ARTICLE 383 ] is (the . . A . A third S. septigena Key words: Rozellida Rozellomycota straminipilous fungi was discovered and as the type species, , and Sparrow (1938) vs polysporangiate) was Foust is dependent upon how # Fungi, G IMA FUNGUS · 9(2): 383–399 (2018) 383–399 · 9(2): FUNGUS IMA R. achlyae, Pleolpidium S. allomycis, and . Subsequently, the developmental the developmental . Subsequently, for the monosporangiate members as the type species for Rozella . Following Foust’s discovery, Sparrow discovery, . Following Foust’s for the polysporangiate form, with R. as a synonym of Rozella P. monoblepharidis P. species, Batko (1977) erected the generic R. allomycis Rozella R. septigena , with as the type species. Clements & Shear (1931:234) as the type species. Clements & Shear and R. allomycis Pleolpidium Rozella Skirgiellia to accommodate the three polysporangiate To accommodate the divergent morphology observed To Foust (1937) discovered a second polysporangiate Foust (1937) discovered a second Rozella " The fourth-named thallus formed a single sporangium. by the absence species was distinguishable from the others by the formation from (or slightness) of host hypertrophy and that were separated the thallus of a linear series of sporangia Thus, at conception, there from each other by cross walls. within Rozella, the were two morphologically distinct forms containing Cornu’s “sporangium” (monosporangiate) form OP containing distinction (monosporangiate (1892) erected regarded as important, such that Fischer the genus Pleolpidium of genus would have to be made for their accommodation.” Sparrow (1938) and Karling (1942b) discussed in detail the # treated polysporangiate species, described by Shanor (1942). among name species, as S. achlyae, and retained septigena reiterated R. septigena species, */@+#$O=BR R. septigena and R. allomycis inconclusively the genus is to be interpreted. If it [Rozella to be restricted to those species in which a single sporangium results from an infection, then [those] two species [R. septigena ) (Cryptomycota Rozella

, R. , provide descriptions of all species, and include a , provide descriptions ), the apodyae Rozella R. apodyae , , and . rhipidii 1 , and the green alga Heterokontophyta Monoblepharidomycota , R. rhipidii spinosi , monoblepharidis polymorphae . Dick (2001: 246) treated the ). Cornu erected the genus , and R. apodyae . Basidiomycota rhipidii spinosi and , R. septigena; he did not hyphenate the apodyae

and Martha J. Powell and Martha 1 , R. rhipidii is a genus of endoparasites of a broad range of hosts. Most species are known by their of a broad range of hosts. Most is a genus of endoparasites , and Charophyta , and and Blastocladiomycota \\" ( %# Rozella

(Cornu 1872) is a genus currently consisting of (Cornu 1872) is a genus currently currently nests among an abundance of environmental sequences in phylum Cryptomycota, superphylum an abundance of environmental sequences currently nests among monoblepharidis <= NOQ=R%U*@@QVBO#+WX@ZZ@XX\^=U^@_Z+WRQU key to the species of Rozella 2018; Published: 16 November 2018. Accepted: 8 November 2018; Article info: Submitted: 18 September Opisthosporidia Abstract: Rozella corresponding author e-mail: [email protected] corresponding author earliest diverging lineage in kingdom in molecular phylogenies to be the genome sequenced. Determined © 2018 International Mycological Association © 2018 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: 1 Peter M. Letcher A taxonomic summary and revision of and revision summary taxonomic A Non-commercial:No derivative works:if you get Any of the above conditions can be waived For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. induced hypertrophy (swelling as a result of an increase in and brachynematis These names names, but these are to be added (Art.23.1). formally described, were those used when the species were short versions of the but Cornu (1872) interchanged long and #$ brachynematis long and short versions as alternative names, and stated that “the selected epithet is therefore determined by accepted usage” and pointed out that Fischer (1892), Minden (1915), Sparrow (1938), and Karling (1942b) had used the shorter versions, and he followed their choice, rather than the longer versions used by Sparrow (1960) or Karling (1977), “because % P ' *+/% species are herein subsequently referred to as R. of Cornu’s monoblepharidis Coleochaete in common: (1) a to describe four species, which had \ through a circular (for three of the species) that escape " of a papilla; and (3) opening that results from the dissolution resting spores with the formation of spherical, thick walled species were described as spiny ornamentations. Cornu’s monoblepharidis polymorphae Fungi phyla Chytridiomycota 27 species of endobiotic, holocarpic, unwalled parasites 27 species of endobiotic, holocarpic, of a variety of hosts in Oomycota ( INTRODUCTION Rozella doi:10.5598/imafungus.2018.09.02.09 VOLUME 9 · NO. 2 Letcher and Powell

type species). Rozella was retained for the monosporangiate of concentric bodies (Figs 2E, 5A, 6A). The cyst produces an forms. Batko (1977) suggested that another of Cornu’s VM„U originally described species, R. monoblepharidis, supposedly # “…the most common species in the New as well as the Old the host cell wall (Fig. 2I). The host plasma membrane is ' B pushed inward as the parasite protoplast is discharged

