Saturday, July 30, 2016

Madia elegans

A honeybee forages on Madia elegans (2016)
Madia elegans, in the field with other wildflowers early in the morning (2015)
Madia elegans D.Don ex Lindl. was new to me when my wife and I moved here over seven years ago. In fact, it was a surprise to me even a few years after we had been living here. It is an annual species native from Southern California north into Western Oregon and perhaps into Washington. The flowers are sensitive to sunlight (or is it temperature?) and respond by closing during the day, only to open after the sun is setting. It is believed that the flowers do this to conserve moisture (prevent the nectar and pollen from drying out, floral segments withering, etc).

Madia elegans is covered in glandular hairs (2016)
It was for this reason that I never noticed them, during the day all you can see are the closed buds. The plants are also covered with resinous hairs that help the plant retain moisture, an important adaptation since they bloom during the hottest part of the year over a long period, roughly starting in July and going til early September in my area. Back to my discovery, it just so happened that for a few years I had not ventured out into the field at or before dawn, so when I decided to go for a walk at the crack of dawn one morning I was in awe of what seemed like a magical phenomenon. I had thought, where had these come from? It was perplexing.

Madia elegans variation
Madia elegans, known as common madia or tarweed (in reference to the odor of the resinous excretion on the stem and leaves), exhibits a great deal of variation in flower forms, plant height, bloom period, number of florets per inflorescence, and number of flowers per plant. They can grow anywhere from six inches to over sixty inches when grown in rich soil and given water, as is the case where I have sown them in garden beds. I would suggest if you are planning on growing these, and I have seen seed available online, that you provide stakes or other plants for them to lean on. The flowers themselves smell of honey, and are probably rich in nectar as evident by my recurring observations of bees on the flowers.

Madia elegans opening (2015)
In the evening, after the sun drops below the horizon, but before the sky goes dark, the flowers start to open. The petals (ray florets) unroll from the center. I have observed few pollinators visiting them at this time, though I have seen a few solitary bees visit at dusk on warm days. Small moths were often present, but I've never seen them land on any of the flowers during a handful of brief observations. I still wonder, do the moths visit the flowers after dark? Probably not so much, I am convinced from three years of observations that bees are in fact the primary pollinators.

Madia elegans closed with Epicauta (blister beetle)
During the day, the flowers appear as unopened buds. The petals are rolled up, and green hairy bracts protect them from drying winds and the heat of the sun. They do not protect the flowers, however, from blister beetles in the genus Epicauta, found on many of the flowers day and night. The small half-inch-long Epicauta beetles linger on the flowers all day, eating the ray florets as they unroll in the afternoon/evening. Perhaps the beetles seek mates and do the deed itself on and around the flowers, something I have observed on the nonnative Daucus carota which shares the field with the Madia and flowers simultaneously.

Madia elegans with Epicauta
Blister beetles are toxic and have been known to give people contact dermatitis when contact to human skin is made with some genera in the Meloidae (blister beetle family). The chemical cantharidin, a poisonous terpene than works by breaking the connections between cellular tissue resulting in blisters, is secreted by the beetles. The amount of the toxin produced by the beetles varies greatly depending n the genus, species, and whether it is a male or female (males produce more). I cannot attest to the toxicity of the genus Epicuata, nor do I intend on testing it for myself.

Madia elegans with Epicauta
Deformed flowers like this are encountered frequently, though new flowers open just as frequently. The fact that blister beetles eat the petals doesn't discount them as pollinators. There have been cases where certain plant species will still set viable seed even after their beetle pollinators eat the petals, parts that are nonessential to seed formation, but instead useful for initially enticing pollinators to visit. However, I do suspect that the chemical cantharidin produced by the beetles must interact with any pollen the beetles inadvertently pick up on their bodies, and I would suspect the interaction to negatively impact the viability of the pollen, though this is based solely on instinct.

Madia elegans with Halictus ligatus ♀
Bees are considered by most to be the primary pollinators of the genus Madia. I can attest to that, during several observations over the past three years bees were the main if not the only floral visitors in the mornings before the flowers closed for the day. Very early in the morning, it may be possible to find some solitary bees, motionless, sitting in the flowers, probably from the preceding evening. Males of some native bees can occasionally be found sleeping on the flowers, as they build no nests to sleep in and instead hang out at the flower looking for a hot date.

