Tuesday, May 14, 2013

The hidden world of the crouching tiger-beetles

An ultrasound detector
Tiger beetles are an iconic group for many entomologists and ecologists. They have been considered to be a good surrogate group for use in biodiversity studies. They are perhaps not as good as the more ecologically dominant ants, but are certainly full of character and action. To add to that, most of them can be found on open paths, saving the need to dash through vegetation and load oneself with ticks. Unfortunately, nobody has quite made an illustrated fieldguide for the Indian region despite the ease of making one especially for someone with access to a good collection. Perhaps the museums have not been welcoming to anyone who might be motivated. A major reference for the Indian region is by R. Acciavatti and David Pearson (Acciavatti, R.E.; Pearson, D.L. 1989: The tiger beetle genus Cicindela (Coleoptera, Insecta) from the Indian subcontinent. Annals of Carnegie Museum, 58). 


C. venus up front
A somewhat less studied aspect of tiger beetles is the production and hearing of ultrasound. Some years ago I went out on a bat-watching outing with the Norwegian Zoological Society to a lake to the east of Oslo. Equipped with bat-detectors, we were able to hear more bats than we could see. It is simply fascinating when a world that our senses do not detect is suddenly made perceptible. I think this is the fundamental allure of photography (showing detail we may not spot), microscopes, binoculars, ultrasound detectors, DNA sequencers and a most other instrumentation. Anyway, I would love to hold a sensitive bat-detector up close to these critters.


Cicindelids apparently hear ultrasound using two "ears" on the dorsal side of the first abdominal segment. They respond to bat pulses with escape behaviour and also produce ultrasound at around 30–35 kHz in non-defensive contexts, perhaps for communication. According to one study the production of ultrasound in flight is possibly through a mechanism that involves the elytra and wings. I however wonder if they may be producing other sounds through mandibular stridulation, when not flying. They often seem to gnash their jaws, mandibles crossing over each other and perhaps the micro-structure of the flat mandibular surface will have something to reveal. Someday hopefully we can hear these beetles, who knows they might even be detected and identified by their sounds.

A regular on forest paths in the Western Ghats
Cicindela (Cosmodela) duponti Dejean, 1826


Cicindela (Jansenia) venus Horn, 1907
another forest species found on leaf-litter
Cicindela (Calochroa) hamiltoniana Thomson, 1857
most times seen on shaded paths close to water
Cicindela aurofasciata Dejean, 1831
a stouter species found on open grassy habitats
Cicindela (Calochroa) flavomaculataHope, 1831
One of the few species more often attracted to light in the night

C. near parvimaculata, Bandipur (April-2013)


To be determined. Mudigere, 11- May 2013
To be determined. Seen in large numbers beside stream.

A feature that makes tiger beetles particularly interesting for those interested in electronic field guides is their patterning. It would appear as if there was a fairly simple archetypal "bodyplan" (what the Germans called Bauplan, but often avoided because of an implicit idea of "design"). An archetypal plan is however possible without any fear of "intelligent design" if one considers that patterns are coded by sections of genetic code, modules of which are shuffled about during the evolution of species. An amazingly old (1917) work on this is by Shelford and he identifies patterns of colour and provides some interesting insights into developmental biology. By subjecting larvae and pupae to specific environmental conditions of temperature and humidity he was able to alter the patterns that form on the elytra of the adults.


"Showing the geographic distribution of types and patterns. The first series at the left are world-wide in distribution, being most generalized in Eurasia and North America. The second group of patterns belong to several groups of species but all are characterized by the presence of three spots at the base and along the elytral suture. They are most numerous in Africa and India. The next group shows the relatively rare type with the pattern oblique but in the opposite direction from the slope of the tip of the elytron. The last type is one showing peculiar joinings of markings characteristic of species found chiefly in • Indo-Australian region." (Shelford, 1917)

Inspired by Shelford, I attempted to look at the patterns on some Indian species. It is however very impossible to see much without access to museums or other collections.

