Sunday, March 27, 2016

Erythronium

Erythronium hendersonii

Erythronium is one of my favorite genera of wildflowers. This year, for some reason, they are blooming prolifically to the delight of bees and myself alike! Honeybees, bumblebees, and anthophorine bees (Anthophora or Habropoda) are visiting the flowers of Erythronium hendersonii, and surely other species, feverously. I expect this year to be one of the best for seed production, the primary mode of reproduction (as opposed to vegetative increase) for E. hendersonii and oregonum, two species which I've been fortunate enough to observe directly in habitat this year.


Erythronium hendersonii
A short introduction for those not introduced to this genus: There are around 20 to 31 species in all, many from the Western US, a few in the Eastern US, and a some in Eurasia. They are in the lily family (Liliaceae), and so related to FritillariaLilium, and Tulipa among others. These are mostly Spring ephemerals, though some of the alpine and subalpine species bloom in the Summer as the snow melts. They grow from bulbs (some say corms), with tunics that don't seem to cover the entire bulb, thus they are susceptible to drying out. Contractile roots pull the bulbs down deeply, perhaps to help them evade herbivores (voles, etc.) and to keep them from desiccation is Summer.


Erythronium hendersonii mottled leaf. E. oregonum is similarly mottled.
Many of the species have mottled or spotted leaves, some plain green. In general, mottled leaves are found in species growing in the West while spotted leaves are found in species in the East and Eurasia. Flowers are often born singly (sometimes in clusters in mature and happy plants, and nearly all species have flowers that face downwards (except Erythronium rostratum, a yellow-flowered American forest dweller of the central-southern states). Most are probably bee pollinated, though they aren't specialized and can probably be pollinated by a variety of insects, including flies.


Erythronium hendersonii, one anther dehisced
The pendant flowers have multiple beneficial consequences. First is to exclude inefficient pollinators, since pendant flowers are difficult for many insects to land on (beetles or butterflies, for example, struggle to land on fully pendant flowers). Bees are the primary pollinators of Erythronium, though flies are certainly capable of pollinating the open flowers and may be of special importance in forest dwelling and alpine species where bees are not always as abundant. Pendant flowers also have the advantage of being protected from rain. The pollen is dehisced over a relatively delayed period since the anthers don't open all at once. In general, three anthers will open first while three will open days after the first three. This lessens the chance that hard rain or wind will damage the pollen. The recurved tepals further protect the flower by closing or bending down in rain or in diminished sunlight, then opening up again when the sun comes out.


Erythronium oregonum ssp. leucandrum
Bulbs always seem to grow deeper than one would think, and digging wild plants is likely to kill them (besides being an unethical practice). Fortunately, Erythronium is relatively easy to grow from seed. Seed of most species can be sown in Autumn on the surface of the potting medium. As per advice from friend and bulb grower extraordinaire Ian Young, soaking the seeds overnight in water improves the first year germ rate. I have found this to be accurate, as seed I had acquired of a handful of species was successfully sown this way. Left outside throughout Winter, all of my Erythronium seed pots have germinated. I will allow them to grow on for another year before repotting them or planting them out into the garden. As stated, neither E. hendersonii or oregonum increase reliably vegetatively and are best increased by seed.


Erythronium hendersonii seedlings
A species from Eastern North America, yellow-flowered Erythronium americanum, has been heavily researched. In a handful of studies, mycorrhizal relationships between E. americanum and associated plants were examined. The most exciting discovery, in my opinion, was that bulbs (or as they say, corms) infected with arbuscular mycorrhizal fungi which enters the roots, unlike ectomycorrhiza which doesn't enter the roots, were found to be nearly twice as large as bulbs treated with fungicide. E. americanum produces roots in the Autumn, as do all Erythronium, and it is then that they are infected. Surrounding plants and trees, such as sugar maple (Acer saccharum) grow in close association with E. americanum and have been found to share mycorrhiza. Another interesting discovery was that E. americanum may help the establishment of young sugar maples by supplying them with carbon through the mycorrhizal network. Ian Young wrote a concise piece on the seeds of Erythronium, including how to grow them, found here: Erythroniums in Cultivation: Seeds (Ch.1). As an aside, the bulbs are supposedly edible, though I couldn't bring myself to eat something so beautiful (and I discourage anyone from eating wild or unfamiliar plants for ethical and safety concerns).


