Agalinis fasciculata (Fasciculate False Foxglove)

Agalinis fasciculata, known as the fasciculate false foxglove and beach false foxglove was one of the more fascinating and unexpected plants I became acquainted with this year. A member of the Orobanchaceae family, this species is an annual hemiparasitic plant that does well in poor and sandy soils. I photographed these plants at the Missouri Mines State Historic Site in St. Francois County.

The genus Agalinis comes from the Greek – agan, meaning ‘very’ and linon, refering to ‘flax’, apparently in reference to the similarity of the flowers to those of flax. The species and common names refer to the fasciculate, or bundled manner in which the leaves are attached to the stem – something I failed to take any photos of this year. In my defense, much of the stem and leaves of these plants in mid-September were beginning to senesce and were not very photogenic.

Agalinis fasciculata (Fasciculate False Foxglove) in glorious bloom at Missouri Mines State Historic Site.
Many species of bees and flies like this syrphid fly act as pollinators of Agalinis fasciculata.
A rare six-lobed corolla of Agalinis fasciculata. This was the only six-lobed flower I found among hundreds I observed on this visit.

Southern Black Widow (Latrodectus mactans)

This southern black widow was found at Sand Prairie Conservation Area in Scott County MO. Quite unusually, she had built a web in the open within the tallest branches of a Polygonum americanum (American jointweed), where she had just dispatched a Dielis plumipes (Feather-legged Scoliid Wasp).

This southern black widow female was found wrapping up her prey, a feather-legged scoliid wasp at Sand Prairie C. A.
My shadow cast as I took photos caused her to head for cover.
I also found a male southern black widow moving about the boundaries of the female’s web.

Photographic Observations of a Communal Nesting Sweat Bee (Agapostemon virescens)

For the past few years I have noticed a good number of native bee nest holes along exposed sections of bare soil at one of my favorite hiking and nature observation sites – August G. Beckemeier Conservation Area in St. Louis Co., MO. This past spring I finally decided to make this a project and set about a quest to make some images of these gals provisioning their nests. As usual, I wound up learning along the way.

An Agapostemon virescens pauses at the entrance of the largest of the communal nest entrances I observed. It is impossible for me to accurately count the number of females using this ~ 10 cm tall conical entrance, but I observed six individuals at one time on or hovering above the entrance.

As is commonly known, many of our native bees are solitary and nest without close contact or cooperation in regards to conspecifics. At the opposite side of this spectrum of sociality in the Hymenoptera are most species of bumble bees and the honeybee. These bees are considered truly social, or, eusocial. The characteristics necessary to be considered a eusocial species are 1) cooperative care of offspring of others within the colony, 2) overlapping generations within a colony of adults, and 3) a division of labor into reproductive and non-reproductive groups. Many of our bee species lie somewhere between these two extremes. The bee of focus here, Agapostemon virescens, lies early in the area we call being presocial, aka parasocial.

Two Agapostemon virescens females exiting a communal nest entrance having dropped off their loads into their individual cells.

Let’s clarify the differences between a presocial species such as A. virescens and the eusocial honeybee. The honeybee shows all three necessary characteristics of a eusocial species. The individual workers obviously care for brood that are not their own – they don’t even have offspring of their own, instead spending much of their lives caring for the offspring of their queen (sisters). They have multiple overlapping generations within the hive in a particular season, as well as across multiple seasons and as just mentioned, there is a division of labor into reproductive and non-reproductive castes. A. virescens on the other hand, is not nearly as cooperative. Individuals of this species share basically just a front door to their brood chambers and nothing more. After entering the communal nest, each female builds their own brood sub-chamber cells and each provisions their own by processing pollen into cakes and leaving them in their respective brood chambers. There is no brood care after the egg is deposited and the sub-chamber sealed. The offspring then emerges later in the summer.

So, what are the pre-conditions necessary for the eventual development of more complicated forms of sociality, i.e. eusociality? Or more directly, what advantages are there in adopting more of a social lifestyle if we assume the starting point was a solitary existence? Scientists consider two important pre-conditions need be met for the evolution of eusociality. First, the species offspring must be altricial, or require a great amount of parental care in order to reach maturity. Second, there need be low reproductive success rates of solitary pairs that attempt to reproduce. Here is what is believed to be the primary driver that pushed A. virescens into this presocial condition.