Patterns on a handful of Indian species - easily drawn using Inkscape


Acknowledgement

Most of the photos were taken on field trips either accompanied or made possible by Dr. Revanna S. Revannavar, entomologist at the University of Horticultural Sciences, Mudigere.
.
References

Postscript
Would be terrific if someone can come up with a low-cost ultrasound monitoring instrumentation in India - based on ideas such as this. Would be very happy to help in field testing for any electronic whizzes out there that might be interested.

Monday, May 6, 2013

Bird fossils from the Indian Subcontinent

I normally desist from the archaic practice of listing things within meaningless political delineations but a need to locate information on bird (and near-bird) fossils and ichnotaxa from the Indian region led me to the discovery that there are no readily available reviews. This is a rather rough working list of taxa and sources. Additions are welcome.
  • Struthio asiaticus A. Milne-Edwards [Siwaliks] (bones, egg-shells from a wider range)
  • Dromaeus sivalensis Lydekker [Siwaliks]
  • Leptoptilos falconeri (=Argala falconeri) A. Milne-Edwards [Siwaliks]
  • Megascelornis sivalensis Lydekker [Siwaliks]
  • Pelicanus cautleyi Davies [Siwaliks]
  • Vastanavis eocaena Mayr et al., 2007 [Vastan, Gujarat]
  • V. cambayensis Mayr et al., 2010 [Vastan, Gujarat]
  • Strigidae [Narmada] Patnaik, R & A. Sahni (1994)
  • Egg clutch [Lameta, MH] Mohabey D.M.; S.G. Udhoji; K.K. Verma (1993)
Postscript: 8-May-2013
In going through an article by Badam (2005) I came across reproductions of a couple of rock drawings from Kathotia and Firengi south of Bhopal. These have apparently been attributed with some doubt to ostriches by paleographers. The illustration from Firengi is pretty poor for a match with an ostrich and on the other hand is a very good match to a Lesser Florican particularly in the fluffy neck and the two plumes exaggerated on the back. I agree with the authors about the inaccuracy in the detail of feet.

A 30,000 year old rock illustration reproduced from Badam (2005).
The bird on the right is a good match for a Lesser Florican.
An old illustration of Lesser Florican (Sypheotides indicus)

Sources

Sunday, April 28, 2013

Be like a bee

I have written in the past about ant-mimicry (Why look like an ant?), and last week I was reminded how good their relatives, the bees and wasps, are as models for Batesian mimicry. It was just by chance that I noticed this yellow spot on a rock beside a stream running through the forest. 
It was evident that this was a moth, a hawk-moth for sure, confirmed by looking at the hooked tip to the antennae. The remarkable similarity in pattern and size to carpenter bees of the genus Xylocopa is readily evident. As kids we used to play a trick with Xylocopa males that would perch atop wooden posts. Toss a pebble in the direction of the male and it would set off and pursue the pebble. Handling a dead carpenter bee was a painfully lesson on how spiny and hard they are. Females can sting but they rarely need to use it. There is probably no predator of adult Xylocopa bees. Little wonder then that several species in the day-flying hawk-moth genus Sataspes have been selected to appear like carpenter bees (some other day-fliers like Cephonodes also look like clear-winged bees). The genus Sataspes does not seem to be well known in southern India and this observation may be among the few reports from southern India.

Species in the genus appear to show considerable variation within populations in the distribution of blue and yellow on their body and wings. Several of these variants have been described in the past as species and the exact determination of species may be reliably established only by dissecting out their genitalia (which show species-specific lock-and-key patterns). In 1900, a year before he died of malaria, Lionel de Niceville described a species from Burma:
S. tagalica (=S. hauxwelli) and model
"I have named this handsome moth (which is unique) after Mr. T. A. Hauxwell, Deputy Conservator of Forests, who is an enthusiastic collector of birds and Lepidoptera. It is a beautiful mimic of the very common large blue carpenter bee, Xylocopa auripennis, Lepeletier, a male of which I have figured-for the first time-for comparison ..." JBNHS 13:173
Some people find the evolution of mimetic forms too implausible to occur by random processes and those that question evolution pose the question as to how the "exact" same pattern could arise. Interestingly enough there is considerable variation within the mimics and it is actually not an exact imitation. Here are a couple of Xylocopas (unidentified) from southern India just for a sampling.

