Coral Fungi – Clavariaceae

Crown – tipped Coral is one of many fungi that have a branching pattern similar to ocean corals

Common Name: Coral Fungus – The branching of the fungal thallus resembles the calcium carbonate structure of  ocean corals. Other common names are applied to differentiated shapes, such as worm, club, or tube fungi for those lacking side branches and antler fungi for those with wider, flange-like appendages. An extreme is cauliflower fungus which looks nothing like coral but is usually included in the coral-like category in field guides. The common Crown-tipped coral is depicted; the ends of the coral segments have tines like miniature crowns.

Scientific Name: Clavariaceae – The family name for the coral fungi is derived from clava, the Latin word for “club;” the type-genus is Clavaria. The coral fungus above was originally Clavaria pyxidata, became Clavicorona pyxidata, and is now Artomyces pyxidatus. Pyx is from the Greek word pyxos meaning “box tree” from which boxes were made (and the etymology of the word box – a pyx is a container for Eucharist wafers). The implication for its use as a name for this species is “box-like.”

Potpourri:  Coral fungi look like coral. The verisimilar likeness can be so convincing that it seems plausible that they were uprooted from a seabed reef and planted in the woods for decoration. The delicate ivory and cream-colored branches rising in dense clusters from a brown-black dead log are one of the wonders of the wooded paths sought by those who wander there. There is an abiding benefit to have some knowledge of the things that nature has created and coral fungi is a good collective mnemonic to apply to the group that surely must be closely related. And so it is  for the traditionalists steeped in the lore of musty mushroom field guides who are referred to collectively as the “lumpers.”  The new world order of DNA has taken the science of biology on a wild ride with many hairpin turns and dead ends; in the case of mycology, the train has left the tracks more than once. Coevolution … that which created a marsupial mouse in Australia unrelated to the placental house mouse everywhere else … globally demonstrates Darwin’s vision. Fungi that branch is a natural evolutionary path for two individual organisms that started at different places and times.  The diaspora of species from one genus to another in search of a home on the genetic tree of life has exploded the coral fungi into fragments. This is the realm of the “splitters,” the subdividers for whom a bar code will become the only true arbiter of species. There is of course a hybrid middle ground, acknowledging the latter but practicing the former, the province of most mushroom hunters.  

Like all epigeal fruiting bodies extending upward above the ground from the main body of a fungus, which is hypogeal or below ground, the branching arms of coral fungi function to support and project the spore bearing reproductive components called basidia. Gilled or pored mushrooms maximize the number of spores they can disperse by creating as much surface area as possible in the limited space beneath the cap or pileus. Similarly, coral fungi branch again and again or extend myriad singular shafts to get as many fingers of spore bearing surface into the air as possible. [1] The topology of using multiple  extensions into a fluid medium is one of the recurring themes of evolution ― coevolution. In this case, it has nothing to do with fungi per se. They look like coral because real coral is doing essentially the same thing; the namesake polyps secrete a type of calcium carbonate called aragonite to form protective exoskeletons in reefs that extend outward into the water where their food floats by.  To extend the analogy to the rest of biology is a matter of observation. Trees send branches covered with photosynthesizing leaves toward the sun and roots toward the water and minerals of the earth where they encounter the branching mycelia of fungi.

Fungi have evolved to distribute reproductive spores with different mechanisms that could only have been naturally selected by the variations in form and function of random mutation. Among the more creative methods are the puffing of puffball spores out a hole in the top by the impact force of raindrops, the odorous spore-laden goo of stinkhorns that attracts insects seeking nutrients, and the redolence of truffles sought by burrowing or digging animals as food digested, their spores excreted intact. The coral fungi are among the most primitive of all basidiomycete fungi in having their club-shaped spore bearing basidia positioned along the upper reaches of each prong so that they can be carried away by either wind or water. [2] Having more fruiting bodies with more branches creates more spores, which is why coral fungi are frequently found growing saprophytically in dense clusters on dead tree logs or growing in mycorrhizal clusters on the ground. Simply sticking indistinguishable club shapes into the air with a bunch of short rods with spores attached to the end is the most straightforward way to disperse them for germination.

The phylogenic diversity of the coral fungi belies their similar ramified appearance. Historically, structure was thought to be the basis for taxonomic classification, an assumption that works reasonably well with plants and animals but not with fungi. The delicate and colorful appearance of the coral fungi brought them to the attention of the earliest naturalists, who grouped them according to shapes. Since fungi were then considered members of the Plant Kingdom (Subkingdom Thallophyta), this was consistent with practice. The French botanist Chevallier placed them in the order Clavariées in 1826 with only two genera, Clavaria and Merisma noting that “se distingue du premier coup d’oeil” – they can be identified with a fleeting glance in having “la forme d’une petit massue” – the form of a little club. [3] The assignment of fungi to families according to form lasted for over a hundred years until the nuances in microstructure and spore appearance initiated cracks in the biological foundation. Toward the end of the last century the fungi were recast as one of five different kingdoms, the foundational genus Clavaria was dissected into six genera with derivative names like Clavulina (little club)  and Clavariadelphis (brother of Clavaria), which is how they appear in the most popular fungi field guides. [4]

In spite of the distinctive shape that suggests a unique origin, coral fungi are agarics, the historical group name for almost all gilled fungi. What is now the order Agaricales is comprised of over 9,000 species, containing over half of all known mushroom forming macrofungi assigned to one of 26 families with about 350 genera that range from Amanita to Xerula. Carl Linnaeas, who established the first taxonomic structure in biology with the publication of Systema Naturae in the 18th century, placed all gilled mushrooms in a single genus that he named Agaricus. One hundred years later, Elias Fries published Systema Mycologicum, which separated the agarics into twelve genera based on macroscopic features such as the structure of the spore bearing surface or hymenium (e.g. gills, pores, teeth, ridges, vase-shaped) and spore color (white, pink, brown, purple-brown, or black). Six groups of basidiomycetes were recognized based on the shape of the sporocarp or fruiting body ― “coral-like fungi” was one of them. While there was some expansion of genera over the ensuing decades, the so-called Friesian approach to gilled mushroom identification has persisted and  is what is still generally in use, spore print color and all. The use of field characteristics is crucial to the practical application of mycology that serves the community of foragers looking for edible species and other aficionados who enjoy their company. [5,6]

Over the last several decades, the use of DNA to map out the true phylogenetic relationships has upended the traditional taxonomy based on macroscopic structure and spore color. Unravelling the complex weave of evolutionary threads from one species to its predecessor  is a monumental task that is just now gaining momentum. The goal is to determine the real or cladistic family tree so that a clade, the term adopted to refer to all species with a common ancestor, can be established with certainty. In one analysis, the agarics fell into six major clades, or single-ancestor groupings named Agaricoid, Tricholomatoid, Marasmioid, Pluteoid, Hygrophoroid and Plicaturopsidoid. The coral fungi are in the latter, which diverged from all the other agarics at the earliest evolutionary branching in the Cretaceous Era some 125 million years ago. It is not unreasonable to conclude from this analysis that the coral fungi evolved a reliable and efficient method of spore dispersal early on and have thrived ever since, branching out to form new species all using the same technique. It is now equally evident that the shape of a fungus does not necessarily establish its proper branch in the family tree. The agarics, now the Eugarics Clade, not only has fungi shaped like mushrooms and coral, but also puffballs like Calvatia and Lycoperdon. Likewise, shapes extend across multiple clades.  For example, coral-shaped fungi also appear in the Russuloid Clade (Russulas)  as Artomyces as pictured above and Sparassis as pictured below in the Polyporoid Clade (Polypores). This is then the dichotomy between the taxonomists of the old school steeped in the Linnaean traditions of field identification and the DNA systematists of the new school for which only the laboratory will do. [5,6]

The new biological life history of coral fungi is still subject to the findings of the most recent research paper devoted to the group and it may be decades before a settled taxonomy emerges. As a brief and incomplete history, in 1999 “four lineages containing cantharelloid and clavarioid fungi were identified,”  with the clavarioid containing most of the corals, but also noting that “Clavicorona is closely related to Auriscalpium, which is toothed, and Lentinellus, which is gilled.” [7] In 2006, it was acknowledged that coral shaped fungi must have evolved at least five times over the millennia and that the “evolutionary significance of this morphology is difficult to interpret because the phylogenetic positions of many clavarioid fungi are still unknown.” The new genus Alloclavaria was added to accommodate the unique fungus Clavaria purpurea  “not related to Clavaria but derived within the hymenochaetoid clade,” which consists mostly of bracket fungi. [8] Seven years later, the coral fungus family was found to consist of four major  clades:  Mucronella,  Ramariopsis-Clavulinopsis,  Hyphodontiella, and Clavaria-Camarophyllopsis-Clavicorona. This thorough phylogenic analysis of 47 sporocarp sequences merged with 243 environmental sequences concluded that “126 molecular operational taxonomic units can be recognized in the Clavariaceae … an estimate that exceeds the known number of species in the family.” [9] Phylogenic studies are continuing.

Returning to the more mundane walk through the woods looking for coral fungi, the two most pressing questions concern edibility and toxicity. Neither of these subjects is broached in the scientific literature, and, like most fungi, data points are empirical, relying on random trial and error anecdote. For coral fungi, this is complicated by the fact that most are small and delicate and therefore rarely sampled by those seeking massive brackets of Chicken-of-the-woods and yellow clusters of chanterelles. Edibility has been a question ever since Chevalier first singled them out in 1826, noting that “Presque tout les clavaires  fournissent a l’homme une nouriteure saine, on mange ordinarements les plus grosses”  –  almost all are good to eat but only pick the big ones,  and “Elles n’ont aucune qualité vénéneuses; quelques-une ont une saveur amèrenone are poisonous but some are bitter. [10] This sweeping assurance cannot have been the result  of a thorough assessment, as there are good and bad corals.  Modern guides are more circumspect, offering a range of information about edibility from choice to poisonous with caveats about having a laxative effect on some people and causing gastrointestinal distress in others. Many are of unknown edibility and likely to remain so. There is one standout worth noting that has the hallmarks of broad acceptability. The Cauliflower Mushroom (Sparassis americana – formerly crispa) is large, unusual, and common. It neither tastes nor looks much like a cauliflower. The “Elizabethan ruff of a mushroom” [11] is hard to miss and there is no doppelganger to fool the hapless hunter.

The Cauliflower Mushroom looks nothing like coral, or cauliflower for that matter. More like a neck ruff of Elizabethan England.

References:

1. Aurora, D. Mushrooms Demystified, Ten Speed Press, Berkeley, California, 1986 pp 630-658

2. Schaechter, E. In the Company of Mushrooms, Harvard University Press, Cambridge, Massachusetts, 1997, p.49   

3. Chevallier F. Flore Générale des Environs de Paris, Ferra Jeune, Paris, France, 1826  p. 102.

4. Lincoff, G. National Audubon Society Field Guide to North American Mushrooms, Alfred A. Knopf, New York, 1981, pp 398-414.

5. Matheny P. et al “Major clades of Agaricales: a multilocus phylogenetic overview”  Mycologia August 2006, Volume 98 Number 6 pp 982–995.

6. https://www.mykoweb.com/articles/Homobasidiomycete_clades.html        

7. Pine E. et al “Phylogenetic relationships of cantharelloid and clavarioid Homobasidiomycetes based on mitochondrial and nuclear rDNA sequences”. Mycologia. 1999. Volume 91 Number 6 pp 944–963.

8. Dentinger B and McLaughlin D. “Reconstructing the Clavariaceae using nuclear large subunit rDNA sequences and a new genus segregated from Clavaria”. Mycologia. Volume 98 Number 5 September 2006 pp 746–762.

9. Birkebak J et al. “A systematic, morphological and ecological overview of the Clavariaceae (Agaricales)”  Mycologia. Volume 105 Number 4, February 2013, pp 896–911.

10. Chevallier, op cit. p.104

11. Lincoff, op cit. p. 412.

Red-spotted Purple Butterfly

The Red-spotted Purple is mostly purple and has red spots at the wing tips

Common Name: Red-spotted Purple and White Admiral – Butterfly names are in most cases descriptive, using color and patterns as leitmotif. The mostly dark blue wings tinged with enough red to produce purple culminating in red wing spots provides one of the more mnemonic names. The alternative name White Admiral is a result of one of the more tantalizing tales of the lepidopterans as they change colors and patterns in mimicry detailed below.

Scientific Name: Limenitis arthemis –  The genus name literally means harbor goddess in Greek. The nautical association is apparently related to or is a result of  the fact that they are called the admiral butterflies, as in White Admiral. The species name is from Artemis (Diana in Roman mythology), the Greek goddess of the hunt and hence the woods. A butterfly as metaphor for a goddess captures the graceful beauty of both.    

The White Admiral has a single broad white stripe – like a US Navy admiral

Potpourri: The  Red-spotted Purple and White Admiral are the same species, Limenitis arthemis. Mimicry, the term for an animal mimicking another object in shape and/or color, is an evolutionary and genetic  response to  the inexorable tug of survival. Although it may seem especially notable in this case because of the striking result afforded by the difference between white striped and stripe-less wings, mimicry in its broadest sense is widespread. Some prey animals change colors according to age and season to provide better camouflage. The spotted fawn turns light tan as a doe or buck in summer and darker in winter to match the scenery. Predators must do the same in order to hide from their quarry long enough effect the coup-de-grace at the last moment. There are no black panthers, only melanoid leopards and jaguars becoming night stalkers (both are in the genus Panthera). Aposematism is similar to mimicry in that coloration is used to ward off predators. But rather than being cryptic, the colors stand out  against the background in sharp contrast, alerting the wary predator that poisons there lurk. The juvenile red eft of the red-spotted newt is a good example of aposematism … a defenseless amphibian that protects itself with vivid orange hues similar to those  used by hunters to accentuate visibility.

