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Welcome to, the website of Bob Klips, a plant enthusiast living in Columbus, Ohio.
(Additional content at flickr Photostream and YouTube Channel)
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  Secular Lotus (Nelumbo lutea)
July 20, 2010
Catawba Island, Portage County, Ohio.

Today we drove to the tip of Catawba island and waved bye-bye to the kid, who caught Miller's Ferry to South Bass Island from which she hopped over to OSU's F.T. Stone Laboratory on wee little Gibraltar Island for five weeks of study during which she changed from someone who knew way less about entomology than me into someone who knows way more about entomology than me. Because that happened by her learning a lot about bugs instead of me forgetting a lot about them (although I think that may have happened also), it's great! Stone Lab is great! Go there and learn about bugs!

On the way back, her happy mom and I drove over the narrow inlet that almost makes Catawba Island an island, an inlet that is home to a huge and striking population of a huge and striking plant, Nelumbo lutea (Nelumbonaceae, the lotus family). Nelumbo, while it is a huge plant, is not a huge genus. There are just two species, constituting one of the many intriguing eastern Asian - eastern North American pairs of closely related but widely disjunct taxa. This peculiar distribution pattern is generally attributed to the fragmentation of a once-contiguous mixed mesophytic forest, the so-called "arcto-tertiary geoflora." This biome spread across the northern hemisphere during a warm period from 65 to 15 million years ago, when the continents were joined by the Bering land bridge. The Asian lotus species, Nelumbo nucifera, is an important spiritual symbol in many eastern religions, hence one prevalent common name for it is "sacred lotus." Because we have a different cultural tradition, based upon the separation of church and stamen, perhaps our species should be called "secular lotus." Perhaps, but it's usually just called "American lotus."

American lotus (Nelumbo luteo)
Secular (American) lotus at Catwaba Island inlet, Portage County, Ohio. July 20, 2010. 

The lotus family, Nelumbonaceae, consists only of the genus Nelumbo. These are perennial herbs with stout fleshy rhizomes, bearing disk-shaped leaves held above the water by a petiole (leafstalk) that is attached in the center of the leaf, an attachment type that is called "peltate."

Nelumbo lutea
Secular (American) lotus. Note peltate leaves held above the water.

The flowers are bisexual (hermaphroditic, i.e., containing both stamens and carpels), radially symmetric, large and showy. They are held stiffly upwards, just overtopping the leaves. The perianth consists of 2-5 outermost sepals, grading into 20-30 spirally arranged petals that are yellow in our species, pink to red in the Asian N. nucifera.

Nelumbo lutea flower
Secular (American) lotus. Note numerous, spirally arranged petals and stamens.

Each flower bears numerous widely separated ovaries loosely embedded in pockets just beneath the flat top of an unique enlarged inverted cone-shaped receptacle that eventually develops into Large Lotus Pods that are perfect for adding texture and a touch of rustic charm to your floral arrangements.

 Nelumbo receptacle
Nelumbo receptacle shortly after petal-fall, but before the development of rustic charm.

There are no styles; the stigma just sits atop the ovary, making the following zoomed-in piece of the picture above look really creepy, like a slew of alien pop-eyes all looking sort of, but not quite, at the same thing.

face of Nelumboi receptacle
Spooky zoomed-in view of Nelumbo receptacle, showing ovaries with sessile stigmas.

Numerous single-seeded fruits sunken into pits atop an enlarged receptacle constitute a peculiar type of "false fruit" that is sometimes called an "accessory fruit."  The only other vaguely similar accessory fruits are the wholly unrelated strawberry-type ones in the rosaceous genera Fragaria and Duchesnia. And yes, these lotus infructescences are those charming things you sometimes see in hobby shopes, for use in flower arrangements.

rustic charm
Arghh. Crass commercialism --the downside of "secular"?

The taxonomic position of Nelumbo has recently been revised. Traditionally, owing to a strong resemblance to the "water-lilies" Nymphaea and Nuphar which are also robust aquatic herbs with circular leaves and showy flowers composed of many separate parts spirally arranged, they were placed along with the water-lilies either in the same family (Nymphaeaceae) or alongside them in a separate family, both within the Order Nymphaeales.