A Xylocopa from the W. Ghats
A Xylocopa from Bangalore

Looking at the variations in Sataspes it is clear that a series of characters are being mixed in various forms and species.Yellow bands, thorax, tufts, bluish wing shine and so on.
Sataspes xylocoparis described by A G Butler in 1875
Writing about Xylocopa aureipennis in 1922, Cedric Dover notes incredulously:
In addition to the localities noticed by Bingham, the Indian Museum also possesses specimens from the Parjiling District, the Naga Hills and Sibsagar in Assam, South India and Nepal. The species is supposed to be mimicked by a Sphingid moth (Sataspes hauxwelli), which according to De Niceville (Journ. Bomb. Nat. Hist. Soc. XIII, p. 174) was "a beautiful mimic of the very common large blue carpenter-bee Xylocopa auripennis, Lepeletier." The wings of the moth are a deep indigo-blue with bronze markings, which scarcely resembles the wings of the bee, and in the cabinet the whole insect seems entirely different.
De Niceville does not say that the bee and the moth were taken together, and in the absence of definite field-observations, the moth has little claims to being a mimic of the Xylocopa.
My own impression was that the moth above was the right size, shape and colour to give the impression of a carpenter bee.

Although carpenter bees are free from predators, they are not free from parasites and their immature stages probably have predators. A peculiar feature of carpenter bee females is a cavity on the first abdominal segment, known as the acarinarium, which is home to mites of the genus Dinogamassus. Nobody quite knows what the role of these symbiotic(?) mites are. Dr B. Mallik, an acarologist (and one of my teachers) at the entomology department in the University of Agricultural Sciences (Bangalore) showed me some of these mites under a microscope a few years ago pointing out that they stack up in a toroidal pattern.
The fact that so few records of these amazing day-flying moths exist is either due to their rarity or the fact that we have so few people keeping an eye on life around them.

References
Postscript
1-May-2013: Ian J. Kitching of the Natural History Museum notes:
... all the evidence now points towards it being the first representative of infernalis from SW India. All other specimens I have seen from the Western Ghats are the relatively non-metallic forms as shown on SEP. None have the strong blue sheen on the wings of your specimen. ..., the moth does look a bit more like S. tagalica but this usually has a greenish metallic sheen and the females (which your moth is) never have a yellow thorax, only the males. That said, there is a definitive feature that could settle the matter BUT it is on the underside of the abdomen. This is a long shot but do you have a photo in which the colour of the underside of the terminal segments of the abdomen is visible, even slightly. These are yellow in infernalis/xylocoparis and black in tagalica.

All that said, no Sataspes in our collection of any species has the subdorsal rows of yellow spots; they either have more extensive yellow bands (see xylocoparis on SEP) or broad central patches (see tagalica on SEP). So I am minded to think that this may be a new species, or at least a distinctive subspecies of infernalis.

Friday, March 22, 2013

Wild laws

An advertisement in the Times of India Delhi (20-March-2013) with a photograph that violates copyright apart from being of the wrong species (showing a Tree Sparrow instead of a House Sparrow)
A newspaper carried a government advertisement last week. A politician declared the house sparrow as the bird symbol for a state but the picture chosen was just picked off the Internet, was of the wrong species and was a case of unethical reuse. A simple image search via Google allowed me to find the person who took that photograph, Dani Studler, and he confirmed that he was never asked for permission to use that image. This is a violation of the law - the Indian Copyright Act 1957 clearly should be something that the government should have known about. Governments however are not known for being law-abiding.

A training in science and biology does not help in making one a fan of the legal system or indeed of traditional governance and authority. One instead seeks to derive one premise from another. One can consider a principle of fairness. Determining fairness in most cases involves examining if you would approve of an action if it was done by someone else. The actions of legal institutions however is not morality based but driven by other forces - those of economics.

