Notes on Florida Wildflowers (Text Only)

Accompanying the Wednesday Wildflower Webinar Series

For Slideshows: 11 December 2024 Botany of Florida Wildflowers; 15 January 2025 Ten Big Ideas; 19 February 2025 Coming to Terms with Plant Names; 16 April 2025 All in the Family; 21 May 2025 Bountiful Jargon; and 18 June 2025 Knowing Keys, Knowing Plants.

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Introduction – Wildflowers and Florida Wildflower Foundation

Florida Wildflower Foundation (FWF) is a non-profit membership-based organization that  “protects, connects, and expands native wildflower habitats through education, research, planting, and conservation.”  Providing information and opportunities for people to study and learn more about the nature of our native wildflowers is basic to advancing that mission, the very purpose of these notes and the Webinars.  

Some people may think of “wildflowers” as wayside elements, cheerful posies along roads and paths, but FWF defines them as Florida’s native herbaceous flowering plants.  With nearly 2800 different kinds of flowering plants native to Florida, most of which are herbaceous or perennial subshrubs, we have to recognize wildflowers are important components of Florida habitats – habitats that are more varied than people might suspect.  Yes, the ground here is sedimentary, a mix of calcareous and deep sands, and the low, rolling topography which tops off at 345 feet elevation in the Britton Hills, (Walton Cty, near Lakewood) means Florida lacks varying habitats of regions with mountainous peaks and gaping valleys.  But extensive lowlands and coastal strands (1,350 miles of coastline that burgeons to 8,436 miles when tidal areas are included) and a very humid climate that stretches from warm temperate to near tropical have gifted this region with a rich variety of habitats and plant life. Our mild, wet climate, which supports extensive swamplands, bogs, and welling springs interrupting extensive flatlands yielding to pleated deep sand, scrub-inhabited ridges and ancient dunes, then ringed by a variable coastline, fosters incredible ecological variation, supporting a native flora acutely responsive to ground water conditions. Florida’s impressive biodiversity is not based on substrate and climate alone, but also historical and geographical.  Areas in the Panhandle and central ridges harbor vestiges of ancient connections with North America’s temperate flora while neotropical connections in the Peninsula introduce elements from the Caribbean Islands as well as Central America.  .  

Covering the diversity and nature of plants around us, something I wish we could undertake fully, would be a greater challenge than most people have time to accept.  Certainly, at retirement age, I will never have the opportunity to appreciate the diversity and ecological interactions that distinguish Florida’s wildflowers.  In this series of discussions, therefore, I’m whittling that challenge down by focusing on topics that may be most useful to people who share an interest in wildflowers.  That goal marks the Wednesday Webinar series that begins in December 2024, presenting information concerning the basic botany, taxonomy, identification, systematics, and ecology of wildflowers

Topics and dates for the series are:

I’m hoping Wildflower enthusiasts will find these informal discussions provide useful information concerning plants they encounter, make literature about wildflowers more accessible, and increase knowledge of native plants to cultivate in their own surrounds.  As importantly, I’m hoping to learn from participants, who will have more detailed and extensive knowledge in many important topics.  

Browse the following notes; don’t feel compelled to read straight through.  They are provided as supplemental information to the webinars, and as encouragement for feedback from other enthusiasts and field professionals, which will encourage me to improve and amplify information provided.  One purpose, of course, is to build a stronger community of people who appreciate and support the general mission of Florida Wildflower Foundation, as well as other organizations dedicated to conservation of nature.  Another is to encourage people to take study into their own hands, learning enough about plants to become citizen scientists, contributing to our community of understanding regarding wildflower biology and conservation.

Basic Botany of Wildflowers

Jump-Links to Contents:

Defining what we mean by “Wildflower.”

Working with the FWF definition of wildflower as any “flowering herbaceous species that grew wild within the state’s natural ecosystems in the 1560s when Florida’s first botanical records were created,”  presents us with a few basic questions:

  • How do we define “plant”? What differentiates “flowering” plants?  What plants do we consider “wildflowers?”
  • Which plants are native, and which are not? 
  • What are useful ways to describe the nature and behavior of flowering plants?
  • How do plants grow & develop?  What do we mean by “herbaceous” – how do herbaceous plants differ from others?
  • How do plants reproduce, disperse, and establish?  What aspects relate most particularly to wildflowers?
  • What are some basic ecological interactions that help us better appreciate  and conserve wildflowers?

What distinguishes Plants from other living beings?  

Answering that question opens the floodgates to technical and arcane jargon, but here it is: Among the many kinds of living things on our planet, several hundred thousand are plants, distinguished as sedentary life forms that grow from points and zones (meristems,) share unique cellular characteristics (division through formation of cell plate, spinning cell walls, reliance on vacuoles), and are basically autotrophic (“feeding” themselves through photosynthesis).    Plants are linear, with growing points creating stems and roots that extend and branch into soil, water, and air. Woody plants manage to thicken stems and roots through blanketing layers of growth formed by lateral meristems; we call this secondary (2º) growth.

Plant cells become stiffened over time, surrounding themselves individually with a fabric of cellulose fibrils which eventually constrains and immobilizes cells in a manner different from animal cells that lack this cellulosic “cell wall”. Inside plant cells, we discover special features not found in other life forms, membrane-bound liquids called vacuoles, as well as immensely important, self-replicating bodies called plastids. These plastids can become chloroplasts, gifting plants the capacity to turn carbon dioxide into food and water into oxygen – substances on which the entire living world depends

Background:  Though plants have forever been part of human experience, ideas as to how one defines a plant have sharpened over the past few decades.  For centuries, people agreed that plants are living organisms.   Along with other sedentary, non-sentient living organisms, such as fungi and algae, they were regarded as the “vegetable kingdom” – condensed in the phrase “animal, vegetable, or mineral,”  a simple formula encapsulating a Western philosophical hierarchy – animals live, reproduce, move, and are sentient.  Plants live and reproduce.  Minerals exist. By default, if something wasn’t an animal or mineral, it was a plant.  Over centuries of study, however, we’ve learned enough about the biologies of living beings to realize life is one grand pageant.  All living creatures share several billion years of convoluted ancestries that diverged into distinct branches in the tree of life, marking the major Kingdoms currently recognized: Bacteria, Fungi, Protists, Plants, & Animals.  

Study of the myriad life forms is the realm of biology, and the realization that Life is Cellular is a foundation of this science.  But our understanding is not ancient; about 200 years passed after the explosion of printing in the West (Gutenberg, c1452), voyages of exploration (Columbus, 1492, de Gama, 1497) , and rise of the Spanish Inquisition (1478) until existence of cells was appreciated.  All of the somewhat quaint herbals  (Bock, Fuchs, even Gerard) were written, illustrated, and published with no understanding that plants and animals are constructed of cells, lacking any knowledge of microscopic life forms, such as bacteria and protozoans.  Jon Baptiste von Helmont, lauded for the first experimental proof that plants did not “eat soil” is also noted for describing a recipe for spontaneous generation of mice that employed grains of wheat and dirty cloth. Even given today’s widespread cultural rejection of science, people do not seem to have reverted to belief that mice and rats arise de novo from old rags.

Not until microscopy extended our eyesight in the second half of the 17th century did Robert Hooke (a contemporary rival to Isaac Newton) publish the first illustrations of cells in his 1665 Micrographia.  In fact Hooke coined the term “cell” based on the resemblance of compartments he examined in a slice of cork to tight quarters in which prisoners were kept.  Even he had no idea that scientists would, eventually, discover all life forms are made of cells.

During the 3+ centuries since that revelation, researchers explored the structure, composition, physiologies, and replication of cells, allowing us to understand how organisms grow, develop, reproduce, and perish.  Beginning in 1802, we employed the new term “biology” as a predictor of the great advances that would come in the 19th and 20th centuries.

By 1850, most notably through work of German scientists Matthias J. Schleiden and Theodor Schwann, our realizations concerning cellular organization of living organisms were consolidated as the “cell theory” – a concept amplified in 1855 by Rudolf Virchow’s explanation “Omnis cellula e cellula” – all cells come from existing cells.  Today we know that all living beings:

  • share distant, common ancestors (an outgrowth of late 19th century evolutionary concepts elucidated by Darwin)
  • are directed through related genetic mechanisms (early 20th century, triggered by Mendel’s 1866 publication summarizing his experiments in hybridizing peas)
  • show similar cell components & physiologies (19th century)
  • utilize glucose as the common currency of energy (mid-20th century) and water as the medium
  • alternate between haploid and diploid states in sexual cycles (20th century)

This means we also understand that, based on cellular organization, living beings:

  • grow & develop – generating & differentiating cells, responding to stimuli, shedding components, apoptosis
  • acquire & process nutrients – interacting with the atmosphere & media, sharing mutualisms and parasitisms, decomposing
  • reproduce & disperse – establishing populations, sharing a gene pool, hybridizing & introgressing
  • interact with the environment, adapting to change, & evolving over generations – reconstructng species limits, speciation & extinction


Plants share significant biologies with all other life forms, but as summarized at the start, plants:

  • show particular and important cellular specializations, such as cell walls, vacuoles, etc.
  • photosynthesize to make their own food, requiring no vitamins or organic compounds
  • grow from localized meristems
  • reproduce through a variety of means, both sexually (sporic meiosis) & asexually
  • generally occupy a set location, being mobile only at certain stages, as spores, pollen, or seed
  • range in size from minuscule to individuals with the greatest age and biomass of any organisms alive on earth today.

What makes a Flowering Plant different from the rest?  

The Plant Kingdom comprises four basic groups gardeners and naturalists will recognize: 

  • Mosses and Liverworts, 
  • Ferns and their very distinct Allies (Equisetum, Quillworts, Psilotum), 
  • Gymnosperms (Conifers), and 
  • Angiosperms (Flowering Plants). 

We distinguish these broadly based on form and lifecycle.  

1.  Mosses and Liverworts are simple, and do not develop complex, vascular systems – tissues that move water, nutrients, and organic compounds through stems and leaves. Since mosses do not make veins, botanists will not recognize them as producing true roots or stems, or even real leaves.  Lacking developed vascular tissue, these “Bryophytes” necessarily keep close to moisture.  Moreover, cells of their green and conspicuous stages bear nuclei with only a single complement of chromosomes – their photosynthetic stages are “haploid”.  These are among the most ancient of plants, and by composition and definition, they are not counted among Florida’s wildflowers.

2.  Ferns and their allies are more dimensional components of the landscape, distinguished from mosses in having true leaves and stems that are of diploid constitution (each cell has two sets of chromosomes), but lacking the structure and biology necessary to produce seed.  Ferns reproduce simply through generating spores that germinate as green, free-living plantlets (gametophytes) which, like mosses, have cells with a single set of chromosomes.  Allied with ferns, you might encounter other native plants that reproduce through dispersal of spores, such as Equisetum (horsetails). Though prominent in the natural landscape and precious in form and function, ferns and horsetails do not generate flowers or seed, and thus fail to qualify as Wildflowers.

3.  Cone-bearing plants are seed producers.  Florida’s flora includes many Conifers, important pines (Pinus) and cypresses (Taxodium, Chamaecyprus), rare Taxus and Torreya, abundant Juniperus, and Zamia – cone-bearing plants all classified as Gymnosperms.  The“sperm” tells us these plants make true seed, but those seed are produced in ovules borne on modified branches and scales, not from ovules formed inside a pistil.  Thus botanists describe the seed as “naked”, as signified by “gymno” being Greek for ‘naked’.  Lacking true flowers, none of our Wildflowers are to be found classified as Gymnosperms.

4.  All Wildflowers will be found among the Angiosperms, the flowering plants.  Representatives of Angiosperms are discovered more recently in fossil records than Gymnosperms. For this and many reasons we consider the flowering plants more recently evolved.  These are, of course seed-bearing plants, those seed being formed through production of specialized structures described as flowers.  We’ll explore flowers later, but in order to differentiate Angiosperms from Gymnosperms, one must consider the presence of anthers and carpels (pistils) as components of flowers, with ovules forming along a fertile zone inside the carpels – loosely to tightly enclosed.  That gives us the term Angiosperm, which derives as seed forming in the chamber (angio) of a carpel that will form a fruit.  Flowering plants, therefore, are also fruiting plants in that a seed-bearing fruit is the mature carpel (pistil) of a flower.  All wildflowers are Angiosperms

Subjects: Sorting Nature’s Cornucopia of Flowering Plants

How do we make sense out of the variation in morphologies (form), biologies (living functions & strategies), distribution (biogeographies), and ecological interactions among the world’s several hundred thousand kinds of naturally-occurring plants?  What sense does this bring to our understanding and appreciation of wildflowers?  Before discussing the life cycle of wildflowers, the following subjects explore different subjects that explain some of the ways botanists group plants to bring order and understanding. Order, of course, begins with a proper name.

I. Nomenclature & Systematics: Taxonomists group and compare plants based on what has been learned about their nature, utility, diversity, ecologies, and evolutionary relationships.  We teach and talk about plants as individuals, and botanists will designate a single plant specimen (or drawing) as the type that represents the species, but taxonomically and ecologically, plants inhabit areas, and function as populations.  That means a plant is understood as a collective, any new specimen examined invites analysis of the broader population that could be scattered through habitats of thousands of acres or very localized. Our discussions almost seamlessly migrate from description and concern over that individual specimen to generalizations that imply or require knowledge of the population as a whole.

That’s because you’ll only learn about the biology of a plant by considering its range, and its nature and variation in that context, what some might call its “autecology”. Widely-occurring plants normally will show variation that has to be explained and dealt with systematically; narrowly-occurring plants, endemics, explain themselves more easily, in that we may more readily identify habitat and history at play. Many Florida wildflowers will be encountered in Wikipedia as endemic (for the ISB table of endemics, follow this link) to the Panhandle and to the Scrub spine that runs down the Peninsula (often segregates and close relatives of more wide-spread forms) speak to ancient connections and periods of isolation. At some point, a name is applied and accepted based on a taxonomist’s concept of what constitutes a population, covering some natural range, reflecting a collective concept of a “species”(or subspecies or variety).  With occasional plants circumscribing a species seems straightforward, but it’s commonly more fluid, exasperating actually.  

Plants botanists designate as distinct entities have valid scientific names, and we have an international convention on botanical nomenclature that formalizes the ways in which a plant name is published and recognized.  The goal of that convention is to ensure everyone in the world uses the same scientific name for every plant, which means names should be stable.  We all know, of course, that’s a pipe dream.

For those who are paying attention, plant scientific names seem to change regularly, but take heart.  There are two useful aspects to this.  The first is that if you learned a plant by its proper scientific name many years ago, that name remains useful, and will be the key to tracking down the current name.  Never be embarrassed that your terminology is “out-of-date”.  There are breadcrumbs that will get you to the current binomial, because Botanists, generating their many indices will have kept records of what various authors called a plant, tucking those names into the seemingly endless “synonyms” that fill the literature.  The “correct” name is not as important as the information you know about the plant tied to its handle and many synonyms. 

The second important reason plant names change is that a name is not simply an identifier, increasingly the name reflects much of our total knowledge of that plant.  Baked into a plant name today are hundreds of characteristics that help define one species as compared to another, give meaning to delimiting genera, and bring order to higher level classification. Thus Nomenclature and Systematics introduce plants to us, providing a wealth of associated information that reflects how we believe plants evolved, how populations vary and should be named, and are the key (the handle) to the heap of literature hiding out in stacks of books and journals, and, increasingly, in the WWW.  

Some History:  By the end of the 1700s, botanists were discussing plants as representing natural orders or families, as today we think of Orchids being a group distinct from the Grasses and Composites.  Two plus centuries later, we interpret the many living plant species as relics of hundreds of millions of years of evolutionary change.  It’s easy to forget that many species, missing from the fossil record and unknown to science, have gone extinct – many more than what’s extant (a nice word that means they haven’t yet gone extinct.) 

The image of a tree of life is fitting, in that the greater biomass of a tree is dead cells, xylem.  What lives is a thin sheath of active cells connecting roots and shoots.  Most shoots were pruned away over the millennia. leaving today’s constellation of root and shoot tips that represent life on Earth. Plant Orders are major trunks, breaking into load-bearing branches we label Families, and finally to smaller branches interpreted as Genera that bear still-living growing tips of populations equivalent to species.  The stems and branching patterns reflect ancestry, with systematists having established as a core goal that each taxonomic grouping represents a “clade” – one of the thousands of boughs on the tree of life, a tree that developed from a single ancestor. 

In botanical context, “wildflower” is not a systematic term. Wildflowers occur throughout the Angiosperms, though we certainly encounter more in families with greater numbers of herbaceous species, such as the Orchids, Composites, Legumes, Mints, Grasses, and Sedges. Higher level plant classification (Systematics) is, however, a helpful tool, formalizing one of the most useful filters, the basic traditional split between Monocots and Dicots. The Monocot clade is coherent, a clear major branch in the tree of life, and one that’s overwhelmingly herbaceous. We find a host of characters that help define this group, such as scattered vascular bundles, parallel veins, sheathing leaf bases, and singular embryonic leaf development.  Those characteristics are a major reason most Florida monocots (Grasses, Sedges, Rushes, Orchids, Lilies & Amaryllids, etc.) are herbaceous, and qualify as wildflowers

II. LIFE-FORM. “A device designed in Hell” – Early in the last century, Danish botanist Christian Raunkiær proposed a system describing plants based on where their growing tips (meristems) hang out during the most adverse seasons.  Though clearly a temperate European approach, Raunkiær’s life form system continues to be useful for ecologists, perhaps most appropriately in comparing physical characteristics of plants in different floras.  For wildflower enthusiasts, the most useful aspect is focusing on how a plant overwinters. What does it die down to?   It’s good to know these terms because you might run into them in ecological literature, and because Raunkiaer’s system has its practical side.  The basic categories, today, are: 

Phanaerophytes – larger plants, rooted in the ground, with growing tips exposed to the elements throughout the seasons), a half meter or taller.

Chamaephytes (shorter plants, specifically plants in which the growing buds are less than ¼ meter from the growing surface), 

Hemicryptophytes (plants with perennating growing tips at ground level, such as rosette-producing  composites, i.e. Chaptalia tomentosa, Helianthus radula, Violets, and most grasses.)

Cryptophytes – plants with growing buds buried in soil or water.  Bulbous wildflowers, such as Crinum, are considered Geophytes, as are rhizomatous perennials such as Lygodesmia.  Hydrophytes are a subcategory of plants with growing points resting underwater (such as Nymphaea), while Helophytes include resting points in mud and marshy soils.  

Therophytes – These are the true annuals and monocarpic plants that live a single growth cycle passing through dormancy as seed.  Almost any native flowering plant Raunkiær would consider a Therophyte would be a wildflower. Sabatia stellaria and Hypericum gentionoides are examples.  Long-lived monocarpic plants, such as many Agaves, are not considered therophytes.

Epiphytes & Aerophytes (plants growing in the air, on materials other than soil)  Herbaceous examples, such as Tillandsia and Encyclia are wildflowers. https://en.wikipedia.org/wiki/Raunkiær_plant_life-form  

III. ANATOMY:  Plant Anatomy gives us many ways to examine and group plants.  A common  differentiator ties to primary versus secondary growth, leading to four basic types of plants: Herbaceous Dicots, Herbaceous Monocots, Woody Dicots, and Woody Monocots.  

For wildflowers defined as herbaceous, this category introduces challenges, generating a middle kingdom of perennials that are “woody” but short-lived, or bear “woody” roots and stem bases that regenerate fresh shoots each year.  Miller et al (2018) describe Hypericum tenuifolium (Atlantic St. John’s wort) as “a semi-woody to herbaceous perennial subshrub,” Iva imbricata (Seacoast marshelder) as a “succulent shrub-like perennial herb,” and Chrysoma pauciflosculosa (Woody goldenrod) as “a multibranched, evergreen sub-shrub.” Hammer (2018) and Taylor (2013) treat all three as “wildflowers.”

Outside that middle world, some plants that are clearly woody (producing secondary growth, i.e. annual rings) are typically included as wildflowers.  Hypericum provides a clear example. Some species, Hypericum gentianoides and H. mutilum, are absolutely monocarpic (annuals) and totally herbaceous.  Others, particularly the long-lived Hypericum fasciculatum and H. chapmanii types, are patently woody shrubs, perhaps even small trees.  Somewhere in the nether-regions, we encounter Hypericum tetrapetalum and H. crux-andrae that perennate as open, hard-stemmed shrubs but would likely be considered wildflowers.  Raunkiaer would consider them large chamaephytes.  Other Hypericums that seem woody are yet closer to herbaceous; as mentioned above, Miller et al (2018) detail the coastal Hypericum tenuifolium as an herbaceous subshrub.  The same reasoning might apply to our spectacular hardy Hibiscus species (Hibiscus moscheutos, H. laevis, and H. coccineus).  Dying to the ground at season’s end and overwintering as underground stem and roots, Raunkiaer would consider these cryptophytes, while most people might think of them summer shrubs.  The distinction proves difficult.  When it comes to structure, perhaps wildflowers turn out to be plants with seasonal stems readily eliminated by a weed trimmer or mower. 

Monocot vs. Dicot Anatomy – The most evident anatomical differentiator for flowering plants is the basic monocot-dicot distinction as to how veins are laid down (generated from growing tips) to form the vascular systems in stems and roots. This is true intro botany; almost anyone who has had a class in plant biology will have seen diagrams or even microscope slides showing primary growth of Sunflower (Helianthus) or Alfalfa (Medicago) with vascular bundles basically in a ring (at least forming a single cylinder), as compared to a stem of Corn (Zea) that shows bundles more evenly distributed throughout the cross-section. If you had an exuberant teacher, you may have even encountered stem sections of Potamogeton, a monocot, that shows scattered bundles appearing to form rings due to bursts of growth similar to the way dicots form annual rings. Generally, teachers would stay away from exceptions, since the singular goal of understanding basic stem anatomy is enough for most beginning students.

Understanding internal organization of stems would be the base on which someone might successfully explain secondary growth, most likely using slides of Linden (Tilia) and Lilac (Syringa). You’d seldom be taught anatomy with native plants because it’s a lot of investment to make and acquire all of those slides. Microtomists have discovered certain plants that can be easily prepped, stained (safranin and fast green), and mounted to model the basic monocot vs. dicot anatomies, and that’s what everyone is taught. It’s good for constructing an understanding of plant anatomy, but remember there are thousands of kinds of native wildflowers, and each might have its own way of doing things.

Pathways:  In another instance, anatomy and physiology provide filters for plants based on photosynthetic specializations.  Botanists will often categorize plants based on whether the the photosystem is C3 (the earliest pathway described), C4 (common to grasses, involving particular leaf anatomy) or CAM (associated with succulents, but identified in a broad range of plants.)  Tremendous effort and the careers of many botanists were linked to decipher this photosynthetic, cytological, and anatomical puzzle.  See Subject XII Physiological Adaptations for further detail on Pathways.