White Admirals, the northern version of the Red-spotted Purple, are named for their prominent white stripes. It is perhaps only coincidence that the progression of officer ranks in the U.S. Navy ranges from an ensign’s single narrow to a single broad stripe for an admiral. The Red Admiral (Vanessa atalanta) which has a similarly placed red stripe is otherwise unrelated. Boldly contrasting prominent stripes on two species suggests purpose and coevolution. While striping may be related to species or mate recognition, it is more likely a matter of predator avoidance, the moving flashes of colored streaks creating a disorienting stroboscopic effect. [1]  On progressing geographically southward, the White Admiral’s broad stripe disappears and the red spots move forward to the edge of the wing tip to become both red-spotted and purple. This rather extraordinary transformation is a combination of the aping of mimicry and the warning of aposematism, a hybrid scheme called apatetic in general or Batesian in particular.  The wing is now more uniformly dark in color, resembling that of a butterfly of a different genus and species ― a poisonous doppelgänger.

The Red Admiral has a single red stripe

It is widely known that the Monarch butterfly is unpalatable to birds because its caterpillars eat milkweed (Asclepias syriaca) that produces cardiac glycosides that are toxic to most animals. It is mimicked by the Viceroy (Limenitis archippus) butterfly, a generic cousin of the Red-spotted Purple, as a matter of enhanced survival. [2] The Green Swallowtail butterfly (Battus philenor) is better known as the Pipevine Swallowtail because its larvae feed on Dutchman’s Pipe (Aristolochia durior), a vine that produces a toxin called aristolochic acid. Since the range of the toxic Pipevine Swallowtail butterfly extends only as far as its namesake food, it is a southern butterfly because that is where the vines are. [3]  The change is not cognitive choice, but rather choice by chance. The White Admirals that ventured south with less prominent stripes survived more frequently since they were more likely to be avoided by predators. Over time and subsequent mating of diminished stripe White Admirals, the stripes disappeared altogether and the Red-spotted Purple became the southern variant, sometimes listed as a separate subspecies Limenitis arthemis astyanax. The name extends the mythological association to include Astyanax, the son of Trojan hero Hector who was defenestrated by the Greek Achilles so he could not avenge the death of his father.

The Pipevine Swallowtail is copied by the Red-spotted Purple to escape predation by birds. It is called Batesian mimicry.

The kaleidoscopic patterns of butterfly wings are among the most artistic creations of nature. Their evolution that began during the Cretaceous Period 150 million years ago was marked by three random mutation “inventions” that radiated in time and space along the way to produce the 18,000 plus extant named species. [4]  The first and defining mutation was wing scales from which the name of the order Lepidoptera was derived, lepis meaning scale and ptera wing in Greek.  Scales are genetically modified sensory bristles, that became flat. senseless, and slippery, probably to avoid capture … the survivors passed the scale genes along. The second invention was changing the scale colors, possible because each scale is from a single cell with control of hue and texture, the combination producing different shadings and sometimes even iridescence. Lastly there was pattern, the genetics of placing colors in ordered arrangement. Spots in general and eyespots in particular start in the caterpillar stage, where an organizer puts them in the right position on the wing, a disc at this point. Colors are added in the chrysalis phase so that the adult butterfly wing emerges after metamorphosis with spots. These are usually at the margins of the wing so that a predator would first strike there, removing only a small portion of the wing as the butterfly flitted to safety. The efficacy of this is demonstrable, as many lepidopterans are found with a bite out of one wing. [5]

Butterflies are among the most studied of all animals, surely more a matter of beauty and ease of net capture than for their scientific import as just another type of insect. Henry Walter Bates spent eleven years in the Amazon rainforest in the mid-nineteenth century, identifying 8,000 species that were then new to science, many of them butterflies. His studies led to the observation that some butterflies had patterns that were quite similar in appearance to unrelated species that were unpalatable to birds. He hypothesized that birds would learn to avoid them after only a few experiences and that this would then perpetuate the verisimilitude. When he returned to England in 1859 to recover from his epic jungle ordeal, he presented a paper on his discovery of butterfly mutations and to what he considered to be one of the best examples of  the “origin of all species and all adaptations.” [6] The phenomenon, known ever after as Batesian mimicry, became one of Darwin’s favorite examples of his epochal Origin of Species which had just been published. The two developed an enduring friendship, corresponding periodically on the new ideas of evolution. Bates became one of the primary adherents to the nascent theory, writing on one occasion that “I think I have got a glimpse into the laboratory where Nature manufactures her new species.” [7] The headwinds of religious dogma required decades to overcome, but gradually and fitfully the theory has gained near universal acceptance excepting those that adhere to biblical literalism.

With the advent of DNA as a roadmap of evolutionary change, Darwin’s insight only remains a theory insofar as it cannot be proven according to the scientific method of testing, which would require going back in time to reset the biological clock. The White Admiral conversion to Red-spotted Purple is one of the most documented of butterfly DNA subjects because of the infraspecific Mason-Dixon line that separates them. Proceeding north, the White Admiral prevails, while the far south is dominated by the mimetic Red-spotted Purple. The validity of the Batesian mimicry has been well established. A thirty year data set of Fourth of July Butterfly Counts confirmed that mimicry occurs even when the population of the unpalatable Pipevine Swallowtail species is low and that a sharp phenotypic geographical transition marks the boundary.[8] Between the two extremes, there is range over which hybridization occurs, affording a singular opportunity to study the interaction between the two variants according to DNA changes. Scientific research has established that the White Admiral variant is monophyletic (single ancestor) and that the hybridization of mimicry occurred just once. The hybrids that exist in the transition zone are thought to be due to mating between the two, producing on occasion a Red-spotted Purple with faint or partial white stripes. [9] More recently the location of the mutation responsible for Batesian mimicry on the genes of two different types of butterflies (Limenitis and Heliconius) that diverged 65 million years ago demonstrates the coevolution of this important survival trait. [10] Genetic confirmation provides the scientific “how” corresponding to the Batesian “why,” proof  for all practical purposes of Darwin’s “theory.”

The employment of Batesian mimicry of Limenitis arthemis in scientific research of butterfly sex  must surely have been considered for the Ig Nobel Prize in biology. One of the more compelling examples of female reproductive choice is sperm retention and storage after mating for fertilization at a later, more auspicious time. In that this would enhance the survival of subsequent generations, it has coevolved across the animal kingdom to include some insects, butterflies among them. It is also the case that many animals mate more than once; males with genetically driven propensity to sire as many offspring as possible and females to ensure successful insemination with the best possible mate characteristics. It is hard to say for sure, but it may also be that both enjoy it. Among the more profound questions facing biology is whether the sperm from a second mating male displaces that of the first or whether the two mix together to produce hybrids. Using the wing patterns that resulted as the biological metric, 17 females were mated with 34 males to conduct the experiment (it was not reported if this was consensual). The results were used to determine “insect mate-seeking strategies and individual fitness.” In that it was the first male’s sperm that prevailed, the conclusion was that it was not in the best interests of either the female or the male to mate multiple times. This then led to the conclusion that “virgin females apparently are sought by males and probably are more receptive to courtship and successful mating than are ones which have mated previously.” [11] This, at least, is the same theory espoused by some college fraternities and numerous religious denominations.

References

1. https://entnemdept.ufl.edu/creatures/bfly/red-spotted_purple.htm    

2. Marshall, S. Insects, Their Natural History and Diversity, Firefly Books, Buffalo, New York, 2006, pp 161-167.

3. Milne, L. and M. National Audubon Society Field Guide to Insects and Spiders, Alfred A. Knopf, New York, 1980, pp718-719.

4. Heikkilä, M. et al. “Cretaceous origin and repeated tertiary diversification of the redefined butterflies”. Proceedings. Biological Sciences. 22 March 2012 Volume 279  Number 1731 pp 1093–1099.

5. Brunetti, C. et al. “The generation and diversification of butterfly eyespot color patterns”. Current Biology. 16 October 2001 Volume 11 (20) pp 1578–1585

6. Bates, H. “Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae”. Transactions of the Linnean Society. 21 November 1861 Volume 23 Number 3. pp 495–566.

7. Carrol, S. Endless Forms Most Beautiful, W.W. Norton, New York, 2005,  pp 197-219.

8. Ries, L. and  Mullen, S.   “A Rare Model Limits the Distribution of Its More Common Mimic: A Twist on Frequency-Dependent Batesian Mimicry” Evolution. 4 July 2008, Volume 62 (7) pp 1798–1803.

9. Savage, W.; Mullen, S. “A single origin of Batesian mimicry among hybridizing populations of admiral butterflies (Limenitis arthemis) rejects an evolutionary reversion to the ancestral phenotype”. Proceedings of the Royal Society B: Biological Sciences. 15 April 2009 Volume 276  Number 1667  pp 2557–2565 at https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2686656/  

10. Gallant, J. et al “Ancient homology underlies adaptive mimetic diversity across butterflies” Nature Communications, 8 September 2014 Volume 5, p 4817.

11. Platt, A. and Allen, J. “Sperm Precedence and Competition in Doubly-Mated Limenitis arthemis-astyanax Butterflies (Rhopalocera: Nymphalidae)”. Annals of the Entomological Society of America. 1 September 2001 Volume 94 (5) pp 654–663.

Brown Thrasher

The Brown Thrasher is a hiker’s bird, searching for food in the wooded thickets we share

Common Name: Brown Thrasher –  There is some conjecture as to the origin of the word thrasher, which could derive from a noun or a verb. The similarity between thrash and thrush, another common bird which is sometimes becomes thrusher in English country dialect, implies a nominal origin. They are erroneously called brown thrushes on occasion. The predicate interpretation calls attention to the long tail that flails about as if thrashing. The reddish-brown upperparts and brown-black stripes across the front both evoke a consummate brownness.

Scientific Name: Toxostoma rufum – The genus is Latin for arched or bowed (toxon) combined with mouth (stoma). This  refers to the long, curved beak that is a common thrasher characteristic. Rufum is Latin for red from which rufous or reddish-brown is derived.    

Potpourri: The Brown Thrasher has a hiker’s perspective, eschewing the cityscapes of pigeons and sparrows and the bird feeders suspended over the manicured lawns of suburbia. They are found mostly strutting through  brushy, unkempt woodland thickets, pausing only to probe for insects with a long piercing bill.  Dressed for the part, the muted brown hues of protective top feathers match the surrounding tree bark. The vertical streaks that extend over the breast from head to toe are like the reedy grasses through which they pass almost invisible.  The cryptic colors are hardly arbitrary, as they share the same wooded areas with scarlet tanagers and  goldfinches with vibrant reds and yellows that are unmistakable even at a distance. Each species follows its own blueprint, hammered out by the evolutionary pressures of survival. All are  songbirds, and the brown thrasher is the most gifted of the lot. 

Thrashers, catbirds, and mockingbirds comprise the family Mimidae, the name originating from the Greek word mimos meaning  to imitate or represent, a word applied equally to mimes as mimics.  Most but not all of the birds in the family are noted for the complexity, variability, and length of their songs, which are not infrequently taken from other birds.  They are even called mimids as a group collective name. The catbird is the least loquacious, preferring a harsh, “downslurred mew,” one of the tortured onomatopoeias favored by birders to use human vocal phonetics to describe what emanates from bird beaks.  This of course sounds like a cat’s meow and what better name for a  bird with uniform dark gray plumage which, admittedly, is not unusual as a cat color. The Northern Mockingbird is incongruously more common in the south-eastern United States. It bears the sobriquet of copying and repeating as if taunting or mocking the sounds made by other birds, the ultimate mimic. [1] The adage that it is a sin to kill a mockingbird has no ethnic roots, but was rather chosen by novelist Harper Lee as a literary metaphor for innocence destroyed by evil.

Mockingbirds are mimics, but they are bested by Brown Thrashers when it comes to repertoire

The Brown Thrasher is adumbrated by its mockingbird cousin. This is due to a combination of familiarity, mistaken identity, and cultural lore. Mockingbirds congregate with relatively high densities near human habitation and are, as mimids, accomplished virtuosos in their own right.  Many songs of the hidden and unappreciated brown thrasher are therefore mistakenly attributed to the better known mockingbirds.  Attention to auditory detail provides a clue to the true troubadour. Mockingbirds normally repeat their purloined song phrases three times in rapid succession. This distinguishes them from both the terse catbird, with its singular tone and the brown thrasher, which repeats each sound just twice. [2] These double tone repetitions follow one set after another in lengthy and involved serenades  that are not only melodically captivating but independently conceived without mimicry. Henry David Thoreau found them boon companions during his philosophical sabbatical at Walden Pond, recording their song as “drop it, cover it up, cover it up – pull it up, pull it up, pull it up.” This was refined to doublets by Cornell’s University Laboratory of Ornithology as “plant a seed, plant a seed, bury it, bury it, cover it up, cover it up, let it grow, let it grow, pull it up, pull it up, eat it, eat it.” [3] None of which makes any sense, but why should it?

