|
2. The genus Hysterographium Corda Fragosa Cif., in Ciferri & Fragoso, 1926 Hysteriopsis Spreg., 1906 Hysterium Tode ex Fries (pro parte), 1823 Polhysterium Speg., 1912
Hysterographium flexuosum (Schw.) Sacc. & Hg. subrugosum (Cooke & Ellis) Sacc.
The genus Hysterographium Corda While Corda (1842) included phragmosporous and dictyosporous species in the genus Hysterographium, De Notaris (1847) restricted the genus to species with muriform ascospores only. Zogg (1962), among others, maintained the genus as Hysterographium Corda emend. De Not., however a change in authorship is not warranted. Clements & Shear (1931) regarded Hg. fraxini (Pers. : Fr.) De Not. as the type. The genus is characterized by pigmented asymmetric dictyospores, with one to several longitudinal septa, ovoid to ellipsoid-fusoid, relatively broad, usually constricted at the first-formed septum. Zogg (1962) extensively revised the synonymy in the genus and recognized four species: Hg. fraxini and Hg. flexuosum (Schw.) Sacc., with large, relatively wide dictyospores, constricted primarily at the medium septum, and Hg. mori (Schw.) Rehm & Hg. subrugosum (Cooke & Ellis) Sacc., with smaller, fewer-celled dictyospores, short and squat in the former, long and slender in the latter, both conspicuously constricted at the median septum. Since then, an additional three species have been described: Hg. minus Amano from Japan, possessing the smallest spores in the genus (Amano 1983), Hg. spinicola Doidge from S. Africa, recollected from the thorns of Acacia and validated by van der Linde (1992), with an atypical brick-red epithecium and spore lengths only slightly larger than those of Hg. mori; and Hg. pulchrum Checa, Shoemaker & Umaña from Costa Rica, also with a red pigment in the hamathecium and spore lengths intermediate between Hg. mori and Hg. subrugosum (Checa et al. 2007). Barr (1975) removed Hysterium nova-caesariense Ellis from synonymy with Hg. flexuosum (Zogg 1962) and placed it as Ostreichnion nova-caesariense (Ellis) M.E. Barr in the Mytilinidiaceae. Likewise, Hysterium formosum Cooke, synonymized by Zogg (1962) under Hg. mori, was accepted as Ostreola formosa (Cooke) M.E. Barr, also in the Mytilinidiaceae (Barr 1990). The genus Gloniopsis Sacc., with hyaline dictyospores, may show a relationship to the genus Hysterographium, as certain series within each possess very similar dictyospores. Molecular data is needed to determine the relationship between these two genera, that differ primarily only in the presence of absence of pigmentation of their dictyospores. Zogg (1962): “Fruitbody a hysterothecium black, brittle, hard, mostly matt, oval to elongated, solitary to parallel gregarious, simple, straight to flexuous, entire, rarely bifurcated, with a sunken, invaginated longitudinal slit running the length of the fruitbody, without a keel. Asci clavate to cylindrical, clearly bitunicate, fissitunicate. Ascospores large pigmented dictyospores: muriform, with cross & vertical septa, parenchymatic in appearance, spindle-shaped, elliptic, oval to long cylindric, clearly