Copyright laws, it turns out grew out of the lobbying first in support of printers. Printing presses came to England in 1476. In later times the booksellers lobbied to change this. There was the Statute of Anne (Wikipedia has an excellent entry on the history of copyright law) in 1710. There were also other controls on the printing press, mostly in the form of censorship. Prior to the Statute of Anne, the printers had a perpetual monopoly on their rights to print copies. The Statute introduced a time limit and this meant that material older than a certain age would go out of control - that is - go into what we call today as the Public Domain. For reasons unknown, the Indian media and government officials routinely use the phrase "public domain" to mean merely that which is publicly available or accessible. Perhaps the government's failure to respect the laws passed by them comes out of this reasoning. Now the legal system spends most of its time taking archaic laws and constantly beaten it into shape to fit the whims-and-fancies of contemporary authorities and cases. The digital revolution however actually requires a complete shake-up of such laws and it was very nice to see an enlightened and refreshing take on the topic.

When material is placed on the Internet - there are so many "copies" - there is one on the server, there are copies on the networks, cached copies in your computer and so on. Search engines like Google makes copies, creates derivatives - indexes that allow searching and so on. So when you browse there are thousands of inadvertent copyright violations happening, if one were to take a strict interpretation. Other sites mix material from multiple sources and completely radical content produces like Wikipedia involve little acts of creativity from large masses of people which make ideas of ownership and authorship increasingly difficult to define. It was therefore of interest to see what William Patry, a Copyright Counsel for Google had to say in How to fix copyright (2012). Like many other things done in Google-style it starts from first principles - his introductory chapter is titled "Unlearning Copyright". Here he notes:
Copyright laws arose out of eighteenth-century markets and technologies, the most important characteristic of which was artificial scarcity. Artificial scarcity was created by a small number of gatekeepers, by relatively high barriers to entry, and by analog limitations on unauthorized copying. Artificial scarcity was important because it created monopoly value: not profits earned above costs of production, but rather profits disconnected to costs.
The spirit of copyright laws is that it protects acts of creativity, however the law does not distinguish the forms of creativity. For the law - a piece of software code, art, music, writing or photography is all creativity. Patry is especially enraged by copyright law advocates who claim that creativity is encouraged by the law. He asks if publishers are involved in creativity when they actually make money out of authors. A devastating case he makes is that of the movie industry and here he cites economist Edward Jay Epstein on the funding of the movie Lara Croft: Tomb Raider. Through international transactions on paper with investments leases and retransfers the movie company makes big money which is completely unrelated to the real process of creativity. However he notes that Copyright law gives the assurance to allow the financing that occurs. With tax deductions and other sops, a lot of the money is actually made through taxpayers! Well that is about the private industry but what about the government itself. Exactly what is the idea behind government?

The normative idea of government is that it enables the welfare of people within its borders. The laws of India are however not ones derived bottom up but those that have been adopted from its "rulers". The Indian Copyright Act 1957 is based on Crown Copyright and one of the most stupid clauses in it is that nearly all work done by the government is copyrighted (for a period of 60 years). That for instance means the taxation rules published by the government may not be legally duplicated by anyone. Maps by the survey of India cannot be recopied or modified - no wonder users who want to use maps or any other useful sources are forced to use freely licensed material such as OpenStreetMaps which are made by volunteers. So all the taxes paid and used by the Survey of India are then a complete waste. Similar can be said of things like the Zoological Survey of India, Botanical Survey of India and the hundreds of other government organizations. Contrast this with the laws of the United States of America - examine 17 USC 105 - Sec. 105. Subject matter of copyright: United States Government works :
HOUSE REPORT NO. 94-1476 Scope of the Prohibition.The basic premise of section 105 of the bill is the same as that of section 8 of the present law [section 8 of former title 17] - that works produced for the U.S. Government by its officers and employees should not be subject to copyright.
The provision applies the principle equally to unpublished and published works.
The general prohibition against copyright in section 105 applies to "any work of the United States Government," which is defined in section 101 as "a work prepared by an officer or employee of the United States Government as part of that person's official duties." Under this definition a Government official or employee would not be prevented from securing copyright in a work written at that person's own volition and outside his or her duties, even though the subject matter involves the Government work or professional field of the official or employee.
Although the wording of the definition of "work of the United States Government" differs somewhat from that of the definition of "work made for hire," the concepts are intended to be construed in the same way. A more difficult and far-reaching problem is whether the definition should be broadened to prohibit copyright in works prepared under U.S. Government contract or grant.
It is hard to see why India cannot follow such a simple bit of wisdom especially since we do have many talented and visionary people in the judiciary.