Measuring bird song tonality is complex, requiring not only frequency band measurements but also statistical calculations. While this may raise doubts about absolute numbers for recorded sounds that comprise a song, comparisons between species using the same methodology are surely valid. Mockingbirds have been recorded and analyzed with a repertoire ranging from 66 to 244 song types, the variation a matter of the individuality of each bird in its selection of songs to replicate and its predilection for singing in general. The songs of the brown thrasher number in the thousands and there is some evidence that improvisation occurs spontaneously in the course of a single burst. One brown thrasher was recorded singing in staccato bursts  for 113 minutes while stationary  at a single perch using  a mixed repertoire of 4,654 separate doublet units. Further evaluation with a spectrum analyzer revealed 45 song segments of which 20 were never repeated and two were repeated seven times. The entire sonata was statistically analyzed and found to consist of 1,805 separate sounds. It is a widely held among ornithologists that brown thrashers are the most accomplished vocalists among the thousands of species of song birds. But there remains a fundamental question: what is all of the singing about? It has been observed that male brown thrashers are mostly vocal in early spring as a part of territorial reckoning. However, once they are selected by a female and begin nesting, the singing stops abruptly like an avian version of musical chairs (nests in this case). While this would suggest that the intricate songfest achieves its intent ― i. e. intimidating rivals and attracting mates, why the complexity? Most birds suffice with a note or two even when there is more competition and closer quarters. There must be another reason. [4]

Why birds sing at all is a matter of conjecture. There are some sounds with obvious purpose and there are some that can only be for an amusement akin to human whistling. Non-songbirds produce mostly instinctual noises that are known  as calls, like the quack of a duck or the honk of a goose. Songbirds also use calls for functional purposes like location and warning but the song is mostly a whimsical trill. From the physiological perspective, songbirds are unique in having a “voice box” called a syrinx which has two sides that can be independently controlled to produce two different tones at the same time. There are many unexplained sonorous behaviors, including why males are the primary singers in temperate climates (as females are in the tropics) and why there is a dawn chorus. Songbirds learn songs from their parents as nestlings in what is called the critical period and practice them after fledging, shortening the years-long human process to weeks. Absent the intricate syntax necessary for human speech, there can be no practical reason for a string of arbitrary sounds extending to the thousands employed by songbirds like  brown thrashers. [5] The dawn’s early light sing-along may provide a clue ― that it has no practical function whatsoever, but is rather an expression of the exuberance in the ineffable beauty of musical tones ― the sine qua non of feeling alive even for creatures that soar over treetops and dart acrobatically from limb to limb. [6]

Brown thrashers pair bond to procreate, incubate, and subsequently feed the nestlings that result. Unlike many birds, however, they are not monogamous, sometimes changing mates even in the middle of a single summer, but only after the teenagers have left the nest for good. The survival of avian species as the only extant relative of the dinosaurs through the Cretaceous – Paleogene extinction 65 million years ago is testimony to the evolutionary resilience of strict behavioral protocols where caring for progeny is concerned.  Once the eggs are laid by the female, they must be kept warm until they hatch two weeks later as helpless altricial chicks that must be fed until fledged. The duties of this months-long endurance test would be impossible without the dedicated support of a mate. In the case of brown thrashers, this does not mean just finding food and guarding against predators, but sharing in chick care as well.  While the numbers vary to some extent, one field observation documented that during one 14 hour period the female sat on the nest for a little under 9 hours and the male just shy of 4 hours, about thirty percent of the time. Once the eggs hatch after about two weeks of brooding time share, the feeding frenzy begins. During one particularly long day, food deliveries began at 3:30 AM and did not end until 9:00 PM during which time the female made 186 sorties and the male 98 for a total of 286, equating to a rate of about one meal every 4 minutes. Since what goes in must come out, the nest would become fouled to overflowing at this rate if the droppings of the sequestered chicks were not removed. Fouling of the nest is a serious taboo among almost all animals. Both male and female adults inspect the nest scrupulously on a regular basis and particularly after feeding to collect the accumulated excrement which they encase in a transparent bag for removal. [7] It is probably not coincidental that birds and mammals are the only two major groups of animals that are warm blooded and that invest long hours of diligent care in overseeing the growth and instruction of their progeny. Both attributes require a lot of time and energy and yield an impressive adaptive result.

Maintaining a constant body temperature inside a thin skin covered in feathers against the onslaught of environmental extremes requires a self-regulating heat engine. Heat requires the oxidation of food, the essence of metabolism. Getting more oxygen to the cells of the body requires a plumbing system that is designed for inhalation and transport ― the four-chambered heart of birds (and mammals) evolved as a result from its three chambered reptilian predecessor. The extra chamber allows for a separate pulmonary loop on one side of the heart to operate at one eighth of normal blood pressure (15 mm HG as opposed to 110 mm HG systolic in humans) so that the hemoglobin of the red blood cells can absorb oxygen in the pressure-limited lungs.[8] To make this work, a steady input of nutrient laden food is needed. Brown thrashers, like all birds, are adept omnivores. In the warmer months,  they stalk resolutely through underbrush, thrashing aside the detritus to reveal the hidden arthropod smorgasbord below.  Since the etymology of thrasher is a matter more of lore than fact, there is no reason to reject the notion that the name arose from this thrashing of the underbrush, akin to threshing grain.  In the less constrained and sometimes exploitive science of the last century, 266 brown thrashers were eviscerated to provide incontrovertible evidence of what they ate. Animal foods comprised 62.22 percent,  consisting mostly of insects (18.14 percent beetles and 5.95 percent caterpillars), a major portion of which was used to feed the gaping maws of the brood. The balance of 37.38 percent was vegetable, mostly wild berries (19.95 percent), constituting the main food source during the  winter months (45 percent in January and February). [9] The berry-bird connection is not coincidental, but rather an evolutionary advance of flowering plants, or angiosperms, allowing them to spread their seeds by embedding them in a nutritive dollop frequently colored red with provocative purpose.  

Birds are intelligent and bird brains are advanced, contrary to the pejorative epithet. Corvids are considered the most intelligent, but mimids cannot be too far behind.  Acorns, the mast of oak trees, are a popular food for brown thrashers throughout the year, but especially in November when the nuts fall to the ground. Acorns are round with hard casings to protect the oak tree seed within from being disturbed. Squirrels pick them up in their hand-like paws and gnaw through the shell with incising teeth, but birds must make do with claws and a beak. Brown thrashers have been observed excavating a shallow hole in the ground as a tool to hold an acorn firmly in place while they hammer away with repeated piecing blows to breach the protective casing. Moving the acorn from one side to the other in the depression for a better angle of attack, the inner nutmeat was fully removed and eaten. Tool-making such as this requires cause and effect reasoning that defines intelligence. [10]  There is also evidence of the employment of props as coaxing tools in the training of fledglings to take the first flying leap. A parent bird was observed with a bit of paper folded to resemble a morsel of food held over the heads of the craning chicks and repeatedly pulling it away. Having inspired rapt attention, a swift move to a nearby branch incited lunging as  the only option. [11]   Taken together, the syntax in song, the resourcefulness in repast, and the complexity in care require a least a modicum of sagacity. The brown thrasher is a very smart bird … avian sapiens.

References:

1. Rosenberg, G. “Mockingbirds and Thrashers” National Geographic Complete Birds of North America, Jonathon Alderfer, editor, National Geographic, Washington DC, pp 495-502.

2. Robbins, C. Bruun, B., and Zim, H. Birds of North America, Western Publishing Company, Racine, Wisconsin, 1983, pp 240-241.

3. Johnson, T. “Out My Backdoor: Brown Thrashers, a Special Songster” available at https://georgiawildlife.com/out-my-backdoor-brown-thrashers-special-songster

4. Kroodsma, D. & Parker, L. (1977). “Vocal virtuosity in the Brown Thrasher”. The Auk Volume 94 Number 4, 1977, pp. 783–785.

5. https://academy.allaboutbirds.org/birdsong/       

6. Hartshorne, C. “The Monotony Threshold in Singing Birds”. The Auk. Volume 77 Number 2. 1956 pp 176–192.

7. Bent, Arthur Cleveland . Life histories of North American nuthatches, wrens, thrashers and their allies. Smithsonian Institution United States National Museum Bulletin 195. United States Government Printing Office Washington, DC, 1948. pp. 351–374. file:///C:/Users/Owner/Downloads/USNMB_1951948_unit.pdf     

8. Needham, W. The Compleat Ambler, Outskirts Press, p. 336.

9. Beal, F.  et al. “Common birds of southeastern United States in relation to agriculture.” U.S. Department of  Agriculture (USDA) Farmer’s Bulletin 755, 1916,  p. 11. available at https://www.biodiversitylibrary.org/page/56848073#page/13/mode/1up      

10. Hilton, B. Jr. (1992). “Tool-making and tool-using by a Brown Thrasher (Toxostoma rufum)” The Chat Volume 56.

11. Bent, op. cit.

Columbine

The Columbine flower has five unusually long tubular channels that lead to the nectar at the very top, an arrangement best suited for its primary pollinator, the ruby-throated hummingbird.

Common Name: Columbine, Rock bells, Meeting houses, Cluckies, Rock-lily, Honeysuckle, Jack-in-trousers – Columba is Latin for “dove” with the implication that dovelike is the intended metaphor for the flower that resembles five doves with uplifted tails and descending wings.

Scientific Name: Aquilegia canadensisAquila is Latin for eagle as the name of the genus. An avian appearance is again the likely etymology, the eagle of war supplanting the dove of peace. The species name attests to its first being identified and classified in Canada.

Potpourri: Like a five-pointed crimson crown for elven kings, the columbine evokes the magical spirits of rocky uplands. It appears as the summer sequel to the spring ephemeral flowers that emerge before the trees leaf out to attract early pollinators,  persisting well into the year to entice more persistent nectar collectors. A splash of red dangling from the end of a two foot long stout stem called a caudex cannot be overlooked even by the most myopic and oblivious of passers-by. It looks like different things to different people according to culture and custom. The original name columbine was assigned to the European variant (A. vulgaris) using the prevailing Romance languages to refer to a cluster of little columbae, doves in Latin. The American variant (A. canadensis) was afforded the rich diversity of colloquialisms that arose locally as pockets of immigrants settled in new communities and independently gave it their own apt mnemonic. Rock bells is perhaps the most obvious, as yellow stamens dangling underneath like a clapper must be  intended for ringing. Since they grow in rocky areas, rock lilies is also useful since lilies are bell-shaped. Meeting houses is a bit more imaginative, and may be a translation from a Native American name as a colorful five-poled tepee for tribal gatherings. Honeysuckle is a compound word for any flower with a long shaft that may be plucked to extract the honey-like nectar. While Jack-in-trousers does have lascivious implications, it is likely innocuous, like Jack-in-the-pulpit. Jack was a common English term for an unkempt young man and donning red trousers to stand out in the crowd with insouciance would make sense. [1] It is not possible to rule out, however, that something Jack concealed underneath might have been the original intent.

Like all things in nature, the peculiar shape of the columbine is not without reason. The only function of a flower is to attract a mobile pollinator to a sessile plant to carry  male anther pollen from one flower to the female pistil ovary of another. The genetic variation that this imbues is why sex evolved, fun has nothing to do with it. Cursory inspection of the wild columbine blossom reveals its most telling feature, the sweet honeypot of nectar at the very top of the “dove tails” is almost impossible to reach. It is apparent that the intent of the gradual natural selection that created this cul-de-sac was to favor a specific pollinator. That this occurs has been demonstrated irrevocably many times. The most famous example is none other than Erasmus Darwin, who had received a shipment of orchids from Madagascar which included one with an exceptionally long nectary; writing to a friend at the Kew Botanical Gardens, he noted that “in Madagascar there must be moths with proboscises capable of extension to a length of between ten and eleven inches.”  A moth with the necessary tongue length was discovered in 1907. [2] The exotic shapes of many orchids not only have guided passageways through which only the desired pollinator can fit, but even go so far as to imitate the female of an insect to incite the male to a lustful assault. Sex is everywhere.

Color is an equally important attribute for selective pollination. It is the evolutionary result not only of the flower as attractant, but also for the visual color perception of the targeted animal population. Color vision arose about 450 million years ago (mya) with the emergence of the agnathan or jawless fish vertebrates whose only modern survivors are the lampreys and hagfish. Color is a matter of wavelength measured in nanometers (nm) ranging from long wavelength red to short wavelength violet. The vestigial color scheme was tetrachromatic, having four cone types categorized as LWS, longwave sensitive, MWS, middlewave sensitive, SWS2 and SWS1, both shortwave sensitive in addition to rods for black and white. The first three cones correspond to the same general red-green-blue spectra that comprise human color perception with the addition of SWS1 that extends well into the ultraviolet range.  The “four color” physiology has been retained for the vast majority of animals, including most fish, reptiles, amphibians, birds and insects, which means, counterintuitively, that they “see” more than we do up to and including ultraviolet light. Almost all land mammals are dichromatic, having lost two of their four cones. The primates evolved from their two-color cone mammalian ancestors about 50 mya, adding a third (red) cone for reasons that are and always will be subject to conjecture. Bees, unlike the majority of tetrachromatic insects, have only three photoreceptors with spectral peaks at 340 nm, 440nm, and 540 nm which means that they can see ultraviolet light quite well but are deficient at the red end of the spectrum. [3]

The wild columbine is scarlet red and has a deep nectary to attract a specific pollinator – the ruby-throated hummingbird. It is certain that the flower and its ecological bird partner coevolved,  the columbine becoming redder and longer and the hummingbird seeking columbines more exclusively as they were much more likely to find nutrition that was, to all intents and purposes, saved for them.  The columbine species that is indigenous to Europe is blue in color, has a shorter stile, and is not as tubular in shape. It is pollinated largely by ultraviolet seeking bees who would miss it if it were red; there are no hummingbirds in Europe.  In the western half of North America, there area about twenty  species of columbine that range from white/yellow to blue with broad, open petals that extend horizontally as a landing pad for flies and bees that alight to crawl down to the nectar. The blue Rocky Mountain Columbine (A. caerulea) was selected as Colorado’s state flower in 1891 based on a vote by school children, its notoriety transfixed by the 1999 columbine massacre, an inaugural to an era of gun violence in schools that has yet to abate. [4] 

There are consequences to hiding nectar at the end of a long and delicate tube for the exclusive use of a chosen species. The struggle for survival is inexorable with hunger as its gnawing impetus. It is not uncommon to find a columbine with bore holes at the top of the dove tails where an enterprising insect has made its way to the ambrosia within. There are ways to stop marauders, and the chemical factory of plant physiology solves these types of problems by evolving countermeasures in the form of repellents that may also be toxic. [5] The compounds that emerged from defensive measures of flora against fauna are nature’s pharmacopeia, coopted by humans over time.  Herbal medicines were the province of medieval apothecaries, dispensing curative roots and fruits based on the collective wisdom of millennial trial and error. The European columbine was apparently not one of them, however, as John Gerard, one of the most notable of the early herbalists, does not prescribe them. Rather, he extolls their uniqueness as “five little hollow horns … of the shape of little birds” and notes that “they are set and  sown in gardens for the beauty and variable color of the flowers.” [6] The wild columbine of North America was another flower altogether.

