priority.

Henneguya odzai Fomena & Bouix 1996 Notes: Narrower spore body than other H. of similar total Host: gills of Marcusenius moorii length, except H. zahoori & gracilis, which are narrower; spore Cyst: 53-125 x 30-90 um; whitish, body is longer than body of other species of similar width. spherical-ellipsoidal; up to 8,000 found Fewer PC coils than H. maculosus. Shorter polar capsule length in gills of one host than for other overlapping species (H. shizodon, mystusia, pungitii, nyongensis, acuta, amiae & esocis) and shorter tail than Type Location: Anga'a River, Cameroon H. eirasi. Length to width ratio: 3.7 Polar capsule coils: 4

Henneguya minuta Cohn 1896 Notes: Longer spore body and longer polar capsule length than Host: gills of Perca fluviatilis other H. of overlapping total length except H. similis, which has a Cyst: oval, 130 x 115 um; up to 6 longer tail. Dimensions overlap with H. creplini, which also cysts/lamellae and 200 per arch infects Percidae, but differences in means for spore width and Type Location: Germany polar capsule length should help separate this species. Polar filament length: 42-45 µm Henneguya vovki Achmerov 1960 Notes: Differentiated from most other H. of overlapping total length by its longer polar capsule and shorter spore body length; Host: body cavity of snakehead mullet differs in body width from H. renalis, percae, dini, (Channa spp.) multiplasmodialis, miyairii, shaharini, gracilis, suprabranchiae, Cyst: not described creplini, salvelini, occulta, rhinogobii, pseudorhinogobii, & acuta. Dimensions overlap with H. nyongensis, ntondei & Type Location: USSR, Amur River electrica, but these infect other fish families. Henneguya maculosus Carriero, Adriano, Notes: Differences in body width, body length, or polar capsule Silva, Ceccarelli & Maia 2013 length separate this from most overlapping species except H. Host: gills of Psuedoplatystoma macropodi, corruscans, assiuti & gigas, which differ in tail corruscans, P. reticulatum length; H. schizodon, nyongenis, cuniculator, ntondei, odzai & Cyst: 0.5-1.5 mm, white, elongated, eirai, which differ in PC coil number; and H. laterocapsulata, polysporous which has polar capsules arranged differently. Dimensions overlap with H. hainanensis, multiplasmodialis, garavelli, Type location: Pantanal National Park, malabarica, leporini, mystusia, australis, pellucida, guanduensis, Brazil shaharini & acuta, of which only H. multiplasmodialis also Polar capsule coils: 6-7 infects Pimilodidae; this species can only be differentiated by GenBank No.: KF296344, KF296345 rDNA analysis (GenBank No. KF296354) and slightly thicker body on average. 46

Henneguya australis Johnston & Notes: Narrower spore body and longer polar capsule than most Bancroft 1919 other H. of overlapping length, except H. suprabranchiae, sinensis & Host: gills of Plectroplites ambiguus rhinogobii, which have longer polar capsules; H. pisciforme, amiae, Cyst: tiny, round, white & esocis, which differ in spore body length; and H. Type Location: Thomson River, multiplasmodialis, eirasi, shaharini & pseudorhinogobii, which differ Central Queensland, Australia in tail length. Dimensions overlap with H. dini, leporini, mystusia, see also Shariff 1982 guanduensis, nyongensis, ntondei, friderici, gracilis, & acuta, which infect different fish families other than Percichthyidae. Henneguya pungitii Achmerov 1953 Notes: Longer polar capsule and narrower body width than other H. of overlapping total length, except H. dini, which has a shorter Host: subcutaneous connective spore body; and H. testicularis, lobosa, vovki, electrica & tissue of Pungitius pungitius pseudorhinogobii, which differ in tail length. Dimensions overlap Cyst: round, white, 1-2 mm with H. leporini, psorospermica, ntondei, doori, sinensis, umbri, Type Location: Kamchatka, USSR rhinogobii, suprabranchiae, acuta, & esocis, but these species have not been reported in fish of Gasterosteidae. Henneguya pseudoplatystoma Notes: Narrower body width and shorter polar capsules Naldoni et al. 2009 differentiate this from most other species of overlapping total Host: gills of Psuedoplatystoma sp. length, except H. renicola, bleekeri, pinnae, travassosi & dini, which Cyst: white, to 0.5 mm have shorter tails, and H. leporinicola & odzai, which differ in spore Type Location: São Paulo & Mato body length. Grosso, Brazil Polar capsule coils: 6-7 Henneguya pellucida Adriano, Arana Notes: Narrower spore body width and shorter polar capsules than & Cordeiro 2005 most other H. of overlapping total length, except H. dini, Host: visceral cavity & tunica malabarica, & garavelli, which have shorter tails; and H. externa of swim bladder of Piaractus leporinicola, laterocapsulata, odzai & mormyri, which differ in spore mesopotamicus body length. Cyst: yellow, round, 0.5-3.0 mm Type Location: Brazil Polar capsule coils: 6-7 Henneguya guanduensis Abdallah, Notes: Differs in spore width and tail length from other H. of Azevedo, Luque, Teresa, & Bomfim overlapping total length, except H. gambusi, latesi, cerebralis, 2007 wenyoni, leporinicola, pisciforme, mormyri, & similis, which differ in Host: gills of Hoplosternum littorale spore body length; H. multiplasmodialis, corruscans, lobosa, Cyst: 0.18 x 0.19 mm (0.06-0.30 x minuta, psorospermica, vovki, pungitii, curimata, doori, salvelini, 0.08-0.38), polysporic sinensis, & rhinogobii, which differ in polar capsule length; and H. Type Location: Guandu River, Brazil ntondei, garavelli, & acuta, which have more PC coils. Dimensions overlap with H. zikaweiensis, tchangi, dini, leporini, daoudi, Polar capsule coils: 3-6 adherens, australis, nyongensis, suprabranchiae, umbri, amiae,

nanhaiensis & tegidiensis, but these species have not been reported to infect fish of Callichthyidae. Henneguya nyongensis Fomena & Notes: Longer polar capsule and differences in spore body width Bouix 1996 separate this species from others of similar total length, except H. Host: gills & muscle of Marcusenius multiplasmodialis, pisciforme, curimata, doori, umbri, amiae, moorii rhinogobii, similis & esocis, which differ in spore body length; H. Cyst: 45-270 x 32-170 μm; whitish testicularis, corruscans, leporini & electrica, which differ in tail ovoid length. Dimensions overlap with H. dini, cerebralis, ntondei, lobosa, vovki, australis, pungitii, guanduensis, friderici, suprabranchiae & Type Location: Nyong & Ntem basins, acuta, but these species have not been reported to infect fish of Cameroon Mormyridae. Polar capsule coils: 4-5

Henneguya friderici Casal, Matos & Notes: Shorter spore body length and differences in spore body Azevedo 2003 width and polar capsule length differentiate this species from Host: gills, gut, kidney, liver of others of overlapping total length, except H. malabarica, salmonis, Leporinus friderici & gigas, which differ in tail length. Dimensions overlap with H. Cyst: whitish, round, 0.5-1.0 mm dini, cerebralis, ntondei, garavelli, australis, guanduensis, Type Location: Amazon River near nyongensis, electrica, sarotherodoni, & cartilaginis, but these Belem, Brazil species have not been reported to infect fish of Anostomidae.

Polar capsule coils:7-8 Henneguya nagelii Azevedo et al. 2013 Notes: Differences in body length and width, as well as tail length, Host: gills of Cyphocharax nagelii differentiate this species from others of overlapping total length, Cyst: white, round or oval, 150-250 except H. ntondei, which has more coils in the polar capsule μm filament, and H. lobosa, adherens, & pungitii, which differ in polar Type Location: Peixe’s River, São capsule length. Dimensions overlap with H. hainanensis, Paolo, Brazil macropodi, fonsecai, dini, garavelli, australis, guanduensis & nyongensis, of which only H. garavelli shares the same host family. Polar capsule coils: 6-8 H. garavelli has 8-9 coils in the polar filament and the cysts are darker and smaller in size.

Henneguya gracilis Johnston & Notes: The very narrow spore body differentiates this species from Bancroft 1919 other H. of overlapping total length, except H. zahoori & mystusia, which have shorter tails; and H. pseudoplatystoma, which has Host: gills of Therapon hillii shorter polar capsules. Dimensions overlap with H. australis and Cyst: narrow, pear-shaped, at right shaharini, which have not been reported to infect fish in angle to gill filament Tetrapontidae. Type Location: Australia

Henneguya shaharini Shariff 1982 Notes: Narrower spore body than most other H. of overlapping Host: gills of Oxyeleotris marmoratus length and longer polar capsule than H of similar width, except H. Cyst: 0.27- x 0.14 mm, ovoid australis & adiposa, which differ in tail length; H. esocis & diversis, Type Location: Selangor, Malasia which differ in spore body length, and H. auchenoglanii & eirasi, GenBank #: EU643630 which have wider polar capsules. Dimensions overlap with H. gracilis, which infects a different fish family. Polar capsule coils: 3-7 Total length by addition. Polar filament length: 16.8(11.2-19.6)µm Henneguya eirasi Naldoni, Arana, Notes: Narrower spore with longer tail than other H. of Maia, Silva, Carreirom Ceccarelli, overlapping total length, except H. leporinicola, laterocapsulata, Tavares, & Adriano 2011 pseudoplatystoma & pellucida, which differ in spore body length; Host: gills of Pseudoplatystoma spp. and H. diversis, mystusia, shaharini, & gracilis, which differ in polar Cyst: white, to 3mm capsule dimensions. Type Location: Pantanal, Mato Grosso, Brazil Polar capsule coils: 12-13 Henneguya curimata Azevedo & Matos Notes: Longer spore body with longer polar capsule and differs in 2002 spore width from most H. of overlapping length, except H. doori, Host: Kidney of Curimata inormata rhinogobii, similis & sinensis, which have a longer PC; & H. pungitii, Cyst: 0.6-1.2 mm diameter which has a wider polar capsule. Dimensions overlap with H. Type Location: Amazon estuary, Brazil umbri, but this species has not been reported to infect fish of Polar capsule coils: 10-11 Curimatidae or kidney tissue.

Henneguya suprabranchiae Notes: Differs in spore body length and width and has a longer tail Landsberg 1986 and polar capsule than most H. of overlapping total length. Host: suprabranchial respiratory Dimensions overlap with H.lobosa, pungitii, electrica, limatula, organ (SRO) & intestine of Clarias ntondei, sinensis, gigas, pseudorhinogobii, diversis & nyongensis; all gariepinus (syn.: C. lazera) except H. sinensis have not been reported in Clariidae. H. sinensis, Cyst: pale yellow to white, to 1 mm from China, tends to have a longer spore body on average (15.1 in SRO; 2-4.5 mm, grayish, ovoid μm), a longer tail (27.6 μm) & 7-8 PC coils. El-Mansy (2002) described immature stages and gives data on H. suprabranchiae (intestine) found in the intestine [ranges of spore body length (12.1-12.7 μm), Type Location: Hula Nature Reserve, spore width (4.5-6.0 μm), & tail (22.9-24.4 μm) are similar, but polar Israel; Nile River, Egypt capsule length is shorter (5.3-6.0 μm). Polar capsule coils: 9-10 Polar filament length: 82.5(55-97)µm Henneguya electrica Jakowska & Notes: Longer tail and differs in spore body width and length from Nigrelli 1953 other H. of overlapping total length, except H. garavelli, malabarica, Host: large electric organs of sarotherodoni, salvelini, occulta, nanhaiensis, cartilaginis, Electrophorus electricus tegidiensis & shackletoni, which differ in polar capsule Cyst: solid white to yellowish-soft measurements. Dimensions overlap with H. miyairii, lobosa, Type Location: Brazil psorospermica, vovki, friderici, ntondei, suprabranchiae, orientalis, & gigas, which have not been reported to infect fish of Polar filament length:44-50 µm Gymnotidae. Henneguya creplini Gurley 1893 Notes: Named for J.C.H. Creplin, who first described it (1842). Metric dimensions by Weltner (1892) were 18.8 μm for body length Host: gills of Gymnocephalus cernua and 6.3 μm at greatest width. This differs slightly from Gurley’s Cyst: to 1 mm (1893) dimensions: length 17.3 μm, breadth 5.8 μm. Greater spore Type Location: Germany body length than most H. of overlapping total length, except H. See also: Hegemann 1958; Lom 1961; percae, neusslini, minuta, occulta, malapteruri, ameiurensis, Haaparanta et al. 1994 diversis, esocis, & acuta, which differ in body width. Dimensions overlap with H. curimata, doori, umbri, amiae, rhinogobii, & similis. H. doori and similis also infect Percidae. Further genetic work is needed to see if these 3 species are synonymous. Henneguya doori Guilford 1963 Notes: Longer and wider spore body with longer polar capsules Host: gills of Perca flavescens than most other H. of overlapping total length, except H. percae, Cyst: 150-500 μm lobosa, creplini, occulta, bopleti, umbri, malapteruri, rhinogobii, & Type Location: Lake Michigan, USA similis. H. percae, creplini, & similis also infect Percidae, but H. Polar filament length: 32-56 μm percae has a more polygonal body shape, & H. creplini on average has a narrower body. H. doori may be a junior synonym to H. Polar capsule coils: 9-14 similis, but were described from different continents. Further GenBank #: HDU37549 (Griffin et al. genetic work is needed. The other overlapping species have not 2009) been found in Percidae. See also Cone 1994. Henneguya nkamensis Fomena, Notes: Shorter spore body and/or shorter polar capsule or body Folefack & Bouix 2008 width differences separate this from other H. of overlapping total Host: gills of Hepsetus odoe length. Cyst: white-yellow, 120-250 x 48-90 µm Type location: Sangé River, Cameroon Henneguya orientalis Nie & Li 1992 Notes: Differs in spore body length and width from other H. of Host: gills of Channa argus overlapping total length, except H. suprabranchiae, electrica, Cyst: 0.12-0.15 mm, round varichorhinusi & cartilaginis, which differ in polar capsule length. Type Location: China Polar capsule coils: 5-6

Henneguya melini Mathews, Maia & Notes: Has shorter polar capsules and differs in spore body Adriano 2016 length and width from other Henneguya of overlapping total length, except H.aequidens, which has a longer tail, and H. Host: gills of Corydoras melini acuta, which has more coils in the polar filament. Cyst: up to 260 µm Type Location: Rio Negro, Amazonas State, Brazil Polar capsule coils: 5-6 Henneguya saratherodoni Fall, Fomena, Notes: Shorter and wider spore body than other H. of Kostoingué, Diebakate, Faye, & Toguebaye overlapping total length, except H. cerebralis & leporinicola 2000 which have shorter spore bodies, and H. lobosa, electrica & Host: intestine of Saratherodon galilaeus gigas, which have longer polar capsules. Dimensions overlap Cyst: free, no cyst with H. miyairii, friderici, tegidiensis, shackletoni & Type Location: Chari River, Chad cartilaginis, but these infect fish of other families. Polar capsule coils: ND Heneguya aequidens Videira, Velasco, Notes: Has shorter polar capsules and differs in spore body Azevedo, Silva, Gonçalves, Matos & Matos length and width from other Henneguya of overlapping total 2015 length, except H. testicularis, which has a shorter tail. Host: gills of Aequidens plagiozonatus Cyst: asynchronous, whitish, ellipsoid

Type Location: Peixe-boi River, Pará State, Brazil Polar capsule coils:4-6

Henneguya curvata Barassa, Adriano, Arana Notes: Narrower and longer spore body than H. of & Cordeiro 2003 overlapping total length, except H. pisciforme & adiposa, which differ in tail length; H. ameiurensis, which has a longer Host: gills of Serrasalmus spilopleura body; H. postexilis, which has a narrower body; H. lobosa, Cyst: white, round to ellipsoid, 0.1 to 0.5 amiae, acuta & esocis, which differ in polar capsule mm dimensions. There is overlap in dimensions with H. sinensis, Type Location: Sao Paulo, Campinas, Brazil limatula, rhinogobii, pseudorhinogobii, & diversis, but these infect other fish families. Polar capsul coils: 10-11

Henneguya varicorhinusi Ma 1998 ( in Chen Notes: Differs in spore body length or width from most other & Ma 1998) H. of overlapping total length, except H. doori, orientalis, bopleti, nanhaiensis, salmonicola, cartilaginis, & shackletoni, Host: gills of Onychostoma simus which differ in polar capsule length. Dimensions overlap with Cyst: not observed H. salvelini, but this infects fish of a different taxonomic order and has sutural ridges lacking in H. varichorhinusi. Type Location: Leshan, China Polar capsule coils: ND

Henneguya striolata Casal, Matos & Azevedo Notes: Differs in spore body width or length from most H. of 1997 overlapping total length, except H. curimata, which has a shorter tail, and H. lobosa, curvata, sinensis, bopleti & Host: gills of Serrasalmus striolatus limatula, which differ in polar capsule metrics. More PC coils Cyst: 60-180 μm than H. umbri, rhinogobii, pseudorhinogobii, diversis, acuta, Type Location: Amazon River near Belem, & adiposa. Dimensions overlap with H. amiae, but this infects Brazil fish of a different family and has a different body shape. PC Coils: 13-14 Henneguya paraensis Velasco et al. 2016 Notes: Wider and shorter spore body than other H. of Host: gills of Cichla temensis overlapping total length except H. electrica which has a Cyst: 800 µm long, white, intrafilamental shorter tail, H. sarotherodoni which has a shorter spore body, Type Location: Tocantins River near Cametá, and H. salmonicola, H. cartilaginis, & H. shackletoni, which Pará State, Brazil have shorter polar capsules. Dimensions overlap with H. salvelini, which infects fish of a different order and tends to PC Coils: 5-7 have higher maxima in body width and length. GenBank: KU535882 Henneguya intracornea Gioia, Cordeiro & Notes: Longer and wider body or longer polar capsules than Artigas 1986 other H. of overlapping total length, except H. minuta which Host: eye of Astynanax scabripinnis is wider. H. umbri has fewer coils in the polar filament. Cyst: 356.65 µm (117-707); 1-16 cysts in Dimensions overlap with H. creplini, doori, occulta, bopleti, both eyes; rhinogobii, similis, & clariae, which infect different fish Type Location: Atibaia & Jaguari rivers, families and have polar capsules that are not set as far back Brazil (posteriorly) from the anterior end of the spore.

PC Coils: 8-9 Polar filament length: 101.9 µm (75.2-134.6)

Henneguya occulta Nemeczek 1926 Notes: Wider and longer body with longer polar capsules Host: gills of Loricaria spp. than other H. of overlapping total length, except H. minuta, Cyst: 75 μm, round which has a shorter tail and longer polar capsule. Type Location: Rio de Janeiro, Brazil Dimensions overlap with H. doori and similis, which infect a Polar filament: 17 μm different fish family. H. occulta is the only Henneguya species parasitizing Loricariidae.

Henneguya salvelini Zandt 1923 Notes: Differs in body length and has a wider body than most other H. of overlapping length; longer polar capsule than Host: skin of Salvelinus spp. spores of similar width. Longer tail than for H. renalis, Cyst: 1-1.5 x 2 mm percae, & guanduensis. Dimensions overlap with H. electrica, Type Location: Germany H. varicorhinusi, H. paraensis & H. malapteruri, but these infect different fish families. Total Length by addition.

Henneguya sinensis Chen & Hsieh 1960 Notes: Longer tail than H. leporini. Differences in body width and length, as well as a longer polar capsule, differentiate this from Host: intestine, ovary of Chana argus, most H. species, but dimensions overlap with many: H. lobosa, C. maculatus, Clarias batrachus, pungitii, nyongensis, curvata, blopleti, limatula, ntondei, Misgurnus anguillicaudatus rhinogobii, pseudorhinogobii, adiposa, sutherlandi & Cyst: 40-280 μm; round to almost round suprabranchiae. H. suprabranchiae also infects intestines of Clarias, but tends to have a shorter body on average and a Type Location: China shorter tail. H. sutherlandi, adiposa, & limatula infect Ictaluridae, Polar capsule coils: 7-8 though sutherlandi has a longer maximum total length (to 90 μm), adiposa has a smaller average width. H. limatula attacks gall bladder and no cyst has been described for it. Henneguya assiuti Mandour et al. 1988 Notes: Long polar capsule relative to spore length (70-80%) and Host: gills and suprabranchial differences in spore body length and width separate this from respiratory organ of Clarias lazera other H. of overlapping total length except H. adiposa, which Cyst: no data tends to have much longer maximum total length and body lengths, and H. sutherlandi, which has fewer coils in the polar Type location: Nile River, Egypt filament. Shorter body than H. sinensis, which infects the same Polar capsule coils: 8-9 host. Data from Rabie et al. 2009; See also Abed 1997. Henneguya bopleti Fomena & Bouix Notes: Differences in body width and length, as well as a longer 1987 polar capsule, differentiate this from most H., except H. Host: gills (rarely in dental palette) of ctenopharyngodoni, which has a longer tail, and H. sutherlandi & Chrysichthys nigrodigitatus umbri, which have fewer PC coils. Dimensions overlap with H. Cyst: 192-450 x 116-303 μm sinensis, limatula, rhinogobii, pseudorhinogobii, & similis, but Type Location: Cameroon these infect fish of other families. Polar filament length: 34.5(20.5-43.2) μm Polar capsule coils: 7-9 Henneguya azevedoi Barassa, Adriano, Notes: Shorter and narrower body with shorter polar capsules Cordeiro, Arana, & Ceccarelli 2012 than other H. of overlapping total length. Host: gills of Leporinus obtusidens Cyst: white, 40-200 μm Type Location: Mogi-Guacu River, Brazil Polar capsule coils: 6-7 Henneguya umbri Guilford 1965 Notes: Differences in body length and width differentiate this Host: gills of Umbra limi from most H., except H. oviperda, which has a shorter tail, H. Cyst: to 190 μm curimata, sinensis, similis, alexeevi, tangschensis, & tunghuensis, Type Location: USA, Great Lakes which differ in polar capsule metrics, and H. astyanax, bopeleti, drainage doori, striolata, pilosa, rhinogobii & pseudorhinogobii, which have more PC coils. Dimensions overlap with H. amiae, limatula, Polar capsule coils: 6 creplini, guanduensis, pungitii, sutherlandi, & clariae, which have Polar filament length: 12-32 µm not been reported to infect fish of Umbridae. Henneguya nanhaiensis Chen & Ma 1998 Notes: Differs in spore body length and width and has shorter polar capsules than most other H., except H. garavelli, Host: skin, kidney of Channa argus salmonicola, cartilaginis & shackletoni, which have even shorter Cyst: nearly round, 100-120 μm polar capsules, and H. ghaffari, which has wider polar capsules. Type Location: China Dimensions overlap with H. guanduensis, which infects different tissues of fish of a different family on a different continent.