Erythronium hendersonii in habitat
I speculate that other species of Erythronium also participate in mycorrhizal networks, and that they benefit from the infection as well. Many of the trees and shrubs that grow in association with Erythronium hendersonii and E. oregonum benefit from or require mycorrhiza (Acer sp., Arbutus menziesii, Arctostaphylos viscida, Ceanothus sp., Quercus sp., Pseudotsuga menziesii, etc.), therefore it is not inconceivable that they too benefit from mycorrhizal associations.


Erythronium oregonum

Erythronium oregonum ssp. leucandrum, differentiated from the type by having white anthers.
I had recently discovered, entirely by accident, a small population of Erythronium oregonum ssp. leucandrum on a nearby mountain above a small creek. It was my first encounter with the species in Oregon, the wild, or anywhere. The air was moist, no doubt in result of the running water complete with a small waterfall, and it was cool. In small clearings of Arbutus menziesiiPseudotsuga menziesii, leafless Quercus, small groups of E. oregonum were awaiting discovery. There were a few individuals of E. hendersonii, though not many. There are conflicting reports of whether the two species hybridize. I didn't see any hybrids, though I did see the species growing in close vicinity. The more adaptable E. hendersonii ventured slightly into the woods, while the more picky E. oregonum was solely found in forest clearings.


Erythronium oregonum ssp. leucandrum, greenish reverse

Erythronium oregonum ssp. leucandrum with a dance fly (Empididae), a potential pollinator
Erythronium oregonum ssp. leucandrum, nectar guides
Erythronium oregonum ssp. leucandrum in habitat
Erythronium oregonum ssp. leucandrum
Erythronium oregonum ssp. leucandrum with Toxicodendron (no leaves yet)

Erythronium oregonum ssp. leucandrum in habitat



Erythronium hendersonii

Erythronium hendersonii and Apis mellifera
Since I have lived in Oregon, Erythronium hendersonii has been a favorite wildflower of mine. Compared to E. oregonum, it is more common in hotter and drier areas, and prefers forest edges to cool shady clearings. This could explain the lack of hybridization, the two seem to prefer distinctly different habitats. This year has been a particularly floriferous year for E. hendersonii, and it is the first year I've seen honeybees visiting the flowers. The typical pollinators, based on my observations, are species of Bombus and anthophorine bees. More photos and information can be found on the Pacific Bulb Society page: Erythronium hendersonii.


Erythronium hendersonii
Erythronium hendersonii
Erythronium hendersonii, an aging flower.
Erythronium hendersonii, one anther just beginning to open.
Erythronium hendersonii
Erythronium hendersonii, aphids on the scape.
Erythronium hendersonii, large and small morphs.
Erythronium hendersonii, anthers shorten after opening.
Erythronium hendersonii, true petals (inner) ridged, sepals (outer) mostly flat.
Erythronium hendersonii leaves.
Erythronium hendersonii, a particularly happy roadside cluster. I find this kind of growth uncommon.
Erythronium hendersonii nectar guides.
Erythronium hendersonii
Erythronium hendersonii
Erythronium hendersonii, interesting reverse coloration.

References:

Alverson, Ed. "My Erythronium "Big Year"" Scottish Rock Garden Club. May 2006.http://www.srgc.org.uk/monthfeature/june2006/content.html 

Lapointe, Line, and Sylvain Lerat. "Annual Growth of the Spring Ephemeral Erythronium americanum as a Function of Temperature and Mycorrhizal Status." Can. J. Bot. 84.1 (2006): 39-48.https://goo.gl/ViL1t3 

Lerat, Sylvain, Rachel Gauci, Jean Catford, Horst Vierheilig, Yves Piché, and Line Lapointe. "14C Transfer between the Spring Ephemeral Erythronium americanum and Sugar Maple Saplings via Arbuscular Mycorrhizal Fungi in Natural Stands." Oecologia 132.2 (2002): 181-87.http://goo.gl/b0RAXU 

Owen, Travis, et al. "Erythronium hendersonii." Pacific Bulb Society. 2015.http://www.pacificbulbsociety.org/pbswiki/index.php/Erythronium_hendersonii 

Young, Ian. "Erythroniums in Cultivation: Erythronium oregonum (Bulb Log 49)." SRGC Bulb Log. 9 Dec. 2015.http://www.srgc.org.uk/logs/logdir/2015Dec091449666371BULB_LOG_4915.pdf 

