Mycol. Res. 103 (4) : 459–467 (1999) 459 Printed in the United Kingdom New and poorly known smut fungi in Cuba* M E I K E P I E P E N B R I NG UniversitaX t TuX bingen, Botanisches Institut, Lehrstuhl Spezielle Botanik}Mykologie, Auf der Morgenstelle 1, 72076 TuX bingen, Germany Recent field work on smut fungi in Cuba yielded one new species and, among many new records, several poorly known species which are described and illustrated. The new species, Cintractia cubensis on Rhynchospora microcephala, is characterized by rather large, dark, and finely foveolate teliospores which can germinate with basidia of only two basidial cells. The poorly known C. samanensis was found on the new host Rhynchospora fascicularis ssp. fascicularis. Entyloma guaraniticum on Bidens pilosa differs from E. bidentis on the same host by thick-walled teliospores embedded in the mesophyll of leaves deforming them. Spores of E. bidentis are smaller and do not deform the leaf. Sphacelotheca everhartii on Andropogon virginicus, S. microspora on Paspalum notatum, and S. panici-leucophaei on Trichachne insularis are transferred into Sporisorium. About 24 species of smut fungi are known for Cuba (Whetzel & Kern, 1926 ; Ling, 1951 ; Zundel, 1953 ; Seidel, 1976 ; Arnold, 1986). Given the high diversity of vascular plants on this largest island of the Antilles (main island ca 105 000 km#) and considering that 54 species of smut fungi are known for the smaller Costa Rica (ca 51 000 km# ; Piepenbring, 1996 and 1996 a, b), Cuba certainly harbours many more species waiting for their discovery. In 1996 I investigated smut fungi in Cuba, resulting in 16 new records for the country. The collected material permits several taxonomic insights which are presented here. MATERIALS AND METHODS Material was collected in Cuba in Aug.–Oct. 1996. The host plants were determined in the herbarium of the Jardı! n Bota! nico Nacional de Cuba and according to the flora of Cuba (Leo! n, 1946). Voucher specimens are deposited in the herbarium of the Jardı! n Bota! nico Nacional de Cuba, La Habana (HAJB), in the Botanische Staatssammlung Mu$ nchen (M), Germany, and my private herbarium (H. U. P.). ‘ MP ’ numbers are my collection numbers. For the determination of the smut fungi, herbarium dried spores were measured by light microscopy (LM) in lactophenol after heating. The values of the size ranges are the means of 50 spores for the new species and at least 20 for previously described species³1 .. Extreme values are given in brackets. Spore measurements include ornamentation and hyaline sheaths. For SEM, dried spores were dusted onto doublesided adhesive tape, fixed on specimen stubs, and sputter coated with gold-palladium, ca 20 nm. The spores were viewed with a Cambridge Stereoscan 250 MK 2. * Part 151 in the series ‘ Studies in Heterobasidiomycetes ’ from the Botanical Institute, University of Tu$ bingen. TAXONOMY Cintractia cubensis M. Piepenbr., sp. nov. (Figs 1–5) Etym. : Being found in Cuba with the help of Cuban people C. cubensis is dedicated to this fascinating island and its inhabitants. Sori in totis spiculis plantae, spiculorum sterilium rhachillae apices circumdantes, 1¬1–2 mm lati. Sporarum massa pulveracea, obscura, sine peridio. Sporae singulares vel laxe connectae, irregulariter angulatae, complanatae, 12–16(–19)¬(15–)16–19(–20) µm, obscure rubellobrunneae, exosporio viso per microscopium electronicum SEM foveolis numerosis, tenuibus, profundis. Sporae modo generis Ustilago germinant. Holotypus in M ; isotypes in HAJB and H.U.P. Type on Rhynchospora microcephala Britton (Cyperaceae), Cuba, Prov. Pinar del Rı! o, municipio Sandino, area Herradura, on a floating mat in the Laguna de la Culebra, alt. ca 10 m, 19 Oct. 1996, C. Landeta, A. Urquiola, & M. Piepenbring (MP 2252). Sori in all the spikelets of an inflorescence (Fig. 