Madia elegans with Halictus ligatus ♀
Halictus bees dig their nests in flat ground and then line the brood cells with a waxy secretion from a special gland at the tip of their abdomen called the Dufor's gland. They are eusocial, meaning they are often solitary in the beginning, then first generation daughters remain in the nest to care for the young (as with honeybees). Depending on the species, a nest may have a single queen and a few workers, or they may have many queens and hundreds of workers in an extensive underground colony. I suspect the climate may also play a role, where food sources are scarce it may be more adventitious to be more or less social. Additionally, cooler climates stimulate solitary behavior while warmer climates motivate eusocialism.

Madia elegans with solitary sweat bee
The flowers of Madia elegans open towards the sun, so bees that spent the night warm up quickly in the morning light. I have found several types of bees perched on the flowers early in the morning, unable to move since the temperature was yet too low.

Madia elegans with Sphaerophoria sulphuripes ♀
Having pollen readily available attracted this small syrphid fly, a Sphaerophoria sulphuripes female. Like many syrphids, Sphaerophoria seek plants with aphids to lay their eggs on. The larvae then seek out and eat the aphids, possibly several per day. Syrphid flies also eat pollen, and are considered important pollinators.

Madia elegans with Mecaphesa schlingeri
An adventitious crab spider awaits a meal, a small bee or fly. According to Bugguide, Mecaphesa schlingeri is only found in California. I have now seen this spider two consecutive years, always on yellow flowers (once on Madia and now on Coreopsis).

Madia elegans with Apis mellifera
Honeybees work the flowers too, to my delight! They visit for both pollen and nectar, spending a generous amount of time on each inflorescence making sure to work every floret. The honeybees spend more time working these flowers than the native bees, suggesting to me that they are either not as efficient at collecting the goods as the natives or not adapted to the local predators, or both.

Apis mellifera with a dusting of pollen on Madia elegans (2016)
Madia elegans may be a very underrated bee plant worth getting to know if you are a beekeeper due to the fact that its natural habit is to grow in large swaths, and that it is worked judiciously by honeybees during what most beekeepers call a dearth. The closely related Madia gracilis has also been worked by honeybees (and halictid bees), though it is likely to be less common than M. elegans. I have not found honeybees to be interested in M. citriodora, though I have only observed one population that was dwarfed by a larger population of M. elegans.

Madia elegans with a male Melissodes
Compared to the honeybees, the long horned bees (Eucerini, so called for the long antennae) spend a few seconds on each flower at the most. Male solitary bees do not provision nests, so do not collect pollen like the females (who gather pollen to feed to their young). Pollen takes less time to collect than nectar, which would account for the quick visits of the females. To collect pollen, a bee must only be required to slam itself into a flower and wiggle around, something aided by sonication or buzz pollination. To collect nectar, focus must be directed to sending the proboscis down the floral tubes, therefore not paying attention to the pesky photographer. Some bees, like the long horned bees and others, collect nectar very quickly as well.

Madia elegans with Melissodes
To take some of these photos of Melissodes, I had to point the camera at a single flower and wait. Fortunately, the bees were very active and it took less than ten minutes to snap these photos. Due to the intensity of light in the morning coming from the East, it is crucial to stand in either a SE or NE position to take decent quality photos of pollinators, especially bees. Standing in an easterly position blocks light, and standing in a westerly position means you are staring at the sun. The issue with Madia is that they tend to face the sun, so if you, too, are facing the sun chances are you cannot see the faces of the flowers, thus you cannot see the bees.

Madia elegans with a female Melissodes long-horned bee
The genus Melissodes is composed of about ninety-seven species, eight of which are in the United States. The can be found in nearly every state as well as Canada, and perhaps Mexico. They are all ground nesting bees, as are all eucerine bees. This is a common trait of native solitary bees here in my area, the vast majority being ground nesters.