IV. Life Strategies & Breeding Systems are varied and complex in the flowering plants.  Our growing understanding of the many strategies has generated various grouping systems.  

Parity (number of births) filters as either Semelparous or Iteroparous.  “Semelparous” is the overarching term for once-flowering plants, equivalent to annual, biennial, monocarpic, &  therophyte, as compared to  “Iteroparous” (reproducing in many episodes over a lifetime) which includes any of of Raunkiær’s life forms that are not monocarpic, and umbrellas the term perennial, as well as the lifestyles of most (but not all) woody plants.  

Isolating Mechanisms (Kimball, 1997): Sexuality – For the sake of the immediate discussion, let’s assume that sexual reproduction and outcrossing benefit plants.  I can explain the reasons, and will touch on that topic later, but that’s a free-standing lesson.  Take my word for it; flowers exist to promote outcrossing.  There is no other basis by which to understand the grand variety of floral form and mechanisms we observe in the natural world.

That position is confounded by realizing the base condition for flowers is the “perfect” state, which tells us an individual flower produces both fertile pollen (the male role) and ovules (the female role).  The perfect flower, then, is bisexual, termed hermaphroditic, with stamens and one or more carpels.  That tells us the closest potential target for the pollen of many flowers is a stigma in the same flower.  With outcrossing a core function of flowers, how do we explain the fact that many wildflowers are perfect, producing their own pollen and ovules?  My answer: “Don’t believe those lying eyes”. 

Plants invest in structure and behavior (shape of the flower, relative positioning of anthers and carpels, moving parts, placement of rewards such as nectar, oils, and pollen, color guides, fragrances, even temperature differences) to manipulate vectors and promote outcrossing – resulting in a variety of breeding systems that achieve seed production and dissemination while mitigating the impacts of inbreeding (self-fertilization).  

We’ll investigate some of these arcane remedies, but I need to begin with an obvious strategy that can be seen easily, producing “imperfect” flowers.  These are flowers that lack one of the two sexual roles, either bearing fertile anthers or fertile carpels, but not both.  If both kinds of flowers are borne on a single plant, we say it’s “monoecious” (mono = one; eco = household).  Many grasses are monoecious, as is Marshelder, Iva imbricata.  Two wetland wildflowers, Sagittaria and Typha, are monoecious.  Euphorbias are cryptically, or technically, monoecious, in that what is perceived as a single flower is a reduced inflorescence of several individual highly reduced flowers. Some plants will border on monoecy, such as Orontium, which produces perfect flowers generally, but staminate flowers near the tip of the spadix. But monoecious plants still confront the proximity challenge; the closest potentially-receptive flowers are likely on the same individual.  

Clearly the absolute answer would be to segregate the sexes, one plant producing male flowers only, with female flowers borne on a separate individual, a condition we do find in nature.  We say plants that produce pollen-bearing flowers (male) on one individual and ovule-bering flowers (female) on another plant are “dioecious” because the sexes are in two houses (eco = household). Numerous understory shrubs and small trees are dioecious – Persimmon, Holly, Ceratiola, Myrica (Morella), Baccharis (https://floridawildlifegardentails.wordpress.com/tag/dioecious/   Among wildflowers, the most commonly cited example is October Flower (Polyganum polygamum), but we also observe dioecy in native Dodders (Cuscuta) as well.

If dioecy is so clearcut and effective at enforcing outcrossing, why don’t we see this in a lot more flowering plants?  There must be a cost, perhaps in lost opportunity.  Outcrossing is vital, but a certain level of selfing is security.  We know many plant strategies bank on both open, outcrossing flowers (chasmogamous flowers) as well as cleistogamy, generating flowers that are closed (cleisto = closed) to outside pollen.  A number of our native violets produce open flowers early in the year, and self-pollinating cleistogamous flowers toward the end of the season.  Some seed are better than no seed.  

Dichogamy & Herkogamy:  Most wildflowers, being bisexual (Hermaphroditic or Monoecious), tells us they could, possibly, allow for pollination within a single individual, i.e. selfing.  Knowing that outcrossing benefits populations through maintaining genetic diversity, we expect to discover mechanisms in bisexual plants that promote outcrossing at some level.  Indeed, these plants have tricks of various sorts that promote outcrossing, limit potential of fertilization, or impact the viability of inbred embryos.  Asynchronous maturation of anthers and carpels (dichogamy) is commonly observed.  Flowers of some plants mature anthers and pollen before stigmas are receptive, which is termed “protandry”.   Flowers in which stigmas mature earlier than anthers are termed “protogynous”.

Flower structure also impacts the potential that pollen might land on a stigma of the same flower – which is the realm of herkogamy.  Sarracenia flowers boast luxuriant, short-lived drooping petals that we are told act to divert visiting insects away from stigmatic surfaces during early flowering when pollen is shed.  Orchids are masters of structural control, positioning pollen such that vectors are unlikely to transfer grains (or pollinia) from an anther to the stigma of the same flower.

A curious but common type of herkogamy is “heterostyly.”  Pontedieria, Lythrum and Oxalis show this strategy, generating flowers on one plant that produce short stamens and long styles, while another plant develops flowers in reverse format.  This mechanism functions to place pollen from flowers of one form in a position more likely to contact the stigma of flowers with the alternative structure, flowers borne on a different individual. Our Lythrums are dystylous, with two forms. But structures may be yet more complex. Other species of Lythrum have been recorded a tristylous, showing 3 distinct forms.

Not surprisingly, many flowers combine dichogamy and herkogamy to limit selfing and promote outcrossing. A recent study of pollinating mechanisms in a Chinese Parnassia species suggests detailed observations of many native plants we have previously considered open to pollination by a wide range of visitors would reveal striking behaviors and juxtapositions at play. I decided to observe our local Parnassia grandifolia over several days, and am intrigued to learn that when flowers first open, the green ovary shows no stigmatic surface; only 3-4 days later does the white, multi-lobed stigma expand – suggesting this plant is protandrous. Such studies take hours of field observation, often over several flowering seasons. The dedication and time required to study the intricacies of every pollination system are well-suited to the efforts of citizen scientists, people who live close to populations and can persist in dedication to study over the time needed to learn the ropes for a particular plant. 

Even seemingly straight-forward tubular flowers, such as those of Salvias and Penstemons, position stamens and styles to take advantage of particular kinds of visitors, sometimes even altering the length and position of the style and stigma over the life of the blossom. It’s a trap to assume that every insect seen on a flower is a pollinator. Close study of each different fkind of lower could help explain how timing and structure might work to limit chances pollen from an individual flower would contact its same stigmatic surface.

Mechanisms & Syndromes: Correlating the various features of floral structure presentation, timing, substance, surface texture, color, fragrance, and reward we realize flowers are truly complex event spaces, co-evolved often to a remarkable degree to harness the wind or secure the services of an insect or animal. We discover it’s often possible to group and predict pollination mechanisms based on characteristics relevant to the vector.. These are well-documented as Anemophily (wind), Hydrophily (water), Entomophily (Bee, Butterfly, Moth, Fly, Beetle), Ornithophily, (Birds) Chiropterophily (Bats), and other types of Zoophily (Possums, Lizards, etc.), and will be discussed when we get to flower pollination. Needless to say, pollination syndrome is yet another way to predict and study something about plants.

Compatability “the Birds and the Bees” – Pollination is not fertilization; it’s simply a shuttle to the chapel (the carpel), with nuptials to follow. There are many stages of potential rejection and failure once pollen has made its journey.   For the groom, there’s work to be done, and hazards ahead.  If you peruse the pollination literature, you’ll run into two kinds of self-incompatability that are often discussed: 1. “sporophytic,” in which genetics of the pollen parent determine whether a pollen grain is allowed to grow on a stigma or not, and 2. “gametophytic,” in which the genetics of nuclei in the pollen grain must be compatible with the carpel. In either case, pollen sperm nuclei will not successfully fertilize ovules on the same plant. The only reason to mention this is to alert readers to the existence of a trove of research and literature that document the ways self-incompatibility might function. It is an important feature related to promoting outcrossing. People growing native plants will discover most Composites, which have very general pollination systems, are self-incompatible; at least 60% of Composites are not self-fertile.

Sexuality vs. Asexuality – Plants do not necessarily require sexual production for dispersal.  Weedy plants, such as Dandelions are noted for “parthenogenesis”, also called “apomixis”.  They can produce seed in which the embryo forms from parental tissues, basically cloning the mother plants.  The shade-tolerant invasive shrub, Ardisia crenata, has been shown to be apomictically clonal  (Noyorui, 2024)   Other plants hedge their chances with vegetative propagules, the typical variety of Allium canadense being a prominent example. In this plant, most floral pedicels develop as bulbils. If you encounter a colony that generates normal flowers, it will be considered Allium canadense var. mobilense.

V. Flowering & Fruiting Structure

Inferior vs Superior vs Perigynous – Flowering plants produce many kinds of fruiting structures, spawning a complex and unsettled terminology.  Perhaps because ovary structure often ties to major taxonomic groupings, we fast-track differences by categorizing an ovary as superior when it’s clearly atop other floral segments or inferior, when the ovary seems buried in the stem, below the point that sepals, petals and stamens attach. That means we often ignore intermediate stages (perigynous) in which the ovary is surrounded but not completely buried below other segments.  In the Rose family, we call the resulting structure a “floral cup.” Regardless, ovary position is one of the most useful characters in determining which plant family to search in identifying an unknown plant.

A group I’ve given some attention, the shrubby Ericads, makes the point on importance of inferior versus superior. Most keys will take you to a point that Gaylussacia and Vaccinium are segregated from other genera based on an inferior ovary that generates a fleshy berry, as contrasted with somewhat similar plants (especially Lyonia) that generate dry capsules from superior ovaries. The trouble, however, is that examining those small flowers with aging eyes, even with aid of a handlens, while being assaulted by mosquitoes and flies, means it can be a challenge to determine whether or not the ovary is inferior. Of course, if last year’s fruit are present, you’re home free. Thankfully, Lyonias tend to hang onto a few of last year’s capsules.

Inflorescence & Display type – Few plants produce just one flower per growth, Lilium catesbaei being an opulent example to cite.  Most plants generate some predictable branching tipped with flowers, which we call an inflorescence (and then, at maturity, an infructescence.) Inflorescence types tend to characterize taxonomic groups, and beg to be categorized and explained.  Historically, botanists worried about which flowers develop first and how to characterize the branching pattern.  This led to comparisons and terminologies such as Determinate (generating a terminal flower that opens earliest) vs. Indeterminate (producing sequential flowers until the growing tip simply peters out). By that system, Determinate was considered essentially equivalent to Cymose and Indeterminate became associated with Racemose, except when we encountered plants that reversed the formula.  Search the web for an herb  with determinate branching, and you’ll likely find Liatris spicata as an example, which would make it a determinate raceme, sort of preposterous when you realize that the side branches themselves are capitulae (headlike inflorescences). 

Since that common system is unworkable, botanists developed the concept of Monotelic vs. Polytelic, suggesting great distinction between development of a single primary side branch versus many side branches. But it’s not so simple. Inflorescence development is highly individualized from one group of plants to another, such that what we might determine to be racemose will be seen to produce secondary branching that appears cymose.  Moreover, in Composites, Grasses, and other groups, branching is compounded, with many flowers united as pseudoflowers (heads) or spikelets that are then displayed in primary and secondary branching patterns.  Botanists have expressed frustration with the unfortunate state of terminology applied to inflorescence structure, and decided it’s much of a mashup, so don’t worry about explanations that create an illusion there’s uniform agreement in this realm of morphology.  (Endress, 2010; Barthélémy and Caraglio, 2007, Barthélémy, Daniel and Yves Caraglio, 2007.) Stay tuned for some future “Come to Jesus” moment on this terminology.

In the meantime, we simply use the more common terms historically employed for descriptions, which means you’ll still want to know the common terms, but will understand each author might have his or her own perspective. Checking Liatris in Flora of North America for example, you’ll find this description: “Heads discoid, in corymbiform, cymiform, racemiform, or spiciform array”, describing Liatris spicata particularly as having “Heads in dense spiciform arrays”, which lets me know the heads will be sessile, thus arranged as a spike, lacking a visible stalk that joins them to the single main stem (the peduncle.) In the field, especially during fruiting season, a character that helps separate Liatris gracilis in overlapping populations with L. spicata, L. tenuifolia, and L. chapmanii, is the evident peduncles (pedicels) that separate heads from the main stem. You’d want to call inflorescences of L. gracilis “racemiform” as distinctive from the spiciform (spike-like) structure of the others. This is just one example of how working descriptions simply avoid assuming an overall organizing system. It’s tacit recognition botanists may never conceive a workable system to describe inflorescence structure.

VI. Architecture –  Plant branching patterns depend on ranking (organization of buds around the stem axis), spacing (distance between nodes), nature of axillary buds, and apical control over the growth of lateral buds.  Strict apical dominance will yield a plant with strong apical growth and few to zero side branches; a modest hormonal shift might produce something highly branched..  Many systems for comparing architecture have emerged, such as Determinate vs. Indeterminate growth, Rythmic vs. Continual Growth, Monopodial vs. Sympodial branching, etc. (Barthélémy et al, 2007) Florida’s native Sabal palms are near perfect examples of trees that show strong apical dominance.  The height and density of branching in different woody Hypericums evidences differing levels of control over growth of side branches.  In totally herbaceous plants, branching usually will shift pattern from that of vegetative growth to characteristic inflorescence patterns.  The Umbels (Apiaceae), almost uniformly, shift to umbrella-like branching, either very loose as in Tiedemannia, or tightly constricted as with Eryngium.  Highly-condensed flowering structures as we see in the heads of Composites, the strict spadix of aroids and the crowded rachis of Typhus are notable extremes. 

VII. Origin – The Institute of Systematic Botany’s Atlas of Florida Plants, an easily accessible and impressively intuitive database (https://florida.plantatlas.usf.edu) indicates whether a plant is Native or Non-native (Alien, Exotic).

Exploring the ISB Atlas you’ll notice terms (and links to definitions) that describe how a plant might be listed, or protected. These include: 1. Florida Listed status, either Threatened or Endangered, 2. Florida Wetland Status, either Obligate, (OBL) Facultative Wetland (FACW), or Facultative (FAC), 3. Federal Wetland status, listed as Obligate Wetland, Facultative Wetland, Facultative, Facultative Upland, or Upland (definitions given in an ISB popup), and Federal Listing, with 943 US plant species designated as Endangered or Threatened. As long as the Fish and Wildlife Service is allowed to track conservation status of plants and animals, that list can be searched through the ECOS (Environmental Conservation Online System) at the following website: https://ecos.fws.gov/ecp/ Internationally, you might encounter a plant listed as Vulnerable (VU), Endangered (EN), or Critically Endangered (CR) in the “Red List” of Threatened Species, a program that began in 1964 under auspices of the International Union for Conservation of Nature (IUCN).

Regardless as to status, I imagine many people are unaware which plants might be native vs. exotic, or when and how non-native plants might cause problems in natural and cultivated landscapes.  Non-native plants would be most common in built landscapes, such as waysides, disturbed sites, gardens, and agricultural lands, all of which would harbor ruderal plants so familiar that many pass as natives. Moreover, our treatment of disturbed and maintained landscapes selects for plants (both native and non-native) that thrive around human settlement, especially weedy plants that establish readily, reproduce and spread quickly in cleared sites, and often benefit from mechanical and chemical intervention. In general literature, you’ll encounter alien plants described variously as Cultivated, Introduced, Escape, Vagrant, Ruderal, Weedy, or Invasive, those varying terms overlapping in use, but implying subtly different outcomes. Vagrant plants are temporary specimens or populations encountered at sites of previous occupation, plants that persist from original cultivation but are unlikely to spread. Ruderals are cosmopolitan, weedy herbs that populate waysides and are unlikely to be extirpated. They prosper along mowed waysides. Weeds are successful ruderals that are problematic to gardeners and farmers. Invasives are weedy non-native plants that move into and modify native habitats, implying various levels of harm to natural ecosystems.   The US Government (as of December 2024) officially defines Invasive Species per Executive Order 13112 (Section 1. Definitions) as being: 1. non-native (or alien) to the ecosystem under consideration and, 2. whose introduction causes or is likely to cause economic or environmental harm or harm to human health.

Problematic non-native plants include grasses adapted to grazing, many of which were introduced for pasturage, legumes that harbor nitrogen-fixing bacteria and thus thrive in our sandy, nutrient-depleted soils, vines that shade out small trees and understory shrubs, and woody plants (trees and shrubs) that occupy canopy and understory, sometimes even altering natural patterns of cover.  A search of all flowering plants reported as wild in Florida (not just wildflowers) using ISB yielded 2766 species considered native and 1513 species designated as non-native.

VIII. Phytochorion (Floristic Region) – Following on work of British Botanist Ronald Good, who studied the geography of plants, Russian-Armenian botanist Armand Takhtajan elaborated a scheme of floristic regions, natural areas considered the ancient cauldrons of plant evolution and diversity.  This imagines the major assemblages of plants that were the basis of regional floras we know today.  North Temperate plants, regardless as to whether Asian, European, North American, or Arctic, evidence an ancient “Holarctic” flora, with Takhtajan’s other major floristic regions being “Palaeotropical”, “Neotropical”, “Australian”, “South African”, and “Antarctic”.  More recently, Yunpeng et al (2024) have retrofitted Takhtajan’s system, recognizing 2 realms,  Gondwanan vs. Laurasian, with 8 regions.  Florida’s native plants are predominantly residuals of the Holarctic flora, with some Neotropical representation in the Peninsula. 

IX. Environmental, such as Habitat (Biomes, Major Habitat types, and Communities), Habitat Zone (Canopy, Understory, Epiphyte, Lithophyte, Strand, Emergent, Floating, Submerged, and at a more detailed level, Niche, and even more specifically Microhabitat: localized but commonly available sites, such as Crevices, Pools & Pocket, Stumps, etc.  The wonderful Pieris phyllyreifolia establishes on root bases of swamp trees, most particularly Cypresses.  In wet prairies, Helianthus heterophyllus is frequently seen establishing on crawdad mounds.

X. Life Zones – Leslie Holdridge devised his system of Life Zones based on work in the tropics, refining concepts first published as early as 1805 by Alexander von Humboldt, then amplified and recast over the 19th century by J. F.  Schouw (1823), Alfonse de Candolle (1855), A. H. R. Grisebach (1872), and C. Hart Merriam (1889).   Correlating belts of latitude with zones of altitude, the Holdridge scheme employs precipitation, temperature, and potential evapotranspiration to chart potential for development of differing sorts of environments (life zones.)  These have been formalized into 38 classes, with the Florida Panhandle as Warm Temperate Moist Forest.   https://en.wikipedia.org/wiki/Holdridge_life_zones

XI. Climate – Similar to the Life Zone concepts of biologists, Geographers have devised many systems to categorize world climate, the most commonly utilized being the Köppen-Geiger Climate Classification system.  In this system, the entire Southeast is “Cfa” (Humid Subtopical), as are Japan and much of Southeast China, southern Brasil, and wetter areas of Southern Europe.  The very southern tip of Florida is classified as “Am” (Tropical Monsoonal)

XII. Physiological Adaptations – Ecologists often frame studies based on which Photosynthetic Pathway (C3, C4, or CAM) characterizes plants in an area. This knowledge was elaborated over the past century. Working in Great Britain, Melvin Calvin, James Bassham, and Andrew Benson are credited with “discovering” the C3 system in 1950, the first and most common photosynthetic pathway described. Sometimes called the Calvin-Benson-Bassham cycle (CBB), the term C3 reminds us the initial product is a 3-carbon sugar (two of which make a glucose). Very soon, however, other botanists were discovering alternative photosynthetic products, which led to the 1966 detailing of a C4 pathway in Australia by Marshall Hatch and Charles Slack. Sometimes called the Hatch-Slack pathway, C4, which particularly characterizes certain grasses, captures carbon dioxide extremely efficiently, generating a temporary 4-carbon compound that can be relocated to specialized zones along veins, where the carbon dioxide is released to pass through the CBB cycle. We simplify this with an understanding that C4 plants move carbon dioxide spatially, creating high concentration levels that improve the photosynthetic efficiency for plants with good access to water and sunlight.

Other plants employ the CAM pathway, utilizing the same enzymes as C4, capturing and concentrating carbon dioxide, but sequestering it over time, banking carbon dioxide for use later. This explains centuries-old observations regarding succulent plants in the Crassulaceae, known to taste acidic early of a morning, but less acidic as the day progressed. Based on studies of those plants, the term Crassulacean Acid Metabolism (CAM) appeared in botanical literature by 1940, even before scientists constructed our more complete understanding of photosynthesis. While first described in succulent plants, CAM has emerged independently across the Angiosperms and has been identified in 35 different plant families, . frequently associated often with plants in water-stressed habitats. Both CAM and C4 exemplify how adaptations to habitats are inseparably physiological and anatomical.

Energy Source: Generally, plants we encounter are photosynthetic “self-feeders,” characterized as “Autotrophic.” But you’ll discover there are many plants, many Florida Wildflowers, that lead alternative Heterotrophic lifestyles, being either Hemi-parasitic or Holo-parasitic, as well as countless other plants exhibiting various symbiotic relationships, such as Myco-heterotrophism (fungal symbiotes) & Nitrogen fixation (bacterial symbiotes).  The great majority of Wildflowers are considered autotrophs, which means those plants take in nutrients and water, and capture sunlight to photosynthesize the sugars and other compounds needed to support life.  Purely parasitic wildflowers, such as vining Cuscuta and the ghostly Monotropa, have long been recognized as “obligate” heterotrophs (which means they have no choice otherwise). They differ, however, in that Cuscuta is parasitic, taking its subsistence from living plants, while Monotropa is saprophitic, living off decomposing life forms through mycorrhizal means.

Increasingly, however, we’ve become aware of relationships that amplify acquisition of nutrients, even carbohydrates.  Many wildflowers, Agalinis (Musselman and Mann, 1979), Gerardia, Seymeria (Mann et al, 1969), Buchnera benefit from hemi-parasitic relationships, relationships in which the host plant is plundered, receiving no known benefit. Though photosynthetic and competent as autotrophs, these hemi-parasitic freeloaders are able to tap into root systems of many other kinds of plants (mostly trees), extracting sugars and possibly other compounds.  We would describe them as “facultative heterotrophs,” the “facultative” implying they will take advantage when given the chance. Another common form of facultative heterotrophism is practised by our many native carnivorous plants, Pinguicula, Drosera, Utricularia, all of which would classify as wildflowers.

At a less extortionate level, many plants benefit from mutualisms, co-evolved relationships in which both parties benefit. Mycorrhizal relationships are increasingly understood to be common in many plant groups, essentially root-fungal entanglements that expand nutrient-gathering capacity of plant roots while providing fungi with needed carbohydrates. Similarly, many Legumes and other plants house nitrogen-fixing bacteria (termed endosymbionts) in specialized root nodules, sheltering and feeding them in an oxygen-regulated environment where the bacterial are able to “fix” atmospheric nitrogen, combining (reducing) N2 , a remarkably inert gas, to ammonia, NH4, which is then converted to various organic amides that are exported to plant tissues.

