Tuesday, May 24, 2016

Welcome to the Jungle: Pursuing the Ancient and Modern Butterflies of Tikal   



It is interesting to contemplate a tangled bank, clothed with many plants of many kinds, with birds singing in the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us.

                                                                     --Charles Darwin, On the Origin of Species, 1859 
  

Frankly, I never thought of letters as a career. Writing has always been for me a blend of dejection and high spirits, a torture and a pastime -- but I never expected it to be a source of income. On the other hand, I have often dreamt of a long and exciting career as an obscure curator of lepidoptera in a great museum.

                                                                                  -- Vladimir Nabokov, Strong Opinions

The jungles surrounding the great Maya archaeological site of Tikal are not only filled with the remnants of ancient civilizations, but also teem with life, much of it existing in much the same way as it did tens of thousands of years ago. 

Author in the forests of Tikal
A great deal of this life consists in the swarms, colonies, and flying insects that cannot help but make a strong impression on any visitor to the surrounding jungles and that must have been a constant aspect of ancient life. In particular, if one subtracts the biting and parasitic flies and the blood drawing mosquitoes, butterflies are perhaps the most notable, constantly flying into and out of one's perceptual frame, especially deep in the most northern parts of the Peten region of Guatemala. During certain times of the year, flashes of blue, red and turquoise light up the jungle canopy, as large morphos and papilios go about their daily  rounds.
Morpho peliedes montezuma. A common butterfly in the bajo forests of Tikal from April to May.
The National Park that includes the ruins of the ancient Maya stronghold of Tikal is large, occupying some 576 sq. km. of seasonal rain forest in northern Guatemala and contains a nearly infinite variety of insect life.
Welcome to the Jungle. The author on one of his many collecting trips.
The vast diversity of the jungle flora making up the park was first characterized by C. L. Lundell in 1937, in a report for the Carnegie Foundation called The Vegetation of Peten, that recognized how difficult a task cataloging tropical life would be. More recent studies have shown the area as once having been very arid and consisting mostly of juniper scrub. It appears that it wasn't until about 11,000 years ago that the rain forest, now dominating the landscape, came into being. The ground cover of the forest today is not completely uniform and is made up of a mosaic of five habitats that support a huge range of butterfly life and composes just a small part the region's complex biogeography. The five kinds of habitat in Tikal can be generally framed as follows:


  1.  Primary upland forest. These are found on well-drained limestone soils and cover about half of the National Park.
  2.  Primary Bajo forest. These are lowland forests that in the rainy season are inundated with water and then become arid in the dry season resulting in extremes in water content. This produces very different vegetation from the more stable upland forests. 
  3.   Disturbed forest. Normally small human settlements and selective logging.
  4.  Fallow fields and second growth. Common just outside the National Park boundaries caused by slash and burn agriculture.
  5.  Edge habitats. 
During the 19th century some of the first explorers to the ruins of Tikal and the surrounding landscapes began to collect and document the huge number of species that inhabit these various micro-environments.  One project in particular stands as a monument to this pursuit, the 51 volume Biologia Centrali-Americana, edited by Osbert Salvin and Frederick Du Gane  Godman. The work, which besides the 51 volumes devoted to zoology, completed its publication run in 1902, with 7 volumes dedicated to archaeology and the photographs of the explorer Alfred Maudslay. Overall the work took more than 37 years to complete.

Alfred Maudslay
Maudslay's Photo of Stele H at Copan
The range of the zoological works alone is dazzling and can be summarized as follows: one volume on on mammals; four on birds; one on reptiles and frogs; one on fish; one on molluscs; four on spiders and scorpions; one on centipedes and millipedes; eighteen on beetles; three on wasps and ants; seven on butterflies; three on flies; two on snouted insects or rhyncota; two on cicadas and chinch bugs and two on grasshoppers and cockroaches. If one includes the five volumes on botany, which are far from a complete survey of the regions flora, the work contains  1677 plates, more than half in color, depicting 18,587 different subjects. All in all about 50,000 species are dealt with and more than 19,000 of them are described in the volumes for the first time. Most of the volumes have been recently digitized by the Smithsonian Institution and are available at the Electronic Biologia Centrali-America site.

Frederick DuGane Godman

Osbert Salvin
The undertaking of a project of this size was inspired by Salvin and Godman's reading of Darwin's Origin of Species, which in the two chapters entitled, Geographical Distribution, Darwin outlines the lack of complete species data for any one region in the tropics that might shed light on the deep biogeographical problem that, according to Darwin can be stated,

 "in considering the distribution of organic beings over the face of the globe, the first great fact that strikes us is, that neither the similarity nor the dissimilarity of the inhabitants of various regions can be accounted for by their climatal [sic] and other physical conditions."


