The genus Ravenelia in Argentina

Mycol. Res. 106 (8) : 954–974 (August 2002).

# The British Mycological Society

954

DOI : 10.1017\S0953756202006226 Printed in the United Kingdom.

The genus Ravenelia in Argentina

Jose! R. HERNA; NDEZ1 and Joe F. HENNEN2 " Systematic Botany and Mycology Laboratory, USDA Agricultural Research Service, 10300 Baltimore Blvd., Beltsville, MD 20705-2350, USA. # Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, USA. E-mail : jose!nt.ars-grin.gov Received 28 February 2002 ; accepted 28 June 2002.

A comprehensive account of the 18 species of Ravenelia from Argentina is presented. Each species is described and illustrated. A key to the species of Ravenelia from Argentina is provided based on micromorphology. Ravenelia echinata var. ectypa and R. macrocarpa are recorded for the first time from Argentina. The name Ravenelia argentinensis was determined to belong to an anamorph, therefore the teleomorph is described as a new species, R. argentinica sp. nov. Several new anamorph\teleomorph connections were made. Names and synonyms for teleomorphs and anamorphs are listed for all species.

INTRODUCTION The genus Ravenelia Berk. 1853 was described to honour Henry William Ravenel (1814–1887). The Rev. M. J. Berkeley of England corresponded extensively with the North American collectors H. W. Ravenel and M. C. Curtis who sent abundant material to him from the New World. Ravenel, in particular, had sent him numerous specimens of novel fungi from South Carolina. Among the rust fungi (Uredinales) Ravenelia is the third most speciose genus with approximately 200 recognized species. Species of Ravenelia are found worldwide predominantly parasitizing members of the Fabaceae (legumes) in all three subfamilies, Caesalpinioideae, Faboideae and Mimosoideae (Cummins & Hiratsuka 1991). Species of Ravenelia occur in warm temperate and tropical regions where members of the Fabaceae are abundant. The largest number of species of Ravenelia is found in the Neotropics, followed by Asia and Africa. No species of Ravenelia are recorded from Europe or Australia. Nearly 20 species of Ravenelia induce abnormal growth of their host, such as galls or witches’ brooms. This symptomatology occurs frequently on fabaceous plants in wooded and shrubby areas in Argentina. All species of Ravenelia are autoecious and some of them are microcyclic. The genus Ravenelia is notable because it produces many-celled teliospores that are among the largest and most complex of fungal spores. Teliospores of Ravenelia are discoid when the adaxial

side is observed and reniform or elliptical in side view. The size and complexity of teliospores varies among taxa and are diagnostic for species of Ravenelia. The teliospores of Ravenelia have been called ‘ capitules composed of pseudospores ’ (Cooke 1880) or ‘ spore heads ’ composed of adherent, one to two celled spores (Cummins 1978, Hennen & Cummins 1990), because they were considered to be a complex structure of many teliospores. Other authors (Gallegos & Cummins 1981), as we do, consider the ‘ spore head ’ to be a single teliospore composed of many pigmented probasidial cells. Hyaline, hygroscopic cysts occur on the abaxial surface of the teliospore and sterile intercalary cells are sometimes present between the probasidial cells and the cysts. Teliospores are produced on a fasciculate pedicel of two to a few hyphae (Figs 38, 71–72). One germ pore or germ slit is present on the adaxial surface of each probasidial cell (Fig. 39). The outer layer of the teliospore is smooth or tuberculate, branched tuberculate or with few to numerous sharply pointed cones. In addition to characteristics of the telia (stage III), the morphological characters of the anamorphs are useful in the taxonomy of the species of Ravenelia. The aecia (stage I) are subepidermal or sometimes subcuticular, becoming erumpent, and have aeciospores that are sometimes sessile and catenulate, but sometimes pedicellate. Uredinia (stage II) are subepidermal or subcuticular, becoming erumpent, and often paraphysate. Urediniospores are produced singly on pedicels, the walls are mostly echinulate and with germ pores that vary in number and position. Anamorph

J. R. Herna! ndez and J. F. Hennen spores of Ravenelia mimosae-sensitivae and Ravenelia spegazziniana have a distinctive ‘ hub and spoke ’ ornamentation composed of knob-like processes interconnected by straight ridges (Gardner & Hodges 1985) (Figs 53, 79–80). In addition to the above stages, four round to ovoid basidiospores (stage IV) are produced on the metabasidia that germinate from the probasidial cells. Those basidiospores reinfect the host and eventually subepidermal or subcuticular spermogonia (stage 0) are produced (type 5 and 7 respectively ; Cummins & Hiratsuka 1983). The Argentinian mycologist Carlos Spegazzini named and described eight species of Ravenelia and a number of anamorphs that were later connected to Ravenelia teleomorphs (Farr 1973). Lindquist (1946, 1951, 1978) described five additional species, and later included 16 species of Ravenelia in his comprehensive treatment of Argentinean rusts (Lindquist 1982). In a recent study of the rust fungi of the ecologically diverse region known as Northwest Argentina, 13 of the 16 previously described species of Ravenelia from Argentina were collected (Herna! ndez & Hennen 2002) ; only eight of the taxa had been collected in the region previously. Two additional taxa unknown from Argentina also were collected (namely R. echinata var. ectypa and R. macrocarpa). In this paper, descriptions and illustrations are provided for the 18 species of Ravenelia known from Argentina based on our recent collections and examination of reference specimens in BPI, LIL, LPS, and PUR. Additional information was obtained from unpublished notes of G. B. Cummins. We follow the Cummins & Hiratsuka (1983) system for terminology of life cycle which is based on the assumed position in the life cycle and comparative morphology. A key is provided to the species of Ravenelia known from Argentina, based primarily on teliospore and anamorph spore morphology. Ravenelia argentinica is described as a new teleomorphic species for R. argentinensis, which was based on an anamorph. TAXONOMY Ravenelia argentinica J. R. Hern. & J. F. Hennen, sp. nov. (Figs 1–11) Etym. : after Argentina, the country from which this species is known. Teliosporae in urediniis 80–120 µm diametro, atro-brunneae, ex 6–10 probasidialibus cellulis in omni directione constatae ; cellulae 15–18 µm diametro, unistratae, utraque 4–7(–8) spinis, 3–6 µm longis ; cystae uniseriales, plus minusve globosae, pendentes, hyalinae ; pedicello ex hyphis multis constato, caduco. Typus : Argentina : Corrientes : on Acacia aroma, 12 July 1944, M. di Fonzo 375 (BPI 149244 – holotypus, LPS 12667 – isotypus) [0–I–II–III].

Anamorph : Caeoma sp. Ravenelia argentinensis Speg., Revista Argent. Bot. 1 : 129 (1925) ; nom. ananams. Type : Argentina : Jujuy, Perico, on A. aroma [orig-

955 inally reported as Vachellia lutea], Jan. 1906, leg. ? (LPS 4941 [4901 sic] – holotype) [I]. Spermogonia not seen. Aecia on abnormal growths on young branches and pods, subepidermal, becoming erumpent, without peridia or paraphyses, yellowish ; aeciospores catenulate, obovoid, ellipsoid to oblongfusiform, sometimes apiculate, 22–26i12–18 µm, wall 1n5–2 µm thick at sides, 2n5–3 µm thick at top, yellowish, densely verrucose, pores mostly 4–5, equatorial. Uredinia 0n5–1 mm diam., epiphyllous, rounded, subepidermal, becoming erumpent ; paraphyses 50– 60i10–22 µm, peripheral, slightly incurved, wall cinnamon-brown, slightly thickened dorsally ; urediniospores 25–29i14–18 µm, pedicellate, obovoid, globose, or broadly ellipsoidal, wall 1n5–2 µm thick, or slightly thicker at an apical papilla, finely echinulate, pores 4, equatorial. Teliospores produced in the uredinia, 80–120 µm diam., dark brown, 6–10 probasidial cells across, cells 15–18 µm diam., 1-layered, each cell with 5–7(–8) spines, 3–6 µm long ; cysts uniseriate, more or less globoid, pendent, hyaline ; pedicel multihyphal, deciduous. Host and distribution : Known only on Acacia aroma from Argentina. Other specimens examined : (all on Acacia aroma) Argentina : Catamarca : N of Catamarca, hwy 38, 28m 22n955h S, 65m 36n918h W, 24 Sept. 1997, J. F. Hennen & J. R. HernaT ndez 097-052 (BPI 841193) [I] ; Jujuy : Parque Nacional Calilegua, 8 Dec. 1997, J. R. HernaT ndez 097-179 (BPI 841267) [I] ; Salta : Meta! n, 2 Apr. 1993, J. F. Hennen & L. D. Ploper 093-103 (BPI 841014, LIL 54823) [I–II–III] ; road to Parque del Rey, 6 Dec. 1997, J. R. HernaT ndez 097-160 (BPI 841256) [I] ; TucumaT n : San Javier, Parque Sierras de San Javier, 27 Mar. 1993, J. F. Hennen, L. D. Ploper & J. R. HernaT ndez 093-032 (LIL 54844, BPI) [I–II–III] ; Cevil Pozo, 30 Mar. 1993, J. F. Hennen & J. R. HernaT ndez 093-051 (LIL 54824, BPI) [II] ; Cruz Alta, San Agustı! n, 30 Mar. 1993, J. F. Hennen & J. R. HernaT ndez 093-059 (LIL 54825, BPI) [II–III] ; J. F. Hennen & J. R. HernaT ndez 093-060A (LIL 54845, BPI) [III] ; Las Cejas, 20 km E. of San Miguel de Tucuma! n, 30 Mar. 1993, J. F. Hennen & J. R. HernaT ndez 093-065 (LIL 54826, BPI) [II–III] ; J. F. Hennen & J. R. HernaT ndez 093-069 (LIL 54847, BPI) [I–II–III] ; J. F. Hennen & J. R. HernaT ndez 093072 (LIL 54848, BPI) [I–II] ; J. F. Hennen & J. R. HernaT ndez 093-74 (LIL 54827, BPI) [II–III] ; J. F. Hennen & J. R. HernaT ndez 093-076 (LIL 54821, BPI) [I–II] ; J. F. Hennen & J. R. HernaT ndez 093-078 (LIL 54822, BPI) [I–II–III] ; Raco, 3 Nov. 1997, J. R. HernaT ndez 097-142 (BPI 841241) [I] ; J. R. HernaT ndez 097-143 (BPI) [II].

The most obvious trait of R. argentinica is the abundant ‘ arachnoid ’ or ‘ spidery ’ abnormal growths produced on the systemically infected inflorescence (Figs 10–11) and the less abundant witches’ brooms produced from systemically infected buds (Fig. 10). In heavily infected trees these can be seen from a distance because of their size and abundance. Infections begin in early spring just as the inflorescences begin to appear and the buds begin to grow. As young pods are forming they grow as a bundle of distorted structures taking on the ‘ spidery ’ appearance (Fig. 11). Aecia soon develop

Ravenelia in Argentina

956

Figs 1–9. Ravenelia argentinica on Acacia aroma. Fig. 1. Caeomoid aecia (BPI 841267). Fig. 2. Aeciospores with typical verrucose ornamentation (BPI 149244). Fig. 3. Surface view of aeciospores showing equatorial germ pores and verrucose ornamentation (BPI 841241). Fig. 4. Median view of aeciospores in Fig. 3. Fig. 5. Uredinium with peripheral paraphyses (LIL 54823, BPI 841014). Fig. 6. Urediniospore with echinulate ornamentation (LIL 54823, BPI 841014). Fig. 7. Urediniospores and paraphyses (LIL 54821). Fig. 8. Uredinium in which teliospores are forming (BPI 149244). Fig. 9. Detail of spines on teliospore probasidial cells (BPI 149244). Figs 1–2, 5–6, 8–9 : SEM. Figs 3–4, 7 : Light Microscopy (LM). Bars : Fig. 1 l 80 µm ; Figs 5, 8 l 20 µm ; Fig. 7 l 13n3 µm ; Figs 3–4 l 12n5 µm ; Figs 2, 6, 9 l 3 µm.

and appear as small dots on young, dark green, infected tissue (Figs 1, 11). As they mature they become erumpent and yellowish. Later uredinia and telia develop on locally infected leaflets (Figs 5, 8). Most of the abnormal growths die during the season but some may remain alive with the fungus continuing to produce spores until the next season. In winter the dead, dry, brittle abnormal growths remain obvious on the leafless trees (Fig. 10). Lindquist (1954) reported that Spegazzini (1925) had erroneously described glandular trichomes from the stem of the host in the type specimen as teliospores of R. argentinensis. Although Spegazzini did not publish any illustrations with his original description, Lindquist (1954) discovered drawings made by Spegazzini in the type packet in LPS that confirmed this mistake.

