"Open House" Field Trip

Betsy and I plan to be at Cranberry Lake from about 12:00 to 2:00 pm, weather permitting, this Saturday and Sunday. We will be looking for dragonflies and damselflies with both binoculars and a spotting scope.

Here's some of what we might be able to see:





Feel free to join us at any time during the "open house."

Species Spotlight: Blue-ringed Dancer

We turn our attention now to a damselfly with beautifully brilliant colors—the Blue-ringed Dancer. This species is widespread in the midwest and along the east coast, but also extends into Arizona and Southern California, which is where we see it. Here's the range map, from OdonataCentral.



One of the best places to find this species, at least in Arizona, is perched on a rock in the middle of a swiftly-flowing stream, as in the photos below:





Identification is easy for this species, with the prominent blue rings along the middle sections of the abdomen. Also be sure to take a look at the thorax and head, where the rich blue colors are particularly vivid:





Notice the tendency of this species to hold the folded wings well above the abdomen when perched—a characteristic of most dancer species. In fact, you can usually tell a dancer from quite a distance just by seeing how it holds its wings. In addition, dancers tend to "flick" their wings open and shut a time or two just after landing.

One individual, seen at the Gilbert Water Ranch in Gilbert, AZ, had a weird looking face. It was kind of pink. At first it brought to mind a scene from the 1958 movie The Fly, where the hero's body has been mixed up with that of a fly during a teleporter experiment—he now has the head of a fly, and the fly has a human head. They try to find the mixed-up fly by looking in the garden for one that has a pink head. Here are some posters and photos from the movie:









Well, that was a terrifying movie for a youngster back in those days. Fortunately, the case I encountered at the Water Ranch wasn't like that at all—it was just a damselfly with a lot of pink on the head. I got closer, and finally had a better look. Here's what was going on with this individual:



Wow, one eye blue, the other pink. You don't see that everyday! Searching the bushes for a damselfly with a pink head sure made me think about the The Fly, though.

What Big Eyes You Have!

Dragonfly eyes are beautiful and impressive. As an example, take a look at the eyes of a young Blue Dasher, shown below.



Notice the large reddish areas on the top of the eyes. These areas are referred to as the dorsal foveae—so named because they are on the top (dorsal) surface of the eyes, and are the parts of the eyes with the sharpest vision (analogous to the vertebrate fovea). The name is ironic because the word fovea is Latin for "pit," and indeed the fovea (area of sharpest vision) of a vertebrate's eye is a pit on the surface of the retina. In the case of a dragonfly, the fovea is not a pit, but it is the area of sharpest vision.

Here's another view of dorsal foveae in dragonflies, this time on a Cardinal Meadowhawk.



One of the interesting aspects of dorsal foveae is that the ommatidia (individual eyes) in them are rather large, and point primarily in one direction. This gives the sharp vision mentioned above. It also means that when you look directly at the dorsal foveae, you see an exceptionally large pseudopupil, as shown below.



My, what big eyes you have!

This effect is perhaps best seen in Blue Dashers. Take a look for the "big eyes" next time you get a good look at a Blue Dasher. You can read more about the eyes of dragonflies in my new field guide: Common Dragonflies and Damselflies of the Pacific Coast.

Species Spotlight: Blue-eyed Darner

The Blue-eyed Darner is one of the easiest darners to identify—even on the wing the intense blue color of their eyes, and the blue stripes on their thorax, are a dead giveaway. Here's the spread on the Blue-eyed Darner in my field guide Common Dragonflies and Damselflies of the Pacific Coast:



The flight picture above shows the blue eyes and the blue thorax stripes that jump out as this dragonfly zips by you on a warm summer afternoon.

Also, notice the photo of the female in the lower right corner of the spread. It shows a "bump," or "tubercle," under segment 1 of the abdomen. This is a distinctive feature of Rhionaeschna darners—commonly referred to as neotropic darners—though it is not always easy to see or photograph. My picture was taken at Butchart Gardens, where a female Blue-eyed Darner flew in and landed on a structure in the middle of a water fountain at just the right angle to show the bump. Mosaic (Aeshna) darners lack the tubercle.

Here's a view of another distinctive feature of Blue-eyed Darners—the forked appendages on the males.



No other male darner has forked appendages, so it's a great field mark—not that it's really needed!

The next photo shows an interesting feature that is present in all darners, but is particularly noticeable in Blue-eyed Darners. It is the small, sharp projection on the front of the thorax, just behind the eyes. This is an "egg tooth" that these darners use to break through their larval skin during emergence.





The Blue-eyed Darner is indeed a study in blue. They perch often in sunlit bushes near the shore, and are very cooperative photographic subjects.



One final feature that distinguishes neotropic from mosaic darners is shown in the next photo. Notice the light area on either side of the black "T-spot". This is found in neotropic darners, but not in mosaic darners.





So many interesting features to look for next time you're identifying darners!