ARTICLE O{PG"P through an opening in the infection tube. Empty cysts (Fig. as pointed out above, a type had already been selected and V\%„= “page priority” is not a consideration in selection. Further, as occupies a compartment within the host cytoplasm (Fig. 2E, Batko included the type of Rozella in his concept of Skirgiella, G, I), then enlarges into an unwalled sporangial plasmodium his generic name is illegitimate as it was nomenclaturally (Fig. 3A) or walled resting spore (Fig. 6). In sporangial \U_*_=N plasmodium development, the plasmodium produces lobed (2014)’s introduction of the family name Skirgielliaceae was # "  @O ZU also illegitmate as it was based on an illegitimate generic often a vacuole occupies the center of the plasmodium (Figs name (Art. 18.3); Doweld also validated two epithets of 3B, 4A). At maturity the multinucleate sporangial plasmodium %Rozella as they had lacked  @O ZU " a Latin diagnosis, achlyae and allomycetis; the necessary  ZO < " % nomenclatural corrections are provided below. (Figs 4B, 5) and released through a discharge pore (Fig. Currently, Rozella consists of 24 monosporangiate and 5B) or tube (Fig. 5C). In resting spore development, multiple \ */W */W@ plasmodia occupy a host compartment (Fig. 6A). Resting 1974, 1975, 1980, 1981) provided seminal insights regarding % the morphology, physiology, and ultrastructure of the become sphaerical (Fig. 6A), and resting spores of most polysporangiate R. allomycetis, and his observations Rozella species have spiny wall ornamentation (Fig. 6B). served as a basis for later comparisons among other Rozella has engendered much interest over the last species. Recent investigations (Letcher et al. 2017, 2018) decade, beginning with two strains in a molecular phylogeny % % " that occurred as the earliest diverging lineage in the fungi monosporangiate and polysporangiate species, indicating (James et al. 2006). One strain (JEL 347, R. rhizoclosmatii) was monosporangiate, while the second (UCB 47-54, "~ R. allomycetis) was polysporangiate. Subsequently, this The Rozella life-cycle has been described elsewhere, lineage came to include not only the two strains of Rozella, "'% but a myriad of novel small subunit ribosomal RNA gene " *M 3  *U sequences (SSU rRNA) derived from environmental "‚ surveys (e.g. Lara et al. 2010, Jones et al. 2011a, Karpov and attach to the host thallus (Fig. 1B). In polysporangiate et al. X*Z ^" Š ‹ X*_ Œ et al. 2016, forms, the parasite induces host hyphae to produce septa, ‘{‹" et al. 2017, Tedersoo et al. 2017). Initially "% referred to as the Rozella clade (James et al. 2006), then into unwalled sporangial plasmodia (Fig. 1C) or walled resting O"P ^ et al. 2010), the lineage is now assigned spores (Fig. 1D). In monosporangiate forms, the parasite to the Cryptomycota (Jones et al. 2011b), and more recently  *VU " has been also referred to as Rozellomycota (e.g. Corsaro et may or may not swarm in the sporangium before discharge, al. 2014) and Rozellosporidia (Karpov et al. 2017). Because and may emerge as a mass and immediately disperse (Fig. of the position of Cryptomycota # *M„ „ "  UMB # *M3{ ‹ et al. (2013) proposed that Cryptomycota \*/W_%# and Microsporidia share a common ancestor. Karpov et al. %"„R. (2014) erected the superphylum Opisthosporidia to include rhizoclosmatii" three phyla: Cryptomycota, Aphelida, and Microsporidia. rods, as shown by serial sections (Letcher et al. 2017) is Opisthosporidia was considered sister to Fungi. More appressed to the surface of the nucleus (Fig. 2D). Posterior to recently, Tedersoo et al. (2017) placed Aphelida as sister of the nucleus is a single sphaerical mitochondrion nestled into Blastocladiomycota in kingdom Fungi, and Cryptomycota + %UMB Microsporidia as a lineage sister to Fungi. Bass et al. (2018), " however, placed Cryptomycota as sister to Microsporidia. In \OMB=\ # # % # = et al. 2018). Obviously, UMB„ Rozella and "{#†^B Cryptomycota. composed of one or more lipid globules, a microbody, and There has been considerable confusion over names to a backing membrane, and ribosomes are dispersed in the be applied to these organisms, and here we aim to provide a UMB %# †" of both the phylum and the genus Rozella based on the VM„R. allomycetis, intact current International Code of Nomenclature for algae, fungi host cytoplasm of Allomyces is distinguishable by the presence and plants (ICN; Turland et al. 2018).