Spegazzini did describe the aecial and uredinial stages present in the type. Because Spegazzini did not describe the teleomorph, the name R. argentinensis can only apply to the anamorph (Art. 59n2). Therefore, the presumed teleomorphic name R. argentinensis applies to the anamorph and should be referred to as the Caeoma state. This is the appropriate genus for the aecial anamorph because the sori are not surrounded by a peridium and the spores are catenulate (Fig. 1). Both Spegazzini (1925) and Lindquist (1954) mistakenly reported peridial cells. Spegazzini did not describe them but Lindquist reported the supposed peridial cells as 18–20i30–36 µm, elongate rectangular, with thick, smooth walls. Our study shows that these supposed ‘ peridial cells ’ are abnormal host cells next to the rust sori.

J. R. Herna! ndez and J. F. Hennen

957

Figs 10–11. Ravenelia argentinica on Acacia aroma. Fig. 10. Witches’ brooms and arachnoid abnormal growth on stems (BPI 841193). Fig. 11. Detail of arachnoid abnormal growth (BPI 841267). Figs 12–13. Ravenellia chacoeW nsis on Prosopis nigra. Fig. 12. Median view of teliospore showing pendent cysts on proximal side of the spore (LPS 11994). Fig. 13. Surface view of smooth teliospore (LPS 11994). Figs 14–15. Ravenelia hieronymi on Acacia sp. Fig. 14. Witches’ brooms on stems (BPI 841267). Fig. 15. Detail of witches’ broom on stem (BPI 841165). Figs 12–13 : LM. Bar l 10 µm.

Lindquist (1954) found abundant teliospores in a specimen from Corrientes that he identified as R. argentinensis because of its anamorph spore traits. He published a description of those teliospores in Spanish, but not Latin, under the name R. argentinensis ; that name also is not available for the teleomorph (Art 59, Ex. 5), therefore we describe it as a new species, providing a Latin diagnosis of the teleomorph and a new type specimen on which the teleomorph is present. Spermogonia were not described by Spegazzini (1925). Lindquist (1954) reported that he did not observe them either, however, he cited this stage (stage 0) for some of the collections examined. We did not find spermogonia in collections from Northwest Argentina. Ravenelia australis Dietel & Neger, Bot. Jahrb. Syst. 24 : 161 (1897). (Figs 16–19) Type : Chile : ConcepcioT n, on Acacia caven (originally reported as A. cavenia), Mar. 1896, Neger s.n. (S F21494, holotype ; S F21495, PUR 68982, BPI 159349, BPI 149348 – isotypes) [III]. Anamorph : Aecidium sp. Uredo hieronymi Speg., Anales Soc. Ci. Argent., Pug. IV, 12 : 73 (1881). Type : Argentina : CoT rdoba : Sierra Chica, near San Jose! , on A. caven, 11 Jan. 1877, Hieronymus s.n. (LPS 9489 – holotype) [I].

Uredo hieronymi Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 23 : 30 (1912), nom. illeg. (later homonym). Type : Argentina : Catamarca, on A. caven (originally reported as A. cavenia), Nov. 1909, Spegazzini s.n. (LPS 9487 – holotype, n.v.). Spermogonia epiphyllous and on stems, 100–120 µm diam, grouped around aecia, without periphyses. Aecia on stems causing witches’ brooms, or amphigenous causing slight localized hypertrophy, 400 µm diam., subepidermal becoming erumpent, pale brown, with rudimentary or inconspicuous peridium ; aeciospores (24–)28–35(–38)i(14–)16–20(–21) µm, catenulate, variable in size and shape, ellipsoid, obovoid or oblongfusiform, sometimes apiculate, wall 1n5–2 µm thick at sides, 3–5 µm at apex, golden yellowish, densely verrucose, pores mostly 5–6(–7) and equatorial, sometimes 6–8 and tending to be bizonate. Uredinia usually amphigenous, sometimes hypophyllous, roundish, 0n5 mm diam., subepidermal becoming erumpent, cinnamon-brown ; paraphyses abundant, peripheral, 50–54i8–10 µm, basally united, brown, incurved, often geniculate, dorsally thick-walled ; urediniospores (23–)25–30(–33)i(14–)16–20(–21) µm, ellipsoid or obovoid, wall uniformly 1n5(–2) µm thick or slightly thicker apically, golden brown, echinulate, pores 4–6, equatorial, with slight or no caps. Telia similar to uredinia ; teliospores often produced within the uredinia

Ravenelia in Argentina

958

Figs 16–19. Ravenelia australis on Acacia sp. Fig. 16. Uredinium with geniculate peripheral paraphyses (BPI 149350). Fig. 17. Detail of basally united paraphyses (Cummins 71-408). Fig. 18. Telium with peripheral paraphyses (BPI 149351). Fig. 19. Teliospore (Long 6151). Figs 20–23. Ravenelia cebil on Anadenanthera sp. Fig. 20. Subcuticular uredinium (LIL 54828, BPI 841029). Fig. 21. Echinulate urediniospores and smooth paraphyses (LIL 54828, BPI 841029). Fig. 22. Capitate, apically thickened paraphyses and urediniospores with equatorial germ pores. Fig. 23. Telium (LIL 54828, BPI 841029). Figs 24–29. Ravenelia cohniana on Fabaceae. Fig. 24. Subepidermal uredinium (BPI 149498). Fig. 25. Urediniospores (H. Sydow 131). Fig. 26. Urediniospores showing echinulate ornamentation and acuminate apex (BPI 149498). Fig. 27. Telium (LIL 54832, BPI 841015). Fig. 28. Median view of teliospore (H. Sydow 131). Fig. 29. Teliospore showing uniseriate, adherent cysts on proximal side (BPI 149512). Figs 16, 18, 20–21, 23–24, 26–27, 29 SEM. Figs 17, 19, 25, 28 : LM. Fig. 22 : LM, Face contrast (FC). Bars : Figs 16, 18, 20, 23–24, 27 l 20 µm ; Figs 17, 19, 22, 25, 28 l 13n3 µm ; Fig. 29 l 10 µm ; Figs 21, 26 l 3 µm.

J. R. Herna! ndez and J. F. Hennen and surrounded by the uredinial paraphyses, (70–)80–110(–135) µm diam., chestnut-brown, smooth, 7–12 probasidial cells across, cells cuneiform, in one layer, central cells (10–)13–18(–20) µm diam. ; cysts biseriate, numerous, more or less globoid, pendent ; pedicel multihyphal, hyaline, deciduous. Hosts and distribution : On Acacia caven (syn. A. cavenia), Argentina (Buenos Aires, Catamarca, Co! rdoba, Santiago del Estero), Chile, USA ; on A. farnesiana, USA ; on A. pennatula, Mexico ; on A. smallii, Mexico, USA. Other specimens examined : Argentina : Buenos Aires : La Plata, Fac. de Agronomı! a, 25 Mar. 1949, on A. caven, J. C. Lindquist s.n. (LPS 16669, BPI 149350) [0–I–II–III]. Chile : ConcepcioT n, on A. caven, Dec. 1995, Neger s.n. (S F21496) [I]. USA : Texas : Uvalde, 7 Nov. 1916, on A. farnesiana, W. H. Long s.n. (BPI 149351) [III].

Uredo hieronymi (Spegazzini 1881) applies to the aecial anamorph of Ravenelia australis. Spegazzini (1912) published the illegitimate homonym Uredo hieronymi and in both descriptions he indicated uncertainty as to whether the spores were catenulate or sessile. The sori are associated with abnormal growths, contain no paraphyses and the spores are catenulate. Spermogonia were later reported by Lindquist (1954) lending even more support to the idea that this is an aecial stage. The epithet ‘ hieronymi ’ cannot be transferred to Aecidium, because A. hieronymi (Spegazzini 1881) is used already for the anamorph of R. hieronymi. We do not provide a new binary name for the aecial anamorph of R. australis because the connection with the teleomorph is firmly established (Art. 59A.3) ; rather we refer to it as Aecidium sp. Ravenelia cebil Speg., Anales Mus. Nac. Hist. Nat. Buenos. Aires 19 : 295 (1909). (Figs 20–23) Type : Argentina : TucumaT n : Parque Roca, on Anadenanthera macrocarpa (originally reported as Piptadenia macrocarpa), 10 Apr. 1906, Spegazzini s.n. (LPS 4950 [4959 sic] – holotype). Anamorph : Uredo vilis (Syd. & P. Syd.) J. W. Baxter, Mycologia 67 : 437 (1975). Ravenelia vilis Syd. & P. Syd., Ann. Mycol. 14 : 68 (1916). Type : Brazil : Estado de CearaT : Serra Maranguape, on Anadenanthera sp. (originally reported as Piptadenia sp.), Oct. 1910, Ule 3408 (S F21395 – lectotypus hic designatus ; S F21396, PUR – isolectotypi) [II]. Uredinia amphigenous, mainly hypophyllous, 1–2 mm diam., scattered or in groups of 4–5, subcuticular becoming erumpent ; paraphyses abundant, light brown, capitate, apically thick-walled, to 70 longi10– 12 µm, intermingled with the urediniospores ; urediniospores (18–)22–30(–32)i(10–)11–14(–16) µm, ellipsoid, elongate-obovoid or oblong-ellipsoid, wall 1n5–2 µm thick or 2–2n5 µm at apex, yellowish to cinnamonbrown, inconspicuously echinulate becoming smooth or with smaller spines apically, pores 4–6 equatorial. Telia amphigenous in small groups, roundish, sub-

959 cuticular, erumpent, blackish brown ; teliospores (60–)75–95(–100) µm diam., chestnut-brown with a pale, thin, outer layer, 5–7 µm wide, smooth or slightly verrucose, (5–)6–8(–9) probasidial cells across, central cells 2-layered, (14–)15–18(–19) µm diam. ; cysts small, globose, multiseriate, pendent, laterally free ; pedicel colourless, multihyphal, short, deciduous. Hosts and distribution : On Anadenanthera colubrina var. cebil, Argentina ; on A. macrocarpa, Argentina (La Plata, Tucuma! n) ; on A. peregrina var. falcata, Brazil ; on A. peregrina var. peregrina, Brazil, Puerto Rico, Virgin Islands ; on Anadenanthera sp., Argentina, Brazil ; on substrate indet., Puerto Rico. Other specimens examined : (all on Anadenanthera sp.) Argentina : Jujuy : Caimancito, 2 Apr. 1993, J. F. Hennen & L. D. Ploper 093-111 (LIL 54831, BPI) [II–III] ; Pampa Blanca, 1 Apr. 1993, J. F. Hennen & L. D. Ploper 093-100 (LIL 54829, BPI) [II–III] ; San Pedro, 2 Apr. 1993, J. F. Hennen & L. D. Ploper 093-107 (LIL 54830, BPI) [II–III] ; TucumaT n : San Pedro de Colalao, 1 Apr. 1994, J. F. Hennen & M. M. Hennen 094-059 (LIL 54828, BPI 841029) [II–III].

