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Script Lichen Lookalike
Morral, Marion County, Ohio March 31, 2010
I'm trying to learn a bit about lichens. Goal: be able to name the very common ones on sight, and gain sufficient facility to run an unknown through the keys. It's great fun and is going quite well, thanks mainly to gracious assistance provided by uber-friendly lichenologist members of OMLA, the Ohio Moss and Lichen Association. If you have an interest in lichens, join the club!
A well-known distinction, and the starting point for lichen identification, is the overall growth form, of which there are three main types: fruticose, foliose, and crustose (i.e., shrubby, leafy, and crusty).
FRUTICOSE lichens are stalked, pendant, or shrubby, with no clearly distinguishable upper and lower surface. The quintessential fruticose lichens in Ohio belong to the genus Cladonia. Most Cladonia species begin development as a scaly (squamulose) primary thallus. ("Thallus" is the general term for a relatively featureless body, the kind had by a fungus, algae, or even some plants, wherein specialized intricate organs are lacking.) Later on, a hollow upright structure, the "podetium" develops. Podetia, depending upon the species, may be pointed, clubbed, or topped by cups and, depending on the species, may bear spore-producting apothecia.
Here are two cladonias seen recently. The first is "peg lichen," C. polycarpoides. This species has fairly long primary squamules that persist after the podetia have formed. Each podetium ends in a large brown apothecium. Peg lichen is common on soil in old fields and roadside banks.
Peg lichen at OSU-Marion Campus Prairie. March 31, 2010
Wand lichen, Cladonia rei, is a cup-forming fruticose lichen that produces apothecia at the tips of numerous proliferations of the cup margins, resembling a little star, giving the overall impression of a magic wand. The primary squamules (not visible in the photo below because it focuses on the upper parts) are small and sparse.
Wand lichen at Dawes Arboretum. April 4, 2010.
FOLIOSE lichens have a more or less "leafy" growth form, distinctly dorsiventral, and varying in the degree of attachement to the substrate, from completely adnate to only centrally attached (umbilicate). Foliose lichens seem to be especially common on tree trunks, comprising most of the ones on this hardwood at Dawes Arboretum in Licking County, Ohio.
Foliose lichens on brak (bark, whatever).
April 4, 2010. Licking County, Ohio.
There are several species here ...it's a vertiable lichen salad! The lowermost large one is "powdered ruffle lichen," Parmotrema hypotropum. Some of the distinctive features of this lichen are its pale greenish gray upper surface, an undersurface that is black with a broad pale margin, the presence of long eyelash-like cilia on the margin, and, also on the margin, localized regions called "soralia" where powdery vegetative propagules called "soredia" are produced. Soralia can be seen along the edge of the thallus in the lower right corner of the picture.
Powered ruffle lichen on bark. April 4, 2010. Licking County Ohio.
CRUSTOSE lichens are in contact with the substratum at all points and lack a lower tissue layer (the cortex) had by the other lichen types. Crustose lichens cannot be removed intact without removing a portion of the substrate as well. Here's the very unremoved substrate of a couple of crustose lichens noted recently at the OSU-Marion campus.
Concrete-based pipe thing where crustose lichens grow. Marion, Ohio. April 2, 2010.
The concrete is substrate for a couple of crustose lichen species, one of which is dimly visible as a yellow patch along the upper edge, seen close-up as presenting a great many bright yellow cup-shaped spore-producing bodies (apothecia). Before testing it with KOH, the best guesses for the yellow one were Candelariella, Candellaria, or Caloplaca. After determining it is K- (no reaction), signs point to "hidden goldspeck lichen," Candelariella aurella, said to be the most common of the goldspeck lichens on calcareous rock. And that odd gray one in the lower left that I initially mistook for a dead old yellow one (!) is a certain call. That's "mortar rim-lichen," Lecanora dispersa, described by Brodo, et al. in "Lichens of North America" as "a survivor," ...going on to say "In New York City, it is the only one to persist in central Brooklyn (on concrete fences in Prospect Park)." They explain that the ability of this lichen to tolerate the urban environment is attributable to the fact that it occupies a calcium-rich substrate. This buffers the effects of acid precipitation, often deadly to lichens.
Crustose lichens on concrete. April 2, 2010. Marion County, Ohio.
An odd little sub-group of crustose lichens are the "crustose script lichens." These produce specialized peculiar elongate, often carbon-black, apothecia termed "lirellae." Frequently found growing on bark (but some species dwell on rock), they can resemble writing in some cryptic Druid language. Here's a hickory tree growing at the edge of a woodland upon which is one of them, apparently made by a cryptic Druid with a messy handwriting, since it's called "scribble lichen." It's Opeographa varia.
