It takes two flowers to make a squash

June 8, 2012

A summer squash plant with both pistillate and staminate flowers. This is a yellow squash, as you can see from the ovaries of the pistillate flowers.

Squashes, melons, pumpkins, cucumbers, and gourds all belong to the squash family, Cucurbitaceae. There is a common pattern of the flowers that children enjoy finding, and that often escapes adults. I’ve pointed it out to many long-time gardeners, who hadn’t noticed it before.

This is a young pistillate flower of a patty pan squash. The ovary is green now, but it will turn white as this squash matures.

Most members of this family are monoecious, which means each plant has flowers with only stamens along with other flowers which are only pistillate. These are commonly called male and female flowers. They are easy to tell apart if you look beneath the corolla. The ovary is inferior (located beneath the other flower parts) or, to put it another way, the other flower parts are epigenous (they sit on top the ovary).

 If you want to find the pistillate (aka female) flowers, just look for a tiny ovary – a baby squash, cucumber, etc. – on the stem under the corolla. You can find the little ovaries well before the flowers open, so it is easy to see which flowers will produce the desired fruit. The mature ovary of a flowering plant is a fruit, so to a botanist, squashes, cucumbers, and melons are all fruits.

The staminate flowers of the squash family have a plain stem beneath the corolla.

The staminate flowers have a plain stem beneath their corolla. Inside the filaments and anthers of their stamens are joined together into a knob-like structure that resembles a pistil. Inside the pistillate flower’s corolla, you can see the three-carpellate structure of the pistil. There are three stigma lobes that have two branches each. The fruit shows the three carpels as well. Look at a cross section of a squash or fruit of other family members to see this.

This is a staminate squash flower that has been split along the corolla and opened to show the fused anthers and filaments of the stamens.

The stigmas of the pistillate flower have several lobes. This flower had bloomed, and its corolla was removed to show the stigmas.

The next question that comes up is often “Why doesn’t my squash plant produce more squashes?” Sometimes the temperature is to blame. It affects the sex of squash flowers in ways that aren’t always obvious. When I lived in the mountains of Colorado, I found that although zucchini plants would grow, they seldom produced fruits. The plants would form female flowers, but seldom have staminate ones, so pollination didn’t happen. The cold soil temperatures were to blame. With other members of this family, cold temperatures cause only staminate flowers to form. You can read more about this on the website of the Ontario Ministry of Agriculture,

Conversely, temperatures above 95 degrees F can also cause flowers to drop instead of developing. There could be a number of factors operating in this case, including moisture stress.

Although squashes and begonias don’t commonly come to mind as relatives, if you look at the flowers of a begonia, you can see the same pattern – monoecious plants with inferior ovaries. The begonia family and the squash family both belong to the squash order, Cucurbitales.

In this view of begonia flowers, the staminate flower is on the top. It has a plain stem. The pistillate flower below has a green, winged ovary.

A front view of begonia flowers. The pistillate flower is on the left. The staminate flower has four tepals; the pistillate has five.


The leaves that didn’t fall in fall

March 14, 2011

Most deciduous trees live up their name and drop their leaves in autumn, but you may have found a curious exception. Some trees or shrubs have leaves that turn brown, but do not fall off. Members of the oak family often exhibit this phenomenon. The leaves that didn’t fall in fall are called marcescent leaves. The term comes from a Latin root that means “withering.”

This Gambel's oak hadn't dropped its leaves in November.

I observed marcescent leaves on the Gambel’s oaks (Quercus gambelii) this past fall. Nearby members of the same species had shed most of their leaves. What survival advantage does this give the plant? Whenever a genetic trait is commonly present, it probably confers some survival advantage. For the oak shrubs along the foothills of the Colorado Rockies, there is a strong threat to survival in the form of mule deer. Cute little bambi grows up to be death to plants on four hooves.

The marcescent leaves on the oaks could help protect them from the onslaught of the large herbivores. In winter, deer are browsers. They eat twigs, buds, and I have even seen them eat the foliage of conifer branches that have blown off the trees or been broken off by heavy snow. The marcescent leaves are thought to deter browsing, probably because they are not tasty and have little food value. The growing points, the meristems, hide behind these dry leaves, protected against drying by their bud scales. In the spring, the old leaves will fall as the new buds start to grow.

This oak, near the first one, had lost most of its leaves by the same time.

However, a study in Denmark showed that the deer there avoided beech and hornbeam branches with marcescent leaves, but not oaks. The dead leaves on the beech and hornbeam branches had a high lignin content. Lignin is a very complex substance that is hard to digest. Perhaps Colorado oaks have less palatable leaves, or the leaves could give a different benefit.