2), young hidden between the glumes, later spores protrude at the top of the spikelets. The glumes of infected spikelets are greener, shorter, appear more shredded, and in denser heads than those of healthy spikelets (compare Fig. 1). Sori globose, ca 1¬1–2 mm, around the tips of the rachillae of sterile spikelets, without peridia or sterile stroma. Spore mass powdery, dark, sometimes mixed with few hyaline cells (Fig. 4). Spores (Figs 3, 5) single or adhering in loose groups, irregularly shaped, angular, sometimes flattened, 12–16(–19)¬(15–)16–19(–20) µm, dark reddish-brown, without hyaline appendages. Teliospore wall 1–2 µm thick, thicker in the angles (this may be difficult to see because of the dark colour of the wall), spore surface granular seen by LM, very densely deeply foveolate seen by SEM (Fig. 5). Germination of teliospores (Figs 6–15) of Ustilago type, basidia with two Cuban smut fungi 460 5 1 2 2 mm Fig. 5. Cintractia cubensis. Teliospore seen by SEM (scale bar ¯ 4 µm) (MP 2252). 16 2 mm 3 4 5 lm 5 lm Figs 1–4. Cintractia cubensis. Fig. 1. Apical part of a healthy inflorescence of Rhynchospora microcephala. Fig. 2. Apical part of an inflorescence infected by C. cubensis. Fig. 3. Teliospores. Fig. 4. Light coloured cells. (MP 2252). basidial cells (Figs 7–9, 13) or with four cells which immediately conjugate (Figs 10–11) and develop large basidiospores. Cintractia cubensis differs from other species of Cintractia by rather large, dark, deeply finely foveolate teliospores without appendages, and sometimes two-celled basidia which are reported here for the first time for a species of Cintractia. The spores of C. major (Desm.) Liro (ca 11–13¬14–17 µm) and C. montagnei (Tul. & C. Tul.) Magnus (ca 11–12¬12–14 µm) are smaller and lighter coloured, those of C. gigantospora Liro (ca 17–19¬19–21 µm) are larger and lighter coloured. Cintractia cubensis does not develop its spores in pockets of a sterile stroma underneath a peridium like the type species Cintractia axicola (Berk.) Cornu (Cornu, 1883), but belongs to 17 Figs 16–17. Cintractia samanensis. Teliospores seen by SEM (bars ¯ 4 µm). Fig. 16. Type. Fig. 17. MP 2103. M. Piepenbring 461 6 7 8 9 10 5 lm 5 lm 13 14 15 11 12 Figs 6–15. Cintractia cubensis. Germinated teliospores after 1–2 d on water agar (MP 2252). the group of species around C. montagnei. Species of this group have sori around the tips of rachillae of sterile spikelets without peridia or sterile stroma, dark teliospore masses, flattened, foveolate teliospores, and Ustilago basidia sometimes with copulating basidial cells (as in C. axicola ; Piepenbring, 1996 : 31). Each pair of conjugated basidial cells produces one large dikaryotic basidiospore. In C. cubensis, a morphologically similar basidiospore can be formed by one basidial cell of a two-celled basidium (Figs 8, 13). These two-celled basidia seem to omit the second division of basidial cells and the conjugation. Cintractia samanensis Cif., Arkiv foX r Botanik 23 : 13 (1931) (Figs 16–21) Type on Rhynchospora oligantha A. Gray, Dominican Republic, Prov. Samana! , Cordillera Central, Sabana de la Mar, at El Valle, 11 July 1930, E. L. Ekman, N. H. 15652 (Herb. Ciferri n. 3549, S). Sori (not illustrated) in spikelets in groups or in all the spikelets of one inflorescence. Sori hidden by stunted curved glumes, glumes with sori greener than in healthy parts of the host, around the tips of rachillae of sterile spikelets, globose, ca 1¬1–2 mm, without peridia or sterile stroma. Spore mass powdery in the upper part, agglutinated at the base of the sorus, dark. Spores (Figs 16–17) single, subglobose or flattened, subangular, in surface view 9–11(–12)¬10–12(–14) µm, in profile 6–8 µm, light olive reddish-brown, rarely with a globose hyaline appendage corresponding to the rest of the hyaline sheath on a flattened side