Saturday, December 24, 2005

Transitional Species in Insect Evolution

In "On the Origin of Species" Charles Darwin listed as one of the gravest objections to his theory the apparent paucity of intermediate or transitional forms, which he attributed (at least in part) to the imperfection of the fossil record. Well over a century later this is still being raised as an objection to evolutionary theory but such objections are increasingly misinformed as a growing number of transitional species, both living and fossil, have been found. Transitional species in the evolution of various vertebrate groups (several of which have a surprisingly good fossil record) have been discussed on the web. Recent molecular studies, combined with some truly exciting fossil discoveries, have illuminated the origins of several major kinds of animals from previously existing groups: tetrapods (4-legged land-dwelling vertebrates) from fish, birds from dinosaurs, and whales from artiodactyls (land-dwelling hoofed mammals with 4 legs) (for more information see the Transitional Vertebrate Fossils FAQ and 29+ Evidences for Macroevolution at Talk Origins). These new discoveries have largely supported hypotheses of evolutionary relationships that were previously based on morphological and other evidence, but provide a much more vivid picture of the evolution of these groups that had previously been largely hypothetical, in many cases providing concrete and compelling fossils of intermediate or transitional species between the major groups.

Similar transitions can be found in invertebrate groups such as insects. Although the origin of insects, the largest group of animals on earth, is still unclear (current evidence suggests they evolved directly from crustaceans) they have left a good fossil record during their subsequent evolution. The evolution of insects is discussed in a recent book titled, appropriately enough, Evolution of the Insects by entomologists David Grimaldi and Michael Engel. If you have any interest in insects and their evolution, this is the book for you. This is a very big and very heavy book, but it is also extremely well-illustrated with both fossils and living insects. The authors present the current state of knowledge of insect evolution as known from morphological, molecular, and fossil evidence. Grimaldi's specialty is fossil insects, particularly those preserved in amber, and the book is abundantly illustrated with diagrams and photographs of fossils, of extinct insects, and of living insects. Detailed phylogenies of all the major groups are presented, with detailed discussion of the evidence upon which those phylogenies are based. The writing is clear and not excessively technical; it should be accessible to anybody with some knowledge of basic biology and/or entomology. One of the most fascinating things I discovered in this book is that current data strongly indicate that several of the distinct, well-defined, and traditionally recognized insect orders have evolved directly from within other existing orders. Examples are Siphonaptera (fleas) from Mecoptera (scorpionflies), true lice (Phthiraptera) from the order containing barklice and booklice (Psocoptera) (with chewing lice providing a transition to sucking lice), and termites (Isoptera) from cockroaches (Blattaria). It is this last transition that I will discuss here.

The differences between cockroaches and termites are dramatic. Cockroaches are more or less omnivorous detritivores and scavengers (as witnessed by the success of several species as household pests) whereas termites eat wood, digesting its cellulose with the aid of symbiotic bacteria and protists. Cockroaches are generally solitary; termites form huge colonies of related individuals founded by a long-lived "queen" and "king". Cockroaches vary little within a species (aside from age and sexual differences); termites have a marked differentiation into winged reproductive and unwinged non-reproductive "castes" (e.g., workers and soldiers). Cockroaches have 2 pairs of wings that are different and are retained through their adult life; termites have 2 pairs of wings that are similar ("isopterous") that are shed after mating. Cockroaches and termites are as well-defined and distinct as any two orders of insects; in colloquial terms they can be considered entirely different kinds of insects. Yet there is now compelling evidence that one of these "kinds"--termites--has evolved directly from the other--cockroaches. This conclusion is both surprising and unsurprising: unsurprising in that termites have always been acknowledged on the basis of internal and external morphology to have some relationship to cockroaches and other orthopteroid orders. What is surprising, beyond the obvious differences between termites and cockroaches, is that termites have evolved from a group within the order Blattaria, and in fact are more closely related to one particular living genus of cockroaches than that genus is to all other cockroaches. In other words, termites ARE cockroaches, albeit highly modified due to their specialized lifestyle that differs from that of their ancestors.