Henneguya amiae Fantham, Porter & Notes: Long body length and unique spore shape, with Richardson 1940 constriction posterior to the capsules, separates this from most other H. except H. esocis, which tends to have longer polar Host: gills of Amia calva capsules, infects a different fish family, and reaches greater Cyst: whitish-yellow, 1 mm diameter maximum total lengths. Based on dimensions only, there is Type Location: Quebec Province, overlap with H. guanduensis, curimata, creplini, striolata, umbri, Canada diversis, acuta, adiposa, sutherlandi, which infect fish of different families. Total length by addition. Polar capsule coils:ND Polar filament length: 66-90 µm Henneguya pethericii Fomena, Folefack & Notes: Differences in spore body length or width, or polar Bouix 2008 capsule length, separate this from other H. of overlapping total Host: gills of Ctenopoma petherici length, except H. rhamdia, which has a wider polar capsule. Cyst: subspherical, polysporous, 80-125 Dimensions overlap with H. astyanax, salmonis, gigas, & x 80-110 μm sutherlandi, which have not been reported to infect fish of Type Location: Littoral Province, Anabantidae. Museum specimen: Univ. Yaounde I, Cameroon. Cameroon Polar capsule coils: ND Henneguya limatula Meglitch 1937 Notes: Differences in spore body length or width, or polar Host: gall bladder of Ictalurus capsule length, separate this from other H. of overlapping total punctatus, I. furcatus length, except H.lobosa, curvata, suprabranchiae, striolata, Cyst: not observed sinensis, bopeleti, rhinogobii, umbri, gigas, pseudorhinogobii, Type Location: Ohio River at arapaima, & sutherlandi. Of these, only H. sutherlandi has Shawneetown, Illinois, USA been found in ictalurids; H. sutherlandi forms cysts in skin and tends to reach much longer total lengths. Total length shown is Polar capsule coils: ND based on addition of tail and body lengths. Henneguya astyanax Vita, Corral, Matos Notes: Differs in spore body length or width, or polar capsule & Azevedo 2003 length from other H. of overlapping total length except H. umbri, Host: gills of Astyanax keithi pethericii, rhamdia, acuta, ghaffari, episclera, esocis, & Cyst: whitish, 250 um sutherlandi. Of these, none have been reported to infect fish of Type Location: Amazon River estuary, Characidae. Brazil Polar capsule coils: 8-9 Henneguya malapteruri Fomena & Bouix Notes: Wider and longer spore body than other H. of 1997 overlapping total length, with differing polar capsule length, Host: muscle of Malapterurus electricus except H. minuta, acerinae & nigris, which have longer spore Cyst: 0.5-0.9 mm, polysporous, bodies; Fewer PC coils than H. doori. Dimensions overlap with spherical to subspherical, white H. guanduensis (slightly), salvelini, ctenopharyngodoni, episclera, Type Location: Lipahé River, Cameroon & clariae, but these have not been reported to infect fish of Malapteridae. Polar filament:40-48 µm. Polar capsule coils: 4-5 Henneguya salmonicola Ward 1919 Notes: Wider and shorter spore body, with shorter polar Host: muscle of Oncorhynchus spp. capsules than other H. of overlapping total length, except H. Cyst: 3-6 mm; to 15 mm (Boyce 1985) mbakaouensis, which has a shorter body. Dimensions overlap Type Location: western USA with H. cartilaginis, shackletoni & mandouri, of which only H. Polar capsule coils: ND cartilaginis also infects fish of Salmonidae. H. cartilaginis can GenBank# AF031411 only be differentiated by host tissue (it infects head cartilage See also Fish 1939; Clouthier et al. 1997 versus muscle in H. salmonicola) or 18S rDNA testing.

Henneguya cartilaginis Yokoyama, Notes: Wider and shorter spore body, with shorter polar Urawa, Grabner, & Shirakashi 2012 capsules than H. of overlapping total length, except H. leporinicola, mormyri & doneci, which have shorter bodies. Host: head cartilage of Oncorhynchus Dimensions overlap with H. latesi, miyairii, cerebralis, neusslini, masou guanduensis, friderici, sarotherodoni, mbakaouensis, Cyst: white, 1-3 mm salmonicola, salmonis, tegidiensis, shackletoni, zschokkei, mandouri, sutherlandi & dogielli; those in bold also infect Type Location: Shubuto River, Japan salmonids. H. cerebralis also infects head cartilage, but is slightly PC Coils: 3 smaller and differs slightly in spore shape. 18S rDNA tests GenBank #JQ411297 separate it from H. zschokkei, neusslini, & salmonicola. H. Total length by addition tegidiensis infects skin, rather than cartilage. Henneguya salmonis Fantham et al. 1939 Notes: Short body, differences in body width and polar capsule Host: muscle of Salmo salar metrics differentiate this from other H. of overlapping total Cyst: present, no size given length, except H. pethericii, zschokkei, schizura, & dogielli. Most Type Location: Gaspe Peninsula, Quebec of these infect other fish families, but H. zschokkei also infects Province, Canada muscles of salmonids, so can only be differentiated by fish host species. Total length by addition. Polar filament length: up to 25 μm Henneguya zschokkei Gurley 1893 Notes: Original descriptions (Zschokke 1884; Gurley 1893) Host: muscle, subdermal tissue of lacked measurements, other than tail (6-8 times body length). Coregonus fera, Prosopium coulteri, P. Doflein (1901) gave the metrics & figure shown here. Spore williamsoni metrics in U.S. Prosopium are in brackets; note unequal tail Cyst: yellow, up to 5-6 mm x 2-4 mm length in some spores [Mitchell 1989]. Differs in spore length or width from other H., except H. cartilaginis, salmonis, Type Location: Lake Geneva, tegidiensis, mbakaouensis, & dogielli; those in bold also infect Switzerland GenBank#: AF378344 salmonids. H. cartilaginis & cerebralis infects head cartilage, See also Batueva et al. 2013( rDNA test), rather than muscle. H. salmonis & tegidiensis may be junior Bykhovskaya-Pavlovskaya et al. 1964; Vik synonyms, but spores in Prosopium tend to have wider PCs. 1960; Davies 1968; Newell & Canaris 1969 Henneguya tegidiensis Nicholas & Jones Notes: Wider and shorter spore body, with different polar 1959 capsule metrics than H. of overlapping total length. Dimensions Host: muscle of Coregonus clupeoides overlap with H. tchangi, miyairii, cerebralis, garavelli, nuesslini, Cyst: cyst 2 mm up to 2 cm zschokkei, cartilaginis, shackletoni, guanduensis, sarotherodoni, Type Location: Lake Bala, North Wales ghaffari, & mbakaouensis; those in bold also infect salmonids. H. Polar capsule coils: 5 cartilaginis & cerebralis infect head cartilage, but H. nuesslini & zschokkei also infect muscle/skin. Longer tail on average than H. neusslini. Unequal tail lengths should separate this from H. zschokkei. Total length by addition.

Henneguya shackletoni Brickle, Kalavati Notes: Wider and shorter spore body, with different polar & MacKenzie 2006 capsule metrics or tail length than H. of overlapping total length. Host: visceral organs of Eleginops Dimensions overlap with H. latesi, miyairii, cerebralis, maclovinus sarotherodoni, cartilaginis, tegidiensis, & mbakaouensis, but Cyst: 500-800 μm, mature spores only these infect fish of other families and a different host tissue. Type Location: Falkland Islands estuary Cysts <200 µm were disporous. Polar capsule coils: 6-7

Henneguya mississippiensis Rosser, Notes: Differs in spore body length or width from other H. of Griffin, Quiniou, Khoo, Greenway, Wise, overlapping total length except H. melini, H. aequidens, H. curvata, Pote 2015 H. striolata, H. sinensis, H. astyanax, H. tangschensis, & H. exilis, Host: gills of Ictalurus punctatus which differ in polar capsule dimensions, H. pisciforme which has a Cyst: 0.3-0.5 mm shorter tail, and H. umbri, H. arapaima, & H. sutherlandi, which Type Location: catfish pond, have fewer PC coils. Dimensions overlap with H. amiae & H. esocis, Washington County, MS, USA which have a different shaped body, and H. rhinogobii and H. pseudorhinogobii, which infect a different fish family and order. Polar capsule coils: 8-9 Among overlapping species infecting Siluriformes, H. limatula, H. GenBank: KP404438 diversis, H. adiposa, & H. gurleyi infect different tissues, and H. Holotype: Smithsonian, Washington, samochimensis & H. acuta have larger cysts. D.C., USNM1270623 Alternate host is aquatic oligochate Dero digitata, which produces an aurantiactinomyxon actinospore (see Bellerud 1993). Henneguya gigas Chen & Hsieh 1960 Notes: More PC coils than H. sutherlandi, guanduensis, but fewer Host: gill, ovary, intestine of Channa than in H. suprabranchiae. Differences in spore body length and argus width, tail length, and polar capsule metrics separate this from Cyst: round or elongate most other H. of overlapping total length, except H. lobosa, friderici, Type Location: China nagelii, electrica, & schizura. These species infect fish of different families. Polar capsule coils: 7-8

Henneguya rhamdia Matos, Tajdari & Notes: Differences in spore length, width, or polar capsule Azevedo 2005 dimensions separate this species from other H. of overlapping total Host: gills of Rhamdia quelen length, except H. sutherlandi & schizura. These infect fish of Cyst: polysporous, spheric to different families. H. sutherlandi also has fewer coils (6) in the polar ellipsoid, to 300 μm diameter filament. Type Location: Peixe Boi River, State of Pará, Brazil Polar capsule coils: 10-11 (rarely 12)

Henneguya samochimensis Reed, Notes: Differences in spore body length or width, or polar capsule Basson & Van As 2003 length, separate this from other H. of overlapping total length Host: gills of Clarias gariepinus except H. ghaffari & H. amazonica, which have wider polar Cyst: polysporous, yellow-white, 2-5 capsules. Dimensions overlap with H. pethericii, H. astyanax, & H. mm sutherlandi; the latter has fewer polar filament coils (6). H. Type Location: Okavango River & pethericii & H. astyanax typically have smaller cysts and have not delta, Botswana been reported to infect siluriform fishes.

Polar capsule coils: 8 Henneguya arapaima Feijo, Arana, Notes: Differences in spore length, width, tail length, or polar Ceccarelli, & Adriano 2008 capsule metrics separate this species from most other H. of overlapping total length. Fewer coils in the polar filament than H. Host: gills, gall bladder of Arapaima pseudorhinogobii, diversis, and sutherlandi. Dimensions overlap gigas with H. limatula, which also infects gall bladder, but this species is Cyst: white,round-ellipsoid, 200-600 not known to infect fish of Arapaimidae. μm Type Location: Araguaia River, Brazil Polar capsule coils: 5

Henneguya bayerii Pronin & Pronin 2002 Notes: Differences in spore body length or width, or the shorter Host: kidney of Comephorus baicalensis polar capsules, separate this species from other H. of & C. dybowskii overlapping total length except H. mbakaouensis & H. Cyst: white, 0.1-1.0 mm cartilaginis, which are thinner bodied. Dimensions overlap with Type Location: Lake Baikal, Russia H. shackletoni, which is also coelozoic, but it reaches wider (to 11.2) and longer (14.4) maximum spore body dimensions and Polar capsule coils: ND infects different fish hosts.

Henneguya ctenopharyngodoni Ma Notes: Differences in spore length, width, or polar capsule 1993 (Chen and Ma 1998) metrics separate this species from most other H. of overlapping total length except H. malapteruri & clariae, which have not Host: gills of Ctenopharyngodon idellus been reported to infect cyprinids; also H. malapteruri infects Cyst: 0.5-1mm; oval muscle, not gill tissue. Type Location: Taihu Lake, China

Henneguya similis Zandt 1924 Notes: Differences in body width, longer spore body, and polar capsule metrics separate this from other H. of overlapping total Host: gills of Perca fluviatilis length except H. intracornea, which is longer bodied and H. Cyst: 0.5 x 1.1-1.7 mm, round, kidney- tunghuensis, which has a longer tail. Dimensions overlap with H. shaped creplini, doori, occulta, bopeleti, alexeevi, sutherlandi, & clariae; Type Location: Germany those in bold also infect Percidae. H. doori tends to reach a No figure in original paper wider maximum body width and shorter maximum length. Maximum body length and width for H. creplini are not as great. Henneguya rhinogobii Li & Nie 1973 Notes: Differences in body length and width, polar capsule Host: intestine & gill of Rhinogobius metrics, and tail length separate this from most other H. of giurinus overlapping total length; differs in PC coil number from H. Cyst: oval to oval elongate, 422-1200 x striolata, sinensis, umbri, pseudorhinogobii, diversis, adiposa, & 292-520 μm; intrafilamental sutherlandi; thicker body than for H. exilis & adiposa, and thinner bodied than H. clariae. Dimensions overlap with H. Type Location: China pungitii, creplini, doori, curvata, bopleti, limatula, similis, Polar capsule coils: 9-10; alexeevi, acuta & tunghuensis; of these, none have been GenBank #AB447992 reported to infect fish of Gobiidae. See also Kageyama et al. (2009) for rDNA differentiation and Chen & Ma (1998). Henneguya pseudorhinogobii Kageyama Notes: Differences in spore length, width, tail length, or polar et al. 2009 capsule metrics separate this species from most other H. of Host: gills of Rhinogobius sp. overlapping total length; differs in PC coil number from H. doori, Cyst: intralamellar, 50-250 μm curvata, striolata, umbri, arapaima, sutherlandi & rhinogobii. Type Location: Nagara River, Japan Dimensions overlap with H. sinensis, bopeleti, limatula, diversis, Polar capsule coils: 8 acuta & adiposa, which have not been reported to infect fish of Gobiidae. H. rhinogobii has larger cysts, infects a different Polar filament length: 29.6(22.9-36.7) µm location of the gill (afferent artery vs. intralamellar), and differs GenBank #: AB447994, AB447995, in rDNA tests. AB447996

Henneguya torpedo Azevedo, Casal, Matos, Notes: Longer spore body or differences in body width or Alves, & Matos 2011 polar capsule metrics differentiate this species from other H. of overlapping total length. Host: brain, spinal cord of Brachyhypopomus pinnicaudatus Cyst: none described Type Location: Peixe Boi River, Brazil Polar capsule coils: 5-6, rarely 7

Henneguya piaractus Martins & Sousa 1997 Notes: Narrower or shorter body than other H. of overlapping total length, except H. diversis & H. adiposa, both of which Host: gills of Piaractus mesopotamicus have greater average and maximum body length. Cyst: dark, ellipsoid, 0.1-0.9 mm

Type Location: Jaboticabal, Sao Paulo, Brazil Polar capsule coils: 8-9 Polar filament length: 28.1 (25.4-31.8)

Henneguya alexeevi Shul’man 1962 Notes: D ifferences in spore body width and a longer body with longer polar capsules, separate this from most other H. Host: gills and ovary of Percottus glehni of overlapping total length, except H. clariae & similis, which Cyst: round to oval have a thicker spore body and infect other fish families. Type Location: Amur River Basin, USSR

Henneguya schizura Gurley 1893 Notes: Differences in spore body width and length separate this from many H. of overlapping total length, but dimensions Host: Circumorbital tissue of the eye of overlap with H. salmonis, tegidiensis, gigas, mandouri, Esox lucius amazonica, dogielli, & sutherlandi. However, these species Cyst: 0.44-1.09 mm infect fish of different families. Type Location: Germany

Henneguya nigris Bond 1939 Notes: Longer body, with differing body width, and shorter polar capsules than other H. of overlapping total length, Host: gills (dermis of inner arch) of Esox except H. intracornea & wolinensis, which are longer bodied, masquinongy, E. niger and H. clariae, which has wider polar capsules. Thinner body than H. episclera. Cyst: to 4-5 x 1-2 mm in E. niger,; typically 2 x 1.5 mm in E. masquinongy Type Location: Maryland & New York, USA Polar capsule coils: 5-6

Henneguya tridentigeri Ozaki & Isizaki 1941 Notes: Differences in body length, the wider spore body, Host: body surface (fins, tongue, operculum, with longer and narrower polar capsules than similarly mouth margin) of Tridentiger obscurus shaped spores, and longer tail, separate this from other H. of Cyst: white, round or ovoid, 1.28 x1 .0 mm overlapping total length. Iodinophilous vacuole was 2.9 x 2.5 Type Location: River Otagawa, Hirosima, µm. See also Hoshina (1952). Japan Polar capsule coils: 6 Polar filament length: 30.4 µm Henneguya pilosa Azevedo & Matos 2003 Notes: Differences in spore body width and length differentiate this species from all but H. umbri & H. clariae, Host: gills of Serrasalmus altuvei which have not been reported to infect fish of Serrasalmidae. Cyst: up to 0.2 mm, white, round to H. umbri also has fewer coils (6) in the polar filament and H. ellipsoidal clariae is thicker bodied. Type Location: Brazil Polar capsule coils: 11-12, rarely 13

Henneguya auchenoglanii Kostoingue, Notes: Narrower or shorter spore body than other H. of Diebakate, Faye & Toguebaye 2001 overlapping total length except H. adiposa & piaractus, which have a narrower polar capsule. Host: gills of Auchenoglanis occidentalis Cyst: ovoid, 1-1.5 mm

Type Location: Chari River, Chad Polar capsule coils: ND

Henneguya doneci Schul’man 1962 Notes: Nearly spherical body, which is shorter than other H. species, except H. zschokkei, which has a slightly wider body, Host: gills of Carassius carassius and H. tegidiensis, which has shorter polar capsules. Cyst: up to 3 mm Described in Bykhovskaya-Pavlovskaya et al. (1964) and Type Location: Amur River Basin, Asia Shul’man (1966). See also Ye et al. (2012) for additional Polar capsule coils: ND molecular and histological data. GenBank #: EU344899 (Griffin et al. 2009)

Henneguya mandouri Rabie, Mohammed, Notes: Shorter polar capsules, differences in body width and Hussein, & Hussein 2009 length, or tail length, separate this from most other H. species; fewer PC coils than H. amazonica & sutherland. Host: gills of Lates niloticus Dimensions overlap with H. schizura, cartilaginis, tegidiensis, Cyst: whitish, oval, 0.2-1 x 0.1-0.5 mm & mbakaouensis. All but H. mbakaouensis infect skin or Type Location: Kena City, Egypt muscle rather than gills and infects different host families. H. Polar capsule coils: 3-5 mbakaouensis infects the same host and tissue, and may be the same species, differentiated only by slightly greater Polar filament length: 12.5-23 μm maximum width and slightly wider polar capsules.