Mount Baker Newcomers

On our recent trip to Mount Baker, Betsy and I enjoyed more than just the scenery.



We also saw some species we had never seen before at that location.

First, we saw the Hudsonian Whiteface. This is a handsome dragonfly with a black body and eyes, dark red markings on the thorax and abdomen, and a chalk-white face.



Notice also the white wing veins that extend beyond the stigmas, giving the wings a whitish tip. This field mark isn't mentioned in other field guides, but you will find it in Common Dragonflies and Damselflies of the Pacific Coast.



Another new species for this location was the Black Meadowhawk, shown below:



It's not a surprise to see this species here, because we've seen it at high altitude before—most notably at the small pond at Rainy Pass.

What a day we had a Mount Baker. We plan to return soon to enjoy the fall foliage.

Wing Grabbing

Betsy and I just returned from Cranberry Lake, where we saw Hermit Thrushes, Varied Thrushes, and Red-breasted Thrushes (American Robins)—a three thrush day. The weather was calm, with temperatures in the mid 60s. A beautiful day at the lake.

We also saw lots of darners there—mostly the Happy-face Darner (Paddle-tailed Darner). Males were patrolling floating logs along the shore, looking for females. We were looking for them too, hoping to observe the behavior that occurs when males attach to females.

As I've described before, when a male darner lands on a female and attempts to attach the tip of his abdomen to the back of the female's head, part of the process involves the male grabbing the female's forewings and pulling them forward—until they're pointing straight ahead. Apparently this induces the female to lower her head, and expose the area where the male attaches, making it easier for him complete the process. 

We discovered this behavior at Beaver Pond in Winthrop, WA in a slow-motion video I took of a male attaching to a female. I published a description of the behavior in the Argia article below, and since then we're always on the lookout for another opportunity to see it. Well, today at the lake we got another chance, and I was able to get the following photo showing the male in the process of attaching to the female.

The female is on the bottom, and the male has his abdomen bent forward to bring the tip in contact with the back of the female's head. Notice also that the male is grabbing the female's forewing—on the left side, where we can see it—and is pulling it forward. The annotated version of the photo, shown below, indicates the key features.

The interesting thing about this photo is that I hadn't realized before that the male grabs the female's wing with all of his legs—not just the rear legs. This adds a new element to the behavior. When I watch the original video again (link below), I can now see that after the rear legs reach down and grab the wings, the middle and front legs join in and help in pulling the wings forward, and giving the female a "bear hug."

Here's a reprint of the Argia article I published describing this behavior for the first time.

Attaching in Tandem: The Role of “Wing Grabbing” and “Wing Pulling”

James S. Walker

One of the most interesting aspects of dragonfly behavior is the fact that they mate like no other creatures on Earth.  They begin by attaching in tandem, and from there they maneuver into the wheel position.  After mating, many species remain in tandem for quite some time as the eggs are laid.  Clearly, the tandem coupling is of crucial importance to dragonflies.

As a result, the actual process of attaching in tandem is of some interest.  It looks fairly straightforward, but the connection usually happens so quickly that few details can be discerned.  In this article, I present observations from a slow-motion video of darners attaching in tandem.  As we shall see, the male “grabs” the female’s forewings, and then “pulls” them forward to an extreme extent as it completes the attachment process.

Before the Attachment

The observations reported in this article come from a slow-motion video (1/4 speed) of a female Paddle-tailed Darner (Aeshna palmata) in the process of laying eggs on a floating log.  She was spending a lot of time probing the log for suitable places to lay eggs.  When satisfied with a location, she would visibly “bear down” as she made a cut into the wood.  

There were several males of the same species patrolling the area.  As I filmed the female, one of the males came swooping in and literally “pounced” on her – landing on her thorax.  Figure 1 shows the male just as he made contact with the female.  It was a sudden impact for the female, and the male immediately began to bend his abdomen downward and forward to bring his appendages toward the female’s head.  It took only 0.67 s to complete this action, and the male then proceeded with the attachment process in earnest.

Figure 1 A male darner makes first contact with a female on a log.

The Attachment Process

The process of attaching in tandem can be thought of as occurring in three phases.  The first phase appears to be a preliminary attachment, the second phase involves “grabbing” and “pulling” of the female’s forewings, and the third phase is the completion of a firm attachment to the female’s head.  In what follows, I flesh out the details of each of these phases.

Phase 1: Preliminary Oscillations (0.36 s) This phase begins when the male first brings his appendages into contact with the back of the female’s head.  After apparently latching onto the female’s head, the male then executes a series of 8 or 9 sideways oscillations of his abdomen with a frequency of roughly 60 Hz.  One might think this would complete the attachment, but there is more to come.

Phase 2: Wing Grabbing and Wing Pulling (0.60 s) At the beginning of phase 2, the female’s wings are in their normal position, as illustrated in Figure 2 (a).  The male now reaches his rear two legs downward, putting them between the forewings and hindwings of the female.  He then begins to “grab” the forewings, and “pull” them forward.  