XIII. Structure, Colloquial – People use a broad vocabulary of loosely-defined terminologies in describing plant structure, such as: Tree, Shrub/Bush, Subshrub, Vine, Wildflower, Weed, Herb, Forb, Graminoid, etc.  Varyingly rigorous, the definitions overlap and change from text to text, speaker to speaker. Some authors define their terms, for example in Biodiversity of the Western Volcanic Plains  we are told what is meant by “graminoid” vs. “herb” vs. “shrub” or “tree.”  The definition of wildflower clearly varies from source to source. Hall, Weber, & Byrd (2010) include trees and shrubs, even common exotics, in Wildflowers of Florida and the Southeast. Hammer (2018) pictures and describes many flowering shrubs, though he admits herbaceous species are “what most people envision when they hear the word ‘wildflower’.” Both authors exclude graminoids (grasses, sedges, rushes) from their photoessays.

As defined by FWF, wildflower includes any native herbaceous flowering plant, which to my mind comprises non-woody vines, herbs, graminoids, and forbs, but could be construed to exclude woody perennial shrubs like Conradina canescens and Chrysoma pauciflosculosa. For this discussion, I’m relating those many terms, and how I define them:

  • Annual – monocarpic plants (semelparous, i.e. once flowering) that play out a generation in a single season of growth
  • Biennial – Monocarpic plants that may require two growing seasons to mature
  • Bulb – a short-stemmed (button-stemmed) perennial monocot that forms nested, green leaves followed by succulent interior scale leaves, the growing point for which typically buries itself in the ground (a geophyte). Hymenocallis and Crinum are perfect examples.
  • Climbing Shrub – Several native plants marry a vining capacity to shrubby flowering and fruiting structure, most notably Eastern Poison Ivy, Rhus toxicodendron, but one might also describe Pieris phyllyreifolia as a climbing shrub.
  • Forb – grazing herbs that are not graminoids, a field and pasture term employed by ecologists
  • Graminoid – Grasses, Sedges, and Rushes, i.e. grass-like plants. Ramet – one leafy segment of a grass or other rhizomatous plant, ecologically functioning as an semi-independent and countable sprig.
  • Herb – a non-woody, short-lived seed plant, may include perennials that do not develop woody basal stock. Sabatia stellaris, Hypericum gentianoides, and other annuals, Bacopa caroliniana and other creeping plants with soft stems, Chaptalia tomentosa and other rosette-forming plants that never develop woody stems or root systems. This botanical use of “herb” is distinct from the culinary use, which pairs with and overlaps the term “spices”
  • Herbaceous – any primary plant growth that has not hardened, or developed woody, secondary growth. You might read about herbaceous young growth of a plant that’s clearly woody at maturity, especially regarding tropical plants.
  • Suffrutescent (sometimes termed Herbaceous shrub or Herbaceous Subshrub) – a perennial that maintains a woody base and varyingly woody or succulent stems that are lost and replaced over the life span.
  • Liana – a woody, long-lived vine, often aggressive enough to live in tree canopies. Our most ever-present liana in the Panhandle is Vitis rotundifolia.
  • Perennial – a long-lived iteroparous (flowering many times over a lifetime) plant, normally an herb or subshrub, but applicable to shrubs and trees also.
  • Shrub – a multi-branched, open to densly foliate, long-lived woody plant. Basically equivalent to the term “bush”.
  • Subshrub – used in tropical habitats to describe plants that persist with a structure of stout, herbaceous stems. Many tropical exotics cultivated in South Florida show this habit.
  • Tree – a woody (usually generating successive rings of secondary xylem), long-lived plant that produces one to very few boles, usually forming canopies. A plant you can walk or sit under. By this definition Sabal palmetto is an honorary tree. lacking rings but having permanence. Elbert Little (1978) sets what is, perhaps, the most technical limits on tree: “Trees are defined as woody plants having one erect perennial stem or trunk at least 3 inches (7.5 centimeters) in diameter at breast height (4.5 feet or 1.3 meters), a more or less definitely formed crown of foliage, and a height of at least 13 feet (4 meters).” This includes a lot of plants we encounter more often as shrubs.
  • Vine – any plant that climbs or depends on other plants and structure for support. An Herbaceous Vine doesn’t develop significant woody tissue, for example the Apocynaceous Swalloworts, Pattalias (syn. Cynancum, Seutera.) The invasive Kudzu, Puerara lobata, is an herbaceous vine that dies back to tuberous roots in cold weather. I’ve chosen to use the term Greenbrier for the many sprawling, viney, basally woody Smilax species that are so common in our flora. For long-lived woody vines, such as Campsis radicans and Vitis, I use the term Liana.
  • Wildflower – In this treatment, speaking to Herbaceous flowering plants, also granting honorary status short-lived subshrubs (i.e. Conradina canescens) and near relatives that generate long-lived woody stems (such as Hypericum fasciculatum). I call them “wildwoodies.


People often discuss plants using overlapping but practical, utilitarian categories.  As comestibles, we speak of vegetables, herbs, fruit trees, nuts, legumes, and grains, even food plants..  In fabrics we talk of bast fibers (linen), leaf fibers, and stem/fruit fibers (cotton), as well as dye plants.  For construction, we discuss hardwoods, softwoods, palms, bamboos, and rattans.  In home gardens we reference trees, shrubs, etc., but also speak of foundation plants, row crops, aquatics, full sun or partial shade, and even 4″ color. Some Florida wildflowers are recognized as medicinal herbs, some as symbolic or iconic. We’ll not worry about these casual nouns, only note that their maleable meanings remind us plant life is part of everyday culture and utility,

XIV. Ecological & Ecosystem Services plants provide range across many dimensions. Plants are Earth’s primary mantle, it’s vegetative cover. In forest layering we recognize differing strata in different biomes, and speak of plants based on which layer we expect them to occupy. For temperate forests, a common approach might be to recognize, from ground up, the rhizosphere, the forest floor, an herb layer, a shrub layer, an understory, and a canopy. The totality of plants in a forest, or a prairie, marsh, or desert create structure, habitat, nutrient cycles, soil development, as well as erosion, temperature, and humidity mitigation.

More specifically, for animal life we classify plants as browse material, seed and fruit sources, hosts for larvae, pollen and nectar providers, nesting materials, litter and humus sources, granary trees, cover, etc. Special words abound, since plants are the source of food, vitamins, and phytochemicals that range from toxic to beneficial. This appreciation should also include realization that plants color the natural world, elaborating a kaleidoscope of greens and every other hue that isn’t merely backdrop, but nature’s surround in which animals display and camouflage.

All of this and we still must remember that photosynthesis by plants, protists, and bacteria is not just the source of food for all life, but has generated the oxygen that makes our Earth so very different from other planets we know. Oxygen, at about 20% of the atmosphere, makes fire and respiration possible. Iron rusts, aluminum oxidizes, ozone blankets. We live in a world ruled by fire and water, alter egos to Oxygen, an element characterized and named by famed chemist Antoine Lavoisier in 1777, but discovered a few years earlier by Joseph Priestley, an English parapatetic polymath, philosopher, and perennial dissident. Burned in effigy as the devil himself, Priestley immigrated to the US, where he spent his final decade. It’s unclear how he’d fare in the contemporary world.

Lavoisier was even less fortunate. Though the greatest chemist the world ever knew, he was beheaded by French Revolutionaries at the age of 50. Mathematician Joseph Louis Lagrange, in translation, remarked: “It took them only an instant to cut off this head, and one hundred years might not suffice to reproduce its like.” Science and Scientists were, and seemingly remain, elite scapegoats to an angry mob.

How do wildflowers grow and develop?

Life cycles of Florida’s wildflowers exemplify what we know about herbaceous plants generally.  Stepping into that cycle with Seed Germination & Establishment, we tackle the first not-so-simple question:  

What is a seed?  I might, incautiously, answer by claiming that a seed is a dormant embryo  tucked inside a mature ovule.  That means a seed develops from an ovule that formed on fertile tissue (placenta) in a carpel.  

At maturity, therefore, we think of a seed as a prepackaged plant, ready to sprout and set up housekeeping.  But there are caveats, as well as differences among plant groups.  

The textbook seed will be that of a dicot, such as a legume.  As with seed of all flowering plants, it will have formed inside a carpel (a specialized leaf) that matured as a fruit.  The fruit might be a dry capsule, a fleshy berry, or whatever specialized structure defines the plant group.  If the fruit becomes a dry capsule, seed may simply be shed, or as in Partridge Pea, there might be some mechanism that sends seed flying a goodly distance from parent plant.  Other plants develop fleshy fruit, such as the berry of Ilex or Vaccinium, typically eaten by birds and other animals, to be dispersed later in time. 

Given appropriate timing and conditions, the seed of our dicot will germinate.  The first visible sign, often, is a pair of small green leaves, completely distinct from mature leaves to follow.  This pair of leaves represents expanded cotyledons (seed leaves), which had formed inside the ovule as the embryo developed.  You’ll see the seed leaves easily in Composites and Mustards, but that isn’t always the case.  In some dicots, the cotyledons remain in the ground, and only a green shoot surfaces.

Whether they emerge or remain underground, cotyledons are usually important sites of energy storage, harboring reserves needed to support root and shoot development, sometimes constituting the great mass of a seed.  Regardless, there’s hardly anything more charming than emerging seedlings, with or without pairs of jewel-green seed leaves that typify Dicots. But when we focus on Monocots, things change.

Seedlings of grasses, lilies, and palms appear completely different because as Monocots the embryo generates a distinctive growth pattern that governs the future life of the plant.  Rather than a pair of lateral seed leaves, the developing growing tip of monocots produces a single encircling tubular leaf, the “scutellum,” which never surfaces. At germination you may notice the second sheathing leaf, the “coleoptile”, which is often thin and frail. Future leaves will follow the same pattern, emerging as a tube around the growing tip, sheathing it.  This is the moment Monocot seedlings become obvious, when grassy blades poke upward through the coleoptile.  These newer leaves Russian-doll their way from inside earlier leaves, each succeeding leaf emerging inside its predecessor – often with hardly any stem between nodes. 

How does this work out?  Let’s take a short break from germination and jump ahead in plant growth sequence.  We teach about plants using a model of leaf and node, spaced by a section of stem simply called the internode.  But that mental image should be paired with a nearly opposite growth form, production of new leaves by a growing tip with hardly any new length of stem.  Clustering leaves tightly means that plants may produce rosettes, or scales surrounding terminal buds, or bulbs.  Most plants alternate these patterns, reaching distances with well-developed internodes, or hunkering down in cluster formation.  Back to our monocot seedling, we now have a stem tip making new leaves one inside the other, each ringing the stem, setting the stage for growth forms particular to this group of plants. That means we might see well-spaced nodes in Canna, or compacted nodes that generate bulbs such as those of Crinum. It also explains cespitose habits of grasses.plants with tall aerial stems rising from tight clumps formed by short rhizomatous side branches, as well as the capacity of palms to remain calm while carrying on with enlarging apical meristems that can form massive trunks.

Though some seed, such as those of Legumes, invest practically all energy storage in embryonic cotyledons, many (most?) seed do not follow this pattern. Particular to flowering plants is a special tissue formed earlier than embryo development.  Initiated at the time of fertilization, seed formation in Angiosperms includes a unique tissue called endosperm, a result of double fertilization that characterizes all flowering plants.  We describe fertilization as “double” because a pollen tube delivers two functional nuclei to the egg sac; one fertilizes the egg while the other joins forces with other nuclei to generate a most curious triploid (to polyploid) mass of cells we call endosperm.  As the zygote develops into an embryo, endosperm grows to fill the rest of the seed, to some extent characteristic of each kind of plant.  Seed of many plants produce significant stores of endosperm.  In grasses, for example, the bulk of each seed is endosperm genetically different from the embryo (the germ).  Thus the edible grain of rice is endosperm, as is the pulverized wheat product we call flour. 

I used the term “seed” for rice, but Grasses and other herbaceous groups, notably Composites and Umbels, pack each single seed inside an indehiscent fruit.  What appears simply as a seed is structurally the entire carpel. Botanists, of course, aware this is different from the textbook representation will avoid the word “seed” and use special names for these propagules, insisting on “cypsela” (or achene) to describe the 1-seeded fruit of Asteraceae, “mericarp”  for each of the two 1-seeded carpels produced in the Apiaceae, and “caryopsis” (grain) for the 1-seeded fruit of grasses (Poaceae).  We encounter this unified structure of seed and fruit wall across the Flowering Plants, such as in oak acorns and maple samaras.

While many flowering plants generate seed that are structurally complex, plants of one monocot family, the orchids, produce seed with neither a developed embryo nor significant reserves of endosperm.  An orchid seed is that are incredibly reduced, such that a single capsule might shed thousands of spore-like seed, each consisting of a small mass of undifferentiated cells enveloped by a fragile seed coat.  Orchid seed have no significant storage resources.  Once dispersed, they will differentiate and germinate only through assistance of a proper fungus that makes contact and shuttles nutrients to the cell mass.

Germination:  Seed of many native plants are considered “orthodox” (read normal), while others may be “recalcitrant” or ‘intermediate”.  There’s no cultural inference here.  Orthodox seed behave the way horticulturists wish all seed do, remaining viable in dry storage conditions and surviving freezing as a storage method.  Recalcitrant seed are typical of many tropical plants in that storage is not easy.  In nature, recalcitrant seed germinate at maturity, perishing otherwise.  Intermediate seed can be stored in refrigerators for limited times.  Seed of the native Asimina tetramera are reported as recalcitrant.

Beyond this, many seed will require some period of dormancy, or perhaps special treatment (stratification), such as chilling, or soaking, or scarification (physically scoring the seed coat), replicating aspects of natural seasonality and exposure.   I’ve learned from personal trials that seed of local Asimina species require cold treatment to unlock germination.  

These particularities remind us that seed flourish when they encounter the most appropriate microhabitat.  Given the numerous hurdles to overcome for successful formation and germination, potential is lost if circumstances do not support establishment.  Flooding or untimely drought, sudden changes in exposure to light, disturbance and even herbivory take their toll.  But eventually, seedlings establish and the race is on to vegetative growth, flowering, and fruiting.  

Growth.  Vegetative growth takes as many paths as there are kinds of plants.  Thankfully, we find similarities.  For example:

  • Plants develop points of growth called meristems, which produce cells that form roots, stems, and leaves.  New root meristems form inside existing roots, but can also form along stems.  Stem growing tips make new stems as well as nodes where lateral buds and leaf “initials” or “primordia” form.  In special cases, older tissue can generate buds.
  • Though roots normally grow down into soil and shoots (stems and leaves) typically grow up,  we distinguish between them based on their origin, development, and internal anatomy.  Roots of epiphytes may orient in any direction; vines often generate roots along nodes that serve for attachment.  Stems of rhizomatous plants will tiller, or even sink lower in substrates.  You’ll find Lygodesmia stems a foot deep in sandy soils, though the seedling certainly established near the soil surface. Many wildflowers in our “fire-maintained savannas” are geophytes, not just bulb-forming monocots, but a range of dicots (Liatris, Centrosema, etc.) generate USOs (Underground Storage Organs) such as corms, tubers, and tuberous roots (Diaz-Toribio and Putz, 2022), preserving resources needed for seasonal regrowth as well as protecting dormant growing points..
  • Nutrient intake depends on many conditions, such as access to soil water, acidity, symbiotic activity.  Though we associate intake with roots, in Tillandsia, specialized leaf scales take in water and nutrients.  Bladderworts (Utricularia) create highly modified leaves (or bladders) that capture small prey (see Wikipedia, also Whitewoods, et al, 2020)
  • Stems take on many roles beyond production of leaves and flowers.  When green, they may constitute the primary photosynthetic surfaces, as in Sabatia decandra or Coreopsis nudata.  Stems may grow out as tendrils (as in Vitis), or thorns (Gleditsia) or tubers (Smilax). 
  • Stem nodes are exciting regions, where leaves are attached, branching occurs, and even roots may form.  Nodes are separated by internodes, relatively quiet sections of stem.  The node/internode complement becomes a basis for iterative (repeated) growth patterns that create plant architecture.  Most herbs will show patterns of repetition and branching for vegetative growth distinct from flowering.  The flower itself is a specialized stem.
  • Plants show characteristic phyllotaxis (leaf arrangement).  As growing tips generate new nodes, the orientation of leaf+bud shifts in predictable ways. Taxonomist use the term ranking, which summarizes how many leaf positions a given plant may spiral before forming a leaf in the same position.  Sometimes this is straightforward, as when leaf arrangement is “decussate” – forming opposite leaves that shift back and forth at right angles. We associate this kind of opposite leaf arrangement with Mints (Salvia, Conradina), Hypericums, and many other plant groups.  Composites, on the other hand, develop both opposite and alternate arrangement.  Leaf arrangement becomes basic to plant architecture, reflecting how leaves are spaced to interact with the environment and establishing orientation of buds that lead to branching. 
  • The fact that one plant species might be herbaceous, while a near relative in the same genus develops woody stems reminds us primary growth is the same for both.  Herbaceous plants shorten the cycle by flowering and fruiting before woody tissues develop.  We also note that monocots, with numerous scattered vascular bundles, are seldom woody, and even when they develop tough, long-lived stems (as in Palms), those tissues are either all primary growth, or in some tree-forms (Yuccas, for example), “anomalous,” producing added layers that replicate the scattered veins observed in standard monocot stems. Monocot stem construction reminds me of buildings made with concrete and rebar. After a big storm, you’ll find few broken palms; their internal anatomy combines strength with flexibility not possible for convential trees with ring growth.
  • Plants produce hormones, chemicals that signal changes in growth patterns and maturation processes.  Hormones function at incredibly low concentrations in diffuse sites, promote complex cascades of change that may initiate or suppress changes in growth and function.  Consider Auxin, which is produced in growth tips but regulates growth of axillary buds further down the stem, thus directing branching patterns.  But Auxin has multiple functions – involved leaf in formation, regulating emergence of lateral roots, and promoting growth in shaded zones that orients plants relative to light sources.  Browsing by animals (as well as pruning or damaging stems) engenders distinctive growth responses, disrupting existing hormonal control mechanisms.  Synthetic analogs to Auxin, such as NAA, promote root formation in cuttings while others, such as 2,4-D  kill vegetation.
  • Leaves and stems show specialized anatomies supporting photosynthesis and epidermal characters that modulate water loss and exchange of gases (stomata).  Plant chemistry and surface features (such as waxes or hairs) protect tissues from the elements as well as well as predators.    
  • Leaves act as filters, capturing, transmitting, and reflecting light based on pigments present.  Typically green, leaves with abundant chlorophylls capture energy of red and blue wavelengths for conversion to energy-rich chemical bonds.  But cells will include other pigments such as carotenes, betalains, and flavonoids (anthocyanins, flavones and flavanols….), which most obviously impact flower coloration and the residual color of dying leaves. A secondary effect is that quality of light (color balance) under a canopy will be different than the color balance experienced in open sunlight.
  • Leaves develop differentially in response to differing conditions, such as full sun versus shade, changes in season, and water conditions. 
  • Though leaf blades do not typically include buds or give rise to stems, under some circumstances leaf tissue may generate roots and shoots. This reminds us of “totipotency”, the capacity of undifferentiated cells to take on new roles, the basis for the modern tissue culture industry.  Though neither native to Florida nor technically a wildflower, the Appalachian walking ferns show “leaf proliferation”, generating new plantlets at leaf-tip veins.  The non-native Mother-of-hundreds (Kalanchoe), common in Florida landscapes, is noted for generating plantlets where veins terminated along leaf margins.
  • Plants have evolved crucial mechanisms that respond to environmental cues.  Quality (color balance) and duration of sunlight regulate plant growth and development in varying ways.  Many plants break dormancy or initiate flowering based on chemical perception of daylength.  Other environmental stimuli include changes in temperature, patterns of rainfall, and even chemical stimuli.

Water Relations:  

Life is cellular and living cells are membrane-bound drops of liquid, inside which molecular robots use chemical templates to manage inputs and exudates, assemble compounds, build useful internal structure to maintain and even replicate the cell, recycle waste, maintain cell osmotic balance and turgor (internal pressure) and even prepare cells for death.  In addition to water that constitutes the greatest mass of cells, nutrients flow in a continuum of water through whole plants, sucrose and other dissolved sugars are translocated in water, water maintains cell connectivity, and water transpiring from surfaces helps cool plants.  Once imported by the roots, water is managed until the moment it is utilized, transpired through pores called stomata or lost through damage. 

Plant anatomy evolved in response to water management.  Though texts describe water intake as passive, it’s not so simple.  Roots actively extend their reach, growing continually to mine water and nutrients.  Internal anatomy concentrates nutrients, and ensures positive flow of water toward the stem, streaming both to above-ground tissues.  Water makes a continuum which can be surprisingly tenuous.  Anyone who has ever uprooted a soft herb will know that wilting can be immediate, even irreparable.  In order to keep a stem alive and fresh, you’ll want to have a deep vessel of water available, something into which the plant can be plunged deeply., or you’ll need to reduce leaf area to lessen water loss through transpiration.  I find some herbs only maintain turgor when the entire plant is quickly inundated.

“Turgor” is positive pressure plants require to keep individual cells intact and functional  Thus the water continuum from root tip to leaf tip is critical.  Mature, living plant cells exist in the water-rich context of surrounding tissues, maintaining their viability and shape based on internal pressure generated by water-filled, balloon-like cellular structures called vacuoles, structurally membrane-bound vesicles that retain enough water to plump up the cell.  We observe any loss of internal water and pressure as wilting, which results from loss of vacuolar pressure called turgidity.  

That nutrients are more concentrated in plant roots as compared to presence in surrounding soil, tells us living cells are at work.  That means they are alive, and life requires inputs of energy necessary to drive reactions, maintain membranes, and build cell components.  Plant sugars manufactured in photosynthetic tissues are translocated to roots, and broken down in a process called respiration.  That is a confusing term, since people often use “respiration” as synonymous to “breathing.” 

But what biologists call respiration is a complex process requiring oxygen. Thus root growth and health depend on import of sugars from stems and leaves as well as access to oxygen.  I would say roots (in fact all underground plant parts) must breathe, if I could avoid any comparisons to animal biology, but it’s better to say that oxygen must be available to living cells.  A sure way to kill most houseplants is to set them in cache pots that fill with water.  Waterlogged soils can be anoxic (lacking oxygen), which is deadly unless plants have adaptations that meet the challenges of a waterlogged environment.  Given Florida habitats, many native plants show adaptations to wet soils.  In fact, many of our most charismatic wildflowers (Sabatia decandra, Coreopsis nudata, Polygala cymosa, Asclepias lanceolata, Lilium catesbaei, Zigadenus glaberrimus, etc.) flourish in wet soils, their very presence at a site alerting us to water-logged conditions.  