A sentry Agapostemon virescens stands guard at the communal nest entrance allowing only conspecifics to enter. This guarding of potential kleptoparastism is regarded as the primary benefit that led to communal nesting in this species.
This sentry Agapostemon virescens closely inspects an incoming conspecific. How it is determined who stands watch while its neighbors forage is not known.

Kleptoparasitism is where one animal takes advantage of the hard work of another by taking their prey or collected foods. In this case, we are primarily concerned with the large group of bees known as cuckoo bees. Kleptoparasitism has evolved numerous times in the Hymenoptera and cuckoo bees lay their egg on or near the host’s provisions. The parasite will hatch first and eat the host’s pollen and will often kill and eat the host’s larvae as well. With such an obviously successful reproductive strategy, it should come as no surprise that there would be a strong selective advantage of finding ways to thwart these parasites. In the case of A. virescens, evidence suggests that by communal living as described here, the rate of kleptoparasitism is much lower when compared to related species that have the completely solitary reproductive strategy.

A busy day of bringing in pollen provisions for these Agapostemon virescens sweat bees.

I guess the obvious next question is how in the world could eusociality evolve from this state? This is a fascinating story that involves terms like kin selection, altruism and haplodiploidy. It also involves a good deal of math and explanation from some of the greatest evolutionary thinkers since the time of Darwin (read anything by William D. Hamilton for example). It is also well out of the scope of this piece. But, I hope it is clear that before getting near the high rung of eusociality on this ladder, that a small first step like seen in this example would be necessary.

Although Agapostemon virescens sweat bees are communal nesters, this photo gives a clue that they are not cooperative foragers like the honeybee. Each of the three returning females is carrying different colored pollen, indicating different pollen source plants for each.

I hope I got most of this correct enough. It’s been a long time since I took Zuleyma Tang-Martinez’s Evolution of Animal Sociality class at University, which I thoroughly enjoyed. Please feel free to leave a comment to correct or clarify or ask a question.

Much of what I covered here and a lot more can be found in Malte Andersson’s The evolution of eusociality (Ann. Rev. Ecol. Syst. 1984. 15:165-89

The evolution of Eusociality

Missouri Orchids – (Platanthera flava var. herbiola) – Tubercled Orchid

Here is my last new orchid for the season. It is also probably the orchid I had to work the hardest to find in this entire project. Platanthera flava var. herbiola is classified as S2 (imperiled) in Missouri due to the very few remaining populations. This is an orchid that likes its feet wet and can be found in a variety of habitats containing moist to wet soils. After trying unsuccessfully in 2020, Pete and I went back to the same location this year – a wet prairie in southern Missouri in early June. This was very tough searching as the high temps, strong sun and saturated air created a potentially dangerous heat index. We tried our best, slowly slogging through the already quite thick prairie. Just when it looked like Pete was wanting to throw in the towel, we came across a patch of less-dense vegetation with water about ankle-high. Here we found young orchids that numbered in the hundreds. Unfortunately, most were on the early side and were not fully flowering but we did find a few that made us happy. We also found that many of the orchids in this group had grazed top leaves – most likely from white-tailed deer that usually find orchids to be very appetizing.

Platanthera flava var. herbiola, the 33rd Missouri orchid species I have been able to see and photograph.

There is another variety of this orchid, P. flava var. flava that is also found in Missouri. Recently Pete and I tried to find this in most of the known locations but came up short. As of now my quest stands at 33 of 36 orchid forms that can be found in the state (36 is my accepted number and others may disagree). In addition to P. flava var. flava, I also need to find Coeloglossum viride (Long-bract Frog Orchid) – this species is known from a single location in the state but apparently does not flower and Epipactis helleborine, the broad-leaved helleborine – the exotic orchid that is becoming naturalized in this state but originates in Asia and Europe. Finding these three remaining orchids should be quite the task and I look forward to attempting these next year.

The longer floral bracts, looser inflorescence and three leaves are among a few of the characteristics that identify this as Platanthera flava var. herbiola and not P. flava var. flava.

I have run into a couple other circumstances in our orchid flora where intermediate forms have caused problems in determining the identity of a plant or population. Where these two varieties overlap, as they seem to do in Missouri, there are intermediate forms between these two varieties as well. I will hope that when I do find a potential flava variety that this will not become a problem.