Hickory tree substrate of scribble lichen.
March 31, 2010. Morral, Marion County, Ohio.
The scribble lichen does indeed resemble little random felt-tip marker markings on the bark. The area between the scribbles is occupied by a thin thallus that, when scratched, exposes green algal cells, constituting the only obvious evidence that this is indeed a lichen.
There are some unlichenized fungi that, while not closely related to the script lichens, constitute an interesting example of convergent evolution because they strongly resemble some script lichens. While out lichen-hunting I found one of these script lichen lookalikes. It's growing on this white oak in an open woodland in Morral, Marion County, Ohio.
Scribble lichen on hickory bark.
White oak substrate of possible Hysterium barrianum.
Morral, Marion County, Ohio.
Like most lichens it's an ascomycete fungus. It's in the family Hysteriaceae, about which there is a wealth of great information on this web site. Its specialized ascoma, termed a "hysterothecium," is elevated above the substate and opens by a narrow longitudinal slit along the top. Based on spore dimensions (45 x 11 micrometers) and morphology (inset; note 7 crosswise septa) this is be the recently described Hysterium barrianum.
Script lichen lookalike, Hysterium barrianum.
Leprose lichen Lepraria lobificans loves limestone
like large leafy liverwort Lophocolea loves logs.
Delaware County, Ohio. March 21, 2010
In woods on the high east bank of the Scioto River are outcroppings of Columbus Limestone of Devonian age upon which grows an exceptional lichen.
Limestone home of the fluffy dust lichen. March .21, 2010
It's a one of a few genera of lichens that are "leprose," i.e. having a thallus (or at least a thallus surface) composed entirely of powdery soredia. Soredia are vegetative propagules consisting of a few algal cells entwined and surrounded by fungal filaments. This odd lichen lacks any well organized tissues --outer cortex and inner medula --had by many lichens. It's just undifferentiated fluff! Considered a crustose lichen because it is in contact with the substratum at all points and lacks a lower cortex, this is the most common of the well named "dust lichens" that constitute the genus Lepraria. It's fluffy dust lichen, Lepraria lobificans.
Fluffy dust lichen. March 21, 2010. Delaware County, Ohio.
Nearby, a community of mosses and liverworts occupies a decorticated log.
Log habitat of mosses and liverworts. March 21, 2010.
One of them is Lophocolea heterophylla (family Geocalycaceae), a fairly large by liverwort standards leafy liverwort, the leaves of which overlap in a succubous manner, i.e., the forward edge of each leaf is situated beneath the base of the leaf ahead of it. Note near the center of the photo, a developing sporophyte. According to the Liverwort Tree of Life evidence is accumulating that liverworts (Phylum Marchantiophyta) were the first land plants to diversify on land some 500,000 million years ago and as such are the oldest living lineage of land plants.
Lophocolea heterophylla. March 21, 2010. Delaware County, Ohio.
Nearby, an American sycamore proclaims "Trespassing Violators will be Ted." That doesn't make any sense. Who's Ted?
Sign-eating American Sycamore. March 21, 2010.
Wee White Wildflower
Delaware County, Ohio March 21, 2010
Snow trillium (Trillium nivale, Liliaceae, the lily family) is our smallest, earliest, and cutest trillium.
Snow Trillium. Delaware County, Ohio. March 21, 2010.
Wee White Weeds
near Terradise Nature Preserve, March 20, 2010
A couple of small annual weeds are exceedingly common and abundant in disturbed open ground, including agricultural land adjacent to Terradise Nature preserve in Marion, Ohio. They're flowering now. How is that possible? Don't flowering plants need time to grow from seed? Yes, they do. And the way these plants accomplish the feat is by being "winter annuals." The seeds of these winter annuals germinate in fall or winter, grow during autumn and also during winter warm spells, remaining dormant but alive when it's really frigid. Come spring, and "ta-da!!," flowers, then fruits, releasing seeds that wait until cool weather to sprout, avoiding summer drought (the adverse season).Harbinger-of-spring (Erigenia bulbosa)
Winter-annual weeds in farmland. Marion, Ohio. March 20, 2010.
Hoary bitter-cress, Cardamine hirsuta (Brassicaceae, the mustard family) is known by its abundant basal pinnatifid leaves that are slightly hairy with long, widely spaced hairs, and small white flowers.
Hoary bitter-cress. Marion, Ohio. March 20, 2010.