Nutrient Acquisition:  In addition to maintaining soil-water relations and anchoring the plant, roots generally are the sites for nutrient uptake, which is my invitation to address the nature of plant “nutrients” as well as an opportunity to share my own prejudices when it comes to loose talk regarding plant nutrition.  To date, physiologists and agronomists have identified 22 elements plants require or utilize at some level.  I remember these using a mnemonic:  “See Hopkins Cafe, managed by my cynical cousin Mona Alcose”, which loosely translates as: C = Carbon, H = Hydrogen, O = Oxygen, P = Phosphorous, K = Potassium, Ca = Calcium, Fe = Iron, Mg = Magnesium, B = Boron, Mn = Manganese, Si = Silicon, Ni = Nickel, Cl = Chlorine, Cu = Copper, Zn = Zinc, Mo = Molydenum, Na = Sodium, Al = Aluminum (very rare, as Al is mostly toxic), Co = Cobalt, Se = Selenium.  Seventeen are considered essential.  Five (Aluminum, Cobalt, Sodium, Silicon, and Sodium) are a bit suss, often referred to as “beneficial” in that some plants may benefit from them, but they are not common to all plants.  We know, for example, that Aluminum is generally toxic to biological systems.

A convenient aspect of this memory device is that “C HOPKNS” introduces the big hitters: Carbon, Hydrogen, and Oxygen, followed by Phosphorous, Potassium, and Nitrogen.  The first three, available in air and water, constitute the great mass of living tissue (96% of dry weight).  The following three elements are among the most common in plants, accounting for about 3% dry weight, but are also the most frequently deficient in soils, which is the reason they have become the basis for commercial fertilizer formulations.  You’ll run into this as as the N:P:K ratio, a specified ratio of available Nitrogen-to-Phosphorous-to-Potassium by weight per 100 lbs of mix.

Photosynthesis and all that Jazz.  The great bulk of plant dry weight (the residual dry material once water has been driven off through heating) is carbohydrate  – sugars, cellulose, starches, compounds made purely of carbon, hydrogen, and oxygen.  Plants are the world’s dealers in carbohydrates, manufacturing them from air and water through photosynthesis, storing the sun’s energy in chemical bonds.  Cellulose holds the golden coconut as the most significant polysaccharide (made of many sugars) in plant tissue. As the principal component of plant cell walls, we are told that cellulose is the world’s most abundant macromolecule.  In fact, it is the world’s most abundant carbohydrate; fully half of all organic carbon on earth is bound up as cellulose.  That build-up tells us cellulose is surprisingly inert, it’s not something plants recycle.  It takes the diligent efforts of special bacteria and fungi to dismantle cellulose.

However, plants make quantities of interconvertible compounds, carbohydrate starches as well as various energy-rich lipids and dimensional proteins that, while functionally-useful, serve both as energy reserves and components to be utilized or retooled.  These are the ingredients of life.  From raw elements, powered by the sun’s energy, plants manufacture every component they need.  And this is the reason I bristle at horticultural marketing terms such as “plant food” and “vitamins”.  Fertilizers are not plant food, they are elemental nutrients.  If you want to claim anything as plant food, it might be air, which is the source of carbon dioxide.  Or perhaps it’s more correct to note that plants make their own food through manufacturing sugars.  While they are at it, plants manufacture compounds that are vitamins for humans.  Vitamins are organic compounds we require in our diets, but plants don’t have diets.  Plants don’t need food.  Plants are food.  Botanists term them autotrophs (self-feeders).

This is the promise of Photosynthesis, the great miracle that changed the nature of Earth, generating enough free oxygen to account for 20% of our atmosphere while creating food reserves sufficient to support the great diversity of life that has evolved and persisted over hundreds of millions of years.  Capturing energy from the sun, photosynthetic systems build order.  Carbon dioxide is taken from the atmosphere and bonded to form carbohydrate chains we call sugars.  Chemists call this a “redox” reaction.  That’s chemical give and take.  Carbon dioxide molecules are “reduced” in being bound together while water is “oxidized” – losing its hydrogen to become free oxygen.  It’s a tradeoff; one compound loses energy, the other gains. There is no free ride.

The summary of Photosynthesis you see most often (6CO2 + 6H2O + energy = C6H12O6 + 6O2) sounds simple, suggesting carbon dioxide and water combine to generate sugars and free oxygen.  In reality photosynthesis is best imagined as a two-step operation.  Light energy is captured chemically, which involves breaking water to harvest electrons, which means oxygen is released to the atmosphere.  This multi-step process is called the Light Reactions.  In a  separate series of reactions, which can occur in light or darkness, the captured energy is invested in bonds that build glucose by assimilating (also called “fixing”) atmospheric carbon dioxide.  We say “fixing” because something that was a free component of air is now tied up (i.e. assimilated) in solids as an organic compound.  You’ll see the same word used when atmospheric Nitrogen is “fixed” as a compound now available for biological use.  Both Nitrogen and Carbon dioxide are otherwise inert. 

There are rapid and incremental ways to reverse the work of photosynthesis.  Take a log that’s 50% cellulose and burn it (fire requires oxygen).  The residual ash has a fraction of the energy that was invested in the log during growth of the tree, that energy being lost to the heat and light of a conflagration, which generated carbon dioxide and water vapor as by-products.

A living system harvests that energy incrementally, through dismantling carbohydrates via the highly controlled process of Respiration, taking in oxygen and releasing carbon dioxide and water piecemeal.  Unlike fire, which releases light, heat, carbon dioxide, and water vapor in rapid combustion, respiration harvests the chemical energy in special compounds that function as common currency, paying the piper to drive life processes.  So I guess you might think of Respiration as the controlled cousin of Conflagration, the drama queen.  From this perspective, we are (as respiring animals) living flames.  That doesn’t mean I believe in spontaneous human combustion, though I’ve certainly seen plenty of headlines promoting this at grocery store magazine racks.

With nutrient acquisition mostly a task of roots, which require sugars that are the product of photosynthesis, we have to give some thought to how these materials move from one place to another, basically from “source” to “sink”.  They are not pumped, there’s no heart or smooth muscle to circulate materials.  There is plant sap moving from roots to stems and leaves in tissues called xylem, and we can measure photosynthate which is “translocated” through conduits called phloem.  Neither xylem nor phloem should be compared structurally to our veins and arteries.  Plant conductive tissues function at a cellular level, forming microscopic tubular networks in dead cells of xylem and networks of living cells that constitute phloem.  Water and nutrients move through the plant in the hollow dead cells of xylem, as well as interstitial (between cells) regions. Sugars (typically as sucrose) are actively “translocated” through living phloem cells, an energy-dependent process. 

Along the way, watch out for casual language.  You’ll hear botanists use a three-letter word “sap” in several ways, but know those liquids are distinct.  Sap from the roots moves in xylem, and is predominantly water and nutrients.  Sap in phloem is dissolved sucrose, better termed “assimilate”.  And you’ll hear vacuolar contents called “cell sap”, which is water with a wide range of solutes and secondary compounds.

Flowering:  At some basically predetermined stage of development, plant growing tips shift gears, producing flowers, usually in branching systems called inflorescences.  This stage in a plant life cycle is more than the culmination of an epoch of growth, and certainly more than a heralding of fruit production.  I say this because for hundreds of years people believed flowers were a prideful way for plants to let us know that fruit would follow. It wasn’t until experiments with monoecious plants (Mulberry, Spinach, Castor Bean) published in 1694 by German scholar Rudolf Jacob Camerarius, who demonstrated pollen was necessary for fruit and seed production. Of course a generation of scholarship was dedicated in an attempt to prove the male component was the vital force while the female was merely the nurturing vessel. But we finally figured that out.

Today we understand a flower is the apparatus for sexual reproduction, which means this is the mechanism through which genetic traits are re-sorted as a new seed crop.  A lot happens in a flower.

Genetics, initially a simple concept as worked out by Mendel in his pea hybrids, has emerged as  a masterfully-complex science.  Keeping to the simplest summary, I think of characters (traits) in a colony of plants like colored marbles in a grab bag, each differently colored marble being equivalent to what a biologist calls an allele (which means “the reciprocal other”).  Players draw two marbles.  After each bout, the marbles are restored to the pot and remain available for the next game.  If one player selfishly grabs a pair of marbles and goes to join another franchise, whatever that player took is lost to the local colony.  That’s what happens when plants self-pollinate exclusively, they are taking their marbles and going home.  It also happens accidentally, when marbles are culled, lost, or destroyed.  Only marbles returned to the pot, restored to what we call the “gene pool”, are available for new combinations.  While selfing and selective pressures deplete the variety of marbles in the pot, restoring marbles and introducing new ones maintains and augments variety.  New colors (alleles) are added when someone joins the game from outside, bringing alleles from another colony, or when an existing marble changes color, which we call mutation.  Flowers are the way plants keep their marbles back together, promoting recycling of alleles to the gene pool through out-crossing.  

 Existence of flowers tells us out-crossing is crucial to sexual reproduction; lacking selective pressure for outcrossing, there’d be no reason for flowers.  This is where the marbles are recombined and re-sorted, keeping or culling the genetic variation available to the population.

Evolutionarily, we focus on new genetic combinations that will impact the future of individuals, their reproductive future (their “success” in contributing to a new generation of seed). So fresh pairings are fodder for natural selection. The sperm nucleus in a pollen grain is, basically, a bundle of genes, and it’s a statement. I’m bringing these marbles to the party. As pollen moves within a population, or to a new population, a next generation moves both physically and genetically, ever so slightly or perhaps great distances.

Flowering leads to fruit production, which is even more impactful. Dispersal of seed is the primary mechanism through which populations spread and new populations establish, not simply over distances but over time.  Orthodox seed may persist several years in soils (or even in dry fruit attached to the parent plant), becoming components of a seed pool that might cover time gaps when an area has been shaded out with canopy or overgrowth.  This potential allows resurgence in populations after fire or other disturbance, like some kind of natural time capsule.

Together, movement of pollen, success or failure of fertilization, generation of viable seed, and dispersal of seed account for change in population. It’s brutally wrote; there’s no emotion involved. Reduction in size, or too much inbreeding will cull variation.  Receiving new alleles (new marbles, i.e. genetic variation) from outside the population might spell change in success. Establishing a new foothold somewhere outside existing populations is especially interesting in establishing fresh frontiers. Like any frontier settlement, success will depend on the nature of the settlers. This is one path for evolution of populations that show different characteristics, because the founders brought their own, distinct subset of genetic material (population biologists call this “founder effect.”)  

Production of seed has other ecological implications, particularly related to productivity of systems.  As natural stores of energy-rich oils, carbohydrates, and proteins, compounds already produced and packaged for dormancy, seed are natural food supplies, useful even for hoarding.  But like shrinkage in a big box store, this loss is baked into the system.  Plant populations survive only when excess production accounts for animal consumption (which we used to call herbivory but now classify as predation).  And of course, this gave rise to many co-evolved systems in which animal consumption of fruit and seed is the primary dispersal mechanism, to the point that some seed require scarification that comes from passing through the gut of a predator. 

Turning our attention back to flowering, we discover development is more complex than textbooks can bear to admit.  We learn a simple model, with 4 basic flower parts – sepals, petals , stamens, and pistils.  Each different group of plants evolved modifications to that basic plan, which then varies from one species to another.  You likely learned to generalize differences in flowers by whether they are complete (having all four basic parts) vs. incomplete, or perfect (having both anthers and pistils) vs. imperfect.  And we even map out floral structure using formulae, to varying degrees of success.  But the world of flowers is not so simple.

Even so, there are generalizations that help us understand the structure and nature of flowers:

  • Flowering plants seem to have a common ancestor, which means botanists continue to interpret flowers as modifications of a basic plan that explains a flower as a stem tip (the receptacle) that produces a series of appendages (equivalent to leaves) in a predictable sequence.  Below the flower (we say “subtending”) there might be nodes with bracts, large or small.  these bracts are considered separate from flower segments because they are more leaflike, and are accompanied by axillary buds.  None of the flower segments bear axillary buds.
  • The flower proper then comprises spirals or whorls of increasingly specialized parts.  Outer, non-fertile segments constitute the perianth, which includes sepals and petals.  In flowers with spirally-arranged segments, distinguishing sepals from the more interior petals usually proves difficult.  You’ll find botanists a bit uncomfortable when asked where sepals end and petals begin in a Magnolia grandiflora flower.  We’ll be resorting to terms like “perigone” or “tepals” rather than clearly identifying sepals versus petals.
  • Plants producing flower segments in whorls have more distinct parts, producing outer segments clearly distinguishable as sepals, each of which will have 3 vascular traces (veins).  Sepals may have arrangement (phyllotaxy) similar to that of vegetative leaves, in which case  you’ll observe a distinct interruption between arrangement of sepals and petals.  Sepals are visible in buds, often green, even leafy, and normally protective.  But they can mature as petal-like, in texture and coloration, such that in some groups, the sepals are the showiest element.  The “wings” of Polygala (Senega) are sepals, as are the floral tubes of Nyctaginaceous plants like Mirabilis and Boerhavia.  Sepals may be reduced, as in Asteraceae (and Caprifoliaceae), where they form bristles or scales.  The collective term for sepals is the calyx.  In some plants, sepals continue to develop during fruit formation, which is what we see in Physalis. They frequently grow to a new position, as with 4-petalled Hypericum.
  • The next inner whorl (or whorls) of the perianth is normally the petals, each segment being served by a single vein – a characteristic marking petals as anatomically distinct from sepals.  Petals will often number the same as sepals, usually alternating in position relative to sepals.  A second whorl of petals will double the number as compared to sepals, alternating in position relative to the first whorl of petals.  Petal color and structure correlates to the pollination system.  But it’s easy to be deceived.  Botanists interpret many plant groups as lacking petals (apetaly), developing petal-like sepals  As mentioned above, the “petals” of Mirabilis and other plants in the Nyctaginaceae are sepals.  
  • Further up the receptacle (more interior), a flower may include a whorl (or two) of stamens.  Many flowers show great numbers of stamens, based usually on a multiplication of stamen primordia during development.  Compare flowers of Hypericum gentionoides (or Hypericum mutilum), which have five stamens, to most of the other Hypericums in our flora, which have multiple stamens (30-160), grouped loosely in fascicles that reveal their multiplication from an a basic whorl of five.  The collective term for anthers is the androecium.
  • Borne at the tip of the receptacle, flowers may develop one or more carpels (fertile leaves).  Those carpels may be distinct and spirally-arranged, as we see in Magnolia. The collective term for carpels is gynoecium.
  • Flowers with parts in distinct whorls are more likely to show fusion and specialization.  Tubular corollas of Rubiaceae with adnate stamens are among the many examples, most of which are Asterids (the clade that includes Asteraceae, Lamiaceae, etc. Cucurbits being a rare exception).  The androphore of Hibiscus, the specialized corona of Passiflora, the multicarpellate pistil we see in Vaccinium and Gaylussacia are possible because floral parts exist in whorls.  You’ll not observe so many specializations in flowers with spiral phylllotaxy, such as Magnolia, Illicium, and Calycanthus.
  • Flowers show many structural adaptations that promote outcrossing.  At one extreme we find dioecious plants, with male flowers borne on separate individuals than female flowers.  Many natives (e.g. Myrica, Ceratiola, Baccharis, Ilex, Diospyros, Smilax, etc) are predominantly dioecious.  Monoecy is common also, present in many herbs, such as Sagittaria, which produces female flowers at the base of flowering stems and male flowers apically.  The reverse is true for Cucurbits, which tend to produce male flowers first, followed by female flowers later in development.  Many grasses are monoecious,, segregating male and female flowers in ways that improve efficiency of wind pollination.
  • Beyond strategies of separating sexes, hermaphroditic (flowers are perfect, with both male and female elements represented) plants promote outcrossing through shifting placement of anthers as compared to stigmas,. In out native wildflowers, Lythrum is among the many heterostylous plants, with what is sometimes called pin and thrum morphology.  Plants in a population will show differing combinations of anther and style length that tends to enforce outcrossing. Temporal separation is common with differential maturation times when pollen is available at one moment while stigmas may be receptive before or after.  Of course we have terms for this – protandrous flowers mature stamens before pistils are receptive, protogynous flowers mature pistils in advance of anther maturity.  
  • Presence, orientation, and structure of flower parts(herkogamy) relates to pollination system.  Commonalities have emerged that allow us to predict the kind of vector that will support the successful pollination, usually referred to as pollination syndromes.  Some crepuscular flowers, such as the native Canna and Ruellia noctiflora, open in the evening and close in the heat of the following day.  Some flowers are open a single day only; others may remain open and fresh for several days.  Many flowers show nyctinasty, opening and closing daily, which makes them difficult to spot early or late in the day.  Some perianth segments are reduced or absent in wind-pollinated plants, notably native grasses, but also in wind pollinated trees, such as Oaks.
  • Some plant groups, Asteraceae notably, have evolved self-incompatability mechanisms that preclude fertilization or reduce viability when self-pollinated.  Evans, et al (2004) report obligate outcrossing for Liatris ohlingerae.
  • “Flowers are seldom solitary.  They are mostly grouped in inflorescences, which can be highly complex.  Several attempts have been made to morphologically interpret inflorescences, mostly with emphasis on describing inflorescences typologically, and more rarely developmentally. However, the complexity of the terminology can be overwhelming.  We’ll cover some of these particulars in Coming to Terms….
  • Ronse (2010) and other authors remind us of the complexities of “pseudanthia,” inflorescences in which clustered flowers take on the presence of a single flower.  Dogwood is a good example, as are cyathia of Euphorbias.  The most ever-present examples come from the largest family of flowering plants, the Asteraceae, in which heads of flowers (capitulae) “function as pseudoflowers.” 

Down the Rabbit Hole:  The section that follows provides more (perhaps too much) detail regarding the biology of flowers.  This story is not crucial to appreciation of plants, but given the focus on “wildflowers” I realized there needs to be a more complete presentation as to what a flower is and how a flower functions to produce seed.  Read along, or skip this.

Pollination is Movement:  Of course a flower is about seed production, which, as a process is a bit of a fantasy.  We’ve already walked through some of the staging involved.  Each kind of plant makes it’s particular performance stage, a flower, or in many kinds of plants two or more different sorts of flowers.  

Flowers will be outfitted so as to produce pollen grains destined to arrive, somehow, in the only place where success is possible, a receptive stigma.  That’s a small target, ranging from smaller than a pin-head to as large as a thumbtack, or in grasses and other wind-pollinated flowers, running along the filamentous margin of delicate airy styles.  Success for a grain of pollen involves an obstacle course that begins with the pollination phase, movement of pollen from an anther to a receptive stigma. Only a few grains will make it to the right stigmatic platform.  The vast majority will wash out.

I say the right stigma because the pollen grain, basically a microscopic plant formed of two or three cells, only makes it through the next phase of obstacles when it lands in a place suitable for germination and growth.  This grain, which is among the tiniest athletes of plants, is also a free-loader, dependent on some vector for aerial transport, and now completely dependent on nourishment provisioned by the carpel (pistil).  Following transport, the new phase of the trial involves germination.  The grain itself, becomes a husk, stuck on the stigmatic surface, and through a system repeated over millions of years, the sperm nucleus must arrive at the egg, which is buried in an ovule on the placenta of a carpel.  The sperm nucleus relies on a tube to extend its reach, a tubular extension of the pollen grain cell that makes me think of one of those long, twisting balloons. But it’s so narrow and delicate as to be practically invisble, a tenth the diameter of a human hair – but this fragile tube relentlessly pushes its growing tip through tissues of the style toward the egg sac, provisioned by local sugars and guided by a chemical gradient. 

As a tiny, dependent plant, the pollen grain is genetically haploid, having a single complement (set) of chromosomes with only two or three nuclei.  Completion of the obstacle course comes when the pollen tube delivers two haploid nuclei to the egg sac.  One of those is the sperm nucleus, the male gamete, which is the reason we call this tiny plant (this pollen grain) a gametophyte (i.e., the gamete-producer).  As the bearer of the sperm nucleus, this gametophyte (pollen grain), with its load exemplifies maleness – dissemination of countless, profligate, highly mobile, indiscriminate, impoverished nuclei, bringing little to the party beyond a set of chromosomes.  

But I’m getting ahead of the story.  A significant goal in this discussion is to make certain I’ve presented pollination as distinct from fertilization.  So we must back up.  Flowers are the unique outposts where plants trade chromosomes to secure sexual out-crossing.  They are the stage, the platform.  Pollination is the performance, the handoff, the process of relocating pollen from source (an anther) to destination (a stigma) usually requires some active outside force.  But for particular instances in which plants self-pollinate by pressing an anther into a stigma, movement of pollen depends on a “vector”  such as wind or water or any number of insects, birds, reptiles, and mammals.  

The systems of pollen movement are driving forces in the story of flowers, generating adaptions that led to evolution of myriad delightful flower forms.  Those flowers…, well they turn out to be quite an investment, costly to make in terms of energy and resources, expensive to maintain and replace.  So why bother?  What’s the benefit to the many flower forms and pollination systems we observe?  The answer is maintenance of genetic diversity, which is lost with inbreeding.  Out-crossing requires movement and mixing, which pollination mechanisms promote.  Lessening self-pollination maintains genetic diversity in a population by continually sharing alleles (different forms of the same gene), thus keeping the skill set fleshed out.  Genetic diversity is the crucial investment a population makes in resilience and adaptability .  Developing efficient mechanisms that solve the pollination obstacle course confers reproductive advantages.

Flower form relates directly to the pollination mechanisms.  Botanical literature is rife with observations and explanations by pollination ecologists who’ve spent countless hours of field study and as many dedicated to lab examination in their efforts to explain the resources a plant invests in flower production (considered “costs”) and the “mechanisms” that ensure pollen grains arrive at their minuscule stigmatic targets.  Simplifying (perhaps over-simpliyfing) the many complex mechanisms, we are able to group flowers by various syndromes, suites of characteristics that allow some prediction about a plant’s effective pollinating system.  All systems involve trade-offs.  Wind pollination is common because air movement is common.  But wind pollination involves a heavy investment in pollen production, so flowers of plants with wind pollination are prodigious pollen parents and excellent pollen capture capacities.  Flowers of wind pollinated plants (which includes almost all grasses and many temperate trees) typically lack showy sepals and petals, but sport freely-moving anthers coupled with elongated, frequently branched stigmatic surfaces.  Flowers of native grasses are typically wind pollinated.

Flowers with showier perianth segments suggest more active pollinators, as we see in co-evolved relationships between many kinds of insects and flowering plants.  Simply imagine this as a natural quid pro quo.  Plants as primary produces have important resources to offer, sugar-rich nectar, lipid-rich pollen, even oils and other organics as well as surfaces for interaction.  Flower visitors mine those resources, unwittingly transferring pollen.  Some mechanisms are very general and messy, though still more efficient that wind pollination, while others are found to be astonishingly precise.  Characteristic behaviors of the various kinds of pollinators (insects, as well as certain birds and a few mammals and lizards) provide the key to understanding how flowers and animals have evolved over millions of years.   We come to appreciate flower color, form, and activity through syndromes – suites of characteristics that reflect those evolved working relationships. 

