[Odonata-l] Odonata phylogeny

[Mike May]

The paper by Hasegawa & Kasuya noted by Roy Beckemeyer and discussed by Don 
Roberson is certainly an interesting paper and a step toward understanding 
odonate phylogeny. However, I'd point out that the taxon sampling is quite 
thin and does not include many taxa that are probably important in 
understanding true relationships. The same criticism applies to Saux, et al. 
(2003). My point is, these papers represent progress, but as Don was careful 
to point out, they don't represent THE answer. It's safe to say, of course, 
given the nature of science, that we'll never really have the whole truth. 
With several groups actively working on this problem, though, I expect we'll 
see a good deal of exciting, and sometimes contradictory and controversial, 
new data and analysis within the next few years, followed eventually by a 
new consensus (maybe actually the first consensus) on the phylogeny of 
Odonata. But don't assume just yet that Lestidae are closet dragonflies.

Mike May

----- Original Message ----- 
From: "creagrus" <creagrus at montereybay.com>
To: <odonata-l at listhost.ups.edu>
Sent: Tuesday, October 24, 2006 8:29 PM
Subject: [Odonata-l] Odonata phylogeny


> In today's postings, Roy Beckemeyer helpfully provided a web link to a
> paper by Hasegawa & Kasuya (2006) that discusses odonate phylogeny. As
> it happens, I had read this paper last week, along with others on this
> general topic, and found it interesting to observe the progress underway
> in this area.
>
> My background is in field ornithology and world birding. I have a
> website that features Bird Families of the World
> [http://montereybay.com/creagrus/list.html  ] and I have been an avid
> reader of taxonomic papers, as well as a correspondent with many of the
> professional ornithologists working in that field.  In ornithology, for
> example, a set of new papers signal a major advance in understanding
> bird phylogeny as it relates to the traditional "Old World Warblers," an
> assemblage of ~700 species that now proves to be paraphyletic. Indeed,
> the "Sylvidae" is actually composed of 8-10 separate lineages. I have a
> 3-web page set on the topic of the "Breakup of the Old World Warblers"
> that summarizes much of the new work, with a host of citations,
> beginning at http://montereybay.com/creagrus/sylvid-intro.html
>
> To comment briefly on odonate phylogeny, let me digress slightly to
> offer a very simplified overview of progress in the avian world. For
> many years, work on avian phylogeny was based on studies of morphology,
> internal structures, behavior, voice, parasites, and numerous other
> lines of evidence. In the last couple of decades of the 20th century,
> the field of modern cladistic studies, based on this type of evidence,
> dominated the literature. In the 1980s, Charles Sibley and colleagues
> used an indirect method for recovering molecular evidence [measuring
> 'melting curves' on lab-created DNA-DNA hybridization strands] that led
> to influential papers in 1990 that upended many classical traditions
> about avian evolution (Sibley &Ahlquist 1990, Sibley & Monroe 1990).
> Many of their findings have proved prescient but others have not been
> supported by later studies. In upsetting the apple cart, they got only
> part of it right. By the late 1990s, studies based on mitochondrial DNA
> became the rage, but these, too, had limitations. While very useful to
> test hypotheses of relationships between closely-related taxa, they
> proved less helpful in looking at larger issues at the family or order
> level, with statistical problems like 'long branch attraction' giving
> misleading 'positives' at unexpected places. Recently, however, the use
> of the sequences of nuclear DNA has proven much more reliable. A growing
> consensus is that nuclear DNA may hold the real key to direct evidence
> of avian relationships. Progress has been unusually rapid in the last
> couple of years, and many suspect that within, say, 10 years, all the
> major issues related to phylogeny may be sorted out. My web pages have
> tried to stay abreast of this progress -- some of which is quite
> surprising, yet quite satisfying. The most satisfying of all the studies
> have been those that use a variety of evidence -- multiple nuclear DNA
> genes, mitochondrial studies, cladistic work on morphology, etc -- that
> support the same phylogeny at a high degree of confidence.
>
> I have no background in the Odonata literature, so these comments may
> reflect some ignorance. As I understand it, it has been generally
> thought that there are three major suborders of the order Odonata: the
> Zygoptera [damselflies], the Anisoptera [dragonflies], and the
> Anisozygoptera [two species of Epiophlebia in Japan and the eastern
> Himalayas]. It has generally been thought that the Anisozygoptera is a
> basal lineage, the two species are often referred to as "living