Now to get back to the spirit of copyright - that of creativity and examining the principles of fairness - it is hard to see what creativity is involved for instance in recording the song of a bird. All that it means is perhaps an investment in equipment which might be related to the quality of the sounds captured. Wildlife photographers often claim to be non-consumptive but even so-called eco-tourism has been shown to have huge negative impacts. Recent studies of Indian wildlife photographers and the destruction caused by them (see for instance this on a grassland near Bangalore and this on lorises ) through intrusions and  uncontrolled use of SUVs (remember that diesel is subsidised by taxpayers) across the few wilderness regions raises concerns. The application of copyrights to photographs of nature is in a way immoral. When a fashion model is photographed, the model too holds rights. The rapaciousness of wildlife  photographers is unlikely to be controllable by law or through self-regulation. Had the Ministry of Environment and Forests gone the American way, it would have made use of its humongous resources to make available photographs in the public domain (and it is well worth examining what that phrase really means ) of such things as House Sparrows. The government being a recalcitrant entity, one can only hope that at least some enlightened photographers (those that are really interested in preserving the environment) will make available (say on flickr/Wikimedia Commons) their best photographs under a Creative Commons attribution license, allowing all forms of use including commercial use, so as to demonetize and destroy the motivation for competitive photography that leads to damage of the environment.

It is very interesting to see that countries like Bolivia and Ecuador have taken steps towards being more moral in their laws. Bolivia passed a Law of the Rights of Mother Earth in 2010. It is only by accepting our biological roots that we can hope for greater morality and ethics. To assume that all humans should have a name that can be transcribed, a surname, gender, an address, a religion, an education and so on is pure arrogance. An organization like the Unique ID authority of India that asks for an address or a fingerprint is just violating human rights - in one case that of nomadic livelihoods and in another those who are  handicapped. One merely needs to see whether a rule can be extended to every other human and every other organism, if it does not, it means that the law is not based on fairness or moral foundations.

Further reading
Postscript: The Government advertisement shown above is copyrighted (in spite of being a violation of Dani Studler's copyright) and will go into the public domain in 2074, however I use this under the fair use claim for the purpose of review/critique that the Indian Copyright Act 1957 allows.

Tuesday, March 12, 2013

More lice on the naked ape

I have written about lice before and some ecological studies they have inspired. There is however a remarkably simple anthropological question about why humans wear clothes and lice crop up prominently in attempts to answer that puzzle. This rather simple and perhaps ubiquitous childhood question is all too often waved away either because it is taboo or because it has the potential to embarrass. Professional researchers, particularly from less liberal countries tend to shy away from such questions. To be fair however, one must note that the question has not been resolved entirely. Biologists typically seek advantages to such behavioural traits and look for forces that might favour the selection of dressing as opposed to the lack of it.
A newspaper article (via Google)
Books that bring human biology out of the covers usually sell well and although most of us would have read the works of Desmond Morris, it is not often that one actually examines the depth of research in popular writing. A bit of browsing led me to a rather remarkable book by Lawrence Langner called The importance of wearing clothes (1959) which is out-of-print (a digital version can be found on the Internet Archive) and what surprised me more was that it finds mention neither in the Naked Ape nor in Peoplewatching. More remarkable is that Langner, as is evident, was a deep thinker, an excellent researcher but mostly known for being a playwright and seems to have moonlighted as a patent lawyer. An excellent writer, a friend of John Steinbeck (the book is a follow up on a play "Lady Godiva" and dedicated to him) and George Bernard Shaw among other freethinkers, he questions what most laypersons take for granted about human clothing and notes the bias in the western view of cultures at his time where nudity was not acceptable as a norm unlike in some hunter-gatherer societies.