Understanding the various systems and syndromes adds value in the study of Florida’s wildflowers, and opportunities for citizen scientists., but there are limitations as to what we might presume.  Without field study, we can’t assume that every insect visiting a flower is a pollinator.  Flowers might be good places to hangout, and catch a meal, but that doesn’t mean the visitor fulfills it’s role in this relationship.  So an offering of resources invites cheating.  Visitors will harvest nectar without transferring pollen to a stigma, but many bees actually rob flowers, piercing into nectaries and defying the system, or taking pollen without contacting a stigma.  And there is some danger, predators abound, especially the green lynx spiders often seen savaging butterflies on Liatris and other inflorescences. There’s much that could be known about natural systems, more than will ever gain the attention of grad students and professional researchers. Lay naturalists, outfitted with basic appreciation of pollination systems, can make and report useful observations that would reinforce or revamp understanding and assumptions.  

A good starting point with insect-pollinated flowers is with Composites. which host an open system.  Observing a population of practically any composite you’ll encounter a range of visitors, from bees to butterflies.  We know Composite flowers are relatively simple and similar in rewards offered.  An inventory conducted by Bennington and May (2020) compared two common wildflowers, the Composite Bidens alba and the Legume Chamaecrista fasciculata.    In their study, xxxx different visitors were noted at xxxxxxxx.  Given the spectrum of visitors, one has to question how they avoid self-pollination.  It seems, in general, they don’t.  The real issue then, is how Composites avoid inbreeding.  The answer seems to be that most Composites practice self-incompatibility.  Though flowers receive pollen from the same plant, fertilization does not follow.  Only pollen from a different plant will successfully deliver a sperm nucleus to an egg.  Assumptions such as this are based on historical studies, but there’s real opportunity for detailed work by citizen scientists who could run simple exclusion trials, comparing outcrossed to self-pollinated Composites.  

Plants with open pollination systems have evolved other, more physical strategies to promote outcrossing, the simplest being dioecy.  Many common native woody plants (Ilex, Ceratiola, Myrica, Baccharis), vines (Smilax), and even wildflowers (October Flower, Polygonum polygamum, Pineland Croton, Croton linearis) avoid inbreeding through isolating pollen-bearing flowers and egg-bearing flowers to different individuals.  Monoecy (producing separate male and female flowers on the same plant) is incredibly common among wildflowers.  As was discussed earlier, Sagittaria and Typha are obvious examples.

Some syndromes are more tailored, and reasonably well-known.  Both wind pollination and open systems are costly, requiring loads of pollen sufficient to offset loss to the winds and incompatible flowers, or consumed by bees.  Many legumes develop specialized structure for managing bees. In Partridge Pea, the flower is asymmetrical, with a cucullate (twisted and cupped) petal that guides visiting bees toward the shock of stamens enveloped by two lower petals.  A similar asymmetry operates for Strophostyles, but in this case a visiting bee exposes the anthers while forcing entry past corkscrewing styles.  

Mints and Scrophs are known for tubular, zygomorphic (bilaterally symmetrical) flowers that provide nectar while positioning pollen above the visitor.  These flowers, often in shades of blue, invite bees to enter for nectar, showing some ingenious mechanisms for depositing pollen atop an insect head and thorax.  Our common Scutellarias show that structure.  Out-crossing is partially enforced by timing differentials in maturity and presentation of anthers and stigmas.

Flowers that develop long, narrowly-tubular nectaries (spurs) are often associated with butterfly and moth pollination, insects that can unfurl a relatively long, narrow, and flexible proboscis.  Here we observe distinctions based on activity.  Butterflies land while feeding, thus the spur (nectary) of a flower of Chapman’s Orchid, Platanthera chapmanii, or the Yellow-fringed Orchid, P. ciliaris, is perfect for butterfly visitation, while fringing along the lip is well-situated as a landing platform.  Moths, many which hover while feeding, come in different flavors; during daytimes and crepuscular hours you might encounter the wonderful Hummingbird Moths, visiting native Rhododendrons for example.  I associate Sphingid Moths with crepuscular and nocturnal activity and a moth syndrome, in which flowers are free of obstructions that would block a hovering insect. Flowers showing this syndrome are often white, or at least pale green, and frequently emanate strong fragrance, especially in the relative coolness of evenings.  A perfect example is the White Fringed to Fringeless Orchid, Platanthera blephariglottis.  To a great extent this flower is a white version of the butterfly-pollinated P. ciliaris, white in color with a longer nectary.  But the main pollinator is a moth. Another would be Florida’s Ghost Orchid, Dendrophylax lindenii (Houlihan, et al, 2019). But you can study flower characteristics and hypothesize moth pollination. Plants with the corresponding suite of features include species of Crinum and Hymenocallis, Canna flaccida, Ruellia noctiflora, Gentiana pennelliana, Ipomoea alba, the weedy Jimsonweed, Datura stramonium and many others. One can’t assume moth pollination, however, without field study.

Even then, these pollination systems are neither absolute nor exclusive.  Natural hybrids between Platanthera ciliaris and P. blephariglottis are well-documented wherever populations overlap, suggesting the butterflies and moths do not follow the rules.  Simply because a flower is white and tubular, predicting moth-pollination is confirmed only through observation.  There are a lot of white flowers out there, and a lot of potential visitors searching for the next calorie.

Around the world, one encounters various examples of bird pollination, but in the US, it’s all about hummingbirds and the suite of floral characters that typify the Hummingbird Pollination Syndrome. Though several hummingbird species have been reported by rare Florida sitings, the Ruby-throated Hummingbird is a seasonal resident throughout Florida and much of the Eastern US.  Annual arrival of Ruby-throated Hummers is presaged by blooming of many plants with tubular, red, nectar-rich flowers, such as the shrubby Aesculus pavia, Erythrina herbacea, and Hibiscus coccineus, vining Campsis radicans, Bignonia capreolata, and Lonicera sempervirens, as well as a host of wildflowers – Lilium catesbaei, Lobelia cardinalis, Calamintha coccinea, Macranthera flammea, Spigelia marilandica, and Salvia coccinea. Once again, you’d want to see honest field studies documenting pollination systems to move from speculation plants (that show this syndrome are truly Hummingbird pollinated) to certainty. Moreover, Hummingbirds visit many kinds of flowers that are not red or orange, and bright red flowers are not a certain predictor the plant is hummingbird-pollinated. 

Fertilization leads to Fruit Formation:  Anthers produce and shed pollen; vectors move pollen; but carpels do pretty much everything else.  The carpel is a fertile leaf formed in a flower, often termed the pistil.  A flower may have one carpel, or many.  Carpels may be separate or connate (grown together as a single structure).  Depending on the kind of plant, you can usually identify three structural components to a carpel – the ovary (a chamber where ovules form along a placenta), the stigma, a tip that receives pollen, and an almost imperceptible to very exaggerated connecting stalk, the style.  As a student, I was gifted a memory device  for this: “though the pistil may have a stigma, it does have style.”

The carpel provides the stigma, a nurturing landing space for pollen capture and germination, one or more ovules, each with its egg, and a nourishing pathway to support and direct growth of a pollen tube from stigma to an ovule.  We find the stigma services its parent carpel, even when several carpels amalgamate to form a single pistil with united stigmas.  The lobes of the stigma usually will reflect the individual carpels involved, and pollen deposited on one lobe is typically directed to ovules in the correlating carpel.  This means when pollen is confined to a single lobe of a multi-carpellate ovary, you might find fertile seed in only one segment of the mature fruit..  

In preparation for fertilization, each developing ovule generates a single cell that forms an egg sac, made of just a few cells.  One of those cells will be the female gamete, the egg.  We label this gamete as female because it’s a completely competent cell, bringing a dowry of thousands of components that maintain life and serve as templates for future cell division.  That includes mitochondria and plastids, as well as the whole internal framework, including endoplasmic reticulum, the microtubular lattice, etc. These components that certify the egg as a fully competent cell perpetuate one thread in the 3+billion year continuum of life.   Recall Virchow’s 1855 cell theory: “omnis cellula e cellula.” – cells only form from existing cells.

Yes, like the sperm nucleus, the egg cell nucleus bears a single set of chromosomes, but chromosomes are simple instructions; they are not alive.  Life is a process, a unique multi-dimensional circus playing out in functioning intact cells.  Plant cells depend on that complex internal rigging of membranes and microtubules, and they require mitochondria for energy processing, as well as plastids that can become chloroplasts.  Both mitochondria and plastids are self-replicating ,independent contractors with their own DNA. 

The starter set of these components as well as many other molecules that will be required by cells of the new plant, is provided, almost exclusively, by the egg cell.  You will hear the phrase “maternal inheritance” applied to plants because mitochondria and plastids in the female egg cell carry memory separate from the nuclear DNA bound up in chromosomes, ensuring cell structure and many cell processes are of exclusively maternal lineage.  It’s like having a complex, self-replicating and multi-capable machine or robot that enters a new generation when it receives fresh instructions, fresh software, fresh nuclear DNA.  All of these self-replicating robots (cells) trace their ancestry to the first ones assembled. It’s the software that changes.

“Egg” therefore, across the plant and animal kingdoms, signifies the carrier of cellular life.  Through fruit development the carpel provides nourishment necessary to bring a new generation to life, to a point of independence, just as we understand the gestation period in animals.  That is what botanists understand and imply when describing the carpel (pistil) as the female component of a flower.

The act:  Everything that has happened by the time the sperm nucleus arrives at the egg is not simply foreplay.  Both sperm and egg will have been through extreme trials,  each having been downsized to a single cell (a single nucleus) in successive stages.  For pollen, individual diploid cells (1st stage) in the anther went through reductive division (meiosis) to produce four spores, each of which (2nd stage) could become a pollen grain.  The spores are haploid, each having inherited one set of chromosomes during meiosis.  

In the following step, each individual pollen grain we are tracking would form a single “cell” (stage 3), that is the sperm nucleus, stripped of property.  The filters acting on the egg are as stringent.  A single diploid cell (1st stage, the mother cell) underwent reductive division from which a single cell (2nd stage) survived.  That cell multiplied to produce a tiny haploid plant inside its ovule.  One cell in that dependent plant would become the egg (3rd stage).  

At fertilization, the sperm nucleus unites with the egg cell to form a single cell, the zygote (4th stage as a single cell).  This is consummation of sexual reproduction, creation of a new genetic being, a process that began with production of male and female gametes, sperm & egg.  But remember, the basic cell itself is a continuation of life; with the new zygote, the software has changed and the plant reboots for the next generation.

As described above, this reboot was an intricate process, four successive shut-downs that reduced the future plant to single nuclei, three stages of single cells in each parental line, and a final single diploid cell (the zygote) – four critical moments at which problematic combinations would be culled before the next generation could come on line.  Given this brutal selection regimen, it’s no wonder plants have been so successful over hundreds of millions of years.  The cell that becomes an individual made of trillions of cells was returned to factory settings and road tested four times, with many trials yet to come.  

Fertilization meant a sperm nucleus had to move from the stigma through tissues in the style and carpel, somehow arriving at an egg.  In small flowers, this involves pollen tube growth of less than a few millimeters   But in large and extravagant flowers, such as Crinum americanum, the distance is considerable, with styles easily surpassing 50 mm.  in the ovary, each ovule that develops has to receive its own sperm nucleus from a separate pollen grain.  It’s quite a feat.  Every seed produced by flowering plants began the journey as a pollen grain in some anther and an ovule in a receptive ovary.  The tiny thousand plus orchid seed maturing in a single capsule reflect an equal number of pollen grains, their tubes having elongated to ready ovules utilizing sugars provided by the style and following a chemical trail emanating from the egg sac.  On arrival, each tube delivers a sperm nucleus to fertilize the egg. Fertilization is consummation, generating the first cell of a new generation – the zygote.  That cell will become the embryonic plant.

But that is not the only union that takes place in the egg sac of flowering plants.  Recalling earlier discussion regarding a tissue people call endosperm, you’ll want to know that a second nucleus from the pollen tube unites with two nuclei in the egg sac, completing a ménage à trois that gives rise to the triploid (or variously polyploid) endosperm.  To botanists, this is the holy grail – double fertilization.  As Dumbledore says, “two lives will be saved tonight.”  Those two lives, the diploid zygote, and the endosperm, evidence a curious peculiarity that unites flowering plants.  Double fertilization characterizes all Angiosperms, and is one pillar in the argument that separates flowering plants from the Gymnosperms.

With fertilization, even at pollination, we typically notice immediate changes in a flower.  There’s no reason to maintain such an elaborate and costly device, the flower, when the job is done.  After all, the flower exists for pollination and reproduction.  When the ovule begins development as a seed and the carpel (the pistil) shunts to its future as a fruit, any floral parts not important to fruit development are off subsidy and will fade.  

The Meaning of Fruit

“Fruit” is one of many plant terms with a much tighter meaning to botanists than in street language, and even botanists might not agree fully.  Some botanists extend the term “Flower” to include male and female cones of Gymnosperms which means the word “Fruit” has to embrace mature cones.  That is not the position I adopt here, because we’d then need to introduce a wholy new and separate set of arcane terminology to distinguish between Gymnosperms and Angiosperms.  In this discussion, “fruit” is the mature stage of a carpel (or multicarpellate structure), which of course includes the mature seed.  In studying the developmental morphology of various fruiting structures, we talk about a fruit wall, which is what happens to tissues comprising the carpel.  That wall can be thin and hard at maturity, or fleshy, or multi-layered, with soft and hard components.  Do not be surprised to learn that fruiting structures have spawned a multitude of terms that carry complex meaning.  And what’s more confounding is that many common words (such as berry and nut) have precise meaning in botanical terminology that’s quite different from street or market language.

Fruit of many wildflowers develop from a superior ovary that was plainly visible at the flowering stage.  Lilium and Hypericum flowers produce superior ovaries, which means the fruit wall that develops consists purely of tissue derived from the carpel (the fertile leaf).  More complex structures might be described as “inferior” because the fruit appears to develop below a perianth and anthers, once again too simple of an interpretation.  Generally, when a fruit is inferior, that tells us the fruit wall is an amalgam of floral tissues, and/or even the stem on which flower segments formed.  Rhexias, Cucurbits, Composites, and Orchids develop inferior ovaries (thus inferior fruit) with walls inconspicuously incorporating sepal, petal, and stamen tissues.  A Cactus imbeds the ovary in stem tissue as well, which is evident from presence of spines on the fruit of some species. 

Either from superior or inferior ovaries, many plants  (and most wildflowers) develop fruit that is hard and dry at maturity, opening naturally or not.  Generally, we think of these as capsules or pods, but there are scores of terms specific to structurally different kinds of dry fruit.  Fabaceae uniformly develop a single carpel as a dry to leathery characteristic pod called a Legume (yes, there are fleshy legumes too).  Also a single carpel, the hard pod of Asclepiads, technically a follicle, splits open to loft plumed seed in the air. Our native Orchids all develop dry capsules, made of 3 carpels, that split open at maturity..   

As was mentioned several sections back, many seed, at maturity, are dispersed while still inside the fruit.  With certain plant families, the dry carpel and seed are inseparable,.  All Composites make an inferior, 1-seeded fruit, a cypsela (also achene) in which the outer layers, technically, are really the fruit wall. The superior 1-seeded Grass fruit is termed a caryopsis.  The paired carpels of Apiaceae are more curious yet, with each separate carpel developing as a 1-seeded fruit termed a mericarp.   By their fruit, you’ll know them.

Other kinds of flowers develop fruit walls that are fleshy, often called berries or drupes.  Once again, those terms have more precise botanical meaning, so there’s no shortage of extra terminology for structurally distinct soft fruit.  If you want to explore this topic more thoroughly, check out my page on Carpology.  Moreover, a list of wildflowers with superior vs. inferior ovaries is appended to this discussion.

This brings us, finally, full circuit.  Fruit develop, bearing mature seed.  Many seed are released, shed or removed somehow from the mature fruit, or shed integral with the fruit wall. Dispersal becomes the second moment in the cycle at which a genotype might find a new destination. Some will lodge in the shadow of the parent plant, others might hitchhike on the feet of waterbird, fur of a bear, in the gut of an herbivore, or through the lofting of a breeze.  Others will be caught by a mower, scooped up in the bucket of earth moving equipment, or purposely collected for propagation. Most will never travel outside the footprint of the established population, an occasional few will land miles, even hundreds of miles distant.

That doesn’t mean plants have failed to explore methods of dispersal. Parallel to study of pollination syndromes, people who study seed and fruit dispersal have explained suites of characteristics associated with the many vectors plants have adapted for hitchhiking. These, of course, glisten with terminology. Autochory indicates a plant takes matters into its own, either simply dropping seed and leaving dispersal to gravity & circumstance, or propelling seed (ballochory). Allochory indicates some vector or dispersal agent is involved, such as wind (anemochory), water (hydrochory), animals (epizoochory & endozoochory), ants (myrmecochory), even humans (anthropochory). The dispersal sweepstakes are taken with occasional feats of long-distance movement. We imagine this can happen, occasionally, when wind-dispersed seed are lofted incredible distances, or when seed in the mud on feet of migratory bird hitchhike their way to a distant wetland. Or perhaps, some animal makes an unexpectedly-long trip with impervious seed in its gut. But the most impactful syndrome has been anthropochory. Human activity, today, outreaches any other means of long distance seed (and plant) dispersal. But for conservation attempts at reintroduction, most of anthropochory has not turned out so well for native habitats.

Despite the many adaptations to promote dispersal, the vast majority of seed that set forth will never germinate and establish successfully. Many that live will be eaten or trampled. A very few will live to reproduce. Those that do may have survived based on some beneficial trait, nature having selected based on a useful character. Others would be stochastic survivers, having no special character or skill other than survivor’s luck at a roll of the dice. Somehow, the balance between extinction, selection, and stochasticism, over hundreds of millions of years, has brought us the wildflowers we study and enjoy today.

Florida Wildflower Plant Lists – Structures, Strategies, & Syndromes. FOF indicates information was extracted from the multi-volume Flora of Florida set, followed by Roman numerals indicating the specific volume. ((STILL UNDER CONSTRUCTION….Feedback welcomed)

Personally, I’m wishing there were some general resource that pointed to what is known about the biologies of Florida wildflowers. So much has been learned, and much has been published, such that the trove of information held by a wide range of practitioners, researchers, horticulturists, naturalists, gardeners, and people who simply love native plants is remarkable. Florida has a wonderful resource in the Atlas of Florida Plants, and active programs in systematics and ecology. But for so many of us, the broad range of information concerning the biologies of Florida wildflowers is difficult to access. There’s no comprehensive summary available that reveals what has been recorded about population biologies, pollination & breeding strategies, microhabitats, germination and establishment, or ecological services. Publications may be difficult to access, many are behind paywalls.

As I encounter information, most importantly when I can cite a reference, I will add information to the following lists. The idea is to concentrate on wildflowers, but I’ll include the entire flora as useful. I’d truly appreciate information from the broader community, and welcome your input through email: planted@huntington.org.

Dioecious Plants: Baccharis, Ceratiola, Cuscuta, Ilex; Myrica (Morella), Polyganum polyganella,

Monoecious Plants: BUXACEAE: Pachysandra procumbens. CUCURBITACEAE: Cayaponia, Cucurbita, Melothria, Sycios.

Annual & Biennial (Semelparous) Wildflowers:

FOF II – CUCURBITACEAE: Cucurbita okeechobeensis, Sycios angulatus. PARNASSIACEAE (CELASTRACEAE): Lupuropetalon.

FOF VI – CONVOLVULACEAE: Most species are perennial vines, and many Convolvulaceae are non-native, several considered invasive. Ipomoea cordatotriloba, I. lacunosa, I. leucantha, I. muricatam, Jacquemontia tamnifolia. SOLANACEAE: Datura stramonium, Physalis cordata, P. pubescens, Solanum americanum. TETRACHONDRACEAE: Polypremum procumbens. PLANTAGINACEAE: Callitriche heterophylla, C. peploides, Gratiola floridana, G. virginiana, Linaria, Plantago aristata, P. heterophylla, P. pusilla, P. virginica, Veronica pregrina (the sole native among 8 species of Veronica). SCROPHULARIACEAE:

FOF VII – APIACEAE: Chaerophyllum, Ptelimnium, Sanicula canadensis, Spermolepis, Trepocarpus. ASTERACEAE: The vast majority of native Composites are perennial herbs or shrubs. Most annual Composites seem to be introduced. Many larger genera of perennial herbs include one or two annuals (i.e. Helianthus, Symphiotrichum). Acanthospermum, Acicarpha,. Ambrosia artemesifolia, A. trifida, Aphanostephus, Artemesia campestris, Balduina angustifolia, Bidens, Bradburia, Chrysopsis (biennial to short-lived perennial), Cirsium (biennial to short-lived perennial), Conyza, Erigeron vernus (biennial to short-lived perennial), Gamochaeta, Helenium amarum, Helianthus agrestis, Iva angustifolia, I. annua, I. microcephala, Krigia cespitosa, K. virginica, Lactuca, Pectis linearifolia, P. prostrata, Pseudognaphalium, Pyrrhopappus, Soliva, Symphiotrichum bahamense, S. subulatum, Xanthium (cosmopolitan). CALYCERACEAE (all annual), CAMPANULACEAE: Campanula americana, Lobelia flaccidifolia, L. hemophylla, Triodanus, OROBANCHACEAE: Agalinus, Aphyllon (holoparasitic), Aureolaria pectinata, Castilleja, Epifagus, Macranthera (?), Orobanche (holoparasitic), Seymeria, Striga.

Herbaceous Families (Families in which all Florida members are herbaceous): Apiaceae, Campanulaceae, Menyanthaceae, Calyceraceae, Calyceraceae, Tetrachondraceae, Parnassiaceae (now commonly subsumed to Celastraceae), Plantaginaceae,

Aquatic Plants: HALORAGACEAE: Myriophyllum, Proserpinaca. MENYANTHACEAE: Nymphoides.