Madia elegans with male Melissodes long-horned bees (2015, inset from 2014)
For the last three years, I've observed male Melissodes bees motionless on the flowers early in the morning. I was able to take the inset photo using my smartphone camera. The recurrent years photos with bees were taken between six and eight o'clock in the morning, thus the increase in bee activity and relative difficulty in photography.

Madia elegans with a male Melissodes
During a recent observation, I witnessed a mating event in which a foraging Eucerini female was, in human terms, raped by a scandalous male. The event was very brief and did not allow me the time to point the camera and focus. This is common in the world of solitary bees, and perhaps other American social bees (like bumblebees) where females mate while foraging. I have witnessed this with a few solitary bee species. Once I witnessed a pair of mating bumblebees, the female laboriously foraging on the flowers of a redbud tree (Cercis canadensis) while the male presumably passed his genes, one last rumpus before breaking off his member inside of her and bleeding to his death (true story).

Agapostemon male spending the night on a Madia flower
This photo marks the sole incident in which I observed Agapostemon on Madia elegans, not to say there aren't a vast variety of other bees which work these flowers. This is a male, evident by the behavior and the lack of scopae, or pollen carrying apparati.

Hesperia juba, Juba skipper (September 2015)
Late in the flowering period, I saw less bees visit the flowers, yet a few skippers were common per observation. I hypothesize that they are less efficient at pollinating the flowers, which I believe to be geared more towards bee pollination, yet butterflies probably do account for some seed production. Rather than spreading pollen by their bodies as with bees, butterflies and moths transfer pollen which adheres to their proboscis or sometimes their feet. On occasion pollen is transferred by the bodies of large Sphingidae moths visiting flowers with exerted stamens such as some species of Oenothera.

Madia elegans with plant bugs (family Miridae)
Not often considered as pollinators, plant bugs (true bugs in the family Miridae) are another insect found to be interested in Madia. It is unclear whether they effectively contribute to pollination, though they are found on many different types of flowers from those in the Asteraceae to Narcissus to Convolvulus. They are often found in small groups or congregations which is a clue that they may be seeking either shelter, mates, or both. They feed on nectar and perhaps pollen to some extent, though the latter is purely conjecture on my part. I am not sure if they frequently move between flowers, yet Madia may be self compatible (pollen from one floret transferred to another in the same inflorescence) so the possibility is valid.

Madia elegans with Daucus carota (2015)
If it hasn't become clear, I love this plant and all of its beneficiaries. Working for a commercial beekeeping operation, I often hear from other beekeepers about the dearth this region experiences after the blackberry bloom concludes. Starthistle (Centaurea solstitialis), another noxious and invasive alongside the blackberry, is highly acclaimed by local beekeepers as the last good nectar flow. I cannot speak with certainty, but it would appear that Madia elegans is at least one native that is perhaps highly underrated for its usefulness to both beekeepers and the native pollinators that are still active at this time of year. Madia grows where it can become very dry, and copes very well I might add, and also spreads to form large populations. Identifying and preserving Madia elegans may go a long way towards helping the local and nonnative pollinators, which will be good for everyone and everything alive around you.

Madia elegans (2016)

Monday, July 18, 2016

Floral Visitors XXI

Allium ampeloprasum var. ampeloprasum with Strymon melinus & Bombus vandykei
If you have a flower garden, it is probably teeming with life right now as mine is. Bumblebees, honeybees, a variety of small solitary bees, large carpenter bees, wasps, skippers, swallowtails, hairstreaks, beetles, and a variety of other beneficial insects are frenzied over the buffet I have planted for them. This is an ideal situation, one that I strive for. The key to my success attracting such a wide array of pollinating insects is the wide array of floral forms presented. Even just within a single genus of flowering plants there will be variation of pollinator types between species, which has been the case for the gray hairstreak butterflies and elephant garlic. I have not seen the hairstreak on the drumstick allium (Allium sphaerocephalon) which is planted nearby, and similar phenomena occur throughout the year with other plants. That is one of the most interesting aspects of planting for pollinators.