384 IMA FUNGUS Taxonomy of Rozella (Cryptomycota) ARTICLE

Fig. 1. Light microscopic images of Rozella. A.†"R. rhizoclosmatii [strain JEL 863, Letcher et al. 2017]. B–D. R. allomycetis and its host Allomyces macrogynus [strain UM690, unpubl.]. B.V"C. Parasite sporangia in septate segments of host hypha. D. Spiny parasite resting spores in septate segments of host hypha. E. Rozella rhizoclosmatii. Uninfected sporangium of host Rhizoclosmatium globosum"“‹V^+”@^ et al. 2017]. F–I. Rozella multimorpha ‚ Pythium catenulatum [strain JEL 883, unpublished; see Letcher et alX*+GU%$BO\ Q Q QOU•*3OMN•*_3V•*XX3M„•*X3

TAXONOMY Circumscription emended by Cavalier-Smith & Chao (1995) and Adl et al. (2005). = # - standing and interpretation of these organisms. Additionally, SUPERKINGDOM: Holomycota Y. Liu, BMC Evol. Biol. 9.272: 3 (2009). without morphological, molecular, or ultrastructural evidence Synonym: Nucletmycea †' O Mol. Biol. Evol,. 26: \- 2706 (2009). late Rozella%%# the genus (Dick 2001). KINGDOM: Fungi R.T. Moore, Bot. Marina 23: 371 (1980).

SUPERGROUP: Opisthokonta Cavalier-Smith, in Rayner et SUBKINGDOM: Rozellomyceta Tedersoo et al., Fungal Divers.: al., Evol. Biol. Fungi: 344 (1987). doi.org/10.1007/s13225-018-0401-0: 13 (2018).

VOLUME 9 · NO. 2 385 Letcher and Powell ARTICLE

Fig. 2. Transmission electron microscopic images of Rozella"A. R. allomycetis [strain CSF 55; Powell et al. 2017]. B. R.multimorpha [strain JEL 883; Letcher et al. 2018]. C–D. R. rhizoclosmatii [strain JEL 863; Letcher et al. 2017]. D. A lattice of perpendicular rods (arrows) about the nucleus. E–I. R. allomycetis“R†”/X ^G"Allomyces macrogynus. E. Early stages of infection. F.V"G.‚H.V"I.B" U%$UB"B%%BOBB" VB"\\„=–^†<  "‚"OUM„•X_3OMN•X_3V•3

386 IMA FUNGUS Taxonomy of Rozella (Cryptomycota)

Diagnosis: “Vegetative cells amoeboid, with pseudopodial THE SPECIES ARTICLE ##" \ " Rozella achlyae (Doweld) Letcher, comb. nov. with a long kinetosome; one single large mitochondrion MycoBank MB827617 (missing in Microsporidea); resting spores thick-walled; Basionym: Skirgiellia achlyae Doweld, Index Fung. 129: 1 chitinous wall present only in some life stages; penetration (2014). of host cells via germ tube; intracellular obligate parasites of Synonyms: Rozella achlyae Shanor, J. Elisha Mitchell Sci. fungi, animals and protists that consume host organelles via Soc. 58: 100 (1942); nom. inval. (Arts. 39.1, 40.1). phagocytosis” (Tedersoo et al. 2018). Skirgiellia achlyae (Shanor) A. Batko, Acta Mycol. 13: 322 Type: Rozella Cornu 1872. (1977); nom. inval. (Arts. 39.1, 40.1).

SUPERPHYLUM: Opisthosporidia Karpov et al., Frontiers Microbiol. Type: Shanor (J. Elisha Mitchell Sci. Soc. 58$*W*MW 5: 8 (2014). 1942; lectotype designated here, MBT 384204).

PHYLUM: Cryptomycota M.D.M. Jones & T.A. Richards, IMA Description: “An endotrophic parasite of Fungus 2: 173 (2011). causing very slight or no hypertrophy. Young plasmodium Synonyms: Rozella clade, James et al., Mycologia 98: 863 hardly distinguishable in the host protoplasm, hyaline and (2006). very nearly optically homogeneous. Sporangia formed in Rozellida Lara et al., Protist 161: 117 (2010). a linear sori, cylindrical to somewhat barrel shaped, length Rozellomycota T.Y. James & Berbee, BioEssays 34: 96 # (2011); nom. inval. (Art. 36.1(a)); D. Corsaro & R. Michel, *_ 3 Parasit. Res. 113: 1916 (2014); nom. inval. (Art. 36.1(a)). "‚‘ Rozellomycota Doweld, Index Fung. 43: 1 (2013). %M@&@MZ3% Rozellosporidia Karpov et al., J. Eukaryotic Microbiol. 64: 573 \*M*_ (2017). 3 Note: The circumscription was emended by Karpov & in host hyphae that resemble sporangial sori, each segment Aleoshin (2014). The name was not validly published earlier containing from one to many resting bodies. Resting bodies in the year by Jones et al. (2011a). As the name is not based % *M”M@W 3 on that of an included genus, no type was required for valid *_+M*W@ 3 % publication (Art.10.10). The name was based on the cryptic *”M@3 nature of the majority of its inclusions, namely environmental in length, wall of mature resting bodies thick, reddish-brown sequences. to amber brown in color. Resting spore germination follows a dormant period and is accomplished by the formation of SUBPHYLUM: Rozellomycotina Tedersoo et al., Fungal Divers. \" doi.org/10.1007/s13225-018-0401-0: 13 (2018). #PQ*/Z Diagnosis: As for subkingdom above. Type: Rozella Cornu 1872. Hosts: , A. proliferoides, Dictyuchus anoma- lus, and D. monosporus (Johnson 1955) (Oomycota). CLASS: Microsporidia Corliss & Levine, J. Protozool. 10: 26 (1963). Rozella allomycetis (Doweld) Letcher, comb. nov. MycoBank MB827616 GENUS: Rozella Cornu, Ann. Sci. Nat., Bot., sér. 5, 15: 148 Basionym: Skirgiellia allomycetis Doweld, Index Fung. 129: 1 (1872). (2014); as “allomycis”. Synonyms: Skirgiellia A. Batko, Acta Mycol. 13: 321 (1977); Synonym: Rozella allomycis Foust, J. Elisha Mitchell Sci. nom. illegit. (Arts. 52.1, 52.2). Soc. 53: 198 (1937); nom. inval. Rozia Cornu, Bull. Soc. Bot. France 19: 71 (1872); nom. (Arts. 39.1, 40.1). illegit. (Art. 53.1); non Rozea Bescherelle, Mém. Soc. Skirgiellia allomycis (Foust) A. Batko, Acta Mycol. 13: 322 Sciences Nat. Cherbourg 16: 241 (1872); nec Rosea (1977); nom. inval. Fabr., Enum.: 47 (1759). (Arts. 39.1, 40.1). Pleolpidium A. Fischer, Rabenhorst’s Kryptogamen-Fl. 1: 43 Rozella allomycetes Nabel, Arch. Mikrobiol. 10: 527 (1939); (1892). nom. inval. (Art. 38.1(a)). Type: Rozella septigena Cornu1872. Note: Dick (2001) designated R. monoblepharidis as the type Type: Foust (J. Elisha Mitchell Sci. Soc. 53$*MW species of Rozella ‘ { @ *MW */@W lectotype designated here, MBT named species (Cornu 1872), but that mechanical method 384205). is contrary to the Code (Arts 10.5, 10.6), and in any case Clements & Shear (1931) had already selected a different Description: “Fungus body parasitic within the distal parts of one of the original species, R. septigena, and that must be the hyphae of AllomycesQ followed (Art. 10.5) unless changed by conservation. *M_ {%"