During the past few decades there's been a revolution, still ongoing, in plant classification fostered both by a  greater acceptance of cladistic systems that are very strict about basing classification on phylogeny, and the wealth of data from DNA sequences. One very good guide to the new view of flowering plant systemetics is a great book called "A Tour of the Flowering Plants" by Priscilla Spears, published in 2006 by the Missouri Botanical Garden Press.

A Tour of the Flwering Plants

The book is based on the classification system of the Angiosperm Phylogeny Group. It includes a simplified tree that Spears calls a "map to help you find your way around the thowering plants." The "map," and accompanying text, explains that there is a small group comprising only about 3% of the flowering plants that diverged early from the flowering plant lineage, appropriately termed "basal angiosperms." Close to the ancestral seed plants, this group includes the water lilies (Nymphaeales), but not the lotuses. The Nelumbonaceae is in the Proteales, located farther up the tree in the "Basal Eudicots."

A map to help you find your way

Spears' treatment of the Nelumbonaceae includes the pleasant surprise that, in our flora, the closest relative of lotus is Platanus, our American sycamore! Look how well it's explained that the similarities between the two families of aquatic herbs is apparently due to convergent evolution. DNA data led to this findking, which has since been corroborated by a closer look at anatomical and developmental traits shared by Nelumbo and Platanus.

relatives do not always look alike

"Just So" Floral Extrafloral Nectaries on Sullivant's Milkweed?
July 19, 2010
Killdeer Plains Wildlife Area, Marion County, Ohio

An interesting feature exhibited by a few plants is the "extrafloral nectary," a glandular structure, usually at the base of a leaf, that produces an energy-rich sugary substance fed upon by some insect that is not involved in pollination. Typically the plant-insect relationship is a mutualism, wherein the insects, most often ants, contribute to the plant's well-being by by acting as bodyguards, fending off potential herbivores. Ants feeding on narrow-leaved vetch extrafloral nectaries last year can be seen here.

However, could there be nectaries that are "extrafloral" in the sense that they serve to attract non-pollinating insects, but that are "floral" in that they are indeed located on the flowers? It seems that maybe there's something like that going on with Sullivant's milkweed.

Sullibant's milkweed and ailanthus webworm moth
Sullivant's Milkweed. June 19, 2010. Killdeer Plains Wildlife Area.
See below for closer look at framed region.

Examining closely these flowers under a dissecting scope, Team Millkweed discovered a peculiarity that doesn't seem yet to have been reported in the fairly exhaustive scientific literature on milkweed flower structure and associated reproductive ecology. This is a set of 10 small, upwardly-projecting swollen flaps of tissue located where the hood rises off the central column. Each flap seems to loosely cover a small opening to the nectar-containing portion of the flower, i.e., the interior base of the hoods and the adjacent stigmatic chambers. Two of these flaps are depicted in the photo below, an enlargement of the boxed area of the photo above. Are they nectaries?

possible floral extra-floral nectaries
Possible floral extrafloral nectaries on Sullivant's milkweed.

One of the tenets of evolutionary theory is, of course, that a novel and intricate aspect of an organism constitutes an adaptation, i.e., a genetically based feature that increases its ability to survive and reproduce in a particular environment. However, while the value of a particular trait may sometimes seem apparent, strong experimental evidence is often lacking concerning the specific way (or ways) in which it benefits the organism. Thoughtful tentative explanations, intruguing but speculative, are often criticized. Such untested claims have been called "Just So Stories" in a derisive homage to Rudyard Kipling's set of childen's stories such as "How the Camel Got His Hump," etc.

Here's a "Just So" story about milkweeds. Perhaps this little set of openings to the nectar at the base of the flower provides a means by which nectar feeding but non-pollinating insects such as various bugs, ants, flies, and very small moths and bees can do their useless nectar feeding in a manner that interferes less with the beneficial pollinating foragers.

Here's a large milkweed bug, Oncopeltus fasciatus, feeding directly from the hood in a manner that might discourage a honeybee or a butterfly (potential pollinators) from occupying that same space.

large milkweed bug
Large milkweed bug drinking nectar directly from the hood.
July 19, 2010. Killdeer Plains Wildlife Area. Marion County, Ohio.

...and here's an instance where the bug seems to have its beak on the auxilliary structure that, if it is serving as a nectary, is doing so in a manner that facilitates feeding from a lower position, freeing the hood area for foraging by insects more likely to transfer pollinia.

large milkweed bug
Large milkweed bug drinking nectar from below.
July 19, 2010. Killdeer Plains Wildlife Area. Marion County, Ohio.