Many people here keep ungulates and ruminants or grow hay for sale as animal fodder, and few if any are likely to be as environmentally conscientious as the readers of this blog. Diverse flower plantings have been demonstrated to increase the amount of beneficial insects (predators, parasitoids, pollinators, etc.) in a farm setting (Altieri & Letourneau 1982Douglas et al. 2000, etc.) and likely in a garden setting as well. The Xerxes Society book Farming with Native Beneficial Insects cites many studies which go further and say that native wildflower plantings attract a higher diversity of beneficial insects while weedy field borders host more pest insects. Additionally, wildflowers may attract more native pollinators which will take some of the pressure off honeybees and simultaneously improve crop yields of crops that require pollination. An initial investment of time and money will pay off in years to come by reducing pest populations and enabling a farmer to reduce or possibly forgo the use of chemical pesticides altogether, which would all increase the profitability of their operation. Why then don't more farms plant flowers?

Allium ampeloprasum var. ampeloprasum with Strymon melinus & Bombus vandykei
Pollinators can be indicative of the area. Gray hairstreaks, occurring across the nation and spanning between Canada and Venezuela, are typically found in nonforested disturbed weedy areas. This is somewhat peculiar since we live basically at the edge of an expansive mixed coniferous forest, though as one goes deeper into the valley or closer to "civilization" there are less trees and more nonnative species, including humans.

Allium ampeloprasum var. ampeloprasum with Apis mellifera
I have mentioned before that alliums are good plants for pollinators. The reason for this is the bloom times between species is incredibly variable, and each bulb flowers over an extended period as new florets continue to open. Elephant garlic (Allium ampeloprasum var. ampeloprasum ) is one of my favorites and is incredibly easy to acquire, though one must procure fresh cloves or else they may never come up. The leaves appear in early spring and look very similar to corn as they clasp the stem, then dry up as the scape elongates up to four feet tall or more. The actual inflorescence itself is close to four inches across and as you can see is composed of hundreds of small lily like florets. Elephant garlic is very closely related to the leek and is only separated taxonomically at the varietal level, with the primary morphological difference being the bulb.

Thread-waisted wasp (Ammophila sp.) on Allium sphaerocephalon
A similarly accessible onion is the drumstick allium, Allium sphaerocephalon. These grow up to three feet, but usually shorter, with flower heads the size of a ping pong ball. They are similarly attractive to a variety of bees as are other alliums. Some find them to be weedy or difficult to remove, but this trait isn't bad for a pollinator garden since the purpose triumphs over the aesthetic, although to me those factors are one in the same. I would suggest planting them in drifts or clumps to enhance the effect, and close to other perennials which will help support the thin yet sturdy scapes.

Ammophila sp. on Allium sphaerocephalon
Large and fast moving thread-waisted wasps (Ammophila sp.) are a good ally in the garden as they parasitize caterpillars and sawfly larvae to feed to their own larvae in underground nests. They are solitary, and much more likely to flee than to be aggressive, although the females can technically sting people (which would probably require one to capture the poor creature and hold it bare handed).

Ceratina sp. on a Hemerocallis hybrid
As far as I'm concerned, daylilies are some of the poorest pollinator plants in my garden. Typically they are either moth or butterfly pollinated (depending on the species) and are only on rare occasions visited by bees. Honeybees? Probably never, but I have heard reports of bumblebees visiting them when planted in masses. I have a few plants, I'll admit, but they were free. I am surprised to find small carpenter bees inside, and on more than one observation. The tiny bees are perhaps able to reach the nectar hidden deep inside, and may even collect some pollen.

Ceratina sp. with pollen on a Hemerocallis hybrid
Unlike the larger carpenter bees (Xylocopa sp.), Ceratina are not large enough to excavate tunnels into wood but rather the dried pith of woody stems from plants like blackberry (Rubus sp.) and the like, though I'm not sure which native plants they prefer to nest in. Females forage for an extended period, pretty much until they can't (mid autumn) and finally rest at the entrance of their linear pithy nest where they die (frosted, I'd imagine), blocking the entrance to potential intruders. The following year the brood pupate and emerge to start the cycle anew.