If termites have indeed evolved from cockroaches, then we should expect to find transitional living or fossil termite-like cockroaches and cockroach-like termites. In addition, other lines of evidence should be consistent with this relationships. For example, the derived taxa (termites) should not predate their ancestors (cockroaches) in the fossil record. Moreover, the earliest members of the derived group--that is, the ones closest to the evolutionary divergence--should be the most similar to the ancestral group. Finally, the phylogenetic relationships of their symbiotic bacteria and protists, which are utterly dependent upon their hosts for survival, should be similar to the phylogenetic relationships of their hosts.

Cryptocercus, a termite-like cockroach

While termites have generally been accepted as being related to other "orthopteroid" orders, different hypotheses had argued for their closest relationship being with either cockroaches or mantids. However, on the basis of morphology and other evidence some authors have suggested a particularly close relationship between one termites and a living genus of cockroaches, Cryptercercus. While typical cockroaches in many ways, the nine species of Cryptocercus share numerous characteristics with termites.

Termite-like characteristics of Cryptocercus:

* exhibit social behavior
* live and nest exclusively in dead wood
* eat and digest wood with the aid of endosymbiotic bacteria and protists
* harbor several genera of symbiotic protists found in termites but not other cockroaches
* live long and pair monogamously (most cockroaches short-lived, mate promiscuously)
* provide parental care for nymphs
* nymphs fed by anal secretions of the adults
* symbiotic protists and bacteria passed to nymphs in anal secretions
* nymphs strikingly termite-like in appearance




Cryptocercus punctulatus adult with nymphs (from Origins of Termites)


Mastotermes, a cockroach-like termite

Are there also termites that retain cockroach-like characteristics? Yes. The Australian "giant termite" Mastotermes darwiniensis, which has long been considered one of the most primitive termites, shares several characteristics with cockroaches that are not found in other termites.

Cockroach-like characteristics of Mastotermes:

* large size (other termites are much smaller)
* expanded pronotum ("shield" on the first thoracic segment, reduced in other termites)
* hind wings with an expanded anal fan (absent in other termites)
* ovipositor present (absent in all other termites)
* eggs laid in oothecae (clusters in a membrane-enclosed capsule; other termites lay solitary eggs)
* feet with 5 tarsal segments (all other termites have 4)
* harbor endosymbiotic bacterium Blattabacterium, a genus found in cockroaches but not other termites

Image hosted by Photobucket.com

Mastotermes darwiniensis, a cockroach-like termite (from Evolution of Insects)



Mastotermes darwiniensis, female reproductive after shedding wings (from Tree of Life)



Mastotermes darwiniensis nymphs, workers and soldier (from Termites as Structural Pests)

Evolution of termites from cockroaches

The molecular, morphological, behavioral and endosymbiont data show that the evolution of termites from cockroaches, while seemingly rapid in geological terms, has gone through several stages, with "typical" termite characteristics being acquired stepwise:

Stage 1: primitive roaches with long ovipositors (earliest roach-like fossils);
Stage 2: ovipositor shortened; typical cockroaches (plenty of those still around);
Stage 3: wood-dwelling cockroaches that mate monogamously, care for their nymphs, and can eat and digest wood. (Represented by the living genus Crytocercus);
Stage 4: development of a true social lifestyle with a caste system consisting of a reproductive "queen" and "king" and sexually suppressed nymphs differentiated into non-reproductive workers and soldiers; acquisition of some "typical" termite characteristics (e.g., forewings membranous and wings shed after mating). (Represented by the living genus Mastotermes);
Stage 5: acquisition of more typical termite characteristics (e.g., ovipositor lost, eggs laid singly, hing wings lose their anal fan, all 4 wings "isopterous"); represented by all other living termites.

Note that the genus Cryptocercus can NOT be considered directly ancestral to termites, as it has specializations of its own (e.g., loss of wings). This is to be expected as the lineage leading to Cryptocercus has had a long evolutionary history of its own since it diverged from the common ancestor with termites, and continued to evolve independently after that divergence. However, the termite-like characteristics of Cryptocercus and cockroach-like characteristics of Mastotermes demonstrate that evolution of termites from cockroaches has been relatively gradual, has occurred stepwise by a progressive loss of cockroach characteristics and accumulation of termite characteristics, and has left evidence of this evolution in the surviving descendants of at least two transitional species that occurred along the way.