VOLUME 9 · NO. 2 387 Letcher and Powell ARTICLE

Fig. 3. Plasmodial development in Rozella. A. Early plasmodial formation of R. multimorpha [strain JEL 883; Letcher et al. 2018] in host Pythium catenulatum. B. Multinucleate plasmodium of R. allomycetis [strain CSF 55; Powell et al. 2017] in host Allomyces arbuscula; plasmodium has a %U%$\ <š%O•_3

Fig. 4. Plasmodial development in Rozella. A. Multinucleate plasmodium of R. rhizoclosmatii [strain JEL 863; Letcher et al. 2017] in host Rhizoclosmatium globosum; plasmodium has a central vacuole. B.V"%R. allomycetis [strain CSF 55; Powell et al. 2017] in host Allomyces arbuscula"\%U%$\ † <š%O•*3

388 IMA FUNGUS Taxonomy of Rozella (Cryptomycota) ARTICLE

VOLUME 9 · NO. 2 389 Letcher and Powell ARTICLE

Fig. 5.B"Rozella. A.‚R. allomycetis [strain CSF 55; Powell et al. 2017] in host Allomyces arbuscula; septum separating host from parasite was induced by parasite. B. ‚ R. allomycetis [strain CSF 55] being released through a discharge pore. C.‚R. multimorpha [strain JEL 883; Letcher et al. 2018] in host Pythium catenulatum; some spores are in the U%$BON N= \\ Q Q‚"O•_3

*MX&XMZX3*_/3 formed later than the sporangia, occurring in the distal part &Z”3= of the host hyphae just behind the sporangia in swollen R#*@3 *M@_ Frequently a primary sporangium may be divided by one or *M*” % @ Z more partitions into several smaller sporangia. ‚ Segments spherical, barrel-shaped, nearly cylindrical, or % @MZ 3 XMZX&XMWX3Q containing one refractive globule and with one posteriorly by resting bodies, usually containing some left-over, dead, \ granular, host protoplasm. Resting bodies with spiny walls, and is directed backward when the spore swims. Swimming *MX 3 % *_/ 3 by darting about with frequent changes of direction as typical (average of 20 measurements) counting the spines which for chytrid spores. The swimming period lasting about an *@3 hour, after which the spores germinate or die. Resting bodies _3

390 IMA FUNGUS Taxonomy of Rozella (Cryptomycota) ARTICLE

Fig. 6. Resting spores of Rozella allomycetis [strain CSF 55; Powell et al. 2017] in host Allomyces arbuscula. A. Resting spore formation. B. QU%$BO\ < QO•_3 hyaline globose mass of material surrounded by a granular Rozella apodyae Cornu, Ann. Sci. Nat., Bot., sér. 5, "Œ 15: 161 (1872). " Synonyms: Rozella apodyae-brachynematis Cornu, Ann. of Allomyces is present. The resting bodies may retain Sci. Nat., Bot., sér. 5, 15: 161 (1872). their vitality either dry or wet for several weeks, perhaps Rozella apodachlyae†'NStramin. Fungi: 373 (2001); ‚ as “M. Cornu”; nom. illegit. (Art. 52.1). from regular sporangia and capable of infecting young host Pleolpidium apodyae-brachynematis (Cornu) A. Fischer, hyphae” (Foust 1937). Rabenh. Krypt.-Fl. 1 (4): 45 (1892); as “apodyae”.