Henneguya tangschensis Wu 1997 Notes: Longer polar capsule, or differences in body length or width separate this species from other H. of overlapping total Host: lips, intestine of Channa argus length, except H. clariae which has a thicker body and tends to Cyst: 0.3-0.6 up to 5 mm reach greater maximum total lengths and body lengths. Type Location: China

Henneguya waltairensis Notes: Longer polar capsules, narrow body, or differences in body Narashimhamurti and Klavati 1975 length separate this species from other H. of overlapping total length. Host: gills of Channa punctatus Cyst: to 120 µm; translucent

Type Location: near Viskhaptanm, India Polar filament length: 15-20 µm Polar capsule coils: 6-7

Henneguya fusiformis Kostoingue et Notes: Longer bodied spore than any other H. species, except H. al.1999 intracornea, which has longer polar capsules that are parallel, rather than one after the other as in H. fusiformis. Host: gills of Clarias anguillaris L. Cyst: ovoid, whitish, 0.5-2.5 mm Type Location: Chari River, Chad Polar capsule coils: 5-6

Henneguya ameiurensis Nigrelli and Notes: Longer bodied than most other H. of overlapping total Smith 1940 length, and more narrow bodied with shorter polar filaments than other long-bodied spores. Dimensions overlap with H. esocis Host: skin of barbels of Ameiurus & amiae, which also feature the characteristic indentation of the nebulosus shell posterior to polar capsule. Differentiation from these two Cyst: 190-342 x 760-1220 μm; species is based on the host family, Ictaluridae. spherical, oval or irregular; Total length by addition. polysporous Type Location: New Hampshire, USA Polar capsule coils: ND Henneguya acuta Bond 1939 Notes: Differences in spore body length, thickness, or width, or polar capsule length separate this species from other H. of Host: gills of Esox masquinongy overlapping total length, except H. australis, pungitii, Cyst: 2-3 x 1-2 mm guanduensis, nyongensis, striolata, amiae, astyanax, rhinogobii, Type Location: Chautauqua Lake, New diversis, pseudorhinogobii, adiposa, esocis, & sutherlandi. H. York, USA guanduensis, nyongensis, striolata, & sutherlandi differ in the number of filament coils. The remainder infect other fish Polar capsule coils: 8-9 families, except H. esocis, which differs notably in spore shape.

Henneguya ghaffari Ali 1999 Notes: Differences in body length or width, polar capsule metrics, separate this species from other H. of overlapping Host: intestinal tract, gills, gill rakers of total length except H. astyanax, which has a shorter tail. Lates niloticus Dimensions overlap with H. zschokkei & H. rhomboides, but Cyst: 1.0 (0.5-1.3) mm in gills, 0.3-1.4 these have not been reported to infect fish of Latidae. H. mm in intestines rhomboides also has a unique, more angular, shape. Type Location: Lake Wadi El-Raiyan, Up to 1400 cysts in one intestine. Egypt Polar capsule coils: 4-5 Henneguya amazonica Rocha, Matos & Notes: Shorter polar capsules or differences in body length or Azevedo 1992 width differentiate this species from other H. of overlapping total length except H. mandouri & shizura. Neither of these Host: gills of Crenicichla lepidota species have been reported to infect fish of Cichlidae. H. Cyst: oval, up to 0.05 x 0.15 mm, mandouri also has fewer coils (3-5) in the polar filament. asynchronous spore development Type Location: Amazon River, Brazil Polar capsule coils: 6 Henneguya mbakaouensis Fomena & Notes: Shorter or wider bodied, with shorter polar capsules Bouix 2000 than most H. of overlapping total length except H. cartilaginis, tegidiensis, shackletoni, & mandouri; most of these species Host: gills of Lates niloticus have not been reported to infect fish of Latidae, except H. Cyst: whitish, ovoid, or subspherical,120- mandouri, which infects the same host and tissue. H. mandouri 470 x 60-230 µm, polysporous may be a junior synonym, differentiated only by a slightly Type Location: Sanaga Basin, Cameroon greater maximum width and slightly wider polar capules. Polar capsule coils: 4-5 Henneguya mailaoensis Kostoingue, Notes: Differences in polar capsule metrics, or body length and Diebakate, Faye, & Toguebaye 2001 width, separate this species from other H. of overlapping total length except H. sutherlandi & clariae, which infect fish of Host: gills of Mormyrus cashive different families. Cyst: ovoid, 0.5-1.5 mm Type Location: Chari River, Chad Polar capsule coils: ND Henneguya mogurndae Fujita 1936 Notes: Longer bodied, with longer polar capsules, than most other H. of overlapping total length except H. wolinensis & Host: gills of Mogurnda obscura clariae, which have a slightly wider body. Cyst: 310 x 270 µm; membrane 5 µm Total length by addition. Type Location: Ogura Pond, Fushimi, Japan Polar capsule coils: ND

Henneguya rhomboideus Ma, Dong & Notes: Unique angular shape of the anterior end of the spore, Wang 1982 as well as differences in body length, width, or polar capsule dimensions separate this species from other H. of overlapping Host: gills of Carassius auratus auratus total length. Dimensions overlap with H. nanhaiensis, Cyst: elongate, 92-62 µm tegidiensis, ghaffari & episclera, but these species infect fish of Type Location: China other families.

Henneguya diversis Minchew 1977 Notes: Narrower body, longer polar capsule, or differences in body or tail length, separate this from most other H. of overlapping total Host: kidney, liver, connective tissue length; fewer PC coils than H. curvata, striolata, rhinogobii, & of muscle and fins of Ictalurus gurleyi. Dimensions overlap with H. sinensis, piaractus, acuta, punctatus esocis, pseudorhinogobii, postexilis, adiposa & sutherlandi; those in bold also infect ictalurids. H. adiposa & postexilis have Cyst: white, polysporous, 250x600 μm monosporous pansporoblasts, whereas H. diversis has a disporous Type Location: southeastern USA pansporoblast (Minchew 1977), and tends to have a slightly thicker Polar capsule coils: 6-8 body. H. sutherlandi tends to reach larger body width & length, & longer maximum total length. Henneguya postexilis Minchew 1977 Notes: Narrower spore body, with differing body length, or shorter Host: gills of Ictalurus punctatus polar capsule than other H. of overlapping total length, except H. Cyst: polysporous, circular to oblong, piaractus, diversis, gurlei, adiposa, & esocis. H. diversis & adiposa 12 x 12-80 x 75 μm, cyst wall about also infect I. punctatus, but H. diversis has a disporous sporoblast 1 μm, monosporous sporoblast and reaches a slightly wider body width and deeper body. H. adiposa has a thicker cyst cell wall (3-4) and larger cysts, protruding Type Location: Saline County, from host tissue, as well as a longer distance from the posterior end Missouri, USA of the polar capsule to the spore cavity-tail junction (8-10 vs. 6-7 Polar capsule coils: 6-8 μm). Henneguya gurleyi Kudo 1920 Notes: Differences in body width or length, or polar capsule Host: fins of Ameiurus nebulosus, A. metrics, separate this species from most H. of overlapping total melas length, except H. acuta, which has a deeper body, and H. amiae, Cyst: white, spherical or which has a shorter tail. Dimensions overlap with H. limatula, subspherical, in clusters, to 1.8 mm astyanax, pseudorhinogobii, diversis, postexilis, adiposa, esocis, sutherlandi, & clariae; those in bold also infect ictalurids. H. gurleyi Type Location: South Creek, Aurora, has more coils in the polar filament than H. diversis, postexilis, N. Carolina, & Storm Lake, Iowa, adiposa or sutherlandi. H. limatula infects different tissue (gall USA bladder) and no cyst has been described. H. clariae tends to reach Polar capsule coils: 9 much longer maximum total lengths. Named for parasitologist R.R. GenBank #: DQ673465 Gurley. Measurements from Iwanowicz et al. (2008). Henneguya exilis Kudo 1929 Notes: More narrow bodied than most H. of overlapping total Host: gills, integument of Ictalurus length, with longer polar capsules (except H. tanschensis, which has punctatus, I. melas Worm host: Dero longer polar capsules), or differences in body length separate this digitata (Lin et al. 1999) species from all overlapping species except H. adiposa. H. adiposa Cyst: 410 x 250 to 825 x 1800 µm also infects ictalurids, and may be a junior synonym, but tends to have a smaller average body length and a lower mimimum body Type Location: Rock R., Illinois, USA width. See also McCraren et al. (1975), Current and Janovy (1978), Genbank #:AF021881 Mitchell (1978), and Duhamel et al. (1986). Henneguya adiposa Minchew 1977 Notes: Narrower body or differences in body or polar capule length Host: adipose fin of Ictalurus separate this species from other overlapping H. except H. punctatus rhinogobii, striolata & gurleyi, which have more coils in the polar Cyst: polysporous, 120 x 290-500 x filament. Dimensions overlap with H. sinensis, gigas, piaractus, 150 µm pseudorhinogobii, acuta, esocis, diversis, postexilis, exilis & sutherlandi; those in bold also infect ictalurids. H. sutherlandi tends Type Location: Mississippi (Lee to have wider spores on average, polar capsules of equal length, & County), USA shorter polar capsules on average. H. diversis also tends to have shorter PCs, infects a wider range of tissues, and has a shorter total Polar capsule coils: 6-8 length minimum and maximum. H. exilis tends to have a larger GenBank #: EU492929 average body length and a higher mimimum body width. On average, H. postexilis tends to have a narrower, shorter body and shorter PCs.

Henneguya wolinensis Romuk-Wodoracki Notes: Longer spore body, with longer polar capsules than other 1990 H. of overlapping total length except H. clariae, which infects Host: skin of Perca fluviatilis fish of another order and reaches a much longer maximum total Cyst: 0.5-1 mm, milky, oval length. Measurements are for fresh spores. Type Location: Dziwna River, Szczecin Firth, Poland Polar filament length: 120.0-163.3 µm

Henneguya esocis Fantham, Porter & Notes: Unique spore shape (the tail about ½ the width of spore Richardson 1939 at the origin, with sporoplasm extending into the caudal fork; sometimes a constriction posterior to polar capsules) separates Host: gills of Esox niger this from all but H. amiae, which has similar dimensions, but Cyst: elongate tends to have shorter polar capsules, infects a different fish Type Location: Brome Lake, Quebec, family, & has a shorter maximum total length. Based on Canada dimensions only, there is overlap with H. curvata, astyanax, pungitii, acuta, diversis, postexilis, gurlei, adiposa, sutherlandi. Polar capsule coils: up to 10 Only H. acuta also infects esocids.

Henneguya dogielli Achmerov 1960 Notes: Differences in polar capsule metrics, or differences in body length or width separate this species from other H. of Host: gills of Siniperca chuatsi overlapping total length, except for H. cartilaginis, which is Cyst: white, 2-6 mm thicker bodied. Dimensions overlap with H. gigas, schizura, Type Location: Amur River, Russia mandouri, ghaffari & sutherlandi, but none of these species has see also: Bykhovskaya-Pavlovskaya et been reported to infect fish of Percithyidae. Total length by al. 1964, Schul’man 1966 addition.

Henneguya episclera Minchew & Sleight Notes: Differences in body length or width or polar capsule 1977 length separate this species from other H. of overlapping total length except H. astyanax & ctenopharyngodoni, which differ in Host: eye episclera of Lepomis gibbosus polar capsule width, & H. nigris, which is not as thick bodied. Cyst: none Dimensions overlap with H. malapteruri, rhomboides & clariae, Type Location: Rhode Island, USA which infect different tissue in different fish families. Polar capsule coils: 5-6

Henneguya sutherlandi Griffin, Pote, Notes: Large variation in dimensions, so lots of overlap with Wise, Greenway, Mauel, & Camus 2008 other species: H. amiae, pethericii, limatula, cartilaginis, astyanax, gigas, rhamdia, arapaima, rhinogobii, Host: skin of Ictalurus punctatus pseudorhinogobii, similis, mandouri, acuta, mailaoensis, diversis, Cyst: round to ovoid, to 2 mm gurlei, adiposa, laseeae, esocis, dogielli, caudalongula, Type Location: Mississippi, USA kwangtungensis, clariae, & pellis; those in bold also infect Polar capsule coils: 6 ictalurids. H. limatula infects a different tissue (gall bladder) and GenBank #: EF191200 (Griffin et al. 2009) does not get as long. H. diversis & adiposa tend to be more narrow-bodied on average. Both H. gurlei & H. pellis have a greater number of coils in the polar filament. Smaller cyst than H. lasseeae.

Henneguya laseeae Leis, Rosser, Baumgartner & Notes: Differs in spore body length and width from Griffin 2016 other H. of overlapping total length except H. Host: gills of Pylodictis olivaris rhomboides & H. tunghuensis, which differ in polar Cyst: 1.2-1.8 mm, asynchronous capsule length; H. esocis, which has more coils in the Type Location: Pool 9, Mississippi River near polar filament; and H. gurleyi, which is thinner bodied. Lansing IA, USA Dimensions overlap with H. sutherlandi, which also infect ictalurids, but creates smaller cysts. Polar capsule coils: 6-7 GenBank: KX354352 Holotype: USNM 1407985, Smithsonian Institute, Washington, D.C. Henneguya acerinae Schröder 1906 Notes: Longer than most H. species, with a wider spore body than other H. that overlap in total length, Host: gill of Acerina cernua; Aspro zingel except H. episclera, which has shorter polar capsules. Cyst: up to 300 µm Dimensions overlap with H. clariae, which differs by Type Location: Hungary having slightly unequal polar capsules and a different Polar filament length: 80-90 µm host family and order. Total length by addition.

Henneguya caudalongula Adriano, Arana, & Notes: Longer than most H. species, that differs in Cordeiro 2005 body length or the narrower body width from other H. of overlapping total length, except H. adiposa, which Host: Prochilotus lineatus has longer polar capsules and H. longicauda, which Cyst: round to ellipsoidal, 0.2-1 mm has a longer tail. Dimensions overlap with H. Type Location: Pirassununga, Sao Paulo, Brazil sutherlandi, but this species infects fish of a different family and has fewer coils in the polar filament. Polar capsule coils: 10-11

Henneguya kwangtungensis Chen 1998 (Chen and Notes: Differences in spore body length or width Ma 1998) separates this from most other H. of overlapping total length except H. limatula, psuedorhinogobii, Host: gills and gall bladder of Mylopharyngodon amazonica, rhomboideus, & dogielli, which differ in piceus, Carrasius auratus, Channa maculatus polar capsule metrics. Dimensions overlap with H. Cyst: not observed sutherlandi, which infects fish of a different family and Type Location: Zhejiang and Guangdong Province, order. China Henneguya tunghuensis Chen 1998 (Chen and Ma Notes: Differences in spore body length or width 1998) differentiates this from most other H. of overlapping total length except H. clariae & similis, which have Host: abdominal cavity of Channa argus thicker bodies, H. rhinogobii, which has a shorter tail, Cyst: not observed & H. psuedorhinogobii, mailaoensis, rhomboideus, Type Location: Wuchang district, Hubei Province, gurleyi & sutherlandi, which have shorter polar China capsules.

Henneguya magna Rice & Jahn 1943 Notes: Longer body and longer total length than most other H. of overlapping total length, except H. ictaluri, Host: gills of Morone chrysops which has a longer polar capsule, and H. episclera, Cyst: not observed which has a wider spore body. Tail is bifurcated for the Type Location: Spirit Lake, Iowa, USA entire length.

Henneguya ictaluri Pote, Hanson, & Shivaji 2000 Notes: Longer spore body, longer polar capsule, or Host: gills of Ictalurus punctatus shorter width than other Henneguya of overlapping Worm host: Dero digitata total length except H. wolinensis, which has an even Cyst: interlamellar, 638 x 661 µm; not visible longer polar capsule. Dimensions overlap with H. clariae, which has slightly unequal polar capsules and Type Location: Mississippi, USA visible cysts. Actinospore: Aurantiactinomyxon Total length by addition. Polar capsule coils: ND GenBank: AFO298320 Henneguya clariae Abolarin 1971 Notes: Longer total length than most spores, with a longer tail or differing in spore body length or width Host: gills of Clarias lazera from H. of overlapping total length except H. striolata, Cyst: 0.5-1 mm, sometimes to 4 mm or more pseudorhinogobii, magna & ctenopharygodoni, which Type Location: Nigeria have narrower or shorter polar capsules; and H. Polar filament length: 22(21.0-22.5) rhinogobii, torpedo, alexeevi, nigris, pilosa, Polar capsule coils: ND tangschensis, gurleyi, & tunghuensis, which are thinner bodied. Dimensions overlap with H. bopleti, umbri, malapteruri, similis, mailoensis, wolinensis, episclera, sutherlandi, acerinae, & ictaluri; of these only H. sutherlandi and H. ictaluri also infect siluriformes, but not fish of Clariidae. Henneguya gigantea Nemeczek 1911 Notes: very long spore, but with a shorter body than any spore that overlaps in total length. Host: gill and skin of Sander lucioperca Cyst: 4-7 x 2-3 mm Type Location: Hungary Polar filament length: 70 Henneguya pellis Minchew 1977 Notes: very long spore with a narrower and shorter body than other H. of overlapping total length except Host: skin of Ictalurus furcatus H. dogielli, which has shorter polar capsules, H. Cyst: white, ovoid, 690 x 2050 µm gigantea, which has a shorter body, & H. longicauda, Type Location: Lee County, Alabama, USA which has a narrower body. Dimensions overlap with Polar capsule coils: 8-10 H. sutherlandi, but it has fewer polar filament coils and GenBank#: FJ468488 (Griffin et al. 2009) does not get as long.

Henneguya longicauda Minchew 1977 Notes: very long spore with a narrower spore body than any Henneguya of overlapping total length. It Host: gills of Ictalurus punctatus has a monosporous sporoblast. Cyst: 130-370 x 110-120 µm, polysporous, whitish, circular to oblong Type Location: Neshoba County, Mississippi, USA Polar capsule coils: 9-12

Marine and estuarine Henneguya species with equal polar capsules

Henneguya multimorpha Li & Wang Notes: Narrower spore body than H. ocellata and H. ogawai, and 1993 shorter tail than H. sichuanensis. Dimensions overlap with H. Host: gills of Acanthogobius hasta periopthalmusi & H. ouakamensis; H. ouakamensis can only be Cyst: oval-elongate to round differentiated by host family (Mugilidae) and the slightly wider Type Location: China sutural ridge around the spore body, which extends into the caudal Polar capsule coils: ND projections of the shell. H. periopthalmusi also infects fish of Gobiidae and may be the same species, but it has a slightly longer polar capsule on average.

Henneguya periophthalmusi Wu, Jiang Notes: Narrower spore body width than H. ocellata, not as thick as & Wang 1993 H. ogawai, and wider polar capsules than H. sichuanensis. Host: gills of Periopthalmus Dimensions overlap with H. multimorpha, which also infects fish of cantonensis Gobiidae, but it has slightly shorter polar capsules on average. H. Cyst: oval-round, 0.2-0.3 mm ouakamensis also overlaps in dimensions, but infects fish of a Type Location: China different family and has a slightly wider sutural ridge around the spore body, which extends into the tail. Polar capsule coils: ND

Henneguya ocellata Iversen & Yokel Notes: Wider spore body than H. multimorpha, H. periohthalmusi & 1963 H. lesteri; longer polar capsule than H. sichuanensis. Dimensions Host: intestinal epithelium, pyloric overlap with H. ouakamensis and H. ogawai. H. ouakamensis ceca of Sciaenops ocellatus infects different tissue in a different family (Mugilidae). H. ogawai Cyst: polysporous, spherical to infects a similar tissue, but tends to have a more oval body shape ellipsoid, 1.2-2.0 mm, whitish-yellow and infects fish of a different family (Sparidae). See also Landsberg 1993. Type Location: Florida, USA Polar capsule coils: not visible Henneguya ouakamensis Kpatcha, Notes: Wider spore body than H. lesteri; longer spore body than H. Faye, Diebakate, Fall & Tobguebaye 1997 latesa, and differs in polar capsules metrics from H. sichuanensis. Host: gill and heart of Mugil cephalus Dimensions overlap with H. multimorpha, H. periophthalmusi, H. Cyst: variable, whitish ocellata, H. ogawai, & H. yokoyamai, so can only be differentiated Type Location: Senegal Coast by fish host or target tissue. Polar capsule coils: ND Henneguya ogawai Li et al. 2012 Notes: Wider spore body than H. multimorpha, thicker body than Host: alimentary tract wall and gall H. periophthalmusi or H. latesa, shorter tail than H. sichuanensis, bladder wall of Acanthopagrus shorter polar capsule than H. lesteri, and shorter body length than schlegelii for H. yokoyamai. Dimensions overlap with H. ouakamensis and H. Cyst: white, 0.78 x 0.58 to 0.66 x ocellata, which infect fish of different families. H. ocellata also infects a similar tissue, but tends to be more oval in shape. 0.66 mm; in gall bladder wall 0.22- Type specimens (NSMT-Pr332, NSMT-Rf331) at Nat. Sci. Mus., 0.71 x 0.17-0.57) Tokyo, Japan. Type Location: Inland Sea off Hofu City, Japan Polar capsule coils: not visible Type specimen: NSMT-Pr332,NSMT- Rr331; Nat. Sci. Museum, Tokyo, Japan

Henneguya sichuanensis Ma 1998 (Chen and Ma Notes: Longer tail than H. multimorpha & H. ogawai; 1998) differs in polar capsule dimensions from H. Host: Urinary bladder of Rhinogobius typus periophthalmusi, H. ocellata, H. ouakamensis, H. Cyst: none described lesteri, & H. yokoyamai; thicker body than H. latesa; Type Location: China. Estuarine. shorter spore body than H. mauritaniensis. Polar capsule coils: ND

Henneguya lesteri Hallett & Diamant 2001 Notes: Total length derived by addition. Spore body is Host: psuedobranch, gill, & oral mucosa of narrower than for H. ocellata, H. ouakamensis, H. Sillago analis latesa, H. yokoyamai, & H. lagodon; Polar capsule Cyst: gill 175-335 µm, psuedobranch 200-270 x length differs from H. ogawai and H. sichuanensis. 210-550 µm This is the only Henneguya species described to date from fish in Sillaginidae. Type Location: Australia, Queensland coast Polar capsule coils: 4 Polar filament length: 23.2(14.7-26.8)µm

Henneguya latesa Wu, Wu & Hua 1994 Notes: Shorter spore body than most H. of Host: gills of Lates calcarifer overlapping total length except H. ogawai & H. Cyst: none reported sichuanensis, which have thicker bodies, H. Type Location: S. China Sea, Guishan, Zhuhai yokoyamai, which has longer polar capsules, and H. Polar capsule coils: ND lesteri & H. lagodon, which differ in spore body width. Total length by addition. Henneguya yokoyamai Li, Sato, Kamata, Ohnishi, & Notes: Slight difference in size between the two polar Suguita-Konishi 2012 capsules; longer polar capsule than for H. sichuanensis Host: gall bladder and peritoneum of & H. latesa; narrower polar capsule than for H. Acanthopagrus schlegelii mauritaniensis; longer spore body than for H. ogawai Cyst: 0.22-0.71 x 0.17-0.57 mm & H. lagodon, but shorter than H. testicularis; shorter Type Location: Inland Sea off Hofu City, Japan tail than H. mbourensis. Dimensions overlap with H. ouakamensis, H. rubicundi & H. akule, which have not Polar capsule coils: not visible been reported to infect fish of Sparidae. H. akule & H. Type specimen: NSMT-Pr334, NSMT-Rr333 in Nat. rubicundi tend to reach a larger total length. Sci. Museum, Tokyo Synchronous spore development.