Langner, Lawrence (1963) G.B.S. and the lunatic. Atheneum, New York. p. 120
Rock paintings (20,000 to 12,000 year old)
of ritual clothing from Langner's book
Playwrights are thinkers who excel in the art of putting themselves in the shoes (or out of them in this case) of others. Some years ago I read an excellent essay on why actors would make excellent software designers for exactly the same reason - they can put themselves in the shoes of others and judge software as if they were persons. (It is not surprising that books on user interface design like About Face by Alan Cooper are well received. I have more to say on this topic, particularly on the personality traits that one could attribute to Indian government websites which shine as hall-of-shame  examples that demonstrate a lack of empathy for potential users and portray the worst of government administration. That post will however have to wait for another day.) Langner, the playwright appears to have begun considering the question of human clothing and nudity after writing a play on the subject "Lady Godiva". Langer considers the "why wear clothes" question in his first chapter and suggests protection of the body (especially of men) so as to allow movement through grass and jungle as a driving force in the warmer regions and the more obvious protection from cold in higher latitudes. He attempts to put an evolutionary structure to clothing - starting with primitive aprons for men going on to animal hides and to the use of stitching. He suggests ornaments as precursors of female clothing. In subsequent chapters he branches off into other cultural and social aspects of clothing. Langner quotes extensively from the writings of M.D.C. Crawford (Philosophy in clothing) and J. Flugel (Psychology of clothing - a title reused by Dearborn in 1918) uses many interesting notes and illustrations. Perhaps it is the lack of biological reasoning that leads to the paucity of citations to Langner's work. Or perhaps it is contempt for easy to read research by and for non-professionals, which is reminescent of the contemporary reception to works like The Origin of Birds (by Gerhard Heilmann). Langner ultimately comes to the odd conclusion that clothing brings us closer to our conception of divinity.


The puzzle of human clothing is also related to the puzzle of (relative) hairlessness. Suggestions for adaptive reasons include ideas that hairlessness may suppress ectoparasites (Pagel & Bodmer, 2003) while others have argued that it may aid thermoregulation. (Jablonski, 2004). Other bizarre ideas like aquatic origins have also be considered but the ectoparasite version gives some interesting options that can be examined using modern tools.
12000 year old non-ritual clothing from Langner's book

The larger ectoparasites of humans that are considered in most of these hypotheses are lice. There are three forms of them which live exclusively on Homo sapiens. Most lice actually specialized and live on very specific hosts. Theirs hosts form the islands on which they survive and the only opportunity to avoid inbreeding on their islands is to hop from one host island to another of the same species. This isolation means that their genomic edit histories can be compared to those of their host. This application is extremely well-known in birds where almost every species has its own specific bird-louse. When a bird species (mammals too) goes extinct, their specific bird-lice species can go extinct as well.* Birds can have several species on them and humans have one that favours the habitat of the hair on the head, another that favours the body and a different species that lives in the pubic region. While the head and pubic louse species hide in hair, the body loving subspecies is actually one that has to seeks shelter in clothes. Attempts to find the age of  divergence of this subspecies from its nearest relative, the head louse yields an age estimate of  72,000 (with quite a lot of room for error, give or take 42,000) years. (Kittler et al., 2003) This then is a surrogate for the age of clothing.

So, naturalists observe, a flea
Has smaller fleas that on him prey;
And these have smaller still to bite ’em;
And so proceed ad infinitum.      -Jonathan Swift

One would think a better estimate might be got if the study was repeated with something that lives within the lice - an obligate endosymbiont bacterium is known (first noticed by Robert Hooke 300 years ago!) - however it turns out that at least the sequences of this bacterium that were examined were almost identical between samples from head and body lice.(Perotti et al. 2007)

* Note: there have been some suggestions that the crab / pubic louse has become endangered due to habitat loss, however there is little serious evidence for it and seems more like newspaper sensationalization.
The crab/pubic louse and the head louse (from Lloyd, 1918)
 
Further reading