Here's a similar set of photos for an unidentified fly. First, plunk down in the hood.

Fly drinking nectar directly from the hood.
July 19, 2010. Killdeer Plains Wildlife Area. Marion County, Ohio.

...and here it is again, possibly exploiting the putative "floral extrafloral nectary," well out of the way of helpful honeybees.

Fly drinking nectar directly from below.
July 19, 2010. Killdeer Plains Wildlife Area. Marion County, Ohio.

Milkweeds are People, Too!
July 19, 2010.
Killdeer Plains Wildlife Area, Marion County, Ohio

One of the most enjoyable arguments you can have with someone, and it really doesn't matter which side you take, is whether or not individual non-human animals are distinctive like people are. The fact that humans are instantly recognizable seems important because, as social animals, we engage in reciprocal exchanges, altruism and trade. Hence we need to keep track of who is trustworthy, owes us a favor, or is a cheater. But what about all those animals that seem to look exactly alike, like toads and birds? The fact that I can't tell one from another might not mean much, because, not being a toad or a bird, I don't have an eye for what makes each one of them unique.

But there shouldn't be any debate about individual Sullivant's milkweeds. I have no idea why, but they're wildly variable and distinctive! Team Milkweed learned this during field work examining the potential for variation in gene flow among plants that consisted either of large multi-stemmed clones, or small few-stemmed ones. Based on appearance, we had to try to guess whether stems ("ramets") standing alongside one another belonged either to the same or to different genetic individuals ("genets").

To get a feeling for phenotypic variation in Sullivant's milkweed, I merrily pranced across the meadow, and snapped pictures of bundled-together pairs of adjacent stems that resembled each other in terms of flower colors, the shapes of their flowers, and the timing of flowering.

Sullivant's milkweed clone1
Two milkweed stems bundled together. July 19, 2010. Killdeer Plains.
Yellow-boxed area encloses flowers from both stems, featured in Gallery 2 below.

Here's  Image Gallery 1, demonstrating among-plant variation in Sullivant's milkweed.


Image Gallery 2 zooms in on the flowers, highlighting within-clone similarity.
Like the gallery above, it's also a good example of that annoying optical illusion where you see little gray squares that aren't really there.

Highslide JS
Uniform color; horns well above column; teardrop-shaped hood rim.
Highslide JS
Pale color; notch along upper outer edge of hood.
Highslide JS
Petals darker than hoods; sharply angled upper inner edge of hood.
Highslide JS
Dark star-shaped top of column.
Highslide JS
Darker pink rim of hoods.
Highslide JS
Round outer edge of hood.
Highslide JS
Horns curve down to touch column..
Highslide JS
Narrow hoods.
Highslide JS
Hoods upright (outer face vertical in side-view)
Highslide JS
Uniform color of petals.
Highslide JS
Darker color between corolla and hoods.
Highslide JS
Slanted hood rims.
Highslide JS
Flat-topped hood rims.
Highslide JS
Hoods short relative to corolla.
Highslide JS
Very short horns.
Highslide JS
Wrinkled petal edges.
Highslide JS
Dark-lined hood.
Highslide JS
Inward-curving hoods.

Many Milkweed Metazoans
Early-Mid July, 2010
Killdeer Plains Wildlife Area, Marion County, Ohio

It's plainly evident that the European honeybee, Apis mellifera, is the principal, indeed essentially the only, pollinator of Sullivant's milkweed here at Killdeer Plains. The bees are abundant, they forage extensively, and can regularly be seen carrying milkweed pollinaria. Here's yet another snapshot of a milkweed umbel featuring a foraging bee. MOUSEOVER the image to see more closely the pollinaria attached to her legs.

bee forages on milkweed
Honeybees forage avidly on Sullivant's killweed at Milkdeer. July 19, 2010.

...and they doubtlessly transfer pollinia from plant to plant as they travel.

bee flies away from milkweed
Honeybee carries pollinia at Killdeer Plains. July 19, 2010.

Here's yet another video of a honeybee foraging on Sullivant's milkweed. I've got dozens of these and have watched them all, closely, in a vain (fruitless vain, not egotistical vain, but maybe some of that too because they are quite nice videos) attempt to catch one in the act of pollinarium withdrawl or pollinium insertion. So far, no luck. Maybe milkweeds like their privacy.

Honeybee beset with pollinaria somehow manages not to insert any, or withdraw more.