Hosts: Allomyces arbuscula, A. javanicus, A. macrogynus, Type: Cornu (Ann. Sci. Nat., Bot., sér. 5, 15$_*XM*Z and A. anomalus (Blastocladiomycota). 1872; lectotype designated by Dick 2001: 373).

Notes: Ultrastructure of the host-parasite interface between Description$OQ R. allomycetis and Allomyces anomalus has been studied " (Powell et al. 2017), as has been ultrastructure of early \ infection stages of R. allomycetis in A. macrogynus (Powell through a small pore resulting from the dissolution of the & Letcher, unpubl.). Foust’s epithet “allomycis” was corrected papilla; resting spore formed in the sporangium of the host, to “allomycetis” by Doweld (2014), as the correct genitive spherical, somewhat thick-walled, brownish (?), covered with case of the host genus Allomyces. Sparrow (1960) noted very short tenuous spines” (Sparrow 1960). that Rozella allomycetes was “A name unaccompanied by a description. Probably referable to Rozella allomycis Foust”. Host: Apodachlya brachynema (Oomycota).

VOLUME 9 · NO. 2 391 Letcher and Powell

Rozella blastocladiae (Minden) Sparrow, Mycologia Description$OQ*XMZX 30: 377 (1938). 3 % *XM*_ 3 & *_M@_ 3 Basionym: Pleolpidium blastocladiae Minden, Krypt. Fl. Mark %# Brandenb. 5 (3): 253 (1911) ["1915"]. " % @@M_ 3 & *+M 3 \ *Z 3

ARTICLE Type: Petersen (Bot. Tidsskrift 29Z$ZZ”M*/X/ # % neotype (as "lectotype") designated by Dick 2001: 374). few seconds. Resting spore faintly yellow, oval, spherical, +M3% Description: “Sporangium assuming the shape of the *M*+ 3 hypertrophied host sporangium, which becomes somewhat *_M@3" broader and more ovoid than normal, with an apical pore, "P–*/Z " " % # { Hosts: Nowakowskiella profusum, N. elegans, N. ramosum, #% Cladochytrium replicatum, C. crassum, and C. hyalinum germination not observed” (Sparrow 1960). (Chytridiomycota).

Hosts: Blastocladia pringsheimii, and B. gracilis (Blastocladi- Note: The original description (Karling 1941) lacked illustra- omycota). later work (Karling 1942) by the same author. Rozella canterae Sparrow, Aquatic Phycom., 2ndedn: 172 (1960). Rozella coleochaetes Sparrow, Papers Mich. Acad. Sci. Arts Letters 50: 118 (1965); as “coleochaetis”. Type: Canter (Trans. Brit. Mycol. Soc. 33$@_W@M Synonym: Plasmophagus coleochaetisQ†'N /ZM_*/_XNXX*$@W_ Straminip. Fungi: 451 (2001).

Description: “Sporangium assuming the shape of the Type: Sparrow (Papers Mich. Acad. Sci. Arts Letters 50: 116, UMV*/”_lectotype designated here, MBT 384207). "%% \ Description$ OQ is dehisced; resting spore somewhat ovoid, with a thick wall, @_{@+3ZXMZ #P &@@MZX3%__&*_M_3 (Canter 1950, Sparrow 1960). *X 3 ‚ ”MW&M_3 Host: Chytridium oedogonii (Chytridiomycota). motile within the sporangium, escaping through a sessile pore upon deliquescence of the papilla. Resting spore not Rozella chytriomycetis Karling, Mycologia 38: 107 observed” (Sparrow et al. 1965). (1946); as “chytriomycii”. Host: Coleochaete (Charophyta). Type: Karling (Mycologia 38$*XZ/M*/*/Z” designated by Dick 2001: 375). Note: Blackwell et al X*” ‘ # R. coleochaetes from the genus Plasmophagus. Description$OQ " Rozella cuculus (E. J. Butler) Sparrow, Mycologia 30: *XMZX 3 # G " 377 (1938). % @ 3 & *_ 3 Basionym: Pleolpidium cuculus E. J. Butler, Mem. Dept. X_MXW3% Agric. India, Bot. Ser. 1: 125 (1907). before emerging; darting about rapidly in swimming, rarely becoming amoeboid. Resting spores partly or almost Type: Butler (Mem. Dept. Agric. India, Bot. Ser. 1: plate VII, %WMX3 M_*/XWNXX*$@WZ large central vacuole, usually coarsely granular cytoplasm; wall dark brown, rarely smooth, usually spiny or echinulate; Description: “Sporangium spherical, subspherical, or germination unknown” (Karling 1946). pyriform, formed in the sporangium of the host or in */MZ Host: Chytriomyces hyalinus (Chytridiomycota). 3"% % % \ Rozella cladochytrii Karling, Torreya 41: 105 (1941). end; resting spore spherical, single, free in the sporangium *M*+3 Type: Karling (Amer. J. Bot. 29$ ” *MZ */Z a smooth pale-yellow somewhat thickened wall, germination lectotype designated here, MBT 384206). not observed” (Sparrow 1960).