> fossils"; e.g., Silsby (2001). The recent genetic analysis by Hasegawa &
> Kasuya (2006), based on nuclear DNA, was intended to consider the
> placement of the Anisozygoptera. Their evidence placed that suborder
> between the other two suborders, consistent with physical characters
> that are "half-damselfly, half-dragonfly." [Much of the paper discusses
> the comparative value of two specific genes for recovering odonata
> phylogeny, rather than the phylogeny itself.]
>
> What I found particularly interesting in Hasegawa & Kasuya (2006) is
> that a Japanese species of spreadwing (Lestes japonicus) also falls
> intermediate between damselflies and dragonflies, and rather close to
> Epiophlebia. The authors do not discuss this finding in any detail, and
> do not express an opinion as to which suborder the spreadwings might
> belong.  This finding, however, is similar with phylogenetic analysis of
> Saux et al. (2003), who looked at mitochondrial DNA evidence. They found
> that spreadwings [family Lestidae] was more closely related to
> dragonflies than to damselflies. The damselflies [suborder Zygoptera]
> are a paraphyletic group until the Lestidae are removed; at that point
> the molecular evidence finds them to be a monophyletic clade. The
> dragonflies are a monophyletic clade, whether of not Lestidae is
> included within the Anisoptera. Saux et al. (2003) did not analyze
> Epiophlebia, so again we do not know whether the spreadwings are (a)
> part of the dragonflies, (b) part of the Anisozygoptera that is
> otherwise restricted to Asia, or (c) should be placed in their own
> suborder. What they found, however, is that spreadwings are not typical
> damselflies.
>
> In avian phylogeny, nuclear DNA evidence is often better to determine
> family or order level relationships than is mitochondrial DNA evidence.
> Therefore, the results of Hasegawa & Kasuya (2006) are an unexpected and
> quite interesting confirmation of what mitochondrial DNA evidence had
> suggested: that spreadwings are not damselflies. This is contrary to
> cladistic studies of morphology, wing venation, and larvae (e.g., the
> extensive work of Rehn 2003). In the avian world, it appears to me that
> properly constructed nuclear DNA studies have overwhelmed even the best
> cladistic work based solely on
> morphology. Of course, studies that use both lines of evidence produce
> the most satisfying results. Quite by accident, today I came across
> reference to Bybee, Ogden, Rehn & Whiting, in prep., "Phylogeny of
> Odonata (Dragonflies and Damselflies) based on molecular evidence."
> Perhaps they are working on such a broad-scale multi-dimensional approach?
>
> The two published papers using mitochondrial and nuclear DNA would not
> seem to be enough to change the classical phylogeny yet, but perhaps the
> handwriting is on the wall. Perhaps it is time for us to consider the
> real possibility that spreadwings are not damsels, and (possibly) adjust
> our checklists accordingly?
>
> Beyond the Lestidae, there is the question of whether the same results
> would apply to all the lestoids in the world. Other questions are
> apparently not yet settled. Are gomphids or aeshnids more basal? Do
> cordulegastrids belong with libellulids, as the Saux paper concluded?
> [The Hasegawa paper hints otherwise.] Dennis Paulson, in litt., points
> out that all the studies so far are holarctic-centric, while the bulk of
> Odonata is tropical. So there is a lot yet to be learned. Indeed, we may
> be comparatively early in the analysis.
>
> In looking through the archives and in general chit-chat, I haven't seen
> much talk on this stuff. I find it fascinating and await the newest
> developments. Does anyone else? How much of our local checklists should
> we be ready to revise? In the bird world, we are just now getting used
> to putting ducks & geese first, then qrouse & quail, before the
> traditional stuff like loons, grebes, and tubenoses.  In the ode world,
> should we be preparing to think of spreadwings -- and perhaps others -- 
> in a different way?
>
> Literature cited:
> Hasegawa, E., and E. Kasuya. 2006. Phylogenetic analysis of the insect
> order Odonata using 28S and 16S rDNA sequences: a comparison between
> data sets with different evolutionary rates. Entomological Science 9:
> 55-66.
> Rehn, A.C. 2003. Phylogenetic analysis of higher-level relationships of
> Odonata. Systematic Entomology 28: 181-239.
> Saux, C., C. Simon, and G.S. Spicer. 2003. Phylogeny of the dragonfly
> and damselfly Order Odonata as inferred by mitochondria 12S ribosomal
> RNA sequences. Ann. Entomol. Soc. Am. 96: 693-699.
> Silsby, J. 2001. Dragonflies of the World. Smithsonian Instit. Press,
> Washington, D.C.
>
>
> _______________________________________________
> Odonata-l mailing list
> Odonata-l at listhost.ups.edu
> https://mailweb.ups.edu/mailman/listinfo/odonata-l