[Odonata-l] Why report hindwing length and not forewing lengthalso?

[George L. Harp]

With respect to Nick's comment about developing a protocol for weight measurement, I went through this with larval chironomids long ago.  I was more interested in energy content, which compounded the problem, as protein/lipid/carbohydrate content varies in response to many factors.  Considering weight alone, one must consider preservation (acetone, other) or not, degree of dehydration, seasonal changes, among I'm sure other factors.  I finally concluded that getting repeatable and/or comparable weight values was beyond my ability.
 
George Harp 
 
 
This interests me a lot, because I have been measuring Odonata for many years.
 
One problem with wing measurement is that it is not certain where the base of the wing is located.  In my recent Orthemis study I have been taking the notch at the base of the costa that seems to correspond to the major point of flexure of the wing.  This is pretty difficult to observe if one is using a millimeter scale on a specimen held beneath the microscope, but it can be done.  Tillyard more or less shows where it is, but he does not figure it worth a damn.  My technique is to scan the wing and scale together, and then to transpose the scale image on to the wing in Photoshop, rotating and translating the scale to make the measurement.  In this way I can get about 0.1 mm repeatability.  This is great for my own exercises, in which I wish to show comparisons with other specimens measured also by me.  But how do I compare my results with those of others?  The simple answer is that I cannot do this, because I have no idea how others measure wings.  So, how do you do it, and from where?
 
And abdominal measurment - don't even talk about that.  Even when the abdomen is still in one piece, and it is straight, not curved, there is the problem of shrinkage during preservation.  This measurement is vastly less certain than wing measurement, and I use the hind wing as the only more or less reliable surrogate for size.  I still am not quite certain what all this is accomplishing.  I found in the Orthemis study one specimen for which I thought I had made a serious mistake - it couldn't really be shaped like that, with such short wings and a long abdomen.  I pulled the specimen out and looked  at it again- It really was shaped like that - long abdomen and short wings.  No mistake at all.  But what was the meaning of its reported size?
 
What I would like to see done is weight measurements.  I once found a species of Nesobasis (damselfly) for which the males tended to be much larger than the females.  I estimated from dimensions alone that the males must weight at least twice what the females weighed, which I think is remarkable.  But a protocol for weight measurement would be very difficult and would have to involve taking a very portable and accurate scale into the field.  
 
A digression.  Years ago I was in my office when the friendly Fishers sales rep wandered in.  "What do you have that is new and interesting?". I asked.  "Why we have just introduced portable, digital, battery powered scales", he said with considerable pride.  "I'll bet the drug dealers are scarfing them up", I replied.  He promptly turned an interesting shade of gray, never having considered this marketing possibility.

________________________________

From: odonata-l-bounces at listhost.ups.edu [mailto:odonata-l-bounces at listhost.ups.edu] On Behalf Of Ola Fincke
Sent: Thursday, March 15, 2007 4:16 PM
To: Mike May
Cc: Odonata-l
Subject: Re: [Odonata-l] Why report hindwing length and not forewing length also?


In many species, it's well-known that size declines over the season, which is one reason that those of us interested in the effects of size on reproductive success take that into account (e.g. Fincke & Hadrys 2001; see also work by Dave Thompson and others, cited in the former). 

There must be a LOT of geographic variation in size; certainly Ischnura verticalis in MI is much larger than it is in OK. Even within species in similar areas, size can vary quite a across populations. I am collecting data on this within Enallagma. 

Ola Fincke
Dept. of Zoology
University of Oklahoma
Norman, OK 73019
Tel: 405-325-5514
Fax: 405-325-6202




On Mar 15, 2007, at 1:52 PM, Mike May wrote:


	Dennis, et al.,
	
	I have to agree that a much more useful set of measurements, regardless of which body parts are selected, would be mean +/- standard deviation (range). Likewise it would be great to include much more precise distribution data (e.g., county rather than state level - hooray again for Nick Donnelly's dot maps) and seasonal data (e.g., flight season at several different latitudes or within regions of similar climate). The reason for not doing so in NWM and W&M is simple, and the same in all these cases - time and effort. To a fairly accurate approximation, one can (read, "Mike can") riffle through the hundreds of specimens of Enallagma civile at FSCA, pick out several of the largest and smallest from different regions, spread them out and compare by eye, select and measure 3-4 at each end of the size range, and write down the maximum and minimum results. That's exactly what I did, and, although tedious enough, it's a lot quicker than selecting a random sample large enough to be statistically relevant (~30), measuring (with a mm ruler, sometimes viewed through a scope) and recording all of them, and calculating the appropriate statistics. That doesn't make the former technique great, especially if one or two atypically large or small individuals are incuded, it's just easier. There's also no doubt that that human error plays a role and is likely to lead to a more misleading outcome in assessing range than aggregated statistics. As an aside, total length and abdominal length are probably much more subject to error than is wing length because of the tendency of heads to fall off, abdomens to curl, etc. 
	
	Anyway, I think a genuinely worthwhile project would be to accumulate and publish careful size measurements of as many odonate species as possible, based on statistically meaningful and geographically diverse samples. This just wasn't feasible in doing the manuals, but it would be very useful in itself and could lead to some interesting questions. E.g., does size vary geographically or seasonally, and is the variation consistent across species? This has been shown to be true for several species, as Dennis notes, but the answer isn't known for most. If such patterns do exist, then you can ask why. If anyone is interested in undertaking such a project, I'd be glad to give them the benefit of my experience, for what it's worth.
	
	Mike May

		----- Original Message ----- 
		From: Dennis Paulson <mailto:dennispaulson at comcast.net>  
		To: Odonata-l <mailto:odonata-l at listhost.ups.edu>  
		Sent: Wednesday, March 14, 2007 12:17 PM
		Subject: Re: [Odonata-l] Why report hindwing length and not forewing length also?

		What is sometimes frustrating is that guidebooks such as these list measurement extremes without giving any idea of means or explaining the significance of the extremes, and the handbooks Alex mentioned are among the best examples of this.

		For example, I opened Needham, Westfall & May to a page at random and found Erpetogomphus bothrops. Length 38-51 mm, abdomen 26-39, hindwing 21-31. This is a common species in Sonora, which I've visited several times in recent years, and I always thought the individuals I saw and collected didn't vary anything like that. So I just measured the hind wings of all my specimens (from throughout the range of the species) and found this:

		males - range 23-26.5 mm, mean 24.9 (n=15)

		Eleven were with n 1 mm of the mean, in the range 24-26 mm.

		females - range 26.5-29 mm, mean 27.5 (n=6)

		Five were within 1 mm of the mean, in the range 26.5-28.5 mm.

		Thus in both sexes, the great majority of individuals measured within a 2-mm range, and that range defines the size of each sex quite well. Combining the sexes would justify listing the range in "usual" hindwing length of 24-28.5 mm. Total length and abdomen length don't seem to vary any more than hindwing length.

		The entire series of 21 individuals ranged from 23-29 mm. Rosser Garrison, in his 1994 revision of the genus, examined 91 specimens. He listed the variation in hindwing length as 27-31 mm, which makes me wonder if we measure wings differently. In fact we must, because some of my specimens were among those he measured. Both of u show a considerably smaller range than that listed in NWM, but between my measurements and Garrison's measurements, we're approaching the range in NWM, so part of the great variation listed in that book could be explained by different measurement techniques and/or acquiring the measurements from all published data. Unfortunately, ranges like that don't give one a very good idea of how big the dragonfly is. Perhaps Mike May can tell us how those figures were obtained. I emphasize that I'm not criticizing NWM, as I'm sure the authors of that book did the best they could in acquiring these figures.

		I do know, from much research on bird size, that guide book writers sometimes get it wrong, and the incorrect figures have a life of their own, copied from book to book. I found total length measurements off by as much as 15% when analyzing measurements in several bird guides, at times a larger species listed as smaller than a smaller species. How many bird-b ok writers nowadays measure birds, live or dead?

