Tuesday, March 15, 2016

Functional Chaos

Erythronium hendersonii, an aging flower
With the arrival of my second daughter (March 3rd, 2016!), life has been chaotic to say the least! Unfortunately for you, this post will likely reflect that in the aimless rambling and random bits of semi-useless information that you are about to encounter. But with chaos comes beauty (and love), both exemplified in the garden. Of course the discovery of a few plants in bloom in my garden hardly compares to the feeling I have for my weeks-old daughter, thus my recent lack of time spent outside, so it is still a joy to find plants in bloom.

Leaves of Erythronium hendersonii, a strongly mottled individual
As may seem apparent by the very title of this website I observe the interactions between pollinators and the plants in my garden, and elsewhere. I also observe, for practical reasons, pest resistance. Insects are generally of little concern as pests for me, my garden supports a diversity of beneficial insects that control and prevent population explosions of insect pests.

Self-sown Scilla seedlings in my rock garden. Last year voles killed nearly every bulb in this bed.
The true pests in my garden are the herbivores, and to a lesser extent the slugs (gastropods, not insects). While slugs have seldom killed any of my plants outright (besides all of my Delosperma!), mammals have seen the demise of many of my bulbs in the past year. Deer are a tragedy, eating the aerial portions of plants to such an extent that they become too weak to flower... ever. But the true bane of my gardening existence are the voles, squirrels, and other digging animals. It would seem the only bulbs in my garden that are truly safe from all those digging bastards are those in the family Amaryllidaceae. A few others, by sheer luck, have managed to go unnoticed and are flowering today. In fact, many of the photos in this post are of solo acts, the lone survivors or freshly planted bulbs awaiting their doom.

Puschkinia scilloides, a traumatized lone survivor of the Bulb Massacre of 2015.
In the past, discovering the loss of a plant due to herbivory would have sent me into an embarrassing fit of rage (and subsequent depression), the consequence of my diagnosis of C-PTSD, first at age fifteen, again last year, age 30. Gardening, photography, writing (this blog), and of course the love I share with my wife and two daughters, have all helped me stay on a healthy path and placed me in a position to perhaps inspire others. Why share this sensitive information of the diagnosis of a mental illness on a blog about plants? Well, I debated on whether or not to delete this paragraph or not, but decided to keep it because this blog is about being honest. There is no need to stigmatize mental illness, after all the brain is another organ that is just as susceptible to malady as any other part of the body. There's no need to perpetuate the stigma by hiding it. Gardening, botany, entomology... these are all things that help me cope. Over time, as life comes into perspective, the loss of a plant is not so important. It simply clears the path for something new that I haven't tried yet, like hardy orchids or Fritillaria.

Puschkinia scilloides detail showing lobed coronas
Last year I wrote a simple dichotomous key highlighting the differences between Scilla sensu stricto (orthodox definition) and its close relatives Chionodoxa (absorbed into Scilla sensu lato as of late), and Puschkinia. No matter what they are called, the species can be difficult to tell apart by the average gardener. This simple guide may be useful in this regard. This was not so much created for taxonomic purposes, but rather for the gardener who wishes to be able to determine what they are growing. This key isn't perfect, and doesn't consider many of the genera once considered to be part of Scilla but since moved into different genera such as Barnardia, Fessia, Hyacinthoides, Merwilla, Nectaroscilla, Oncostema, Othocallis, Prospero, Schizocarphus, Schnarfia, or Tractema to name a few. Perhaps in the future I will expand this key to include more genera, but for now I am satisfied with what I have included since they seem to be the most commonly grown of the entire group.

1a. Filaments are round - Scilla sensu stricto (Scilla in the strict sense)
1b. Filaments are flattened - see "2"
2a. Filaments are fused forming a corona - see "3"
3a. Corona is lobed - see "5"
5a. Flowers appear before leaves are fully developed - Puschkinia scilloides
5b. Flowers appear after leaves are fully developed - Puschkinia peshmenii
3b. Corona margin is entire - see "6"
6a. Anthers inserted at middle of corona - Puschkinia bilgineri
6b. Anthers inserted at apex of corona - Scilla vardaria
2b. Filaments are independent and distinct - see "4"
4a. Filaments are even lengths, usually forming a cone - Chionodoxa "Forbesii" group
4b. Filaments are differing lengths, usually forming a tube - Chionodoxa "Luciliae" group

† Chionodoxa "Forbesii" group contains at least Chionodoxa forbesiiC. sardensisC. lochiae, and C. siehei (recently considered a form of C. forbesii). This group often has many small flowers with spherical or pyramidal unfused coronas formed by the flattened filaments of mostly equal lengths.
‡ Chionodoxa "Luciliae" group contains at least Chionodoxa luciliae and C. nana. This group often has fewer large flowers and cylindrical to slightly pyramidal unfused coronas formed by the flattened filaments, often of visibly different lengths. The anthers themselves are of equal length, and they are all attached to the apex of the filament (which differ in length) giving the impression that they too differ in length.

