LICHEN FLORA OF EASTERN NORTH AMERICA
THE GENUS OPHIOPARMA NORMAN
Philip F. May
20 Divinity Avenue, Cambridge, MA 02138
Abstract: Descriptions, keys, illustrations, and distribution maps are provided for the two species of Ophioparma Norman found in eastern North America (eNA), both saxicolous. O. lapponica, with simple to one-septate ellipsoidal spores less than 30 µ in length, is known from only one locality in the Flora area. The divaricatic/usnic chemotype of O. ventosa is found in alpine and boreal settings as far south as New England and New York. Occasional eNA specimens of O. ventosa contain psoromic acid, salazinic acid, atranorin, zeorin, or other triterpenes as accessory compounds.
Keywords: Ophioparma / ventosa / lapponica / North America / key
This paper on the lichen genus Ophioparma Norman is another in the series of generic reviews by various authors for the Flora of eastern North America (eNA). The Flora covers the part of the continent bordered by the Gulf of Mexico, the Atlantic Ocean, the 97°W meridian, and the 54°N parallel. The 97°W meridian was chosen as the approximate boundary between the midgrass and tallgrass prairies. The 54°N parallel was chosen as the approximate southern boundary of the arctic floristic zone.
The present paper is based on a detailed study of Ophioparmalapponica (May 1997) in which 470 mostly North American, mostly saxicolous Ophioparma specimens from CANL, COLO, FH, F, H, US, AMNH, and NY were examined. Since then, all eNA specimens at WIS and DUKE have also been examined. A total of 70 specimens from eNA were found among these collections. The species descriptions are based on these eNA specimens, supplemented by other NA specimens in the case of O. lapponica. The description of the genus, on the other hand, is based on published data (Printzen & Rambold 1996; Kalb & Staiger 1995; Rogers and Hafellner 1988; Ekman 1993; Staiger and Kalb 1995; May 1997), with additional specimen-based information provided for eNA.
Previously published distribution maps (Thomson 1968, 1997) and discussions
of saxicolous Ophioparma in NA (Culberson 1963) confound the two
morphological species now known to be on the continent (May 1997). The
distribution maps presented here are based on examination of the specimens
mentioned previously. The synonyms listed later for O. ventosa represent
only a subset of the many found in Zahbruckner (1928), namely those commonly
encountered on NA herbarium specimens and in the literature of the last
OPHIOPARMA Norman, Nytt Mag. Naturv. 7:230-231, 1853.
ETYMOLOGY: from the Greek noun ophis, snake (combined form, ophio-); and the Latin noun parma, small round (military) shield. The meaning of the name is unclear, but perhaps has something to do with the sinuous shape of the spores in O. ventosa when seen in the ascus.
TYPE SPECIES: Ophioparma ventosa (L.) Norman (Hafellner 1984).
KEY REFERENCES for NA: May 1997; Kalb & Staiger 1995; Rogers & Hafellner 1988; for Europe, South America, and Asia: (Kalb and Staiger 1995; Printzen & Rambold 1996; Leukert and Meinel 1981; May 1997; Skult 1997).
SELECTED PREVIOUS ILLUSTRATIONS: Illustrations in the literature have not distinguished between the two morphological saxicolous Ophioparma species now known and thus may not be identified correctly at the species level. However, they can be used to get a sense of what the genus looks like.
Color illustrations of the thallus: Wirth 1995, p. 630 (almost certainly O. ventosa, based on the locality); Hansen & Andersen 1995, p. 91. Black and white illustrations of thallus: Dopson 1992, p. 220 (probably O. ventosa); Thomson 1997, p. 440 (definitely O. lapponica, based on recent reexamination of the actual specimen). Drawings showing anatomical characters of the genus: Rogers and Hafellner 1988, p. 170; Staiger and Kalb 1995, p. 15.
HISTORY OF GENUS: Presently six species are known, four of which are corticolous, two saxicolous. Until recently the saxicolous species of Ophioparma were included in Haematomma as the "ventosum group" (Culberson 1963). In segregating the ventosum group, Rogers & Hafellner (1988) revived the 1853 generic name Ophioparma Norman. Since 1988, four corticolous species--not present in eNA--have been transferred from other genera to Ophioparma (Kalb and Staiger 1995; Printzen and Rambold 1996; Ekman 1996). In the process, the definition of the genus was emended to include simple-spored taxa (Printzen & Rambold 1996). Until recently, species concepts for saxicolous Ophioparma were primarily based on chemistry. May (1997) and Skult (1997) re-examined saxicolous Ophioparma and found there was little basis for erecting taxa based on chemistry, but that two saxicolous species existed based on spore type. May (1997) realigned the names O. ventosa and O. lapponica to refer to these morphological species. (Previously, O. lapponica had been used in NA and elsewhere as the name for the divaricatic/usnic chemical strain of O. ventosa.)