Mustard fruits are modifed capsules --dry many-seeded fruits. Some genera produce long narrow ones called "siliques," while others produce short squat fruits called "silicles." Cardamine fruits are siliques. This is just beginning to develop in the leftmost flower in the picture below. Other mustard family traits that can be seen here are the 4 separate sepals, and 4 separate petals, equal in size.
Cardamine flower cluster. March 20, 2010, Marion, Ohio.
The other winter-annual weed here today is common chickweed, Stellaria media, a member of the Caryophyllaceae (pink family). This family consists mainly of opposite-leaved herbs with radially symmetric flowers having 5 petals and sepals, and 5 or 10 stamens. The garden carnation is a pink, as are sweet William and baby's-breath.
Common chickweed. Marion, Ohio. March 20, 2010.
Each of chcikweed flower's five petals are deeply bifid, giving the appearance of a 10-petaled flower.
Common chickweed flower. Marion, Ohio. March 20, 2010.
Sharon Woods MetroPark, Westerville, Ohio
March 19, 2010
Excepting oddities like skunk cabbage and various flowering trees that are oblivious to pollinators (e.g., silver maple, see below), our earliest wildflower is one aptly dubbed "harbinger of spring," Erigenia bulbosa (Apiaceae, the carrot family). The species may be fairly frequent in our deciduous forests, but is easily overlooked because it protudes only a mini-bit above the leafy ground.
Harbinger of spring at Sharon Woods, Franklin County, Ohio. March 19, 2010.
Erigenia is also known locally as "pepper and salt" owing to the sharp contrast between the black stamens and the white petals. Although perhaps not obvious at first glance, Erigenia is indeed a fairly typical representative of its family. Note first the leaves; they're quite lacy, thrice compound in fact, a common leaf complexity in the family. Moreover, the family is also known as the "Umbelliferae" because of its characteristic inflorescence type, the umbel. An umbel is a flower cluster having stalked flowers that are all attached at one point on the stem, suggestive of an umbrella. Many umbelliferous plants, including Erigenia, bear compound umbels consisting of little umbels ("umbellets") all of which are attached to one spot. The compound umbels of Erigenia, including the one shown here, typically have three such umbellets.Corticolous Cryptogams (Epiphytic Excitement)
If you examine the flowers closely (mouseover to see), several more apiaceous features jump out: an inferior ovary (evident on the old petal-less flower on the left), a bicarpellate ovary (evident by the dual style-branches on that flower), pale-colored radially symmetric flowers with sepals "obsolete" (so small they are essentially absent) ...an overall form not much different from, say, Queen Anne's lace.
(A Lichen and Liverwort on American Elm)
Delaware County, Ohio. March 18, 2010
An American elm tree at a a research preserve (Kraus Woods) in Delaware County, Ohio is the substrate for several delightful epiphytes.
Elm bark habitat of liverworts and lichens. Delaware , Ohio. March 18, 2010.
The liverwort is Frullania eboracensis (family Frullaniaceae), with smallish ovate, distinctively wide-spaced leaves and a pattern of spawling tree-like doodles on bark, with stems that are tightly adherent to it.
Frullania eboracensis on elm bark. Delaware County, Ohio. March 18, 2010.
Leafy liverworts, which are more numerous than the relatively featureless thallose types chosen to depict liverworts in textbooks, can be disturbingly moss-like in appearance. Leafy liverwort leaves are much more intricate than those of mosses, the leaves of which are simple, all alike or nearly so, and arranged in three rows (ranks) around the stem. By contrast, the leafy liverwort stem typically lays flat, and the upper and lower leaves are differentiated. There is a pair of upper leaves which in many genera have their rear edges folded under (conduplicate, forming a dorsal lobe ventrsl lobes) and/or are erose or divided (sometimes elaborately so). There is nearly always a single row of much smaller lower leaves.
Microscopically, the leaf-lobing of Frullania is most distinctive. The large dorsal lobe, the only leaf part that you see when the plant is growing on the tree, is fairly ordinary --ovate, wide-spaced, and incubously overlapping. Here's a fairly low power microscopic picture of this species taken a few years ago, wheren the large dorsal leaf lobes are blocking a clear view of the reduced ventral leaf lobes.
Frullania eboracensis, dorsal view. Note large dorsal lobes overtopping small ventral lobes.
The Frullania ventral lobe is extremely differentiated. Usually referred to as a "lobule," it is so very concave that is resembles a cup. Its function is not known with certainty, but is suspected to aid in water retention or absorption. Here's a photo of the underside of Frullania.