Selasphorus rufus ♀ on Crocosmia 'Lucifer'
A few other plants in my garden are also marginally good pollinator plants. Irids (aka plants in the Iridaceae, or Iris family) such as Crocosmia and Gladiolus attract few insect pollinators, yet are excellent for the hummingbirds. Crocosmia is a fairly weedy genus, particularly annoying to gardeners in mild coastal climates as the plants can easily escape and become nearly impossible to remove. Crocosmia grow from corms, which unlike true bulbs are technically a part of stem modified for storing starches and nutrients. Corms of these offset freely, and any attempt to remove them must require removal of all the offsets.

Selasphorus rufus 
The Rufous hummingbird is a migratory species which spend spring and summer in my garden, and even as far north as Alaska. In the fall, they migrate as far south as Southern Mexico. They feed primarily on nectar, but are also insect predators, and I suspect bees and other pollinators to be included in their diet. I suspect they nectar on natives such as the locally common Lonicera hispidula as well as Arbutus menziesii which flowered earlier in the year. Crocosmia is one of their favorites in my garden, though I have also seen them visit some of the taller alliums and a few of the plants in the Boragnaceae including Anchusa azurea, not the typical hummingbird plant. They may also be one of the visitors of the Salvia sclarea in my garden, though I haven't directly observed this.

Gladiolus hybrid reproductive structures
For the first time this year, I have observed a hummingbird visiting the tall hybrid Gladiolus growing alongside the Crocosmia. This would be the first time I have ever witnessed any pollinator visiting the Gladiolus. If space is limited and you would like to plant for pollinators, I would suggest adding Gladiolus to the list of plants not to use along with Hemerocallis.

Sedum album with Apis mellifera
Fortunately there are plenty of other plants in the garden for bees to party on, including various sedums. The most floriferous, and prolific, is Sedum album. This is one of the easiest plants to propagate, one could simply tear a stem or few leaves off and toss them wherever there is soil (or sand, or gravel) and they'll form roots and grow. I wonder if I could pin it to a wall and see if it would root. Little to no water is required, and it flowers prolifically regardless.

Sempervivum sp. with Apis mellifera
I suspect flowers in the Crassulaceae such as Hylotelephium, Rhodiola, Sedum, and Sempervivum to be prolific nectar producers under conditions that would be considered harsh for other plants, or produce nectar with a very high sugar content, or both. In my experience these are nearly literal bee magnets, bees of many types being highly active on the blooms. While the flowers of Sedum tend to be small individually, the flowers of Sempervivum are much larger, and quite stunning.

Sempervivum sp. with Bombus vandykei
Sempervivum is a monocarpic genus, which means they do not flower for a long time (perhaps years), then use all of their energy to flower prolifically and then die. Fortunately they usually produce many offsets which live on after the "mother" dies. I am also hoping that this plant and another that are in flower will set seed for me to attempt. The advice I have been given is to surface sow, though there have been conflicting suggestions as to whether they need cold stratification or not.

Magnolia sp. with Apis mellifera
Another prolific nectar producer is the magnolia, typically a beetle pollinated genus. The genus is very ancient and evolved when beetles were the primary pollinators of insect pollinated plants. The flowers likely provided shelter, a place to seek mates, and as a source of food (pollen, primarily). The large waxy petals close up at night, enhancing the flower as a protective shelter. Beetles pollinate the flowers by climbing up the center structure, the base of which is lined with the anthers which are topped by a crown of stigmata (plural of stigma, not crucifixion wounds). Waxy petals at a certain stage, I surmise, may be more difficult for a beetle to climb, and so the beetle would be forced to cross first the anthers and then the stigmata to escape, transferring pollen on the way.

Lathyrus latifolius with Apis mellifera
Outside the garden in weedy sites like roadsides and forgotten pastures there are a number of weeds proliferating in the aftermath of the destruction of the native ecosystem many decades ago when the first settlers arrived to this area and "improved" it. One such weed is the common perennial sweet pea, a vine once planted to reduce erosion around roadside ditches and similar areas. It has few, if any, natural predators, and spreads primarily by seed. The vines have tendrils that wrap around nearby plants, including blackberry, smothering everything below.