VEGETATIVE STRUCTURES: Thallus crustose, dispersed areolate, rimose-areolate, granular, or subsquamulose, up to 15 cm in diameter, ochre, orange brown, gray, gray-olive, or greenish yellow (in eNA usually gray-olive to greenish yellow). Thallus 0.2 mm to 8 mm thick. Photobiont trebouxiod.
REPRODUCTIVE STRUCTURES: Apothecia sessile to 3 mm in diameter, proper exciple well developed, thalline exciple (with algae) present or not (lacking in eNA specimens), hypothecium hyaline, hymenium of mostly simple, strongly agglutinated, fairly thick paraphyses. Epihymenium encrusted with haemoventosin, appearing deep red from above, K+ deep blue, then violet, and finally dissolving.
Asci of the "Ophioparma-type" sensu Rogers and Hafellner (1988), that is, with an amyloid tholus lacking axial mass, ring structure, and ocular chamber (Fig. 7-8). Here the "tholus" means the thickened part of the ascal tip that lies within the ascal wall. There is also a thin coating of KI+ dark blue material on the outside of the top of ascus wall, so that the actual appearance in specimens is of a sandwich of blue separated by the unstained cell wall (Fig. 8). The shape of the KI+ dark blue-stained portion of the tholus varies considerably, from a thin crescent at the top, to a filled-in arch, to a circle with a crescent removed from the bottom A late-developing blue-hyaline-blue sandwich completely within the tholus, as described by Ekman (1993), is seen only rarely.
Spores eight, hyaline, simple to transversely multiseptate, 3.5-9 mm wide and 9-70 mm long, with a length-to-width ratio of 2.2 to 18, irregularly ellipsoid to asymmetrically bi-fusiform, sometimes become spatulate at one end (having a "tail"), lacking a gelatinous epispore.
Conidiomata immersed, having a green or red pigment in the ostiole. Conidiophores similar to type VI sensu Vobis (1980). Conidia bacilliform, 7-11 x ca. 1 mm.
CHEMISTRY: Haemoventosin, usually granular, as an epihymenial, hymenial, and sometimes excipular pigment and, for one Asian species, also present in the ostioles of the conidiomata. For most species the ostioles of the conidiomata contain an unidentified green pigment. In the thallus, one or more of the following: usnic acid, divaricatic acid, siphulin, lecanoric acid, thamnolic acid, hypothamnolic acid, various B-orcinol depsidones, atranorin, zeorin and other triterpenes, and various fatty acids. The two species occurring in eNA always have usnic and divaricatic acid. They may also have accessory compounds (see the species descriptions).
DISTRIBUTION AND SUBSTRATES: Ophioparma grows on wood, bark, and silicic rock, but only on rock in eNA. All Ophioparma species but one have a north temperate to arctic distribution. O. araucariae grows in the temperate forests of Chile and Argentina.
LICHENICOLOUS FUNGI: Muellerella and Sphinctrina (Staiger & Kalb 1995).
RELATED GENERA: Haematomma has the following generic characters that differ from Ophioparma (Staiger & Kalb 1995): The ascus has both an axial mass and a distinct ocular chamber. The red pigment in the hymenium is not haemoventosin; it reacts KOH+ red or KOH-. The proper exciple of the apothecia is thin; a thick thalline margin is always present (but may be aspicilioid). The paraphyses are thin, individually covered with a gelatinous layer, and highly branched and anastomosed. The conidiomata lack a green pigment. The conidiophores are Vobis (1980) Type V. Some species are sorediate. Specimens are seldom found on rock. Most species have atranorin, sphaerophorin, or placodialic acid as major constituents. The thalli host different lichenicolous fungi than Ophioparma.