Frullania eboracensis ventral view. Note cup-like lobules.
The underleaves of Frullania eboracensis are small, wider than long, and bidentate. The photo below is a micrscopic ventral view focussing on a lower leaf, showing also the cup-like lobule in the mid-background, and the large dorsal lobe in the distance. Some of the the cup-like lobules of the Kraus woods Frullania are home to an amazing little animal --a rotifer (Phylum Rotifera) which I am assuming is one of the "bdelloid" rotifers (Class Eurotatoria, Subclass Bdellloidea) famous for consisting only of asexually reproducing females. Rotifers are surprisingly complex animals, considering they are microscopic. The are filter-feeders that attach themselves to the substrate and draw in their food --bacteria, algae, and organic debris --by means of water currents established by two helicopter-like sets of whirling cilia.
MOUSEOVER the IMAGE to see ROTIFER
Frullania underleaf and (mouseover to see) rotifer-inhabited lobule. Delaware, Ohio. March, 2010.
Here's a video of the rotifer in action.
Rotifer filter-feeds from within Frullania lobule.
Seeing those little critters snuggled comfortably in those extraordinary little cups, it's tempting to think that this might be an elegant mutualism like the one where aggresive bodyguard ants live in swollen hollow Acacia thorns in the tropics. Alas, probably not, or at least not in an obvious way, is this a mutualistic symbiosis. Rotifers in Frullania lobules were the subject of interesting study by Mary Puterbaugh and colleagues that was published in The Bryologist in 2002. Noting that rotifers are frequently found in the lobules, they propose three plausible hypotheses for the effect of the rotifers on the liverwort, and four hypotheses for vice versa effects.
Here's Table 1 from Puterbaugh, M.N., J.J. Skinner and J.M. Miller. 2002. A nonrandom pattern of rotifers inhabitating lobules of the hepatic, Frullania eboracensis. The Bryologist 107:524-530.
In an attempt to lend strength to a working hypothesis that rotifers might benefit the liverwort, and that liverworts helped thusly would be more likely to reproduce, these investigators tallied rotifer presence/absence on thousands of lobes of 84 plants from a site in New York State, recording whether the plants were male-expressing, female-expressing, or nonexpressing, as well as plant size and position on the plant. There was no difference with respect to the plant's reproductive status, but there was a strong tenedency for rotifers to be more frequent in lobules at the edges of plants (nearly 50% of the lobules occupied) compared with central ones (approx. 20% of lobules occupied). Their data "provide tantalizing evidence that the microhabitat throughout a single epiphytic plant is not the same for the rotifer and the explanation for this pattern may provide insight into relationships between bryophytes and invertebrates in general."
Frullania has barkmates. Notable for its abundance and ubiquity on hardwood trees and rocks throughout the northeastern U.S. and adjacent Canada, even in shady areas, but inconspicous due to its small size and brown color is an elegant foliose lichen, orange-cored shadow lichen, Phaeophyscia rubropulcra. Along with the brownish Phaeophyscia is a bit of a gray apothecium-bearing foliose lichen I'm guessing is Physcia stellaris.
Foliose lichens on elm bark in Delaware County, Ohio. March 18, 2010.
A diagnostic character of Phaeophyscia rubropulcra is the color of the interior layer --the "medulla" is orange-red! This is evident if you carefully scrape away the upper layer (the cortex). Ray Showman and Don Flenniken in their excellent manual "The Macrolichens of Ohio" tell us that this lichen is frequently grazed by slugs that eat the upper cortex, exposing the orange medulla.
Phaeophyscia rubropulchra, with cortex removed to show orange-red medulla.
Silver maple (Acer saccharinum, family Sapindaceae) is flowering. The flowers are in unisexual clusters. At the Marion (Ohio) Cemetery, some trees seem to be entirely staminate (male), and some entirely pistillate (female). Other trees are mixed, with both male clusters and female ones. Here's a staminate flower cluster. Note the long slender filaments, an adaptation to cast pollen into the wind.
Silver maple staminate flower cluster. March 17, 2010.
The female flowers present rabbit-ears-antennae-like stigmas that snag pollen drifting about.
Silver maple pistillate flower cluster. March 17, 2010.
A friend in Delaware, Ohio invited me to visit her profuse and lively display of early-blooming garden flowers, including these crocuses. Crocuses are in the Iris family (Iridaceae). The expensive spice saffron is the dried styles and stigmas of an autumn-flowering Mediterranean species of Crocus.
Crocus in bloom in Delaware, Ohio. March 17, 2010.