 A. canadensis was widely used as an herbal remedy by Native Americans who were equally adept in the use of plants for medicinal purpose as their counterparts in Europe. The difference between the two columbines may stem from the need for chemical protection in the parlous American wilderness or perhaps from the welcoming of insect pollinators by A. vulgaris that its American cousin restricts in favoring the hummingbird. Whatever the reason, wild columbine has a rich history of practical applications that differ according to tribal custom, as there was limited cultural interchange. Among the more interesting formulations were a wash made from columbine leaves to treat poison ivy itch by Iroquois (a confederation of six tribes), an infusion for “heart troubles” by Cherokee, and as perfume for smoking tobacco by Meskwaki. Young bachelors of the Omaha tribe chewed columbine seeds into a paste to spread on their blankets and bodies as perfume for prospective mates. [7] It would follow that the suggestive red flowers would be used as an adornment to further the intent of the tryst although there is no record of this (there being no medicinal purpose). There is no evidence that any of these uses resulted in actual medical benefit other than what might have been conveyed by the placebo effect. A popular field guide to medicinal plants ascribes vague astringent, diuretic, and anodyne uses for the wild columbine with the provocative warning that it is “potentially poisonous.” [8] Since all medicine is governed by dose, this could be said for nearly anything if used to excess.

European naturalists followed in the footsteps of early explorers and colonists to study and eventually classify and catalogue the cornucopia of the New World. The variety of new plants and animals each carefully described in Latin overwhelmed established European biology, such as it was. To restore order out of the ensuing chaos, Carolinus Linnaeus, a Swedish botanist and physician, devised a taxonomic system based on genus and species published as Systema Naturae in 1735 that is still in use today. The red columbine from the Americas was one of the earliest plants to be identified and exported, probably due in part to its physical resemblance to the European flower of the same name and only accentuated by its flame red florescence.  The Canadian species name provides its provenance … it was first sent by French explorers to the noted Parisian botanist Jacques-Phillipe Cornut who wrote a treatise on Canadian plants in 1635  without ever travelling there to see them for himself. Cornut sent a specimen to John Tradescant in Lambeth, London who renamed it Virginia columbine in 1656. [9] The botanical dispute was a microcosm of the global struggle between France and England that was culminated in the French and Indian or Seven Years’ War one hundred years later. Tradescant and his son were the first English horticulturalists to collect extensively in the Jamestown colony as members of the Worshipful Company of Grand Gardeners. Captain Bligh of HMS Bounty fame was commander of many of their global collecting expeditions. During one of the three trips that the younger Tradescant made to Virginia between 1637 and 1654, he allegedly fell in love with a Powhatan girl and promised to marry her on his return. The Smith-Pocahontas affair was evidently no mere fluke. However, when he sailed back to England, he was introduced to the wife that his father had chosen.  Although reportedly devastated, he catalogued the contents  of what became known as the Lambeth Ark in 1657. It became the British Museum of Garden History in 1981. [10]

The columbine has not escaped the notice of poets, its beauty the inspiration for lofty verse and mellifluous metaphor. John Burroughs, the notable literary naturalist of the nineteenth century was especially fond of it. In his poem Columbine, he extolled its many virtues:

            I strolled along the beaten way, Where hoary cliffs uprear their heads, And all the firstlings of the May Were peeping from their leafy beds,

            When, dancing in its rocky frame, I saw th’ columbine’s flower of flame.   Above a            lichened niche it clung, Or did it leap from out a seam?–

            Some hidden fire had found a tongue And burst to light with vivid gleam.   It thrilled the   eye, it cheered the place, And gave the ledge a living grace.  

There is indeed something almost surreal about an encounter with the crimson columbine along a rocky trail where few things grow and the palette of green, gray, and brown prevails. It is nature’s way of reminding awkward bipeds stumbling along uneven trails that we are also its product ―  there is room for both beauty and the beasts.

References

1.  https://www.fs.fed.us/database/feis/plants/forb/aqucan/all.html   

2. Needham, W. The Compleat Ambler, Outskirts Press, pp 80-81.

3. https://hikersnotebook.blog/other-articles/geology-and-earth-science/colors-of-nature/

4. Sanders, J. Hedgemaids and Fairy Candles, Ragged Mountain Press, Camden, Maine, 1995. pp 23-25.

5. Adkins, L. Wildflowers of the Appalachian Trail, Menasha Ridge Press, Birmingham, Alabama, 1999, pp 138-139

6. Gerard, J. Gerard’s Herball – Or, Generall Historie of Plantes, John Norton, London, 1597 pp 69-70.

7. Ethnobotanical data base http://naeb.brit.org/uses/search/?string=columbine    

8. Duke, J. and Foster, S. Peterson Field Guide to Medicinal Plants and Herbs, Houghton Mifflin Company, Boston, 2000, p 153.

9. Drake, J. “Growing from Seed” The Seed Raising Journal from Thompson & Morgan, Winter 1987-88 Vol. 2 Number 1. https://www.thompson-morgan.com/aquilegias-article     

10. https://people.elmbridgehundred.org.uk/biographies/john-tradescant/   

11. https://poems.one/poem/john-burroughs-columbine    

Vernal Pools

Vernal Pool, ephemeral pool, vernal pond – Literally, an area of pooled water associated with the spring months that begin with the vernal equinox (equal night ~ 21 March) and end with the summer solstice (sun standing [still] ~ 21 June).

Vernal pools are seasonal wetlands that result from the accumulation of snowmelt and spring rains in low lying areas that provide a unique habitat for many plants and animals, notably amphibians. In general, the aquatic habitat is ephemeral or short-lived, as the heat and reduced precipitation of summer promote evaporation.  Eventually, only the cracked mud and silt remains, a condition  that can persist through the autumnal months. As the annual progression of seasons continues due to earth’s tilted-axis solar orbit, cooler and wetter conditions refill the pool. Many species have adapted to this cycle, living and procreating as the aqueous environment allows … estivating, desiccating, or migrating in the dryness of the summer months. Depending on local environmental factors and weather patterns, some pools may go through several cycles of filling and draining in a single year. When overall precipitation is above normal, they may persist through an entire year; areas with rock or clay substrates reduce drainage losses. Conversely, during extended droughts, vernal pools may fail to fill, drying out for the duration. [1]

The raison d’être for the coevolution of many species to a precarious reliance on an unreliable habitat like a vernal pool is the absence of fish predators. Life in the aquatic habitats of oceans, seas, lakes, and streams is a hierarchy of predation that is the epitome of survival of the fittest. At the top of the food pyramid are the streamlined, killing-machine sharks with rows of dagger teeth that are fearful even to humans, the most predatory of all animals. That fish eat other fish caught the eye of Benjamin Franklin as he travelled by ship, noting that when cod were hauled on board “I saw smaller fish taken out of their stomachs.” His on and off vegetarian regimen inspired by the works of Thomas Tryon thereafter included fish, since “ … if you eat one another, I don’t see why we mayn’t eat you.” [2]  The oceans abound with clever tactics to find food but avoid becoming food that range from the secretive lives of octopodes with eight suckered arms surrounding a beaked mouth that escape in a cloud of opaque black ink to the blue whale, a literal leviathan that consumes upwards of one ton of small crustaceans called krill every day. The cleverest of all of  the aquatic animals were those that moved from the littoral onto the shore to escape the fish carnage as amphibians during the Devonian Period about 400 million years ago.  The vernal pools are their palladium for the same reason.

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Wood frogs are the most visible, audible, and frequently encountered denizens of vernal pools.  The reason is that they are obligate … they require an aqueous habitat for egg deposition and tadpole maturation. The sexual positioning of the male and female frogs depicted above is called amplexus (Latin for embrace) and can persist for days.  Eggs are fertilized individually by male spermatozoa activated by contact with the water so they gradually float up and around the embracing couple. The protective and nutritive eggs are round sacs of yolk surrounding the frog embryo protected by a surface coat. They coalesce into softball size globules each with about a thousand eggs that float just beneath the pool surface like billowing clouds. The eggs, once adrift,  are entirely on their own. [3]  After several weeks, they hatch into tadpoles (Middle English for “toad head” as that is what they seemed to be). With a “beat the clock” mandate, tadpoles must mature to adulthood, develop lungs and venture ashore before seasonal heat turns the vernal pool into a slough of despond.

Obligate vernal pool fauna can be subdivided into four basic categories according to the timing and duration of habitat occupation. Those that arrive in the spring, mate and move on are non-wintering spring migrants, like wood frogs. The other three categories all involve over-wintering, surviving the dry period in the form of eggs or cysts that are drought resistant or as larvae or juvenile adults capable of burrowing to residual moisture. Some are year round as perpetual denizens and some are either spring or summer recruits, leaving the pool area during a portion of the annual cycle. [4] And then there are the facultative animals that use the vernal pools as a matter of choice but are not obligated to do so.  They are drawn to the vernal pools for the same reasons that animals are drawn to water generally. Some come to drink the essential elixir of life. Herbivores come to feed on the plants growing where there is water and carnivores come to prey on anything looking for food or water. This group would include the primarily aquatic green and bull frogs that will eat anything smaller than their considerable bulk, and the primarily terrestrial American toads that use their extended sticky tongues to zap the insects that swarm there. Adult eastern, red-spotted newts are the most notable of this category, swimming through the water to consume the eggs of other amphibians after wandering about the forest as juvenile red efts looking for a new home.  

Vernal pools, like any ecological niche habitat, operate according to nature’s law of supply and demand where everything has its place in the food chain. In this the invertebrate world, the amphibian vertebrates are the apex species that must be eluded whenever possible. Bipedal terrestrial humans are myopic megafauna, oblivious to the teeming masses of the miniscule that exist sight unseen. Adult and larval insects of all sizes and shapes, crustaceans  such as fairy shrimp and tiny copepods, and a variety of freshwater hydrozoa coexist by feeding on each other, and in many cases on amphibian eggs, prolific for that reason. How they manage to survive and replicate is exemplified by fairy shrimp, vernal pond obligates whose eggs are transported unwittingly by migrating waterfowl like mallards who eat gravid females in one pond and then fly to another. Their eggs are called cysts as they form a protective sac that can survive for decades even after being subjected to extremes of temperature ranging from near absolute zero to boiling. [5]  Dragonfly, damselfly, and caddisfly larvae occupy the middle ground as food for the adult frogs and toads and as predators of amphibian eggs and smaller invertebrates and crustaceans. [6] The real survivors are the hydrozoan hydras that can regenerate incised body parts and appear to be immortal, a matter of some interest to the study of human senescence and longevity. Since they reproduce solely by asexual budding, their stem cells must be continually renewed to sustain unmutated genetic function. [7]  The vernal pool may then also be the fountain of (eternal) youth.

Wetlands in general and vernal pools in particular are genetic repositories, a fact that has elevated their importance as habitats in need of protection. As is the case with many environmental concerns, wetlands are vulnerable to the tragedy of the commons … any resource  used by everyone and owned by everyone in common will become despoiled and no longer usable absent some usage rules. In the case of water, tragedy may not be a strong enough word as it is fundamental to almost all living things. The only recourse is government regulation to prevent their witting or unwitting annihilation, an intrusion seen by some as encroaching on the private property rights of landowners. The generic term wetlands came into use in the mid 1970s to definitize a panoply of colloquialisms such as bog, fen, swamp, and marsh. According to the Federal Geographic Data Committee standard FGDC–STD-004-2013, wetlands are “lands transitional between terrestrial and aquatic systems where the  water table is usually at or near the surface or the land is covered by shallow water.”  However, there is no absolute line of demarcation from either the physical or the ecological perspective between wet and dry areas as there is a continuous gradation from one to the other. This is particularly true of vernal pools that are on the fringe by their very nature; any mud puddle on a rutted dirt road could qualify … occasionally tadpoles can be seen squirming in them. The issue of what is or is not a wetland is generally moot until a change to the land is proposed. Not infrequently this involves developments like housing tracts or roads to access them. The generally accepted though far from ideal work around is called mitigation: wetlands can be destroyed as long as new wetlands are artificially created to compensate for the loss of habitat. [8]

The destruction of wetlands due to urban sprawl is one of the more persistent aspects of the human footprint that defines the Anthropocene.  The extent of the problem is at best a rough estimate due to wetland diversity, the lack of historical record, and inadequate survey data due in no small part to their ephemeral nature. Historical soil data in the Central Valley of California indicate that vernal pools occupied over 4 million acres a century ago. Less than one fourth of that amount remains ― one third of obligate crustacean species are thought to have become extinct. [9] The problem is most evident in the northeast and mid-Atlantic states where land is largely owned by  private parties of expanding populations. As a case in point, the town of Orono hosts the University of Maine and must account for housing needed for students, faculty, and support staff. A fifty seven home project on a forty acre tract on undeveloped land that had been approved by the town planning board ran afoul of the Army Corps of Engineers ruling that a vernal pool  could not be filled in.  The reduction in the allowable number of houses rendered the project uneconomical and it was cancelled. While this was a victory for the environment, it surely did not go over all that well with the Orono town council nor with the local chamber of commerce. [10]

The counterpoint to the tragedy of the commons is the erosion of democracy as citizens become frustrated with regulations they perceive as intransigent and arbitrary. Mitigation seeks to balance the commonweal of natural resources with the shared benefits to society of economic prosperity. But for the mitigation replacement of wetlands to count, it must be with an alternative that sustains the unique plants and animals that live there. An excavated depression filled with rocks and surrounded by berms that sometimes passes as ersatz wetland is not a habitat for wildlife but an opportunity for invasive species. The United States Department of Agriculture (USDA) Natural Resources Conservation Service (NRCS) established the Wetland Mitigation Banking Program (WMBP) to incentivize a better tradeoff. Using the concept of credits like that implemented to reduce acid rain, the system works by allowing the purchase of offsets in the form of wetland restoration, enhancement, or creation by any commercial or personal entity that seeks to fill one in somewhere else. Restoring or enhancing extant wetlands are the easiest to implement, mostly by establishing perpetual conservation easements. Creating a new wetland is a formidable task; engineering the ecology that nature does naturally requires planning and attention to detail. The plan must include the hydrology of water flow  management and  the establishment of soil type and consistency to ensure that fundamental qualities of a wetland habitat are retained. Flora and fauna considerations follow with the planting or  promotion of wetland vegetation to grow and attract amphibians, crustaceans, and hydras to the proposed oasis. [11] Joyce Kilmer poetically observed that only God can make a tree. This applies equally to aboriginal wetlands but mitigation is better than nothing.