Loxospora has the following generic characters that differ from
(Staiger & Kalb 1995): The ascus has a wide axial mass. The pigment
in the hymenium is not haemoventosin; it reacts KOH+ yellow. The paraphyses
are more frequently anastomosed, but are not stuck together with hymenial
gelatin. The spores gain their traverse septation only in old age. The
conidiomata lack a green pigment. The conidiophores are Vobis (1980) Type
V. Many species are sorediate or isidiate. Specimens are rarely found on
rock. Most species have thamnolic acid as the main constituent, whereas
only one species of Ophioparma sometimes contains this compound.
The thalli host different lichenicolous fungi than Ophioparma.
KEY TO SPECIES OF OPHIOPARMA IN EASTERN NORTH AMERICA
1a. Spores 3-7 septate at maturity, more than 30 µm long (even when immature), with a length-to-width ratio greater than 6:1, asymmetrically tapered, with one end blunt or fusiform and one end subulate (broadly acicular) to acicular, or sometimes becoming narrowly spatulate (i.e., with a "tail"), often helically arranged in the ascus, but also often arranged subparallel to the long axis of the ascus..........................................O. ventosa
1b. Spores simple to one-septate at maturity, less than 30 µm long, with a length-to-width ratio less than 6:1, narrowly ellipsoid to subfusiform (when immature sometimes asymmetrically tapered, irregular, or curved), arranged sub-parallel or diagonally to the long axis of the ascus..................O. lapponica
Ophioparma lapponica (Räs.) Hafellner & R. W. Rogers, Lichenologist 20: 173. 1988.
Haematomma lapponicum Räs., Ann. Acad. Sci. Fenn, Series A, 34(4): 67. 1931. TYPE: Finland, Ob. Alkkula (Ylitornio), Aavasksa, Kvartsiitilla, 20 VII 1915, Veli Räsänen (H, lectotype, designated by May, 1997) [chemistry: usnic and divaricatic acids, haemoventosin].
Thallus: crustose, pale yellow to pale green-yellow or gray green-yellow, rimose-areolate, occasionally with a few dispersed areoles at the margin, evenly thickened to quite variable in thickness within a single thallus, to 7 mm thick (including the hypothallus) where groups of areoles form convex heaps, but more typically 2-5 mm thick. Areoles usually moderately convex, sometimes almost flat, to 2.5 mm across, corticate partway into the cracks, generally matte in luster, with a smooth, finely verruculose (0.1 mm) or micro-rugulose surface. Areoles sometimes resting directly on the substrate, especially near the thallus margin; but more typically areoles and mounds of areoles resting on a thin to thick layer of fungal tissue (which might be considered a hypothallus), often incorporating grains of rock. Hypothallus and the lowest part of the areoles white to dark gray where exposed.
Apothecia: sessile, attached over entire diameter or partly to mostly constricted at base, to 3.2 mm in longest dimension, round when solitary, but often crowded together, then slightly compressed to highly irregular in shape, sometimes partially subdivided. Disks plane to moderately convex, deep red brown to very deep red. Margin 100-500 m m thick, sometimes raised, smooth to finely rugose, often strongly flexuose, shiny to matte, pale-yellow to orange-yellow, almost thalline-appearing, or occasionally the same color as the thallus. Margin, however, lacking algae at every stage of development except occasionally a few in the outer basal area of mature apothecia where the margin meets the thallus, usually separated from these algae by a small infold of cortex. A blue-green pigment (same as that in the ostioles of conidiomata) sometimes present in the margin of apothecia, giving the false impression of an algal layer. The interior part of the margin sometimes covered with red pigment, and the outer part of the hymenium sometimes with very few asci, giving the false impression of a two-layered margin. Hymenium and epihymenium together 60-80 mm high, inspersed at the surface and part or all the way down with crystals of deep orange pigment, strongly agglutinated, hyaline and I+ blue in the lower part. Subhymenium hyaline, 60-95 mm thick, I+ blue. Hypothecium: hyaline, I-, continuous with and grading into the thick fungal layer (hypothallus) in some specimens; but in other specimens, more compact than the thick fungal layer and separated from it by cortex. Exciple of radiating, highly branched, ± anastomosing hyphae originating from the hypothecium, cells 7-12 ´ 3 m m, the outer, shorter cells forming a compact gelatinous cortical layer interspersed with granules, algae generally lacking. Paraphyses: ± simple above, moderately branched halfway down, slightly anastomosing, uniformly thickened to some-what beaded, multiseptate, cells 7-13 ´ 1-2 mm, not or slightly swollen at the tips. Asci: clavate, 50-65 x 14-18 µm, tholus of Ophioparma-type, containing 8 spores Ascospores: hyaline, simple to one-septate, not ornamented, lacking a halo, very narrowly ellipsoid, or slightly asymmetrically subfusiform, less commonly fusiform or teardrop shaped, occasionally slightly bent, always lacking a tail, 12-25 ´ 4-6.7 mm, frequently found immature, but with a few mature spores almost always present, arranged longitudinally to diagonally in the ascus.