pigmented shades of brown.” Amano (1983): “Hysterographium Corda emend de Notaris. Icon. Fung. 5: 34, (1842), emend., de Notaris, Giorn.. bot. ital. II, 2 (7-8): 21, (1847). Type species: Hysterographium fraxini (Pers.) de Not., Giorn.. bot. ital. II, 2 (7-8): 21, (1847). Corda (1842) included two species in Hysterographium: H. pulicare (Pers.) Corda with phragmosporous ascospores & H. elongatum (Wahlenb.: Fr.) Corda with muriform ascospores.. Five years later, de Notaris (1847) restricted the genus to species with muriform ascospores, namely H. fraxini (Pers.) de Not. & H. elongatum (Wahlenb.: Fr.) Corda. The former species was selected as the type of the genus by Clements & Shear (1931). Since de Notaris emendation of the genus, more than 70 epithets have been assigned to the genus. Zogg (1962) accepted only four species.” Barr (1990): “Ascomata erumpent to superficial, separate or gregarious, elongate, hysterothecioid; apex rounded, opening by long slit; surface smooth or roughened or longitudinally striate; peridium wide, composed of small pseudoparenchymatous cells, heavily pigmented at surface. Asci basal, clavate. Pseudoparaphyses narrowly cellular. Ascospores brown, obovoid or ellipsoid fusoid, ends obtuse or acute, straight or inequilateral, several septate, with one to three longitudinal septa, usually constricted at first-formed septum, less so or not constricted at later septa; wall thin or thick, smooth or verruculose with one globule per cell; overlapping biseriate in the ascus. Anamorphs coelomycetous where know. Saprobic or hemibiotrophic on woody plants. Type: H. elongatum (Wahl..:Fr.) Corda = H. fraxini (Pers.:Fr.) De Not. (Zogg, 1962). The extensive lists of synonyms presented by Zogg (1962) is accepted, except for a few. Hysterium nova-caesariense Ellis is removed from synonymy of Hysterographium flexuosum & is accepted as Ostreichnion nova-caesariense in the Mytilinidiaceae (Barr, 1975, 1990). Hysterium formosum Cooke, considered by Zogg (1962) to be a synonym of Hysterographium mori, is Ostreola formosa (Cooke) Barr, also in the Mytilinidiaceae.” Vasilyeva (1999): “Hysterographium Corda, Ion. Fung., 5: 34, (1842), emend. De. Not., Giorn. Bot. Ital., 2(1), 2: 21, (1847). Hysteriopsis Speg., Rev. Fac. La Plata, 2: 308, (1906). Polhysterium Speg., Anal. Mus. Nac. Buenos Aires, 23: 87, (1912). Fragosoa Cif. in Cif. et Gonz. Frag., Bol. R. Espan. Hist. Nat., 26: 192, (1926). Hysterothecia erumpently superficial, densely aggregated or scattered, oblong, straight to subflexuous, sometimes stellate, black, carbonaceous. Asci clavate or cylindrical, short-stipitate. Ascospores oblong-elliptic or fusiform, muriform, colored. Type species: Hysterographium elongatum (Wahlenb.) Corda.” Wehmeyer (1975): “This genus also has ellipsoid to elongate ascocarps, but he ascospores are muriform & brown. A number o species of Hysterographium have been reported as having pycnidial states. Hilitzer (1929) placed these in Hysteropycnis. Lohman (1933) reported superficial pycnidial forms as in Aposphaeria & sunken ones as in Dendrophoma.”