Ravenelia cebil differs from R. henningsiana Dietel 1906 because of the mostly smooth probasidial cells, teliospore size and absence of paraphyses (Lindquist 1954) of the former. Sydow & Sydow (1916) described only uredinia and urediniospores in their publication of R. vilis. Baxter (1975) transferred R. vilis to the anamorph genus Uredo. Ravenelia chacoe$ nsis J. C. Lindq., Bol. Soc. Argent. Bot. 1 : 300 (1946). (Figs 12–13) Type : Argentina : Chaco : Laguna Blanca, on Prosopis nigra, 15 June 1944, di Fonzo 108 (LPS 11994 – holotype) [III]. Spermogonia, aecia, and uredinia unknown. Telia hypophyllous and on rachis’, subepidermal, 0n5–1 mm diam., becoming erumpent, blackish brown, rounded, grouped in circles ; teliospores (80–)100–120(–130) µm diam., chestnut-brown, smooth, with a thin but discrete, nearly colourless outer wall layer, 7–12 probasidial cells across, central cells 17–22 µm diam., cells 1-layered, wall 2–3 µm thick at sides and 3–7 µm thick at the apex ; paraphyses incurved, slightly clavulate, intermingled with teliospores, 30i10 µm, wall thin, colourless, later cinnamon-brown ; cysts multiseriate, pendent, globoid, colourless, swollen in water ; pedicel colourless, short, multihyphal, mostly deciduous. Hosts and distribution : On Prosopis nigra, Argentina, Chaco (Laguna Blanca). Known only from the type. Four species of Ravenelia have been described on Prosopis in the Americas, namely R. arizonica Ellis & Everhart 1895, R. chacoeW nsis, R. holwayii Dietel 1894 and R. prosopidiicola J. C. Lindq. 1951. Although Herna! ndez & Hennen (1999) proposed treating these four taxa as varieties of the same species, they are maintained herein as separate species based on a reevaluation of the morphological characters. The telia of the apparently microcyclic R. chacoeW nsis are very similar

Ravenelia in Argentina to those of the macrocyclic R. holwayii, but the latter can be distinguished because it also produces spermogonia, aecia and uredinia. R. chacoeW nsis differs from R. arizonica (Ellis & Everhart 1895) in that the latter produces echinulate teliospores and host hypertrophy, and from R. prosopidiicola because the teliospores of the latter are smaller and lighter in colour (Figs 71–72). Ravenelia cohniana Henn., Hedwigia 35 : 246 (1896). Type : Brazil : Rio de Janeiro : on Caesalpinia sp., July 1887, E. Ule 703 (B – holotype, BPI 149496, HBG, PUR, – isotypes) [II–III]. Ravenelia distans Arthur & Holw., in Arthur, Amer. J. Bot. 5 : 424 (1918). Type : Guatemala : Retalhuleu : on Mimosoideae, 26 Feb. 1916, Holway 535 (BPI 149511 – lectotypus hic designatus ; BPI 149410, BPI 149512, BPI 149513, PUR 6118 n.v., NY 332844 n.v., NY 28480 n.v. – isolectotypi) [II–III]. Ravenelia concinna Syd., Ann. Mycol. 28 : 46 (1930). Type : Venezuela : Caguita, on Acacia sp., 29 Dec. 1927, H. Sydow 135 (BPI 149498 – lectotypus hic designatus ; MINN, PAC, PUR – isolectotypi) [II–III]. Ravenelia lindquistii J. F. Hennen & Cummins, Rep. Tottori Mycol. Inst. 28 : 7 (1990). Type : Argentina : Salta : Alemania, Quebrada del Rı! o, on Acacia praecox, 12 Aug. 1936, Cabrera 3753 (LPS 12626 – holotype ; PUR F11379 – isotype) [II– III]. Anamorph : Uredo imperspicua Speg., Rev. Arg. Bot. 1 : 136 (1925). Type : Argentina : Salta : North of Ora! n, Rı! o Pescado, on A. praecox, Feb. 1905, Spegazzini s.n. (LPS 9534 – holotype) [II]. Ravenelia imperspicua (Speg.) J. C. Lindq., Bol. Soc. Argent. Bot. 1 : 301 (1946). Spermogonia and aecia unknown. Uredinia 0n5–1 mm diam., hypophyllous and caulicolous, subepidermal, erumpent, cinnamon-brown, scattered, without paraphyses ; urediniospores (12–)20–28(–32)i(11–)13– 17(–19) µm, variable in shape, mostly ellipsoid or ovoid, commonly pointed apically and acute towards the base of the spore, wall 1n5–2n5(–3) µm thick at sides, 3–5(–7) µm at apex, finely and rather sparsely echinulate, pale golden brown, pores (3)4(–6) equatorial. Telia as the uredinia except blackish brown ; teliospores (39–)45–73(–74) µm diam., (3)4–5(6) probasidial cells across, 16–22i13–15 µm, 1-layer, clear chestnut brown with an outer hyaline thin layer bearing (2–)3–8(–10) sharp cones 3–5 µm and 2–3 µm wide at base, central probasidial cells, 14–18(–25) µm diam. ; cysts uniseriate of same number as peripheral probasidial cells, globoid, coherent, appressed or semipendent ; pedicel colourless, mostly deciduous, few hyphae (two hyphae in larger teliospores but one in small teliospores, fide Dietel 1906). Hosts and distribution : On Acacia praecox, Argentina (Buenos Aires, Catamarca, Salta, Tucuma! n) ; on Acacia

960 sp., Venezuela ; on Caesalpinia sp., Brazil ; on Mimosa sp., Guatemala ; on Mimosaceae indet., Guatemala ; on Fabaceae indet., Argentina. Other specimens examined : Argentina : Buenos Aires : La Plata, Fac. de Agronomı! a, 16 May 1945, on Acacia praecox, J. C. Lindquist s.n. (LPS 12599) [II–III] ; Catamarca : Rı! o Las Juntas, camping area, 28m 25n949h S, 65m 50n739h W, 23 Sept. 1997, on A. praecox, J. F. Hennen & J. R. HernaT ndez 097-039 (BPI 851185) [II–III] ; bank of Rı! o Las Juntas, 28m 25n103h S, 65m 50n978h W, 23 Sept. 1997, on A. praecox, J. F. Hennen & J. R. HernaT ndez 097-041 (BPI 841187) [II–III] ; Jujuy : Chalica! n, 2 Apr. 1993, on Fabaceae indet., J. F. Hennen & L. D. Ploper 093-105 (LIL 54832, BPI 841015) [II–III] ; Salta : Pichanal, 2 Apr. 1993, on A. praecox, J. F. Hennen & L. D. Ploper 093-108 (LIL 54850, BPI) [II] ; J. F. Hennen & L. D. Ploper 093-109 (LIL 54834, BPI) [II–III] ; TucumaT n : San Pedro de Colalao, 2 Apr. 1994, on A. praecox, J. F. Hennen & M. M. Hennen 094-067 (LIL 54833, BPI) [II–III].

Spegazzini (1925) described Uredo imperspicua as a new anamorphic species. Lindquist (1946) found teliospores on other U. imperspicua collections, which he considered to belong to the same rust life cycle. He transferred the anamorph epithet to the teleomorphic genus Ravenelia but did not provide a Latin description or indicate a teleomorph type. Hennen & Cummins (1990) considered R. imperspicua a nomen nudum because it lacked a Latin description and therefore it was not validly published as a teleomorphic name (Art. 36). They published the new name R. lindquistii for the teleomorph, provided a Latin description of the teleomorph, and designated Lindquist’s (1954) type as the holotype. We consider R. concinna, R. distans, and R. lindquistii to be taxonomic synonyms of R. cohniana because they all have similar spiny teliospores, urediniospores with pointed or apiculate apices, (3–)4(–5) equatorial germ pores, and subepidermal sori. The sori of R. concinna are subepidermal in origin, not subcuticular as originally reported (Sydow 1930). Teliospores of R. distans have spines that are slightly more pointed than the rounded spines of R. concinna, but we do not consider these to be significantly different. Ravenelia cohniana is unusual because it infects hosts in two legume subfamilies, the Caesalpinioideae and Mimosoideae. Ravenelia echinata var. ectypa (Arthur & Holw.) Cummins, Bol. Soc. Argent. Bot. 18 : 85 (1977). (Figs 30–34) Ravenelia ectypa Arthur & Holw., in Arthur, Mycologia 10 : 120 (1918). Type : Costa Rica : San JoseT : on Calliandra gracilis, 26 Dec. 1915, Holway 296 (PUR 6160 – holotype ; BPI 149523, BPI 149525, BPI 149527, MINN n.v., NY 28483 n.v., isotypes) [II–III]. Anamorph : Uredo sp. Spermogonia amphigenous, subcuticular, type 7. Aecia amphigenous surrounding the spermogonia, subcuticular becoming erumpent, cinnamon-brown, with few, thin-walled, hyaline, cylindrical or clavate, per-

J. R. Herna! ndez and J. F. Hennen

961

Figs 30–34. Ravenelia echinata var. ectypa on Calliandra spp. Fig. 30. Urediniospores (BPI 149524). Fig. 31. Surface view of urediniospores showing scattered germ pores (BPI 149524). Fig. 32. Median view of urediniospores shown in Fig. 31. Fig. 33. Telium (LIL 54837, BPI 841034). Fig. 34. Teliospore with conical spines and 4 central surrounded by 6 peripheral probasidial cells (LIL 54837, BPI 841034). Figs 35–39. Ravenelia hieronymi on Acacia sp. Fig. 35. Aecium (BPI 841165). Fig. 36. Densely and finely verrucose aeciospores (BPI 841224). Fig. 37. Telium (BPI 149789). Fig. 38. Teliospore showing cysts and multihyphal pedicel on proximal side (Hennen 76-310). Fig. 39. Surface view of teliospores showing pairs of germ slits (BPI 841165). Figs 30, 33–37 SEM. Figs 31–32, 38–39 LM. Bars : Fig. 33 l 20 µm ; Figs 31–32, 38–39 l 12n5 µm ; Figs 30, 34–35, 37 l 10 µm ; Fig. 36 l 3 µm.

ipheral paraphyses ; aeciospores (18–)19–26(–28)i (16–)17–19(–20) µm, pedicellate, mostly broadly ellipsoid, wall 1–1n5 µm thick, golden or pale brown, echinulate, pores scattered (5)6–8(–10) with slight caps. Uredinia amphigenous, scattered or grouped, subcuticular becoming erumpent, cinnamon-brown, without paraphyses ; urediniospores similar to aeciospores. Telia amphigenous, subcuticular, becoming erumpent, 0n5–1 mm diam., pulverulent, blackish brown ; teliospores (46–)52–66(–68) µm diam., chestnutbrown, (3)4(5) probasidial cells across, mostly 4 central cells surrounded by 6 peripheral cells, each with 3–9(–12) conical spines 2–2n5 µm wide at base and 3–5 µm long as part of an outer pale layer covering the spores, central cells (17–)20–25(–28) µm diam., cells 1-

layered ; cysts uniseriate of same number as peripheral probasidial cells, appressed to the underside of the spores or becoming pendent ; pedicel hyaline, of few (often 2) hyphae, mostly deciduous. Hosts and distribution : On Calliandra calothyrsus, Guatemala ; C. formosa (syn. Acacia formosa), Costa Rica, Mexico ; on C. gracilis, Costa Rica, Guatemala, Mexico ; on C. laxa (syn. Acacia laxa), Bolivia ; on C. marginata, Mexico, Trinidad & Tobago ; on C. penduliflora, Guatemala, Mexico ; on Calliandra sp., Costa Rica, Guatemala, Mexico, South America. Other specimens examined : Argentina : Salta : Pichanal, 2 Apr. 1993, on Calliandra formosa, J. F. Hennen & L. D. Ploper 093-102 (LIL 54835, BPI) [II–III] ; TucumaT n : San Pedro de Colalao, on C. formosa, 1 Apr. 1994, J. F. Hennen

Ravenelia in Argentina & M. M. Hennen 094-054 (LIL 54836, BPI) [II–III] ; 2 Apr. 1994, on C. formosa, J. F. Hennen & M. M. Hennen 094-073 (LIL 54837, BPI 841034) [II–III]. Bolivia : Sorata : 14 Apr. 1920, on C. laxa, E. W. D. & M. M. Holway 520 (BPI 1495128) [II–III]. Guatemala : Palin : Amititlan, 24 Dec. 1916, on C. gracilis, E. W. D. Holway 633 (BPI 149524) [II]. Mexico : Sinaloa : Mex. Hyw 40, 4 miles E. of Concordia, 4 Dec. 1970, on C. penduliflora, G. B. Cummins 70-207 (BPI 196226) [III].