Lathyrus latifolius with Apis mellifera
My previous assumptions were that nonnative sweet peas in general were not visited by honeybees, and were only visited by large bumblebees and carpenter bees. This year I have been forced to reassess my assumptions, and from the photos it is clear that honeybees do in fact visit sweet peas. Small Megachile sp. were also seen visiting the flowers. All of the bees access the nectar from the top of the flower, negating the need to move any petals.

Daucus carota with Apis mellifera
Years ago I had attempted to learn if honeybees visited the wild carrot, Daucus carota, that grows rampant in the area. I had been told by many beekeepers that honeybees either never visit the flowers, only visit them once every ten years, and that if they do visit the carrot it ruins the honey. This is the second year in a row I have observed honeybees on the carrot flowers, so the ten year wives tale, suspicious to begin with, goes out the window. Last year's honey from my own hive tastes great, and obviously they visit Daucus, so hereby goes the death of the other two assumptions.

Daucus carota with Apis mellifera
This particular plant, clearly in fact, particularly attractive to honeybees as for a two hour period while I was in this yard (an apiary, in fact) there were at least four honeybees each time I looked. Variation between individual plants doesn't need to be restricted to their appearance, although that is what is most apparent to us. Fitness and general hardiness will vary between individuals, as will the production of certain constituents inside the plant. This is apparent to breeders of plants used for their medicinal properties, such as Echinacea which have varying amounts of phenolic compounds, alkamides and polysaccharides between individuals not to mention species. It is not a great leap to consider nectar production to vary with sugar content or other factors.

Dipsacus fullonum with Bombus vosnesenskii
More common in agricultural sites than elsewhere, common teasel is an abundant biennial European native that is in bloom now. In my observations they were highly attractive to both bumblebees and honeybees, with emphasis on the former. Other observers have reported bumblebees being the most frequent visitors along with some dipterans in a site in Canada, while observations in a site in England reported the majority of floral visitors to be dipterans, primarily Syrphidae, followed by visitation by lepidopterans and bumblebees. (See Gucker et al. 2009). Studies also show that the plants are highly dependent on insect pollination and will not set seed when pollinators are excluded.Considering honeybees, another nonnative species, this may be a case of an invasive mutualism where two nonnative species support each others proliferation in the foreign land. Such a scenario was found to be the case with yellow star-thistle (Centaurea solstitialis), an important late season honey crop for beekeepers in my region. (See Barthell et al. 2001) I suspect this type of mutualism is much more common than we currently know.

Dipsacus fullonum with Bombus vosnesenskii
The plants are quite interesting, and if it weren't for their invasive nature I would grow them myself. They are taprooted with some roots reaching nearly two feet deep and an inch in diameter at the crown. Leaves are opposite and sessile, clasping the stem to form cups which can fill with water when it rains. This is hypothesized to protect the flowers from crawling nectar thieves or perhaps sap-sucking insects from climbing up the stem. The plants are biennial or monocarpic perennials, which means they die after flowering. They may have been introduced intentionally for use in the textile industry as early as the 1700s, or perhaps it was introduced accidentally or brought as an ornamental. In either case it is probably here to stay.

Hypochaeris radicata
Another invasive, not solely subjected to agricultural land, catsear (or false dandelion) is an abundant European native. They are perennial growing from thick fleshy taproots that break easily when pulled and regenerate into multiple crowns when disturbed. The seed goes airborne when it is ripe, or is perhaps distributed by birds. The flowers open when the sun comes up and close if it gets hot out, though I'll admit to not knowing the exact threshold at which this occurs. Collectively they bloom for the entirety of summer and into the first half of autumn, and are highly attractive to bees of various types. My employer has a sizable apiary located in this field, and I have personally observed multitudes of honeybees visit the flowers.