Dichotomous Key to species of Hysterographium Corda A. Ascospores dictyosporous, clearly vertical- & cross-septate, & pigmented shades of brown. Typically centrally constricted at the first-formed septum. 1. Dictyospores 26 – 38 x 10 – 15 μm. = Hysterographium minus N. Amano 2. Dictyospores 30 – 51 x 12 – 22 μm. = Hysterographium fraxini (Pers. : Fr.) De Not. 3. Dictyospores (40) 45 – 65 (70) x (8) 9 – 17 (19) μm. = Hysterographium flexuosum (Schw.) Sacc. B. Ascospores dictyosporous as above, but either smaller & with fewer cells, or with fewer vertical septa. Typically, constricted at each septum, rather than just at the central septum. 1. Dictyospores mostly with three- to four-cross septa & one- to two- vertical septa. Dictyospores (12) 15 – 23 (25) x (5) 7 – 10 (11) μm. = Hysterographium mori (Schw.) Rehm 2. Dityospores with (five-) six cross septa & with one vertical septum in either cell adjacent to the primary septum, or in both cells. Dictyospores 22 –25 (-27) x 5 - 6 μm. Red pigment in hamathecium. Neotropical (Costa Rica). = Hysterographium pulchrum Checa, Shoemaker & Umana 3. Dictyospores with five- or six-cross septa & mostly with one longitudinal septum. Dictyospores 25 – 28 x 11 – 13 um. = Hysterographium spinicolum Doidge 4. Dictyospores mostly seven- to eleven-cross septa but with only one- to two-vertical septa; vertical septa mainly restricted to central cells. Dictyospores (22) 25 – 34 (45) x (6) 8 – 12 (17)μm. = Hysterographium subrugosum (Cook & Ellis) Sacc. Detailed Dichotomous Key to species of Hysterographium Corda A. Ascospores dictyosporous, clearly vertical- & cross-septate, & pigmented shades of brown. Typically centrally constricted at the septum. 1. Dictyospores 26 – 38 x 10 – 15 μm: = Hysterographium minus N. Amano N. Amano. 1983. Trans mycol Soc Japan 24: 283-297. Amano (1983) : “Ascomata aggregated or dispersed, superficial with immersed base, elongated with acute ends, straight or curved, rarely branched, opening by a narrow longitudinal slit, black, carbonaceous 0.5 – 1.0mm long, 130 - 180 μm wide, 180 – 230 μm high. Tissues of ascomatal walls of textura angularis close to textura epidermoidea, 20 – 50 μm tich, composed of thick-walled, brown cells, 3 – 5 μm in diameter. Pseudoparaphyses filiform, branched, anastomosed, hyaline, 1.0 – 1.5 μm thick, enlarged to 2.0 – 2.4 μm in diameter at the apex, in 2% KOH staining greenish white. Asci bitunicate, broadly clavate, stipitate, 8-spored, in 2% KOH staining pale green or greenish white, staining more intensely above, 80 – 105 x 23 – 34 μm. Ascospores irregularly biseriate, ellipsoid, with 6 – 13 transverse septa & 1 – 3 longitudinal septa, slightly constricted at the transverse septa, straight or curved, 26 – 38 x 10 – 15 μm. On decorticated branch of broad-leaved tree. Japan. H. minus resembles H. fraxini but differs in size of ascospores. Taxonomic significance of color reaction of asci & pseudoparaphyses in 2% KOH is not known. It has not been reported before for hysteriaceous fungi.” 2. Dictyospores 30 – 51 x 12 – 22 μm: = Hysterographium fraxini (Pers. : Fr.) De Not. de Notaris. 1847. Giorn. Bot. Ital. II, 5 – 52. Cosmopolitan. On live bark & dead wood of: Populus, Juglans, Corylus, Fagus, Quercus, Cotinus, Acer, Cornus, Fraxinus, Syringa, Olea, Ligustrum, Jasminum, Phillyraea, Bignonia, Catalpa, Viburnum. Hysterium fraxini Pers., 1801 Ellis & Everhart (1892): “Hysterothecia scattered or gregarious, erumpent, elliptical, black, obtuse above, 1-1.5mm long, 0.5-0.75mm wide, lips swollen, smooth, partially open so as to expose the narrow disk. Asci clavate rounded above, 150-200 x 30-40 μm, eight spored with filiform