References:

1. https://www.epa.gov/wetlands/vernal-pools    

2. Stuart, T. The Bloodless Revolution, A Cultural History of Vegetarianism form 1600 to Modern Times, W. W. Norton and Company, New York, 2006, pp 243-244.

3. Book, R. The Frog, Its Reproduction and Development, The Blakiston Company. Chapter 3 https://embryology.med.unsw.edu.au/embryology/index.php/Book_-_The_Frog_Its_Reproduction_and_Development_3    

4. https://mnfi.anr.msu.edu/abstracts/ecology/vernal_pool.pdf   

5. Brennan, D. “Vernal pools: Rains bring to life mini-ecosystem of button celery, Otay Mesa mint and fairy shrimp” The San Diego Union-Tribune, 1 April 2019.

6. Johnson, S. Vernal Pools, Documenting Life in Temporary Ponds, North American Nature Photography Association, 2021, pp 46-55.

7. Boehm, A. “FoxO is a critical regulator of stem cell maintenance in immortal Hydra” Proceedings of the National Academy of Sciences Volume 109 Number 48, 27 November 2012 pp 19697 – 19702. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3511741/  

8. https://www.fws.gov/wetlands/documents/Classification-of-Wetlands-and-Deepwater-Habitats-of-the-United-States-2013.pdf   

9. Brown K. “Vanishing Pools take Species with them” Science  31 Jul 1998: Vol. 281, Issue 5377, p 626.

10. Adams J. “Pooling Resources” Science  2 Oct 2015: Vol. 350, Issue 6256, pp. 26-28

11. https://www.nrcs.usda.gov/Internet/FSE_DOCUMENTS/nrcs143_025863.pdf

Cicadas

Common Name:  Cicada, Periodical cicada, 17-year Locust, Harvest fly – The common name cicada is from Latin meaning ‘tree cricket’ dating in usage from the 14th Century. In all probability it is in part onomatopoeia in reference to the sonorous noise produced by cicadas.

The cicadas oversized and starkly red compound eyes that protrude ominously from the sides of the foreboding black head in combination with the mysterious “W” embroidered near the trailing edge of the wing suggests Stygian origin.

Scientific Name: Magicicada septendicum The preternatural appearance of the cicada after an absence of seventeen years is so unusual as to seem to have an occult origination and therefore magical. The species name is Latin for seventeen to indicate the number of years in the cycle.

Potpourri: The emergence of up to 1.5 million cicadas per acre over an expansive area encompassing hundreds of square miles every 17 years is unique among the many curiosities of nature; the cacophony a crescendo to a long wait in darkened silence. The oversized and starkly red compound eyes that protrude ominously from the sides of a foreboding black head in combination with the mysterious ‘W’ embroidered near the trailing edge of the wing suggests Stygian origin. This is perhaps the etymology of the mistaken name locust, a type of grasshopper that swarms in scourging numbers bent on crop destruction so devastating as to evoke the biblical ‘plague of locusts.’ There are few events of nature which result in a perturbation so profound; the epitome of phenology, the study of natural phenomena that occur periodically.

Cicadas abound geographically due to their adaptable success in filling an ecological niche that protects them from predation with sufficient nutrition for growth and procreation.  Species numbers vary according to reference, but there are about 3,000 individual cicada species worldwide of which 1,000 occur in the Western Hemisphere; mostly in tropical environments. The periodical cicadas are among the 180 species endemic to the United States and southern Canada. Cicadas are in the order Hemiptera (half-wing in Latin) known as the ‘true bugs’ and are differentiated taxonomically into a suborder Homoptera (same-wing) due to their characteristic membrane-thin wings held aesthetically in lengthwise alignment with the plump body. The homopterans include aphids, leafhoppers and spittlebugs. True bugs suck. They are grouped according to the evolution of mouth parts in a beak-like rostrum ideally suited for imbibing the liquefied contents of other organisms; the rostrum contains two mandibles for making a hole and two maxillae that form a tube for alternatively injecting digestive saliva and ingesting nutritive fluids. Cicada nutrition is derived from the xylem sap that flows in the roots of a host tree. [1]

The egg-laying may result in the death of the branch; it is called “flagging.”

As is the case with all arthropods of the Class Insecta, cicadas have a life cycle that metamorphoses at least in part. A post-coital female deposits 400 to 600 eggs in 40 to 50 different nests consisting of drilled slots in small branches about one half inch in diameter.  The egg-laying may result in the death of the branch, known as “flagging.”  The eggs hatch after about two months into ant-size larvae that fall to the ground and begin digging to access the tree root food source; their fore legs are especially adapted for digging. Burrow depth varies according to reference and possibly by species from about one foot to eight feet. This has apparently been an historical conundrum; Moses Bartram writing in 1767 notes “I have not yet been able to discover the full depth to which these little animals descend. Some, I have heard, have been found thirty feet deep. I myself have seen ten.” [2] It is quite possible that this variability is correct, and that they keep digging until they literally hit roots.  For almost all cicadas, the subterranean sabbatical that lasts for a few years is punctuated by emergence from the deep and metamorphosing to adults. This occurs randomly in any given year toward midsummer; they are therefore sometimes called dog-day cicadas in reference to the period of the ecliptic cycle during which the Dog Star Sirius (the alpha star of the constellation Canis Major and the brightest in the night sky) is visible; Sirius is best seen in the starry nights of mid-winter January and not mid-summer July [3].  The periodical cicadas, meanwhile, are just settling in for the long haul.

There is considerable conjecture about the provenance of the 17 year cicada cycle even as there is no debate about its efficacy; the sheer number of progeny is proof of the latter. As the genus Magicicada is indigenous to eastern North America, there was no a priori knowledge of the phenomenon outside Native American experience until the European colonists geographically expanded and experienced its repetition. While their appearance in any given year could hardly be missed, the periodicity would not be noted for at least one and probably two cycles; what an average person would live to experience. Peter Kalm was sent by the Royal Swedish Academy to North America to study flora from 1748 to 1751; he recorded in the Swedish Transactions of 1756 “There is a kind of locusts which about every seventeenth year come hither in incredible numbers. They come out of the ground in the middle of May and make, for six weeks, such a noise in the trees and woods, that two persons that meet in such places, cannot understand each other.”  The genus Magicicada was officially instated by his noted Swedish countryman Carolinus Linnaeus in 1758. But that was just the beginning of a century of observation and recordation necessary for a full understanding of the recondite life cycles of the periodical cicadas.  The first noted anomaly was that some of the periodical cicadas have a life cycle of 13 years, an observation first documented in 1845 by D. Phares of Woodville, Mississippi in the Woodville Republican newspaper. It was not until 1907 with the publication of a USDA Bulletin by the government entomologist C. L. Marlatt entitled ‘The Periodical Cicada’ that the full range and scope of cicadas became manifest. Marlatt proposed thirty geographic groups called broods designated by Roman numerals I – XXX; seventeen broods of 17-year cicadas and thirteen of the 13-year variant.[4] The symmetry of the assignations was surely the result of apophenia, finding patterns in random data, as subsequent field evaluations established only 17 broods of which two have since become extinct; the original Roman numerals are still in use. As of now, there are twelve 17-year and three 13-year cicada broods, separated by both geographic location and by timewise sequencing.  Brood X is the largest of the 15 extant groupings; known as the Great Eastern Brood, it covers areas in fifteen states that include the mid-Atlantic region. [5]

The phenology of the periodical cicada is a matter of scientific interest due to complex and multi-species interaction with different environmental factors; it is also a matter of public interest due to the auditory variance from the quotidian when they emerge. An entomologist at the Maryland Department of Agriculture measured the din of cicada mating calls in the mid – 90 decibel range, about the same as a lawn mower, noting that “it got to the point where your skin started to crawl” [6]. There are three different species of Magicicada designated decim, cassini and decula that independently evolved a 17-year life cycle and subsequently to the 13-year life cycle, the decim having done so twice. While cicada fossils date from the late Permian Period about 300 million years ago (mya), phylogenetic studies indicate the divergence of the periodical cicadas about 4 mya with subsequent branching in the intervening years to the extant seven species of Magicicada: the 17-year M. septendicum, M. septendecula, M. cassini; and the 13-year M. tredicum, M. neotredicum, M. tredecula and M. tredecassini. DNA studies completed in 2012 provided the first real scientific understanding of the transitions that occurred over the last four million years. The overarching hypothesis is that predator satiation and the glacial cycles of the Pleistocene Epoch were the forcing factors for population survival and geographic dispersion.

What prompted the original lengthening of the underground period from several years to seventeen is a matter of conjecture, but survival from predation would clearly be a key factor. The term predator satiation refers to the simultaneous emergence of so many individuals that predators become satiated so that there is a concomitant high rate of cicada survival; the long, prime number periodicity ensures against a cyclical predator multiplying to meet the food supply. The glacial cycles of the Pleistocene Epoch resulted in the formation of periodic areas of suitable habitat during the warmer interglacial periods; it is these glacial basins that coincide with the current boundaries of the periodical cicada broods. In other words, cicadas evolved independently to the 17-year cycle at least eight times to form the regional broods we they currently occupy, a phenomenon called convergent evolution. The different species individually adapted to the cyclic pattern of the original species thereby becoming synchronized to the same cycle.  The subsequent divergence of the 13-year species occurred about 530,000 years ago when the southern M. tredicum diverged from the northern M. septendicum. The prevailing theory is that the warmer temperatures of the south during interglacial periods moved their emergence up by four years. It has been empirically established that a ground temperature of 64°F triggers cicada pupal ascent and this would occur consistently earlier to the south. [7]

Fully metamorphosed adult cicadas live for several weeks, the culmination of years of growth and survival punctuated by the primordial drive to create as many offspring as possible. The Strum und Drang of over three hundred insects per square meter advertising their virility with lawn mower-like 90 decibel mating calls can only be experienced; words do not suffice. The mating call is produced by two specialized organs called tymbals that each consist of a thin membrane with transverse struts – something like a snare drum. The contraction of the struts inward creates a clicking noise that repeats with every cycle with the mostly hollow abdomen as the echo chamber at a rate of about 7,000 cycles per second or hertz (by comparison, the whine of a mosquito is 600 hertz). Physically, this is accomplished by having as many as 1,500 scolophores, the organs that attach the membrane to the rigid exoskeleton of the body; by comparison, moths have at most four scolophores. [8] Each species has its own signature sound so that the corresponding female can make her choice, presumably based on tonal quality; the sounds of the three 17-year cicada species have been characterized as the word ‘pharaoh,’ a rotating lawn sprinkler, and a sizzling skillet. Close scrutiny has revealed that the female’s acceptance signal is a ‘finger snap’ sound that is made within a half-second of the completion of the male’s song. It is not as dour as it seems, as the subsequent courtship can be of some duration, and, in the compressed grow-mate-die cycle, a male may wait for a female to mature, chasing off other suitors with an interference noise to mask out the intruder. Given the stentorian sound, it stands to reason that the cicada has a means of blocking its own ears. The French nature writer Jean-Henri Fabre was so incensed with the noise of cicadas that he came to the conclusion that they must be deaf. To test the hypotheses, he borrowed two field guns used for local celebrations and fired them under a cicada infested tree next to his house in full song; the cicadas were completely unaffected. This rather obscure experiment found life in the poetry of  the Pulitzer Prize winning Richard Wilbur, who concluded his poem ‘Cicadas’ with:

This thin uncomprehended song it is

springs healing questions into binding air.