This is the first report of R. echinata var. ectypa for Argentina. Ravenelia ectypa was described as a species distinct from R. echinata (Arthur 1918), but Cummins (1977) considered R. ectypa a variety of R. echinata, both of which occur on species of Calliandra. Ravenelia echinata var. ectypa produces larger urediniospores and smaller teliospores than R. echinata var. echinata. Both varieties are reported in South, Central, and North America, although not in the United States or Canada. Ravenelia hieronymi Speg., Anales Soc. Ci. Argent., Pug. IV, 12 : 66 (1881). (Figs 14–15, 35–39) Type : Argentina : CoT rdoba : Sierra Chica, San Jose! , near La Calera, on Acacia caven, 11 Jan. 1877, Hieronymus s.n. (LPS 4930 – holotype ; NY 28492 n.v. – isotype) [I–III]. Pleoravenelia hieronymi (Speg.) Long, Bot. Gaz. (Crawfordsville) 35 : 127 (1903). Cystingoephora hieronymi (Speg.) Arthur, N. Am. Fl. 7 : 131 (1907). Ravenelia mimosae Henn., Hedwigia 34 : 95 (1895). Type : Brazil : GoiaT s : Paranahyba, on Acacia farnesiana (originally reported as Mimosa sp.), July 1892, E. Ule 1942 (NY 28462 n.v. – isotype). Ravenelia acaciae-farnesianae Henn., Hedwigia 34 : 321 (1895). Type : same as Ravenelia mimosae. Pleoravenelia deformans Maubl., Bull. Soc. Mycol. France 22 : 73 (1906). Type : Mozambique : Mhalume, on Acacia sp. (? A. arabica), LeTestu 7616 (PUR 69001 – isotype). Ravenelia deformans (Maubl.) Dietel, Beih. Bot. Centralbl. 20 : 404 (1906). Cystingophora deformans (Maubl.) H. Syd., Ann. Mycol. 19 : 165 (1921). Anamorph : Aecidium hieronymi Speg., Anales Soc. Ci. Argent., Pug. IV, 12 : 78 (1881). Type : Argentina, CoT rdoba : Sierra Chica, near San Jose! , on A. caven, 11 Jan. 1877, Hieronymus s.n. (LPS 9771 n.v. – holotype). Spermogonia observed once (Pitta 76-310). Mycelium systemic. Aecia densely scattered over witches’ brooms on young branches, conspicuous, with long cylindrical peridia, peridial cell walls striate-verrucose ; aeciospores (18–)20–26(–28)i(13–)15–20 µm, catenulate, variable in size and shape, often angular, mostly broadly ellipsoid or oblong-ellipsoid, pale yellow or pale golden in mass, wall 2–3 µm thick, densely and finely verruculose, pores 7–10, scattered, relatively obscure. Telia scattered among or produced in aecia, sub-

962 epidermal becoming erumpent, blackish brown ; teliospores (65–)75–120(–140) µm diam., clear chestnut brown or dark golden-brown, smooth, variable in shape, flat or often concave on the under side and strongly convex on the upper side, thus appearing to be thick centrally, 5–9(–12) probasidial cells across but the number difficult to determine at times, two layers of probasidial cells (as observed in longitudinal section), probasidial cells (18–)21–25(–27)i(14–)16–22 µm diam., one germ slit per probasidial cell, pairs of germ slits observed on adaxial spore surface, one belonging to upper probasidial cell and the other to lower probasidial cell which extends to the spore surface through the upper layer of probasidial cells ; cysts apparently uniseriate, of same number as peripheral probasidial cells, appressed to underside of the spore, coherent, but often swelling to appear pendent ; pedicel multihyphal, colourless, usually deciduous. Hosts and distribution : On Acacia caven (syn. A. cavenia), Argentina, Brazil, Chile, Paraguay, Uruguay ; on A. farnesiana, Brazil, Chile, Uruguay ; on Acacia sp., Mozambique. Other specimens examined : Argentina : Catamarca : Rio Las Juntas, camping area, 28m 25n949h S, 65m 50n739h W, 23 Sept. 1997, on Acacia caven, J. F. Hennen & J. R. HernaT ndez 097038 (BPI) [I] ; W. of Catamarca, 28m 22n488h S, 65m 52n683h W, 23 Sept. 1997, on A. caven, J. F. Hennen & J. R. HernaT ndez 097-044 (BPI 841188) [III] ; CoT rdoba : Cuesta del Cerro, 18 July 1876, on A. caven, G. Hieronymus s.n. (BPI 149785) [I–III] ; TucumaT n : SW of J. B. Alberdi, 27m 40n633h S, 65m 46n824h W, 22 Sept. 1997, on A. caven, J. F. Hennen & J. R. HernaT ndez 097-006 (BPI 841165) [I–III] ; J. F. Hennen & J. R. HernaT ndez 097-007 (BPI 841166) [I–III] ; J. B. Alberdi, S. of Escaba Dike, 28 Nov. 1997, on A. caven, J. R. HernaT ndez 097-108 (BPI 841224) [I–III]. Brazil : Sa4 o Paulo, city of Sa4 o Paulo, Rua Timor 96 (Ibirapuera), 24 Aug. 1976, on A. farnesiana, G. B. Pitta 76-310 (Hennen, pers. herb.). Chile : Bang os de Apoquindo, 1905–1906, on A. caven, R. Thaxter s.n. (BPI 149781) [I–III] ; Santiago : Taltil, May 1920, on A. caven, M. R. Espinosa 4 (BPI 149782) [I–III] ; Bang os de Cauquenes : 13 Jan. 1920, on A. farnesiana, E. W. D. Holway & M. M. Holway 295 (BPI 149791) [I–II]. Uruguay, Dept. Artigas : Loc. Santa Rosa Cuareim, Nov. 1927, on A. farnesiana, Herter 82584 (BPI 149789) [I–III].

Spegazzini (1881 : 73) published the anamorph name Uredo hieronymi and listed Ravenelia hieronymi as the ‘ status stylosporicus. ’ Five pages later (op.cit. : 78) he published Aecidium hieronymi and indicated R. hieronymi as the ‘ status hymeniferus. ’ The name that refers to the anamorph of R. hieronymi is A. hieronymi (Lindquist 1954). Uredo hieronymi is an anamorph of R. australis (Lindquist 1954). R. acaciae-farnesianae was an unnecessary new name for R. mimosae. Hennings (1895) made this change because the host, originally identified as Mimosa, was reidentified as Acacia farnesiana. Additionally, Hennings mistakenly described the anamorph of R. mimosae as uredinial. R. hieronymi produces anamorph sori and telia on witches’ brooms and branches. We consider this an

J. R. Herna! ndez and J. F. Hennen aecial stage because spermogonia were observed once and morphologically the anamorph is an Aecidium. The role of the Aecidium spores in the life cycle is only speculative. Presumably they could be aeciospores, urediniospores, or teliospores, or even a combination of these depending on the environmental conditions. The aecial sori of R. hieronymi are abundant, longcylindric, and yellowish, and because of their large size, are conspicuous compared to the other species of Ravenelia causing witches’ brooms on Acacia in the region of Northwest Argentina (Figs 15, 35). R. subtortuosae Long 1921, R. australis, and R. hieronymi appear to be closely related as evidenced by their systemic anamorph (Aecidium) infections causing witches’ brooms, cupulate anamorph sori with peridial cells, catenulate spores, and teliospores with two cell layers. The peridium in the Aecidium sori of R. australis is often poorly developed. The oblique distal cell walls of the lower level teliospore cells of R. hieronymi extend up between the cells of the upper level to the outer surface of the spore. Two germ slits occur in this region, one from the lower cell and one from the upper cell. These can be seen as a pair of slits when the spore is viewed from the top (Fig. 39). This teliospore feature distinguishes R. hieronymi from R. subtortuosae and R. australis. Dietel (1906) suggested a strong relationship between R. deformans, known from Africa (Mozambique), and R. hieronymi reported from South America. We confirm that R. deformans is a synonym of R. hieronymi based on the morphology of the teliospores and the observation of spermogonia in a collection from Brazil (Pitta 76-310). R. hieronymi is found in the Southern Hemisphere of the New World. Lindquist (1954) listed specimens from Argentina, Chile, Paraguay, and Uruguay, and Hennen, Hennen & Figueiredo (1982) listed this species from Brazil. Although Arthur (1907) included Mexico and the USA in the distribution of R. hieronymi, those collections from North America are actually Spumula heteromorpha J. W. Baxter 1966. The witches’ brooms on which anamorph spores of R. hieronymi and S. heteromorpha are produced are similar macroscopically. However, R. hieronymi also produces telia on the witches’ brooms whereas S. heteromorpha produces telia on leaves. Additionally, S. heteromorpha produces spermogonia with aecia on the witches’ brooms (Arthur 1907, Cummins 1978). We examined type material of S. heteromorpha as well as specimens identified as R. hieronymi from North America and Mexico housed in BPI. That Spumula heteromorpha has spermogonia and aecia on witches’ brooms, and telia on either or both leaf surfaces (Cummins 1978), as we observed on BPI collections, confirms our conclusion that the collections of R. hieronymi from Mexico and USA actually are S. heteromorpha. Specimens examined of S. heteromorpha : Mexico : Durango : Highway 45, South of Durango, on Acacia sp., 28 Nov. 1963,

963 Cummins 63-617 (BPI 150955, type) [III] ; Cummins 63-622 (BPI 150954) [0–I]. Specimens examined of S. heteromorpha identified as Ravenelia hieronymi in BPI ; Durango : Tepehuanes, 4–25 July 1906, on A. farnesiana, E. Palmer 267 (BPI 149799) [0–I] ; Hidalgo : Valley of Tula, 2 July 1901, on A. farnesiana, C. G. Pringle 19 (BPI 149793) [0–I] ; Morelia : 22 Sept. 1893, on A. farnesiana, C. G. Pringle s.n. (BPI 149788) [I] ; (BPI 149790) [I] ; (BPI 149797) [0–I] ; 1848–1849, on Acacia sp., J. Gregg 852 (BPI 149778) [I]. USA : Texas : Eagle Pass, 18 July 1931, on A. farnesiana, Gardner s.n. (BPI 149800) [I].

Ravenelia indigoferae Tranzschel, in Dietel, Hedwigia 33 : 369 (1894). (Figs 40–44) Type : Mexico, Jalisco : Rocky hills near Guadalajara, 1 Oct. 1889, on Indigofera palmeri, Pringle s.n. (LE 75241 – holotype, S F21394 – isotype) [II–III]. Pleoravenelia indigoferae (Tranzschel) Long, Bot. Gaz. 35 : 129 (1903). Ravenelia schroeteriana Henn., Hedwigia 35 : 245 (1896). Type : Argentina : Salta : Campo Santo, Apr. 1873, on Indigofera sp., Lorentz & Hieronymus s.n. (BPI 149966 – isotype) [II–III]. Ravenelia antiguana Cummins, Bull. Torrey Bot. Club 67 : 608 (1940). Type : Guatemala : Dept. Sacatepeteguez : near Antigua, on Indigofera sp. (originally identified as Cassia biflora), Nov. 1938–Feb. 1939, Standley 63356 (PUR 49070 – holotype). Anamorph : Uredo anilis Henn., Hedwigia 38 : 68 (1899). Type : Brazil : Santa Catarina : Sa4 o Francisco, on Indigofera anil, May 1884, Ule 145 (S F21398 – lectotypus hic designatus). Spermogonia few in a group, amphigenous, subcuticular, type 7. Aecia mostly hypophyllous and on rachis’, usually circinately grouped around the spermogonia, otherwise as the uredinia. Uredinia amphigenous or mostly hypophyllous, subepidermal becoming erumpent, yellowish brown ; paraphyses mostly capitate, 80–100 µm long, 15–30 µm diam. apically, wall brown apically and mostly 2–3 µm thick, colourless and thinner below ; urediniospores (20–)21–25(–27)i (18–)19–21(–24) µm, mostly broadly ellipsoid, golden or pale cinnamon-brown, wall 1n5(–2) µm thick, echinulate, pores 9–12, scattered. Telia mostly hypophyllous, but also on rachis’ and stems, subepidermal, becoming erumpent, blackish ; teliospores (70–)85– 110(–130) µm diam., chestnut-brown, without a thin, pale outer layer, (4)5–7(8) probasidial cells across, each cell with 3–7(–10) cylindrical tubercles 3–6 µm long, central probasidial cells (18–)20–27(–30) µm across, central probasidial cells 2-layered ; cysts uniseriate of same number as peripheral probasidial cells, appressed to spore ; pedicel multihyphal, colourless, deciduous. Hosts and distribution : On Indigofera anil, Brazil, Cuba, Puerto Rico ; on I. conzattii, Mexico ; on I. cuernavacana, Mexico ; on I. emarginella, Malawi, Tanzania, Uganda ; on I. galegoides, China ; on

Ravenelia in Argentina

964

Figs 40–44. Ravenelia indigoferae on Indigofera sp. Fig. 40. Uredinium with a teliospore emerging from the center of the sorus (BPI 149978). Fig. 41. Echinulate urediniospores and smooth, capitate paraphyses (BPI 149558). Fig. 42. Detail of echinulate urediniospore (BPI 149958). Fig. 43. Telium (BPI 149958). Fig. 44. Detail of teliospore with cylindrical tubercles on probasidial cells. Figs 45–46. Ravenelia lonchocarpicola var. lonchocarpicola on Lonchocarpus sp. Fig. 45. Surface view of teliospores with twice the number of peripheral as central probasidial cells (LPS 4926). Fig. 46. Median and surface view of teliospores (LPS 4926). Figs 47–51. Ravenelia macrocarpa on Cassia sp. and Senna sp. Fig. 47. Subcuticular uredinium with urediniospores intermingled with clavate to capitate paraphyses (BPI 841195). Fig. 48. Echinulate urediniospores with fewer echinulations or nearly smooth on proximal " of spore. Fig. 49. Subcuticular $ telium, note pedicels from which teliospores have become detached (BPI 191302). Fig. 50. Teliospores with smooth probasidial cells and globoid, pendent cysts (BPI 191302). Fig. 51. Teliospores (NY 31520). Figs 40–44, 47–50 SEM. Figs 45–46, 51 LM. Bars : Fig. 49 l 80 µm ; Fig. 51 l 22 µm ; Figs 40–41, 43, 47, 50 l 20 µm ; Figs 45–46, 48 l 13n3 µm ; Fig. 44 l 10 µm ; Fig. 42 l 3 µm.

