My much-loved,  old, heavy botanical manuals (e.g. Fernald and Britton and Brown)  always include a sentence or two about the habitat where a plant is found, as well as exceedingly detailed morphological descriptions.  “Found in rich soil” is a frequent description that can apply to fallow farmland, alluvial  floodplains, a bouldery forest at the base of a hillside,  or  a rocky summit with two inches of mineral-rich soil , covered with red columbines.  I used to think rich soil was rich soil, no matter where it was, with other ecological factors making the plant communities so different from each other. But I’ve learned that is only partly true.

Fallow Farm

A fallow, fertile crop field supports a rank stand of annual weeds like pigweed, ragweed, and giant foxtail, that have the genetic capacity to grow tall in response to high levels of  the three basic nutrients (especially nitrogen, but also phosphorus, and potassium).  The field produces abundant birdfood, but  weed competition excludes all the wildflowers and ferns than are genetically programmed to stay short for their whole lives. Frequent plowing  also excludes native perennials.  Soils may or may not be “rich” in ninerals like magnesium and calcium.  The probability of finding rare species is very low.

Floodplain

The dominant understory vegetation in a “rich” floodplain of a large river is also thick and lush, mostly annuals like jewelweed and false nettle, though flooding and ice, not plowing excludes perennials.  Frequent deposits of fresh silt and organic matter provide an abundant supply of the three, basic common  nutrients. Especially if the watershed has traprock ridges, alluvial soil is also rich in other minerals like calcium, magnesium, and manganese, and subacidic.

Such  soil is well-suited to late-season farming.   It also can support uncommon, minerotrophic (mineral-loving)  plants, where growing space and light is available. Floodplain annuals start growing only after floodwaters  recede, and tree falls and thick deposits of sediment often  open up new bare soil patches.

I have found delicate wildflowers only in early spring, before they are shaded by the rank annuals.  Dutchman’s Breeches (Dicentra cucullaria)  and spring beauty (Claytonia virginica) do grow on the banks of the Farmington River in Simsbury, with much traprock in its watershed. I also know a few rare floodplain sedges, like Davis sedge (Carex davisii),  with a vigorous,  tall growth form,  that can compete with the dense floodplain annuals – though not invasive shrubs like Euonymus alata.  Do  these uncommon floodplain plants need soil with high concentrations of minerals, with or without high availability of nitrogen, phosphorus and potassium?  Has any  research on this been published?

Base of a hillside

Another  place to find  “rich site” wildflowers, ferns  and sedges  in Connecticut is  the base of  a  hillside, among the boulders. Soil water at the base of a hill has been seeping slowly  downhill for hundreds of feet,and for many centuries,  dissolving minerals from the surfaces of soil particles and rocks. Topsoil has  also slowly washed downhill over the centuries. Slope-base soil  typically  has ample minerals and enough of the three basic nutrients, and is moist as well.  Stately bottomland trees grow in this rich, rocky soil:  sugar maple, red oak,  tulip poplar,  ironwood, and occasional basswood.   Spring ephemerals like red trillium (Trillium erectum) , bloodroot,  and trout lily  (Erythronium americanum) do most of their growing  before the trees leaf out.  However, some shade-tolerant minerotrophic plants  can keep growing  through the summer, like red elderberry and  broad beech fern – and other much rarer ones, like Goldie’s fern (which I have yet to find.)   The  understory is  less dense than in the floodplain, with less competition, and greater diversity. It still rankles me that a Target big box store was built  in this habitat at the base of a Meriden traprock cliff, without any ecological survey beforehand. It was over ten years ago, but I still boycott the store!

At the base of very long  seepage hillsides, soil water has the highest mineral concentrations,  and the slope-base plant community is potentially most diverse. The reason is simple, as I was taught by my major professor Ton Damman: the further the groundwater travels, the more minerals are dissolved.  I  recall an amazingly diverse  swamp at the base of a great hill in Winsted, Connecticut, west of Route 8.  We measured the nitrogen  levels, and they were quite low. Vegetation was low in density and stature, however, not a rank, impenetrable  thicket.  This allowed diverse, minerotrophic plants to coexist, including  melicgrass (Glyceria melicaria), chestnut sedge (Carex brunnensis), and a dwarf raspberry called Rubus pubescens.

Rocky Outcroppings

This year I  found these same plant species – and also Dutchman’s breeches  – on several  shale outcrops  in the  East Berlin geologic formation.  The laminated  shale rock structure increases surface area available for mineral dissolution. Positively charged cations (e.g. calcium and magnesium)  enter the soil water and increase the pH.  Subacidic soils derived from traprock, limestone, or shale  have the highest mineral levels, and  support diverse and interesting  plant communities.  Minerotrophic   plants are most likely in areas with subacidic soils, in large part because higher pH makes minerals more available to plants.  (This is the reason that farmers apply lime.)

Characteristic plant species, uncommon  in other habitats, as well as truly rare, state-listed species,   are also often associated with  ledges, outcroppings, and  crevices of  rock formations, regardless of bedrock type. Botanizing is always rewarding in such habitats! I often find red columbine (Aquilegia canadensis) on traprock summits,  but sometimes also in areas with  a metamorphic  outcrop of gneiss or schist.     Dwarf saxifrage, Dutchman’s Breeches, Canada moonseed (Menispermum canadense), and red elderberry (Sambucus racemosa), and brittle fern  grow at the base of a low traprock cliff  near Kensington Road in Berlin.  The first four species I  often see in  undisturbed trap habitats, but rarely elsewhere.  The brittlefern (Cystopteris  fragilis) is rarer, but less tied to traprock.  The only orange-fruited horse gentian (Triosteum aurantiacum)   I have ever seen was at the base  of  a sandstone shale rock face, but they have also been found on coastal bedrock outcrops in Branford.  Rare prickly pears  (Opuntia humifusa) have been found in Old Saybrook on bedrock outcrops on “The Preserve” property; this is  probably because the open, southern exposure mimics the warmer growing conditions at the center of its range., and may – or may not – be also related to mineral-rich soil.

 

Questions

The term minerotrophic is widely used, but solid data is lacking as to exactly which minerals are needed by which plant species, and at what levels.  What are the relative roles of microclimate and soil mineral needs, as they affect plant  distributions on rocky summits and outcrops?    How often is the distribution of a “rich site”  species limited, not by soil composition, but rather  by competition with other plants?   Many  uncommon plants are known to be characteristic of rocky habitats.  How often is this  due to the role of rocks and boulders in reducing competition, rather than mineral availability?  To what extent are “rich site” plants found along slope bases or on “rocky site” plants on summits because  the areas were historically too bouldery for farming, so that the plants remain there, but were long since eliminated elsewhere by  agriculture?   A telling comparison is the nearly pristine, and botanically diverse, forested  north slope of the traprock ridge at Dinosaur State Park, versus the depauperate east slope, which has been farmed for over a century.  Parts of this this field  are  infested by invasive Japanese barberry and burning bush, and the dominant ground cover is the prickly dewberry, a very common dry-site plant.  But even this field, also supports populations of uncommon plants  like Carolina rose and panicled bush clover, growing in sweet (subacidic), mineral-rich soil with traprock near the surface.

These are opportunities  for interesting ecological research!  We really have not advanced very much past the “rich site”  or “rocky site”  habitat  characterizations in the old botany manuals.