Fabre, by firing all the municipal cannon

under a piping tree, found out

cicadas cannot hear [9]

The emerging adult cicada

The periodical cicada is a long-lived insect that thrives in an underground Palladium where its many mammalian and avian predators cannot intercede. It emerges in finality as an adult only to take advantage of mobility and the full range of auditory and visual senses to find a mate and consummate a relationship. In the anthropocentric world of the epigeal, this short burst of activity constitutes its life, when in reality it is the other way around; from the ground up. However, understanding subterranean activities is impeded by access, and it was not until 1889 on the Washington D.C. grounds of the Department of Agriculture that Chief Entomologist C. L. Marlatt instituted a test that lasted until the adults emerged in 1906. Based on digging up various parts of the habitat over the 17 year test cycle, it was determined that the cicada larva molted four times until pupation and then an additional two times before the final adult metamorphosis. The total of six molts at intervals of two to three years was necessary to account for the growth of the larva and then the pupa; the most notable changes were the increasing complexity of the anterior legs, with enormous enlargement of the femora and tibia. In the opinion of Marlatt, these legs were specifically designed for “digging, tearing and transporting earth in the course of the insect’s subterranean life.” [10] It is the adaptation to a long life underground that is the sine qua non of the periodical cicada, and not its dystopian emergence after nearly two decades. The plump white cicadas that emerge from the last pupal shell are easy prey for birds until the predators are satiated; Melanism rapidly turns them black as camouflage against decimation.

The plump white cicadas that emerge from the last pupal shell are easy prey for birds until the predators are satiated.

Cicadas have figured prominently in folklore and fairy tale on a global scale. In ancient China, they were regarded as symbols of immortality in a manner similar to the deification of the scarab by the ancient Egyptians. Depictions of cicadas have been found on funerary vessels and carved bone spatulas dating from 1500 BCE. The symbolism of cicadas was persistent up to and including the Han Dynasty of 200 CE, their use having evolved to an amulet that was placed in the mouth as a part of the burial rite. Some studies have intimated the usage of similar jade cicada amulets by the Mayans and other Native American peoples suggesting some prehistorical cultural association that spanned the Pacific Ocean. The natural history of the cicada was metaphor for early Buddhist writings, the cast off carapace of the pupa in the metamorphosis to adult serving as a symbol of the hollow nature of human existence absent spiritualism. [11] Aesop’s classic Ant and the Grasshopper fable was originally written as the Ant and the Cicada. The storyline contrasts the diligent ant toiling to store food while the blithe cicada spends the entire summer singing. With the advent of winter, the starving cicada beseeches the ant and is rebuffed; the moral of hard work triumphing over idleness is a recurrent theme in Western Civilization. This was at one time much more prevalent in cultural associations than today. The late nineteenth century French artist Jules-Joseph Lefebvre named his painting that depicted a young, idle and artistically nude young woman among fallen leaves ‘La Cigale,’ the French word for cicada. That it was a metaphor for improvidence was implicit. [12]

References:  

  1. Marshall, S. Insects, Their Natural History and Diversity, Firefly Books, Ontario, Canada, 2006.  pp. 101-104.
  2. Bartram, Moses (1766). “Observations on the cicada, or locust of America, which appears periodically once in 16 or 17 years.” Communicated by the ingenious Peter Collinson, Esq. The Annual Register, or a View of the History, Politicks, and Literature, for the Year 1767. London: Printed for J. Dodsley (1768). pp. 103–106.
  3. Rey, H. The Stars, Houghton Mifflin Co. Boston, 1976.  The classic book to understand the constellations and where to find them in the night sky.
  4. Marlatt, C. L. The Periodical Cicada, USDA Bureau of Entomology Publication, 1907. Accessed https://archive.org/details/periodicalcicad00marlgoog  – An excellent resource for the history and scientific explanation of the periodical cicadas.
  5. http://www.cicadamania.com/ – is dedicated to cicadas “the most amazing insects in the world” and provides details of brood sizes, locations, and phenology.
  6. Maloney, B. “Cicadas’ Serenade a Source of Dread and Inspiration” Washington Post – 16 May 2004. The article was during the emergence of Brood X in the area.
  7. http://magicicada.org/magicicada/ “Magicicada species”. National Geographic Society. Retrieved June 2017.
  8. Encyclopedia Britannica, 15th ed Volume II pp 933-934.
  9. http://thedabbler.co.uk/2015/07/song-of-the-cicada/
  10. Marlatt Op. cit.
  11. Riegel, G. Cicada in Chinese Folklore, Entomological Society of Pennsylvania. Retrieved from https://www.insects.orkin.com/ced/issue-3/cicada-in-chinese-folklore/
  12. https://en.wikipedia.org/wiki/The_Ant_and_the_Grasshopper

Nodding Wild Onion

Common Name: Nodding Wild Onion, Lady’s leek – The umbel at the top of the leafless stem is bent at the top so that the flowers bend in a  downward nod, probably to enhance the presentation of the blossoms to pollinators.

Nodding Wild Onion in full flower with pollinators at right and lower left approaching

Scientific NameAllium cernuumAllium is the Latin word for garlic, which is one of the notable species of the  “onion” genus. The Latin form evolved from the more ancient Sanskrit word aluka meaning edible root, suggesting the importance of onion-type plants to the earliest hunter-gatherer peoples. Cernuus is probably the most descriptive species name of all time. It means “falling headlong, with the face toward the earth” in Latin.

Potpourri: The onion is one of nature’s most commendable creations. It is also one of its most prolific. There are 2,685 listed species names of which 969 are accepted as singular for the genus Allium ranging from A. aaseae to A. zergericum in a list of plants that is still growing. [1] The onion’s most notable but not universally shared attribute is the hypogeal bulb, which consists of thickened leaf bases that grow radially outward in layers from the stem plate at its base. The bulb shape is metaphor for the prominent domes of Russian architecture. Shallow, fibrous roots grow from the bottom of the plate as anchor to the smooth, linearly veined stalk that grows upward to produce an umbel or other florescence. Onion layers take on the meaning of getting to the heart of the matter by being peeled back. The ultimate onion encomium is attributed to Carl Sandberg,  “Life itself is like an onion; it has a bewildering number of layers. You peel them off one by one, and sometimes you cry.”

The redolence, ubiquity, and edibility of the many onion variants must surely have been noted and exploited by the earliest hominid gatherers supplementing the hunt on which they were dependent for sustainment. The shift from the collection of wild foods to the intentional planting and harvesting of selected cultivars took place over thousands of years in eight principal regions. The onion originated in central Asia and extended across Asia Minor into the Mediterranean region.  By about 3,000 BCE, vegetable cultivation in general and onions in particular was certainly practiced in the Nile River Valley. Egyptian tombs dating from the fist and second dynasties of 3200 to 2780 BCE depict laborers eating them. [2] In Numbers, one of the five books of the Pentateuch, the people of Israel complained to Moses that “We remember the fish we ate in Egypt for nothing, the cucumbers, the melons, the leeks, the onions, and the garlic; but now … there is nothing but this manna to look at.” [3] It is evident that, even then, onions and their more pungent leek and garlic relatives were both memorable, desirable, and even favored over what has been traditionally proclaimed “manna from heaven” which may actually have been a type of desert fungus.

The onions, leeks, and garlic of the Allium genus are similar in form to the grasses in having leaves with parallel veins extending along the longitudinal or blade axis. In the 16th century, it was recognized that these plants were characterized by a single embryonic leaf or cotyledon in their seeds and were called monocotyledons to distinguish them from the majority of flowering or angiosperm plants which are dicotyledon. Monocots and dicots are still in use as broad descriptive terms. Formal taxonomy started with Carolinus Linnaeus in the 18th century as the flood of new species from the Americas overwhelmed any attempt to sort them out according to their Latin descriptions. In that sex producing progeny of the same species was the essence of biology, the number and configuration of identifiable male and female organs offered a reasonable starting point. Linnaeas assigned 30 species to the genus Allium to a grouping named Hexandria monogynia for the six male anthers and the one female pistil. [4] The relationships among the many Allium species were confusing from the start and they were included in the diverse and overly large lily family until the end of the last century.  When the DNA vocabulary code  of nucleobase letters A,C,G, and T was deciphered, an understanding of the real genetic history connecting Egyptian tomb onion sculptures to cultivated fields of shallots and chives became possible.

The genus Allium is taxonomically situated (at least for now) in Asparagales, the largest order of the monocots, in the family Amaryllidaceae with the additional hierarchical categories  of  subfamily Alliodeae and tribe Allieae. However, there are many gaps in  knowledge that remain and a final accounting is years away. One recent scientific paper proposed “six subgenera, 50 sections and subsections for 600–700 species based on a multidisciplinary approach including morphological, anatomical, karyological, serological and numerical investigations as well as studies of life cycles, distribution, ecology and isozyme data.” What may be gleaned from this is that evolution could be too complex to go into unreasonable detail and that broad brush groupings may be better. It is certain that Allium is monophyletic or single ancestor and that there are two geographically distinct centers of diversity from which species radiation occurred, one in central Asia and the other in western North America. It is probable that the “first onion” originated at about the time of the Cretaceous-Tertiary extinction in Asia and that the Beringian Land Bridge was the conduit for intercontinental dispersion eastward. The adaptive Allium characteristics that were retained produced a series of herbaceous perennial plants with tunicate or layered bulbs,  narrow leaves emanating from the base, a stalk topped by umbellate flowers, and an aroma nonpareil. The complex chemistry of sulfur is responsible for the lachrymose vapors and distinct taste that can only be described as onion-like. [5] It is interesting to note that the word onion is derived from the Latin unio, meaning oneness…. there is nothing like an onion.

The bulb is composed of overlapping leaf bases anchored by roots

Representative species in the diaspora of the Allium genus as it spread eastward across North America from its western origins include wild (nodding) onion (A. cernuum), wild garlic (A. canadense) and wild leek or ramp (A. tricoccum). They are similar in habitat, preferring woods, thickets and meadows, and in phenology, emerging in spring and early summer growing in clusters convenient for harvest. [6]  The entire plant is edible, including the green stalk as long as it is still relatively young and tender. Most wild food guides prescribe chopping stems and bulbs as an ingredient to salad or boiling the bulbs as either an independent side dish or in combination with broth for soup or meat and vegetables as flavoring. [7]  The bulb that is the essence of the onion is also the repository of the chemical engineering plant that produces its beguiling taste and smell that has been described as “zesty, lusty, assertive, piquant, and distinctive.” It all starts with sulfur in combination with an allyl group, a specific combination of hydrocarbons  (methylene bridge and vinyl group) that is named for the Allium genus.  The resultant organosulfur compounds are diverse and offer “striking physiological activity as well as culinary appeal” that challenge analytical chemists. Depending on the species, 1 to 5 percent of the dry weight of the bulb is comprised of sulfur amino acid secondary metabolites. [8] That these compounds appeal to Homo sapiens is immaterial. They came about to protect onions from insects and other herbaceous animals at the lower end of the food chain.

The medieval medical practitioner Paracelsus is considered the Martin Luther of medicine for his notion that the healing power of nature could be brought to bear in the treatment of disease.[9] Notwithstanding the fact that native peoples have employed local herbs and tonics globally for millennia, his blinding flash of the obvious precipitated a sea change in what had been the ignorance of alchemy in the “civilized” world. One of his contributions was the notion that a poison could become a medicine at a reduced dosage. The sulfur compounds of onions are exemplary. Onion toxins are so successful at chemical warfare in garden trenches that they are planted in intercropping systems to protect vulnerable food crops, taking advantage of the excretions of the bulbs and roots and from the apparently disturbing smell of their leaves. [10] It would certainly not be beyond the perception of Native Americans, to cite just one well documented indigenous group, to note that any plant that kept bugs away would be useful either as a potential medicine, or at least to deal with what can only have been the maddening ordeal of biting flies and sucking ticks endemic to the woods and wetlands.  One widely known ethnobotany database [11] contains over three hundred separate uses for various onion species by widely dispersed tribal groups that range from asthma to scurvy and include being rubbed on the body to protect against insects.

The use of onions as both food and medicine was widespread in Europe by the seventeenth century as espoused by the numerous herbalists then popular. John Gerard wrote that the “juice of onions snuffed up into the nose, purgeth the head, and draweth forth flegmaticke humors,” in addition to providing a cure for baldness, a salve for burns, a treatment for ague, and even to prevent the scourge of rabies after the bite of a mad dog. His culinary admonitions probably reflect a personal distaste for onions that “causeth headache, hurteth the eyes, and maketh a man dim sighted, dulleth the senses, and provoketh overmuch sleep.” [11] Recent research has revealed that onion therapy is not as outlandish as it sounds. Sixteen species of Allium  have been shown to have anti cancer properties. The sulfur and organic compounds interfere with the “formation, growth, differentiation, and metastasis of cancer cells.” [12] The Mediterranean diet is one of the gold standards for a healthy life; that onions are a key component may not be coincidental.

Onion as weed

One of the consequences of any species that has mastered the art of procreation and the science of self defense is overabundance whenever ecological conditions permit. The suburban lawn and garden is the ideal habitat for a member of the Allium genus that has earned the weed sobriquet wherever it gains a foothold. Field Garlic (A. vineale) is a native of Eurasia but has become naturalized globally, especially in North America. It is characterized by numerous hypogeal bulblets, supporting a tuft of leafless flower stalks called scrapes that usually have no flower. It grows in clumps that are similar in appearance to tufts of grass that may escape detection except for a whiff of onion while mowing the lawn. It spreads both by seed, aerial bulblets, and offset bulblets that multiply during the winter like narcissus.  With the bulblets buried about three inches below the surface as both anchor and spreader, field garlic is almost impossible to eradicate without major excavation. [13] If it is of any consolation, the piles of green shoots and clusters of white bulblets are edible once you clean off the fist sized clumps of dirt that come with them.

References:

1. Missouri Botanical Garden Plant List at  http://www.theplantlist.org/tpl/search?q=Allium

2. Warid, W. A. “Vegetables and Vegetable Farming,” Encyclopedia Britannica Macropedia, William and Helen Benton Publishers, Chicago, 1972, Volume 19, pp 43-53.

3. The Holy Bible, Revised Standard Version, Thomas Nelson and Sons, Camden, New Jersey, 1952. Book 4, Chapter 11, Verse 5.