Fossil record

Grimaldi & Engel place the first appearance of true cockroaches in the fossil record as early Cretaceous, and termites at about the same time. Although appearing at about the same time in the fossil record, the Cretaceous termite fossils belong to the primitive (although still extant) families Mastotermitidae, Hodotermitidae, and Termopsidae. However, the earliest cockroach fossils are already diverse and represent several modern families, suggesting a much earlier origin and evolutionary divergence. This earlier origin is also suggested by "roachoid" fossils that occur from the Carboniferous into the Jurassic. Although it is often claimed that cockroaches first appeared in the Paleozoic era in more or less modern form and have changed little since, these early "roaches", dubbed "roachoids" by Grimaldi, while similar to cockroaches general appearance and sharing many characters with them, differ in having elongate ovipositors more like those found in the closely related order Orthoptera, rather than the reduced ovipositors of cockroaches (and Mastotermes)--in other words, they comprise an entirely different set of transitional species between primitive orthopterans and true cockroaches. Some of these "roachoid" fossils are so roachlike that the difference is little more than semantic, and there can be little doubt that modern cockroaches evolved directly from one of the "roachoid" lineages sometime during the Jurassic. The discovery of trace fossils interpreted as termite nests in the early Jurassic also suggests that the evolution of both true cockroaches and termites may have occurred considerably earlier than the Cretaceous although this is consistent with both the appearance of diverse cockroach and termite body fossils in the early Cretaceous, as well as the occurrence of "roachoid" fossils into the Jurassic.



A Carboniferous "roachoid" (from LARGEST FOSSIL COCKROACH FOUND)


Molecular evidence

Molecular phylogenies have supported morphological and other evidence (e.g., social behavior) in showing that Cryptocercus is more closely related to termites than it is to other cockroaches, and that Mastotermes is primitive with respect to all other termites. An interesting test of the phylogenetic relationships between cockroaches and termites is the phylogenetic relationships of their symbionts. As stated above, there is a strongly suggestive overlap of symbiotic protists and bacteria between cockroaches, Cryptocercus, Mastotermes, and other termites. These bacteria and protists are incapable of living apart from their hosts, and are passed from one generation of hosts to the next via anal secretions. Their utter dependence upon their hosts means that they have evolved in parallel with their hosts, and indeed the molecular phylogenies of the symbionts have proven to almost perfectly parallel the host phylogeny: the symbionts of Mastotermes and other termites are most closely related, are next most closely related to the symbionts of Cryptocercus, and are next most closely related to those of other cockroaches.

Summary

1. Cryptocercus is a cockroach with social characteristics of termites.
2. Cryptocercus shares endosymbionts with termites that are not shared with other cockroaches.
3. Molecular evidence shows that Cryptocercus is more closely related to termites than it is to other cockroaches.
4. Molecular evidence shows that the endosymbionts of Cryptocercus are more closely related to those of termites than to those of other cockroaches.
5. Mastotermes is a termite with numerous morphological characteristics of cockroaches that are not shared with other termites.
6. Mastotermes shares certain endosymbionts with cockroaches that are not found in other termites.
6. Molecular evidence shows that Mastotermes is primitive with respect to other termites.
7. The oldest termite fossils belong to Mastotermitidae and other primitive families.

Creationists might make the typical claim that "Cryptocercus is still just a cockroach, and Mastotermes is just a termite" but this would be a gross oversimplification that conveniently ignores the evidence. It does not address why Cryptocercus has so many termite-like characteristics, and Mastotermes has so many cockroach-like characteristics, when the molecular evidence indicates that Cryptocercus is the closest living relative of termites, that Mastotermes is the most primitive living termite, and even the relationships of their endosymbionts support this relationship. In other words, the data from several different kinds of evidence point toward the conclusion that termites evolved directly from cockroaches. Taken individually these data could be argued or interpreted in different ways; taken together they point to the unavoidable conclusion that, as Grimaldi says in a review of insect evolution, "Termites are highly modified, social, myopic, wood-eating roaches."

References available on the web (in no particular order):

Colony composition, social behavior and some ecological characteristics of the Korean wood-feeding cockroach (Cryptocercus kyebangensis)

Origins of Termites

Termites and trees: a review of recent advances in termite phylogenetics

Evidence from multiple gene sequences indicates that termites evolved from wood-feeding cockroaches

Phylogeny of Mantodea based on molecular data: evolution of a charismatic predator (supports paraphyly of roaches with respect to termites)

Mantophasmatodea and phylogeny of the lower neopterous insects

Insect evolutionary history from Handlirsch to Hennig, and beyond

Evidence for Cocladogenesis Between Diverse Dictyopteran Lineages and Their Intracellular Endosymbionts

Advanced early Jurassic termite (Insecta: Isoptera) nests: evidence from the Clarens Formation in the Tuli Basin, southern Africa.