of a spore. Wall ca 1 µm thick, thicker and darker in the edges, inconspicuously pitted as seen by LM, finely foveolate with a granular surface as seen by SEM. Germination (Figs 18–21) of the Ustilago type, with mostly four-celled basidia forming basidiospores, sometimes basidia with more than four basidial cells (Fig. 20). Hitherto, C. samanensis has been known only from the Dominican Republic on R. oligantha (Cyperaceae). In Cuba, it Cuban smut fungi 462 18 20 21 5 lm (11–)13–17¬(13–)15–19(–24) µm, fresh hyaline, dried often yellowish-brown. Wall two-layered, the inner layer ca 0±7 µm, the outer (sheath) of irregular thickness 2–6 µm, often ca 4 µm thick, smooth. Germination (Figs 24–26) of the Tilletia type with immediate conjugation of the sessile basidiospores and development of sickle-shaped and filiform conidia. Known on Bidens pilosa L. (B. leucantha Willd.) (Asteraceae), from the U.S.A. (Florida), Paraguay and Taiwan. This is a new record for Cuba. Other specimens examined : On Bidens pilosa, Cuba, Prov. Cienfuegos, Atkinson Botanical Garden, alt. ca 70 m, 18 Aug. 1996, M. Piepenbring (MP 2143). Prov. Ciudad de La Habana, Jardı! n Bota! nico Nacional, alt. ca 75 m, 9}10 Sep. 1996, M. Camino & M. Piepenbring (MP 2174). And other collections. 19 Figs 18–21. Cintractia samanensis. Germinated teliospores after 1 d on water agar (MP 2106). was found as a new record on the new host R. fascicularis (Michaux) Vahl ssp. fascicularis. Other specimens examined : On R. fascicularis ssp. fascicularis, Cuba, Prov. Pinar del Rı! o, municipio Sandino, close to the protected area of San Ubaldo, white sands, Laguna del Toro, alt. ca 5 m, 14 Aug. 1996, J. Gutierrez, A. Urquinola & M. Piepenbring (MP 2106). Prov. Pinar del Rı! o, municipio Sandino, protected area of San Ubaldo, white sands, alt. ca 5 m, 14 Aug. 1996, J. Gutierrez, A. Urquinola & M. Piepenbring (MP 2114). On R. cf. fascicularis, Prov. Pinar del Rı! o, municipio Sandino, protected area of San Ubaldo, white sands, alt. ca 5 m, 14 Aug. 1996, J. Gutierrez, A. Urquiola & M. Piepenbring (MP 2103). Ling (1950 : 506) considers the name C. samanensis a synonym of C. montagnei but the spores of C. samanensis are smaller, more densely finely pitted, and rarely carry hyaline appendages. Entyloma guaraniticum Speg., Anales de la Sociedad CientıU fica Argentina 17 : 127 (1884) (Figs 22–26) Type on Bidens pilosa L., Paraguay, close to Guarapı! , Dec. 1882, B. Balansa 3731 (in herb. LPS 3361). Sori (Fig. 22) as leaf spots, circular, prominently convex pustular towards the abaxial (rarely adaxial) side of the leaf, heavy infection deforming the leaf. Sori mostly 1–1±5 mm diam., sometimes confluent, when fresh, yellow-greenish, when old, brown in the centre, when dry, ochre to brown. Spores (Fig. 23) densely packed between the host cells, globose, often subangular and irregularly shaped, Infection by Entyloma guaraniticum is macroscopically different from that of E. bidentis Henn. (Hennings, 1895) by smaller, usually more numerous spots which are pustular and deform the leaves while spots of E. bidentis usually do not protrude from the leaf surface. The spores of E. bidentis are smaller [(10–)11–13¬(12–)13–15(–18) µm] with a more regularly and less thickened (up to 3 µm) outer spore wall layer (sheath ; Fig. 27). The thick outer wall layer of the spores of E. guaraniticum probably causes the swelling of the leaf spots. Spore sizes given for E. guaraniticum in literature are very variable (Spegazzini, 1884 : 10–14 µm ; Clinton, 1906 : 11–20 µm ; Savile, 1947 : 8±5–15±0¬9±8–18±3 µm ; Ling, 1953 : 10±5–13±5 or 8–11¬12–18±5 µm). The spores of the type studied by me, however, are as large as those collected in Cuba. The spores of E. spegazzinii Sacc. & Syd. in Saccardo (1902) (3 E. bidentis Speg., later homonym of E. bidentis Henn.) on Bidens bipinnata