Henneguya lagodon Hall & Iversen 1967 Notes: Caudal end of spore body has an extended Host: skin of head, Lagodon rhomboides shell forming a stiff flap. Shorter spore body length Cyst: 0.2-1.4 mm (mean 0.70 mm), whitish than other H. of overlapping total length, except H. yellow; spherical to ellipsoidal lesteri, H. latesa, H. brachydeuteri, cynoscioni, H. Type Location: Everglades, Florida, USA cutanea, H. clini & H. yoffensis, which differ in spore body width. Dimensions overlap with H. priacanthi Polar capsule coils: ND and H. pagri; H. priancanthi infects a different tissue Type specimen:U.S. Nat. Museum, No. 23757 (gill) of fish from a different family; H. pagri also infects fish of Sparidae, but infects heart and gills, not skin.

Henneguya testicularis Azevedo, Corral, & Matos Notes: Longer spore body length than other 1997 Henneguya of overlapping total length. See also the Host: testes of Moenkhausia oligolepis freshwater section for this species. Cyst: yes, no size given Type Location: Brazil, Amazon estuary near Belem Polar capsule coils: 12-13

Henneguya malabarica Azevedo and Matos 1996 Notes: Narrower spore body width than other Host: gill of Hoplias malabaricus Henneguya of overlapping total length except H. Cyst: polysporic, irregular rubicundi which has slightly wider polar capsules. Type Location: Brazil, Amazon estuary near Basal portion of spore and tails covered with a Belem homogeneous sheath. Polar capsule coils: 6-7

Henneguya mbourensis Kpatcha, Faye, Diebakate, Notes: Wider spore body than H. latesa, H. malabarica Fall, & Toguebaye 1997 & H. adherens; longer tail than H. yokoyamai; differs in spore body length from H. lagadon, H. testicularis, H. Host: kidney of Dentex canariensis curimata, H. rubicundi & H. mauritaniensis. Cyst: none Dimensions overlap with H. akule, which has longer Type Location: Senegal maximum tail lengths and can also be differentiated by the different host family (Carangidae) and tissue Polar capsule coils: ND (heart).

Henneguya rubicundi Haldar & Mukherjee 1985 Notes: Differs in spore body length from other H. of overlapping total length, except H. cynoscioni & H. Host: fins of Gobioides rubicundus cutanea, which differ in spore body width; H. Cyst: oval, creamy-yellowish, 1.5 x 2.0 mm brachydeuteri & H. pagri, which have a longer tail; H. Type Location: West Bengal, India malabarica, H. akule, & H. vitiensis, which have Polar capsule coils: 6-8 narrower polar capsules; and H. adherens, which has Type specimen: GR/Henn/Myxo-77, Univ. Kalyani fewer coils in the polar filament. Dimensions overlap with H. yokoyamai & H. mauritaniensis, which can only be differentiated by the different host families and tissues.

Henneguya adherens Azevedo & Matos 1995 Notes: Differs in spore body length from other H. of Host: gills of Acestrorhynchus falcatus overlapping total length, except H. mbourensis, H. Cyst: asynchronous, polysporous mauritaniensis, H. cynoscioni, and H. vitiensis, which Type Location: Brazil, Amazon estuary near have wider spore bodies; and H. pagri, which has a Belem longer tail. Dimensions overlap with H. rubicundi & H. akule, which can only be differentiated by the Polar capsule coils: 3-4 different host families and tissues infected. Sheath Type specimen: Accession No. 47799, Internat. around whole spore, thinner around body (~0.1 µm) Protozoan Type Slide Collection, Smithsonian Inst., than tail (0.3-0.4 µm). Washington D.C., USA

Henneguya curimata Azevedo & Matos 2002 Notes: Longer spore body length than other H. of Host: kidney of Curimata inormata overlapping total length except H. striolata, which has Cyst: 0.6-1.2 mm diameter a longer tail, and H. rhinogobii & H. vitiensis, which Type Location: Brazil, Amazon estuary differ in body width. PC coils: 10-11

Henneguya brachydeuteri Kpatcha, Faye, Diebakate, Notes: Shorter spore body than other H. of Fall & Tobguebaye 1997 overlapping total length, except H. priacanthi, which Host: heart of Brachydeuterus auritus has a shorter body; H. lagadon, & H. malabarica, Cyst: variable, in bulbus arteriosis or in ventricle which differ in spore body width; H. rubicundi which Type Location: Senegal has a shorter tail, H. cynoscioni, H. cutanea, H. pagri, & PC coils: ND H. akule, which differ in polar capsule dimensions. Dimensions overlap with H. mauritaniensis, which also infects heart tissue, so can only be differentiated by the different fish host family.

Henneguya mauritaniensis Khlifa, Miller, Adlard, Notes: Differences in spore body length or the longer Faye, & Sasal 2012 tail differentiate this from other H. of overlapping Host: heart arterial bulb of Pagrus caeruleostictus total length, except H. malabarica, H. adherens, H. Cyst: 0.1 mm diameter, white subspherical to tunisiensis, which differ in body width; and H. spherical cynoscioni, H. cutanea, H. pagri, H. akule, & H. Type Location: Mauritania vitiensis, which differ in polar capsule dimensions. The measurements overlap with H. rubicundi, H. PC coils: ND brachydeuteri & H. yoffensis; H. rubicundi & H. Type specimen: ZS108 to ZS111, Mus. Nat. History, brachydeuteri can only be differentiated by the Paris different host family or tissue (fins for H. rubicundi). H. yoffensis also infects fish of Sparidae, but infects gills, not heart tissue. Total length by addition. Henneguya cynoscioni Dykova, deBuron, Roumillat, Notes: Wider spore body or shorter polar capsule & Fiala 2011 length than other H. of overlapping total length except Host: heart of Cynoscion nebulosus H. priacanthi, H. yoffensis, H. rhinogobii, H. vitiensis, & Cyst: polysporic, large (size not given) H. astyanax, which differ in spore body length. Type Location: coastal S. Carolina, USA Dimensions overlap with H. akule & H. miyazakii; H. PC coils: 3-4 akule also infects heart tissue; both can only be differentiated by the different fish hosts or, in the case GenBank#:JN017203 of H. miyazakii, by the different host tissue (skin). SSU rRNA data has shown only 86.7% similarity with H. akule. Henneguya cutanea Dogiel & Petruschewsky 1933 Notes: Wider spore body or longer polar capsules than Host: fins, skin of Abramis brama, Leuciscus idus, other H. of overlapping total length except H. Pelecus cultratus priacanthi & H. neopolitana, which have a shorter Cyst: white, spherical to oval, to 1.8 x 1.3 mm spore body. Dimensions overlap with H. akule & H. Type Location: Neva Gulf, USSR sebasta. H. sebasta reaches much longer total lengths and longer maximum body lengths and infects fish of a PC coils: ND different family. H. akule also infects a different fish GenBank#: AY676460 host family and tissue (heart). See also Schul’man 1966, Yokoyama et al. 2005

Henneguya priacanthi Kpatcha Faye, Notes: Shorter spore body length than other H. of overlapping Diebakate, Fall & Tobguebaye 1997 total length, except H. lagodon which overlaps in Host: gills of Priacanthus arenatus measurements. H. lagodon can only be differentiated by the Cyst: 2-3 per gill arch, no size given different host family and host tissue (skin). Type Location: Senegal coast Polar capsule coils: ND Henneguya pagri Yokoyama et al. 2005 Notes: Differs in body length or width from other H. of Host: heart, gills of Pagrus major overlapping total length, except H. rubicundi & H. adherens, Cyst: none which have shorter tails; H. brachydeuteri, H. mauritaniensis, & Type Location: Japan, Mie Prefecture H. clini, which differ in polar capsule metrics; H. otolithi, which is Polar capsule coils: 3 thinner bodied; & H. texana, which has a longer tail. Dimensions overlap with H. lagodon & H. akule; H. lagodon also infects fish Polar filament length: 20.5(16.3-31.3) of Sparidae, but infects a different tissue (skin). H. akule also GenBank: AB183748 infects heart tissue, but infects fish of a different family. Plasmodia mature in bulbous arteriosus, then migrate to the gills. Hairlike extension of caudal appendages. Total length shown is by addition of maxima or minima of body and tail lengths. GenBank data indicated that the closest relative is H. lateolabracis. Henneguya akule Work, Takata, Whipps, & Notes: Narrower polar capsule or differences in spore body Kent 2008 length separate this from other H. of overlapping total length Host: aortic bulb, Selar except H. tunisiensis, which is thicker bodied, & H. yokoyamai, crumenophthalmus which has a shorter tail. Dimensions overlap with several Cyst: pleomorphic, 10-700 µm species, but all infect fish of other families; H. mbourensis & H. adherens tend to have a shorter tail. H. cynoscioni also infects Type Location: S. Oahu, Hawaii, USA heart tissue, but tends to be shorter bodied on average. H. Polar capsule coils: 3-4 cutanea infects a different tissue and has markings on suture Type specimen: USDA, Nat. Parasite rim. H. jocu infects a different tissue (gill). H. pagri, lateolabracis Collection, Beltsville, Maryland, USA & clini are very similar, but have shorter maxima in body and GenBank: EU016076 total length. H. astyanax tends to be less rounded, with a longer body on average. H. vitiensis also infects heart tissue, so can only be differentiated by host family. H. sebasta tends to reach larger total and body lengths. Henneguya tunisiensis Bahri et al. 2010 Notes: 4-6 marginal markings on suture rim. Wider spore body Host: gills, gill arch of Symphodus tinca than other H. of overlapping total length except H. Cyst: white, 1.0-1.5 x 1.5-2.0 mm brachydeuteri, H. priacanthi & H. neopolitana, which differ in Type Location: near Kerkennah Islands, spore body length; H. cutanea, H. yoffensis, which differ in polar Tunisia capsule metrics; and H. akule & H. miyazakii, which are thinner bodied. Shorter tail than for H. sebasta, which also has longer Polar capsule coils: 4-5 maximum body and total lengths. Type specimen: ZS53, Natural History Mus., See also Bahri and Marques 2008. Paris GenBank: GQ340975 Henneguya striolata Casal, Matos & Notes: Longer spore body than other Henneguya of overlapping Azevedo 1997 total length except H. curimata & H. astyanax, which differ in Host: gills of Serrasalmus striolatus tail length; H. cutanea, H. yoffensis & H. vitiensis, which differ in Cyst: 60-180 µm spore body width, and H. rhinogobii, which has longer polar Type Location: Brazil, Amazon estuary capsules. near Belem Polar capsule coils: 13-14

Henneguya clini Reed, Basson, Van As & Notes: Differs in spore body length from other H. of overlapping Dyková 2007 total length except H. lagadon, H. cynoscioni, H. cutanea, H. Host: gills, gill arches of Clinus miyazakii, H. tunisiensis & H. texana, which differ in spore body superciliosus width; and H. priacanthi, H. pagri, & H. otolithi, which differ in Cyst: 1-3 mm, white, polysporous polar capsule metrics. Dimensions overlap with H. akule & H. Type Location: Africa, southern coast lateolabracis, which infect different families; H. akule tends to have longer maxima in polar capsule length, body length, & tail Polar capsule coils: 4-5 length. H. lateolabracis tends to have a slightly narrower body, narrower polar capsule with only 3 coils, and a longer tail.

Henneguya jocu Azevedo, Rocha, Notes: Differs in spore body width or length from other H. of Matos, Matos, Oliveira, Al-Quraishy, overlapping total length except H. pagri, H. miyazakii, H. clini, H. Casal 2014 lutjani, which differ in polar capsule dimensions, and H. Host: gills of Lutjanus jocu shackletoni, which has a thinner spore body. Dimensions overlap Cyst: up to 260 x 130 µm with H. akule, which has more variability in spore size, fewer coils, Type Location: Atlantic coast near and infects heart tissue, not gills. Algodoal, Pará State, Brazil Polar capsule coils: 4-5 Henneguya yoffensis Kpatcha, Faye, Notes: Wider spore body than other H. of overlapping total length Diebakate, Fall, & Toguebaye 1997 except H. brachydeuteri, cynoscioni, priacanthi, pagri, clini, Host: gills, heart of Sparus rhinogobii, otolithi, & H. neopolitana, which differ in spore body caeruleostictus length; and H. cutanea, akule, tunisiensis & H. tachysuri, which Cyst: whitish cysts, semispherical differ in polar capsule metrics. Dimensions overlap with H. Type Location: Senegal mauritaniensis, miyazakii, & H. sebasta; H. mauritaniensis also infects fish of Sparidae and heart tissue, but has not been reported Polar capsule coils: ND in the gills and tends to have a slightly narrower body on average. H. sebasta tends to have a longer tail, longer polar capsules, and wider spore body on average. H. miyazakii infects a different tissue (skin) & tends to reach longer total lengths. Henneguya astyanax Vita, Corral, Matos Notes: Longer spore body than other Henneguya of overlapping & Azevedo 2003 total length except H. cutanea, tunisiensis, & yoffensis, which are Host: gills of Astyanax keithi wider bodied; and H. rhinogobii, vitiensis, tachysuri, amazonica & Cyst: cyst; whitish 250 um H. sebasta, which differ in polar capsule dimensions. Type Location: Amazon Estuary, Brazil Polar capsule coils: 8-9 Type specimen: No. 1004430, Internat. Protozoan Collection, Smithsonian Inst., Washington DC Henneguya vitiensis Laird 1950 Notes: Total length derived by addition. Differs in body length or Host: heart muscle; blood of width from other H. of overlapping total length except H. Leiognathus fasciatus rubicundi, mauritaniensis, cutanea, astyanax, tachysuri & H. Cyst: none described sebasta, which differ in polar capsule dimensions; and H. lutjani & Type Location: Fiji H. tridentigeri, which have a shorter tail. Dimensions overlap with H. akule, which also infects heart tissue, so can only be Polar capsule coils: 3-4 differentiated by the differing fish host taxon.

Henneguya miyazakii Hoshina 1952 Notes: Total length shown is by addition of spore and tail length. Host: skin; corium of Chaenogobius Wider spore body than other H. of overlapping total length except annularis H. cutanea & H. akule, which differ in polar capsule metrics; H. Cyst: 1.0-3.8 x 1.0-2.2 mm tunisiensis, which is thicker bodied; H. striolata, rhinogobii, Type Location: Japan, Kanazawa Bay astyanax & neopolitana, which differ in body length. Dimensions overlap with H.cynoscioni, yoffensis & sebasta, which infect a Polar capsule coils: 4 -5 different tissue (heart, gill). H. sebasta tends to reach longer total Polar filament length: 18.6 µm lengths. H. yoffensis tends to have longer maximum and mean Spore length/width ratio: 1.24(1.07-1.42) body lengths, whereas H. cynoscioni tends to have shorter maximum and total lengths.

Henneguya rhinogobii Li & Nie 1973 Notes: Longer spore body or longer polar capsule than other Host: intestine & gill of Rhinogobius Henneguya of overlapping total length except H. curimata, cutanea, giurinus tunisiensis & vitiensis, which are wider bodied; and H. striolata, Cyst: oval to oval-elongate, 422-1200 astyanax & sebasta, which have shorter polar capsules. x 292-520 µm Type Location: China Polar capsule coils: 9-10 GenBank No. AB447992

Henneguya lateolabracis Yokoyama, Notes: Differs in body width and length from other Henneguya of Kawakami, Yasuda,& Tanaka 2003 overlapping total length except H. otolithi, which is thinner bodied; Host: bulbus arteriosis; gills of and H. lutjani, which has a wider polar capsule. Dimensions overlap Lateolabrax spp. with H. akule & H. sebasta, which also infect heart tissue, but in a Cyst: rounded, irregular different host family; both species also tend to have greater Type Location: Japan, Mie Prefecture maximum width and length of the body. Polar capsule coils: 3 polar filament length:22.4(15.9-26.7 µm) Henneguya otolithi Ganapati 1941 Notes: The dimensions overlap with H. akule, clini & lutjani. Host: heart muscle of Otolithus spp. However, the unique shell thickening which forms a band around Cyst: whitish, opaque the middle of the spore, perpendicular to the long axis of the spore, Type Location: India distinguishes this species from all other Henneguya. Polar capsule coils: 5-6 Total length by addition.

Henneguya texana Joy 1972 Notes: Total length by addition of spore and tail length. Shorter Host: gills of Pogonais cromis spore body than other Henneguya of overlapping total length Cyst: variable size except H. neopolitana, which is wider bodied; and H. kayarensis, Type Location: USA, Texas coast which has a wider polar capsule. Polar capsule coils: 3 Type specimen: Cat. No. USNM 24121, US Nat. Mus.

Henneguya shackletoni Brickle, Notes: Wider and shorter spore body, with different polar capsule Kalavati, & MacKenzie 2006 metrics or tail length than H. of overlapping total length. H. Host: visceral organs of Eleginops cynoscioni & miyazakii have fewer coils. Dimensions overlap with maclovinus H. latesi, miyairii, cerebralis, sarotherodoni, cartilaginis, Cyst: white, spherical-ellipsoid, 500- tegidiensis, & mbakaouensis, but these infect freshwater fish of 800 μm, mature spores only; <200 other families and a different host tissue. Also overlaps with H. um, disporous brachydeuteri, mauritaniensis, yoffensis, lutjani, & sebasta, which can only be differentiated by host species differences. Type Location: Falkland Islands estuary Polar capsule coils: 6-7

Henneguya lutjani Kpatcha, Faye, Notes: Differences in spore body length and width separate this Diebakate, Fall,& Toguebaye 1997 species from other Henneguya of overlapping total length except Host: gills of Lutjanus agennes H. akule, vitiensis, lateolabracis, tridentigeri & H. tachysuri, which Cyst: whitish, 1.8 x 0.8 mm differ in polar capsule metrics. Dimensions overlap with H. Type Location: Senegal otolithi, which can be differentiated by its unique band around the Polar capsule coils: ND spore.

Henneguya tridentigeri Ozaki & Isizaki Notes: Total length by addition of spore and tail lengths. 1941 Differences in body length separate this from other Henneguya of Host: skin of Tridentiger obscura overlapping total length except H. miyazakii & H. amazonica, Cyst: variable, up to 1.28 x 1 mm which differ in spore body width; and H. lutjani & H. tachysuri, Type Location: Japan, Kanazawa Bay which differ in polar capsule length. Polar capsule coils: 6 polar filament length: 30.4 µm

Henneguya tachysuri Menon 1979 Notes: Longer polar capsule than other Henneguya of overlapping Host: subcutaneous muscle of total length except H. cutanea & H. rhinogobii, which differ in Tachysurus thalassinus spore body width. Cyst: white to light brown, round- oval, 1-3 x 0.5-1 mm, 50-100/fish Type Location: Indian Ocean Polar capsule coils: 3-5

Henneguya neopolitana Parisi 1912 Notes: Shorter spore body length than other Henneguya of Host: kidney of Box salpa overlapping total length except H. texana & H. kayarensis, which Cyst: 40-50 µm have narrower body widths. Type Location: Italy, Napoli Polar capsule coils: ND

Henneguya amazonica Rocha, Matos & Azevedo Notes: Differences in spore body length separate this 1992 species from other Henneguya of overlapping total Host: gills of Crenicichla lepidota length except H. miyazakii & H. tachysuri, which differ Cyst: oval, up to 0.05 x 0.15 mm, asynchronous in spore body width, and H. sebasta, which has longer Type Location: Amazon estuary, Brazil polar capsules. Polar capsule coils: 6

Henneguya kayarensis Kpatcha, Faye, Diebakate, Notes: Shorter spore body length than other Fall,& Toguebaye 1997 Henneguya of overlapping total length except H. Host: liver of Galeoides decadactylus neopolitana, which has a wider spore body, and H. Cyst: none texana, which has a narrower polar capsule. Type Location: Senegal Polar capsule coils: ND

Henneguya sebasta Moser & Love 1975 Notes: Differences in spore body length separate this Host: heart (bulbus and truncus arteriosus) of from other Henneguya of overlapping total length Sebastes spp. except H. tunisiensis, which has a shorter tail; H. Cyst: 1-5 mm by 2 mm, white to light brown, rhinogobii, H. tachysuri, & H. amazonica, which differ round or irregular clumps in polar capsule length, & H. astyanax, which has a thinner body. Dimensions overlap with H. cutanea, Type Location: USA, central and s. California akule, yoffensis, vitiensis & miyazakii, so can only be coast differentiated by the different fish host family and the Polar capsule coils: 4-6 longer maximum total length. H. akule, yoffensis & vitiensis also infect heart tissue.