376 Comments:

Blogger Jay said...

It would be a shame if no one congratulated you on the nice post. I found it fascinating, and a little more interesting that the traditional Christmas eve reading of "The Night Before Christmas".

12/25/2005 12:25 PM  
Blogger Jay said...

Manual trackback... I mentioned your excellent post on my site:

http://www.ocellated.com/2005/12/25/which-would-you-rather-read/

12/25/2005 12:56 PM  
Blogger Chief said...

I had no idea. Excellant post, altho I must admit it was mostly over my head. Termites from roaches - wow!!

12/25/2005 3:45 PM  
Anonymous Dior said...

As a high school bio teacher, the more transitional examples laid out neat and understandable to high school students (and some teachers) will ultimatly make our job easier. Thank you for the post and I challenge all other science bloggers to continue. With the lack of money to take kids to the Nat. Hist.Museums, I need all the help I can get; and it will help us all. TY again,

12/25/2005 10:20 PM  
Blogger MrDarwin said...

One of these days I'll figure out how to do trackbacks. In the meantime, this article has been featured on The Panda's Thumb at http://www.pandasthumb.org/archives/2005/12/transitional_sp.html

12/27/2005 4:28 PM  
Anonymous Steviepinhead said...

This was a great post, which I also found via Panda's Thumb.
I had had Evolution of the Insects in my amazon.com "wish list" for some time, prompted in part by a plug from PZ Myers, but you're article--and a gift certificate from a generous sister!--finally pushed me over the edge, and I ordered it!
A couple of non-substantive critiques: I found the white lettering on black background a bit of an eyestrain. Some of your paragraphs needed subdivision into shorter units.
Otherwise, great, and I read every word with relish!

12/28/2005 6:57 PM  
Anonymous Anonymous said...

Just one dumb question if I may:

Why aren't there tons of living transitional forms? That is, if humans came from apes, or whatever, why don't we see, now, living, every version in between? If the thing just "above" ape was "better", why can't it still be alive? and why IS the ape still alive? I would think we'd see literally hundreds of living intermediates; and the same with cats, dogs, elephants (everything from the mamoth down) and every other animal on earth. Why so much apparent "stasis", if that's the right term.

I tried to find a link that addressed this question first, but with no success.

Thanks for your reply.
es

1/18/2006 1:01 PM  
Anonymous Gregor Samsa Mendel said...

I'll hazard a response to es's post...

The vast majority of species that have lived on the Earth are now extinct. Not just "transitional forms" (although they didn't know that they were in transit to anywhere) but "finished products," like trilobites and dinosaurs. Those "living intermediates" you think should be here aren't around because they died out long ago, erased like footprints in the sifting sands of time. Some of these intermediates were kind enough to leave fossils of themselves, though.

Just because something has evolved from something else doesn't make the ancestral form "inferior" or the derived form "superior," in some sort of absolute sense. Just better adapted to the local conditions. For example, people are generally considered more "advanced" than bacteria, but hurricane Katrina changed the environment in New Orleans in way so that people suffered but many bacteria flourished. Although living things are often considered to be "primitive" or "advanced," as far as I'm concerned, if it's around now, it's advanced enough.

Back to termites...I happen to be interested in them at the moment. Even with an obvious transitional form like Mastotermes, things are never as straightforward as they seem. In terms of social structure, Mastotermes is more advanced than members of the family Termopsidae, which form smaller colonies and have a less "rigid" delineation of castes. (For example, Termopsids occasionally form fertile soldiers, whereas the soldiers of all other termites are sterile.) The current idea is that Mastotermes retained much of its ancestral morphology but went on to evolve a more complex social system than that of the original ancestral termites. Things are always messier than they ought to be...

3/15/2006 11:29 PM  
Anonymous Anonymous said...

i dont really believe in evolution....i think charles darwin just made things up..there's no such thing as evolution....humans didnt come from apes because there is tremendous or significant difference between us and apes......
there's no way that apes could evolve that dramatically from banana eating animals to sophisticated humans with a capacity to advance...