When the forewings reach the position shown in Figure 2 (b), the legs are pulling quite hard on the forewings, causing a considerable distortion in their membranes.  In fact, this particular female had a tear in the middle of her left forewing, and the male’s rear leg fit nicely into the tear.  It appears that tears like this on a female’s forewing, which might ordinarily be chalked up to a bird attack, could instead be a sign of the rough handling that occurs during attachment.

Figure 2 Positions of the female’s wings during attachment.  (a) Normal position of wings.  (b) Wing grabbing begins, and forewings are pulled forward.  (c)  Final phase of wing pulling, resulting in forewings that point straight forward.

The male continues to pull the female’s forewing toward her head.  Eventually, the forewings are pointing directly forward, as in Figure 2 (c), and the plane of their membranes is vertical rather than horizontal.  The male now does a “bear hug” on the female, pressing her forewings firmly against her thorax.  The male holds the wings stationary against the thorax during this “bear hug”, and remains stationary himself, for about 0.22 s.

Phase 3: Final Oscillations (0.29 s) In phase 3, the male executes a series of roughly 15 side-to-side oscillations of his abdomen, like he did in phase 1.  The oscillations are again at about 60 Hz.  At the end of phase 3 the male has a good attachment, and he releases the female’s forewings.  He then begins to flap his wings.

After the Attachment

I mentioned that at the end of phase 3 the male begins to flap his wings.  I didn’t say he takes off, because in this case the female held on tightly to the log and prevented him from flying off with her, as shown in Figure 3.  The male tried and tried to dislodge the female – a couple times he even fell into the water briefly during his efforts.  After 7.5 s of futile attempts to takeoff, the male finally detached from the female and departed the scene.

Figure 3 After attachment the male darner tries to takeoff, but the female holds on to the log.

Additional Observations

After observing this “wing grabbing” and “wing pulling” behavior, I looked more carefully at other female darners that were potential mating partners for a roving male.  I have seen the same behavior in 3 additional cases now, clearly indicating that grabbing the forewing and pulling it forward are standard parts of their attachment process.

I haven’t had much of a chance to observe attachment in other families of dragonflies yet.  However, I looked back over earlier slow-motion videos to see if the behavior appeared in any of them.  In fact, one of my videos did show wing grabbing.  This involved a pair of Black Saddlebags (Tramea lacerata) that I filmed doing their egg-laying behavior of releasing the female, letting her drop to the water to lay an egg, and then immediately reattaching in flight.  The video showed clearly that the male grabbed the female’s forewings and pulled them forward as he reattached.  He pulled the forewings into the position shown in Figure 2 (b), which prevented them from flapping until attachment was accomplished.

I’ve also found a few published results that may bear some relation to the behavior reported here.  For example, Corbet reports that “… when a male Anax junius in tandem is attacked (bitten) by another male, the tandem male shakes its abdomen in a convulsive movement detectable to the human observer only when portrayed in slow motion” (Corbet, page 276).  This sounds similar to the oscillations observed in phase 1 and phase 3, though perhaps performed for a different reason.

In terms of interactions with the female’s wings, the following may be somewhat related.  Corbet states that when a male Ischnura graellsii detects a female he rushes at her, “… grasping and sometimes even biting her wing bases and simultaneously beating her wings with his abdomen …” (Corbet, page 485).  Similarly, he reports that a male Hemiphlebia mirabilis “… flew to a perched female and straddled her, holding her wings over her abdomen with all his legs for one or two minutes …”, (Corbet, page 485).  Though certainly quite different from the “wing grabbing” and “wing pulling” described here, these observations may be part of a larger repertoire of wing interactions during attachment.

Conclusions

It will be of some interest to see just how widespread the wing grabbing and wing pulling behavior is among dragonflies, and whether it is also done by damselflies, as suggested by the above observations.  

Some of the questions that can be addressed in future studies include:

•  Is wing grabbing done intentionally, or does the forewing just “get in the way” of the male’s hind legs?

•  If wing grabbing is intentional – and it looks that way in the video – what role does it play in attaining a solid attachment?  Does grabbing the forewings give the male a better grip on the female?  Does pulling on the wings trigger a response that helps to induce a female to mate?

These and many other questions can be addressed with additional observations over the years.  I know I’ll be looking more carefully at the attachment process in the dragonfly seasons to come.

Acknowledgements

I would like to thank Dennis Paulson and Betsy Walker for helpful conversations and feedback.

Literature Cited

Corbet, P. 1999.  Dragonflies: Behavior and Ecology of Odonata. Ithaca, New York: Cornell University Press.

Online Material

A male Paddle-tailed Darner (A. palmata) attaches to a female.

http://www.youtube.com/watch?v=p_ap7Aex9Y4

Black Saddlebags (T. lacerata) laying eggs and reattching.

http://www.youtube.com/watch?v=wukDS-1shAU