J. R. Herna! ndez and J. F. Hennen I. mucronata, Brazil, Dominican Republic, Guatemala ; on I. palmeri, Brazil, Mexico ; on I. sphaerocarpa, Mexico, USA ; on I. subulata, Colombia ; on I. suffruticosa, Argentina, Barbados, Brazil, Colombia, Cuba, Dominican Republic, England, Guatemala, Jamaica, Puerto Rico, Trinidad & Tobago, Venezuela, Virgin Islands ; on I. tinctoria, China, Cuba ; on Indigofera sp., Argentina, China, Cuba, Dominican Republic, Guatemala, Malawi, Puerto Rico, Republic of South Africa, Uganda, USA, Virgin Islands. Other specimens examined : Cuba : Baracoa : 9 Mar. 1903, on Indigofera sp., leg. ? (BPI 149963) [I–III]. Guatemala : Panajachel : Solola, 3 Jan. 1917, on Indigofera sp., E. W. D. Holway s.n. (BPI 149958) [II–III]. Mexico : Cuernavaca : 30 Oct. 1903, on I. cuernavacana, Holway 5296 (BPI 149978) [II–III].

Lindquist (1954) listed this species as R. schroeteriana but indicated that it was similar to R. indigoferae. However, Lindquist (1982) considered R. schroeteriana a synonym of R. indigoferae, which the results of this study support. Ravenelia lonchocarpicola Speg., Rev. Argent. Bot. 1 : 131 (1925). var. lonchocarpicola (Figs 45–46) Type : Argentina : Salta : Ora! n, on Lonchocarpus nitidus, 11 Feb. 1905, Spegazzini s.n. (LPS 1926 n.v. – holotype ; LPS 4926 – isotype) [0–III]. Spermogonia epiphyllous, grouped, subcuticular in origin, 100–120i70–80 µm, without periphyses. Aecia and uredinia not produced. Telia mostly epiphyllous around the spermogonia, subepidermal becoming erumpent, blackish-pale, without paraphyses ; teliospores (50–)60–80(–90) µm diam., dark chestnut-brown with no discrete, paler, outer wall layer, smooth, 3–6 probasidial cells across with twice as many peripheral cells as central cells, cells 1-layered, 18–24 µm diam., cuneiform ; cysts uniseriate, of same number as peripheral probasidial cells, coherent and adherent to spores from margin to pedicel, conspicuous, as thick as the probasidial cells when inflated ; pedicel multihyphal, hyaline, mostly persistent, to 100–120 µm long. Hosts and distribution : On Lonchocarpus lilloi, Argentina (Jujuy) ; on L. nitidus, Argentina (Salta). Other specimens examined : Argentina : Jujuy : Ledesma, from Rı! o Agua Negra to Alba de Can4 as, 29 May 1977, on Lonchocarpus lilloi, Legname & Cuezzo 8247c [III] ; Ledesma, between Valle Grande to Rı! o Agua Negra, 8 Apr. 1971, on L. lilloi, Legname & Cuezzo 7970c [0–III].

Spegazzini (1925) differentiated R. lonchocarpicola from R. lonchocarpi Lagher. 1894, because the first is microcyclic, while the second is macrocyclic. Ravenelia mera Cummins 1943 is similar to R. lonchocarpicola, but the former is differentiated by its smaller probasidial cells. Baxter (1968) reduced R. mera to a variety of R. lonchocarpicola and also described R. lonchocarpicola var. robusta J. Baxter. The two varieties were established on the basis of differences in pedicel length, diameter of teliospores, thickness of teliospore

965 walls, and diameter of the spermogonia (Baxter 1968). Neither variety is known from Argentina. Spegazzini (1925) published this name as R. lonchocarpicola but Lindquist (1954, 1982), and Baxter (1968) spelled this epithet ‘ lonchocarpiicola ’. According to Article 60n1 ‘ The original spelling of a name or epithet is to be retained, except for the correction of typographical or orthographical errors and the standardizations imposed by Art. 60n5, … ’. This case does not fit those exceptions and therefore the epithet should be ‘ lonchocarpicola ’. Ravenelia macrocarpa Syd. & P. Syd., Ann. Mycol. 1 : 329 (1903). (Figs 47–51) Type : Brazil, on Cassia bicapsularis, F. Sellow s.n. (S F21392 – lectotypus hic designatus ; S F21393, NY 31520 n.v., PUR – isolectotypi) [II–III]. Haploravenelia macrocarpa (Syd. & P. Syd.) Syd., Ann. Mycol. 19 : 165 (1921). Anamorph : Uredo cyclogena Speg., Anales Soc. Ci. Argent. 9(4) : 172 (1880). Type : Argentina : Buenos Aires : Capital Federal, Boca del Riachuelo, on Senna corymbosa, Feb. 1880, Spegazzini s.n. (LPS 9408 – holotype ; BPI 154597 [C. Spegazzini exsiccata, Hongos Sud-Americanos, no. 21.] BPI, PUR F2119 – isotypes) [II]. Spermogonia and aecia not seen. Uredinia amphigenous, mostly hypophyllous, in concentric circles on small, circular, pale cinnamon-brown chlorotic spots, subcuticular becoming erumpent ; paraphyses hyaline, clavate to capitate, thick-walled ; urediniospores (20–)22–27(–30)i(15–)18–20(–22) µm, mostly obovoid or broadly ellipsoid, wall 2n5–3(–3n5) µm thick, pale cinnamon-brown or golden brown, echinulate but less so to nearly smooth on proximal 1\3 of spore, pores 6–8(10), mostly 8, tending to be bizonate around the equator. Telia amphigenous, subcuticular becoming erumpent, blackish brown ; teliospores (80–)93–120(–145) µm diam., dark chestnut-brown, smooth, with no discrete outer wall layer, (6)7–10(11) probasidial cells across, central probasidial cells (13–)17–18(–20)i(12–)13–17 µm, variable both in size and shape, cells in one layer ; cysts multiseriate, numerous, globoid, pendent ; pedicel multihyphal, fragile, deciduous. Hosts and distribution : On Cassia bicapsularis (syn. Senna pendula), Brazil ; on C. coluteoides, Brazil ; on C. corymbosa (syn. S. corymbosa) ; Argentina ; on C. hookeriana, Argentina ; on C. laevigata, Brazil ; on Cassia sp., Brazil ; on Senna araucarietorum, Brazil ; on S. birostris, Argentina ; on S. birostris var. hookeriana, Argentina ; on S. hookeriana, Argentina ; on S. pendula (syn. C. pendula), Brazil ; on S. pendula var. glabrata, Brazil ; on S. subulata, Argentina ; on S. tropica (syn. C. tropicana), Brazil ; on substrate indet., Argentina. Other specimens examined : Argentina : Catamarca : E. of La Vin4 a, 28m 03n735h S, 65m 35.324h W, 24 Sept. 1997, on Senna

Ravenelia in Argentina subulata, J. F. Hennen & J. R. HernaT ndez 097-056 (BPI 841195) [II] ; TucumaT n : Dept. Trancas, Cuesta Grande, 22 Apr. 1926, on S. birostris var. hookeriana, S. Venturi s.n. [III] ; Tafı! del Valle, 6 Apr. 1993, on S. birostris var. hookeriana, J. F. Hennen & J. R. HernaT ndez 093-117 (LIL 54838, BPI) [II–III] ; Tafı! , 19 Mar. 1944, on S. hookeriana, A. Courteig 516 [II–III] ; Tafı! del Valle, Apr. 1950, on Cassia hookeriana, R. Singer s.n. (BPI 191302) [III] ; (BPI 191303) [III] ; Dept. Tafı! , 7 Feb. 1924, on S. subulata, S. Venturi s.n. [II–III] ; San Javier, Parque Sierras de San Javier, 23 July 1994, on S. subulata, J. R. HernaT ndez 094-156 (LIL 54839, BPI) [II]. Brazil : Rio de Janeiro : Sylvestre, 15 Aug. 1921, on C. bicapsularis, E. W. D. & M. M. Holway 1038 (BPI 191301) [II].

The urediniospores of Uredo cyclogena are characterized by the bizonate germ pores in two wide supraand subequatorial areas, and the echinulate ornamentation with an area less so to nearly smooth on the proximal " of spore of the spore wall (Fig. 48). $ This is the first time the teleomorph and anamorph stages of this rust have been connected. This also is the first record of the teleomorph R. macrocarpa in Argentina. Ravenelia mimosae-sensitivae Henn., Hedwigia 35 : 246 (1896). (Figs 52–55) Type : Argentina : TucumaT n, on Mimosa sensitiva, Lorentz s.n. (B – holotype ; BPI 191437 – isotype) [II– III]. Ravenelia mimosae-caeruleae Dietel, Bot. Centralbl., Beib. 20 : 378 (1906). Type : Mexico : Morelos : Cuernavaca, on Mimosa caerulea, 30 Oct. 1903, Holway 5290 (BPI 191411 – lectotypus hic designatus ; BPI 191413, BPI 191410, isolectotypi ; NY 31524 n.v., NY 31525 n.v., syntypi) [II–III]. Ravenelia mimosae-albidae Dietel, Beih. Bot. Centralbl. 20 : 378 (1906). Type : Mexico : Morelos : Cuernavaca, on Mimosa albida var. floribunda, 24 Sept. 1898, Holway 3125 (BPI 191398 – isotype) [II–III]. Ravenelia mimosicola Arthur, N. Am., Fl. 7 : 137 (1907). Type : Mexico : Guerrero : Iguala, on Mimosa stipitata, 4 Nov. 1903, Holway 5326 (BPI 191447 – lectotypus hic designatus ; BPI 191449, PUR 6269 – isolectotypi) [II–III]. Anamorph : Uredo sensitiva Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 6 : 236 (1898). Type : Argentina : TucumaT n, Valley of Rio Chico near Tucuma! n, on M. sensitiva, Jan. 1895, Spegazzini s.n. (LPS 9455 – holotype) [II]. Ravenelia sensitiva (Speg.) Speg., Revista Argent. Bot. 1 : 132 (1925). Uredo assumptionis Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 31 : 392 (1922). Type : Paraguay : near Asuncio! n (Recoleta and Trinidad), on Mimosa balansae, July 1919, Spegazzini s.n. (LPS 9504 – holotype) [II].

966 Ravenelia mimosae-pudicae F. Kern, Thurst. & Whetzel, Mycologia 25 : 483 (1933). Type : Colombia : Itagui, on Mimosa pudica, 17 Aug. 1930, Archer H-108 (PAC n.v. – holotype) (described only an anamorph). Spermogonia and aecia unknown. Uredinia amphigenous, and on pods, subcuticular, erumpent, 1n5–2 mm diam., scattered or confluent, yellowish brown ; paraphyses golden brown, abundant, mostly spathulate, or clavately capitate, 8–12(–20) µm wide and 45–55 µm long, the apical wall to 8 µm thick, the stalk usually solid ; urediniospores (17–)18–21(–25)i(13–)14– 18(–21) µm, mostly broadly ellipsoid, wall 1n5–2n5 µm thick, golden or pale cinnamon-brown, closely verrucose-echinulate with hub and spoke pattern, pores 8–10, scattered. Telia as the uredinia except blackish brown and without paraphyses ; teliospores (55–)50– 90(–100) µm diam., (3)4–5(7) probasidial cells across, chestnut-brown with an outer thin but discrete pale layer bearing more or less cylindrical tubercles 2n5–3 µm wide and (2–)5–7 µm long, with (3)4–8(–10) tubercles on each probasidial cell, all probasidial cells in one layer, triangular or pentagonal, wall 4–5 µm thick at the top, central cells (16–)19–25(–28) µm diam. ; cysts uniseriate of same number as peripheral probasidial cells, not coherent, pendent ; pedicel multihyphal, short, colourless, deciduous. Hosts and distribution : On Mimosa albida, Costa Rica, Guatemala, Mexico, Peru ; on M. albida var. floribunda, Guatemala, Mexico, Venezuela, unknown ; on M. argentinensis, Argentina ; on M. balansae, Paraguay ; on M. biuncifera, Mexico : on M. caerulea, Mexico ; on M. debilis var. debilis, Argentina ; on M. floribunda, Ecuador, Venezuela ; on M. galeottii, Mexico ; on M. leptocarpa, Mexico ; on M. polyanthoides, Mexico ; on M. pudica, Colombia ; on M. sensitiva, Argentina, Colombia, Venezuela ; on M. sinaloensis, Mexico ; M. stipitata, Mexico ; on Mimosa sp., Costa Rica, Guatemala, Mexico ; on Schrankia distachya, Mexico. Other specimens examined : Argentina : Salta : between Rosario de la Frontera and Horcones, 11 Apr. 1994, on Mimosa debilis var. debilis, J. F. Hennen & L. D. Ploper 094120 (LIL 54843, BPI 841052) [II–III] ; TucumaT n : Las Cejas, 30 Mar. 1993, on M. debilis var. debilis, J. F. Hennen & J. R. HernaT ndez 093-073 (LIL 54840, BPI) [II–III] ; Dept. Cruz Alta, San Agustı! n, Monte Redondo, 29 Mar. 1994, on M. debilis var. debilis, J. F. Hennen, M. M. Hennen & J. R. HernaT ndez 094-027 (LIL 54841, BPI) [II–III] ; 30 Mar. 1994, on M. debilis var. debilis, J. F. Hennen, M. M. Hennen & J. R. HernaT ndez 094-035 (LIL 54842, BPI) [II–III] ; Mexico : Oaxaca : 10 Nov. 1903, on M. albida, E. W. D. Holway 5868 (BPI 191425) [II–III] ; Chihuahua ; 22 Miles West of Chihuahua, 21 Oct. 1923, on Mimosa sp., G. B. Cummins 63403 (BPI 191417) [III].