		In many taxonomic papers, the author measures a series of specimens and gives ranges and means, but for many species there are no such papers, so writers of guide books have a difficult time getting measurements, with two options. Either they have access to a collection and spend long hours measuring specimens, or they take the figures from other publications, the research literature and/or other guide books. One of my colleagues recently confessed to me that when he listed measurements for a book on dragonflies, he just used the midpoint of the measurements in NWM and Westfall & May, which seems as good a way of doing it as any. I think having a single measurement in a guide book makes it easier to picture the size of the animal than when there is a rather substantial range.

		Some species vary in size over the season. Pachydiplax longipennis t at emerge in summer in southern Florida have hind wings about 3 mm shorter than those that emerge in the winter, and the size varies seasonally at least as far north as the US Gulf states. I am not aware if seasonal variation has been established for other species. Just from eyeballing it, it seems to me that some species vary more in size than others, but perhaps that just depends on how many of them you see. I was just examining two Sympetrum costiferum, one of which seemed no more than 2/3 the bulk of the other, and two Enallagma carunculatum, one of which was surely only half the bulk of the other and about 2/3 its length. These are unusual extremes, of course. When I look at my collection or at odonates in the field, individuals of a species mostly look around the same size.

		Another source of variation in odonates is temperature. Damselflies in Mexico and Central America tend to be larger at higher elevations, presumably something about temperatu e vis-a-vis development time of the larvae. I don't know if there is latitudinal variation as well, but there should be. Nick Donnelly called attention to the very small size of some damselflies in eastern Texas, and I have seen these same populations of impressively tiny creatures. I don't know whether these populations are genetically distinct, or does some environmental factor of the area have them emerging at a smaller size? I collected tiny Perithemis mooma in Yucatan, as small as or smaller than Nannothemis bella.

		There are probably many more examples of interesting size variation in odonates, and it would be good to know more about this.

		Dennis Paulson

		On Mar 14, 2007, at 6:58 AM, Mike May wrote:


			Certainly part of the reason is tradition. Once a large body of data is 
			available for a particular measurement, the value of that measurement for 
			comparative study is enhanced. I suspect, also, that the hindwing may have 
			been selected originally because, when a dragonfly is held in the hand with 
			the wings pressed together above the back, the hindwing is positioned 
			outside the forewing. Although the forces produced by fore- and hindwings in 
			flight are certainly somewhat different, I feel pretty confident that had 
			nothing to do with the decision.

			The forewings are nearly always slightly longer than the hindwings. My 
			impression is that the proportionate difference does not vary greatly with 
			size, taxon or sex, but I don't know whether this has been investigated 
			rigorously. Females do typically have longer wings than males in comparison 
			to other measures of body size such as total length or thoracic mass 
			(probably because abdominal mass becomes greater in mature females because 
			of the mass of eggs), but I don't know that the differences between fore- < SPAN>
			and hindwings is affected by sex. Maybe Roy Beckemeyer can illuminate that 
			question a bit more.

			Mike May

			----- Original Message ----- 
			From: <aardila at uoguelph.ca>
			To: "Odonata-l" <odonata-l at listhost.ups.edu>
			Sent: Wednesday, March 14, 2007 4:28 AM
			Subject: [Odonata-l] Why report hindwing length and not forewing length 
			also?



				Hello everyone,

				I have the books "Dragonflies of North America"(Needham, Westfall ,and
				May, 2000) and "Damselflies of North America" (Westfall and May,
				1996). These books are excellent. They report the body size lenght,
				abdomen lenght, and hindwing length for every Odonata species in North
				America.

				However, I don't understand why the size of the forewings is not
				reported. Is this by convention? What is the history behind this? Why
				were the hindwings and not the forewings chosen? Is there a
				statistically and biologically insignificant difference between the
				forewings and the hindwings within species (and sex) in Odonates? Is
				this selection based on the role of the hindwings in flight? I would
				appreciate any guidance that could help me find an answer to these
				questions. Thank you.

				Alex

				-- 
				Alex Ardila-Garcia


		
		-----
		Dennis Paulson
		1724 NE 98 St.
		Seattle, WA 98115
		206-528-1382
		dennispaulson at comcast.net




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