Geophytes with USOs (Underground Storage Organs): AMARYLLIDACEAE: Crinum, Hymenocallis, Zephyranthes. ASTERACEAE: Liatris (corm), COMMELINACEAE: Commelina erecta (tuberous roots). MELASTOMATACEAE: Rhexia. XYRIDACEAE: Xyris (bulbs). (Diaz-Toribio & Putz, 2022)

Heterostylous Wildflowers: Heteranthera, Lythrum, Oxalis, Pontederia

Obligately Heterotrophic – Parasitic Lifestyle (Holoparasite): Cuscuta, Phoradendron (because Mistletoe photosynthesizes, these plants are usually considered hemi-parasites, but they cannot live independently, which in my mind moves the arc toward obligate parasitism); OROBANCHACEAE: Aphyllon (FOF, VII), Conopholis (Squawroot, FOF VII), Epiphagus (annual, FOF VII), Orobanche (annual, FOF VII)

Obligately Heterotrophic – Saprophitic Lifestyle: Monotropa, Monotropsis, Burmannia,

Obligately Myco-Heterotrophic Lifestyle – Corallorhiza, Hexalectris

Facultatively Heterotrophic – Hemi-parasitic Lifestyle: OROBANCHACEAE: Agalinus, Aureolaria, Buchnera, Castilleja, Macranthera, Pedicularis, Schwalbea, Seymeria, Striga (annual, FOF VII)

Flowers lacking Petals – NYCTAGINACEA: Mirabilis, Boerhavia. Euphorbia, RANUNCULACEAE: Actaea, Anemone, Clematis, Enemion, Thalictrum, (in Aquilegia, Consolida, Delphinium , and others the sepals are evident & colorful but petals still present),

Pollination Syndrome – Hummingbird – Plants that match the syndrome, though I’ve seen no publications confirming Hummingbird pollination: BIGNONIACEAE: Campsis radicans, Bignonia capreolata,. CAMPANULACEAE: Lobelia cardinalis, CAPRIFOLIACEAE: Lonicera sempervirens. FABACEAE: Erythrina herbacea. HIPPOCASTANACEAE: Aesculus pavia,LAMIACEAE: Calamintha coccinea,. Salvia coccinea, LILIACEAE: Lilium catesbaei. MALVACEAE: Hibiscus coccinea (and possibly others). OROBANCHACEAE: Macranthera flammea,

Pollination Syndrome – Hovering Moth – ACANTHACEAE: Ruellia noctiflora. AMARYLLIDACEAE: Crinum, Hymenocallis. CANNACEAE: Canna flaccida. GENTIANACEAE: Gentiana pinelliana. ORCHIDACEAE: Platanthera blephariglottis, Platanthera nivea.

Pollination Syndrome – Wind (Anemophily) –

Pollination Syndrome – Beetle –

Pollination Syndrome – Bee –

Pollination Syndrome – Butterfly –

Seed/Fruit Dispersal – Wind (Anemochory) – Asclepias, Asteraceae (most), Orchidaceae, Poaceae (most)

Seed/Fruit Dispersal – Animal fur, hide, or feet (Epizoochory) – Bidens, Cenchrus, Desmodium

Seed/Fruit Dispersal – Animal by consumption (Endozoochory) – Asimina,

Seed/Fruit Dispersal – Ants (Myrmecochory) – Asarum, Sanguinaria, Trillium,

Seed/Fruit Dispersal – Water (Hydrochory) – Hibiscus coccineus

Seed/Fruit Self-dispersal – Ballistic (Ballochory) – Chamaechrista, Justicia,

Florida Endemic Species list from ISB Atlas of Florida Plants

Extracted from the Atlas and organized by Family



Justicia crassifoliaTHICKLEAF WATERWILLOWACANTHACEAE
Ruellia succulentaTHICKLEAF WILD PETUNIAACANTHACEAE
Agave decipiensFALSE SISALAGAVACEAE
Amaranthus floridanusFLORIDA AMARANTHAMARANTHACEAE
Hymenocallis franklinensisCOW CREEK SPIDERLILYAMARYLLIDACEAE
Hymenocallis gholsoniiGHOLSON’S SPIDERLILYAMARYLLIDACEAE
Hymenocallis godfreyiGODFREY’S SPIDERLILY; ST. MARK’S MARSH SPIDERLILYAMARYLLIDACEAE
Hymenocallis henryae var. glaucifolia
AMARYLLIDACEAE
Hymenocallis henryae var. henryaeHENRY’S SPIDERLILY; GREEN SPIDERLILYAMARYLLIDACEAE
Hymenocallis palmeriALLIGATORLILYAMARYLLIDACEAE
Hymenocallis puntagordensisSMALLCUP SPIDERLILYAMARYLLIDACEAE
Hymenocallis rotataSPRING-RUN SPIDERLILYAMARYLLIDACEAE
Hymenocallis tridentataFLORIDA SPIDERLILYAMARYLLIDACEAE
Riccardia stricta
ANEURACEAE
Asimina manasotaMANASOTA PAWPAWANNONACEAE
Asimina obovataBIGFLOWER PAWPAWANNONACEAE
Asimina pulchellaPRETTY FALSE PAWPAW; ROYAL FALSE PAWPAW; WHITE SQUIRREL-BANANAANNONACEAE
Asimina rugeliiRUGEL’S FALSE PAWPAW; YELLOW SQUIRREL-BANANAANNONACEAE
Asimina tetrameraFOURPETAL PAWPAWANNONACEAE
Asimina x bethanyensis
ANNONACEAE
Asimina x colorata
ANNONACEAE
Asimina x kralii
ANNONACEAE
Asimina x oboreticulata
ANNONACEAE
Asimina x peninsularis
ANNONACEAE
Eryngium cuneifoliumWEDGELEAF ERYNGO; SCRUB ERYNGIUMAPIACEAE
Tiedemannia filiformis subsp. greenmaniiGIANT WATER COWBANE; GIANT WATER-DROPWORTAPIACEAE
Asclepias curtissiiCURTISS’ MILKWEEDAPOCYNACEAE
Asclepias feayiFLORIDA MILKWEEDAPOCYNACEAE
Ilex opaca var. arenicolaSCRUB HOLLYAQUIFOLIACEAE
Sabal etoniaSCRUB PALMETTOARECACEAE
Sabal x miamiensis
ARECACEAE
Asplenium x biscaynianumBISCAYNE SPLEENWORTASPLENIACEAE
Asplenium x curtissiiCURTISS’ SPLEENWORTASPLENIACEAE
Asplenium x plenumRUFFLED SPLEENWORTASPLENIACEAE
Arnoglossum albumWHITE INDIAN PLANTAINASTERACEAE
Arnoglossum floridanumFLORIDA INDIAN PLANTAINASTERACEAE
Berlandiera subacaulisFLORIDA GREENEYESASTERACEAE
Berlandiera x humilis
ASTERACEAE
Bigelowia nudata subsp. australisPINELAND RAYLESS GOLDENRODASTERACEAE
Brickellia mosieriMOSIER’S FALSE BONESET; BRICKELLBUSHASTERACEAE
Carphephorus carnosusPINELAND CHAFFHEADASTERACEAE
Carphephorus odoratissimus var. subtropicanusPINELAND PURPLE; FALSE VANILLALEAFASTERACEAE
Chromolaena frustrataCAPE SABLE THOROUGHWORTASTERACEAE
Chrysopsis delaneyiDELANEY’S GOLDENASTERASTERACEAE
Chrysopsis floridanaFLORIDA GOLDENASTERASTERACEAE
Chrysopsis highlandsensisHIGHLANDS GOLDENASTERASTERACEAE
Chrysopsis lanuginosaLYNN HAVEN GOLDENASTERASTERACEAE
Chrysopsis latisquameaPINELAND GOLDENASTERASTERACEAE
Chrysopsis linearifolia subsp. dressiiDRESS’ GOLDENASTERASTERACEAE
Chrysopsis linearifolia subsp. linearifoliaNARROWLEAF GOLDENASTERASTERACEAE
Chrysopsis subulataSCRUBLAND GOLDENASTERASTERACEAE
Coreopsis bakeriBAKER’S COREOPSISASTERACEAE
Coreopsis floridanaFLORIDA TICKSEEDASTERACEAE
Eupatorium mikanioidesSEMAPHORE THOROUGHWORTASTERACEAE
Eurybia spinulosaAPALACHICOLA ASTER; PINEWOODS ASTERASTERACEAE
Flaveria floridanaFLORIDA YELLOWTOPSASTERACEAE
Garberia heterophyllaGARBERIAASTERACEAE
Hasteola robertiorumHAMMOCKHERB; GULF HAMMOCK INDIAN PLANTAINASTERACEAE
Helianthus carnosusLAKESIDE SUNFLOWER; FLATWOODS SUNFLOWERASTERACEAE
Helianthus debilis subsp. debilisEAST COAST DUNE SUNFLOWERASTERACEAE
Helianthus debilis subsp. vestitusWEST COAST DUNE SUNFLOWERASTERACEAE
Liatris gholsoniiGHOLSON’S GAYFEATHERASTERACEAE
Liatris ohlingeraeFLORIDA GAYFEATHER; SCRUB BLAZING-STARASTERACEAE
Liatris provincialisGODFREY’S GAYFEATHER; GODFREY’S BLAZING-STARASTERACEAE
Liatris savannensisSAVANNA GAYFEATHERASTERACEAE
Melanthera parvifoliaSMALL-LEAF SQUARESTEMASTERACEAE
Palafoxia feayiFEAY’S PALAFOXASTERACEAE
Pectis linearifoliaFLORIDA CINCHWEEDASTERACEAE
Phoebanthus grandiflorusFLORIDA FALSE SUNFLOWERASTERACEAE
Pityopsis flexuosaZIGZAG SILKGRASSASTERACEAE
Pluchea longifoliaLONGLEAF CAMPHORWEEDASTERACEAE
Rudbeckia graminifoliaGRASSLEAF CONEFLOWERASTERACEAE
Symphyotrichum fontinaleFLORIDA WATER ASTERASTERACEAE
Symphyotrichum plumosum
ASTERACEAE
Verbesina chapmaniiCHAPMAN’S CROWNBEARDASTERACEAE
Vernonia x concinna
ASTERACEAE
Nasturtium floridanumFLORIDA WATERCRESSBRASSICACEAE
Tillandsia floridanaFLORIDA AIRPLANTBROMELIACEAE
Tillandsia simulataFLORIDA AIRPLANTBROMELIACEAE
Consolea corallicolaSEMAPHORE PRICKLYPEAR; SEMAPHORE CACTUSCACTACEAE
Harrisia aboriginumPRICKLY APPLECACTUS; WEST COAST PRICKLY-APPLECACTACEAE
Harrisia fragransCARIBBEAN APPLECACTUS; INDIAN RIVER PRICKLY-APPLE; SIMPSON’S APPLECACTUSCACTACEAE
Opuntia abjectaKEYS SPANISH LADYCACTACEAE
Opuntia austrinaDEVIL’S-TONGUECACTACEAE
Opuntia ochrocentraBULLSUCKERSCACTACEAE
Campanula floridanaFLORIDA BELLFLOWERCAMPANULACEAE
Campanula robinsiaeROBINS’ BELLFLOWER; CHINSEGUT BELLFLOWERCAMPANULACEAE
Lobelia apalachicolensisAPALACHICOLA LOBELIACAMPANULACEAE
Lobelia homophyllaPINELAND LOBELIACAMPANULACEAE
Paronychia chartaceaPAPER NAILWORT; PAPERY WHITLOW-WORTCARYOPHYLLACEAE
Paronychia discoveryiFLORIDA NAILWORTCARYOPHYLLACEAE
Paronychia minimaPAPER NAILWORTCARYOPHYLLACEAE
Lechea cernuaNODDING PINWEED; SCRUB PINWEEDCISTACEAE
Lechea divaricataDRYSAND PINWEED; SPREADING PINWEEDCISTACEAE
Lechea lakelaeLAKELA’S PINWEEDCISTACEAE
Hypericum chapmaniiAPALACHICOLA ST.JOHN’S-WORTCLUSIACEAE
Hypericum cumulicolaHIGHLANDS SCRUB ST.JOHN’S-WORT; HIGHLANDS SCRUB HYPERICUMCLUSIACEAE
Hypericum edisonianumARCADIAN ST.JOHN’S-WORT; EDISON’S ST.JOHN’S-WORT; EDISON ASCYRUMCLUSIACEAE
Hypericum exileFLORIDA SANDS ST.JOHN’S-WORTCLUSIACEAE
Hypericum lissophloeusSMOOTHBARK ST.JOHN’S-WORTCLUSIACEAE
Callisia ornataFLORIDA SCRUB ROSELINGCOMMELINACEAE
Bonamia grandifloraFLORIDA LADY’S NIGHTCAP; FLORIDA BONAMIACONVOLVULACEAE
Jacquemontia curtissiiPINELAND CLUSTERVINE; PINELAND JACQUEMONTIACONVOLVULACEAE
Jacquemontia reclinataBEACH CLUSTERVINE; BEACH JACQUEMONTIACONVOLVULACEAE
Stylisma abditaSHOWY DAWNFLOWER; HIDDEN STYLISMA; AUSTIN’S DAWNFLOWERCONVOLVULACEAE
Carex paeninsulaePENINSULA SEDGECYPERACEAE
Carex vexansFLORIDA HAMMOCK SEDGECYPERACEAE
Rhynchospora megaplumosaLONGBRISTLE BEAKSEDGECYPERACEAE
Drosera filiformis var. floridanaTHREADLEAF SUNDEWDROSERACEAE
Drosera x californica var. arenaria
DROSERACEAE
Monotropsis reynoldsiaePIGMYPIPESERICACEAE
Rhododendron minus var. chapmaniiCHAPMAN’S RHODODENDRONERICACEAE
Eriocaulon nigrobracteatumBLACK-BRACTED PIPEWORTERIOCAULACEAE
Croton glandulosus var. floridanusVENTE CONMIGOEUPHORBIACEAE
Euphorbia confertaEVERGLADE KEY SANDMATEUPHORBIACEAE
Euphorbia cumulicolaCOASTAL DUNE SANDMAT; SAND DUNE SPURGEEUPHORBIACEAE
Euphorbia deltoidea subsp. deltoideaWEDGE SANDMAT; ROCKLAND SPURGEEUPHORBIACEAE
Euphorbia deltoidea subsp. pinetorumPINELAND SANDMATEUPHORBIACEAE
Euphorbia deltoidea subsp. serpyllumWEDGE SANDMATEUPHORBIACEAE
Euphorbia garberiGARBER’S SANDMAT; GARBER’S SPURGEEUPHORBIACEAE
Euphorbia hammeri
EUPHORBIACEAE
Euphorbia inundata var. garrettii
EUPHORBIACEAE
Euphorbia pinetorumPINELAND SPURGE; EVERGLADES POINSETTIAEUPHORBIACEAE
Euphorbia polyphyllaLESSER FLORIDA SPURGEEUPHORBIACEAE
Euphorbia porterianaPORTER’S SANDMAT; PORTER’S SPURGEEUPHORBIACEAE
Euphorbia rosescensSCRUB SPURGEEUPHORBIACEAE
Euphorbia telephioidesTELEPHUS SPURGEEUPHORBIACEAE
Tragia saxicolaFLORIDA KEYS NOSEBURN; ROCKLANDS NOSEBURNEUPHORBIACEAE
Aeschynomene pratensis var. pratensisMEADOW JOINTVETCHFABACEAE
Amorpha herbacea var. crenulataMIAMI LEAD PLANTFABACEAE
Baptisia calycosaFLORIDA WILD INDIGOFABACEAE
Baptisia simplicifoliaSCAREWEEDFABACEAE
Centrosema arenicolaPINELAND BUTTERFLY PEA; SAND BUTTERFLY PEAFABACEAE
Chamaecrista lineata var. keyensisNARROWPOD SENSITIVE PEA; KEY CASSIAFABACEAE
Chapmannia floridanaFLORIDA ALICIAFABACEAE
Clitoria fragransSWEETSCENTED PIGEONWINGSFABACEAE
Crotalaria avonensisAVON PARK RATTLEBOX; AVON PARK HAREBELLSFABACEAE
Dalea adenopodaSUMMER FAREWELLFABACEAE
Dalea floridanaFLORIDA PRAIRIECLOVERFABACEAE
Galactia pinetorumPINE ROCKLAND MILKPEAFABACEAE
Galactia smalliiSMALL’S MILKPEAFABACEAE
Lupinus aridorumBECKNER’S LUPINE; MCFARLIN’S LUPINEFABACEAE
Lupinus westianus var. westianusGULF COAST LUPINEFABACEAE
Rhynchosia cinereaBROWNHAIR SNOUTBEANFABACEAE
Tephrosia angustissima var. angustissimaNARROWLEAF HOARYPEAFABACEAE
Tephrosia angustissima var. curtissiiCURTISS’ HOARYPEAFABACEAE
Tephrosia mysteriosaSANDHILL TIPPITOESFABACEAE
Tephrosia rugeliiRUGEL’S HOARYPEAFABACEAE
Tephrosia x varioforma
FABACEAE
Vachellia farnesiana var. pinetorumPINELAND ACACIAFABACEAE
Vicia ocalensisOCALA VETCHFABACEAE
Quercus inopinaSCRUB OAKFAGACEAE
Scientific NameCommon NameFamily
Frullania sabaliana
FRULLANIACEAE
Frullania taxodiocola
FRULLANIACEAE
Gentiana pennellianaWIREGRASS GENTIANGENTIANACEAE
Halophila johnsoniiJOHNSON’S SEAGRASSHYDROCHARITACEAE
Trichomanes punctatum subsp. floridanumFLORIDA BRISTLE FERNHYMENOPHYLLACEAE
Calydorea caelestinaBARTRAM’S IXIAIRIDACEAE
Nemastylis floridanaCELESTIAL LILY; FALLFLOWERING IXIA; HAPPYHOUR FLOWERIRIDACEAE
Carya floridanaSCRUB HICKORYJUGLANDACEAE
Conradina brevifoliaFALSE ROSEMARYLAMIACEAE
Conradina cygnifloraFALSE ROSEMARYLAMIACEAE
Conradina etoniaETONIA FALSE ROSEMARYLAMIACEAE
Conradina glabraAPALACHICOLA FALSE ROSEMARYLAMIACEAE
Conradina grandifloraLARGEFLOWER FALSE ROSEMARYLAMIACEAE
Dicerandra christmaniiLAKE WALES BALM; CHRISTMAN’S MINTLAMIACEAE
Dicerandra cornutissimaLONGSPUR BALM; ROBIN’S MINTLAMIACEAE
Dicerandra densifloraFLORIDA BALMLAMIACEAE
Dicerandra frutescensSCRUB BALM; LLOYD’S MINTLAMIACEAE
Dicerandra immaculata var. immaculataLAKELA’S BALM; OLGA’S MINTLAMIACEAE
Dicerandra immaculata var. savannarumSAVANNA BALM; DICERANDRA-OF-THE-SAVANNASLAMIACEAE
Dicerandra modestaBLUSHING SCRUB BALMLAMIACEAE
Dicerandra thinicolaTITUSVILLE BALMLAMIACEAE
Macbridea albaWHITE BIRDS-IN-A-NESTLAMIACEAE
Physostegia godfreyiGODFREY’S FALSE DRAGONHEAD; APALACHICOLA DRAGONHEADLAMIACEAE
Scutellaria floridanaFLORIDA SKULLCAPLAMIACEAE
Stachydeoma graveolensMOCK PENNYROYALLAMIACEAE
Persea borbonia var. humilisSILK BAYLAURACEAE
Cheilolejeunea polyantha var. polyantha
LEJEUNEACEAE
Cololejeunea subcristata
LEJEUNEACEAE
Lejeunea minutiloba var. heterogyna
LEJEUNEACEAE
Pinguicula ionanthaVIOLET BUTTERWORT; PANHANDLE BUTTERWORTLENTIBULARIACEAE
Linum arenicolaSAND FLAXLINACEAE
Linum carteri var. carteriCARTER’S FLAX; EVERGLADES FLAXLINACEAE
Linum smalliiSMALL’S FLAXLINACEAE
Micranthemum glomeratumMANATEE MUDFLOWERLINDERNIACEAE
Spigelia loganioidesFLORIDA PINKROOT; LEVY PINKROOTLOGANIACEAE
Cuphea asperaTROPICAL WAXWEEDLYTHRACEAE
Lythrum flagellareFLORIDA LOOSESTRIFE; LOWLAND LOOSESTRIFELYTHRACEAE
Lythrum nieuwlandiiNIEUWLAND”S LOOSESTRIFELYTHRACEAE
Magnolia macrophylla var. asheiBIGLEAF MAGNOLIA; ASHE’S MAGNOLIAMAGNOLIACEAE
Schoenocaulon dubiumFLORIDA FEATHERSHANKMELANTHIACEAE
Cartrema floridanumSCRUB WILD OLIVEOLEACEAE
Chionanthus pygmaeusPIGMY FRINGETREEOLEACEAE
Govenia floridanaFLORIDA GOVENIA; GOWEN’S ORCHIDORCHIDACEAE
Spiranthes igniorchisFIRE LADIESTRESSESORCHIDACEAE
Spiranthes triloba
ORCHIDACEAE
Triphora craigheadiiCRAIGHEAD’S NODDINGCAPS; CRAIGHEAD’S ORCHIDORCHIDACEAE
Triphora rickettiiRICKETT’S NODDINGCAPSORCHIDACEAE
Agalinis flexicaulisHAMPTON FALSE FOXGLOVE; SPRAWLING FALSE FOXGLOVEOROBANCHACEAE
Phyllanthus liebmannianus subsp. platylepisFLORIDA LEAFFLOWER; PINEWOODS DAINTIESPHYLLANTHACEAE
Plagiochila invisa
PLAGIOCHILACEAE
Mecardonia acuminata subsp. peninsularisAXILFLOWERPLANTAGINACEAE
Andropogon cumulicola
POACEAE
Aristida patulaTALL THREEAWNPOACEAE
Aristida rhizomophoraFLORIDA THREEAWNPOACEAE
Coleataenia abscissaCUTTHROATGRASSPOACEAE
Dichanthelium ensifolium var. breveDWARF CYPRESS WITCHGRASSPOACEAE
Digitaria floridanaFLORIDA CRABGRASSPOACEAE
Digitaria gracillimaLONGLEAF CRABGRASSPOACEAE
Digitaria paucifloraTWOSPIKE CRABGRASS; FLORIDA PINELAND CRABGRASSPOACEAE
Digitaria simpsoniiSIMPSON’S CRABGRASSPOACEAE
Echinochloa paludigenaFLORIDA COCKSPURPOACEAE
Eragrostis pectinacea var. tracyiSANIBEL ISLAND LOVEGRASSPOACEAE
Eriochloa michauxii var. simpsoniiSIMPSON’S CUPGRASSPOACEAE
Piptochaetium avenacioidesFLORIDA NEEDLEGRASSPOACEAE
Schizachyrium niveumPINESCRUB BLUESTEMPOACEAE
Schizachyrium rhizomatumSOUTH FLORIDA LITTLE BLUESTEMPOACEAE
Schizachyrium sericatumSILKY BLUESTEMPOACEAE
Sorghastrum apalachicolenseAPALACHICOLA INDIANGRASSPOACEAE
Sporobolus vaseyiFLORIDA SANDREED; CURTISS’ SANDGRASSPOACEAE
Polygala lewtoniiLEWTON’S MILKWORT; LEWTON’S POLYGALAPOLYGALACEAE
Polygala rugeliiYELLOW MILKWORTPOLYGALACEAE
Polygala smalliiSMALL’S MILKWORT; TINY POLYGALAPOLYGALACEAE
Eriogonum longifolium var. gnaphalifoliumLONGLEAF WILD BUCKWHEAT; SCRUB BUCKWHEATPOLYGONACEAE
Polygonum basiramiaFLORIDA JOINTWEED; TUFTED WIREWEEDPOLYGONACEAE
Polygonum delopyrumHAIRY JOINTWEEDPOLYGONACEAE
Polygonum dentocerasSMALL’S JOINTWEED; WOODY WIREWEED; SANDLACEPOLYGONACEAE
Polygonum nesomiiLARGEFLOWER JOINTWEED; SANDHILL WIREWEEDPOLYGONACEAE
Polygonum polygamum var. brachystachyumOCTOBER FLOWERPOLYGONACEAE
Potamogeton floridanusFLORIDA PONDWEEDPOTAMOGETONACEAE
Clematis baldwiniiPINE-HYACINTHRANUNCULACEAE
Pseudoziziphus celataFLORIDA JUJUBE; SCRUB ZIZIPHUSRHAMNACEAE
Prunus geniculataSCRUB PLUMROSACEAE
Spermacoce neoterminalisEVERGLADES KEY FALSE BUTTONWEEDRUBIACEAE
Nolina atopocarpaFLORIDA BEARGRASSRUSCACEAE
Nolina brittonianaBRITTON’S BEARGRASSRUSCACEAE
Sideroxylon reclinatum subsp. austrofloridenseFLORIDA BULLYSAPOTACEAE
Sideroxylon rufohirtumRUFOUS FLORIDA BULLYSAPOTACEAE
Sarracenia rubra subsp. gulfensisGULF COAST REDFLOWER PITCHERPLANTSARRACENIACEAE
Illicium parviflorumYELLOW ANISETREE; STAR ANISESCHISANDRACEAE
Taxus floridanaFLORIDA YEWTAXACEAE
Harperocallis flavaHARPER’S BEAUTYTOFIELDIACEAE
Glandularia maritimaCOASTAL MOCK VERVAINVERBENACEAE
Glandularia tampensisTAMPA MOCK VERVAINVERBENACEAE
Lantana depressa var. depressaROCKLAND SHRUBVERBENA; PINELAND LANTANAVERBENACEAE
Lantana depressa var. floridana
VERBENACEAE
Lantana depressa var. sanibelensis
VERBENACEAE
Lantana x floridana
VERBENACEAE

Florida Endemics – directly transcribed from Wikipedia:

WIKIPEDIA Category: Endemic flora of Florida

The following 109 pages are in this category. This list may not reflect recent changes.