Hypochaeris radicata in the bee yard
I have mentioned in previous posts (see Apiaries, Apiaries II, & Apiaries III) that I work for a commercial beekeeping operation, and thus I am in the delightful position (literally) to view the bees at work. I have noticed that where there are many hives, there does not seem to be an abundance of native bees (with the exception of bumblebees). I am slightly at odds with honeybees, though I cannot argue their importance to us as we are today. The problem, I find, is that far too much prestige is placed on the honeybee while little to no attention or thought is given to the native bees. The reason we need honeybees to the extent that we do is because we have in many cases destroyed the habitat and food sources of many of the native bees (not to mention the other pollinators and other beneficial arthropods) in many parts of the country, creating places that are largely devoid of naturally occurring life (what does live there, we have put there). Honeybees directly compete for resources with the native pollinators, something that should be intuitive since plants are limited to the quantity of pollen and nectar they can produce per day.

The preeminence of honeybees is not without its benefits, as they are the poster children of environmental consciousness today. They are the martyrs of the current cultural trend towards pollinator conservation, and for that they deserve great credit. I myself would not be writing this today if it weren't for the honeybee, nor would I have one of the best jobs in the world. And I truly love my job!

Another day at the office!

Saturday, July 2, 2016

Floral Visitors

Bombus sp. on Echinacea purpurea
The following series of photos (twentieth post in the Floral Visitors series) will be short on commentary. This is a mixture of native pollinators (and honeybees which are nonnative) on both native and nonnative (garden) plants. The title, Floral Visitors, is descriptive of the fact that pollinators may visit a flower without necessarily contributing to pollination, thus they are merely floral visitors gaining sustenance from the floral rewards (nectar and/or pollen) while the plant gets nothing. This is a concept of anthecology (literally translating to flower ecology) in which plants support pollinators which may be important for the reproduction of other plant species. With enough biodiversity, the hope is that things balance out and everything gets what it needs. When diversity decreases or foreign species are introduced (introduction of nonnatives, benign or noxious) the system is thrown out of balance. I may sound hypocritical when saying this since I plant a lot of nonnative flowers in my garden, but my mere presence is disrupting the natural balance by taking up space (houses, roads, etc.) as well as byproducts of my presence (pollution from light, etc.). Perhaps I'm just being cynical, but when I started seeing life on the scale of my photography I started thinking about life in a new light, and that we ourselves are an invasive species.

My hope is that be illuminating the subject, and not only focusing on honeybees (no doubt paramount to our proliferation as a species), others may see the small life that I see which is more complex than we can know at our current level of awareness. Many of the smaller species of bees do not travel long distances, some no greater than 200ft, and so require food sources within that small span of their nest, or vice versa. Predatory and parasitic wasps, the vast majority are of little to no threat to people, require specific hosts of prey to survive. Either by pollination or by pest reduction, these small creatures are doing us a great service where they are prevalent, but the use of pesticides and the lack of nutrition (flowers) degrade their numbers thus requiring the use of honeybees, which is expensive.

A study by the Xerxes Society (Losey et al. 2006) reported an estimated $57 billion in ecological services provided by insects in the United States. Why would we not want to take advantage of these free services, use less pesticide, and take some of the pressure off of honeybees? The manufacturers of pesticides would not like to see this happen, and so the stereotype of insects being malicious and evil, the fear of these very small creatures, is propagated and shoved down our throats. Surely the number of benign, even beneficial arthropods far outweighs the ones harmful to us and our things, so long as a balance is maintained. What can you do? Plant flowers (preferably native to your region), and use less pesticide.

Apis mellifera on Navarretia squarrosa, an Oregon native.
Apis mellifera on Holodiscus discolor (oceanspray). Though the flowers of this native shrub were visited by a variety of small native bees and bumblebees, the honeybees practiced the peculiar behavior of "swimming" through the inflorescences.
Apis mellifera on Holodiscus discolor
Calliopsis sp. on Horkelia daucifolia
Calliopsis sp. on Horkelia daucifolia