Descriptions of milkweed pollination sometimes include a scenario wherein a foraging insect will get its foot caught in the stigmatic groove and, while trying to extricate itself, withdraw a pollinarium and then fly off with it. That may be happen, but it seems the pollinaria can be withdrawn cleanly, without all that drama. Nonetheless, bees often do get caught, and stay that way. Here's a honeybee with several feet snared.

bee stuck on milkweed
Honeybee fast on milkweed. July 10, 2010.

Here's the video they show in at scary special assembly every year at Honeybee High, to frighten those crazy kids, and keep them from playing on the milkweeds. But they never listen.

Honeybee snagged by milkweed flower. Sad, so sad.

But its not all honeybees all the time. Other insects visit the flowers, although they don't seem to carry many (any) pollinia. A terrific milkweed associate is the large milkweed bug, Oncopeltus fasciatus. This is a type of "seed bug" that does, indeed, feed largely on milkweed seeds. They are often accessed by piercing the wall of a milkweed fruit. But they also sip nectar, passing time until fruits arrive.

Oncopeltus fasciatus, the large milkweed bug
Large milkweed bug sips nectar. Killdeer Plains. July 19, 2010.

The large milkweed bug is an especially charismatic insect, as evidenced by the fact that it's all by itself on the cover of Eaton and Kaufman's excellent "Insects of North America."

Oncopeltus book cover
Large milkweed bug is a star! (This is very "meta.")

The milkweed bug is a good example of a true "bug" in the entomologist's sense, i.e., a member of the order Hemiptera, a large and diverse group with sucking beak-like mouthparts tucked beneath the body when not in use, and an incomplete metamorphosis wherein immatures resemble miniature adults, but with incompletely developed wings. In their suborder, Heteroptera (the "original" true bugs), the wings are oriented to give the insect a very flat-backed appearance, and each forewing is thickened in its forward half, giving it a somewhat two-toned appearance.

Here's an action-packed video of a large milkweed bug sipping nectar.

Buggy buggy slurpy slurpy.

Another brightly colored asclepiavore is a type of longhorn beetle, the red milkweed beetle, Tetraopes tetropthalamus. Eaton and Kaufman tell us that are 13 species of Tetraopes in North America, and that "most of the species studied so far specialize on one or a few species of milkweeds, the larvae mining in the roots, the adults feeding on the upper foliage and blooms." They do so in a manner the authors rightly characterize as "boldly in the open by day, their bright colors advertising their toxic nature." This species, they note, is the most numerous milkweed beetle in our region, and it feeds chiefly on common milkweed, Asclepias syriaca, but has been noted on at least two other Asclepias species. It is interesting to note that these two extremely bold beetles have evidently read the book and accordingly selected one of the few A. syrica plants (note the velvet underleaf) in a sea of Sullivant's.

Another Tetraopes peculiarity that Eaton and Kaufman mention is that "each eye is divided by the antennal socket, hence they are often called 'four-eyed beetles'." It's true.

MOUSEOVER the IMAGE for a ZOOM-CROP showing the nifty FOUR-EYE-NESS
red milkweed beetle
Red milkweed beetles mating and feeding on common milkweed. July 19, 2010.

One of the few easily identified small moths sometimes called "microlepidoptera" is the Ailanthus webworm moth, Atteva punctella. Their larvae occur in communal webs on tree-of-heaven (Ailanthus) trees.

Ailanthus webwormAilanthus webworm moth sips milkweed nectar. July 19, 2010.

Here's another apparently non-pollinating insect, a fly, dipping deeply onto a Sullivant's milkweed hood.

fly dipping into milkweed hood
Fly feeding on mikweed nectar at Killdeer Plains. July 14, 2010.

...and here's a video of the fly feeding.

Fly feeds on milkweed nectar at Killdeer.

It was an rare treat to see a lightning bug wandering about on a milkweed umbel. It looks like Photinus pyralus, a common species.

Photinus firefly on milkweed umbel at Killdeer.

Moth Mullien and Syrphid Fly
July 4, 2010
Miami County, Ohio

I saw a pretty weed driving to the milkweed meadow. Who knew a flower could drive? This roadside plant, a native of Eurasia, is moth mullen, Verbascum blattaria (Scrophulariaceae, the figwort family). It comes in two colors: yellow and white. Last year at this time I snapped some pics and a little video of the white form. According to some sources, the yellow form is generally more common, but it seems the white ones are more frequent in central Ohio. Here's yellow moth mullien.
yellow moth mullien
Yellow moth mullien. July 4, 2010. Miami County, Ohio.