4. Linnaeus, C. Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Stockholm: Impensis Laurentii Salvii.  1753.

5. Li, Q. t al “Phylogeny and biogeography of Allium (Amaryllidaceae: Allieae) based on nuclear ribosomal internal transcribed spacer and chloroplast rps16 sequences, focusing on the inclusion of species endemic to China” Annals of Botany 21 October 2010 Volume 106 (5) pp 709–733.

6. Niering, W. National Audubon Society Field Guide to North American Wildflowers, Alfred A. Knopf, New York, 1998, pp 591-594.

7. Elias, T. and Dykeman, P. Edible Wild Plants, Sterling Publishing Company, New York, 1990, pp 58-61.

8. Block, E. “The Organosulfur Chemistry of the Genus Allium – Implications for the Organic Chemistry of Sulfur” Angewandte Chemie International Edition English 1992, 31, pp 1135-1178 http://ramsey1.chem.uic.edu/chem494/downloads-2/files/Block%201992.pdf    

9. Needham, W. The Compleat Ambler, Outskirts Press, 2020, pp 52-53.

10. Debra, K. and Misheck D. “Onion (Allium cepa) and garlic (Allium sativum) as pest control  intercrops in cabbage based intercrop systems in Zimbabwe” IOSR Journal of Agriculture and Veterinary Science March 2014 Volume 7, Issue 2 Ver. II, pp 13-1

11. http://naeb.brit.org/uses/search/?string=allium&page=6

12. Gerard, J.  Herball or General History of Plantes, London, 1633 pp 176-178.

13. Asemani, Y. et al “Allium vegetables for possible future of cancer treatment” Phytotherapy Research December 2019 Volume 33(12) pp 3019-3039.

14. https://plants.ces.ncsu.edu/plants/allium-vineale/

Eastern Fence Lizard

A wary Eastern Fence Lizard ponders flight or freeze as a defensive measure. The tail can be sacrificed if necessary to escape.

Common Name: Eastern Fence Lizard – The eastern geographic and fence habitat descriptions are used to identify the lizard according to where it may be found. There are many subspecies named according to different regions and forage preferences including both Northern and Southern Fence, Prairie, and Plateau in addition to the more specific White Sands Prairie, and Red-lipped Prairie. Lizard is derived from  lacerta, Latin for “upper arm” or lizard. It is not clear if there is any association between the two but it is suggested that it may derive from the notion of a serpent with legs i.e. arm-like  appendages.

Scientific Name: Sceloporus undulatus – The genus is from the Greek skelos meaning leg and poros meaning pore or passage. This refers to the anatomical arrangement of femoral (the femur is the upper leg bone) pores which extend along the inner side of the thighs that are believed to be used for pheromonal chemicals used for territorial marking. Undulate means “to form or move in waves” from the Latin undula, referring here to the wavy dorsal patterns of the reptilian scales.

Potpourri:  The Eastern Fence Lizard is the most prototypical of all reptiles, fleetingly seen darting across the rocks or up a tree trunk on four splayed legs covered with scales from the  head to the end of a long, narrow tail.  Lizards are relict reminders of the passage of geologic time from the dominant  “terrible lizard” dinosaurs of the Mesozoic Era to the modern Holocene Epoch that has become the Anthropocene of Homo sapiens everywhere. Even though the birds are the closest dinosaur relatives, it is the lizards and snakes of the Order Squamata named for there scale-covered bodies that evoke some sense of what the earth might have been like with tank-sized predators as the dominant life form. Reptiles evolved from amphibians in the late Carboniferous Era in part due to  the availability  of high quality protein from the burgeoning insect populations as terrestrial, lunged carnivores reproducing with amniotic eggs. What is left of this once apex class of animals is now divided into the clade Archcosauromorpha (first lizard body) comprised of crocodilians, turtles, and birds and the Lepidosauromorpha (scaly lizard body) which includes snakes, lizards and tuataras. [1]

While a scaly, slender tetrapod like S. undulatus is what comes to mind when a lizard is imagined, the Lacertilian suborder is globally quite diverse and expansive.  Unlike their snake cousins in the Squamate order all of which slither, most lizards come with legs though some are legless and some can run on just two. Some are massive and some are miniscule and many sport a variety of spines, crests, horns, and expanding plates that mostly evolved for defensive purposes. Lizards are also distinguished from snakes in having movable eyelids and external ear openings. Because they are cold-blooded, most of the roughly 3,000 lizard species live in  warmer temperate and tropical climates where the necessary external thermoregulation is   sufficient. For the most part, they still live on insects and run away or hide from nearly everything else. Eastern Fence Lizards were originally in the overly large Iguanid family with 14 genera and 44 species in North America and fifteen times that many when the tropics of Central and South America were included. As a part of the DNA biological revolution, Iguania is now an infraorder with 14 families of which Phrynosomatidae, meaning “toad body”  is the most recent, but probably not the last, taxonomic host family of the genus Sceloporus. [2]

The complicated relationships of the various lizards that live on trees (or fences), plateaus, and prairies across the United States and into northern Mexico provide an illustrative case study of the limits of phylogeny, the lines of descent or evolutionary development of all living things. Based on a mitochondrial DNA dataset of 56 populations of the “wide ranging and geographically variable” S. undulatus, it was concluded that the species was really a group that was monophyletic (single ancestor) comprised of at least four separate lineages each recognizable as an evolutionary species. It was further concluded that the variability among S. undulatus populations was “remarkable” with stark contrasts in “behavior, morphology and color pattern, sexual dimorphism, life history, demography, reproductive ecology, and chromosome structure.” But this is only the beginning of what promises to be an even more granular breakdown of species according to their DNA. It is not unreasonable to conclude that any classification apparatus intended to provide a better human understanding of the complexities of living things but instead offers only a death spiral of over-speciation is in sore need of rethinking. A general geographic  tripartite division of the ten-odd subspecies of Sceloporus undulatus has been proposed thatlumps them into  eastern woodland, central grasslands, and western canyon types to address the most prominent differences to impose some sense of order. [3] This is likely to be a more enduring outcome.

The life cycle of the Eastern Fence Lizard and its cohorts lends itself to the genetic diversity and dispersion that DNA changes chronicle. Two physiological factors stand out. One is that cold-blooded carnivore reptiles can subsist on a fraction of the food that a warm-blooded bird or mammal requires. This has its downside in geographic restrictions to warmer climates where freezing is not a risk factor and the need to locate and capture other animals that are just as adept at not being eaten as any other extant surviving species. The second factor  is the ability to excrete uric acid directly without the copious amounts of water needed by most other animals to expel nitrogenous wastes in the form of urine. This allows for the exploitation of desiccated terrestrial habitats like rocky outcrops, arid wastelands, and scruffy canyons that are anathema to mammals and amphibians. Taken together, a behavioral pattern that sustains population divergence and adaptation emerges with individuals spreading radially in search of food and seeking shelter in literal niches between the rocks. Specific adaptation is not uncommon among lizards. The anole populations of a number of small Caribbean islands consist of as many as ten different species each specializing in particular prey and habitats within the severely constrained ecological confines.

Sex is the engine of  speciation which would otherwise result only from random mutation. Any reptilian diaspora must therefore account for the essential meetup. The colocation of a male and a female of the same species is one of evolution’s most interesting and enduring outcomes. This is especially true in the case of widely dispersed carnivores hunting for food with only individual satiation in mind like lizards.  Territory and smell are the key factors. Male Sceloporus lizards establish boundaries to their domains using a series of aggressive displays that rarely involve physical aggression. Each species has a somewhat different variant that generally includes some combination of tail waving, head bobbing, and gaping. [4] There is a tradeoff between the establishment of territory and exposure to lurking predators like birds for whom such movements are an invitation. Those males that successfully survive while maintaining larger areas of control will have access to more females who are attracted by the pheromones that are distributed around the perimeter through the leg (skelos)  pores (porus) for which they are scientifically named. Evolutionary success favors those males that balance the hazards of exposure with the benefits of reproductive monopoly which would make them genetically fittest to survive. Field studies have found that male body size is the only parameter that correlates to home-range area and that the size and the shape of the area is determined by the location and number of females.  [5] Life then imitates nature as it is demonstrably true that jocks get the most dates.

Squamates (snakes and lizards) are endowed with forked male genitalia called hemipenes that are held internally, everted and erected when needed for copulation. The redundancy is neither a matter of improved fecundity nor a matter of multiple insertion but in all probability a matter of genetic happenstance. Amphibians have no need for a sexual insertion device since everything happens in the water. The hypothesis is that terrestrial reptiles required some new “hardware” and the parsimony of genetic creativity repurposed the leg forming genes to that end … two new legs became two proto-penises. By the same logic, the mammalian penis evolved from the tail forming genes and therefore retained its singularity. When a successful lizard male lures an interested female into his territory, one of the hemipenes is inserted into the cloaca with the ultimate result of the fertilization and deposition of about half a dozen eggs. A fertilized egg surrounded and protected by the amniotic sac is a second terrestrial adaptation necessary for nonaqueous survival. Amphibian eggs are suspended in water and don’t risk desiccation; on land it is a different story.  The penis and egg innovations coevolved so that the embryotic union would occur before the release of the fertilized egg with the protective, amniotic coating. [6] Live young came later on the way to us.

Brown hues enhance survival on tree trunks (lichens don’t help)

The realities of the natural food chain, as brutal as they frequently are, are no less a matter of evolutionary fitness. Cinematic excess from Jaws to Jurassic Park exploit the grim result. Just as lizards seek a meaty meal they may become one. Cryptic coloration is one of the main defenses employed to confuse the offense. It is also effective as an agent of stealth in duping nonobservant prey into a false sense of security ― nothing there does not always mean that nothing is there. Lizards are masters of the palette, notably chameleons  and anoles. Pigment cells called melanophores contain granules that are able to migrate based on a complex combination of hormones, temperature, and nerve signaling. The many lizards of the woodlands, grasslands, and canyons that are genetically related but distinct display a variety of bars, hues, and splotches that distinguish and protect them. Running and hiding under a rock, down a burrow, or into a tree knothole is a backup option if the disguise doesn’t work. The Gadarene leap would of course be headfirst leaving the tail as the last thing to disappear… which makes it a particularly vulnerable appendage. To a lunging fox or grasping bird the tail is a perfect target. Which is why most lizards have evolved with a tail that is not only detachable, but even wriggles suggestively as a decoy. Special muscles constrict at the connection point to minimize blood loss and a new tail grows back in about three months. [7] The would-be predator is at least left with something, albeit a small, bony snack.

The wavy scale pattern of S. undulatus is cryptic

Since lizards are masters of water management and prefer warmer temperatures, they are prolific in desert wastelands. One would think that the increasing temperatures of climate change would therefore be a boon and leaping lizards more than a dated Orphan Annie epithet. Maybe yes and maybe no. Nest temperature is correlated to the sex ratio of the hatchlings. Male lizards are more cold tolerant than females so cooler temperatures produce more males according to evolutionary pressures. Conversely,  a warmer climate would gradually shift the ratio toward more females, which would mean more eggs and more lizards. A recent study of Australian skinks found that the average nest temperature had increased by 1.5 degrees Centigrade between 1997 and 2006. The study also showed that lizards incubated in warmer nests were smarter based on laboratory testing of escape route detection. [8] So there should eventually be a surplus of  smarter females and fewer dumb males and the world will presumably be a better place according to a certain mindset. However, this does not seem to be the case, as lizard populations are declining. Since 1975, Mexico has experienced a 12 percent decline and reductions in lizard sightings have been reported elsewhere. Experiments have revealed that the temperature in the desert areas where lizards are no longer found is significantly hotter than areas where they remain. The hypothesis is that foraging time becomes restricted by excessive temperatures to the extent that survival is threatened. Extrapolations to 2080 result in a 20 percent lizard population decline.[9] By then, perhaps another meteor will crash into the earth near Mexico like the last one and all biological clocks will be reset, perhaps making the reptiles top dog again.

References: 

1.  Starr, C. and Taggart, R. Biology, The Unity and Diversity of Life 5th Edition, Wadsworth Publishing Company, Belmont, California, 1989, pp 683-687.  

2. Behler, J. and King, F. National Audubon Society Field Guide to North American Reptiles and Amphibians, Alfred A. Knopf, New York, 1979, pp 497-538.

3. Leaché A., Reeder T. “Molecular systematics of the eastern fence lizard (Sceloporus undulatus): A comparison of parsimony, likelihood, and Bayesian approaches”. Systematic Biology. 2002, Volume 51 (1) pp 44–68.  

4. Gorman, G. “Sauria” Encyclopedia Britannica Macropedia, William Benton, Publisher, Chicago, 1974, Volume 16 pp. 282-288.

5. Haenel, G. et al  “Home-Range analysis in Sceloporus undulatus (eastern fence lizard). I. Spacing patterns and the context of territorial behavior”. Copeia. 2003 pp 99–112.

6. Laslo, M. “When two become one: the evolution and development of external genitalia on land” 8 April 2015,   https://evobites.com/2015/04/08/laslo_external_genitalia/    

7. Missouri Department of Conservation https://web.archive.org/web/20120702004844/http://mdc.mo.gov/discover-nature/outdoor-recreation/nature-viewing/amphibians-and-reptiles/lizards

8. Youngsteadt, E.  “Could Climate Change Alter Lizard Learning?” Science 10 January 2012.

9. Price, M. “Climate Change Causing Lizards to ‘Wink out of existence’.” Science, 13 May 2010

Royal Fern Family (Osmundaceae)

The Royal fern (Osmunda regalis) is bipinnate, or twice divided as each pinna that extends from the central stem or rachis has individual pinnules. Royal ferns can grow to the height of a small tree.