Ravenelia mimosae-sensitivae is found throughout the neotropics from the subtropical areas of Argentina to Mexico (Lindquist 1982). The SEM photomicrographs illustrate the hub and

J. R. Herna! ndez and J. F. Hennen

967

Figs 52–55. Ravenelia mimosae-sensitivae on Mimosa sp. Fig. 52. Subcuticular uredinium (Holway 3528). Fig. 53. Detail of hub and spoke pattern of ornamentation on urediniospore (Holway 3528). Fig. 54. Subcuticular telium (LIL 54843, BPI 841052). Fig. 55. Detail of proximal side of teliospore with pendent cysts (Holway 3528). Figs 56–60. Ravenelia monosticha on Acacia sp. Fig. 56. Subepidermal uredinium with abundant peripheral paraphyses (BPI 191457). Fig. 57. Detail of sparsely echinulate urediniospore with 4–5 equatorial germ pores (BPI 191457). Fig. 58. Median view of urediniospore (BPI 191456). Fig. 59. Surface view of urediniospore in Fig. 58. Fig. 60. Surface view of inconspicuously ornamented teliospores (PUR 2090). Figs 61–63. Ravenelia oligotheles on Cathormion sp. Fig. 61. Polyhedric-ellipsoidal, echinulate urediniospores. Fig. 62. Teliospore with conspicuous conical or apically rounded ornamentation. Fig. 63. Teliospore in lower right corner showing cysts and pedicel on proximal side. Figs 52–57, 61–63 SEM. Figs 58–60 LM. Bars : Figs 54, 56 l 20 µm ; Fig. 60 l 13n3 µm ; Figs 52, 55, 61–63 l 10 µm ; Figs 57–59 l 3 µm ; Fig. 53 l 1 µm.

Ravenelia in Argentina

968

Figs 64–65. Ravenelia oligotheles on Cathormion sp. Fig. 64. Surface view of teliospore (LPS 4940, type). Fig. 65. Median view of teliospore in Fig. 64. Figs 66–68. Ravenelia papillosa. Fig. 66. Anamorph galls on stems of Parapiptadenia sp. (BPI 841191). Fig. 67. Detail of anamorph gall (BPI 841191). Fig. 68. Teliospores (PUR 89579, type). Figs 69–70. Ravenelia platensis on Erythrina crista-gallii. Fig. 69. Spermogonial galls on leaflets (BPI 841204). Fig. 70. Spermogonial galls on deformed petioles (BPI 841204). Figs 71–72. Ravenelia prosopidiicola on Prosopis cfr. alpataco. Fig. 71. Median view of smooth teliospore with pendent cysts and pedicel on proximal side (LPS 13740, type). Fig. 72. Surface view of teliospore shown in Fig. 71. Figs 64–65, 68, 71–72 LM. Bar l 10 µm.

spoke pattern of ornamentation of the urediniospores (Figs 52–53). Also, the early subcuticular development of the telia can be seen on the distal layer of a very young sorus. The cylindrical tubercles on the teliospores become almost full size while the spore is immature (Fig. 54). The uniseriate cysts are pendent (Fig. 55). Ravenelia monosticha Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 31 : 379 (1922). (Figs 56–60) Type : Paraguay : AsuncioT n, on Acacia bonariensis, July 1919, Spegazzini s.n. (LPS 4943 – holotype) [II–III]. Anamorph : Uredo sp. Spermogonia and aecia unknown. Uredinia hypophyllous, subepidermal becoming erumpent, cinnamon-

brown ; paraphyses hyaline, abundant, peripheral, capitate, 25–35 µm long, 8–12 µm wide apically and apical wall to 5 µm thick ; urediniospores (23–)26– 30(–33)i(8–)12–14(–15) µm, ellipsoidal, elongate obovoid or fusoid, wall 1n5(–2) µm thick at sides, same or to 3 µm at apex, golden to pale yellowish or nearly hyaline, sparsely echinulate, pores equatorial 4–5(6). Telia mostly hypophyllous, without paraphyses, amber and shiny as seen with a stereoscope ; teliospores tending to be ellipsoidal, (50–)53–55i65–70 µm, 4 probasidial cells wide and 6 probasidial cells long, with a thin, colourless outer wall layer with 4–8 hyaline tubercles per probasidial cell that are sometimes obscure or lacking, especially in mature spores, probasidial cells 16–19i13–15 µm diam., 1-layered ; cysts uniseriate, of

J. R. Herna! ndez and J. F. Hennen

969

Figs 73–74. Ravenelia papillosa on Parapiptadenia sp. and Acacia sp. Fig. 73. Echinulate urediniospores and smooth paraphyses (BPI 841270). Fig. 74. Urediniospores with numerous scattered germ pores (BPI 841191). Figs 75–76. Ravenelia platensis on Erythrina sp. Fig. 75. Ostiolate spermogonium with spermatia (BPI 841204). Fig. 76. Echinulate urediniospores with smooth areas around germ pores on two flattened sides (BPI 841204). Figs 77–78. Ravenelia sarmientoi on Bulnesia sarmientoi. Fig. 77. Surface view of spinulose teliospores (PUR 89743). Fig. 78. Median view of teliospores shown in Fig. 77. Figs 79–84. Ravenelia spegazziniana on Acacia sp. Fig. 79. Urediniospores (BPI 191761). Fig. 80. Detail of hub and spoke pattern of ornamentation on urediniospores (BPI 191761). Fig. 81. Surface view of urediniospores with germ pores in two bands (LIL 54821, BPI 841005). Fig. 82. Median view of urediniospores shown in Fig. 81. Fig. 83. Smooth teliospore. Fig. 84. Clavate paraphyses and part of teliospore (BPI 191755). Figs 73, 75–76, 79–80, 83–84 SEM. Figs 74, 77–78, 81–82 LM. Bars : Figs 77–78 l 20 µm ; Figs 81–82 l 13n3 µm ; Fig. 74 l 12n5 µm ; Figs 73, 75, 83–84 l 10 µm ; Figs 76, 79–80 l 3 µm.

Ravenelia in Argentina same number as peripheral probasidial cells, coherent, appressed ; pedicel multihyphal, brownish next to the spore, colourless below, deciduous. Hosts and distribution : On Acacia bonariensis, Argentina, Paraguay ; on Acacia sp., Mexico. Other specimens examined : Mexico : Cd. Mante : Tamaulipas, 24 Sept. 1963, on Acacia sp., G. B. Cummins 63-165 (BPI 191456) [II] ; (BPI 191457) [II].

Spegazzini (1922) described the teliospores of R. monosticha as smooth (‘ … lisas y sin ape! ndices ’ …), whereas Lindquist (1954, 1982) described them as having 4–8 hyaline tubercles per probasidial cell, basing his description on the type collection from Paraguay and also on di Fonso collections from Argentina (Chaco) on the same host (LPS 12253, LPS 12263). We observed tubercles on many probasidial cells (Fig. 60), especially on young spores. In older spores the tubercles are often difficult to observe and the spores appear to be smooth, possibly because the outer, hyaline layer becomes thinner as the spore matures. Lindquist (1954, 1982) specifically stated that there were no uredinial paraphyses. However, Spegazzini (1922) had mentioned ‘ … ciertos corpu! sculos casi fusariformes bacilares ma! s o menos arqueados … (28–32i3– 4 µm) … ’ which could refer to paraphyses. We observed numerous capitate paraphyses in the type material (LPS 4943) and in the two collections on Acacia from Mexico (BPI 191456, BPI 191457). Ravenelia oligotheles Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 19 : 296 (1909). (Figs 61–65) Type : Argentina : Formosa, on Cathormion polyanthum (originally reported as Arthrosamanea polyantha), Jan. 1904, Spegazzini s.n. (LPS 4940 – holotype ; PUR F11240 – isotype) [III]. Anamorph : Uredo sp. Spermogonia and aecia not seen. Uredinia epiphyllous, small, cinnamon brown, subepidermal in origin, without paraphyses ; urediniospores (22–)23–27(–29)i (14–)17–20(–22) µm, more or less polyhedric-ellipsoid, with 3 small equatorial lobes each with one pore, wall 2 µm thick, golden brown, echinulate. Telia epiphyllous, blackish, 1–2 mm diam. ; teliospores (50–)55–70(–75) µm diam., rounded as seen from the top, hemispherical as seen from the side, typically 4 probasidial cells across (4 central and 6 peripheral probasidial cells) ; probasidial cells mostly 18– 22i17–36 µm diam., wall thicker at the apex, 3–5 µm, each central probasidial cell with 4–6 and each peripheral probasidial cell with 6–12 narrow cones or apically rounded spines 2–4 µm long, these on a thin but discrete, nearly colourless layer overlaying the chestnut-brown spore, cells 1-layered ; cysts uniseriate of same number as the peripheral probasidial cells, globose, appressed but swelling to appear semipendent, swollen slightly in water ; pedicel hyaline, of few coarse hyphae, usually deciduous.

970 Hosts and distribution : On Cathormion polyanthum (syns A. polyantha, Pithecellobium multiflorum), Argentina (Chaco, Formosa), Paraguay, Uruguay. Other specimens examined : Argentina : Chaco : Resistencia, on Cathormion polyanthum, 15 Nov. 1943, Di Fonzo 346 (LPS 11960).

Only teliospores were found in the type collection. The description of uredinia and urediniospores was based on a collection from Chaco (LPS 11960) and Lindquist (1954). The three small, sometimes indistinct equatorial bulges or lobes associated with the pores in the urediniospore walls are characteristic of this species. Ravenelia oligotheles, R. echinata var. ectypa and R. lonchocarpicola var. lonchocarpicola are examples of rust species with a consistent and characteristic number of central and peripheral probasidial cells in the teliospores, and this number varies with the species (Fig. 64). This species is known from Argentina, Paraguay and Uruguay. A report of R. oligotheles from Costa Rica (Cummins & Stevenson 1956) is a mistake ; it is actually Ravenelia havanensis on Enterolobium cyclocarpum (Cummins 1978). Ravenelia papillosa Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 6 : 229 (1898). (Figs 66–68, 73–74) Type : Argentina : La Plata : horto botanico, on Acacia visco (originally erroneously identified as Albizia julibrissin), 8 June 1897, Spegazzini s.n. (LPS 4956 n.v. – holotype ; PUR 89579 – isotype) [III]. Anamorph : Uredo leguminicola Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 19 : 317 (1909). Type : Argentina : Buenos Aires : horto botanico bonaerensi, on A. visco [originally erroneously identified as Acacia lophantha, and reidentified by Spegazzini as Acacia platensis fide Lindquist (1954) which is a synonym of A. visco], 6 Mar. 1906, E. Autran s.n. (LPS 4001 – holotype) [II]. Spermogonia and aecia not seen. Mycelium systemic. Uredinia amphigenous and on rachis’ and branches on conspicuous galls and witches’ brooms, subepidermal, rounded, becoming erumpent, brown, 0n5–1 mm diam. on leaves, confluent and larger on abnormal growths ; paraphyses yellowish to brown, intermixed, numerous, clavate or clavate-capitate, 40–50i14–16 µm, pedicel 5 µm across, thick-walled ; urediniospores (18–)20– 24i(12–)14–19 µm, broadly ellipsoid or broadly obovoid, wall pale brownish or golden, 2n5–3 µm thick, echinulate, pores 10–12, scattered. Telia following uredinia or formed de novo, blackish ; paraphyses as in the uredinia ; teliospores 88–120(–130) µm diam., dark chestnut-brown, 7–10 probasidial cells across, wall 7–8 µm thick at the top, with 3–5 cones or conical papillae 2–3 µm high on each probasidial cell, especially obvious peripherally, cells 1-layered ; cysts coherent, hyaline, uniseriate, of same number as peripheral

J. R. Herna! ndez and J. F. Hennen probasidial cells, swollen in water ; pedicel multihyphal, hyaline, deciduous. Hosts and distribution : On Acacia visco, Argentina (Buenos Aires) ; on Parapiptadenia excelsa, Argentina (Catamarca). Other specimens examined : Argentina : Catamarca : Dept. Ambatos, El Rodeo, El Nogalal, 14 Nov. 1999, on Acacia visco, J. R. HernaT ndez 099-001 (BPI 841270) [II] ; J. R. HernaT ndez 099-009 (BPI 841277) [II] ; Dept. Ambatos, El Rodeo, El Nogalal, 23 Sept. 1997, on Parapiptadenia excelsa, J. F. Hennen & J. R. HernaT ndez 097-049 (BPI 841191) [II–III] ; TucumaT n : Las Salas, 5 May 1995, on A. visco, J. R. HernaT ndez 095-019 (BPI) [II–III] ; Dept. Trancas, Vipos, 28 Mar. 1996, on A. visco, J. R. HernaT ndez 096-007 (BPI 841119) [II–III].