392 IMA FUNGUS Taxonomy of Rozella (Cryptomycota)

Hosts: Pythium intermedium, and P. monospermum (Oomy- Hosts: Pythium (?) vexans, and Pythium sp. (Oomycota). ARTICLE cota). Note: Karling (1981) observed a species he tentatively Note: Sparrow (1960) putatively included Pleolpidium R. irregularis in Pythium debaryanum, and on tuberculorum Vuillemin (1909) and Chytridium simulans the basis of his observations and those of others (Butler Dangeard (1896-97) as referable to R. cuculus. */XW Q Š Q */Z+ # the species. Rozella diplophlyctidis Karling, Nova Hedwigia 45: 529 (1987). Rozella itersoniliae D. J. S. Barr & Bandoni, Mycologia 71: 1261 (1979). Type: Karling (Nova Hedwigia 45$ _@Z *M” */+W M Synonym: Pleotrachelus itersoniliae (D. J. S. Barr & Bandoni) holotype). †'NStramin. Fungi: 453 (2001).

Description$OQ Type: Barr & Bandoni (Mycologia 71$*”*MZ*/W/ *+M+3G*MZ# M ‚%*+M_3 with a small hyaline refractive globule. Resting spores partly Description$ OQ *”M3 ‚ XM_ 3 Germination unknown” (Karling 1987). \ \ **M*@ 3  spores not seen” (Barr & Bandoni 1979). Host: Diplophlyctis intestina (Chytridiomycota). Host: Itersonilia perplexans (Basidiomycota). Rozella endochytrii Karling, Torreya 41: 106 (1941). Rozella laevis –#^% Type: Karling (Amer. J. Bot. 29$@X_M@_*/Zneotype MycoBank MB828495 designated here, MBT 384208). Synonym: Rozella laevis Karling, Mycologia 34: 201 (1942); nom. inval. (Arts. 39.1, 40.1). Description$ OQ *_M XX 3 % Type: Karling (Mycologia 36$”Z*M*/*/ZZM " %#M”3 Description: “Sporangium solitary, partly or completely ‚%@ZMZ3&*_3 % with optically denser apical and basal regions which give them "XM_3%*XMX3& a characteristic appearance; swirling in the sporangium before @XM**3*@# dehiscence; emerging in a stream and becoming actively motile @MZ3M@3‚ in a few seconds. Resting spores unknown” (Karling 1942a). hyaline, with a globular spot that is not markedly refractive, %*_M*+3&/M@@3G\ Host: Endochytrium operculatum (Chytridiomycota). *XM*3**M*+3% elongate or obpyriform with a large central vacuole and Note: The original description (Karling 1941) lacked *_M 3P–*/Z from a later work (Karling 1942) by the same author, of the original material. Hosts: Pythium gracile, and Pythium sp. (Oomycota).

Rozella irregularis (E. J. Butler) Sparrow, Mycologia Note: The original description (Karling 1942) lacked illustra- 30: 377 (1938). Synonym: Pleolpidium irregulare E. J. Butler, Mem. Dept. work (Karling 1944) by the same author. Agric. India, Bot. Ser. 1: 123 (1907). Rozella longicollis Karling, Sydowia 19: 218 (1965). Type: Butler (Mem. Dept. Agric. India, Bot. Ser. 1$š„„„ *M**/XWNXX*$@WZ Type: Karling (Sydowia 19$ ^š„ *M/ */”_ lectotype designated by Dick 2001: 375). Description: “Sporangia formed in the hyphae of the host, %@3 Description$ OQ "% XM”X3%*_M*+&M a single cilium borne posteriorly, [resting] spores single, free @_3*M_%#WM*_3 %{ @M@W3‘ **M*_ 3 ‚*_M&_M@ yellow color, with a moderately thick wall, provided with short 3 regular spines, germination not observed” (Butler 1907). \*M*Z3

VOLUME 9 · NO. 2 393 Letcher and Powell

 " % {%*M*W3 brown, the thickened wall densely covered with tenuous diameter, wall punctate to spiny, rarely smooth; germination spines, in intercalary or lateral swellings of the host hyphae, unknown” (Karling 1965). germination not observed” (Sparrow 1960).

ARTICLE Host: Pythium sp. (Oomycota). Hosts: Monoblepharis polymorpha, and M. macrandra (Monoblepharidomycota). Rozella longisporangia'ŠVNova Hedwigia 37: 378 (1983). Rozella multimorpha Letcher & Longcore, J. Euk. Microbiol. 64: 65: 183 (2018). Type$ ' Š  Nova Hedwigia 37$ @WW * 1983; lectotype designated here, MBT 384209). Type: Letcher et al. (J. Euk. Microbiol. 65$*+*X*+ M Description$ OQ *_MZ 3 _XMW3& Description: Sporangia are monosporangiate, spherical, *XM* 3 # *_MX 3 _X 3 _@3&_@MW3*WX pyriform, or irregularly-shaped, with (typically) one to & _@MW 3 Q *MZ @+M ž 3 # _MZX 3 ”+3‚~ the discharge tube terminating with a constriction and ‚ {‚*M*3 ‚ # *”M*+ 3 { \*X”M**Z3% {# @MZ3 single mitochondrion, a pair of lipid globules, a microbody, %P'Š*/+@ ‚ sporangium prior to release, and are released as a mass Host: Pythium oligandrum (Oomycota). of motile spores that rapidly disperse. Resting spores not observed (Letcher et al. 2018). Rozella marina (Sparrow) Sparrow, Mycologia 30: 377 (1938). Host: Pythium catenulatum (Oomycota). Basionym: Pleolpidium marinum Sparrow, Biol. Bull. Mar. Biol. Lab. Woods Hole 70: 256 (1936). Rozella parva Canter, Bot. J. Linn. Soc. 62: 275 (1969).