Young, Ian. "Erythroniums in Cultivation: Erythronium hendersonii (Bulb Log 51)." SRGC Bulb Log. 23 Dec. 2015.
http://www.srgc.org.uk/logs/logdir/2015Dec231450868200BULB_LOG_5115.pdf

Saturday, March 19, 2016

Floral Visitors 18

Muscari armeniacum with a honeybee
Pollinators of all kinds have been out, particularly bees who are taking advantage of the introduced species in my garden. Muscari is in full bloom, attracting gangs of honeybees. Last year I witnessed a variety of other floral visitors such as plume moths (Pterophoridae), geometrid moths (Geometridae), anthephorine bees (tribe Anthophorini), and other small arthropods. They don't seem to increase very well here, perhaps the soil is not to their liking or it dries up too quickly after the Spring rains cease. Most of them are planted out in native soil, rather than a raised bed where I grow most of my perennials. Perhaps they would benefit from a fall application of leaves to encourage some microbial activity (thus stimulating quicker nutrient cycling).

Scilla siberica
The bulbs of Scilla are sometimes considered to be irritating when handled by some people, and my assumption was that the assumed irritating constituents would also deter animal pests. Deer surely ignore them, yet voles are seemingly unaffected. Last year, I had a lot of Scilla siberica planted around and it had been a reliable resource for honeybees and bumblebees. However, this year has seen a drop in the number of flowering plants and honeybees have not been observed visiting them. Still, this wont keep me from attempting to grow more, next time I'll just use mesh baskets or wire cages, a necessary pain in the a**.

Chionodoxa luciliae (syn. Scilla luciliae)
The Chionodoxa, now widely accepted as Scilla, are like S. siberica and now are few in the garden. Yet, these tiny blue flowered bulbs have swooned me and I will continue to plant them into the garden, next time with underground protection. The pollen of Chionodoxa is not as exposed as Scilla, perhaps an adaptation to prevent damage or loss of the pollen from rain. It may just be a coincidence, but it seems bees are less interested in Chionodoxa than Scilla (sensu stricto) which have highly available pollen and nectar. Bumblebees were spotted visiting Chionodoxa last year, perhaps because they possess probosces that are robust enough to push between the anthers to reach the nectaries within.

Hyacinthus orientalis with Mecaphesa, a bee predator.
Hyacinthus, the common hyacinths sold alongside tulips and daffodils in the fall, are highly attractive to honeybees even when planted in small quantities, unlike the other scillas. I have a handful of hyacinths planted under a native wild lilac, aka deerbrush (Ceanothus integerrimus). Ceanothus are good for beneficial soil dwelling microbes, and are known to fix nitrogen, a result of a symbiotic relationship with some type of soil dwelling fungi. The plant trades something (root exudates), and in return receives nutrients or something similarly useful from the fungi. This in turn benefits everything growing in the vicinity because nutrients that would otherwise be unavailable to plants for one reason or another are being made continuously available by microbial activity. Hyacinthus orientalis are known to be heavy feeders, and will dwindle over time in less than adequate soil. Yet here under the Ceanothus, they are thriving! (Learn more about nutrient cycling here.)

Anemone blanda
Similar to the Hyacinthus, Anemone blanda is similarly attractive to honeybees even though there are relatively few of them and they are not planted near each other. Unlike hyacinths, the main appeal here is for the pollen. Besides honeybees, a variety of bumblebees and flies (Syrphidae, probably other families too) also visit these flowers. It is uncertain as to whether or not they produce nectar, or if they do it may be in a very small quantity and highly dependent on the growing conditions (including aspect). In general, when plants are grown in less than optimal conditions, the first thing they'll do is cease nectar production, survival comes first. Pollen, however, is much more important to a plant since it is required to create the next generation of plants. Pollen is also highly important to bees this time of year because it is the sole source of protein for developing larvae, nectar and honey being reserved for adults who require high amounts of carbohydrates to fuel flight and other tasks important to survival. (A fun/appetizing fact: wasps utilize other insects as the protein source for their larvae instead of pollen. Bees are basically vegans while wasps are largely predators/carnivores, though the adults of both feed on nectar.)