5/09/2006 5:16 AM  
Anonymous Anonymous said...

As a creationist - I am unimpressed - are the termites that live in large - constant temporture regulated - 6 foot nests from cockroaches too and did they have similiar dwellings?

I read this thing and said to myself - what a stretch!

6/27/2006 3:45 PM  
Anonymous Alaysian said...

having no real side to this arguement, all i can say to some of you creationists:

Do some reading

Plain and simple. The Theory of Evolution by Natural Selection doesn't say humans evolved from apes, only that they had a common ancester somewhere back along the line.

And as far as transitional species goes, while not very read up on the subject, i would think that there really is no such thing. Since evolution is constant, the only reason that "transitional species" exist in that catagory is that it makes sorting species and their evolutionary path easier. If on ever had all the data on a species evolution, most of the "transitional species" would be shown to be purely subjective, barring major mutations (such as changes in the methods of bearing offspring)

7/02/2006 12:44 PM  
Anonymous quixote8 said...

An excellent post: comprehensive roundup of cogent factors in a clear and logical presentation.
Creationists seem to have several personality deficiencies among which are:-
1) A biggoted predisposition to handed down *authority* of a 2,000 year old 'world view' (rather like the Taliban's 800 year old world view)
2) A confrontational denial of their own (and others) reasoning capacities.
3) A lack of courage to face the uncertaities of knowledge and its aquisition ie. a pathogenic need for absolutes and certainty in a threateningly uncertain world.

Whilst having a sympathy for their timidity in the face of change, I dispise and abhore their raucous agressiveness of assertions of unreasond righteous certainty.

Regards to all
harry

7/25/2006 1:00 AM  
Anonymous Doug said...

Hello!

I found your insightful post while searching for information about "transitional species." I had been told that no examples had been discovered; I find that there are many examples presented on the internet.

One statement in your post does need clarification. Fairly early on it says, "One of the most fascinating things I discovered in this book is that current data strongly indicate that several of the distinct, well-defined, and traditionally recognized insect orders have evolved directly from within other existing orders." Evolution within Orders is one thing, transitional Species is another. Does the book in question deal with species, or orders?

2/07/2007 11:15 AM  
Anonymous Anonymous said...

Great post! Actually, I had a few questions about some of the species you discussed and was wondering what would be the best way to get in contact with you? Thanks!

Best,
Bobby
bmpham@gmail.com

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小吃餐車加盟連鎖「一條龍的創業模式」複製成功開店

經驗
更新日期:2009/08/25 17:19

(中央社訊息服務20090825 16:19:17)大環境不景氣,工作難找,有不少人就想乾脆自己創業當老闆。為

強化國內創業能量,五路財神開店總部於8月20

日-9月20日舉辦「2009夢幻小餐車創業成果

展」歡迎蒞臨參觀。針對國內一窩蜂創業加盟潮,餐車

達人陳滌五總監提醒創業者,加盟不失為一條創業捷徑

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近年來以成本低的小吃餐車最受創業者青睞。沒經驗的失業族 大部份都選擇餐車擺攤,最多人都想在創

業市場闖出一片天。在台中成立的五路財神開店總部,小吃餐車用品牌方式來營運,總經理陳滌五用自己

的經驗來現身說法。以國人創業首選的小吃餐車來說,如何在同質性高的市場中殺出重圍?雖然是路邊攤

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道。
陳總分享成功關鍵,「五路財神餐車與其他路邊攤最大的差異化,在於餐車外觀亮眼與眾不同的加分效果

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品,用創新工法為策略,顛覆傳統路邊攤的概念,長期下來在台灣建立起最多元品牌的餐車王國。
很多創業者最憂心沒經驗,業界首創從餐車打造、生財器具、原物料配方、技術教學、促銷廣告、開幕指

導,「一條龍的創業輔導」模式,使得五路財神餐車品牌特色更為彰顯。陳總說,不景氣下小吃餐車經營是加倍艱辛,建議創業者不但要瞭解

小吃經營的核心價值,凸顯產品獨特性及差異化之外還要定期和你商圈的客人互動,「好餐車、好商品、

好客人」,三個好,如此才有可能成為「不敗的小吃攤」。
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