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Ten Big Ideas about Plants

You may not be able to see the forest because of the trees….  As I continue to learn more about plants, some of my greatest teachers have taken the time to help me build a framework that provides themes common to the plant kingdom.   Those overarching themes are described below, as big ideas.  Just as a curator would tie disparate elements of an exhibition together with some unifying thought, you might find a larger picture useful in making sense of details…, staying out of the weeds.

I.  PLANTS ARE ALIVE

At its most elementary level, life is a process, one in which remarkable molecular robots capture and conserve energy and manage water.  Moreover, those molecules are part of a  self-replicating system we call LIFE.  Life is:

Cellular – compartmentalized, regulated by membranes that permit or prohibit passage of materials. A great vat of organic soup could not control it’s internal contents, harness energy, or replicate itself. Cells must be small enough to move components and respond to stimuli nearly instantaneously. Cells work together to form complex organisms, and we are talking about millions and trillions of cells.

We are told the largest single cell is an Ostrich egg, but the nucleus and capacities of that “cell” are limited. Immediately, with fertilization, the zygote that forms is miniscule. An entire bird egg does not divide to form two huge cells, so I’d place a disclaimer on this claim. The egg is a small somewhat-cell sitting atop a huge reservoir of nutritional matter that sustains development of a hatchling.

Energy-dependent – capturing and managing energy to create order and organization.  Without an external source of energy (such as the sun), there can be no life.

Organic – crafted of carbon-based molecules that are non-random and modular.  The ability of carbon to form chains of atoms means that countless energy-rich kinds of molecules are possible, such as proteins & enzymes, carbohydrates, fats, DNA. 

Self-contained & Interactive – bounded by a self-generated organic membrane or interface that maintains internal structure, process, and equilibrium, while keeping the physical world at bay.   Each life is its own tiny universe, competent to carry on process and rejuvenate itself.

Water-mediated – hoarding a precious film or droplet of water and solutes partitioned and surrounded by specialized organic membranes.  Living cells depend on water as the solvent in which reactions occur and equilibrium is maintained.

Iterative – marked by rhythmic repetition at every molecular, cellular, and organismal level.  Plants cycle and recycle materials and process, building on what has been built, replacing what was expended, expanding through repetition – in response to internal drive and greater natural patterns, such as days and seasons.

Self-perpetuating and Extinguishable – actively constructing and deconstructing itself, through innate control/intelligence.  Cells and living beings constantly change, rebuilding and reconstituting systems.  When membrane or process is destroyed, or the surrounding environment becomes too hostile, cells are denatured.  They die and that spark of life is extinguished.  At a fundamental level, we realize that any reactant inimicable to the basic components of life, such as DNA, proteins, or glucose, would eradicate living beings.

Successive – originating from living ancestors, in branching chains that form direct lineages back to those for all other life forms on earth, establishing unbroken cadence to all cells.  Living cells and beings are not generated spontaneously.  All cellular life forms derive from creation’s ground zero, a point when the earliest life forms amalgamated in testy confederations we think of as cells, which then went on to build living beings of nearly countless form.  

BEING Alive means that plants actively:

  • Grow and develop
  • Acquire and process light energy, water, nutrients, and gases
  • Reproduce and disperse
  • Interact and evolve

II. PLANT CELLS ARE DIFFERENT FROM ANIMAL CELLS:  Plant cells are membrane-bound real time spaces, delimited and functioning in a challenging and unrelenting world.  All living plant cells have discernible components, such as a nucleus (or nuclei), membranes, plastids, and mitochondria, each managing its own vital function.  Some cells survive singularly, completely capable of carrying out the functions that define life.  Others live communally as organisms or tissue systems within organisms, where many kinds of cells take on distinctive form and function. 

In most cases, unlike other organisms, plant cells are encapsulated by  a network of cellulose microfibrils that constitute a cell wall (which becomes impregnated with other compounds.)  This fabric-like wall is produced from inside the individual cell; many kinds of specialized cells thicken the wall and even deposit other sorts of compounds, such as water-repellent suberin or gummy, protective lignin. The presence and resilience of this wall brings opportunity and character basic to the nature of plants.

Various types of plant, especially those depositing materials in the cell wall, grow to a certain state, at which point they perish so as to take on a lifeless destiny. Vessel elements and trachieds in xylem tissue mature as microscopic hollow tubes through which sap flows. Fibers convey toughness and protection, but to do so they will so completely thicken the cell was as to perish. The same is true for external tree bark, such as cork oak, cells of which become so waterproof as to succumb.

Except for sap you see draining from wounded tissue and waxes deposited over surfaces, almost everything else about a plant is made of nearly countless cells. Plant fibers, long, tough cells that often accompany veins find many uses. Even plant “hairs” (better called trichomes) are cells projecting from the epididermis. Cotton “fibers” are long trichomes with cellulose walls that sufficiently resilient to be spun into thread.

III. THE CHAIN OF LIFE IS UNBROKEN – Life Without End.  As was said in Big Idea #1, life is successive.  For over 3 billion years life has existed as a continuum.  Every organism on earth today is a moment in that drama, each being a composite of cells that trace to some unheralded origin, to some very point when a soup of biochemical reactions was formalized in a protective and regulating cellular envelope, a cell that managed to reproduce itself.  Cell formation and replication was the miracle that stuck.  This was creation that took a billion years to evolve, and happened in a very different world, a world without free atmospheric oxygen. But cells emerged none-the-less, most certainly in a reticulate and complex manner over millions of years.   

Today’s world is pervaded with the most rudimentary (yet marvelous) kinds of cells, like bacteria and blue green algae.  But at some halcyon moment, there came the genesis of a true, nucleus-bearing, mitochondrial-powered cell which is the basis for all complex forms of life on earth today.  The resulting web of life includes plants, which at their furthest point of origin share a single, simple-celled primordial ancestor with all of the world’s animals.  We are one, but prefer to taxonomize ourselves into species, seemingly separate beings.

Biologists have determined there was an ancient progenitor to all cellular life, which has been designated LUCA (the Last Universal Common Ancestor), but I believe such a primordial being should have a Linnaean name,, perhaps the binomial Principium principale.

IV. Elements  are BASIC – Of the many elements in the periodic table, about 19 participate in assembling the beautiful molecular building blocks that make plants, and another 3 show up as incidentals.  There’s a simple mnemonic botany students use to recall this list:  “See Hopkin’s Cafe, Managed by my cynical cousin Mona Alcose”, which is transliterated to C HOPKNS CaFe, Mg B Mn SiNiCl CuZn MoNa AlCoSe (for the 19, leave off the last name “AlCoSe” – Aluminum, Cobalt, & Selenium, which have no known physiological role).  Thus we have the list of elements that have been discovered naturally and functionally present in plant materials: Carbon, Hydrogen, Oxygen, Phosphorus, Potassium, Nitrogen, Sulfur, Calcium, Iron, Magnesium, Boron, Manganese, Nickel, Chlorine, Copper, Zinc, and Molybdenum are considered essential, while Silicon, Sodium, Aluminum, Cobalt, and Selenium are termed “beneficial”, in that some plants sequester or benefit from those elements.  (Aluminum is a true outlier, being generally toxic to biological systems.)

The most abundant elements in plant tissues, Carbon, Oxygen, and Hydrogen are available to plants from air and water.  The rest are harvested from earth.  Using energy from the sun, those elements create the form and process we call life.  Anciently, we think of life’s raw ingredients as Earth – Air – Fire – Water.

Earth – Soil is the principle rooting medium and source of water and nutrients for plants.  Ancient herbalists thought roots could “eat” soil, a logical but inaccurate assumption that persists today in the unfortunate marketing of fertilizer as “plant food.”   Van Helmont and others [TimeLine 1648] demonstrated that plants retrieve only minuscule quantities of nutrients from soil, and that the great proportion of plant fresh weight is water.  It would be a bit more accurate to say that plants consume water, but plants do not “drink” and despite current street talk, they are never “thirsty.”

Air – Between 1727 and 1804 [see TimeLine] a succession of early scientists – Hales, Priestley, Senebier, Ingen-Housz, and de Saussure – convinced fellow scholars that:

  • Some principle (carbon) from the atmosphere is used by plants to build themselves.  If we want to talk about a “food” source for plants, it is the atmosphere.
  • Dephlogisticated air (oxygen) in the atmosphere was liberated from water as part of the “light reactions” during photosynthesis. https://library.si.edu/digital-library/book/experimentsuponv00inge

The significance cannot be better stated than by Ingen-Housz, who wrote in 1779: “When I first found in the works of that excellent philosopher and inventive genius, the reverend Dr. Priestley, his important discovery, that plants wonderfully thrive in putrid air; and that the vegetation of a plant could correct air fouled by the burning of a candle, and restore it again to its former purity and fitness for supporting flame, and for the respiration of animals; I was struck with admiration…

The discovery of Dr. Priestley, that plants thrive better in foul air than in common and in dephlogisticated air (pure oxygen), and that plants have a power of correcting bad air, has thrown a new and important light upon the arrangement of this world. It shews, even to a demonstration, that the vegetable kingdom is subservient to the animal; and, vice versa, that the air, spoiled and rendered noxious to animals by their breathing in it, serves to plants as a kind of nourishment. But in what manner this faculty of plants is excited remained still unknown.”

Fire – We focus on the sun’s energy as the lightning bolt over the arrow in photosynthesis, but in that simplification there is much inherent confusion. 

        CO2 + H2O + light     C6H12O6 (glucose) + O2

Plants do capture energy from sunlight, but not in a single step as implied by the summary formula that almost any book gives for photosynthesis.  Additionally, many plants also track relative quantities of different colors of light to gauge daylength and seasons.  Without the qualities of natural light there would be little potential for flower color to guide pollinators.  And it is oddly easy to forget that the sun’s energy heats the land, water, and atmosphere, which drives all of our climate cycles.

Water constitutes the great bulk of fresh plant weight, and is of course basic to the emulsion that is the arena of life.  But the movement of water through a plant also carries nutrients from soil to foliage and flowers.  And evaporation of water from leaves aids in the flow of water through plants, as well as cooling foliage and habitat.  

The ancient elements remain basic to plant life.  Students are often taught to remember SWAN (Soil, Water, Air, Nutrients) or LAWN (Light, Air, Water, Nutrients) as memory aids, but I am fine with Earth-Air-Fire-Water.

V.  THE RAINBOW (ROYGBIV) is THE USEFUL COLOR RANGE.  When thinking about light, I always have to stop myself in order to recall that longer wavelengths are lower energy and shorter wavelengths are more powerful.  Wavelengths longer than Red (we call this infrared, which carries a lower energy level than red light) come across as heat, important in creating living conditions.  Wavelengths shorter than Violet (the ultraviolet range) are increasingly energetic, and thus increasingly destructive to organic molecules.  It is UV light that sunburns skin and increases chances of skin cancers. 

Biochemical reactions involving light necessarily center on wavelengths that are sufficiently energetic to activate or change an organic molecule, but not so energetic as to destroy that molecule.  Light meeting those strictures is the light we can see, i.e. the visible spectrum – or in other words, the rainbow.  And for us humans, green is at the center.  It is logical that the wavelengths of light animals can utilize for vision (which is, in the end, a series of molecular interactions) are wavelengths that can interact nondestructively with plants.  Since photosynthetic organisms are the primary food producers for the planet, we eat the rainbow. But how are sugars spun from gas, water, and light?

Anciently, people thought plants “ate soil,” somehow turning soil or humus into flesh. By 1648 that viewpoint was challenged through a famous trial (published posthumously) by Jan von Helmont, a study from which he concluded that a willow branch barely extracts any mass from soil as it grows. Some scholars turned their thoughts to water as the source of plant substance. But little was understood about water until after 1780 when Henry Cavendish and other scholars revealed it’s a compound that can be condensed from two gases (not yet named). Even as those discussions advanced, “humus” theory persisted – plants were thought to build themselves from carbon compounds taken in by roots. Beginning in 1827 understanding of soil chemistry matured as work by Carl Sprengel, and later Justin von Liebig established that plants take minerals from soil. Interestingly, had their studies involved hemi-parasites like Agalinus, the conclusions might have been different.

Stephen Hales in his 1727 xxxx, had suggested the atmosphere has a role in plant growth. Characterization of elements that constitute the atmosphere would await the epoch of Black, Priestley, and Lavoisier, when precise instrumentation became the hallmark of chemistry. By 1788 Jean Senebier reported that “fixed air” (the term Black had used for carbon dioxide) is consumed in production of plant matter. The following year, 1789, Jan Ingen-Housz established the relationship between sunlight and generation of oxygen. By the turn of the century, Senebier had established the role of carbon dioxide, followed shortly, in 1804, when Nicolas-Théodore de Saussure published elegant experiments confirming the combined roles of water and atmosphere in plant growth.

More than a century would pass, however, before modern research began to unravel the secrets of photosynthesis. Beginning with a crucial publication by C. B. van Niel in 1923, followed by detailing of the Light Reactions (the Hill Reaction) between 1937 and 1939, and elucidation of the Dark Reactions (the Calvin-Benson-Bassham Cycle) in 1950 the basic process was established.

Here is my take on the simplest formulation. Carbon dioxide diffuses into leaves (and green stems) through stomata, dissolves in water around cells, and (through a complex series of interactions) is attached to an existing 5-carbon compound (a “skeleton”). The new 6-carbon compound is not a glucose; it’s not the final product. Rather, two 3-carbon skeletons result, one of which is refitted as a 5-carbon skeleton to repeat the cycle. The net result is a single carbon has been added to the carbohydrate load.

Bonding a free carbon (from carbon dioxide) to a carbon chain is called “fixing” – a free gas is now fixed, tied down, as a solid. The most celebrated player in this important reaction is RuBisCO (Ribulose-1,5-bisphosphate carboxylase/oxygenase), the fixer, the enzyme that facilitates bonding the carbon. RuBisCO is said to be the most abundant enzyme in nature; it works day or night, but that work isn’t free. There is a price to be paid to fix carbon; chemical energy must be invested in the carbon-to-carbon bond. There’s no free ride. This simple connection is where plants store energy. It’s the reason sugars are a quick energy source, the reason starches are energy reserves.

That chemical energy is generated in daylight through what are called Light Reactions. It’s the light reactions that require chlorophyll, and a host of other receptors. It’s the Light Reactions that play hot potato with electrons, electrons energized when light rays, particularly red and blue spectra, are harvested. Those energized electrons will be scraped of energy that’s invested in chemical currency, leaving the chlorophyll reaction centers unstable, in need of electrons. The centers are “quenched” with electrons stripped from water. In a unique reaction, water is split, generating free oxygen, protons (i.e. hydrogen), and electrons.

Given this basic, though elaborate schema, scientists were excited to discover there are variations that improve the efficiency of this C3 system….

VI. WATER IS THE MEDIUM OF LIFE – Life  as we know it does not seem possible without water.  Water is called the universal solvent because so many compounds are soluble in water.  But whole classes of organic molecules (like fats) do not dissolve readily, and that quality allows these molecules to construct membranes and partition water into zones where different reactions can be organized.  Basically, that membrane-generating capacity means organic molecules can form cells, and the immiscibility between water and membranes allows plants to create concentration gradients (situations in which solutions can be more or less concentrated), as well as generate flows that carry nutrients and produce internal pressure that makes young stems erect and leaves crisp.  

Early plants clearly came into being in a wet world, in moist niches that did not brew the challenges faced by land dwelling plants that are surrounded by air and the chance of desiccation.  Most critically, the sex cells (gametes) of early plants required water for reproduction.  Botanists trace the evolution of land plants from primordial organisms that are completely dependent on water for the sexual cycle (green algae) to those that produce pollen that can be carried through the air to a stigmatic surface.  Conquering terrestrial life is one of many clear evolutionary trends, for both plants and animals.

VII. PLANTS ALTERNATE between Diploid and Haploid stages in each Life Cycle.  A life cycle begins with a diploid cell that forms when a sperm unites with an egg.  The next life cycle begins when a succeeding generation of sperm and egg unite.  But in a single plant life cycle, there are two phases, distinct generations.  With flowering plants, we readily see the diploid phase, which began as a fertilized egg, grew to form an embryo inside a seed, and then into a fully mature plant.  But the second phase, the haploid generation, is practically invisible to us, with pollen being as much as we usually notice. 

The situation looks very different when you study mosses and ferns, and then compare their life cycles to those of cone-bearing or flowering plants.  The moss stage that is chromosomally-equivalent to a flowering plant is a crafty-looking hatpin growing out of the top of a green moss plant.  This clever hatpin is all there is to the diploid generation, emerging atop the green moss from a fertilized egg.  The head of the pin forms a capsule where haploid spores are produced, spores that are dispersed, germinate and grow into the green haploid moss plant. That plant, which we think of as the moss is therefore a great curiosity, being basically equivalent to a pollen grain, in that its cells have only one set of chromosomes (they are haploid) .  Just like the pollen grain, it produces sperm nuclei through simple cell division.  Unlike pollen, the same moss plant can produce eggs just as easily.  

So with the moss, the diploid phase of the life cycle is small and dependent, while the haploid phase is photosynthetic and much more elaborate.  Just as with flowering plants, the two phases (generations) make up a single life cycle, a diploid phase followed by a haploid phase.  Things are kind of reversed as to which phase predominates.

We call this behavior “alternation of generations” – because a complete life cycle requires a diploid generation and a haploid generation.  Botanists are highly amused by this secret life of plants, especially with mosses, in which we can easily see the two distinct phases.  Studying plants of differing levels of complexity, from simpler mosses to ferns, to conifers, and finally to flowering plants, we see growing predominance of the diploid generation and reduction of the haploid phase to just a few cells.  But those few haploid cells, vestigial though they may be, document the stages through which plants alternate, from diploid to haploid in each complete sweep of the lifecycle. 

VIII. Plants Use Vectors for Dispersal – Male gametes have to get around, minimally to fertilize the egg.   The male gamete (the sperm nucleus) is produced in the pollen of seed-bearing gymnosperms and angiosperms, which has to lodge on a receptive surface for the sperm nucleus to access an egg.  This pollination can be close (within the same plant) or over fairly long distances.  In flowering plants, pollination often evolves as elaborate co-adaptations with visiting insects and animals. In fact, the structure of most flowers tells the story of a relationship with certain kinds of insects or other animals, a relationship in which flowers reward visitors for their service (usually).

Another shot at dispersal comes when seed (or specialized vegetative propagules) are moved by wind, water, gravity, or animals to establish in new sites. The various “syndromes” were covered earlier.

In non-flowering plants, spores disperse in the air, and grow into gametophytes that produce sperm which actually swim through liquid to reach an egg.  Though simple, this mechanism has proven highly effective, in that mosses and ferns can have very large ranges.

IX.  Plants develop from POINTS OF PRIMARY GROWTH – Across the board, plants grow in straightforward ways.  The first kind of development, called primary growth, begins with the embryo, in which two growth points originate.  One point is the shoot apical meristem, and this cell-producing zone will give rise to all of the components that form the stem and leaves.  The other point is the root apical meristem, and this will grow to form the root complex in its wake.  The resulting plant has ARCHITECTURE with predictable structure, including future stem development and arrangement of appendages, such as bracts, green leaves, and flower parts.  Each kind of plant has its own intrinsic growth habits based on the ways these meristems behave.  Armed with this simple appreciation of growth patterns, you can decipher almost any plant and discover how it grew to its current state.

Result: Plants only make ROOTS, STEMS, & LEAVES –   It is useful for students of plants to construct their understanding of plants based on this simple concept.  Knowing how to examine a plant and determine what is leaf, what is stem, and what is root empowers you to analyze any plant and make sense of the structure   Comparing these observations for a plant new to you to a plant you truly understand allows you to draw useful ideas about form, function, and relationships.  In the Gardens, we say you can “read” something about the life of a plant through understanding special aspects of its structure.

Let’s carry that a bit further. Leaves show incredible specialization, morphologically (the external form), anatomically (internal cellular organization), and physiologically (biochemical functionality). We certainly see this in photosynthesis, given that three systems have been identified (so far): C3, C4, and CAM. Though CAM (Crassulacean Acid Metabolism) has an older history, the first system basically characterized was C3, which is the base-line for interpreting others. And, C3 is the most straightforward.