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Key to the identification of Henneguya species with unequal polar capsules 1a Maximum total spore length (spore + tail) ≥ 50 µm…..2 1b Maximum spore length < 50 µm…….7 2a Minimum spore body length >20 µm; polar capsules averaging 10-11 µm ……Henneguya theca 2b Minimum spore body length <20 µm…….3 3a Spore body width ranging from 6 to 7.5 µm……4 3b Spore body width range is less than 6 µm……….6 4a Total length (spore + tail) ≥ 56 µm……..Henneguya paranaensis 4b Total length < 56 µm……..5 5a Polar filament length about 10 µm…………Henneguya basifilamentalis 5b Polar filament length >10 µm, typically >26 µm………Henneguya ophicephali 6a Anterior end of spore is pointed (acuminate), larger polar capsule wider than smaller one….Henneguya notopteriae 6b Anterior end of spore is rounded (see figures for comparison), larger polar capsule narrower than smaller one……Henneguya singhi 7a Spore body length ≤ 9 µm, Marine fish host……….Henneguya joalensis 7b Spore body length >9 um…….8 8a Spore body length > 16 µm…….9 8b Spore body length ≤ 16 µm……...10 9a Spore body length about 20 µm, with small polar capsules (about 3 µm)…….Henneguya wenyoni 9b Spore body length smaller (17-19 µm), with larger polar capsules (4.4-6.0 µm)…. Henneguya namae 10a Tails typically of unequal length (about 3 µm difference)………Henneguya chrysichthyi 10b Tails typically of equal length……11 11a Minimum tail length ≥ 15 µm……..12 11b Minimum tail length < 15 µm………14 12a Spore body width ≥ 7 µm……….Henneguya chongqingensis 12b Spore body width < 7 µm…….13 13a Polar capsule lengths average ≤ 4 µm…….Henneguya logonensis 13b Polar capsule lengths average > 4 µm, typically 4.5-5.2 µm ……..Henneguya cyphocharax

14a Total length (spore + tail) averages < 20 µm…….15

14b Total length averages ≥ 20 µm……..16

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15a Polar capsule width typically ≥ 2.8 µm……. Henneguya markewitschi

15b Polar capsule width < 2.8 µm, typically 1.1-1.9 µm …….. Henneguya qadrii

16a Spore body width ≥ 9 µm……. Henneguya chalbulaevi

16b Spore body width < 9 µm……..17

17a Smaller polar capsule is wider than larger polar capsule……. Henneguya dispar

17b Smaller polar capsule is as wide or not as wide as the larger polar capsule…….18

18a Spore body shape is symmetrical, fusiform; gill parasite……Henneguya hemibagri

18b Spore body shape as asymmetric, broadly rounded at the anterior end; brain parasite…..Henneguya thermalis

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Henneguya species with Unequal Polar Capsules Henneguya markewitschi Allamuratov 1967 Notes: Shorter tail or shorter spore body length than other Host: skin, fins of Gobio gobio lepidolaenus Henneguya with unequal PCs, except H. qadrii, which had Cyst: 0.4-0.6 mm narrower polar capsules. Type location: Russia PC coils: ND Henneguya qadrii Lalitha Kumari 1965 Notes: Differences in spore body width separate this from Host: intestine of Ophicephalus gachua other Henneguya with unequal PCs, except H. markewitschi, Cyst: 1 mm which has wider polar capsules, H. dispar, which has shorter Type location: Hyderabad, India polar capsules, and H. hemibagri, wenyoni & chrysichthyi, Polar filament length, bigger PC: 45.4 (41.8- which have a longer spore body. Named for author’s 53.1); smaller PC: 34.5 (28.0-46.2) supervisor.

PC coils: 9 Henneguya chalbulaevi Gasimagomedov 1970 Notes: Wider spore body than other Henneguya with Host: Muscle of Rutilis rutilis unequal PCs, except H. markewitschi and H. chongqinensis, Cyst: 0.5 mm which differ in tail length and H. dispar which has shorter Type location: Russia polar capsules. PC coils: ND Henneguya dispar Ma, Wang, & Cai 1986 Notes: Longer tail than H. markewitschi, shorter polar capsules than H. qadrii & H. chalbulaevi, shorter spore body Host: gills of Channa striatus than H. hemibagri, and narrower body width than other H. Cyst: 167-158 μm, round of overlapping length with unequal polar capsules. Type Location: China

Henneguya hemibagri Zhang & Ma 1993 Notes: Shorter spore body than other overlapping Host: gills of Mystus macropterus Henneguya with unequal polar capsules except H. dispar, Cyst: 573 x 350 µm which is not as long bodied, H. chalbulaevi & H. Type location: China chongqingensis, which are wider bodied, H. logonensis, which has longer tail, & H. chryichthyi, which has a thicker PC coils: ND body. Dimensions overlap with H. thermalis, which has a See also Molnar et al. 2006 different spore shape and host tissue. Henneguya thermalis Seenhappa, Manohar, & Notes: Shorter tail than other overlapping Henneguya with Prabhu 1981 unequal polar capsules, except H. qadrii & H. chalbulaevi, Host: Brain of Lepidocephalichthys thermalis which differ in spore body width, H. chrysichthyi, which has Cyst: none reported a narrower polar capsule (longer one), & H. wenyoni, which Type location: near Honnenahalli, India is longer bodied. Dimension overlap with H. hemibagri, PC coils: ND which differs in body shape and host tissue. Similar in shape Museum specimen: Fish Pathology Unit, College to Hennegoides, but the axis of the polar capsules is not of Fisheries, Mangalore, India oblique to longitudinal axis.

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Henneguya chrysichthyi Obiekezie & Enyenihi 1988 Notes: Differences in spore body length or width Host: gills of Chrysichthys nigrodigitatus separate this species from other overlapping Cyst: polysporous, white, oval, 530(510-540) x Henneguya with unequal polar capsules except H. 265(260-280)µm; disporous pansporoblast thermalis, which has a wider polar capsule Type location: Cross River Estuary, Nigeria (longer one), H. cyphocharax, which has a longer PC coils: 9-11 tail, and H. hemibagri, which in thinner bodied. Data from formalin-fixed spores.

Henneguya wenyoni Pinto 1928 Notes: Pinto (1928) notes “The presence of polar capsules of large differences within the same Host: gills of Tetragonopterus sp. spore is relatively common”, but does not Cyst: 2 x 1 mm provide measurements. The species is listed here Type Location: Sao Paulo, Brazil as well as in the freshwater Henneguya. Longer PC coils: ND spore body than other H. of overlapping total length, except H. theca, minuta, pisciforme, doori, & creplini, which have longer polar capsules.

Henneguya chongqingensis Ma 1998 (Chen and Ma Notes: Wider spore body than other overlapping 1998) Henneguya with unequal polar capsules except H. markewitschi, which has a longer spore body, H. Host: ureter and urinary bladder of Carassius chalbulaevi & H. thermalis, which differ in tail auratus auratus length, and H. wenyoni, which is longer bodied. Cyst: not observed Type location: Chongqing City, Sichuan Province, China PC coils: ND

Henneguya logonensis Kostingue, Diebakate, Faye, Notes: Narrower spore body than other & Toguebaye 2001 overlapping Henneguya with unequal polar Host: gills of Citharinus citharus capsules except H. markewitschi, hemibagri, & Cyst: ovoid, 0.5-1.5 mm thermalis, which have shorter tails, H. theca, Type location: Logone River, Chad which has a longer spore body, and H. PC coils: ND cyphocharax, which has longer polar capsules. Museum specimen: Univ. C.A. Diop, Dakar, Senegal Henneguya cyphocharax Abdallah, Azevedo, Luque, Notes: Longer tail than H. thermalis & H. Teresa, & Bomfim 2007 chrysichthyi; shorter spore body than for H. Host: gills of Cyphocharax gilbert wenyoni & H. theca, but longer bodied than H. Cyst: 0.10-0.32 x .12-0.35 mm joalensis; narrower bodied than H. chongqinensis; Type location: Guandu River, Brazil differs in polar capsule metrics from H. Polar filament lengths: 48.2(23.1-55); 38.6(27.5- logonensis, namae, ophicephali & singhi. 48.4) Dimensions overlap with H. basifilamentalis, PC coils: 7-9 which has shorter polar filaments and infects a different host famliy.

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Henneguya namae Haldar, Das & Sharma 1983 Notes: Differences in spore body width separate Host: gill of Ambassis nama this species from other overlapping Henneguya Cyst: 180-220 x 150-190 µm; oval,cream-white, with unequal polar capsules except H. wenyoni, polysporous which is longer bodied, H. joalensis & H. Type location: Krishnagar, West Bengal, India basifilamentalis, which are shorter in body Polar filament lengths: 48.2(23.1-55); length, H. cyphocharax, which has wider polar 38.6(27.5-48.4) µm capsules, and H. ophicephali, which has longer PC coils: 9-10 in larger, 8-9 in smaller polar capsules. Museum specimen: Slide H/3, National Collection of the Zoological Survey of India, Calcutta

Henneguya joalensis Kpatcha, Faye, Diebakate, Fall, Notes: Shorter spore body than other & Toguebaye 1997 overlapping Henneguya with unequal polar capsules; H. ophicephali has longer polar Host: kidney of Cephalopholis taeniops capsules. Cyst: none observed Type location: Senegal coast PC coils: ND

Henneguya basifilamentalis Molnár, Székely, Notes: Wider spore body than other overlapping Mohamed, & Shaharom-Harrison 2006 Henneguya with unequal polar capsules except H. Host: gills of Hemibagrus nemurus wenyoni, which has a longer body, & H. Cyst: 580-720 x 240-320 µm chongqingensis, which has a wider body. Type location: Terengganu River, Malasia Dimensions overlap with H. cyphocharax & H. Polar filament length: 10 µm ophicephali, which have longer polar filaments PC coils: 7 and infects different host families. Museum specimen: HNHM-69910, Hungarian National History Museum Henneguya theca Kent & Hoffman 1984 Notes: Longer or thinner bodied than other Host: brain (usually medulla) of Eigemannia overlapping Henneguya with unequal polar virescens capsules. Cyst: none described Type location: Brazil Polar filament length: 19.9(17.1-23.4) PC coils: ND

Henneguya ophicephali Chakravarty 1939 Notes: Differences in spore body width separate Host: gills of Channa (syn. Ophiocephalus) this from other overlapping Henneguya with punctatus unequal polar capsules except H. cyphocharax, Cyst: oval to spherical; to 2 mm joalensis & namae, which have shorter polar Type location: Calcutta, India capsules. Dimensions overlap with H. Polar filament length: 26-32 µm basifilamentalis which has shorter polar filaments PC coils: ND and infects a different host family.

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Henneguya notopteriae Qadri 1965 Notes: Differences in spore body width separate this species from other overlapping Henneguya Host: gills of Notopterus notopterus with unequal polar capsules except H. joalensis, which has a shorter body, H. paranaensis & H. Cyst: none observed or found theca, which have a longer spore body. Dimensions overlap with H. singhi, which is not as Type location: near Hyderabad, India pointed at the anterior end. Also, the longer polar capsule of H. notopteriae has a longer duct, PC coils: ND appears golf-club shaped, and is wider than shorter PC.

Henneguya singhi Lalitha Kumari 1969 Notes: Narrower body than other overlapping Henneguya with unequal polar capsules except H. Host: gills of Notopterus osmanii cyphocharax, which has a wider polar capsule Cyst: none visible (longer one), and H. joalenis, H. theca, and H. Type location: Hyderabad, India paranaensis, which differ in body length. Polar filament length in longer PC: 33.5(26.0-38.5) Dimensions overlap with H. notopteriae, which is Polar filament length in shorter PC: 10.1(7.0-16.5) more pointed at the anterior end and its longer PC coils: ND polar capsule has a longer duct, is golf-club shaped, and wider than shorter PC.

Henneguya paranaensis Eiras, Pavanelli, & Notes: Longer total length than other Henneguya Takemoto 2004 with unequal polar capsules. Differentiated from H. notopteriae and H. singhi by the longer body Host: gills of Prochilodus lineatus length. Cyst: 0. 1 mm, rounded to oval Type location: Parana River, Brazil PC coils: 10-12 Museum specimen: Dept. Zoology & Anthropology, Faculty of Sciences of Porto, Portugal

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Laterocaudata Chen & Hsieh 1984 Ellipsoid Myxobolus-like spores with two polar capsules and a caudal appendage that arises laterally from the shell valve, rather than terminally as in Henneguya or Unicauda species. Type species is Laterocaudata mastacembala (Chen and Hsieh 1984) found in the gills of the freshwater fish Macrognathus aculeatus. The only other species in the genus is one transferred to this genus in this work. Gupta and Khera (1987) described Unicauda armati, but the drawing the myxospore indicates that the parasite found in the gills of Mastacembelus armatus belongs to Laterocaudata instead of Unicauda. So, herein the species is recognized as Laterocaudata armati comb. nov.

Laterocaudata armati comb. nov. Notes: syn. Unicauda armati. Tail stains pink Gupta & Khera 1987 with Giemsa stain whereas spore wall stains Host: gill filaments of Mastacembelus blue. Shorter tail than L. mastacembelus. armatus Cyst: Light yellow, almost rounded, 510 x Polar filament length: 12-17 µm

391 µm Type Location: Nangal & Perozepur, Punjab, India Polar filament length:12-17 µm

Laterocaudata mastacembela Chen & Notes: longer tail than L. armati. Hsieh 1984 Host: skin, fin, & gills of Mastacembelus aculeatus Cyst: round-elliptical, 13.6(8.1-18) x 12.0(8.4-15.6) µm Type Location: river near Tangkou, Yangjiang, Guandong, China

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Neohenneguya Tripathi 1952 Spindle-shaped spores with four filiform extensions, two extending from the anterior end and two from the posterior end of the shell. In the anterior end are two spherical polar capsules, one behind the other. The openings of the capsules are in different directions, the anterior polar capsule extruding to the front side, the other to the sutural side. Polysporous cysts. Histozoic.

Only one species, Neohenneguya tetraradiata (Tripathi 1952), described to date (Figure 12). The parasite was found in gill cysts (0.5 mm diameter) in the estuarine fish Odontamblyopus rubicundus from the Hooghly River, India. The shell valves are smooth and slightly unequal in size. The tails are equal in length. Iodiniphilous vacuole situated in posterior end of the body.

The measurements are: spore body length: 16.2-21.6 µm Figure 12. Neohenneguya tetraradiata Tripathi 1952 body width and thickness: 5.4 µm round polar capsules: 2.0-2.7 µm in diameter iodinophilous vacuole: 1.5-2.5 µm tail lengths: 63-72 µm

Phlogospora Qadri 1962 Flame-shaped spore with a single bottle-shaped polar capsule. Caudal projection is forked and its anterior end fits closely with the posterior end of the spore body. However, there is a distinct demarcation between the tail and spore body. No sutural line or ridge observable and the shell is smooth, without striations. The polar filament is coiled perpendicular to long axis of the spore and is symmetrical throughout. One known species, Phlogospora mysti (Qadri 1962; Fig. 13), has the following metrics:

Spore body: 14.0-18.0 µm x 3.5-5.0 µm Polar capsule: 7.0-9.0 x 1.75-2.5 µm tail length: 21- 24 µm Polar capsule coils: 10-11 (filament length: 65-70 µm) Total length: 35-42 µm Sporoplasmic nuclei: 2, which vary in location; 0.75-1.0 µm diameter Figure 13. Phlogospora mysti Qadri 1962

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Tetrauronema Wu, Wang & Jiang, 1988 Myxobolus-like spores with four thin ‘tails’ projecting posteriorly, two arising from each shell valve, each arranged symmetrically on either side of the longitudinal axis of the spore. Histozoic in freshwater & estuarine fishes. The type species is Tetrauronema macropodus (Wu et al. 1988), found in the gills of Macropodus opercularis (syn. M. chinensis) in China. The only other species known to date is T. desaequalis (Azevedo and Matos 1996). Wu et al. (1988) created a new family for this species, Tetrauronematidae, but Lom and Dyková (1992, 2006) included this genus within Myxobolidae. Given recent genetic studies that indicate that structures arising from the spores have less relationship to the relatedness among species than hosts and host tissues, I have also included this genus within Myxobolidae.

Tetrauronema macropodus Wu, Wang & Jiang, Notes: easily differentiated from T. desaequalis 1988 by the shorter spore body, equal tail lengths, Host: gills of of Macropodus opercularis (syn. M. shorter polar capsule, and fewer coils in the polar chinensis) filament. Freshwater. Cyst: greyish-white, round or ellipsoid, ~0.2 mm in diameter Type location: Fujian Province, China Polar capsule coils: 4-5 Tetrauronema desaequalis Azevedo et Matos, 1996 Notes: easily differentiated from T. macropodus Host: base of ventral fins of Hoplias malabaricus by the longer spore body, unequal tail lengths, Cyst: oval to ellipsoid, 1-2 mm in diameter, longer polar capsule, and more coils in the polar polysporous filament. Estuarine. Type location: Amazon River near Belém, Brazil Polar filament length: 50-60 µm PC coils: 9-11

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Trigonosporus Hoshina 1952 Overall spore shaped like an ‘A’. In valvular view, spore body is broadly triangular, with rounded anterior tip and a posterior end flattened perpendicular to the longitudinal axis. In sutural view, the spore is spindle-shaped. Each shell valve extends posteriorly on each side into two long filiform processes, creating a total of four tails. Each pair of tails is connected by a filament that parallels the base of the spore. Two equal polar capsules in anterior end of the spore are pyriform and convergent. Polysporous cysts. Histozoic in estuarine fish.

Only one species, Trigonosporus acanthogobii (Hoshina 1952) found in the gills of Acanthogobius flavimanus from Kanazawa Bay, Japan. Polysporous cysts were nearly spherical or ellipsoid, 560-760 µm x 410-650 µm. The spores were as described above, measuring 50-99 µm overall. The transverse filament was 13-19 µm long. Sutural ridge is not distinct; sutural line coincides with the largest diameter of the spore. The two symmetrical shells are smooth and about a micron thick. Polar filament has 4 coils within the polar capsule. Two spherical nuclei in the sporoplasm, 1.6-1.7 µm in diameter.

Measurements are {mean (range)}: spore body length: 8.7 (7.3-10.5) µm spore body width: 13.9(11.0-18.8) µm spore body thickness: 5.5(4.7-6.1) µm tail length: 68.3(50.4-99.1) µm transverse filament: 15.4(13.0-19.3) µm polar capsule length: 3.8(3.1-4.5) µm polar capsule width: 2.5(2.0-2.9) µm

Figure 3. Trigonosporus acanthogobii Hoshina iodinophilous vacuole diameter 3.3(2.7-4.8) µm 1952

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Unicauda Davis 1944 The genus Unicauda was described by Davis (1944), who divided the genus Henneguya into Henneguya (two-tailed, smooth shell), Unicauda (single tail), and Myxobilatus (two-tailed, striated shell, suture line lies between polar capsules). As described by Davis (1944), Unicauda features

“Spores ovoid with two polar capsules at anterior end. Posterior end of spore with single tail-like appendage which is not an extention of shell valves and composed of different material. Body of spore biconvex and compressed parallel to sutural plane. An iodinophile vacuole in sporoplasm. Tissue parasites. Polysporous.”

Davis (1944) transferred 10 Henneguya species to Unicauda: U. brachyura Ward 1919, U. carassii Fujita 1924; U. clavicauda Kudo 1934 (type species for the genus); U. crassicauda Kudo 1934; U. fontinalis Fantham, Porter, & Robertson 1939; U. macrura Gurley 1894; U. monura Gurley 1893; U. plasmodia Davis 1922; U. spatulata Fujita 1924; and U. strongylura Gurley 1894. To date, a total of 24 species have been described from North America, Eurasia, Japan, and Africa. The majority of the hosts are in the minnow family Cyprinidae, though Salmonidae, Mormyridae, Channidae, and three siluriform (catfishes) families also have Unicauda parasites (Table 1). A dichotomous key to the identification of the Unicauda species is provided at the end of this section.