Chionodoxa (Scilla) luciliae
Contrary to my previous assumptions, Scilla, Chionodoxa, and Puschkinia are quite susceptible to herbivory by voles, and perhaps gophers or ground squirrels. This is highly unfortunate because I am quite fond of them, and had assumed them to be inedible by any of the pests in my yard. So, I was quite wrong. This is part of the education I am receiving as a grower of plants, and part of life. Gardening here is challenging, but overcoming the challenge is part of the fun, and success will be sweeter than ever. One observation I've made is that when planted in raised beds with rich soil, the voles find the bulbs much easier. When they are planted out in the landscape, unamended, rocky alluvium, they remain [mostly] ignored or undiscovered by herbivores.

Narcissus 'Jetfire'
In the wake of my article on Narcissus pollination ecology, Natural Pollination of the Genus Narcissus, I am continuing to observe my plants for new correlations and evidence to support my arguments, and the arguments of the authors of the studies I had sited. As insinuated in my article, Narcissus 'Jetfire' is prone to bee pollination based on the parentage (N. cyclamineus was the pollen parent) as well as the floral characteristics ("daffodil" form), and as stated I have observed bumblebees visit the flowers. This doesn't mean that large hawk moths wont visit the flowers, but rather that bees are the primary pollinators of the "daffodil" form, while the "paperwhite" form is prone to moth pollination2.

Narcissus 'Jetfire' has a comparatively short corolla tube
Another consideration when hypothesizing pollinator type is the depth at which the nectar is secreted. The nectaries of Narcissus are located at the ovary, the relatively dark green swollen part near the bend atop the stem behind the perianth (often partially covered by the spathe, see a diagram here). Now observe the photo and notice the corolla, the part between the perianth and the ovary, is flared and relatively short.

Narcissus 'Minnow' has a relatively long corolla tube
Narcissus 'Minnow,' possibly a hybrid of N. tazetta1 or perhaps N. canaliculatus, is a typical "paperwhite" form which is pollinated most effectively by Lepidoptera (butterflies and moths). The constricted corolla tube is too long for bees and other short tongued insects to reach the nectar unless they (bumblebees or other large bees, typically) cheat and bite a hole in the side, thus evading the work of fertilization and going straight for the reward.

Narcissus tazetta, however, exhibits floral dimorphism which means that there are two distinct floral forms (two distinct morphs, thus di-morphic) that occur in separate wild populations. This was studied by Juan Arroyo and Amots Dafni in Israel (Arroyo et al 1995). Populations in low swampy locations had longer corolla tubes and were visited primarily by large hawk moths, while hill populations in relatively drier conditions had shorter corolla tubes and were frequented by bees and other short tongued insects such as flies. Of course, N. 'Minnow' is a clone so by definition does not exhibit dimorphism of any kind, regardless of whether or not it was bred from N. tazetta.

Narcissus jonquilla (or assoanus?) hybrid
This is an unnamed hybrid in my garden, a reliable single-flowered performer. It is strongly scented, and is most likely a hybrid involving both "daffodil" and "paperwhite" forms. The stamens do not appear to be constricted in the tube, and unlike N. jonquilla and N. assoanus (possible parents) it blooms singly and has strap-shaped leaves rather than thin tubular leaves.

Narcissus jonquilla (or assoanus?) hybrid, relatively long corolla tube.
A good sign that it is moth pollinated is the length of the corolla, with nectar quite out-of-reach for even the largest bees. No matter, I still like it and it is very vigorous though it has not increased by offsets or seed.

A beautiful nameless Narcissus hybrid, with a dance fly (Empididae)

Muscari armeniacum, the beauty is in the details
Muscari are nice in detail, but the common Muscari armeniacum are easily lost in the landscape unless they are planted in high density. Bees and moths visit the flowers feverously when the sky is clear, but heavy nonstop rain has halted any pollinator activity over the past week. The bulbs of Muscari are occasionally eaten by rodents and deer seem to have similar intermittent interest, perhaps because they do not stand out very much. I have found bulbs far from where they were planted, suggesting they were moved by rodents (squirrels maybe?) in an attempt to store them, resulting in a blooming plant in a really odd location much to my amusement. The leaves are an eyesore, however, and thus I do not recommend planting these in any prime garden location, but rather in the peripheral or where they will be hidden from sight from other herbaceous perennials.