He's cute, but he is murder on plants. Twigs on trees and shrubs are among his favorite foods.

Another theory is that the leaves help hold more wind-blown snow, and therefore bring more moisture to the shrub. In this semi-arid climate, every bit of moisture is important. While this may be a part of why Colorado scrub oaks keep their leaves, it doesn’t really fit the picture for taller oak trees.

Some oak species, notably pin oaks (Quercus palustris) keep their petioles alive through the winter and shed their leaves in spring. In other cases, the petiole appears to be brown and dead. Another factor could be sudden fall freezes that kill the petiole before it has completed the abscission process. There are many steps to abscission, the “cutting away” of the leaves. They involve enzymes, and these protein molecules, as well as cell structures, may be damaged by sudden cold.

It is likely that many factors play into the puzzle of the leaves that didn’t fall in fall. They will, however, fall in spring.

A Rare Plant Treat

July 6, 2010

I was in Seattle recently and enjoyed a visit to the greenhouses at the University of Washington. I was delighted to find a collection of Welwitschia plants there, one of which was forming cones. This bizarre plant of the Namibian Desert belongs to the gnetophyte lineage of seed plants. It is one of three wildly different branches of this clade.

Welwitschia plants in the UW greenhouse have tall pots to accommodate their long tap roots.

The gnetophytes are one of five extant lineages of seed plants. The others are the cycads, the ginkgo, conifers, and the angiosperms, aka flowering plants. The angiosperms greatly outnumber the others, but that has become the case in the Cenozoic Era. In the Mesozoic Era, there were many other types of gnetophytes, but presently the three lineages are the genus Ephedra, genus Gnetum, and genus Welwitschia.

Although there are about 60 species of Ephedra and about 30 of Gnetum, there is only one species of Welwitschia, W. mirabilis. Ephedra species are native to arid environments in North and South America, Africa, and Eurasia. They have greatly reduced leaves and look like a bush of tough, narrow stems. Gnetum is the odd one on environment. It grows in the tropics of Indonesia, the Philippines, and parts of Africa. Most of its members are vines, although a few are trees or shrubs. Their leaves look very much like those of angiosperms.

Ephedra, a relative of Welwitschia, growing in Utah. The leaves of this bush are reduced to tiny scales.

Gnetophytes were once thought to be closely related to flowering plants, but the DNA told a different story. They are now considered to be more closely related to the conifers. Characteristics that all gnetophytes share include opposite leaves and having staminate and ovulate reproductive structures on separate plants, i.e. they are dioecious.

Welwitschia grows only two leaves, not counting its two seed leaves, in its whole life time. The leaves arise from a woody stem that has a sort of upside down cone shape. The plants are estimated to live about 1000 years, judging from their growth rates and the length of the leaves on wild plants. The leaves grow from their bases and often split so that it isn’t easy to see that there are only two.

The stem of Welwitschia is woody and has a roughly inverse cone-shape.

The two leaves of this Welwitschia plant are split into several strips. The top of its woody stem is visible.

One of the plants in the UW greenhouse had formed cones. It is a male plant, and there are stamens showing near the bottom bracts. The branch that bears the cones grows from the top of the woody stem, near the base of the leaves. I had read about their cones, but never seen them “in person” before, so finding them was a real treat. 

These are staminate cones of Welwitschia. Their stalk grows from the top of the woody stem.

What color is your Doug fir?

June 13, 2010

When you read the title of this post, did you say to yourself “Well, it’s green, of course, that is if it is alive.” Doug firs (Douglas fir, Pseudotsuga menziesii) have some parts that aren’t green, however, and those parts are a visual confirmation of an important property of life – genetic variation in a population.

If you look at the branch tips of a Douglas fir early in the spring, you may notice its cones. This is provided that it is making cones that year. These trees don’t make cones every year. Each year some will form a few cones, but in a cone year, almost all the trees in an area grow cones at the same time. I’m not sure how they manage this trick, but it is a good one for a wind-pollinated species.

The bright pink structures are the ovulate cones of this Doug fir. The pollen cones are the smaller ones below.

Like other conifers, Doug firs have pollen cones and ovulate (aka seed) cones. Both pollen and seed cones grow on the same tree – the species is monoecious. I’ve noticed that the Doug firs where I live have different colors of cones. Some of my trees have deep rose ovulate cones, while others have lemon yellow ones. The pollen cones on the tree match the color of the ovulate cones, but they aren’t quite as intensely colored.

This Doug fir has yellow cones. The ovulate cone is on the upper right.


As the ovulate cones develop, their characteristic three-pointed bracts protrude from between the cone scales. Most Doug firs around here have straight bracts, but some have curly bracts. The straight or curly feature remains after the cones have dried and fallen from the tree.