Common Name: Royal Fern Family – Named for its most prominent member. The Royal Fern is relatively large, up to ten feet tall, evoking the grandeur that royalty implies. In addition to the royal fern, there are two other prominent species in the family: Cinnamon Fern, named for the light brown color of its fertile frond; and Interrupted Fern, named for the fertile fronds that “interrupt” the branches or pinna along the stem or rachis.

Scientific Name: Osmundaceae – The etymology of osmund (aceae is the  standard suffix to indicate family in taxonomy, ‘belonging to’ is a synonym) is not established. Several references cite  Osmunder as the Saxon name for the Norse god Thor, which would suggest a similar rationale as god-like or royal. However, the Saxon name for Thor is Thunor or Thonar [1] which is the origin of Saxon-English Thursday (vice Osmunday).  Other suggestions include os + mund as god + protector in Saxon or os + mundus as bone + clean in Latin. A more imaginative account comes from Harpers Magazine for Young People published on 9 December 1879 that relates the story of Osmund of Loch Tyne who hid his wife and daughter from marauding Danes in medieval Great Britain on an island covered with the large royal ferns.  His daughter who was thus saved commemorated the event by naming the ferns after her father. Referring to this story as a “fanciful tale,” the writer proffers a second option  as  “derived from an old Saxon word signifying strength, the specific name indicating its royal or stately habit of growth.”

Potpourri: Fern-like plants were among the original land biota, appearing with liverworts and mosses in the Devonian Period some 390 million years ago. They were the first vascular plants in having vessels or ducts to carry water and minerals upward and sustaining sugars downward to extend and expand photosynthetic tissues toward solar energy, unlike their ground hugging cohorts. They reached prominence in number and size during the succeeding Carboniferous Period, sometimes called the “Age of Ferns,” forming the coal beds that powered the Industrial Revolution of the 19th Century that created the carbon dioxide problems of the present (ferns are not at fault). While the primeval ferns appear only in the fossil record, their ancestors are evident to this day wherever the combination of wetness and sunlight meet the necessary and sufficient ecological requirements for their growth. [2] The osmundaceous ferns are among the oldest of those that still prevail, their longevity a result of robust physiology and adaptability, even to meteoric cataclysms like the Cretaceous – Paleogene extinction that extirpated all dinosaurs except birds. The species named royal fern, Osmunda regalis, is the only vascular plant to have  been found on all seven continents.

The royal fern family is the sole member of the order Osmundales, which is the oldest and smallest of the seven orders in the subclass Polypodiidae, which includes most of the 10,560 species of extant ferns. They are also called leptosporangiates on account of their characteristic  one cell thick spore cases, which differ from other ferns and seed plants which have multiple cell layers (lepto means ‘peeled’ in Greek and is used to connote weak or thin i.e. easy to be peeled). For comparison, as of 2016 there were 374,000 species of plants comprised of 295,383 flowering angiosperms, 44,000 algae, and 12,700 mosses, the ferns and several other groups like conifer gymnosperms ― about 500 new plant species are being added annually. [3] Osmundales is the smallest fern order since there are only 20 species in 6 genera, which would equate to one fifth of one percent of all ferns. Paradoxically, they are at the same time the most common fern fossil, with over 100 species in 20 genera that date to the early Mesozoic about 220 million years ago, which makes them the oldest. [4]  The paradox of becoming small in number (or stature) and old in age over time is something that every living thing must grapple with, both in the lifetime of one individual and in the span of a species.

The anatomy of a fossilized fern specimen recently unearthed provides some rationale to the conundrum of why there are so many fossil species with only a few survivors. A two inch long segment of fern stem surrounded by frond bases and rootlets which was removed from 220 million year old mafic volcanoclastic rocks in Sweden. Due to its apparent submergence in hydrothermal brines and consequent rapid permineralization by calcite, the preservation was complete to the subcellular level of the nucleus and even its entrained nucleoli. The detail that this uniquely preserved fossil provided allowed for thorough characterization. The physical measurements of stem size, nuclear parameters, DNA and chromosome count were essentially identical to those of a living royal fern family member. The incontrovertible conclusion was that the genome had been static for over 180 million years. [5] Why? One reason is that the royal ferns, both ancestral and recent, have a tough, tree-like constitution.

Ferns grew to great heights in the Carboniferous and tall tree ferns persist to the present,  primarily in tropical and subtropical regions. Deciduous and coniferous trees can grow only with the support of the cellulose and lignin of their woody trunks. Arborescent ferns  evolved two non-wood mechanisms to support their weight. One method employs a hardened tissue called sclerenchyma that extends along the length of the stem and provides support against buckling. The reinforced fern “trunks” are so hard that they cannot be cut down even with a chainsaw. A second method to support a lofty frond expanse is to surround the stem with a mantle of dense roots that provide rigidity by interlocking like rebar used in concrete. With up to five times the girth of the bare stem, the matrix is an unbending sheath. Osmundaceae is an ancient lineage, one that originally consisted  of several genera of tree ferns. The root support structure that evolved in the Mesozoic to provide for greater height (and   limit browsing by herbivores) was retained in the  family members that survived as royal, cinnamon, and interrupted ferns.  A cross section inspection of one of these three fern species requires a heavy saw, revealing a small central stem surrounded by black leaf bases, a structure that is similar to its ancient relatives. The retention of this feature may well have been the key to survival beyond the extinction at the end of the Cretaceous,  and why they never changed. [6]

Interrupted fern fertile frond grows on the rachis, interrupting the pinna.

There are several other features that distinguish the members of the royal fern family. Most notable are the distinctive sporangia, the structures in which spores are produced. Most ferns have spores in small round structures called sori that are visible as brownish dots usually on the underside of the fronds. The royal ferns have separate structures called fertile fronds  with modified pinnae to which the sporangia are attached singly in clusters.  When mature, each sporangium opens with a long slit to allow the spores to escape, turning brown when empty before eventually disintegrating. The spores are green in color as opposed to the more common brown to black and are also relatively short-lived. Like all other ferns, the spores fall to the ground where they germinate if conditions are suitable into either male or female gametes whose sub rosa sexual union produces a new sporophyte frond. The fertile fronds take on a variety of shapes that distinguish the three main species:  Royal fertile fronds are at the end of the pinnae like crowns; Interrupted fertile fronds are intermittent along the rachis, interrupting it; and Cinnamon fertile fronds are on a separate stalk, turning cinnamon colored when  mature.[7]

Cinnamon fern fertile fronds grow from a separate stem, turning cinnamon brown when they release the spores

The use of plants as herbal remedies for human ailments is not without reason, although modern science in medication is usually a better course. Phytochemicals are produced by plants through evolutionary mutation to ward off predatory animals from microbes to moose. Plant toxins that repel soil microbes can and have been used to prevent human microbes from gaining purchase, with the proviso that dose matters and poisons can kill. Those plants that have been around for millennia like the Osmundaceae are a good place to look for natural cures. One rarely sees a royal, interrupted, or cinnamon fern with the holes of leaf-cutter insects or the dark spots of fungal invaders. Native Americans put their sapience to good use in identifying curative concoctions mostly through trial, error, and observation. The six nations of the Iroquois Confederacy of the Northeast (who called themselves Haudenosaunee for the long houses they built) used all three royal family ferns for a wide variety of conditions including various “women’s problems” and joint pains. The Cherokee of the middle Atlantic region used  the cinnamon fern for snake bite and an unspecified spring tonic while the southern Seminoles used royal ferns for so-called  chronic conditions one of which was insanity. [8]

The peoples of Eurasia were no less adept at herbalism which started similarly as tribal remedies and succeeded to an organized doctrine with the advent of the written record. By the sixteenth century, a consensus emerged in the form of a number of publications listing various plants and their purported curative properties. Among them was The Herball of John Gerard who listed “Water-Fern or Osmund the water-man” as “a great triangle stalke two cubits high” which is surely Osmunda regalis, the royal fern. The root is described as  “great and thicke” with a hard woody part named “the heart of Osmond the water-man.” One must conclude from this account that the Osmund folk tale concerning the origin of the genus Osmunda has some basis in fact, however romanticized. The “Osmund” root, and especially its knotty core, was prescribed to be pounded and mixed with liquor and used for  “those that are wounded, dry-beaten and bruised.” These “wound-drinkes” would “dissolve cluttered blood in any inward part of the body.” The tender sprigs of spring were equally beneficent, especially when made into healing plasters for the “aforesaid wounds, punctures, and suchlike.” [9]  Hyperbole aside, palliative properties of royal fern have been demonstrated in the inhibition of head and neck squamous cell carcinoma, one of the most common forms of cancer. [10]

Linnaean taxonomy of the 18th century based on extrinsic characteristics has been obsolesced  by phylogeny based on DNA in the 21st century. This revolution in biology has had limited effects on animal classification … but plants and especially fungi have been mercilessly rearranged. The ferns were originally assigned to the class Cryptogamia (Greek for “hidden marriage”)  with 16 genera and 174 species which included fungi, algae and bryophytes. Since the arrangement of plant sexual organs (stamens and pistils) was the original basis for plant classification, the flowerless cryptogams were cryptic. A default fern  classification system was based on the location and shape of the spore-bearing sori on the fronds (which are absent in Osmundaceae), the first of  numerous reassessments over the next two centuries. The fact that fern spores germinate to form sexual gametes that mate in wet soil was not recognized until a British surgeon in Jamaica named Lindsay studied the development of “fern dust” he had planted in a flowerpot in 1794. The guessing game came to an end at an international symposium of pteridologists (fern botanists) in 1972 where it was settled that it  would be impossible to establish a proper taxonomy without monophyletic studies, which are really only possible by DNA comparison. [11]

Cinnamon ferns (Osmundastrum cinnamomeum) have pinnate-pinnatifid fronds

The Pteridophyte Phylogeny Group (PPG) was established to bring DNA order out of the Linnaean farrago. Even though this seminal 2016 classification lists 11,916 species, 337 genera, 51 families, 14 orders, and 2 classes of ferns and fern allies (the lycophytes, mostly club mosses), it carries the caveat of “not intended as the final word … but rather a summary statement of current hypotheses.” This caution relates to complexity in evolutionary change in general and to the peculiarities of ferns in particular. The PPG restricted monophyletic lineage (single ancestor) only at the genus level and above because of fern polyploid speciation, which is a mutation where the number of chromosomes is increased. Humans have two sets of 23 chromosomes (46 total)  which is referred to as 2n; polyploidy would make this 3n or more. It is estimated that more than a third of all fern species are correlated to a change in ploidy. Even within the strictures of DNA analysis, the royal fern family Osmundaceae retained its coherence and relevance as the most ancient of the extant ferns. However, the royal, cinnamon and interrupted ferns, which used to all be in the genus Osmunda (and are still listed there in most field guides and in Encyclopedia Britannica) are now, tentatively in three separate genera. [12]

Interrupted ferns (Claytosmunda claytoniana) are nearly identical to cinnamon ferns except for the interrupting fertile frond.

The royal fern as Osmundaceae  type species retained its position in the genus as Osmunda regalis with several  variations like spectabilis. It is bipinnate, which means that the pinna that grow from the stem are fully divided or cut into separated pinnules; this is sometimes called two-cut or twice-divided. The cinnamon fern became Osmundastrum cinnamomeum as the only species in a new genus. Its fronds are  pinnate-pinnatifid, which means that the pinnules are only partial and not separated. This is also the case with the interrupted fern which is almost indistinguishable from the cinnamon fern without the very distinct difference in their fertile fronds, one a cinnamon brown stalk and the other interspersed along the main stem. The interrupted fern, O. claytoniana, became Claytosmunda claytoniana paying double homage to the noted colonial botanist John Clayton of Virginia. This is probably not the end of the names of the members of the royal fern family, but it is the end of this particular discussion of them.

References:

1. Encyclopedia Britannica Micropǣdia “Thor” Volume IX, William Benton publisher, University of Chicago, 1974  p.967.

2.  Wilson, C. and Loomis, W. Botany, Fourth Edition, Holt, Rinehart. and Winston, New York, 1967, pp 499-532.

3. Christenhusz, M. and Byng, J. “The number of known plants species in the world and its annual increase”. Phytotaxa. 2016  Volume 261 (3) pp 201–217.

4. Bomfleur, B. “The fossil Osmundales (Royal Ferns)—a phylogenetic network analysis, revised taxonomy, and evolutionary classification of anatomically preserved trunks and rhizomes”. PeerJ. 11 July 2017 Volume 5: e3443

5. Bomfleur, B. et al, “Fossilized Nuclei and Chromosomes Reveal 180 Million Years of Genomic Stasis of Royal Ferns,” Science, Volume 343 21 March 2014, pp 1376-1377.

6. Moran, R. A Natural History of Ferns, Timber Press, Portland, Oregon, 2004, pp 140-146.

7. Cobb, B. Farnsworth, E., and Lowe, C. Ferns of Northeastern and Central North America, Houghton Mifflin Company, Boston, 2005. pp.170-177.

8. The ethnobotany database at http://naeb.brit.org/uses/search/?string=osmunda

9. Gerard, John, The Herball, or Generall Historie of Plantes, John Norton, Publisher, London, 1597.

10. Schmidt M, et al “The influence of Osmunda regalis root extract on head and neck cancer cell proliferation, invasion and gene expression. BMC Complementary and Alternative Medicine. 4 December 2017, Volume 17(518).

11. Christenhusz, M.; Chase, M. (2014). “Trends and concepts in fern classification”. Annals of Botany 13 February 2014, Volume 113 (4): pp 571–594.

12. Pteridophyte Phylogeny Group. “A community-derived classification for extant lycophytes and ferns”. Journal of Systematics and Evolution. November 2016 Volume 54 (6) pp 563–603.