Type: Sparrow (Biol. Bull. Mar. Biol. Lab. Woods Hole 70: Type: Canter (Bot. J. Linn. Soc. 62: 275, type material IMI @M@@*/@”lectotype designated here, MBT 384211). *@*”W@M

Description$ OQ Description: “Sporangium solitary assuming the shape of the @XMZ_3 unhypertrophied host sporangium or young resting spore. " Q ZM/_ 3 "@33 GR*" \ emerging singly on dehiscence and immediately swimming spore not observed” (Sparrow 1960). ‚ _ 3 *+ & @_3' Host: Algochytrops polysiphoniae (Chytridiomycota). % % \*_3 Note: In addition to Sparrow’s observations (Sparrow 1936b, _M*X”3 1938), R. marina was observed in the same host from Iceland refractive wall, on the outside of which may be deposited lumps (Johnson 1966). of solid material, or an apparently thin corrugated undulate #' Rozella monoblepharidis Cornu, Ann. Sciences Nat., and many small refractive globules” (Canter 1969). Bot. sér. 5, 15: 150 (1872). Synonyms: Rozella monoblepharidis-polymorphae Cornu Host: (Chytridiomycota). Ann. Sciences Nat., Bot. sér. 5, 15: 150 (1872). Pleolpidium monoblepharidis-polymorphae (Cornu). A. Fischer, Rozella polyphagi (Sparrow) Sparrow, Mycologia 30: Rabenh. Krypt.-Fl. 1 (4): 44 (1892). 377 (1938). Basionym: Pleolpidium polyphagi Sparrow, Trans. Brit. Mycol. Type: Cornu (Ann. Sci. Nat., Bot. sér. 5, 15$Z*@M*+ Soc. 18: 215 (1933). 1872; lectotype designated by Dick 2001: 373). Type: Sparrow (J. Linn. Soc., Bot. 50$ *Z */MX 1936; neotype designated by Dick 2001: 374, as “lectotype”, Description: “Sporangia formed in intercalary swollen parts drawn from the same material used in Sparrow 1933). of the hyphae, ovoid … with a single small lateral discharge

394 IMA FUNGUS Taxonomy of Rozella (Cryptomycota)

Description$OQ Host: Araiospora spinosa (Oomycota). ARTICLE XMZ+ 3# Rozella rhizoclosmatii Letcher & Longcore, Fungal ZM+ 3 Biol. 121: 5 (2017). \ % " {@ 3 *_M 3 Type: Letcher et al. (Fungal Biol. 121$ Z V X*W M discharged; resting spore not observed (Sparrow 1960). holotype).

Host: Polyphagus laevis (Chytridiomycota). Description$ OQ G " Note: Ultrastructure of the thallus of R. polyphagi has been Z_ 3 % % studied (Powell 1984). *_3%‚ motile in the sporangium before they release through a single Rozella pseudomorpha (Scherff.) Sparrow, Aquatic #‚%*M*@3&*+M Phycom.: 124 (1943). 3 \ *XM* 3 Synonym: Olpidium (?) pseudomorphum Scherff., Arch. ‚ % Rozella{ \ */W_ Protistenk. 54: 510 (1926). plus a lattice of perpendicular rods surrounding the nucleus. Resting spore not observed” (Letcher et al. 2017). Type: Scherffel (Arch. Protistenk. 54$+*M_*/” lectotype designated here, MBT 384214). Host: Rhizoclosmatium globosum (Chytridiomycota).

Description$ OQ %% Rozella rhizophlyctidis Karling, Amer. J. Bot. 29: 32 " (1942); as “rhizophlyctii”. " ellipsoidal, ovoid, or plump and rod-like, somewhat arched, Type: Karling (Amer. J. Bot. 29$ @X @”MZW */Z % % \ lectotype designated by Dick 2001: 375). long, trailing, attached at the concave side of the body, " % Description$OQ R"XM**X3% amoeboid change in shape, movement hopping; resting *Z#‘ spore unknown” (Sparrow 1960). G " _M@3&*_M3 Host: Lagenidium rabenhorstii (Oomycota). the anterior end, with a minute globule near the posterior end; \G\*”M*+3 Note$ Q */”X Ÿ in the sporangium before emerging; darting about rapidly in %% Ÿ ' R. pseudomorpha swimming, occasionally becoming amoeboid. Resting spores \"Ÿ % *ZM*+ 3 with a large central vacuole and coarsely granular cytoplasm; Rozella rhipidii Cornu, Ann. Sci. Nat, Bot., sér. 5, 15: *+ 3 *_M 3 153 (1872). " Synonyms: Rozella rhipidii-spinosi Cornu, Ann. Sci. Nat, Bot., "P–*/Z sér. 5, 15: 153 (1872). Pleolpidium rhipidii-spinosi (Cornu) A. Fischer, Rabenh. Hosts: Rhizophlyctis petersenii, and R. rosea (Chytridiomy- Krypt.-Fl. 1 (4): 44 (1892). cota). Pleolpidium araiosporae (Cornu) Minden, Krypt.-Fl. Mark Brandenb. 5: 252 (1911) ["1915"]. Rozella rhizophydii Karling, Mycologia 36: 645 (1944).