Why should Doug firs have different colors of cones and different types of bracts? I certainly can’t tell you the exact advantage, but I do know that variations in every population are important. They are that species’ library of solutions to life’s problems. Life tinkers – tries this and that. Some traits may work better for some situations, while others may be an advantage in different conditions.

Developing Doug fir cone with curly bracts.

Most Doug firs have cones with straight bracts, like this one. The photos of this one and the curly bract one were taken on the same day.

No organism can predict the future, so the best survival strategy is to have lots of genetic diversity. This is the reason for cross pollination and sexual reproduction. The species that shuffles its genes and deals them out in all combinations has the best change of continuing on, whatever the environmental conditions.

Certainly there are some species that don’t seem to change, at least on the surface, but they just change more slowly. I prefer the term “slow-evolving species” to “living fossil” because the latter gives the false impression that the species doesn’t have variations and doesn’t change. Today’s ginkgoes are not the same as the Triassic ones, even if the leaves look the same.

When you are out in the field with your children, point out the variations that you can see. There may be an albino among a stand of blue flowers or some that have different shades of color. These are outward manifestations of genetic diversity in the population. Many more traits that we can’t see have variations in a natural population, and that’s a good thing for long term survival.  Hurray for being different – diversity is an important characteristic of life.

This rose-colored sugarbowl or leather flower (Clematis hirsutissima) is very unusual.

This purple is the usual color for sugarbowls. It was growing near the pink one.

Botany marches on – Part 1, basal angiosperms and monocots

April 13, 2010

Warning! I’m going to get into some detailed botany here, so if you are not deeply into the flowering plants and their classification and you do not own my book, A Tour of the Flowering Plants, you may not wish to wade through all this. It is basically for advanced elementary and secondary Montessori botany studies and for anyone else that owns the book. A Tour of the Flowering Plants is still quite useful to show the appearance of angiosperm families, whether or not you are concerned about the latest classification.

Last fall, the Angiosperm Phylogeny Group published a third report concerning the classification of the orders and families of flowering plants. The report, published in the Botanical Journal of the Linnean Society, is called APG III. There is a summary of APG III on Wikipedia. My book, A Tour of the Flowering Plants, is based on the 2003 report from this group, which is called APG II. The book has some further advances that were published on the Angiosperm Phylogeny Website of Peter Stevens.

I’ll go over the APG III changes, beginning with this post. If you own my book, you can decide if you want to add notes to it. You may contact me via my website ( if you would like a list of the pages and changes for bringing A Tour of the Flowering Plants in line with APG III.

Will the changes keep coming over the years as we acquire more and more DNA data? I think that future changes will be modest and will not affect the general structure of the angiosperm tree. There aren’t that many unplaced groups left. There will likely be little surprises, like one I give below for Nymphaeales.   

For now, let’s start with the first branches of the angiosperms, grouped as the basal angiosperms in A Tour of the Flowering Plants. The Nymphaeales got another family, Hydatellaceae, which was formerly placed in the grasses. This shows how much DNA studies can reveal, and what studies of morphology may not be able to distinguish. This family is tiny, both in size and numbers of species. The only reason I mention it is the idea that when plants adapt to living in water (or any other extreme environment), they often change form so much that they don’t resemble even their closer relatives. You can see the plant here:

The order Chloranthales was previously unplaced – no one was sure what its closest relatives are. Now it is considered to be a sister group to the magnoliids. The magnoliids themselves have not changed in APG III. This branch of the flowering plants includes the laurel and black pepper families, as well as the magnolias. Although their seeds have two cotyledons, they are not closely related to the eudicots. The eudicots are the traditional dicots minus the magnoliids and the basal angiosperm lineages.

In A Tour of the Flowering Plants, I used terms for branches of monocots that have since disappeared. You don’t have to worry about whether to call the Liliales and Asparagales “lilioid monocots” or “petaloid monocots.” Just call them monocots and go on. The only lineage of monocots that gets a special name now is the commelinids. “Lilioid” and “petaloid” should be understood as informal terms that refer to plants that were traditionally lumped in the lily family. Most of them have large, showy tepals.

The major change in the monocot is the grouping of several small families as subfamilies under the enlarged families Amaryllidaceae, Asparagaceae, and Xanthorrhoeaceae. This is all within the order Asparagales. Here’s the breakdown:

The enlarged Amaryllidaceae has the agapanthus subfamily (Agapanthoideae, equivalent to the former Agapanthaceae), the onion subfamily (Alliodeae, equivalent to the former Alliaceae), as well as the amaryllis subfamily (Amaryllidoideae, equivalent to the former Amaryllidaceae). The members of this enlarged family have their flowers in umbels that are enclosed by two bracts when the flowers are in bud.