The difference in the composition of the tail was noted early on. Gurley (1894) observed that the tail of U. macrura was completely dissolved in sulfuric acid and was almost invisible in balsam, whereas the shell valves that encase the spore body were not dissolved. Using Giemsa stains, Ward (1919) and Kudo (1934) observed that the tail of U. brachyura, U. clavicauda, and U. crassicauda stained pink or rose-red, whereas the body stained a clear blue. Cone and Melendy (2000) summarized the species of Unicauda, but did not include species for which the tail was not of a different material. This raises the issue of where should species that have one tail, but it is composed of the same material as the shell, be assigned? Recent genetic analyses (Smothers et al. 1994; Kent et al. 2001; Fiala and Bartosová 2010) indicate that some Henneguya species are more closely related to other Myxobolus species than to other Henneguya species. Liu et al. (2010) observed that Myxobolus turpisrotundus occurred as both tailed (10%) and untailed myxospores within the same cyst. More than a century ago Gurley (1894) reported the same observation of tailed and untailed forms in the same cyst. Dong et al. (2011), using 28S rDNA analysis, found that Unicauda pelteobagrus grouped with Myxobolus longisporus in a Myxobolus-Henneguya-Unicauda clade. This suggests that Henneguya and Unicauda species should be assigned to Myxobolus due to the priority of the latter (Bütschli 1882) or that a different classification is needed that better reflects the evolutionary relationships among myxozoans.

Nonetheless, the presence or absence of the tail has been a useful character for differentiating among the many species of both Henneguya and Myxobolus. So Henneguya has persisted in myxosporean taxonomic works (Kudo 1920; Lom and Dykova 1992), including this guide. Until the great majority of species are genetically analyzed using the same rDNA strands, and the evolutionary relationships sorted out, the data in this guide should still be useful for identifying tailed species. So, given the genetic overlap among the three genera and the observations of tailed and untailed forms of Myxobolus, I have chosen to provisionally maintain the current classification of species with a single tail, arising symmetrically from the posterior center of the spore body, within the genus Unicauda, whether the tail is made of a different material or not. Until genetic analyses indicate otherwise, it does not make sense to create a new genus for those species with tails that are of the same material as the spore shell, as in

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U. basiri (Bhatt and Siddiqui 1964) and the other species for which the tail composition is uncertain (Cone and Melendy 2000).

Table 1. List of Unicauda species sorted by fish host taxon. Host Order, Family Host species Host tissue Unicauda species Cypriniformes Cyprinidae Barbus arypus Liver U. lumae Barbus lacerata, Skin, under U. caudatus Barbus bocagei scales Campostoma anomalum Fin, skin U. crassicauda Carassius auratus U. wuhanensis Carassius carassius Muscle U. xihuensis (syn. C. vulgaris) Carassius carassius Kidney U. carassii Carassius carassius Kidney, U. spatulata spleen Hybognathus nuchalis Skin of head U. macrura Hypothalmichthys (syn. U. aristichthydis Aristichthys) nobilis Notropis anogenus Fins U. brachyura Notropis blennius Skin U. clavicauda Pimephales promelas Fins U. magna Osteoglossiformes Gut, kidney, U. trentinii sp.nov. Mormyridae Gnathonemus petersii liver Perciformes Channidae Channa (syn.Ophicephalus) Gill U. andhrae gachua Channa punctata Mouth cavity U. basiri & U. bengalensis Channa punctata Gill U. bicornuata & U. chaudhuryi Salmoniformes Salmonidae Salvelinus fontinalis Fins U. fontinalis Siluriformes Bagridae Rita rita Gill U. ritae Tachysurus (syn. Pelteobagrus) Muscle, gill U. pelteobagrus fulvidraco Ictaluridae Ictalurus punctatus Gill U. plasmodia Intestine U. fimbrethilae Mochokidae Synodontis schall Skin of head U. strongylurus

Some additional taxonomic issues have been discussed in the literature. Tripathi (1952) suggested that the form of Henneguya acerinae Schroeder 1906 described by Nemeczek (1911) should be transferred to Unicauda, as well as H. oviperda and H. limatula. However, Nemeczek (1911) noted that the tail was ‘cleaved’, so H. acerinae does not have a single tail. The tail of H. limatula is forked as well. H. oviperda was originally drawn as having a single tail, but there is no mention in the text by Cohn (1896) as to tail number or whether it was composed of different material. In a photomicrograph by Tyutyaev (2008), H.

117 oviperda appears to have a single tail, but it appears to be a continuation of the shell valve. So, until further data on the genetic relationship of H. oviperda to Unicauda is available, I have left the species within Henneguya. Tripathi (1952) found a muscle parasite in Ophicephali gachua that was very similar to Henneguya ophicephali, described by Chakravarty (1939) in the muscle of O. punctatus; Tripathi (1952) named the parasite Unicauda ophicephali and suggested that the form described by Chakravarty (1939) in the muscle, should also be considered U. ophicephali. Both Gupta and Khera (1987) and Cone and Melendy (2000) have questioned the validity of Unicauda ophicephali. I have also left U. ophicephali Tripathi 1952 as a junior synonym of Henneguya ophicephali. Unicauda monurus Gurley 1893, was originally described by Ryder (1880), who found polysporous cysts in muscle tissue of Aphredoderus sayanus near Woodbury, New Jersey, USA. However, no dimensions were provided in the description, so it has not been included in this guide. Alvarez-Pellitero and Gonzalez-Lanza (1985) described a single-tailed myxosporean, Myxobolus cutanei, that infects Barbus barbus bocagei in River Esla, Spain. Barbus lacerata cyri from the Kura River, Georgia, (former U.S.S.R.), had a very similar myxozoan parasite of the skin that was described as Myxobolus caudatus by Gogebashvili 1965. Myxobolus cutanei was considered the same species (junior synonym) as M. caudatus by Landsberg and Lom (1991), who reclassified the parasite as Unicauda caudatus.

Caffara et al. (2007) described a Unicauda species parasitizing the elephantnose fish Gnathonemus petersii, but did not name the species. It is unique from other Unicauda species, similar only to U. strongylura Gurley 1893, which was poorly described. U. strongylura is histozoic, whereas the form described by Caffara et al. (2007) is coelozoic, infecting a much different host and host tissue. I believe the species to be distinct from the current list of described Unicauda species. In consultation with the authors, the species has been affectionately named Unicauda trentinii sp. nov., in honor of the late Massimo Trentini, an Italian parasitologist, zoologist, and entomologist.

Key to species of Unicauda 1. a. total spore length ≤70 µm…..2 b. total spore length >70 µm…..23 2. a. minimum tail length < 40 µm….3 b. minimum tail length ≥ 40 µm……U. crassicauda 3. a. polar capsules lengths are generally equal …4 b. polar capsule length unequal (mean difference about 1 µm)…U. wuhanensis 4. a. tail not forked ….5 b. tail with small fork at the distal end….U. bicornuata 5. a. spore body longer than wide…..6 b. spore body wider than long….U. plasmodia 6. a. tail length less than spore body length….7 b. tail length as long or longer than body length…..10 7. a. tail is >2 µm wide, making tail look like a short ‘handle’…..8 b. tail narrow, ≤ 2 µm wide…….22

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8. a. spore body thick, ≥ 8.5 µm, coelozoic (internal) cysts to 600 µm…U. lumae b. spore body < 8.5 µm; cysts in skin, under scales, or fins …9 9. a. tail is a continuation of shell body, without clear separation, salmonid host…U. fontinalis b. tail is differentiated from spore body, cyprinid host….U. caudatus 10. a. tail <5 µm wide….11 b. tail about 5 µm wide, spore shaped like ‘tennis racket’…..U. spatulata 11. a. spore width < 6.5 µm wide ……12 b. spore width ≥ 6.5 µm……..20 12. a. tail wth cup-like receptacle attaching to spore body….13 b. tail without cup-like receptacle attaching to spore body…..14 13. a. tail length ≥ 30 µm……U. macrura b. tail length < 30 µm …..U. andhrae (U. trentinii may key out here; shorter tail than U. andhrae) 14. a. maximum spore body width ≤ 4.4 µm ….15 b. maximum spore body width 4.5 to 6.4 µm …….17 15. a. total spore length ≤ 30 µm …….16 b. total spore length > 30 µm ----U. chaudhuryi 16. a. polar capsule length ≥ 5 µm……U. basiri b. polar capsule length < 5 µm…….U. bengalensis 17. a. polar capsule length ≥ 5 µm……U. ritae b. polar capsule length < 5 µm…….18 18. a. total spore body length >32 µm……U. xihuensis b. total spore body length ≤ 32µm…….19 19. a. spore body length ≥ 9.6 µm ……U. aristichthydis b. spore body length < 9.6 µm ……U. strongylura 20. a. (from 11b) spore body length ≥ 12.5 µm …….U. pelteobagrus b. spore body length < 12.5 µm ……….21 21. a. tail ≥ 20 µm long…….U. clavicauda b. tail < 20 µm long…….U. brachyura 22. a. spore body width averages about 10 µm……..U. carassii b. spore body width ranging from about 4 to 6 µm …..U. trentinii 23. a. spore body length <7.9 um, caudal end of tail dissected into root-like branches…………………………….U. fimbrethilae b. spore body length > 7.9 um, caudal end of tail not dissected…….U. magna

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Unicauda species

Unicauda lumae Rahemo 1976 Notes: Wider spore body than other Unicauda of Host: liver of Barbus grypus overlapping total length, except U. brachyura, U. carassii, U. Cyst: 100-600 to 80-430 µm clavicauda, U. caudatus, and U. crassicauda, which are not Type Location: Tigris River, Iraq as thick-bodied. Tail shape varies from pointed to knob-like Polar capsule coils: 5 or club shaped; typically 2-4 µm long, but can reach up to See also Rashan et al. 1987 8.4 µm. Tails stains differently than spore. Unicauda trentinii sp. nov. Notes: Tail either twists away from the long axis of the spore Host: intestinal wall, kidney, gall or is straight. Differs in body width from other overlapping bladder of Gnathonemus petersii Unicauda except U. strongylurus, which is histozoic rather Cyst: small plasmodia, no size than coelozoic. Also differentiated by the different host. given Poor description of U. strongylurus makes comparison Type Location: Nigeria difficult, but drawing indicates U. trentinii has longer polar Polar capsule coils: 6 capsules too. See original description by Caffara et al. 2007. Unicauda strongylurus Gurley 1893 Notes: Tail twists away from the long axis of the spore. Host: skin of head of Synodontis Differs in body width from other overlapping Unicauda schall except U. trentinii, which infects a different host and host Cyst: >2.18 mm tissue; U. strongylurus also has macroscopic cysts. Type Location: Nile River Polar capsule coils: ND Unicauda plasmodia Davis 1922 Notes: Spore body wider than long. Shorter or wider spore Host: gills of Ictalurus punctatus body than other overlapping Unicauda species except U. Cyst: microscopic amoeboid brachyura, which has wider polar capsules, and U. plasmodia pelteobagrus, which has longer polar capsules, U. brachyura, Type Location: Mississippi River at which has shorter polar capsules, and U. fontinalis, which is Fairport, Iowa, USA wider bodied. Polar capsule coils: ND Unicauda carassii Fujita 1924 Notes: Total length by addition. Increase in prevalence with Host: kidney of Carassius vulgaris lower latitudes. Wider spore body than other Unicauda Cyst: 20 µm species of overlapping total length except U. crassicauda Type Location: Okayama, Japan and U. spatulata, which have longer tails, and U. lumae Polar filament length: 20-23 µm which has a thicker body. Shorter spore body than U. Polar capsule coils: ND brachyura. Dimensions overlap with U. fontinalis, which can be differentiated by the cyst size, host tissue, and different host. Unicauda caudatus Gogebaschvili Notes: Wider spore body than other Unicauda of 1965 overlapping total length, except U. lumae and U. Host: scales, skin of Barbus pelteobagrus, which differ in body thickness. Dimensions lacerata cyri & Barbus barbus overlap with U. fontinalis, but the tail in this species appears bocagei to be a continuation of the shell and not clearly separated as Cyst: whitish, under scale in U. caudatus. Syn. Myxobolus cutanei, Myxobolus Type Location: Krura River, Georgia caudatus (Landsberg and Lom 1991; see Unicauda text). Polar capsule coils:8-10, usually 9 Dimensions from Alvarez-Pellitero & Gonzalez-Lanza (1985). Polar filament length: 31.1(25.0- 39.5)

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Unicauda fontinalis Fantham, Porter & Notes: Longer or wider spore body than overlapping Richardson 1939 species of Unicauda except U. crassicauda, which has Host: fin of Salvelinus fontinalis a longer tail, and U. spatulata, which has a shorter Cyst: 1500 µm, disporous sporoblast polar capsule. Type Location: Quebec Province, Canada Polar capsule coils: ND

Unicauda aristichthydis Zhao & Ma 1995 Notes: Wider or shorter spore body than other Host: nephretic ducts of Aristichthys nobilis Unicauda of overlapping total length except U. Cyst: none described macrura which is not as thick-bodied, and U. Type Location: Chongqin, China wuhanensis, which has unequal polar capsule lengths

Unicauda wuhanensis Xiao & Chen 1993 Notes: Unequal polar capsule lengths separate this Host: nose, skin of Carassius auratus from all other Unicauda species. Differences in spore Cyst: none described body length or width also separate this from other Type Location: Wuhan, Hubei Province, China Unicauda of overlapping total length except U. Polar capsule coils: 3-4 aristichthydis, and U. macrura. The latter can also be differentiated by the longer tail.

Unicauda brachyura Ward 1919 Notes: Sutural edge with distinct folds (8-10). Not as Host: fin rays Notropis anogenus thick as U. lumae and U. clavicauda, with a shorter Cyst: 160-360 x 240 µm body length than other overlapping Unicauda species Type Location: Lake Erie, Ohio, USA except U. plamodia, which has a wider polar capsule, Polar capsule coils: ND and U. strongylurus and U. macrura, which are not as wide-bodied. Tail stains differently than spore body.

Unicauda bengalensis Chaudhuri & Notes: Narrower spore body than most other Chakravarty 1970 Unicauda species of overlapping total length except U. Host: mouth cavity of Channa punctatus bengalensis, which has shorter polar capsules, U. Cyst: oval, whitish, 1.0-1.25 mm bicornuata, which has a small fork at the end of the Type Location: Naihati, India tail, and U. andrhae, which has a cup-like shape to the Polar capsule coils: ND tail where it joins the spore body. Polar filament length: 21.7(18-27) µm

Unicauda xihuensis Wu & Li 1986 Notes: Shorter spore body or differences in body Host: muscle of Carassius carassius width separate this from other Unicauda of Cyst: none described overlapping total length except U. ritae, which has Type Location: China (West Lake, Hangzhou) longer polar capsules. Polar capsule coils: 4-5

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Unicauda basiri Bhatt & Siddiqui 1964 Notes: Narrower bodied than other Unicauda Host: root of buccal cavity of Channa punctatus species of overlapping total length, except U. Cyst: white, oval, 0.5-1.0 x 0.1-0.3 mm basiri, U. bicornuata, U. chaudhuryi and U. Type Location: India andhrae, which have longer polar capsules. Both Polar capsule coils: ND tail and body stain the same.

Unicauda clavicauda Kudo 1934 Notes: Differences in spore body length or width Host: skin of Notropis blennius separate this from other Unicauda species except Cyst: 1.0-1.5 mm U. lumae, which is thicker bodied, and U. Type Location: Rock River, Rockford, Illinois, USA brachyura, which is thinner bodied. Tail stains Polar capsule coils: ND differently from spore body.

Unicauda pelteobagrus Chen & Ma 1998 Notes: DNA analysis shows overlap with Host: gill, muscle of Pelteobagrus fulvidraco Henneguya & Myxobolus (Dong et al. 2011). Cyst: 500-1000 µm in muscle; 250-280 x 310-330 Differences in spore body length or width µm in gill separate this from other Unicauda of overlapping Type Location: Sichuan, China total length except U. plasmodia, which has Polar capsule coils: 4-7 shorter polar capsules. GenBank: HQ230327 Unicauda ritae Bajpai & Haldar 1982 Notes: Differences in spore body width separate Host: gills of Rita rita this from most other Unicauda of overlapping Cyst: none observed total length except U. xihuensis which has shorter Type Location: India polar capsules, and U. ahdhrae, which has a Polar filament length: mean 18.6 µm, range 16.7- shorter tail. 23.2 µm Polar capsule coils: 6-7

Unicauda bicornuata Chaudhuri & Chakravarty 1970 Notes: Narrower spore body than other Unicauda Host: gills of Channa punctatus species except U. andhrae, which has a cup-like Cyst: oval, white, 1.2-1.5 mm receptacle on the tail where it attaches to the Type Location: India spore body, U. bengalensis, which has shorter Polar filament length: mean 30.5 µm, range 26.8- polar capsules, and U. chaudhuryi and U. basiri, 35.6 µm which lack the fork in the tail. Polar capsule coils: ND Type specimen: Protozool. Lab., Calcutta Univ. Unicauda andhrae Kalavati & Narasimhamurti 1981 Notes: Narrower spore body than other Unicauda Host: gills of Channa gachua species of overlapping total length except U. Cyst: no cyst chaudhuryi, U. bicornuata, U. bengalensis and U. Type Location: freshwater ponds on way to basiri, which lack the cup-like receptacle on the Anantagiri, Visakhapatnam Dt., India tail where it attaches to the spore body. Polar filament length: mean 28.2 µm, range 25.4- 30.4 µm. Polar capsule coils: ND

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Unicauda spatulata Fujita 1924 Notes: tail is 5 µm wide and the ‘tennis racket’ shape of it separates this species from other Host: kidney, spleen of Carassius vulgaris Unicauda species. Cyst: 60-100 µm Type Location: Fukuoka & Shimane, Japan Polar capsule coils: ND Polar filament length: 60 µm

Unicauda macrura Gurley 1893 Notes: syn: Myxobolus macrurus. Differences in spore body width or length separate this Host: skin of head of Hybognathus nuchalis species from other Unicauda of overlapping Cyst: pinhead size total length except U. aristichthydis, which is Type Location: Neches River, Texas, USA thicker bodied, and U. wuhanensis, which has a Polar capsule coils: ND shorter tail. Tail is made of different material than spore.

Unicauda chaudhuryi Bajpai & Haldar1982 Notes: Longer spore body than most other Unicauda of overlapping total length; narrower Host: gills of Channa punctatus spore body than U. ritae and U. pelteobagrus, Cyst: no cyst and shorter tail than U. andhrae. Dimensions Type Location: Kalyani, West Bengal, India overlap with U. bicornuata, but its tail is not Polar capsule coils: 6-7 forked.

Unicauda crassicauda Kudo 1934 Notes: Longer tail than any other Unicauda species except U. magna, which is wider bodied Host: fin, skin of Campostoma anomalum and has a longer tail. U. crassicauda is similar Cyst: ovoid, 200-400 x 100-250 µm to U. macrura, but is wider and thicker bodied. Type Location: creek near Rockford, Illinois, USA Tail stains differently from spore body. Polar capsule coils: ND

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Unicauda fimbrethilae Rosser, Alberson, Notes: Larger than other Unicauda except U. Baumgartner, Mauel, Pote, and Griffin 2016 magna, which has a larger spore body. Host: intestinal tract of Ictalurus punctatus Projections on the end of the caudal process are Cyst: none observed variable in length 26.8(12.3-42.3) and width (0.9 Type Location: Catfish pond, Brooksville, (0.5-1.6). Immature spores lack these Mississippi, USA projections. Polar capsule coils: 4 GenBank: KT072742 Holotype: USNM 1283045, Smithsonian Institute Unicauda magna Minchew 1981 Notes: The large size (total length and body width) of this species differentiates it from all Host: fins of Pimephales promelas other Unicauda species. Tail stains differently Cyst: 1.0-1.5 mm from spore body. Type Location: Pennsylvania, USA Polar capsule coils: 5-6

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Appendix A List of Henneguya species of the world and the host tissue where they were found, organized by fish host taxon. The host’s habitat is also indicated, F for freshwater, M for marine, E for estuarine.