L. are even larger than those of E. guaraniticum according to Spegazzini (1899) (16–20¬20–25 µm). Type material of E. spegazzinii could not be located. A specimen of E. spegazzinii on B. sub-alternans DC. [La Plata, Dec. 1936(?), Lindquist, LPS 2971] studied by me has spores measuring 11–16(–19)¬(13–)14–20(–23) µm, being as large as those of E. guaraniticum. Further studies are necessary to decide whether E. spegazzinii is a synonym of E. guaraniticum. Sporisorium everhartii (Ellis & Galloway) M. Piepenbr., comb. nov. (Figs 28–35) Basionym : Sorosporium everhartii Ellis & Galloway, Journal of Mycology 6 : 32 (1890). Type on Andropogon virginicus L., U.S.A., N. J., Newfield, Oct. (N.A.F., 2265 b). Synonym : Tolyposporium everhartii (Ellis & Galloway) Dietel, in Die NatuX rlichen Pflanzenfamilien. I (ed. Engler & Prantl) : 14 (1897). (For further synonyms see Zundel, 1953 : 59.) Sori (Fig. 28) in all the spikelets of one or several pairs of racemes in an inflorescence, other parts of the inflorescence healthy. Sori usually formed by ovaries of sessile spikelets (compare Hansing & Lefebvre, 1941), narrow cylindrical, with cream coloured peridium of sterile fungal cells covered by few host tissue cells. The peridium ruptures exposing the spore M. Piepenbring 463 22 24 25 5 lm 1 cm 23 27 26 5 lm 5 lm Figs 22–26. Entyloma guaraniticum. Fig. 22. Leaf of Bidens pilosa deformed by pustular spots (MP 2174). Fig. 23. Teliospores (type). Figs 24–26. Germinated teliospores after 2–3 d on water agar (MP 2281). Fig. 27. Teliospores of Entyloma bidentis (on Bidens pilosa, Costa Rica, Prov. Alajuela, 2 km west of San Ramo! n, 8 Nov. 1992, M. Piepenbring, MP 616). mass and a, when dry, twisted, when wet, extended, flat columella of axial host tissue, 1–2±5 cm long, with adhering spore balls. Mass of spore balls powdery, dark. Spore balls (Figs 29–31) permanent, of irregular shape, globose to angular, 40–70¬70–90 µm (Ellis & Galloway, 1890 : 32 : up to 120 µm), composed of central light coloured cells and an outer layer of spores (Fig. 31). Spores 8–10¬10–12 µm, dark reddish brown. Spore wall of medium thickness warty only on the spore ball surface, as seen by LM, large warts and interspersed small warts are evident only on the spore ball surface, as seen by SEM. Light coloured cells (Fig. 31) probably sterile (no germination was observed), polyangular, hyaline, smooth. Germination (Figs 32–35) of the Ustilago type, with up to 110 µm long, curved basidia whose 4–7 basidial cells produce 5–15 µm long basidiospores. Known on many different species of Andropogon (Poaceae), from the U.S.A. and Mexico, and on Gayana densiflora (name not found in the Index Kewensis) and Hyparrhenia ruprechtii Fourn. from Africa (Zundel, 1953). This is a new record of this species for Cuba. Specimens examined : On Andropogon virginicus, Cuba, Prov. Pinar del Rı! o, municipio Sandino, area Herradura, white sands close to the Laguna de la Culebra, alt. ca 10 m, 19 Oct. 1996, C. Landeta, A. Urquiola, & M. Piepenbring (MP 2244). Prov. Pinar del Rı! o, municipio Pinar del Rı! o, S Las Ovas, Reserva Natural El Punto, alt. ca 30 m, 30 Oct. 1996, A. Urquiola, & M. Pipenbring (MP 2270). Further specimen examined for comparison : On Andropogon scoparius (?), U.S.A., Alabama, Lee Co., Auburn, 14 Nov. 1897, F. S. Earle (sub Sorosporium everhartii in Sydow, Ustilagineen no. 190, in herb. M). The determination of this species is based on the host species which is similar to the host species of the type specimen, the presence of permanent spore balls, and a germination similar to the one reported for this species (as Sorosporium everhartii ) by Dura! n (1987). Sorosporium Rudolphi seems to be restricted to the type Cuban smut fungi 464 29 28 2 mm Fig. 28. Sporisorium everhartii. Two sessile spikelets with sori (one sorus only partly shown). Note the twisted stripes of the peridia and the dark columellae bearing spore balls 2MP 2244). 