Awkward Botanical Sketches #5: Leaves of Yellowstone Edition

Earlier this month, I met up with Eric LoPresti and others at Yellowstone National Park to help take a census of Abronia ammophila, a rare plant endemic to the park and commonly referred to as Yellowstone sand verbena. Abronia (a.k.a. the sand-verbenas) is a small genus of plants in the family Nyctaginaceae that is native to western North America. Several species in the genus have fairly limited distributions, and as the common name implies, members of this genus generally occur in sandy soils. A. ammophila is no exception. A report written by Jennifer Whipple and published in 2002 described it as “restricted to stabilized sandy sites that lie primarily just above the maximum splash zone along the shoreline of [Yellowstone Lake].” Despite the large size of the lake, A. ammophila is not widespread. Most individuals are found along the north shore of the lake, and even there it has been declining. According to Whipple’s report, “Yellowstone sand verbena has been extirpated from a significant portion of its original range along the shoreline of the lake due largely to human influences.”

Like other sand verbenas, A. ammophila has sticky leaves to which sand particles easily adhere, a phenomenon known as psammophory and an act that may help in defense against herbivory. The plant grows prostrate across the sand and produces attractive, small, white, trumpet-shaped flowers in groups of up to 20 that open wide when light levels are low, such as in the evening and in times of heavy cloud cover. The flowers are self-fertile, but insects may also play a role in pollination. It is imperative that questions surrounding its pollination biology, seed dispersal, and other factors regarding its life history are answered in order to halt any further decline of the species and ensure its survival for generations to come.

While in Yellowstone, I enjoyed looking at the all plants, several of which were new to me. I decided to sketch a few of the leaves that I found common around our campsite. I was particularly interested in discolored, diseased, drought-stressed, and chewed-on leaves, since they are more interesting to sketch and color. While I was at it, I attempted to draw a Yellowstone sand verbena seedling as well.

wild strawberry (Fragaria sp.)
Richardson’s geranium (Geranium richardsonii)
lodgepole pine (Pinus contorta)
veiny dock (Rumex venosus)
cinquefoil (Potentilla sp.)
seedling of Yellowstone sand verbena (Abronia ammophila)

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Inside of a Seed: Gymnosperms

“Every tree has to stay where it put down roots as a seedling. However, it can reproduce, and in that brief moment when tree embryos are still packed into seeds, they are free. The moment they fall from the tree, the journey can begin.” — The Hidden Life of Trees by Peter Wohlleben

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Seed plants – also known as spermatophytes – make up the largest group of plants on earth. Seed plants consist of five divisions, and among them the angiosperm division (a.k.a. flowering plants) dominates in its number of species. The four remaining divisions are referred to collectively as gymnosperms. This incudes the cycads (Cycadophyta), Ginkgo biloba (the only living species in the division Ginkgophyta), gnetophytes (Gnetophyta), and the conifers (Coniferophyta). Conifers are by far the largest and most widespread gymnosperm division.

Angiosperms and gymnosperms have different evolutionary histories, resulting in their distinct genetic and morphological differences. That being said, an overly-simplistic way of differentiating the two groups is to say that, while both groups produce seeds, angiosperms produce flowers and fruits while gymnosperms produce pollen cones and seed cones. There are always exceptions (Ginkgo biloba, for example, doesn’t produce cones), but for the most part, this is the case.

Pollen cones (top) and seed cones (bottom) of mugo pine (Pinus mugo) via wikimedia commons

Sexual reproduction in gymnosperms follows a familiar pattern. Pollen, which contains the male sex cells, is produced in pollen cones, which are essentially miniature branches with modified leaves called scales that house the male reproductive organs. Mature pollen is shed and carried away by the wind. Lucky pollen grains make their way to the female cones, which are also modified branchlets, but are a bit more complex. Scales sit atop bracts, and on top of the scales are ovules – the female reproductive structures. During fertilization, the bracts open to collect pollen and then close as the seed develops.

When pollen lands on an ovule it forms pollen tubes that help direct the male sex cells to the egg cells inside. The process is similar to pollen tubes extending down the style of a flower. In flowering plants, additional pollen cells combine with cells in the ovule to produce endosperm, a storage tissue that feeds the growing embryo. This doesn’t happen in gymnosperms. Instead, haploid cells within the ovule develop into storage tissue and go on to serve the same role.

The ovule eventually matures into a seed, and the cone opens to release it. The seed sits atop the scale rather than enclosed within a fruit, as it would be in an angiosperm. For this reason gymnosperms are said to have naked seeds. The development of seeds can also be much slower in gymnosperms compared to angiosperms. In some species, seeds don’t reach maturity for as long as two years.

Seed cones and winged seeds of mugo pine (Pinus mugo) via wikimedia commons

Seeds in the genus Pinus are excellent representations of typical gymnosperm seeds. Their basic components are essentially identical to the seeds of angiosperms. The seed coat is also referred to as an integument. It was once the outer covering of the ovule and has developed into the seed covering. A micropyle is sometimes visible on the seed and is the location where the pollen cells entered the ovule. The storage tissue, as mentioned above, is composed of female haploid cells that matured into storage tissue in the ovule. Like angiosperms, the embryo is composed of the radicle (embryonic root), the hypocotyl (embryonic shoot), and cotyledons (embryonic leaves).

Angiosperms can be divided into monocotyledons and dicotyledons according to the number of cotyledons their embryos have (monocots have one, dicots have two). Gymnosperms are considered multi-cotyledonous because, depending on the species, they can have a few to many cotyledons.

Seedling of Swiss pine (Pinus cembra) showing multiple cotyledons via wikimedia commons

For the sake of this introduction to gymnosperm seeds, I have offered a simple overview of the production of seeds in the conifer division. Sexual reproduction and seed formation in the other three gymnosperm divisions is a similar story but varies according to species. Even within the conifers there are differences. For example, the “seed cones” of several gymnosperm species can actually be quite fruit-like, which serves to attract animals to aid in seed dispersal. Also, the pollen of gymnosperms is often thought of as being wind dispersed (and occasionally water dispersed in the case of Ginkgo biloba and some cycads); however, researchers are continuing to discover the pivotal role that insects play in the transfer of pollen for many cycad species, just as they do for so many species of angiosperms.

All of this to say that Botany 101 is simply a window into what is undoubtedly an incredibly diverse and endlessly fascinating group of organisms, and that, as with all branches of science, there is still so much to discover.