Hepatica acutiloba
Closely related to the genus Anemone, and often considered to be included in it, the single Hepatica in my shade garden attracts few to no pollinators. Though beautiful, it has some factors working against it when it comes to pollinator attraction. First, it is grown in shade while bees prefer to work flowers in the sun, especially this early in the year when the days are still very cool. Second, there is only one. A single flower of any plant is highly unlikely to attract anything, except for a photographer. Bees and other pollinators are more attracted to groups of plants (Xerxes Society studies suggest single species plantings of 3×3' or larger to be most effective), and are most attracted to groups of single species rather than mixes. However, I do find that having a mix of flowers does attract a wide range of pollinators nonetheless. I do plan on acquiring more Hepatica for my shade garden, as well as more flowering plants for my shade garden to entice more pollinators into the dark.

Crocus vernus
A lone Crocus stands under Pinus ponderosa, where dozens of other crocuses were planted and many were uprooted by both deer and turkeys. As the year progresses, fewer and fewer crocuses are to be seen and fewer and fewer are likely to be visited by pollinators in my garden. This is probably due to the increase in competition and the decrease in the population of the crocuses rather than their general appeal. I strongly believe that if I had a field of the latest blooming Crocus they would be crawling with pollinators (and probably also filled with deer, voles, turkeys, and other herbivores). This is probably the last look at Crocus for the year, unless my saffron decided to bloom this year.

Viola odorata and Vinca minor
I have worked hard to build up a thriving community of plants in the small raised bed around my well house, and the work has paid off. Vinca minor, a close relative to the highly invasive Vinca major (which coats entire riparian banks and shady hillsides that are too dark for Rubus armeniacum), grows happily along with dozens of other small plants. Much of what grows here now was sown as seed, some planted as offsets. Roots from nearby Pseudotsuga menziesii invaded these beds quickly after they were established, and brought with them myriads of fungal hyphae. At first I thought this was a horrible thing, and I had attempted to eradicate the beds of the intruders, a futile task since there is a 100' tree less than ten feet away. Now I see that the plants growing there now have no issue competing with the tree roots, and I hypothesize that the fungal network has brought with it a diversity of other microbes which are supporting the growth of the plants.

Microbes sequester nutrients which could otherwise be lost due to watering or rain and delve deep out of the reach of plant roots. These nutrients are eaten by microbes, sequestered in their bodies, and released when the microbes either die of natural causes or are eaten by other microbes, thus making them continuously available to plant roots. So a diversity of microbes is a good thing. The microbes are supported by a diversity of plants, all of which release different exudates (the closest thing to plant poop) into the rhizosphere (the area just around the roots) and feeding the microbes. Many microbes occupying the rhizosphere feeding on exudates can crowd out pathogenic microbes and prevent disease in plants. The benefits go on and on.

Now back to Vinca, they are reportedly bee pollinated, though I have made no such observation. The blue flowers are nice though, and with all of the competition I doubt the plant will get too out of hand.

Viola odorata
Sweet violets are very satisfying to me. They grow very well here, require no Summer irrigation (though they flower better with it), and are happy in sun or shade. Bees appreciate the flowers, and where they are prevalent can be detected from afar by their sweet scent. They also like to spread, so it is worth taking the time to consider their placement before introducing them to your garden. If you have a lawn or a large patch of bare ground, say, under a tree, they'd be perfect! They don't require pollination to set seed, and will spread by runners, so a sizable patch may be formed in a short period of time.

Lamium maculatum
Another very easy plant that is attractive to bees, Lamium maculatum grows in the sunny corners of the raised beds surrounding my well house. It is a low growing maintenance-free creeper that can tolerate a lot of drought when established, is ignored by deer, and unaffected by mole/vole tunneling. Bumblebees are most interested in the odd reptilian-reminiscent flowers, though honeybees will occasionally frequent them.



Further Reading:

Lee-Mäder, Eric, and Marla Spivak, eds. Attracting Native Pollinators: Protecting North America's Bees and Butterflies: The Xerces Society Guide. North Adams, MA: Storey Pub., 2011. Print.
http://goo.gl/cAMJvR

Walker, Travis S. et al. "Root Exudation and Rhizosphere Biology". Plant Physiol. Vol. 132, 2003.
http://goo.gl/A2vWLh

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
http://goo.gl/miBzlr

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.
http://goo.gl/4Xzrnc

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

http://goo.gl/CkzIzG

Owen, T. "Piperia Transversa." Pacific Bulb Society Wiki. 9 July 2015. Web.
http://goo.gl/0JwjPD

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.
http://goo.gl/3IUwlG

Erythronium hendersonii with a muscid fly (Muscoidea)