Conidiomata: immersed pycnidia, of variable shape, often forming in compact groups with up to 20 or more closely adjacent ostioles, the group sometimes occurring in distinctive tubercles up to 2.5 mm in diameter, sometimes occurring in "disks" which resemble apothecia, sometimes in concave hollows; lateral and basal walls of pycnidia hardly pigmented or pigmented strong brown to medium red brown, intensifying in KOH; ostioles pigmented dark green (KOH intensifying, 50% HNO3+ strong red brown), appearing black from above, the groups of ostioles often forming circular or lengthwise elongated gray to black patches on the surface of areoles, also pigmented dark green, lacking algae. Conidiophores: similar to Vobis (1980) Type VI, i.e., highly branched with both terminal and lateral conidiogenous cells. Conidia: bacilliform, straight to slightly curved, 7-9 (-11) ´ 1.0-1.4 mm.
Chemistry: the only eNA specimen has usnic acid and divaricatic acid by TLC. Elsewhere in NA occasional single specimens have been found with psoromic acid, stictic acid, norstictic acid, or atranorin as accessory compounds. Hymenial pigment (haemoventosin) deep orange in section, KOH+ strong blue, slowly becoming strong purple. Spot tests for eNA: cortex and medulla KOH-, C-, Pd-. Medulla UV+ whitish.
Ecology and distribution: Saxicolous, holarctic or subarctic in NA, barely extending south into the eNA floristic area at 53.85 N in Labrador.
Remarks: See the remarks under O. ventosa.
Ophioparma ventosa (L.) Norman, Nytt Mag. Naturv. 7:230-231, 1853.
Lichen ventosus L., Sp. pl. 2, 1753, p. 1141. TYPE: locality apparently unknown (LINN 1273:15, Ehrhart exs. 30, neotype, designated by Jorgensen et. al., 1994) [chemistry: usnic, divaricatic, thamnolic, gyrophoric (tr.) and ?psoromic acids, haemoventosin].
Parmelia ventosa Ach., Methodus, 1803, p. 166.
Lecanora ventosa Ach., Lichenogr. universalis, 1810, p. 399.
Haematomma ventosum Mass., Ric. auton. lich. cost., 1852, p. 33.
Thallus: crustose, light gray-yellow, medium green-yellow, or gray-olive (to gray or brown in European specimens), usually rimose-areolate, sometimes plicate-areolate (i.e., having strongly convex ridges with cracks in between), typically quite variable in thickness within a single thallus, to 8 mm thick (including the hypothallus) where groups of areoles form convex heaps, but more typically 2-5 mm thick; the larger areoles usually moderately convex, but sometime flat, 2-6 mm across, corticate partway down into the cracks, sometimes subumbilicate, generally shiny in luster, verrucose, microsquamulose, or cracked within, the verrucae sometime subdivided into verrucules; the areoles and mounds of areoles typically resting on a thin to thick layer of fungal tissue (which might be considered a hypothallus), which often incorporates grains of rock. Hypothallus and the lowest part of the areoles white to dark gray where exposed. Thallus occasionally partly (near the margin) or entirely of dispersed areoles resting directly on the substrate, then as thin as 0.2 mm and lacking a hypothallus.