Plantago lanceolata with a honeybee, making it rain pollen.
Apis mellifera on Allium cepa, aka common onion.
Lucilia sericata on Allium cepa
Gasteruption sp. on Allium cepa. Wasps in this family feed on flowers as adults, but the larvae feed on other hymenopteran larvae or their food stores.
Stenodynerus, a predatory mason wasp which preys primarily on moth larvae. The mason wasps are not very aggressive and are solitary rather than social, meaning that each female provisions their own nest. On Levisticum officinale.
Stenodynerus on Levisticum officinale
Stenodynerus on Levisticum officinale
Dolichovespula arenaria, the common aerial yellowjacket. This is a predator which hunts for other insect as protein rich prey for their young. Yellowjackets are social wasps, infamous for their defensive behavior. When they are not near the nest they can be quite docile.
Polistes dominula on Salvia sclarea. A European invader, this paper wasp is one of the few pollinators to be interested in the clary sage in my garden. Despite their nasty reputation (not unlike that of yellowjackets or other social vespids) they are docile when visiting flowers.
Xylocopa tabaniformis on Salvia sclarea. Large carpenter bees like this one are the most conspicuous visitors to the clary sage which has seeded around my garden. This species sometimes has a brownish coat of hair on its thorax, but the one in these photos is entirely black with a coating of pollen. Note that the stigma is in contact with the bees back in this photo.
Xylocopa tabaniformis on Salvia sclarea. Other bees, like carder bees (Anthidium sp.) have been busy on the clary sage.
Xylocopa tabaniformis on Salvia sclarea
Bombus vandykei  on Lavandula angustifolia. The males of B. vandykei are a completely different color than the females. Males are mostly yellow while the females are mostly black except for yellow on their faces and a few bands on their thorax and abdomen. They are particularly attracted to the English lavender.
Bombus vandykei  on Lonicera hispidula, a native honeysuckle.
Machimus sp. with prey on a tomato leaf. Robber flies are predatory and prey mostly on other dipterans and perhaps small hymenopterans or other small flying insects.
Blepharipappus scaber, visited by honeybees and bumblebees in Central Point in one of our apiaries.
Madia gracilis, visited by honeybees and small solitary bees in Ashland in another apiary. The flowers are only open in the morning, like other Madia species.
Apis mellifera with a dusting of pollen on Madia elegans in our Central Point apiary.
Apis mellifera on Madia elegans. M. elegans is a native annual which has flowers that are open in the morning but close as the sun and heat rise. This is an adaptation to prevent the nectar and flower itself from drying up. By the looks of it this is an effective strategy, bees were all over these flowers.
Apis mellifera on Madia elegans.
Madia elegans with plant bugs (Miridae)
Apis mellifera on Trifolium arvense in Gold Hill. This native clover blooms over a long period, yet honeybees seem only marginally interested in it. As other floral sources dry up, I may be seeing the bees work these flowers more and more.
Solitary bee on Coreopsis lanceolata in the garden.
Clockwise from left: Apis mellifera, Bombus vandykei ♂, and an unidentified Bombus sp. on Allium sphaerocephalon
Bombus vandykei ♂ on Allium sphaerocephalon. Male bumblebees are easy to photograph early in the morning or in the evening. Most males don't live in nests, and rather wait by flowers to meet a mate (or several) and ensure the survival of their genes. Therefore, I find them often just clinging to flowers over the course of the night, too cold to move, and thus excellent subjects for photography.
Bombus vandykei ♂ on Allium sphaerocephalon
Bombus vosnesenskii  on Dasiphora fruticosa 'Abbotswood'
Bombus vosnesenskii ♂ on Dasiphora fruticosa 'Abbotswood'
Bombus vosnesenskii ♂ on Hyssopus officinalis
Bombus vosnesenskii ♀ on Lathyrus latifolius. Females generally differ from males by having corbicula, or pollen sacks, which the males will not have.
A mated Apis mellifera queen, held by the thorax (and never by the abdomen). I work for a comercial beekeeper, and one of my duties is to catch queens for sale or our own use. Queen honeybees, unlike the working females of most other bee genera, only leave the nest to mate and so do not collect pollen or nectar or anything besides semen. This is on stark contrast to bumblebee queens which in the beginning of their life have to collect food and build a nest before they build up a work force. Bumblebee colonies are usually diminutive with up to 400 individuals while a productive honeybee hive can have upwards of 50,000. Honeybee queens can lay up to 2,000 eggs per day! Think about that.
Apis mellifera queens have large elongated abdomens, a result of numerous mating flights.