Host Order Host Host Henneguya Hab- Host Family Species Tissue Species itat Amiiformes Amiidae Amia calva gill amiae F

Characiformes Acestrorhynchus falcatus gill adherens E Acestrorhynchidae Characiformes Leporinus copelandi fins-cutaneous fonsecai F Anostomidae tissue Leporinus friderici gill, gut, kidney, friderici F liver Leporinus lacustris gill caudicula F Leporinus macrocephalus gill leporinicola F Leporinus mormyrops urinary ducts leporini F Leporinus obtusidens gills azevedoi F Schizodon fasciatus kidney schizodon F Leporinus copelandi muscle , skin travassosi F Characidae Astyanax altiparanae gill chydadea F Astyanax fasciatus peritoneal cavity bergamini F Astyanax fasciatus branchial cavity cesarpintoi F Astyanax fasciatus gill hoimba F Astyanax fasciatus; muscle , skin travassosi F Astyanax keithi gill astyanax E Astyanax scabripinnis gill artigasi F Astyanax scabripinnis cornea intracornea F Hyphessobrycon anisitsi gill pisciforme F Moenkhausia oligolepis testes testicularis E Tetragnopterus santae gills, branchial santae F arches Tetragonopterus sp. gills wenyoni F Citharinidae Citharinus citharus gill logonensis F Curimatidae Curimata inormata kidney curimata E Cyphocharax gilbert gill cyphocharax F Cyphocharax nagelii gill nagelii F Cyphocharax nagelli gill garavelli F Erythrinidae Hoplias malabaricus gill malabarica E Hemiodontidae Hemiodopsis microlepes gill hemiodopsis F (currently Hemiodus microlepis) Hepsetidae Hepsetus odoe gill nkamensis F Prochilodontidae Prochilodus lineatus gill caudalongula F Prochilodus lineatus gill paranaensis F Serrasalmidae Piaractus mesopotamicus visceral cavity& pellucida F

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tunica externa of swim bladder Piaractus mesopotamicus gill piaractus F Serrasalmus altuvei gill pilosa F Serrasalmus spilopleura gill curvata F Serrasalmus spilopleura gill iheringi F Serrasalmus striolatus gill striolata E Cypriniformes Lepidocephalichthys brain thermalis F Cobitidae thermalis (loach) Misgurnus front intestine hainanensis F anguillicaudatus Cyprinidae Abramis brama fins, skin cutanea M Campostoma anomalum fins, skin crassicauda F Carassius auratus ureter; urinary chongqingensis F bladder Carassius auratus subcutaneous miyairii F tissue of head Carassius auratus gills, skin, cornea of zikaweiensis F eye Carassius auratus auratus swim bladder rhomboides F Carassius auratus gibelio gill doneci F Cirrhinus mrigala gill bicaudi F Ctenopharyngodon idellus gill ctenopharyngodoni F Gobio gobio lepidolaenus skin, fins markewitschi F, E Mylopharyngodon piceus gills, gall bladder kwangtungensis F Notropis blennius skin (subdermal clavicauda F connective tissue) Rutilus rutilus muscle chalbulaevi F Schizothorax davidi urinary bladder tchangi F Silver crucian carp gill doneci F Varichorhinus gill varichorhinusi F (Onychostoma) sinus Nemacheilidae Cobitis barbatula urinary bladder legeri F Nemacheilus (syn. gill, kidney, liver obpyriformis F Noemachellus) yingjiangensis Cyprinodontiformes Gambusia affinis skin gambusi F Poeciliidae Umbridae Umbra limi gill umbri F Esociformes Esox lucius ovary oviperda Esocidae Esox lucius muscle, eye schizura F Esox lucius (syn. Lucius gill psorospermica F lucius) Esox lucius, Esox reichertii gill lobosa F Esox masquinongy gill acuta F Esox masquinongy dermis of inner gill nigris F arch

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Esox niger gill esocis F Gadiformes Lota lota kidney diplurus F Lotidae Gasterosteiformes Gasterosteus aculeatus kidney gasterostei F Gasterosteidae Gasterosteus aculeatus kidney tubules, media F ovary Gasterosteus pungitius kidney renicola F Pungitius pungitius skin pungitii F Gymnotiformes Electrophorus electricus large electric electrica F Gymnotidae organs Electrophorus electricus Kidney, liver, heart, visceralis F mesentery Hypopomidae Brachyhypopomus brain, spinal cord torpedo F pinnicaudatus Rhamphichthyidae Gymnorhamphichthys peripheral nervous rondoni F rondoni system Sternopygidae Eigemannia virescens brain, usually in theca F medulla Mugiliformes Mugil cephalus gill and heart ouakamensis F,E,M Mugilidae Osteoglossiformes Arapaima gigas gill, gall bladder arapaima F Arapaimidae Heterotis niloticus gill dini F Mormyridae Brienomyrus brachyistius gall bladder wall, ntemensis F kidneys and spleen Marcusenius moorii gill and muscle nyongensis F Marcusenius moorii gill odzai F Mormyrus cashive gill mailaoensis F Mormyrus cashive gill mormyri F Notopteridae Notopterus notopterus gill notopteriae F Notopterus osmanii gill singhi F Perciformes Ambassis nama (currently gill namae F Ambassidae Chanda nama) Anabantidae Anabas testudineus muscle manipurensis F Ctenopoma kingsleyae fins pinnae F Ctenopoma nanum gill ctenopomae F Ctenopoma petherici gill pethericii F Carangidae Selar crumenophthalmus aortic bulb akule M Centrarchidae Lepomis gibbosus eye episclera episclera F Channidae Channa (syn. viscera near spleen qadrii F Ophicephalus) gachua Channa argus gill, ovary, intestine gigas F Channa argus skin; kidney nanhaiensis F Channa argus gill orientalis F Channa argus abdominal cavity tunghuensis F Channa argus lips, intestine tangschensis F

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Channa argus; Clarias intestine, ovary sinensis F batrachus, Misgurnus anguillicaudatus, Channa maculatus Chana maculata body cavity vovki F Channa marulius kidney renalis F Channa punctata, C. gill & muscle ophiocephali F,E,M maculata Channa punctata gill waltairensis F Channa punctata gill zahoori F Channa striatus gill dispar F Channa striatus intestine ovaliformis F Cichlidae Aequidens plagiozonatus gill aequidens F Cichla temensis gill paraensis F Crenicichla lepidota gill amazonica E Saratherodon galilaeus intestine sarotherodoni F Clinidae Clinus superciliosus gills, gill arches clini M Eleginopsidae Eleginops maclovinus visceral organs shackletoni E, M Eleotridae Mogurnda obscura gill mogurndae F Oxyeleotris marmoratus gill shaharini F Gobiidae Acanthogobius flavimanus gill tridentigeri F Acanthogobius hasta gills multimorpha M, F Chaenogobius annularis skin; chorium miyazakii M Odontamblyopus (syn. fins rubicundi M,E Gobioides) rubicundus Periopthalmus gill periophthalmusi E,M cantonensis Periopthalmus gill periophthalmusi M cantonensis Rhinogobius giurinus intestine rhinogobii F,E,M Rhinogobius giurinus intestine rhinogobii M Rhinogobius sp. gill pseudorhinogobii F Rhinogobius typus urinary bladder sichuanensis F,E Rhinogobius typus urinary bladder sichuanensis M,E Tridendiger obscurus skin tridentigeri M Haemulidae Brachydeuterus auritus heart brachydeuteri M Labridae Symphodus tinca gill, gill arch tunisiensis M Lateolabracidae Lateolabrax sp. bulbus arteriosis; lateolabracis M gill Latidae Lates calcarifer gill latesa F,E Lates calcarifer gills, buchal mucosa latesi F,E Lates niloticus intestinal tract; gill; ghaffari F gill rakers Lates niloticus gill mandouri F Lates niloticus gill massii F

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Lates niloticus L. gill mbakaouensis F Leiognathidae Aurigequula (syn. heart muscle; blood vitiensis M Leiognathus) fasciatus Lutjanidae Lutjanus agennes gill lutjani E Lutjanus jocu gill jocu M Moronidae Morone chrysops ( syn. gill magna F Lepibema chrysops) Nandidae Nandus nandus Gill and heart nandi F Odontobutidae Perccottus glenii gill, ovary alexeevi F Osphronemidae Macropodus opercularis skin, fins macropodi F (syn: M. chinensis) Trichogaster trichopterus gill daoudi F Percichthyidae Plectroplites ambiguus gill australis F Siniperca chuatsi gill dogielli F Percidae Gymnocephalus (syn. gill acerinae F Acerina cernua; Aspro zingel; Lucioperca lucioperca; L. sandra Gymnocephalus cernua gill; eggs, ovary creplini F Gymnocephalus cernua connective tissue of tenuis F alimentary tract Lucioperca sandra gill acerinae F Lucioperca sandra gill gigantea F Perca flavescens gill doori F Perca flavescens gill percae F Perca fluviatilis gill minuta F Perca fluviatilis gill psorospermica F Perca fluviatilis gill similis F Perca fluviatilis skin wolinensis F Sander canadensis gill creplini F Polynemidae Galeoides decadactylus liver kayarensis M Priacanthidae Priacanthus arenatus gill priacanthi M Sciaenidae Cynoscion nebulosus heart cynoscioni F,E Otolithus sp. heart muscle otolithi M Pogonias cromis gill texana M Sciaenops ocellatus intestinal ocellata M epithelium, pyloric ceca Sebastidae Sebastes sp. heart, bulbus and sebasta M truncus arteriosus Serranidae Cephalopholis taeniops kidney joalensis M Sillaginidae Sillago analis psuedobranch; gill; lesteri M oral mucosa Sparidae Acanthopagrus schlegelii alimentary tract ogawai M wall

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Acanthopagrus schlegelii gall bladder wall yokoyamai F,E Dentex canariensis kidney mbourensis M Lagodon rhomboides skin of head lagodon M Pagrus caeruleostictus heart arterial bulb mauritaniensis Pagrus major heart,gill pagri M Sarpa (syn. Box) salpa kidney neopolitana M Sparus caeruleostictus gill, heart yoffensis F,E,M Tetrapontidae Therapon hillii gill gracilis F Percopsiformes Aphredoderus sayanus brevis F Aphredoderidae Salmoniformes Coregonus muscle kolesnikovi F Salmonidae Coregonus clupeoides muscle, skin tegidiensis F Coregonus fera, muscle, skin, fin zschokkei F Leucociscus leucociscus; Prosopium coulteri; P. williamsoni Oncorhynchus gorbusha, muscle salmonicola F O. kisutch, O. tschawytscha Oncorhynchus masou head cartilage cartilaginis F Salmo salar muscle salmonis F Salvelinus fontinalis muscle, fins nüsslini F Salvelinus sp. skin salvelini F Thymallus arcticus cranial cartilage; cerebralis F connective tissue Scorpaeniformes Comephorus baicalensis, kidney bayerii F Comephoridae C. dybowskii

Siluriformes Tachysurus thalassinus subcutaneous tachysuri M Ariidae muscle Bagridae Hemibagrus nemurus gill basifilamentalis F Mystus bleekeri kidney bleekeri F Mystus macropterus gillraker hemibagri F Mystus sp. gill mystusia F Tengra tengra (Mystus gill ophiocephali F bleekeri or M. tengara) Callichthyidae Hoplosternum littorale gill guanduensis F Corydoras melini gill melini F Clariidae Clarias anguillaris L. gill fusiformis F Clarias batrachus suprabranchial sinensis F organs, gills, intestine, ovary Clarias gariepinus gill assiuti F Clarias gariepinus gill samochimensis F

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Clarias gariepinus intestine, outer suprabranchiae F layer Clarias lazera gills clariae F Clarias lazera skin laterocapsulata F Clarias lazera suprabranchial suprabranchiae F respiratory organ Claroteidae Auchenoglanis gill auchenoglanii F occidentalis Chrysichthys gills, rarely dental bopeleti F nigrodigitatus palette Chrysichthys gills chrysichthyi E nigrodigitatus Heptapteridae Rhamdia quelen gill rhamdia F Rhamdia quelen (syn. linearis F Pimelodus sebae) Ictaluridae Ameiurus melas fins, base of 2nd gurleyi F dorsal Ameiurus nebulosus skin of barbels ameiurensis F Ictalurus furcatus skin, body wall of pellis F peritoneal cavity Ictalurus punctatus adipose fin adiposa F Ictalurus punctatus kidney, liver, diversis F connective tissue of muscle and fins Ictalurus punctatus gill exilis F Ictalurus punctatus gill ictaluri F Ictalurus punctatus gall bladder limatula F Ictalurus punctatus gill longicauda F Ictalurus punctatus gill mississippiensis F Ictalurus punctatus gill arch postexilis F Ictalurus punctatus skin sutherlandi F Ictalurus punctatus; I. gill; integument exilis F melas Loricariidae Loricaria sp. gill occulta F Malapteruridae Malapterurus electricus muscle malapteruri F Mochokidae Synodontis batesii gill camerounensis F Pangasiidae Pangasius hypophthalmus gill shariffi F Pimelodidae Platystoma fasciatum . linearis F Pseudoplatystoma gill corruscans F corruscans Pseudoplatystoma gill cuniculator F corruscans Pseudoplatystoma gill maculosus F reticulatum Pseudoplatystoma sp. gill eirasi F Pseudoplatystoma sp. gill multiplasmodialis F

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Psuedoplatystoma sp. gill pseudoplatystoma F Zungaro (syn. gall bladder lutzi F Pseudopimelodus) zungaro Schilbeidae Schilbe mystus gill ntondei F Sisoridae Glyptothorax trilineatus abdomen; tail yunnanensis F Sisoridae Pseudecheneis sulcata gill tengchongensis F

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Appendix B Illustration Credits Nearly all the illustrations of myxospores in this volume are redrawn from original figures. The source reference and publisher follow each species.

Dicauda Henneguya bayerii, Pronin and Pronin 2002, Russian Acad. Nauk Dicauda atherinoidi, Hoffman and Walker 1978, Henneguya bergamini, Gioia and Cordeiro 1996, Wiley & Sons Jagiellonian Univ., Krakow, Poland Hennegoides Henneguya bicaudi, Kaur & Attri 2015, Springer Hennegoides berlandi, Molnár et al. 2006, Inter- Henneguya bleekeri, Haldar and Mukherjee 1985, Research Sci. Center Elsevier Hennegoides longitudinalis, Lom et al. 1991, Inter- Henneguya bopleti, Fomena and Bouix 1987, Royal Research Sci. Center Musuem for Central Africa, Belgium Hennegoides malayensis, Molnár et al. 2006, Inter- Henneguya brachydeuteri, Kpatcha et al. 1997, Research Sci. Center Masson et Cie, Paris Hennegoides obpyriformis, Ma et al. 1986, Chinese Henneguya camerounensis, Fomena and Bouix 1987, Acad. Sci. Royal Musuem for Central Africa, Belgium Hennegoides pangasii, Molnár et al. 2006, Inter- Henneguya cartilaginis, Yokoyama et al. 2012, Research Sci. Center Elsevier Henneguya Henneguya caudalongula, Adriano et al. 2005, Inst. Henneguya acerinae, Schroder 1906, Elsevier Oswaldo Cruz, Brasil Henneguya acuta, Bond 1939, Am. Soc. Parasitol. Henneguya caudicula, Eiras et al. 2008, Jagiellonian Henneguya adherens, Azevedo and Matos 1995, Univ., Krakow, Poland Wiley & Sons Henneguya cerebralis, Pronin 1972, Propodniey Henneguya adiposa, Minchew 1977, Wiley & Sons uslovia I resersy Prikhubsugulya Henneguya aequidens, Videira et al. 2015, Springer Henneguya cesarpintoi, Gioia and Cordeiro 1996, Henneguya akule, Work et al. 2008, Am. Soc. Jagiellonian Univ., Krakow, Poland Parasitol. Henneguya chongqingensis, Chen and Ma 1988, Nat. Henneguya alexeevi, Schul’man 1966, U.S. Natural Sci. Found. China Government Henneguya chrysichthyi, Obiekezie and Enyenihi Henneguya amazonica, Rocha et al. 1992, Elsevier 1988, Royal Musuem for Central Africa, Belgium Henneguya ameiurensis, Nigrelli and Smith 1940, Henneguya chydadea, Barassa et al. 2003, Inst. New York Zool. Soc. Oswaldo Cruz, Brasil Henneguya amiae, Fantham et al. 1940, Cambridge Henneguya clariae, Abolarin 1971, E. Aftrican Univ. Press Literature Bureau Henneguya arapaima, Feijo et al. 2008, Elsevier Henneguya clini, Reed et al. 2007, open access Henneguya artigasi, Goia and Cordeiro 1987, Cong. Henneguya corruscans, Eiras et al. 2009, Elsevier Brasileira Zool. Henneguya creplini, Gurley 1893, U.S. Government Henneguya astyanax, Vita et al. 2003, Inter-Research Henneguya ctenopharyngodoni, Ma 1993, China Sci. Center Acad. J. Electronic Publ. House Henneguya auchenoglanii, Kostoingue et al. 2001 Henneguya ctenopomae, Fomena & Bouix 1996, Jagiellonian Univ., Krakow, Poland Royal Musuem for Central Africa, Belgium Henneguya australis, Johnston and Bancroft 1919, Henneguya cuniculator, Naldoni et al. 2014, Inter- Royal Soc. New S. Wales Research Sci. Center Henneguya azevedoi, Barassa et al. 2012, Springer Henneguya curimata, Azevedo and Matos 2002 Henneguya basifilamentalis, Molnar et al. 2006, Henneguya curvata, Barassa et al. 2003, open access Inter-Research Sci. Center Henneguya cutanea, Schul’man 1966, U.S. Government

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Henneguya cynoscioni, Dyková et al. 2011, open Henneguya hemiodopsis, Azevedo et al. 2009, Inst. access Oswaldo Cruz, Brasil Henneguya cyphocharax, Abdallah et al. 2007, open Henneguya hoimba, Cordeiro and Gioia 1987, Soc. access Brasiliera do Parasitol. Henneguya dini, Kabre et al. 1997, Eur. Assoc. Fish Henneguya ictaluri, Pote et al. 2000, Am. Fish. Soc. Pathol. Henneguya iheringi, Gioia and Cordiero 1996, Clube Henneguya dispar, Ma et al. 1986, Chinese Acad. Sci. Zoologico do Brasil Henneguya diversis, Minchew 1977, Wiley & Sons Henneguya intracornea, Gioia et al. 1986, Inst. Henneguya dogielli, Schul’man 1966, U.S. Oswaldo Cruz, Brasil Government Henneguya joalensis, Kpatcha et al. 1997, Masson et Henneguya doneci, Schul’man 1966, U.S. Cie, Paris Government Henneguya jocu, Azevedo et al. 2014 Henneguya doori, Guilford 1963, Am. Soc. Parasitol. Henneguya kayarensis, Kpatcha et al. 1997, Masson Henneguya doudi, Székely et al. 2009, Inter- et Cie, Paris Research Sci. Center Henneguya kwangtungensis, Chen and Ma 1998, Henneguya eirasi, Naldoni et al. 2011, Elsevier Nat. Natural Sci. Found. China Henneguya electrica, Jakowska and Nigrelli 1953, Henneguya lagodon, Hall and Iversen 1967, open New York Zool. Soc. access Henneguya episclera, Minchew and Sleight 1977, Henneguya lateolabracis, Yokoyama et al. 2003, Am. Soc. Parasitol. Wiley & Sons Henneguya esocis, Fantham et al. 1939, Cambridge Henneguya laterocapsulata, Landsberg 1986, Univ. Press Springer Henneguya exilis, Kudo 1929 , Elsevier Henneguya latesa, Wu et al. 1994, S. China Sea Inst. Henneguya fonsecai, Gioia and Cordeiro 1996, Oceanol. Jagiellonian Univ., Krakow, Poland Henneguya latesi, Tripathi 1952, Indian Museum, Henneguya friderici, Casal et al. 2003, Cambridge Kolkata, India Univ. Press Henneguya leporini, Nemeczek 1926, Elsevier Henneguya fusiformis, Kostoingue et al. 1999, Henneguya leporinicola, Martins et al. 1999, Acad. Jagiellonian Univ., Krakow, Poland Brasileira de Ciencias Henneguya gambusi, Parker et al. 1971, Am. Soc. Henneguya lesteri, Hallett and Diamant 2001, Inter- Parasitol. Research Sci. Center Henneguya garavelli, Martins and Onaka 2006, Henneguya limatula, Meglitch 1937, Am. Soc. Elsevier Parasitol. Henneguya ghaffari, Ali 1999, Inter-Research Sci. Henneguya lobosa, Cohn 1896, Fischer, Jena Center Henneguya logonensis, Kostoingue et al. 2001, Henneguya gigantea, Nemeczek 1911, Elsevier Jagiellonian Univ., Krakow, Poland Henneguya gigas, Chen and Hsieh 1960, Chinese Henneguya longicauda, Minchew 1977, Wiley & Acad. Sci. Sons Henneguya gracilis, Johnston and Bancroft 1919, Henneguya lutjani, Kpatcha et al. 1997, Masson et Royal Soc. New S. Wales Cie, Paris Henneguya guanduensis, Abdallah et al. 2007, open Henneguya macropodi, Shiba 1934, Keijo Teikoku access Daigaku Igakubu Henneguya gurleyi, Kudo 1920, Univ. Illinois, USA Henneguya macrura, Gurley 1893, U.S. Government Henneguya hainanensis, Chen and Ma 1998, Nat. Henneguya maculosus, Carriero et al. 2013, open Natural Sci. Found. China access Henneguya hemibagri, Tchang and Ma 1993, Nat. Henneguya magna, Rice and Jahn 1943, Iowa Acad. Natural Sci. Found. China Sci.