paraphyses. Ascospore biseriate, oblong-elliptical, scarcely constricted in the middle, seven- to nine-septate & muriform, dark yellow brown measuring 30-40 x 15-18 μm. On dead limbs of Fraxinus, NY, PA, IA & Canada. Probably common throughout the USA.” Bisby (1944): “Hysterothecia about 1 – 2 x 0.5mm, erumpent through the cortex, rarely superficial on the wood, dull black, sometimes forming ‘fairy rings’; asci 125 – 200 x 25 – 40 μm, with eight (or fewer) spores; the ascospores are irregularly biseriate, 30 – 50 x 12 – 20 μm, golden to chestnut brown, surrounded by a gelatinous envelope, ellipsoid with ends somewhat narrowed, with or without constriction at the center, having five to ten cross-septa, each locule with none to three longitudinal divisions; paraphyses filiform, branched above & forming a brown epithecium. Nearly all British collections from ash (Fraxinus) branches. European & North American collections appear identical.” Barr (1990): “Ascomata 1 – 2 mm long, 330 μm dia., ellipsoid, widely erumpent; surface smooth, lacking longitudinal striae, longitudinal slit usually depressed; peridium 40 – 45μm wide. Asci 90 x 25 - 30 μm. Ascospores (25-) 30 – 45 (-51) x (10-) 13 – 15.5 (-22) μm, obvoid, ends obtuse, 7 – 9 septate, with 2 – 3 longitudinal septa, constricted at median first-formed septum. Anamorphs coelomycetous; conidiomata developed within periderm of steam-sterilized ash twigs; conidiogenous cells 8 – 12 x 1 – 2 μm, as short branches; conidia 4.5 – 5 x 0.1 – 1.2 μm, hyaline, cylindric (Hysteropycnis fraxini Hilitzer, Lohman, 1932). On woody branches, chiefly Fraxinus, cosmopolitan. Zogg (1943, 1962) studied the development of this species in culture. Infection studies, utilizing pure cultures derived from single ascospores, resulted in the formation of the anamorph &/or the telomorphs on a wide range of woody plants, both gymnosperms & angiosperms. Zogg concluded that H. fraxini was a non-specialized facultative saprobe that was able to spread from dead to living tissues.” van der Linde (1992): “Fruitbodies single or in small groups, superficial, elliptical with rounded ends, straight or slightly curved, usually not branched, 1.5 – 2.0 x 0.2 – 0.5mm. These dimensions are slightly smaller than those given by Zogg (1962) for this species. Pseudoparaphyses hyaline, septate, thickend apically, branched to form an epithecium. Asci irregularly biseriate, cylindric-clavate, 8-spored, 110 – 120 x 30 – 35um. Ascospores dark olive brown, dictyosporous with 5 – 8 transverse septa & 3 longitudinal septa, soemtines slightly curved, 40 – 45 x 12 – 15um.” 3. Dictyospores (40) 45 – 65 (70) x (8) 9 – 17 (19) μm: = Hysterographium flexuosum (Schw.) Sacc. Saccardo. 1883. Syll. Fung. II: 781. Somewhat rare. Living bark & old wood of: Pinus, Salix, Betula, Fagus, Quercus, Prunus, Gleditschia, Acer, Ceanothus, Vitis, Garrya, Cornus, &romeda. Uncommon. North America, Europe (Germany), & South Africa. Hysterium flexuosum Schw., 1822 Note*: Hysterographium nova-caesariense (Ellis) Ell. & Ev. was transferred by Zogg (1962) to Hysterographium flexuosum (Schw.) Sacc. However, Barr (1975) transferred Hysterium nova-caesariense to Ostreichnion nova-caesariense (Ellis) Barr. Barr adds an additional synonym Hysterographium acerinum Peck, Bull. NY State Mus. 167: 43 (1913). Ellis & Everhart (1892) for H. vulvatum: “Hysterothecia erumpent-superficial, scattered or gregarious, 1-3mm long, straight or flexuous; lips at first closed, then open so as to leave a wide furrow between them, mostly with 1-2 deep striae on each side, often apparently double i.e., with one set within another. Asci oblong, clavate, obtuse, paraphysate, 