Platanthera flava var. herbiola is the more southern variety. Here another distinguishing characteristic can be observed – the lip of variety herbiola is longer than it is broad, whereas the lip of variety flava is as broad as it is long, being almost square or circular in appearance.

Synchlora aerata (camouflaged looper)

Back in mid-June I discovered a number of Synchlora aerata (camouflaged looper, wavy-lined emerald moth) that were using our coreopsis as host. Not only are these spectacular adult moths in the family Geometridae, but they are obviously special while in the larval phase as well. These caterpillars are known for attaching bits and pieces of the plant tissues they feed on (often flower petals) to their backs as means of camouflaging against their predators.

The Synchlora aerata, on Coreopsis sp. in a suburban wildflower garden in St. Louis County, MO, USA
I often find these guys with their camouflage dull, dry and not very attractive. You can change this pretty easily by placing them in a container with a fresh native flower of your choice. Hopefully within a day or two the caterpillar will have adorned itself with a fresh and colorful coat!

From the Home Garden – Callirhoe bushii (Bush’s poppy mallow)

Photographed on June 11th of this year. I chose this species of Callirhoe because it is the most likely to stay small and behave in the front yard beds. However, I fear that the primarily clay soils will get the worst of it and it won’t stick around long. Of the three plants, only one made it to flower. These guys are adapted to more xeric and well-draining soils.

Leaf and flower of Callirhoe bushii (Malvaceae)

A Lizard Beetle

The Languria bicolor (Erotylidae) is placed in the tribe Languriini (lizard beetles). Larvae of lizard beetles develop within the stems of plants and adults feed on the tissues and pollen of the same or nearby plants. This individual was found in July 2021 at the Beaumont Scout Reservation, St. Louis County, Missouri.

Small Carpenter Bee (Ceratina sp.)

Here we have a few shots of a small carpenter bee that was very cooperative this past April at Beckemeier Conservation Area as it nectared from a spring beauty blossom. This is one of the bees that nests and overwinters in old broken pithy stems that it excavates. So here is who you might be helping by leaving your dead stems sit through the winter.

The Beardtongues?

Here is a genus that I find interesting. The Penstemon is made up of approximately 270 species and is the largest genus of flowering plants that are endemic to North America. Now classified in the plantain family (Plantaginaceae), this is a very diverse genus found across a variety of habitats and altitudes. Most species should be readily identified as a Penstemon due to their unique flower morphology. The corolla is a fused tube, comprised of five petals that can be identified as lobes in a two on top, three on the bottom configuration. Inside the corolla you will find two pairs of stamens with anthers pushed towards the top of the open mouth. In between the fertile stamens is a staminode that lies towards the bottom of the tube. This sterile modified stamen usually ends in a brush-like structure. This is the eponymous “beardtongue”. The generic name, Penstemon, meaning “stamen-like”, also refers to this staminode.

I got to meet four species of Penstemon in bloom this year – two of which I planted in the garden. I was happy to see them bloom in their first season.

Penstemon pallidus (pale penstemon) from my front garden in St. Louis Co., MO.
A closeup of Penstemon pallidus flowers. Note the yellow beard (staminode) that is thought to aid in pollination by pushing hymenopteran pollinators towards the stigma and anthers located at the top of the corolla tube. Also note the dark nectar guides that point towards the back of the tube.
Penstemon digitalis (foxglove beardtongue) is the most common and least particular member of the genus in eastern Missouri. These plants were found in a field at Beckemeier Conservation Area in St. Louis County.
The flowers of Penstemon digitalis are mostly white in color and have a relatively long blooming period compared to other local members of the genus.
Penstemon tubaeflorus is a showy white penstemon that is found primarily in the southwestern quadrant in Missouri. These photos were taken at Tingler Prairie Natural Area near West Plains, MO.
Here you can see why this species gets its common name of ‘trumpet beardtongue.’
The large and showy flowers of Penstemon cobaea (prairie beardtongue). These were photographed from the author’s front garden in St. Louis Co., MO.
I used focus stacking to capture the details in the flower of this Penstemon cobaea (prairie beardtongue). Note the two pairs of stamens that wrap around the inside of the corolla and present their pollen-filled anthers at the top. The stiff brush-like beard of the staminode pushes would-be pollinators towards these reproductive organs.

-OZB