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Henneguya mailaoensis, Kostoingue et al. 2001, Henneguya ntondei, Fomena et al. 2008, Royal Jagiellonian Univ., Krakow, Poland Musuem for Central Africa, Belgium Henneguya malabarica, Azevedo and Matos 1996, Henneguya nyongensis, Fomena and Bouix 1996, Springer Royal Musuem for Central Africa, Belgium Henneguya malapteruri, Fomena and Bouix 1996, Henneguya nüsslini, Schuberg and Schroeder 1905, Royal Musuem for Central Africa, Belgium Elsevier Henneguya mandouri, Rabie et al. 2009, open access Henneguya occulta, Nemeczek 1926, Elsevier Henneguya manipurensis, Hemanand et al. 2008, Henneguya ocellata, Iversen and Yokel 1963, Univ. Turkish Soc. Parasitol. Miami Press Henneguya massii, Kostoingue et al. 2001, Henneguya odzai, Fomena and Bouix 1996, Royal Jagiellonian Univ., Krakow, Poland Musuem for Central Africa, Belgium Henneguya mauritaniensis, Khlifa et al. 2012, Henneguya ogawai, Li et al. 2012, Springer Springer Henneguya ophicephali, Chakravarty 1939, Elsevier Henneguya mbakaouensis, Fomena and Bouix 2000, Henneguya orientalis, Nie and Li 1992, Chinese open access Acad. Sci. Henneguya mbourensis, Kpatcha et al. 1997, Masson Henneguya otolithi, Ganapati 1941, Indian Acad. Sci. et Cie, Paris Henneguya ouakamensis, Kpatcha et al. 1997, Henneguya melini, Mathews et al. 2016, Springer Masson et Cie, Paris Henneguya minuta, Cohn 1896, Fischer, Jena Henneguya ovaliformis, Ma et al. 1986, Chinese Henneguya mississippiensis, Rosser et al. 2015 Acad. Sci. Henneguya miyairii, Kudo 1920, Univ. Illinois, USA Henneguya oviperda, Weltner 1892, Gesellschaft Henneguya miyazakii, Hoshina 1952, Tokyo Univ. Naturforschender Freunde zu Berlin Fisheries Henneguya pagri, Yokoyama et al. 2005 Henneguya mogurndae, Fujita 1936, Zool. Soc. Japan Henneguya paraensis, Velasco et al. 2016, Springer Henneguya mormyri, Kostoingue et al. 2001, Henneguya paranaensis, Eiras et al. 2004, Eur. Jagiellonian Univ., Krakow, Poland Assoc. Fish Pathol. Henneguya multimorpha, Li and Wang 1993, China Henneguya pellis, Minchew 1977, Wiley & Sons Ocean Press Henneguya pellucida, Adriano et al. 2005, open Henneguya multiplasmodialis, Adriano et al. 2011, access Elsevier Henneguya percae, Fantham et al. 1939, Cambridge Henneguya mystusia, Sarkar 1985, Jagiellonian Univ. Press Univ., Krakow, Poland Henneguya periophthalmusi, Wu et al. 1993, Henneguya nagelii, Azevedo et al. 2013, Springer Chinese Acad. Sci. Henneguya namae, Haldar et al. 1983, Elsevier Henneguya pethericii, Fomena et al. 2008, Royal Henneguya nandi, Gupta and Khera 1987, Punjab Musuem for Central Africa, Belgium University Henneguya piaractus, Martins and de Sousa 1997, Henneguya nanhaiensis, Chen and Ma 1998, Nat. Acad. Brasileira de Ciencias Natural Sci. Found. China Henneguya pilosa, Azevedo and Matos 2003, open Henneguya neopolitana, Parisi 1912, Soc. Ital. Sci. access Natur. Henneguya pinnae, Schubert 1967, Springer Henneguya nigris, Bond 1939, Am. Soc. Parasitol. Henneguya pisciforme, Cordeiro et al. 1983, Inst. Henneguya nkamensis, Fomena et al. 2008, Royal Butantan, Sao Paolo, Brasil Musuem for Central Africa, Belgium Henneguya postexilis, Minchew 1977, Wiley & Sons Henneguya notopteriae, Qadri 1965, Liepzig: Henneguya priacanthi, Kpatcha et al. 1997, Masson Akadamische Verlaggesellschaft Geest and et Cie, Paris Portig Henneguya pseudoplatystoma, Naldoni et al. 2009, Henneguya ntemensis, Fomena and Bouix 1996, Elsevier Royal Musuem for Central Africa, Belgium

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Henneguya pseudorhinogobii, Kageyama et al. 2009, Henneguya sutherlandi, Griffin et al. 2008, Am. Fish. Springer Soc. Henneguya psorospermica, Thélohan 1892, Soc. Henneguya tachysuri, Menon 1979, Mar. Biol. Assoc. Philomatique de Paris Henneguya tangschensis, Chen and Ma 1998, Henneguya pungitii, Schul’man 1966, U.S. Chinese Acad. Sci. Government Henneguya tchangi, Chen and Ma 1998, Nat. Natural Henneguya qadrii, Lalitha Kumari 1965, Univ. Sci. Found. China Messina, Italy Henneguya tegidiensis, Nicholas and Jones 1959, Henneguya renalis, Sarkar et al. 1985, Elsevier Cambridge Univ. Press Henneguya renicola, Stekhoven 1920, Elsevier Henneguya tengchongensis, Ma et al. 1986, Chinese Henneguya rhamdia, Matos et al. 2005, Wiley & Acad. Sci. Sons Henneguya tenuis, Vaney and Conte 1901, Soc. Henneguya rhinogobii, Li and Nie 1973, Nat. Natural Lineenne de Lyon Sci. Found. China Henneguya testicularis, Azevedo et al. 1997, Henneguya rhomoboideus, Ma et al. 1982, Chinese Springer Acad. Sci. Henneguya texana, Joy 1972, Wiley & Sons Henneguya rondoni, Azevedo et al. 2008, Wiley & Henneguya theca, Kent and Hoffman 1984, Wiley & Sons Sons Henneguya rubicundi, Haldar and Mukerjee 1985, Henneguya thermalis, Seenhappa et al. 1981, Elsevier Current Science Assoc. Henneguya salmonicola, Ward 1919, Am. Soc. Henneguya torpedo, Azevedo et al. 2011, Inter- Parasitol. Research Sci. Center Henneguya salmonis, Fantham et al. 1939, Henneguya travassosi, Guimaraes and Bergamin Cambridge Univ. Press 1933, Dept. Industria Animal, Brasil Henneguya salvelini, Zandt 1923, Elsevier Henneguya tridentigeri, Ozaki and Isizaki 1941, Henneguya samochimensis, Reed et al. 2003, open Tokyo Univ. Fisheries access Henneguya tunghuensis, Chen and Ma 1998, Nat. Henneguya santae, Guimarães and Bergamin 1934, Natural Sci. Found. China Dept. Industria Animal, Brasil Henneguya tunisiensis, Bahri et al. 2010, Springer Henneguya sarotherodoni, Fall et al. 2000, Elsevier Henneguya umbri, Guilford 1965, Wiley & Sons Henneguya schizura, Gurley 1893, U.S. Government Henneguya varicorhinusi, Chen and Ma 1998, Nat. Henneguya schizodon, Eiras et al. 2004, open access Natural Sci. Found. China Henneguya sebasta, Moser and Love 1975, Am. Soc. Henneguya visceralis, Jakowski and Nigrelli 1953, Parasitol. New York Zool. Soc. Henneguya shackletoni, Brickle et al. 2006, Springer Henneguya vitiensis, Laird 1950, Am. Soc. Parasitol. Henneguya shaharini, Shariff 1982, Wiley & Sons Henneguya vovki, Schul’man 1966, U.S. Government Henneguya shariffi, Molnar et al. 2006, Inter- Henneguya waitairensis, Narashimhamurti and Research Sci. Center Klavati 1975, Soc. Ital. Parassitol. Henneguya sichuanensis, Chen and Ma 1998, Nat. Henneguya wenyoni, Pinto 1928, Soc. Biol, (Paris) Natural Sci. Found. China Henneguya weishanensis, Hu 1965(Nie 1996), open Henneguya similis, Zandt 1924, Elsevier access Henneguya sinensis, Chen and Hsieh 1960, Nat. Henneguya wolinensis, Romuk-Wodoracki 1990, W. Natural Sci. Found. China Pomeranian Univ. Technol. In Szczecin Poland Henneguya singhi, Lalitha Kumari 1969, Univ. Henneguya yoffensis, Kpatcha et al. 1997, Masson et Messina, Italy Cie, Paris Henneguya striolata, Casal et al. 1997, Springer Henneguya yokoyamai, Li et al. 2012, Springer Henneguya suprabranchiae, Landsberg 1986, Henneguya yunannensis, Ma et al. 1986, Chinese Springer Acad. Sci.

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Henneguya zahoori, Bhatt and Siddiqui 1964, Wiley Unicauda brachyura, Ward 1919, Am. Soc. & Sons Parasitol. Henneguya zikaweiensis, Sikama 1938, Shanghai Sci. Unicauda carassii, Fujita 1924, Nihon Gakujutsu Inst. Kaigi Henneguya zschokkei, Gurley 1893, U.S. Unicauda caudatus, Alvarez-Pellitero and Gonzalez- Government Lanza 1985, Jena Fischer Laterocaudata Unicauda chaudhuryi, Bajpai and Haldar 1982, Univ. Messina, Italy Latercaudata armati, Gupta & Khera 1987, Punjab Unicauda clavicauda, Kudo 1934, Univ. Illinois, USA University Unicauda crassicauda, Kudo 1934, Univ. Illinois, Latercaudata mastacembela, Chen and Hsieh 1984, USA Nat. Natural Sci. Found. China Unicauda fimbrethilae, Rosser et al. 2016, Am. Soc. Neohenneguya Parasitol. Neohenneguya tetraradiata, Tripathi 1952, Indian Unicauda fontinalis, Fantham et al. 1939, Museum, Kolkata, India Cambridge Univ. Press Phlogospora Unicauda lumae, Rahemo 1976, Springer Phlogospora mysti, Qadri 1962, Elsevier Unicauda macrura, Gurley 1893, U.S. Government Unicauda magna, Minchew 1981, Wiley & Sons Tetrauronema Unicauda pelteobagrus, Chen and Ma 1998, Nat. Tetrauronema desaequalis, Azevedo and Matos Natural Sci. Found. China 1996, Am. Soc. Parasitol. Unicauda plasmodia, Davis 1922, Am. Soc. Tetrauronema macropodus, Wu et al. 1988, Chinese Parasitol. Acad. Sci. Unicauda ritae, Bajpai and Haldar 1982, Univ. of Trigonosporus Messina, Italy Trigonosporus acanthogobi, Hoshina 1952, Tokyo Unicauda spatulata, Fujita 1924, Nihon Gakujutsu Univ. Fisheries Kaigi Unicauda Unicauda strongylurus, Gurley 1893, U.S. Government Unicauda andhrae, Kalavati and Narashimhamurti Unicauda trentinii, Caffara et al. 2007, Eur. Assoc. 1981, Springer Fish Pathol. Unicauda aristichthydis, Zhao and Ma 1995, Chinese Unicauda wuhanensis, Xiao and Chen 1993, Nat. Acad. Sci. Natural Sci. Found. China Unicauda basiri, Bhatt and Siddiqui 1964, Wiley & Unicauda xihuensis, Wu and Li 1986, Chinese Acad. Sons Sci. Unicauda bengalensis, Chaudhuri and Chakravarty

1970, Jagiellonian Univ., Krakow, Poland Unicauda bicornuata, Chaudhuri and Chakravarty 1970, Jagiellonian Univ., Krakow, Poland

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Species Index Dicauda Henneguya clini………………………….……..94 Henneguya corruscans……………..……….40 Dicauda atherinoidi………………13 Henneguya creplini…………………….….….52 Hennegoides Henneguya ctenopharyngodoni…..…….66 Hennegoides berlandi…….……..14 Henneguya ctenopomae……………...……26 Hennegoides longitudinalis……14 Henneguya cuniculator…………….……….38 Hennegoides malayensis……….14 Henneguya curimata………………………50, 90 Hennegoides obpyriformis…….14 Henneguya curvata……………….…….…….54 Hennegoides pangasii……………14 Henneguya cutanea……………….….….…..90 Henneguya Henneguya cynoscioni…………….….……..90 Henneguya adherens…………….46, 88 Henneguya cyphocharax………………….106 Henneguya acerinae…………………80 Henneguya daoudi………..…………………..42 Henneguya acuta……………………. 72 Henneguya dini…………….…………….…….38 Henneguya adiposa………………….76 Henneguya dispar………….…..……………104 Henneguya aequidens………………54 Henneguya diversis……………………………76 Henneguya akule…………….………. 92 Henneguya dogielli………………..……….…78 Henneguya alexeevi………….………68 Henneguya doneci…………………….………70 Henneguya amazonica……………74, 100 Henneguya doori……………………….……..52 Henneguya ameiurensis……………72 Henneguya eirasi……………………….…….50 Henneguya amiae…………………….60 Henneguya electrica………………….…….52 Henneguya arapaima………………64 Henneguya episclera…………….….………78 Henneguya artigasi……….…………28 Henneguya esocis………………….…………78 Henneguya assiuti ……………………58 Henneguya exilis……………………………..76 Henneguya astyanax ……………60, 94 Henneguya fonsecai…………………….….34 Henneguya auchenoglanii…………70 Henneguya friderici……………………….…50 Henneguya australis…………………48 Henneguya fusiformis……………..……….72 Henneguya azevedoi…………………58 Henneguya gambusi………………………..36 Henneguya basifilamentalis……108 Henneguya garavelli………………………..40 Henneguya bayeri…………………….66 Henneguya ghaffari…………………….…..74 Henneguya bergamini………………26 Henneguya gigantea………………………..82 Henneguya bicaudi…………………..42 Henneguya gigas………………………..……64 Henneguya bleekeri……….…………28 Henneguya gracilis……………………..……50 Henneguya bopleti…………….…….58 Henneguya guanduensis………………..…48 Henneguya brachydeuteri……..…90 Henneguya gurleyi………………………..….76 Henneguya brevis…………….………22 Henneguya hainanensis……………………34 Henneguya camerounensis………24 Henneguya hemibagri………………..…..104 Henneguya cartilaginis…….………62 Henneguya hemiodopsis…………….….…26 Henneguya caudalongula…………80 Henneguya hoimba……………….………….32 Henneguya caudicula……….………22 Henneguya ictaluri……………….…………..82 Henneguya cerebralis………………38 Henneguya iheringi……………….………....32 Henneguya cesarpintoi………….…22 Henneguya intracornea……………...…….56 Henneguya chalbulaevi………….104 Henneguya joalensis……………….………108 Henneguya chongqingensis……106 Henneguya jocu…………………………..……94 Henneguya chrysichthyi…………106 Henneguya kayarensis……………..…….100 Henneguya chydadea………………24 Henneguya kwangtungensis……….…….80 Henneguya clariae………………….82 Henneguya lagodon………………….………86 Henneguya laseeae…………………….…….80

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Henneguya lateolabracis………………….96 Henneguya ogawai………………………84 Henneguya laterocapsulata……………..46 Henneguya ouakamensis…..………..84 Henneguya latesa……………………..……..86 Henneguya occulta……………..…..….56 Henneguya latesi…………………….……….36 Henneguya ocellata…………………….84 Henneguya legeri……………………….……28 Henneguya odzai…………………………46 Henneguya leporini………………….………40 Henneguya ophicephali….………….108 Henneguya leporinicola……………………42 Henneguya orientalis…………….…….52 Henneguya lesteri……………………….……86 Henneguya otolithi……………….……..96 Henneguya limatula……………….……….60 Henneguya ovaliformis………….…….26 Henneguya lobosa…………………..………44 Henneguya oviperda…………………….44 Henneguya logonensis…………………..106 Henneguya pagri…………………….……92 Henneguya longicauda……………….….82 Henneguya paraensis……………….….56 Henneguya lutjani……………………..…..98 Henneguya paranaensis…………….110 Henneguya macropodi…………….…….32 Henneguya pellis………………………….82 Henneguya maculosus…………………..46 Henneguya pellucida…………………...48 Henneguya magna………………………..80 Henneguya percae……………………….36 Henneguya mailaoensis………………….74 Henneguya periophthalmusi…….26, 84 Henneguya malabarica……………….40, 88 Henneguya pethericii…………….…….60 Henneguya malapteruri………………….60 Henneguya piaractus…………….…….68 Henneguya mandouri……………….…….70 Henneguya pilosa…………………..……70 Henneguya manipurensis………….……34 Henneguya pisciforme…….……..……42 Henneguya markewitschi………….…102 Henneguya pinnae………………………30 Henneguya massii…………………….……30 Henneguya postexilis……………….….76 Henneguya mauritaniensis………….…90 Henneguya priacanthi………………...92 Henneguya mbakaouensis………..……74 Henneguya pseudoplatystoma….…48 Henneguya mbourensis…………...….…88 Henneguya pseudorhinogobii……...66 Henneguya melini…………………….……54 Henneguya psorospermica………..…44 Henneguya minuta…………………………46 Henneguya pungitii………………………48 Henneguya mississippiensis……………64 Henneguya qadrii…………………….…104 Henneguya miyairii…………………….….38 Henneguya renalis………………………..34 Henneguya miyazakii…………………….96 Henneguya renicola…………….………..22 Henneguya mogurndae…………………74 Henneguya rhamdia…………….……….64 Henneguya mormyri……………………..44 Henneguya rhinogobii………………..66, 96 Henneguya multimorpha…………….24,84 Henneguya rhomoboideus…………….74 Henneguya multiplasmodialis….……38 Henneguya rondoni………………………24 Henneguya mystusia……………………..44 Henneguya rubicundi…………………...88 Henneguya nagelii…………………………50 Henneguya salmonicola……………..…60 Henneguya namae………………………108 Henneguya salmonis…………………….62 Henneguya nandi………………………….28 Henneguya salvelini………………………56 Henneguya nanhaiensis………….…….58 Henneguya samochimensis……………64 Henneguya neopolitana………….….…98 Henneguya santae…………………………26 Henneguya nigris………………….………68 Henneguya sarotherodoni………………54 Henneguya nkamensis………………….52 Henneguya schizura…………….…………68 Henneguya notopteriae……………...110 Henneguya schizodon…………….………36 Henneguya ntemensis……………..……24 Henneguya sebasta………..……………100 Henneguya ntondei………………………40 Henneguya shackletoni………………62, 98 Henneguya nyongensis…………………48 Henneguya shaharini………….………..…50 Henneguya nüsslini…………………..….42 Henneguya shariffi………………………….30

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Henneguya sichuanensis…………………28, 86 Neohenneguya Henneguya similis………………….….………66 Neohenneguya tetraradiata………….113 Henneguya sinensis………………….….…...58 Henneguya singhi……………………………110 Phlogospora Henneguya striolata………………………..56, 92 Phlogospora mysti…………………………113 Henneguya suprabranchiae……………...52 Tetrauronema Henneguya sutherlandi……………..……..78 Tetrauronema desaequalis…………….114 Henneguya tachysuri…………….….………98 Tetrauronema macropodus………..….114 Henneguya tangschensis………………….72 Trigonosporus Henneguya tchangi………………………….38 Trigonosporus acanthogobii……….….115 Henneguya tegidiensis………………..…..62 Henneguya tengchongensis…………….22 Unicauda Henneguya tenuis………………………..….22 Unicauda andhrae…………………………124 Henneguya testicularis……………….….40, 88 Unicauda aristichthydis………………...122 Henneguya texana………………..…..…….96 Unicauda basiri………………………….….124 Henneguya theca……………………………108 Unicauda bengalensis…………………...122 Henneguya thermalis………….………….104 Unicauda bicornuata……………….….…124 Henneguya torpedo……………….….…….68 Unicauda brachyura………………...……122 Henneguya travassosi…………….……….36 Unicauda carassii………………….……….120 Henneguya tridentigeri………..……….70, 98 Unicauda caudatus…………….………….120 Henneguya tunghuensis………………….80 Unicauda chaudhuryi…………………….126 Henneguya tunisiensis…………………….92 Unicauda clavicauda……………………..124 Henneguya umbri…………………………..58 Unicauda crassicauda………………….. 126 Henneguya varicorhinusi…………….….54 Unicauda fimbrethilae…………………..128 Henneguya visceralis……………….…….30 Unicauda fontinalis…………………….…122 Henneguya vitiensis……………….………94 Unicauda lumae………………………….…120 Henneguya vovki…………………….…..…46 Unicauda macrura………………………...126 Henneguya waitairensis……………..….72 Unicauda magna……………………………128 Henneguya wenyoni………………….42, 106 Unicauda pelteobagrus………………….124 Henneguya wolinensis……………….….78 Unicauda plasmodia………………….…..120 Henneguya yoffensis………………….….94 Unicauda ritae……………………….………124 Henneguya yokoyamai………………....86 Unicauda spatulata………………………..126 Henneguya yunannensis……………….32 Unicauda strongylurus………………..….120 Henneguya zahoori……………………….32 Unicauda trentinii…………………….…… 120 Henneguya zikaweiensis……………….30 Unicauda wuhanensis…………………... 122 Henneguya zschokkei………………..….62 Unicauda xihuensis………………………...122 Laterocaudata

Latercaudata armati……………………112 Latercaudata mastacembela…….…112

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