There is an interesting variety of winter aconite growing in my friend's garden. According to the ancient "Doctrine of Signatures," whereby a plant's utility in treating ailments of a particular body part is revealed by its resemblance to that body part, these plants can fix eye problems!
Winter aconite, variety "ocularis" in Delaware, Ohio. March 17, 2010.
A fabulous filbet shrub is in bloom. This member of the birch family (Corylaceae) exhibits a fairly common nut-tree pollination syndrome, that of being monoecious, with staminate flowers uber-numerous and teeny-tiny, presented in long drooping catkins. The female flowers are fewer in number. In this case the female flowers are in minuscule ovoid catkins, one of which is seen in the picture below cradled by the peducle of the terminal male catkin.
MOUSEOVER the IMAGE to see ZOOM-CROP of FEMALE CATKIN
Tales from the Cryptogam
Columbus, Ohio. March 14, 2010)
An especially natural-looking mausoleum at Greenlawn Cemetery is the substrate for a sweet suite of mosses and lichens, organisms that, along with ferns and other primitive vascular plants, are sometimes called "cryptogams," a term that literally means "hidden marriage" because of the secrecy, relative to that of seed plants, of their reproductive methods.
Cryptogam substrate at Greenlawn Cemetery. Columbus, Ohio. March 14, 2010.
This is very spooky. There's something incubous here! Is is a male evil spirit, or a vexing problem? No, it's just an incubous liverwort. "Incubous," not quite the same word as "incubus," is a leaf-arrangement term that describes flat-laying liverworts the leaves of which overlap (Venetian blind-like) such that the forward edge of each leaf lays on top of the back edge of the leaf in front of it. Incubus leafy liverworts are less numerous than succubous (not succubus, the female evil spirit) ones.
Here's a picture. The liverwort is Porella platyphylla (family Porellaceae), a very widespread species of leafy liverwort. Leafy liverworts are more common than the flat and relatively featureless thallose ones most often shown in textbooks.
Porella platyphylla, an incubus leafy liverwort. March 14, 2010.
Alongside the liverwort are some mosses. Distintively dark, almost blackish when dry but looking quite green today, patches of Schistidium rivulare (family Grimmiaceae) are abundant, along with a very bright green moss that is "julaceous,"i.e., smoothly cylindric like a worm or a catkin --Entodon seductrix (family Entodontaceae).
Two mosses mingle. March 14, 2010.
There's a ghostly white foliose lichen here. Was it scared by the incubus (liverwort) perhaps? No, it's actually white because it is extremely "pruinose" (having a powdery bloom) all over the upper surface. This is probably Physconia leucoleiptes.
Ghostly white foliose lichen at Greenlawn Cemetery. Columbus, Ohio. March 14, 2010
Mildflowers (urban garden plants)
Columbus, Ohio. March 10, 2010
Early-blooming garden plants might be called "mildflowers." While not wildflowers, they are nonetheless intriguing and welcome evidence that spring has sprung. Winter aconite (Eranthis hyemale, family Ranunculaceae) is a Eurasian species that reportedly also occurs sparingly in wild places, but it is not regarded as invasive.
Winter aconite. Columbus, Ohio. March 10, 2010.
A common asscociate of the aconite is snowdrops, Galanthus nivale. This is a member of the Amaryllidaceae, a monocot family closely related to the Liliaceae, differing from the lilies primarly in its inferior ovary. While we have no native members of the Amaryllidaceae in Ohio, we do have two genera that were formerly placed in it: Hypoxis (star-grass, family Liliceae) and Manfreda (American aloe, Agavaceae).
Snowdrops. Columbus, Ohio. March 10, 2010.
Iconically Poking Through The Snow
Skunk Cabbage in Delaware, Ohio. March 8, 2010
Skunk cabbage, Symploparpus foetidus (Araceae, the arum family) is one of only a few plants that exhibit thermogenesis --they get hot! The utility of generating heat is apparently to increase the volatility of odoriferous compounds that attract early-emerging pollinators.
Skunk cabbage has melted the snow around it. March 8, 2010. Delaware, Ohio.
It's intriguing to see circular patches of melted snow surrounding the infloresences.
Skunk cabbages have melted the snow around themselves. March 8, 2010. Delaware, Ohio.
While the primary function of the thermogenesis is to facilitate pollination, it seems reasonable to assume that the snow-melting effect is beneficial also. Plants buried beneath snow would be inaccessible to insects.
Skunk cabbage in a little snow cave. March 8, 2010. Delaware, Ohio.