The enlarged Asparagaceae is really big. It has the Brodiaea subfamily (Brodiaeoideae), the scilla subfamily (Scilloideae, which includes the former hyacinth family, as the tribe Hyacintheae), the agave subfamily (Agavoideae, which includes the former Agavaceae), the Nolina subfamily (Nolinoideae, equivalent to the former Ruscaceae), as well as the asparagus subfamily (Asparagoideae). Yet another subfamily holds several Australasian species such as the cabbage tree, Cordyline.  

Asparagaceae members have flowers in racemes or in umbels that have three or more bracts at their base. The umbels, if present, do not have the pair of enclosing bracts seen in Amaryllidaceae.

The enlarged Xanthorrhoeaceae (the grass tree family) includes the daylily subfamily (Hemerocallidoideae) and the asphodel subfamily (Asphodeloideae), as well as the grass-tree subfamily. I did not include the grass tree family in A Tour of the Flowering Plants because it is native to Australia and not commonly used in North American landscaping.

The cattail family, Typhaceae, got a second genus, Sparganium, the bur-reeds. It’s not hard to see these two aquatics as relatives.

I’ll address the APG III changes to the eudicots in another post.

Pollen cones

April 13, 2010

I was so concerned with getting the photos of the seed cones (ovulate cones) into my last post that I forgot to show the pollen cones of the ponderosa pine. Here are a couple of views.

The young pollen cones of the ponderosa pine.


One pink seed cone and many pollen cones ready to release their pollen.

A pine cone tale

April 7, 2010

A major goal of Montessori botany studies is to help children learn to observe and understand plant structures. There are a lot of things going on in the plant world that take a sharp eye and careful observation to find. The life cycle of pines is one of them. It is important for the teacher/guide to show children inconspicuous plant structures such as pine cones throughout the year and explain to them what is happening.

Most people are familiar with conifer cones, although they tend to call all of them “pine cones.” Few have followed the development of a cone through the year – or two years in the case of pines – that it takes for a cone to mature. I have been photographing the development of pine cones and here’s a look at their life cycle.

Pines have two kinds of cones on the same tree, pollen cones and seed cones. The latter are formally called ovulate cones. The trees don’t usually form cones every year. In cone years, the cycle begins as the new shoots elongate in the spring. The seed cones form at the end of the new growth. They look like tiny pink-to-purple bristles.

These are young seed (ovulate) cones on the new shoot of a ponderosa pine.

The pollen cones cluster at the base of the new shoots, beneath the terminal bud. Most of the pollen cones form on the lower branches of the tree, away from the seed cones, but sometimes they form on the same shoot as the seed cones. The wind usually won’t take pollen from the base of the tree to its upper branches, so the arrangement of seed and pollen cones encourages cross-pollination. 

These two cones formed in the previous spring. The one on the right died during the winter. The left one is starting to grow.


By early July, the living seed cone has quadrupled in size. Its scales are noticeably green.

Conifers use wind pollination, which requires a lot of pollen to work, and in cone years the trees produce an amazing amount of pollen. Pollen cones tremendously outnumber seed cones. After they release their pollen, most of the spent pollen cones drop off the tree. You can sometimes find dried ones in the branches later in the summer, however.

It takes careful observation to find the budding ovulate cones, even though they can be colorful. They hide among the new needles and are most easily seen from above, the bird’s eye view we don’t usually have. It doesn’t help that the ovulate cones usually form on the higher branches. You may need to pull a branch down so that the children can see the tiny new cones. The little cones of pines don’t grow much during their first year. In the fall, they have become browner and drier looking, but are nearly the same size as they were in the spring.

In the second spring the pine seed cones rapidly enlarge. A shoot I photographed had a pair of seed cones, but one of them had died. It provides a size scale to show how much the live cone has grown. Fertilization is a slow process in pines. It takes about 15 months for the egg cells to form and the pollen tube to grow and deliver the sperm to the eggs. The scales of the seed cones are green until late fall. By that time the seeds are mature. The cone dries and the scales spread apart, releasing the winged seeds. The dried cone may remain on the tree for months or years, until a strong wind brings it down.

In the fall, the seed cone has dried. Its brown scales spread apart and the seeds are released.

In case you need help finding your local pines, look for a conifer tree with needles in bundles of two to five. Other conifers, such as firs or spruces bear their needles singly. Their cones mature in one year, but they can be even harder to see because the seed cones form in the tops of the tree.

Take a look around this spring and see if you can locate some young cones to show your children and to follow through the cone life cycle.

The winged seeds of the pines blend in with the soil and rocks very well.