The only published records of this species were from La Plata (Lindquist 1954). There has been confusion over the types of symptoms produced by R. papillosa. Spegazzini did not mention the association of sori with abnormal growths in his original description of either the telial (Spegazzini 1898) or the anamorph stage (Spegazzini 1909). Lindquist (1954) reported the association of sori with witches’ brooms and leaves and stated ‘ Es una de las especies ma! s perjudiciales en el paı! s pues provoca la formacio! n de escobas de brujas con gran desmedro para el a! rbol. Todos los ejemplares cultivados en ciudad Eva Pero! n se hallan intensamente atacados. ’ Lindquist (1982) later cited both ‘ tumors ’ and witches’ brooms as symptoms, and illustrated tumors in his fig. 7. Those tumor-like symptoms look the same as those of our collections from Catamarca where we observed only conspicuous galls (Figs 66–67). Even though we did not observe witches’ brooms as mentioned by Lindquist, these collections from galls are microscopically identical to R. papillosa so we apply that name to our material. Parapiptadenia excelsa is a new host record for R. papillosa. Distribution and frequency of infection could not be determined because we have only one collection on this host. Ravenelia platensis Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 6 : 228 (1898). (Figs 69–70, 75–76) Type : Argentina : Buenos Aires : Santa Catalina, on Erythrina crista-galli, summer of 1890, Spegazzini s.n. (LPS 4958 – lectotype) [II–III]. Anamorph : Caeoma sp. Spermogonia subcuticular, amphigenous, also on stem and petiole galls, 75–115i40–70 µm. Aecia amphigenous and associated, often circinately, with the spermogonia, subepidermal becoming erumpent and often confluent, without paraphyses ; aeciospores pedicellate, (28–)30–40(–46)i(22–)24–29(–33) µm, flattened ellipsoidal or obovoid or occasionally reniform, wall (3–)4–5 µm thick, pale golden, strongly and rather sparsely echinulate, the spines 2–3 µm long, with smooth areas around pores on the two flattened sides, pores 3(4) equatorial. Uredinia and urediniospores similar to aecia and aeciospores. Telia also amphigenous or caulicolous on galls, blackish brown, roundish, with-

971 out paraphyses ; teliospores 100–120 µm diam., dark mahogany, smooth, 5–6 probasidial cells across, cells 1-layered ; cysts 15–20, adherent, circling the pedicel as a collar, swelling slightly in water ; pedicel hyaline, composed of numerous hyphae, persistent. Hosts and distribution : On Erythrina crista-galli, Argentina (Buenos Aires, Misiones), Brazil, Uruguay ; on Erythrina malungu, Brazil ; on Erythrina sp., Brazil. Other specimens examined : Argentina : Buenos Aires : La Plata, 29 Sept. 1997, on E. crista-galli, J. F. Hennen & J. R. HernaT ndez 097-073 (BPI 841204) [0]. Brazil : Rio de Janeiro : Therezopolis, 28 Sept. 1921, on Erythrina sp., E. W. D. Holway & M. M. Holway 1157 (BPI 191556) [II]. Uruguay : Montevideo, Nov. 1925, on E. crista-galli, Herter s.n. (BPI 191557) [I].

Spegazzini (1898) did not specify a type and listed the locality as ‘ … prope La Plata, in Santa Catalina, in Isla de Martı! n Garcı! a et in Chaco, per ann : 1883–98. ’ Lindquist (1954) cited the type as above which we consider to be a lectotype. Lindquist (1954) reported this rust fungus as widespread and recognizable from a distance because of the large galls on the branches. Lindquist & Garcı! a Zorro! n de Rosengurtt (1967) reported it from Montevideo, Uruguay and as widespread in the La Plata River area of Argentina and Uruguay. The galls produced by R. platensis appear to be annual. We observed numerous dead galls that had been produced the prior season in a city park in La Plata, Argentina in October 1997 (spring). No new aecial sori were forming on those galls but numerous spermogonia that were oozing honey-dew with a detectable odor were seen on young stems, petioles and leaflets, producing large hypertrophied areas (Figs 69–70) which would eventually develop into galls on which aecia are produced. The host is the ‘ national flower ’ of Argentina (Cabrera 1967), widespread in southern Brazil, Paraguay, and northeastern Argentina, and a widely cultivated ornamental that has various other folk uses. Ravenelia prosopidiicola J. C. Lindq., Revista Fac. Agron. Univ. Nac. La Plata 28 : 82 (1951). (Figs 71–72) Type : Argentina : San Luis : Alto Pencoso, on Prosopis cfr. alpataco, 17 Nov. 1944, Ruiz Leal 9600 (LPS 13740 – holotype) [0–I–III]. Anamorph : Aecidium prosopidicola Speg., Revista Argent. Bot. 1 : 98 (1925). Type : Argentina : San Luis, Piscu-yacu! , on Prosopis algarobilla (probably P. nigra), Nov. 1921, J. BretheT s s.n. (LPS 9835 – holotype) [I]. Spermogonia on deformed pods and woody stem galls, subcuticular, 70i30–35 µm. Aecia associated with the spermogonia, only on young pods, cupulate, with large, finely striate, verrucose peridial cells ; aeciospores catenulate, (25–)28–35(–38)i(15–)18–24(–26) µm, variable but mostly broadly ellipsoidal or oblong-ellipsoidal, wall 1–1n5 µm at sides, same or thicker at apex

Ravenelia in Argentina and sometimes at base, pale yellowish or nearly colourless, finely striate verrucose, pores 6–7, equatorial with slight caps, obscure. Uredinia apparently not produced. Telia subepidermal, becoming erumpent, densely grouped on large stem galls, light brown ; teliospores 65–90 µm diam., pale dark golden-brown or chestnut-brown with no discrete outer wall layer, smooth, 5–7 probasidial cells across, central probasidial cells variable mostly (14–)16–20(–23) µm diam., walls 2n5–3 µm at sides, 5–7 µm at the apex, germ pores obscure, cells 1-layered ; cysts globoid, pendent, uniseriate of same number as peripheral probasidial cells, colourless ; pedicel colourless, of few hyphae, 200 µm or more in length, persistent. Hosts and distribution (known only from Argentina) : On Prosopis alba, Santiago del Estero ; on P. algarobilla, San Luis ; on P. cfr. alpataco, Mendoza, San Luis ; on P. caldenia, La Pampa ; on P. chilensis, Catamarca ; on P. flexuosa, Mendoza ; on P. nigra, Mendoza, San Luis. The epithet prosopidicola was used by Spegazzini (1925) in the original description of the anamorph Aecidium prosopidicola. Lindquist (1951) described Ravenelia prosopidiicola, using a double ‘ i ’ and in the same and later papers (Lindquist 1954, 1958, 1982) referred to Aecidium prosopidicola as A. prosopidiicola, thereby changing the spelling of that epithet. According to Article 60n1 of the ICBN, Spegazzini’s epithet should be ‘ prosopidicola ’ and that of Lindquist ‘ prosopidiicola ’. Lindquist (1958) was the first to connect A. prosopidicola with R. prosopiidicola. Ravenelia prosopidiicola is morphologically similar to R. holwayi but can be distinguished by the fact that the teliospores of R. holwayi are larger (85–125 µm across) than those of R. prosopidiicola (65–90 µm across). Additionally, teliospores of R. holwayi have 7–12 probasidial cells across and it produces uredinia, whereas those of R. prosopidiicola have 5–7 probasidial cells across (Fig. 71) and uredinia have not been observed. Ravenelia sarmientoi J. C. Lindq., Hickenia 1 : 152 (1978). (Figs 77–78) Type : Argentina : Salta : Meta! n, on Bulnesia sarmientoi, 1 Apr. 1974, Cordo 5861 (LPS – holotype) [III]. Spermogonia, aecia and uredinia unknown, possibly not produced. Telia amphigenous, subcuticular becoming erumpent, chestnut-brown, without paraphyses ; teliospores 80–90 µm diam., 4(5) probasidial cells across, all spores 1-celled chestnut-brown overlaid with a pale, discrete, outer wall layer bearing 4–5 colourless spines on each probasidial cell ; cysts of same number as peripheral probasidial cells, appressed to the underside of the teliospore and adherent to each other, colourless ; pedicel colourless, short, multihyphal, fragile. Hosts and distribution : On Bulnesia sarmientoi, Argentina (Salta). Known only from the type. Ravenelia sarmentoi was reported on Bulnesia (Zygophyllaceae) (Lindquist 1978). It is likely that the host was misidentified since species of Ravenelia are

972 found mainly on Fabaceae. Lindquist stated that the type specimen was composed of only three leaves (‘ El material se compone de so! lo tres-hojas ’). Bulnesia leaves have two opposite leaflets somewhat similar to some species of Cassia. Therefore we suspect that the host is in fact Fabaceae, possibly in the Caesalpinoideae. Ravenelia spegazziniana J. C. Lindq., Bol. Soc. Argent. Bot. 1 : 298 (1946). (Figs 79–84) Type : Argentina : Buenos Aires : La Plata, Jardı! n Bota! nico de la Facultad de Agronomı! a (Eva Pero! n), on Acacia aroma, 16 June 1945, J. C. Lindquist (LPS 12604 – holotype) [II–III]. Anamorph : Uredo sp. Ravenelia siliquae Long, Bot. Gaz. (Crawfordsville) 35 : 118 (1903). Type : Mexico : Oaxaca : Etla, on A. farnesiana, 25 Oct. 1899, E. W. D. Holway 3841 (BPI 191725 – holotype) [II]. Haploravenelia siliquae (Long) H. Syd., Ann. Mycol. 19 : 165 (1921). Spermogonia unknown. Aecia on pods and deformed young branchlets, deep-seated, without peridium or with a few peridial cells, opening by a rift in the epidermis ; aeciospores catenulate, 30–36i12–18 µm, oblong fusoidal, pentagonal or irregularly polygonal, the base usually flat, the apex acute, densely echinulate with fine spines, pores 3 or 4, equatorial (from Lindquist, 1954). Uredinia on pods and leaflets, subcuticular in origin, in large, confluent groups on pods, small and discrete on leaflets, rachis’ and petioles, cinnamon-brown ; paraphyses few or numerous, uniformly thin-walled, intrasoral, mostly clavate, nearly colourless ; urediniospores (20–)23–28(–30)i(12–)14– 16(–18) µm, mostly ellipsoidal or narrowly obovoid, wall 1n5–2 µm thick at sides, 2–3 µm at apex, pale and rather dull cinnamon-brown, closely verrucose with fine basal connections, hub and spoke pattern of ornamentation, pores in 2 bands of 4 each, one above and one below the equator. Teliospores in uredinia, spores (65–)70–95(–100) µm diam., chestnut-brown with a thin but discrete pale outer layer, appearing smooth, (4)5–7 probasidial cells across, cells in one layer, central cells (13–)17–22(–24) µm across ; cysts uniseriate mostly of same number as peripheral cells, globoid, pendent ; pedicel colourless, multihyphal, deciduous. Hosts and distribution : On Acacia aroma, Argentina ; on A. catechu, USA ; on A. farnesiana (syn. Vachellia farnesiana), Cuba, Guatemala, Honduras, Mexico, Nicaragua, Puerto Rico, Virgin Islands, USA ; on A. pennatula, Mexico, unknown ; on A. smallii, Mexico ; on Acacia sp., Mexico. Other specimens examined : Argentina : Salta : Meta! n, 2 Apr. 1993, on Acacia aroma J. F. Hennen & L. D. Ploper 093103A (LIL 54823, BPI) [III] ; TucumaT n : Dept. Cruz Alta, San Agustı! n, 30 Mar. 1993, on A. aroma, J. F. Hennen & J. R. HernaT ndez 093-059A (LIL 54825, BPI) [III] ; J. F. Hennen &