Apothecia: sessile, usually attached over most of its diameter, to 3 mm in longest dimension, sub-round to more typically highly irregular in shape, sometimes partially subdivided. Disks gently concave to gently convex, often undulate, deep red brown to very deep red. Margin ca. 100 mm wide parallel to the excipular hyphae, and up to 400 mm thick perpendicular to the hyphae, generally not raised, smooth to finely rugose, often strongly flexuose, shiny to matte, pale-yellow to orange-yellow, almost thalline-appearing, or sometimes pigmented the same color as the disk. Margin, however, lacking algae at every stage of development except occasionally a few in the outer basal area of mature apothecia where the margin meets the thallus, usually separated from these algae by a small infold of cortex. A blue-green pigment (same as that in the ostioles of conidiomata) sometimes present in the margin of apothecia, giving the false impression of an algal layer. An excipular cortex of gelatinized cells present, 7-12 mm thick. Hymenium and epihymenium together 35-90 mm high, strongly agglutinated, inspersed at the surface and part or all the way down with crystals of deep orange pigment, hyaline, I+ blue in lower part. Subhymenium hyaline, 15-105 mm thick, I+ blue. Hypothecium: hyaline, I-, continuous with and grading into the thick fungal layer (hypothallus) in some specimens; but in other specimens, more compact than the thick fungal layer and separated from it by cortex. Exciple of radiating, highly branched, ± anastomosing hyphae originating from the hypothecium, the outer cells forming a compact gelatinous cortical layer, algae generally lacking in NA specimens. Paraphyses: ± simple above, sometimes moderately branched halfway down, slightly anastomosing, multiseptate, cells 5-13 ´ ca. 2.5 mm, mostly uniformly thickened, not or slightly swollen at the tips (to 4.5 mm). Asci: clavate, 45-60 x 14-22 µm, tholus of Ophioparma-type, containing 8-spores. Ascospores: hyaline, 3-7 septate at maturity, 35-61 x 3.3-6.1 µm, (more than 30 µm long even when immature), asymmetrically tapered, with one end blunt or fusiform and one end subulate (broadly acicular) to acicular, or sometimes becoming narrowly spatulate (i.e., with a "tail"), often helically arranged in the ascus, but also often arranged subparallel to the long axis of the ascus.
Conidiomata: immersed or often partly raised above the thallus, of variable shape, the ostioles obvious, the exposed part up to 0.3 mm in diameter, sometimes resembling black apothecia with a thalline margin, often in loose groups with up to eight adjacent conidiomata; lateral and basal walls of conidiomata not pigmented, ostioles pigmented dark green in thin section (KOH intensifying, 50% HNO3+ strong red brown), lacking alga. Conidiophores: similar to Vobis (1980) Type VI, i.e., highly branched with both terminal and lateral conidiogenous cells. Conidia: bacilliform, 6-9 ´ 1 mm.
Chemistry: all specimens with usnic acid and divaricatic acid by TLC (one or the other may occasionally be present only in trace amounts); three eNA specimens also with psoromic acid, one with salazinic acid, three with atranorin, one with zeorin, five with unidentified triterpenes. Hypothamnolic, stictic, and norstictic acid have not been found yet in eNA, but do occur in other parts of the continent (May 1997). Hymenial pigment (haemoventosin) deep orange in section, KOH+ strong blue, slowly becoming strong purple.Spot tests: cortex and medulla KOH-, C-, Pd- (or if accessory compounds are present: KOH+ yellow or orange-yellow and/or Pd+ yellow or orange). Medulla UV+ whitish.
Ecology and distribution: Saxicolous; arctic, subarctic-boreal, and alpine in NA, in eNA extending south into alpine regions of northern New England and northern New York.
The southern limit of the species is supported by published and unpublished data that is partially independent of the herbarium collections studied. O. ventosa has not been reported for the southern Appalachian Mountains (Degelius 1941; Ciegler 1997) or for the Ozarks of Missouri (Ladd 1996); and it has been found neither in the highest Berk-shire Hills of Massachusetts nor the Catskills of New York (first and second Tuckerman Workshops, unpublished lists of species).
Remarks: O. ventosa and O. lapponica are similar species, differing mainly in spore type. Although there are differences in thalline and apothecial characters, these are not always easy to use nor completely reliable. However, O. ventosa is more variable than O. lapponica. Some variants—for example, those with relatively dark thalli, sub-squamulose areoles, or thin apothecial disks—are different from any NA specimen of O. lapponica and therefore can, with experience, be identified on sight.
In NA, O. lapponica appears not to occur as far south as O.
ventosa (May 1997). If this proves correct, specimens from all but
the most northern localities in the eNA Flora region will prove to be O.
ACKNOWLEDGEMENTS: I would like to thank the curators of BM, CANL, COLO, DUKE, F, H, US, AMNH, NY and WIS for loans of material; Paula DePriest for hosting me at US; John W. Thomson for help in locating the specimen illustrated in Arctic American Lichens; Richard C. Harris, for advice and comments on the manuscript; James Hinds for advice and a draft of his own Flora paper; Scott LaGreca for comments on the manuscript; Bill Buck for general support; Arlene Olivero for assistance with computer mapping; and Donald Pfister for kindly allowing me to use the facilities at FH for my research.
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