8-spored, 110-112 x 20-25 μm, with stipe 30-35 μm long. Ascospores irregularly biseriate, broad-fussoid, slightly curved, multi-(10 -15) septate & muriform, strongly constricted in the middle, the upper part broader, olive-brown measuring 50-62 x 15-20 μm. Common on dead, dry mostly decorticated limbs of oak & other deciduous tress.” Barr (1990): “Ascomata 1 – 2 (-3) mm long x 330 - 385 μm wide x 330 – 440 μm high; surface longitudinally striate; peridium 52 – 75 μm wide, to 90 μm wide at base. Asci 100 – 130 (-180) x 20 – 30 (-38) μm. Ascospores (30-) 40 – 65 (-80) x 10 – 18 μm, ellipsoid fusoid, ends acute, 7 – 15 septate, with 1 – 3 longitudinal septa, wall usually verruculose. Anamorph in culture coelomycetous or conidiogenous cells developing on surface hyphae, 5 – 8 x 1.5 μm; conidia 3 – 4 x 1.3 – 1.5 μm (Lohman, 1932, as Hysterographium vulvatum). On decorticated wood, cosmopolitan. Although Zogg included H. acerinum as a synonym of H.. fraxini based upon the type description, the original collection has narrower ascospores than those of H. fraxini & the ascomata are longitudinally striate, characteristic of H. flexuosum.” Messuti & Lorenzo (2003): “Hysterothecia gregarious, closely associated, superficial, straight to flexuous, elliptical with blunt to pointed ends, longitudinally striated, 0.5 – 3.5 x 0.2 – 0.8mm; pseudoparaphyses hyaline, septate, flexuous, branched near the tips to form an epithecium; asci clavate-cylindrical, 8-spored, 100 – 122 x 20 – 40 μm; ascospores yellowish to dark brown, biseriate, with 7 – 15 transverse & 1 – 4 longitudinal septa, muriform, ellipsoid to broadly ellipsoid, constricted at the medium septum, 36 – 65 (-70) x (8-) 9 – 20 μm. Cosmopolitan. In Patagonian Nothofagus forests, H. flexuosum is very frequent. Collected on decorticated trunks or wood of Nothofagus alpine, N. antarctica & N. pumilio & on bark of Escallonia serrata, N. antarctica & N. dombeyi.” Messuti & Lorenzo add the following as synonyms: Hysterographium magellanicum Speg., H. fuegianum Speg. & H. fuegianum f. intermedium Rehm. van der Linde (1992): “Fruitbodies arranged in groups, not closely associated, embedded or ermpent, long, linear with tapering ends, straight, not branched, up to 2.0 x 0.4mm. Pseudoparaphyses hyaline, filiform, branched near tips to form an epithecium. Asci not seen because the material was very old. Ascospores golden to dark brown, usually constricted at median septum, 9- or 10-transverse septa & 1- to 2-longitudinal septa, measuring 52 – 57 x 17 – 20um.” Hysterographium flexuosum (Schw.) Sacc. B. Ascospores dictyosporous as above, but either smaller & with fewer cells, or with fewer vertical septa. Typically constricted at each septum, rather than just at the central septum. 1. Dictyospores mostly with three- to four-cross septa & one- to two- vertical septa. Dictyospores (12) 15 – 23 (25) x (5) 7 – 10 (11) μm: = Hysterographium mori (Schw.) Rehm Rehm. 1881. Ascom. Fasc. 7, 26. Bericht d. Nat. hist. Vereins Augsburg: 76-87. Cosmopolitan. Gregarious but highly variable. On aged wood of: Pinus, Juniperus, Salix, Ostrya, Castanea, Quercus, Ulmus, Morus, Pyrus, Amelanchier, Crataegus, Rubus, Cercocarpus, Prunus, Gleditschia, Fabaceae, Melia, Pistachia, Cotinus, Rhus, Acer, Zizyphus, Vitis, Fraxinus, Olea, Aspidosperma. Hysterium mori Schw., 1834 Ellis & Everhart (1892): “Hysterothecia erumpent-superficial, elliptical, oblong, linear or cylindrical, 1-3 mm long by 0.5-1 mm wide, mostly straight & lying parallel, gregarious & often crowded so as to cover the matrix more or less completely for some extent, more or less distinctly longitudinally striate; lips mostly closed at