Moth mullien has exceptionally long-lived seeds. This was decisively shown through a wonderful  long-running plant ecology experiment in Michigan, where, in 1879, William J. Beals buried 20 vials of soil containing seeds of 21 different weeds. He developed a protocol for their excavation and viability-testing.  The most recent withdrawls --there are 5 bottles remaining and the next will be dug up in 2020 --have consistently yielded only moth mullien, plus, in some instances, traces of a few other species. 

It's called "moth" mullien because its filaments (the anther-supporting stalk portion of the stamens) are plumose, like a moth's antennae. The specific epithet "blattaria" is a reference to roaches, which at one time were thought to be repelled by the plant.

moth mullien with syrphid fly
Moth mullien with syrphid fly. July 4, 2010. Miami County, Ohio.

Midnight and Morning Milkweed Marathon
Late June and early July, 2010
Killdeer Plains Wildlife Area, Marion County, Ohio.
Team Milkweed is hard at work, discovering what pollinates Sullivant's milkweed (answer: honeybees), when the  pollination occurs (answer: daytime), and whether nectar-foraging bees on small clones and large clones differ in whether they are carrying pollinia from the same, or different plants (to be determined).
Team Milkweed
Team Millweed at Killdeer Plains. June 27, 2010.

Since Milkweed Midsummer is a longstanding tradition, a description of Sullivant's milkweed can be found here, and more detailed information about its pollnation is here.

This summer, the focus is more on the broader pollination ecology of the species rather than just its breeding system and potential for hybridization, so we are noticing more insect species exploiting these beautiful plants. This, for example, is not what it looks like, unless it looks like a fly that looks like a bee. It's a type of flower-fly (Order Diptera; Family Sirphidae) that is a bumblebee-mimic, most likely Mallota bautias.

Mallota on Asclepias sullivantii
Mallota flower fly forages on milkweed at Killdeer. June 27, 2010.

Milkweed flowers don't ever close, so do night-time nectar feeders seek nectar from them? Indeed, there have been studies showing a small but substantial amount of nocturnal pollination of the closely related common milkweed, Asclepias syriaca. Our day/night study involves placing mesh bags over the flower clusters that prevent insect access to them during either the daytime or the nightime. Here's a night-flying moth that seemed more interested in the bagged flowers than all the others nearby. It may be a type of owlet moth, i.e., a member of the lepidopteran Family Nolidae (formerly included in Noctuidae).

moth on netting
Nolid moth on nylon pollinator exclusion bag. July 1, 2010.

While generally the blossoms are ignored at night, a few visitors show up. Here's another nolid moth and, if you look closely, there are at least two of what I guess are male mosquitoes, because they have plumose antennae and are feeding on nectar.

moth on milkweed
 Nolid moth and male mosquitoes sup on milkweed nectar. July 1, 2010.
Note also a snagged honeybee in the background at the upper edge of the photo.

While prowling around at night it was nice to see a monarch chrysalis on yellow fox sedge (Carex annectans). These "emerald houses with golden nails" are quite common this summer.

monarch chrysalis
Monarch butterfly chrysalis. July 1, 2010.

Here's the same sedge in the daylight.

Carex annectans
Yellow fox sedge with monarch chrysalis. July 1, 2010.

It is intriguing and informative to observe a variety of insects, day and night, on Sullivant's milkweed blossoms. Nonetheless, many hours of observation indicate that just one insect, a non-native species, the honeybee, is practically its only pollinator at Killdeer Plains. Here's one foraging; note the little yellowish pollinaria attached to her feet.

Honeybee forages on Milkweed at Killdeer. July 10, 2010.

There are a few other milkweed species in the area. One of them, whorled milkweed, Asclepias verticillata, has such small flowers and a small stature, that it is easy to miss.

Whorled milkweed at Killdeer. July 2, 2010.

Whorled milkweed, while scarce at Killdeer, is exceedingly abundant in some parts of northwestern Ohio, where it is a weed, for example along interstate highway I-75 in Wood County. The blossoms are tiny, but with the same flower structure as its larger congenors.

Asclepias verticillata
Whorled milkweed at Killdeer. July 2, 2010.