Type: Cornu (Ann. Sci. Nat, Bot., sér. 5, 15$*_@_ Type: Karling (Mycologia 36$”Z_XM+*/ZZ *M/*+WNXX*$@W@ designated by Dick 2001: 375).

Description$ OQ Description$ OQ swollen and obpyriform usually smooth sporangium of the R"*_M@X "% 3%*XM*3&*@MX3*M*_3& shape, reniform, spherical or ellipsoidal, with a long posterior *”M_ 3 *M@ # G" \ % M_ 3 & @MZ 3 \ *M*Z 3 spherical, yellowish brown or reddish, with dense contents, Resting spores unknown” (Karling 1944). wall slightly thickened, covered with tenuous spines, germination not observed, predominantly formed in the spiny Host: Rhizophydium globosum (Chytridiomycota). sporangia of the host” (Sparrow 1960).

VOLUME 9 · NO. 2 395 Letcher and Powell

Rozella septigena Cornu, Ann. Sci. Nat., Bot., sér. 5, Hosts: Achyla, Saprolegnia (Oomycota). 15: 163 (1872). Synonym: Skirgiellia septigena (Cornu) A. Batko, Acta Mycol. Note: This species was designated as the type of the genus by 13:322 (1977); nom. Ilegit. (Arts. 52.1, 52.2). Clements & Shear 1931: 234). “Cornu’s name was applied, in error, by Fischer (1882: 365) to a similar-appearing parasite in

ARTICLE Type: Cornu (Ann. Sci. Nat., Bot. sér_$*_$”*{*W Saprolegnia\"PQ*/”X 1872). Doubtful and excluded species Description: “Sporangia possibly formed by successive fractionation of one thallus, in transversely or obliquely Rozella barrettii Karling, Mycologia 34: 202 (1942). walled-off segments of the sometimes slightly swollen host “Based on an incompletely known form described by Barrett % */@Z$**@+Q\" " known, the fungus cannot be placed generically” (Sparrow \ 1960: 180). spherical, with a slightly thickened wall covered with short tenuous spines, brownish, with dense contents, formed in Rozella maxima Karling, Amer. J. Bot. 29: 24 (1942); spherically swollen short lateral branches of the hyphae, as “maximum”. # by a cross wall, germination not observed” (Sparrow 1960). Apparently a typographical error for R. marina (q.v.).

KEY TO THE SPECIES OF ROZELLA

1 Thallus polysporangiate ...... 2 Thallus monosporangiate ...... 4

* \$Oomycota ...... 3 \$Blastocladiomycota: Allomyces ...... allomycetis

@ \$Saprolegnia………………………………………………………..………. ……………………… ...... septigena \$Achlya and Dictyuchus ...... achlyae

Z* \$Blastocladiomycota: Blastocladia ...... blastocladiae \$Monoblepharidomycota: Monoblepharidis ...... monoblepharidis \$Basidiomycota: Itersonilia ...... itersoniliae \$Coleochaetophyta: Coleochaete ...... coleochaetes \$Oomycota ...... 5 \$Chytridiomycota ...... 11

_Z \$Araiospora ...... rhipidii \$Apodachlya ...... apodyae \$Pythium ...... 6 \$Lagenidium ...... pseudomorpha

6 (5) Resting spore present ………………………………………………………………………………………… ...... 7 Resting spore absent ...... 10

7 (6) Resting spore smooth ...... 8 Resting spore spiny ...... 9

+W \$P. gracile; resting spore hyaline ...... laevis \$P. intermedium, P. monospermum; resting spore pale yellow ...... cuculus

/W \$P. vexans; resting spore pale yellow ...... irregularis \$Pythium sp., resting spore dark brown ...... longicollis

*X” \$P. oligandrum ...... longisporangia \$P. catenulatum ...... multimorpha

396 IMA FUNGUS Taxonomy of Rozella (Cryptomycota)

**Z \$Diplophlyctis ...... diplophlyctidis ARTICLE \$Rhizophlyctis ...... rhizophlyctidis \$Chytriomyces ...... chytriomycetis \$Polyphagus ...... polyphagi \$Endochytrium ...... endochytrii \$Cladochytrium, Nowakowskiella ...... cladochytrii \$Rhizophydium ...... rhizophydii \$Zygorhizidium ...... parva \$Rhizoclosmatium ...... rhizoclosmatii \$Chytridium ...... canterae \$Algochytrops ...... marina

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