30 species, S. saponariae Rudolphi on Caryophyllaceae, with spore balls developing on the surface of floral organs of the host (Langdon & Fullerton, 1975 ; Va! nky, 1987 ; personal observations). Smut species in Sporisorium have sori with peridia and columellae on Poaceae (Langdon & Fullerton, 1978). Their teliospores are single, in loose groups, or form permanent teliospore balls. Sporisorium everhartii has to be included in Sporisorium because of its sori with peridia and columellae and because of its teliospores in permanent balls. Balls with inner smooth, hyaline cells and outer ornamented germinating teliospores are known from other species of Sporisorium, for example S. heteropogonicola (Mundk. & Thirum.) Va! nky, in Shivas & Va! nky (1997). 31 Sporisorium microsporum (J. Schro$ t. & Henn.) M. Piepenbr., comb. nov. (Figs 36–41) Basionym : Ustilago microspora J. Schro$ t. & Henn., in Hennings, Hedwigia 35 : 215 (1896). Type on Paspalum sp., Brazil, St Catharina, Itajahy, Nov. 1885, E. Ule (1621). Synonyms : Sphacelotheca microspora (J. Schro$ t. & Henn.) Cif., in Ciferri & Herter, Botanisches Archiv 34 : 532 (1932). Ustilago paspali-notati Henn., in Herb. Holway (see Clinton, 1902 : 140). Sphacelotheca paspali-notati (Henn.) G. P. Clinton, Journal of Mycology 8 : 140 (1902). Type on Paspalum notatum, Mexico, Guadalajara, 15 Sep. 1899 (in herb. Holway, BPI 165251). Sori (Figs 36–37) in one host plant as two linear bodies corresponding to the two racemes of the inflorescence, initially enclosed by peridia of host tissue (sometimes with remains of glumes attached) and few sterile fungal cells, columellae formed by axial tissue of the racemes. Spore mass powdery, with few sterile cells, dark brown. Spores (Figs 38–39) single, globose to subglobose, often polyhedral, 7–9¬8–9(–10) µm, light golden brown to reddish-brown. Figs 29–31. Sporisorium everhartii. Spore balls seen by SEM. (bars ¯ 10 µm). Fig. 29. One spore ball (MP 2270). Fig. 30. Part of a spore ball (MP 2270). Fig. 31. Broken spore ball showing inner smooth cells (Sydow, Ust. no. 190). M. Piepenbring 465 (ii) Sporisorium microsporum (as Sphacelotheca microspora) on Paspalum spp. : sori in the inflorescence ; teliospores 7–10 µm, minutely verruculose ; (iii) Sphacelotheca panici-leucophaei on Trichachne insularis and Trichachne californica (Benth.) Chase : sori with numerous columellae ; teliospores 6–8 µm, smooth to very minutely granular. These characteristics are confirmed for Sporisorium microsporum (see above) and for Sphacelotheca panici-leucophaei which is cited for Cuba by Whetzel & Kern (1926) and Zundel (1953) and for which material from the Dominican Republic was available for study. These species have to be classified in Sporisorium because of the presence of columellae of host tissue in the centre of the sori and peridia of more or less persistent host tissue with at least some sterile fungal cells on the inner side of the peridium (Langdon & Fullerton, 1978). Based on these characteristics species of Sporisorium are distinguished from those classified in Ustilago. While species of Sporisorium and most species of Ustilago are found on Poaceae, Sphacelotheca seems to be restricted to smuts on Polygonaceae with columellae formed mainly by sterile fungal cells (Langdon & Fullerton, 1978). 32 33 5 lm 34 35 Sporisorium panici-leucophaei (Bref.) M. Piepenbr., comb. nov. (Figs 42–43) Basionym : Ustilago panici-leucophaei Bref., Untersuchungen aus dem Gesammtgebiete der Mykologie. XII : 114 (1895). Type on Panicum leucophaeum H.B.K. [¯ Trichachne insularis (L.) Nees], Brazil, Rio de Janeiro, E. Ule. Figs 32–35. Sporisorium everhartii. Fig. 32. Basidiospores. Figs 33–35. Germinated teliospores after 1 d on water agar. (MP 2270). 