Pseudomuscari azureum (syn. Muscari azureum)
One of my favorite "muscaris" is the inconceivably unnoticeable Pseudomuscari azureum. This diminutive plant is from the mountains of Turkey, a hot spot for Mediterranean bulbs. The genus Pseudomuscari is differentiated from Muscari by the relatively open mouth of the flowers (bell-shaped rather than urn-shaped). This species is easy from seed, I have a full pot of seedlings which I have no plan for as of now (trading, perhaps?)

Fritillaria persica
This is my first attempt at growing Fritillaria, a genus that has evaded my interest until recently. The genus seems to be somewhere between Lilium, Calochortus, and Tulipa in texture and flower shape. If I can keep the deer from eating this one I will show you the flowers when they appear. The flowers of this species are black, something which I look forward to with great anticipation.

Platanthera ciliaris
I have been feeling adventurous, so expanding my plant horizons by trying to grow more plants like Fritillaria and now hardy orchids. We have a few native orchids here, one being Piperia (Platanthera) transversa and surely a number of others that I haven't encountered. Platanthera ciliaris, the yellow-fringed orchid, is a butterfly pollinated species from the Southeastern US, including the Gulf Coast and in boggy conditions inland. I am growing it in my forest/shade garden with some shade and some filtered sunlight close to one of the main soaker hoses which I use throughout the entire garden. It is fenced from deer, but not squirrels, so I hope it will flourish, and maybe even flower!

Goodyera pubescens
Another orchid, a genus I have encountered both in coastal Oregon and in Northern California (see post), Goodyera is grown as much for its flowers as it is for the decorative leaves. It is rhizomatous, and so will perhaps spread out a bit if it is happy. Both the Platanthera and the Goodyera were inoculated with a generalist mycorrhizal fungi mix which will hopefully improve their performance if a mutualism is formed. Goodyera pubescens is another Eastern US orchid in cultivation. It is known as the downy rattlesnake plantain due to the fine pubescence covering the leaves and flowering stalk.

Piperia (Platanthera) transversa
A small native orchid, this species grows in filtered light under a canopy of Pinus, Arbutus, and Quercus. In this area, the flowering stalks (one per plant) appear around June. In the drier part of the yard, under a stand of Pinus ponderosa, they do not seem to flower, or at least have not flowered in the last six years since we have lived here. Under a mix of oak and Douglas fir, they seem to flower more reliably. They are perhaps pollinated by moths, possibly flies, though I have made no such observations unfortunately.

Pulmonaria officinalis, seed grown
Last year I showcased some of my seed grown Pulmonaria (Boraginaceae) plants. Here they are again! Please compare with some of the cultivated selections photographed below. I hope that they will someday hybridize, though I am not sure they are known to self sow in my area. Perhaps someday they will set a good crop of seed that I can grow on in pots. These were sown aout an inch deep in the shade in gallon pots, covered with a thin layer of fungus dominant compost (aka leaf mold). I believe this genus would also benefit from a mycorrhizal inoculant.

Pulmonaria officinalis, seed grown
Pulmonaria officinalis, seed grown
Pulmonaria longifolia hybrid
Pulmonaria saccharata 'Sissinghurst White'

Further reading:

Acton, Hamilton. "On the formation of sugars in the septal glands of Narcissus." Ann Bot (1888) os-2 (1): 53-63.
doi: 10.1093/aob/os-2.1.53

Arroyo, Juan, and Amots Dafni. "Variations in habitat, season, flower traits and pollinators in dimorphic Narcissus tazetta L.(Amaryllidaceae) in Israel." New Phytologist 129.1 (1995): 135-145.

Owen, T. "Natural Pollination of the Genus Narcissus." The Amateur Anthecologist. 7 Mar. 2016.


Owen, T. "Piperia Transversa." Pacific Bulb Society Wiki. 9 July 2015. Web.

Ugiansky, R. 2010. Plant fact sheet for downy rattlesnake plantain (Goodyera pubescens). USDA-Natural Resources Conservation Service, Norman A. Berg National Plant Materials Center. Beltsville, MD 20705.

Erythronium hendersonii with a muscid fly (Muscoidea)


  1. Those little white dots in the Frit are interesting. That's where the flowers come from, but sometimes they don't develop beyond white dot stage.

    Interesting to see the Puschkinia, I've grown some this year, have to see if the pictures make it to the PBS wiki.

    Nice to see your soldiers and sailors (pulmonaria), presumably because British soldiers wore redcoats (to give the other chaps a chance). I used to worry which came first blue or red.

  2. thanks so much for the beautiful photos & great info!


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