first, finally more or less open, exposing a narrow, linear or lanceolate disk. Asci cylindrical, about 100 x 12 μm (including short stipitate base); paraphysate, eight spored. Ascospores uniseriate or sub-biseriate above, ovate, varying to oblong or ovate-elliptical, three- to five-septate, constricted at the middle septum, one or two of the cells divided by a longitudinal septum, brown, 15-25 x 7-8 μm. On decorticated exposed wood, also, but less frequently, on dead limbs still covered with bark. Common.” Barr (1990): “ Ascomata 1 – 2 (-3) long, 220 – 275 (-440) μm wide, 190 – 330 μm high; surface usually longitudinally striate; peridium 35 – 52 μm wide, to 100 μm at base; Asci (55-) 80 – 120 x 11 - 16 μm, Ascospores 14 – 22 (-26) x 6.5 – 9 (-10.5) μm, obovoid, ends obtuse, 3- (5 – 7) septate, with one longitudinal septum, rarely partial second septum in one or two cells; wall finely verruculose. Anamorph coelomycetous, Aposphaeria – like in nature, in culture condidiomata as irregular locules; conidiogenous cells 8 – 10 x 1.5 - 2 μm; conidia (2-) 2.5 – 3.5 (-4) x 1 – 2 μm (Lohman, 1932). On periderm & decorticated wood, cosmopolitan.” van der Linde (1992): “Fruitbodies single or in small groups, sometimes closely associated, superficial when mature with embedded base, long, linear with tapering ends, most commonly straight, not branched, up to 1.5 x 0.26mm. Pseudoparaphyses hyaline, thickened apically, septate, branched, forming an epithecium. Asci irregularly biseriate, 8-spored, 150 – 170 x 12 – 13um.Ascospores brown, most commonly 5 or 6 transverse septa & 1 longitudinal septum, constricted at the median septum, dictyosprorous 18 – 23 x 7 – 10um.” Hysterographium mori (Schw.) Rehm 2. Dityospores with (five-) six cross septa & with one vertical septum in either cell adjacent to the primary septum, or in both cells. Dictyospores 22 –25 (-27) x 5 - 6 μm. Red pigment in hamathecium. Neotropical (Costa Rica). = Hysterographium pulchrum Checa, Shoemaker & Umana Checa, Shoemaker & Umana. 2007. Mycologia 99(2): 285-290. Neotropical. Collected from Guanacaste, Tempisque, National Park Palo Verde, Palo Verde, Costa Rica. Checa, Shoemaker & Umana (2007): "Ascomata in bark, semi-immersed, hystereiform, 0.5 - 2mm long, 250um wide, 250um high, smooth, glabrous. Sulcus central. Ascocarp wall in section mostrly 20-30um thick, of about seven layers of rectangular, brick-like thin-walled cell 8-20 x 2-4um, with occasional lateral excrescences of 20-25 layers of similar cells and 50um wide. Paraphyses numerous, 1um wide, aseptate, without guttules, without slime coating, outermost centrum tissue red. Asci bitunicate, numberous, in a basal cluster, somewhat clavate, 80-90 x 13-15um (par sporif.), long-stalked (ca. 50um) with eight, over-lapping linearly tetraseriate ascospores above to uniseriate below. Ascospores terete, ellipsoid, L/W 3.7, straight to inequilateral, 22-25 (-27) x 5-6um, transversely (5) 6-septate, in centrifugal sequence (4):3:2:1:2:3:(4), mostly with one vertical septum in both cells next primary septum (absent in some spores), first-forme3d septum supramedian (0.40), constricted at first-formed septum and slightly so at others, without dots at ends of septa, septa thin, sidest at third cells from apex, pale reddish brown, without guttules, smooth, without sheath." The authors further state that Hysterographium pulchrum differs from H. mori in having six transverse septa and usually one vertical septum in one or both cells next to the supramedian primary septum. The red pigment is also a distinguishing feature (found also in Hysterium asymmetricum).
Hysterographium subrugosum (Cooke & Ellis) Sacc.
|
|
|
|