J. R. Herna! ndez and J. F. Hennen

973

J. R. HernaT ndez 093-060 (LIL 54845, BPI) [II–III] ; J. F. Hennen & J. R. HernaT ndez 93-076A (LIL 54821, BPI 841005) [II] ; 30 Mar. 1994, on A. aroma, J. F. Hennen, M. M. Hennen & J. R. HernaT ndez 094-038 (LIL 54846, BPI) [III] ; Trancas, Vipos, 30 Mar. 1995, on A. aroma, J. R. HernaT ndez 095-044 (BPI 841087) [III] ; Monte Redondo, 31 Mar. 1996, on A. aroma, J. R. HernaT ndez 096-089 (BPI) [II–III]. Guatemala : Lake Amitlan : Laguna, 8 Feb. 1915, on A. farnesiana, E. W. D. Holway 199 (BPI 191761) [II]. Mexico : Guaymas : Sonora, 17 Jan. 1962, on Acacia sp., Cummins 62-20 (PUR 60086, BPI 191755) [III].

referred to as the ‘ hub and spoke ’ sculpture pattern. The ‘ hub and spoke ’ pattern of R. spegazziniana is different from that of other species of Ravenelia with this type of ornamentation : the ‘ hub ’ is flat topped with a central ‘ dimple ’ (Fig. 80). The ‘ hub and spoke ’ pattern of the eight other species of Ravenelia in the New World known to have that sculpture pattern (Lo! pez-Franco & Hennen 1990) has a ‘ hub ’ that is more or less pointed or narrowly rounded and subtended by one or two ring-like extensions. Ravenelia spegazzinia is similar to five of the other nine species of Ravenelia with the ‘ hub and spoke ’ pattern in that the urediniospore walls usually have two bands of germ pores (Figs 81–82). Long (1903) based his description of R. siliquae on the uredinial stage found on the collection made by H. W. D. Holway in 1899. According to Art. 59.2 (ICBN), R. siliquae can only be applied to an anamorph. We do not provide a new binary name for the uredinial anamorph of R. siliquae because the connection with the teleomorph is firmly established (Art. 59A.3) ; rather we refer to it as Uredo sp.

Ravenelia spegazziniana is known from Argentina (Buenos Aires, Co! rdoba, Salta, Santa Fe and Tucuma! n provinces), and also from some South, Central, and North American countries, all reported on species of Acacia. Teliospores are known only from the type (La Plata) and recent collections from Tucuma! n and Salta provinces, Argentina. As seen under SEM or using an oil immersion objective, the ornamentation of the urediniospore walls is distinctive in having knob-like processes interconnected by straight ridges (Gardner & Hodges 1985)

Key to species of Ravenelia in Argentina based on teliospores and anamorph spores 1

Teliospores smooth . Teliospores ornamented

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. 2 . 11

2(1)

Teliospores with fewer than 7 probasidial cells across Teliospores with more than 7 probasidial cells across

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

3(2)

Only telial and spermogonial stages known (microcyclic) . . . . .lonchocarpicola var. lonchocarpicola Another anamorph stage known in addition to telial and spermogonial stages . . . . . . . 4

4(3)

Anamorph spores (urediniospores) with hub-and-spoke type of ornamentation Anamorph spores echinulate or verrucose . . . . . . .

. .

. .

. .

. spegazziniania . . . 5

5(4)

Anamorph spores echinulate but smooth around the equatorial pores Anamorph spores not smooth around the equatorial germ pores .

. .

. .

. .

. .

. .

. .

6(5)

Anamorph sori with capitate paraphyses . Anamorph sori without paraphyses . .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. monosticha . prosopidiicola

7(2)

Only telial stage known (microcyclic) . . Telial and one or more anamorph stages known

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

chacoe$ nsis . . 8

8(7)

Teliospores with germ slits Teliospores with germ pores

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

hieronymi . . 9

9(8)

Uredinia with geniculate paraphyses Uredinia with capitate paraphyses .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

. australis . . 10

10(9)

Urediniospores with 6–8(–10) equatorial pores, most frequently eight, then tending to be bizonate . Urediniospores with 4–6 equatorial pores . . . . . . . . . . . .

macrocarpa . cebil

11(1)

Uredinia with paraphyses . Uredinia without paraphyses

. .

. .

. .

. .

. .

. .

. .

. .

. .

. platensis . . 6

. .

. .

. .

. .

. .

. .

. .

. .

12(11) Urediniospores with hub and spoke type of ornamentation Urediniospores echinulate . . . . . .

. .

. .

. .

. .

. .

. .

. mimosae-sensitivae . . . . 13

13(12) Urediniospores with 4 equatorial pores . Urediniospores with 9–12 scattered pores .

. .

. .

3 7

. .

. 12 . 15

. .

. .

. .

. .

. .

. .

. .

. .

. .

. .

argentinica . . 14

14(13) Urediniospores (20–)21–25(–27)i(18–)19–21(–24) µm Urediniospores (18–)20–24i(12–)14–19 µm . .

. .

. .

. .

. .

. .

. .

. .

. .

. .

indigoferae . papillosa

15(11) Urediniospores with 5–10 scattered pores . . . Urediniospores with fewer than 5 equatorial pores .

. .

. .

. .

. .

. .

. .

. .

.echinata var. ectypa . . . . 16

16(15) Urediniospores polyhedric-ellipsoid, with 3 equatorial lobes each with one pore Urediniospores ellipsoid to limoniform, with (3)4(–6) equatorial pore . .

. .

. .

. .

. .

oligotheles . cohniana

Ravenelia in Argentina A C K N O W L E D G E M E N TS We are grateful to the Herbaria BPI, LIL, LPS and PUR for the loan of collections, and S for providing type specimen information. Sincere thanks to Christian Feuillet for the Latin description and to Alberto Slanis (LIL) for host plant identification. The first author expresses his appreciation to Amy Y. Rossman for encouragement that contributed to the completion of this paper, and to Jim Plazkowitz for his help on photomicrographs and plates. Finally he thanks his wife, Mary E. Palm-Herna! ndez, for her invaluable motivation and support.

REFERENCES Arthur, J. C. (1907–27) Uredinales. North American Flora 7 : 83–848. Arthur, J. C. (1918) Uredinales de Guatemala based on the collections by E. W. D. Holway. American Journal of Botany 5 : 320–446. Baxter, J. W. (1968) The species of Ravenelia occurring on Lonchocarpus. Mycologia 60 : 42–48. Baxter, J. W. (1975) Notes on Brazilian species of Ravenelia. Mycologia 67 : 436–437. Cabrera, A. L. (1967) Flora de la Provincia de Buenos Aires. Parte III. Instituto Nacional de Tecnologı! a Agropecuaria, Argentina. Cooke, M. C. (1880) The genus Ravenelia. Journal of the Royal Microscopical Society 3 : 384–389. Cummins, G. B. (1977) A synopsis of the species of Ravenelia on Mexican Legumes. BoletıT n de la Sociedad Argentina de BotaT nica 18 : 83–90. Cummins, G. B. (1978) Rust Fungi on Legumes and Composites in North America. University of Arizona Press, Tucson. Cummins, G. B. & Hiratsuka, Y. (1983) Illustrated Genera of Rust Fungi. Revised edn. American Phytopathological Society Press, St. Paul, MN. Cummins, G. B. & Hiratsuka, Y. (1991) Illustrated Genera of Rust Fungi. Revised edn. American Phytopathological Society Press, St. Paul, MN. Cummins, G. B. & Stevenson, J. (1956) A check list of North American rust fungi (Uredinales). Plant Disease Reporter 240 (Suppl.) : 109–192. Dietel, P. (1894) Die gattung Ravenelia (Nachtra$ ge). Hedwigia 33 : 367–371. Dietel, P. (1906) Monographie der gattung Ravenelia Berk. Beihefte zum Botanischen Centralblatt 20 : 343–413. Ellis, J. B. & Everhart, B. M. (1895) New fungi, mostly Uredineae and Ustilagineae from various localities, and a new Fomes from Alaska. Bulletin of the Torrey Botanical Club 22 : 362–364. Farr, M. L. (973) An Annotated List of Spegazzini’s Fungus taxa. 2 vols. J. Cramer, Lehre. Gallegos, H. M. & Cummins, G. B. (1981) Uredinales (royas) de MeT xico. Vol. 2. Instituto Nacional de Investigaciones Agrı! colas, Culiaca! n, Sinaloa, Me! xico. Gardner, D. E. & Hodges, C. S. jr (1985) Spore surface morphology of Hawaiian Acacia rust fungi. Mycologia 77 : 575–586. Hennen, J. F. & Cummins, G. B. (1990) New species and nomenclature of Ravenelia in Neotropica. Report of the Tottori Mycological Institute 28 : 1–14.

974 Hennen, J. F., Hennen, M. M. & Figueiredo, M. B. (1982) I; ndice das Ferrugens (Uredinales) do Brasil. Arquivos do Instituto BioloT gico, Sao Paulo 49 : 1–201. Hennings, P. (1895) Nachtra$ ge zu den Fungi goyazenses. Hedwigia 34 : 319–324. Herna! ndez, J. R. & Hennen, J. F. (1999) Ravenelia species (Uredinales), parasites of Prosopis species (Leguminosae) in South and North America. Abstracts, XVI International Botanical Congress, St Louis, Missouri, August 1–7 : 318 Herna! ndez, J. R. & Hennen, J. F. (2002) Rust fungi (Uredinales) of Northwest Argentina. Sida 20 : 313–338. Lindquist, J. C. (1946) Tres especies, nuevas o crı! ticas, de Ravenelias en Argentina. BoletıT n de la Sociedad Argentina de BotaT nica 1 : 298–302. Lindquist, J. C. (1951) Uredinales de la Provincia de Mendoza. Revista de la Facultad AgronomıT a de la Universidad Nacional de La Plata 28 : 65–114. Lindquist, J. C. (1954) Las especies argentinas de Ravenelia. Revista de la Facultad AgronomıT a de la Universidad Nacional de La Plata 30 : 103–128. Lindquist, J. C. (1958) Notas uredinolo! gicas V. Revista de la Facultad AgronomıT a de la Universidad Nacional de La Plata 34 : 209–222. Lindquist, J. C. (1978) Especies nuevas de Uredinales. Hickenia 1 : 152–155. Lindquist, J. C. (1982) Royas de la RepuT blica Argentina y Zonas LimıT trofres. Instituto Nacional de Tecnologı! a Agropecuaria, Buenos Aires. Lindquist, J. C. & Garcı! a Zorro! n de Rosengurtt, N. (1967) Uredinales del Uruguay. Revista de la Facultad AgronomıT a de la Universidad Nacional de La Plata 43 : 187–214. Long, W. H. (1903) The Ravenelias of the United States and Mexico. Botanical Gazette (Crawfordsville) 35 : 111–133. Lo! pez-Franco, R. & Hennen, J. F. (1990) Spore wall sculpturing patterns of Ravenelia anamorphs as determined by scanning electromicroscopy. IMC4 Abstracts : 1A-34\3. Spegazzini, C. (1881) Fungi Argentini, additis nonnulis Brasiliensibus Montevideensibuque. Anales de la Sociedad CientıT fica Argentina, Pug. IV, 12 : 64–81. Spegazzini, C. (1898) Fungi Argentini, novi vel critici. Anales del Museo Nacional de Historia Natural de Buenos Aires 6 : 81–368. Spegazzini, C. (1909) Mycetes Argentinenses (series IV). Anales del Museo Nacional de Historia Natural de Buenos Aires 19 : 257– 458. Spegazzini, C. (1912) Mycetes Argentinenses (series VI). Anales del Museo Nacional de Historia Natural de Buenos Aires 23 : 1–146. Spegazzini, C. (1922) Fungi Paraguayenses. Anales del Museo Nacional de Historia Natural de Buenos Aires 31 : 355–450. Spegazzini, C. (1925) Uredı! neas argentinas (nuevas o crı! ticas). Revista Argentina de BotaT nica 1 : 93–145. Sydow, H. (1930) Fungi Venezuelani. Annales Mycologici 28 : 29–224. Sydow, H. & Sydow, P. (1916) Fungi amazonici a cl. E. Ule lecti. Annales Mycologici 14 : 65–97. Corresponding Editor : D. L. Hawksworth