36 37 Spore wall ca 0±5 µm thick, seen by LM minutely granularverrucose to almost smooth, seen by SEM covered by small and somewhat larger warts. Sterile cells subglobose, sometimes in chains, larger than the spores, hyaline, thin-walled. Germination (Figs 40–41) of Ustilago type with mostly fourcelled basidia carrying basidiospores. Known on Paspalum dilatatum Poir., P. notatum Flu$ gge, P. plicatulum Michaux, P. proliferum Arech., P. urvillei Steudel, and P. vaginatum Swartz, Poaceae, from several countries of Latin America. This is the first record of this species for Cuba. Specimen examined in addition to the type of U. paspali-notati : On Paspalum notatum, Cuba, Prov. Pinar del Rı! o, Sandino, in front of the Instituto Pedago! gico Superior, alt. ca 10 m, 14 Aug. 1996, J. Gutierrez, A. Urquiola & M. Piepenbring (MP 2101). Fischer (1953 : 133) considers Sporisorium microsporum (as Sphacelotheca microspora) together with Sphacelotheca panicileucophaei (Bref.) G. P. Clinton (and other names) a synonym of S. cordobensis (Speg.) Cif. These species are treated as distinct by Zundel (1953). He uses the following distinctive characteristics : (i) Sphacelotheca cordobensis on Trichachne insularis (L.) Nees : sori with numerous columellae at the place of the inflorescence ; teliospores 9–11(–14) µm, distinctly echinulate ; 1 cm Figs 36–37. Inflorescences of Paspalum notatum infected by Sporisorium microsporum. Fig. 36. Young sori. Fig. 37. Old sori. (MP 2101). Cuban smut fungi 466 38 40 41 39 5 lm Figs 40–41. Sporisorium microsporum. Germinated teliospores after 2 d on water agar (MP 2101). Figs 38–39. Sporisorium microsporum. Teliospores seen by SEM. (bars ¯ 4 µm). Fig. 38. Type. Fig. 39. MP 2101. 43 Fig. 43. Sporisorium panici-leucophaei. Teliospores seen by SEM (scale bar ¯ 4 µm) (N. H. 15969). Synonym : Sphacelotheca panici-leucophaei (Bref.) G. P. Clinton, North American Flora 7 : 28 (1906). (For further synonyms see Zundel, 1953 : 103.) Sori (Fig. 42) preventing the development of an inflorescence, the axis of the plant is shredded with vascular strands forming numerous columellae, sori surrounded by peridia made of host tissue and some sterile fungal cells. Spore mass powdery, with few sterile cells, dark brown to black. Spores (Fig. 43) single, globose to subglobose, occasionally polyhedral, 6–7¬(6–)7–8 µm, light golden brown to reddish-brown. Spore wall ca 0±5 µm thick, as seen by LM minutely granularverrucose to almost smooth, as seen by SEM sparsely warty. Sterile cells subglobose, sometimes in chains, larger than the spores, hyaline, thin-walled. Germination of Ustilago type with four-celled basidia bearing basidiospores (Brefeld, 1895). Known on Trichachne californica (Benth.) Chase (T. saccharata Nash., Panicum saccharatum Buckl.), T. insularis (L.) Nees (Panicum lanatum Rottb., P. leucophaeum H.B.K., P. jaboncillo Hieron. nom. nud.), Poaceae, from many countries of Latin America including Cuba (Whetzel & Kern, 1926) and the U.S.A. Specimen examined : On Trichachne insularis, Dominican Republic, Prov. Santiago, Valle del Cibao, Santiago, Hatillo, in fields, 17 Sep. 1930, E. L. Ekman (Plantae Hispaniolae Exp. III. Regnellianae, N. H. 15969 ; in herb. S). M. Piepenbring 42 1 cm Fig. 42. Trichachne insularis infected by Sporisorium panici-leucophaei (N. H. 15969). I thank the Cuban mycologists and botanists M. Rodrı! guez, M. Camino, J. Gutierrez, A. Urquiola, and many more, who helped as guides in the field, in identification, and organisation of the research which was supported by the DFG (Deutsche Forschungsgemeinschaft, III 06-477}1161}96). Z. L. Yang and M. Weiß are thanked for improving the manuscript, H. Schoppmann for technical assistance, curators of the herbaria LPS, M, and S for loans of specimens, and V. 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