X.  Character Boils Down to NATURE & NUTURE – Plant development follows genetic programs, which we call the genotype (or genome).  But life happens and every plant thrives under its own particular circumstances.  Some plants grow in brighter light than others.  Some have better soil, or more protected sites.  A plant may get browsed by a deer, while its sibling is untouched.  Circumstantial differences will engender somewhat distinct growth patterns and responses, yielding a plant that is distinctive from the way it might have grown otherwise or causing one part of a plant to grow differently.  Researchers have cultivated genetically identical plants under differing circumstances and documented the extent to which they respond differently.   But you do not have to go that far.  Just look at the plants you grow.  Cryptanthus (a nice and curious houseplant) will develop more colorful leaves in full sunlight than when it is grown in the shade, as will the cultivated Crotons.  On almost any tree, you can even observe differences within a single plant by plucking a leaf that is exposed to full sun and comparing it to a leaf that formed in the shade of the canopy.  The full sun leaf, generally, will be smaller and thicker than the shade leaf.  These different expressions of genetic potential are responses to nurture (i.e. the environment).

It was stated that genotype refers to the plant genetics, but the exact expression (brightly colored or thicker leaves) is called a plant’s phenotype (pheno refers to how something appears – as in phenomenon). That appearance (expression) can vary depending on circumstances, but it can only vary so much.  The range within which a plant can differ in growth characteristics is called its phenotypic plasticity.   The long-lived question of Nature vs. Nurture has meaning with plants, just as it does with humans, but botanists say Genotype vs. Phenotype.

P.S.  The American Society of Plant Biologists also has published a list of 12 Principles.  https://aspb.org/education-outreach/k12-roots-and-shoots/the-12-principles-of-plant-biology-2/

  1. Plants use the same biological processes and biochemistry as microbes and animals. Yet plants are unique because they mix the sunlight’s energy with chemicals for growth. This process of photosynthesis makes the world’s supply of food and energy.
  2. Plants require certain inorganic elements from soil for growth. Plants play an essential role in the circulation of these nutrients within the biosphere.
  3. Land plants evolved from ocean-dwelling, algae-like ancestors, and plants have played a role in the evolution of life, including the addition of oxygen and ozone to the atmosphere.
  4. Reproduction in flowering plants takes place sexually, resulting in the production of a seed. Reproduction can also occur via asexual propagation.
  5. Plants, like animals and many microbes, respire and utilize energy to grow and reproduce.
  6. Cell walls provide structural support for the plant and also provide fibers and building materials for humans, insects, birds and many other organisms.
  7. Plants exhibit diversity in size and shape ranging from single cells to giant trees; there are ~350,000 plant species.
  8. Plants are a primary source of fiber, medicines, and countless other important products in everyday use.
  9. Plants, like animals, are subject to injury and death due to infectious diseases caused by microorganisms. Plants have unique ways to defend themselves against pests and diseases.
  10. Water is the major molecule in plant cells and organs. It’s essential to plant structure, growth, and the internal circulationof organic molecules and salts.
  11. Plant growth and development are under the control of hormones and can be affected by external signals such as light, gravity, touch or environmental stresses.
  12. Plants live in and adapt to a wide variety of environments. Plants provide diverse habitats for birds, beneficial insects and other wildlife in ecosystems.

References:

Armbruster, W. Scott, Sarah A. Corbet, Aidan J. M. Vey, Ahu-Juan Liu, and Suang-Quan Huang, 2014.  In the right place at the right time: Parnassia resolves the herkogamy dilemma by accurate repositioning of stamens and stigmas, Annals of Botany, 113:97-103. doi:10.1093/aob/mct261

Bennington,Cindy and Peter May, 2016.  Volusia Sandhill Pollinator Project https://www.stetson.edu/other/gillespie-museum/vse/pollinator-project.php

In the summer of 2016, Professors Bennington and May recorded more than 1300 insect visitors to two plant species (Spanish needles, Bidens alba, and Partridge pea, Chamaecrista fasciculata) in two sites. In both sites, flowers of Spanish needles, attractive to generalist insect pollinators, were visited by a variety of bees, wasps, flies, butterflies and beetles. 

Bennington, Cynthia . and Peter May, 2020.  Pollinator Communities of Restored Sandhills: a Comparison of Insect Visitation Rates to Generalist and Specialist Flowering Plants in Sandhill Ecosystems of Central Florida April 2020 Natural Areas Journal 40(2):168 DOI: 10.3375/043.040.0208    This article, in the April 2020 issue of Natural Areas Journal summarizes Bennington and May’s findings, based on research from three years (2016-18), assessing the ability of the Volusia Sandhill Ecoystem to support wild insect pollinators. Comparing visitation to flowering plants in two sites—our campus urban restoration and a sandhill site at nearby Heart Island Conservation Area—they found that total insect visitation rates were not significantly different between years or sites, suggesting that even a small urban fragment is capable of maintaining abundant pollinators.  Check out the abstract of their article, “Pollinator Communities of Restored Sandhills: a Comparison of Insect Visitation Rates to Generalist and Specialist Flowering Plants in Sandhill Ecosystems of Central Florida.”

 Bennington has created a how-to video on how you can help support pollinators by incorporating native plants into your own yard or garden: How-to video on Attracting Pollinators with Native Florida Wildflowers.

Blackwell, Amy Hackney and Patrick D. McMillan, 2013. Collected in South Carolina 1704-1707: The Plants of Joseph Lord, Phytoneuron 2013-59L 1-15. http://4.namethatplant.net/PDFs/PhytoN-JosephLord.pdf

Bridges, E.L. and S.L. Orzell. 2024. Systematics of the unifoliolate Floridian Lupinus clade (Leguminosae: Papilionoideae). Phytoneuron 2024-04: 1–61. Published 15 January 2024. ISSN 2153 733X  https://www.phytoneuron.net/wp-content/uploads/2024/01/04-PhytoN-UnifoliolateLupinus.pdf, also see FNPS Blog https://fnpsblog.blogspot.com/2024/04/study-names-three-new-species-of.html

Clewell, Andre F.  1985.  Guide to the Vascular Plants of the Florida Panhandle, University Presses of Florida, Tallahassee.

Coogan, Kenny, 2022. Florida’s Carnivorous Plants, Pineapple Press, Palm Beach.

Evans, Margaret E. K., Eric S. Menges, and Daaria R. Gordon, 2003.  Reproductive biology of three sympatric endangered plants endemic to Florida scrub, Biological Conservation, 111: 325-246.

Evans,  Margaret E. K., Eric S. Menges, and Doria R. Gordon, 2004.  Mating systems and limits to seed production in two Dicerandra mints endemic to Florida scrub, Biodiversity & Conservation. 13, 1819–1832 (2004).     https://link.springer.com/article/10.1023/B:BIOC.0000035869.12388.0f

Fishman, Gail, 2000.  Journeys through Paradise – Pioneering Naturalists in the Southeast, University Press of Florida.

Gann, George D., Keith A. Bradley, and Steven W. Woodmansee, (2002) 2016 EV.  Rare Plants of South Florida: Their History, Conservation, and Restoration.  The Institute for Regional Conservation.  https://www.regionalconservation.org/ircs/pdf/Gann_et_al._2002.pdf

Gilbert, Katherine M., John D. Tobe, Richard W, Cantrell, Maynard E. Sweeley, and James R. Cooper, 1995. The Florida Wetlands Delineation Manual, Department of Environmental Protection. https://floridadep.gov/sites/default/files/delineationmanual.pdf

Godfrey, Robert K., Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia and Alabama, The University of Georgia Press, Athens.

Hall, David W. 2020.  Illustrated Plants of Florida and the Coastal Plain, 2n Ed., Univ. of Florida Press, Gainesville.

Hall, David, William J. Weber, and Jason H. Byrd, 2010. Wildflowers of Florida and the Southeast, Self-published by Hall and Byrd.   ISBN 978-0-615-39502-9 s

Hammer, Roger L., 2018.  Complete Guide to Florida Wildflowers, Falcon Guides, Guilford, Connecticut.

Hiatt, Drew and S. Luke Flory, 2020. Populations of a widespread invader and co-occurring native species vary in phenotypic plasticity, New Phytologist 225; 584-594. https://doi.org/10.1111/nph.16225

Houlihan, Peter R., Mac Stone, Shawn E Clem, Mike Owen, Thomas C Emmel, 2019.  

Pollination ecology of the ghost orchid (Dendrophylax lindenii): A first description with new hypotheses for Darwin’s orchids.  Sci Rep. 2019 Sep 6;9:12850. doi: 10.1038/s41598-019-49387-4  PMCID: PMC6731287  PMID: 31492938

Hoyer, Mark V., Daniel E. Canfield, Jr., Christine A. Horsburgh, and Karen Brown, 1993.  Florida Freshwater Plants – A Handbook of Common Aquatic Plants in Florida Lakes, University of Florida Institute of Food and Agricultural Sciences.

Kalaman, Heather, Rachel E. Mallinger, Gary W. Knox, Kim Taehoon, Kevin Begcy, and Edward van Santen, 2021.  Evaluation of Native and Nonnative Ornamentals as Pollinator Plants in Florida: II. Floral Resource Value HortScience 57(1):137-143. Department of Environmental Horticulture, IFAS, University of Florida, P.O. Box 110670, Gainesville, FL 32611  Online Publication Date: 21 Dec 2021  Volume/Issue: Volume 57: Issue 1  DOI: https://doi.org/10.21273/HORTSCI16124-21

Kimball, John W., 1997. Unit 16 – The Anatomy and Physiology of Plants, Libre Texts.  https://bio.libretexts.org/Bookshelves/Introductory_and_General_Biology/Biology_(Kimball)/16:_The_Anatomy_and_Physiology_of_Plants/16.03:_Reproduction_in_Plants/16.3E:_Self-incompatibility_-_How_Plants_Avoid_Inbreeding)

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Musselman, Lytton J. and  William F. Mann, Jr, 1979.  Agalinis fasciculata (Scrophulariaceae) A Native Parasitic Weed on Commercial Tree Species in the Southeastern United States, American Midland Naturalist 101(2): 4589-464      https://www.jstor.org/stable/2424616

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Pitts-Singer, T.L., Hanula J.L. and J.L. Walker (2002) Insect pollinators of three rare plants in a Florida longleaf pine forest. Florida Entomologist 85(2): 308-316.   https://www.srs.fs.usda.gov/pubs/ja/ja_pitts-singer001.pdf

Raven, Peter – Evert, Ray F. and Susan E. Eichorn, 2013. Biology of Plants, W. H. Freeman and Company.

Simons, Robert W., 2021.  The Ecology of Trees, Shrubs, and Woody Vines of Northern Florida, Univ. of Florida Press, Gainesville.

Small, John Kunkell, 1929 (2004 reprint with editorial Preface by Bill Bleville, et al). From Eden to Sahara – Florida’s Tragedy., Seminole Soil & Water Conservation District, Sanford, FL

Sommers, Kristen Penny, Michael Elswick, Gabriel I. Herrick, and Gordon A Fox, 2011,  Inferring microhabitat preferencs of Lilium catesbaei (Liliaceae), American Journal of Botany, 98(5):819-828  https://doi.org/10.3732/ajb.1000250

Whitewoods, Christopher D., Beatriz Gonçalves, Jie Cheng, Minlong Cui, Richard Kennaway, Karen Lee,  Claire Bushell, Man Yu, Chunlan Piao,  and Enrico Coen, (3 January 2020). “Evolution of carnivorous traps from planar leaves through simple shifts in gene expression”. Science. 367 (6473): 91–96. Bibcode:2020Sci…367…91W. doi:10.1126/science.aay5433. ISSN 1095-9203. PMID 31753850. S2CID 208229594

Wunderlin, Richard P. and Bruce F. Hansen, 2011.  Guide to the Vascular Plants of Florida, 3rd ed., The University Press of Florida, Gainesville.

Wunderlin, Richard P., Bruce F. Hansen, and John Beckner, 2000.  Botanical Exploration in Florida, in Flora of Florida, Vol. 1, pp. 35-99.  University of Florida Press.

Taylor, Walter Kingsley, 2013.  Florida Wildflowers – A Comprehensive Guide, University Press of Florida, Gainesville.

Yunpeng Liu, Xioting Xu, Dimitrt Dimitrov, Carsten Rahbek and Zhiheng Wang have retrofitted Takhtajan’s system, recognizing 8 regions grouped as either Gondwanan or Laurasian realms.  https://en.wikipedia.org/wiki/Phytochorion  https://www.nature.com/articles/s41467-024-47544-6

BLOGS & Websites:

Butterfly Host Plants for Southeast Florida, https://miamiblue.org/plantlist/

Heather Holm – a resource for publications and events related to insect pollinators:  https://www.pollinatorsnativeplants.com/

Link to this Page that you can share – Notes on Florida Wildflowers: https://botanyincontext.com/notes-on-florida-wildflowers-text-only/

Additional References:

Amasino, Richard M., 2013. My favourite flowering image: Maryland Mammoth tobacco., Journal of Experimental Botany 64: 5817-5818. https://doi.org/10.1093/jxb/ert083

Barrett, Spencer C. H., 1998.  The evolution of mating strategies in flowering plants, Trends in Plant Science 3(5): 335-341 https://doi.org/10.1016/S1360-1385(98)01299-0

Barthélémy, Daniel and Yves Caraglio, 2007.  Plant Architecture: A Dynamic, Multilevel and Comprehensive Approach to Plant Form, Structure and Ontogeny; Annals of Botany, 99(3): 375–407, https://doi.org/10.1093/aob/mcl260

de Craene, Louis P. Ronse, 2010.  Floral Diagrams – An Aid to Understanding Flower Morphology and Evolution, Cambridge Press

Diaz-Toribio, Milton and F.E. “Jack” Putz, 2022. Time to Dig: Underground Storage Organs of Plants in Fire-Maintained pine Savannas, Palmetto 38:10-12.

Endress, Peter K, 2010.  Disentangling confusions in inflorescence morphology: Patterns and diversity of reproductive shoot ramification in angiosperms, Journal of Systematics and Evolution  Free access:   https://doi.org/10.1111/j.1759-6831.2010.00087.x

Gest, Howard, 2002.  History of the word photosynthesis and evvolution of its definition, Photosynthesis Research 73:7-10.  https://www.life.illinois.edu/govindjee/Part1/Part1_Gest.pdf

Graça, José, 2015. Suberin: the biopolyester at the frontier of plants, Frontiers Chemistry vol 3. https://doi.org/10.3389/fchem.2015.00062

Gundersen, Alfred, 1918. A Sketch of Plant Classification History, Torreya 18: 213-219 https://www.jstor.org/stable/40595853

Ingen-Housz, John, 1779. Experiments upon Vegetables, Discovering Their great Power of purifying the Common Air in the Sun-Shine…., P. Elsly, in the Strand. https://library.si.edu/digital-library/book/experimentsuponv00inge

Judd, Walter S., Christopher S. Campbell, Elizabeth A. Kellogg, Peter F. Stevens, Michael J. Donoghue, 2016. Plant Systematics – A Phylogenetic Approach, 4th ed. electronic ed. Sinauer Associates, Sanderland.

Liu, Hong, Robert W. Pemberton, and Peter Stiling, 2006..  Native and introduced pollinators promote a self-incompatible invasive woody vine (Paederia foetida L.) in Florida (Rubiaceae)  J Torrey Botanical Society 133(2): 304-311  https://www.jstor.org/stable/20063841

Ma, Suhui Feng He, Di Tian, Dongting Zou, Zhengbing Yan, Yulong Yang, Tiancheng Zhou, Kaiyue Huang, Haihua Shen, and Jingyun Fang , 2018.  Variations and determinants of carbon content in plants: a global synthesis, Biogeosciences, 15, 693–702, 2018  Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China  https://doi.org/10.5194/bg-15-693-2018

Norman, Jaimie M Van , Wei Xuan, Tom Beeckman, Philip N Benfey, 2013. To branch or not to branch: the role of pre-patterning in lateral root formation. PubMed 140(21):4301-10. PMID: 24130327 PMCID: PMC4007709 DOI: 10.1242/dev.090548

Noyori, Wataru, Yuji Isagi, Naoto Nakamura, Shuichiro Tagane, Gerardo Celis, and Kaoru Kitajima, 2024.  Phylogeography of apomictic and outcrossing individuals in invasive and native populations of Ardisia crenata (Primulaceae), Plant Species Biology, Early View.  https://doi.org/10.1111/1442-1984.12482

Silva, N.F. and D. R. Goring, 2001.  Mechanisms of self-incompatibility in flowering plants, Cellular Molecular Life Science (CMLS) 58:1988-2007.  https://pmc.ncbi.nlm.nih.gov/articles/PMC11337325/pdf/18_2001_Article_CMLS_101114.pdf

Spies, Thomas A., 1998.  Forest Structure; A Key to the Ecosystem, Northwest Science 77, Special Issue 2.  https://andrewsforest.oregonstate.edu/sites/default/files/lter/pubs/pdf/pub2564.pdf

Taiz, Lincoln, Ian Max Moller, Angus Murphy, and Wendy A. Peer, 2018. Fundamentals of Plant Physiology, Enhanced E-book, Oxford University Press,

Zuntini, A.R., Carruthers, T., Maurin, O. et al.  2024. Phylogenomics and the rise of the angiosperms, Nature (open access) 629:843-850. https://www.nature.com/articles/s41586-024-07324-0

The Angiosperm Phylogeny Group,  M. W. Chase,  M. J. M. Christenhusz,  M. F. Fay, J. W. Byng,  W. S. Judd,  D. E. Soltis,  D. J. Mabberley,  A. N. Sennikov,  P. S. Soltis, et al., 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV, Botanical Journal of the Linnean Society, Volume 181, Issue 1, May 2016, Pages 1–20, https://doi.org/10.1111/boj.12385

Citations to Research on Native Families & Genera:

CELASTRACEAE (including PARNASSIACEAE): Parnassia: Armbruster, W. Scott, Sarah A. Corbet, Aidan J. M. Vey, Ahu-Juan Liu, and Suang-Quan Huang, 2014.  In the right place at the right time: Parnassia resolves the herkogamy dilemma by accurate repositioning of stamens and stigmas. See References for complete citation.

DROSERACEAE

Drosera: Brewer, J.S. 1998. The effects of competition and litter on a carnivorous plant, Drosera capillaris (Droseraceae). Amer. J. Bot. 85:1592–1596; Krupa, J.J., K.R. Hopper & M.A. Nguyen, 2021. Dependence of the dwarf sundew (Drosera brevifolia) on burrowing crayfishdisturbance. Pl. Ecol. 222:459–467.

FABACEAE:

Lupinus: Bridges, E.L. and S.L. Orzell, 2024. Systematics of the unifoliolate Floridian Lupinus clade (Leguminosae: Papilionoideae). See References for entire citation.

GENTIANACEAE:

Bartonia: Clotir, Claudia, Chris Yesson, and Joanna Freeland, 2013. The evolutionary history and conservation value of disjunct Bartonia paniculata subsp. paniculata (Branched Bartonia) populations in Canada, Botany, https://doi.org/10.1139/cjb-2013-0063; Gillett, John M., 1959. A Revision of Bartonia and Obolaria (Gentianaceae), Rhodora 61 : 43-62. https://www.jstor.org/stable/23306375 Mathews, Katherine G., Niall Dunne, Emily York,and Lena Struwe, 2009. A Phylogenetic Analysis and Taxonomic Revision of Bartonia (Gentianaceae: Gentianeae), Based on Molecular and Morphological Evidence, Systematic Botany 34: 162-172. https://www.wcu.edu/WebFiles/biology_Bartonia_Mathews-etal_Systbot.pdf

HYDROLEACEAE:

Hydrolea: Erbar, C., S. Porembski, and P. Leins, 2005. Contributions to the Systematic Position of Hydrolea (Hydroleaceae) based on floral development, Plant Systematics and Evolution 252:71-83; Davenport, L. J., 1988. A monograph of Hydrolea (Hydrophyllaceae), Rhodora 90 A#862.

LAMIACEAE: Evans,  Margaret E. K., Eric S. Menges, and Doria R. Gordon, 2004.  Mating systems and limits to seed production in two Dicerandra mints endemic to Florida scrub. See References for entire citation.

Physostegia: Cantino, P. D., 1980. The Systematics and Evolution of the Genus Physotegia, Harvard University Herbaria; Cantino, P.D., 1982. A monograph of the genus Physotegia (Labiatae). Harvard University Herbaria, http://www.jstor.org/stable/41764739; Lersten Nels R. and John D. Curtis, 1998. Foliar idioblasts in Physostegia virginiana (Lamiaceae), Journal Torrey Botanical Society 124: 133-137. https://www.jstor.org/stable/2997300 https://doi.org/10.2307/2997300 Kelly, John W. and Terri W. Starman, 1990. Postharvest Handling of Physostegia purpurea Cut Flowers, HortScience 25: 552-553 https://www.google.com/url?sa=t&source=web&rct=j&opi=89978449&url=https://journals.ashs.org/hortsci/downloadpdf/journals/hortsci/25/5/article-p552.pdf&ved=2ahUKEwiS24jN0KSKAxWJSTABHVDKAWkQFnoECBUQAQ&usg=AOvVaw38Gq47Z6bY9_U6_t2GlmLc; Middleton, Beth and Casey R, Williams, 2023. Seed Banks of Rare Physostegia correllii (Lamiaceae) in Lady Bird Lake, Austin, Texas, U.S.A., Journal of teh Botanical Research Institute of Texas, 17: 363-368, https://doi.org/10.5066/P9Z2KNGL.

NYCTAGINACEAE: Goodson, J.J.* and P.S. Williamson. 2011. Germination of seeds in the endangered Abronia macrocarpa. Southwestern Naturalist 56:141-146.

NYMPHAEACEAE:

Nuphar: Lippok, B., A.A. Gardine*, P.S. Williamson, and S.S. Renner. 2000. Pollination by flies, bees, and beetles of Nuphar ozarkana and N. advena (Nymphaeaceae). American Journal of Botany 87:898-902.; Schneider, E.L., S.C. Tucker, and P.S. Williamson. 2003. Floral development in the Nymphaeales. International Journal of Plant Science 164:5279-5292; Moseley, M.F., E.L. Schneider, and P.S. Williamson. 1993. Phylogenetic interpretations fromselected floral vasculature characters in the Nymphaeaceae, sensu lato. Aquatic Botany 44:325-342.

OROBANCHACEAE:

Agalinus: Musselman, Lytton J. and  William F. Mann, Jr, 1979.  Agalinis fasciculata (Scrophulariaceae) A Native Parasitic Weed on Commercial Tree Species in the Southeastern United States

SOLANACEAE: Zhang, Jingbo, Peter F. Stevens, and Wenheng Zhang, 2022. Evolution and development of inflorescences and floral symmetry in Solanaceae, Americdan Journal of Botany 109:746-767 https://doi.org/10.1002/ajb2.1864

Link to this Page: https://botanyincontext.com/notes-on-florida-wildflowers-text-only/