The plates and volumes relating to the butterfly fauna found in the Biologia are particularly interesting to anyone studying the lepidoptera of the region today. The species attributions were mostly completed by Godman, whose own butterfly collection numbered more than 107,000 specimens. To work on the volumes both Godman and Slavin traveled to Guatemala, Honduras and Belize, and employed groups of both professional and amateur collectors. In the introduction to the entire series Godman writes of his collecting experiences and the impression the butterfly fauna made on him,

"In August 1861, I joined Salvin on his third expedition to Guatemala, and, after spending three weeks in Jamaica, en route, we landed at Belize; thence taking our passage in a coasting schooner we arrived at Yzabal on the Golgo Dolce. Here we remained a few days, making preparations for our journey [...] This place will, however, always be associated in my mind with my first sight of a living example of one of the most striking and gorgeous of all butterflied, Morpho peleides. I was sitting on the trunk of a fallen tree in the forest, when it came floating past me, but I was so overcome with astonishment and delight at this wonderful vision that, although I had butterfly net in hand, I was utterly unable to rise in pursuit until it was too late to capture it."

In the Biologia three species of Morpho butterflies are mentioned and pictured, but the plate gives little impression of the difficult sorting out of the many subspecies of this genus that have been identified by modern collectors and entomologists.


Morpho Plate from the Biologia Centrali-Americana
The three species included by Godman have slightly different names in modern terminology due to the complexities associated with the many subspecies of Morpho that have been identified. The three represented in the Biologia plates are shown below by their type specimens ( "a type" is the single example used by entomologists to define a species):

Type specimens from the Biologia.  From the top Morpho deidamia granadensis, Morpho theseus justitae, and Mopho helenor octavia

For Godman, the helenor group must have been extremely difficult to identify, and although he must have noticed a wide variety of variations of this butterfly in the field, he makes no attempt to sort out the subspecies, of which the Morpho peleides that he was so taken with in his introduction is one. Today the complexity of the Morpho genus in Central America is an ongoing area of research with the following subspecies recognized so far:

  • M. h. achillaena (Hübner, [1823])
  • M. h. achillides C. & R. Felder, 1867
  • M. h. anakreon Fruhstorfer, 1910
  • M. h. charapensis Le Moult & Réal, 1962
  • M. h. coelestis Butler, 1866
  • M. h. cortone Fruhstorfer, 1913
  • M. h. corydon Guenée, 1859
  • M. h. guerrerensis Le Moult & Réal, 1962
  • M. h. helenor (Cramer, 1776)
  • M. h. insularis Fruhstorfer, 1912
  • M. h. leontius C. & R. Felder, 1867
  • M. h. macrophthalmus Fruhstorfer, 1913
  • M. h. maculata Röber, 1903
  • M. h. marajoensis Le Moult & Réal, 1962
  • M. h. marinita Butler, 1872
  • M. h. montezuma Guenée, 1859
  • M. h. narcissus Staudinger, 1887
  • M. h. octavia Bates, 1864
  • M. h. papirius Hopffer, 1874
  • M. h. peleides Kollar, 1850 now in Morpho peleides
  • M. h. peleus Röber, 1903
  • M. h. pindarus Fruhstorfer, 1910
  • M. h. popilius Hopffer, 1874
  • M. h. rugitaeniatus Fruhstorfer, 1907
  • M. h. telamon Röber, 1903
  • M. h. theodorus Fruhstorfer, 1907
  • M. h. tucupita Le Moult, 1925
  • M. h. ululina Le Moult & Réal, 1962
  • M. h. violaceus Fruhstorfer, 1912
  • M. h. zonaras Fruhstorfer, 1912

With all the amazing butterfly life existing in the Maya region it is strange that we have few identifiable references to these particular kinds of insects in Maya iconography and painting. Many of the precise representations that we do have from native peoples of the region come from the Aztecs. One identifiable example is found in the sixteenth century Kingsborough Codex that details the history of Tepetlaoztoc (translated Cave-place).


Page from the Kingsborough Codex

According to Karl Taube, writing in the Oxford Encyclopedia of Mesoamerican Cultures, butterflies, as creatures associated with daylight, flowers and warmth, were commonly identified with gods of fertility and abundance in Postclassic Central Mexico. The fertility goddess, Xochiquetzal, or "flower quetzal, often, according to Traube, wears a butterfly nose piece as well as butterflies in her headdress.

In the Kingsborough Codex, shown above, we see a feather mosaic shield containing Xochiquetzal portrayed as an upward flying butterfly.  Because of the two tails shown in the image the butterfly can be readily identified as a drawing of Papilio multicaudatus, one of the largest common butterflies found in the Valley of Mexico.
Papilio multicaudatus in the wild
Besides the shield, the Codex has other renditions of lepidoptera throughout its pages and even though they are less realistic, nonetheless, the drawings still give us a sense of the variety of these insects noticed by indigenous artists.


Today, research on the butterflies of Central America and the region surrounding Tikal and other ancient sites continues, with new species being found, and the genetic variations amongst known varieties being worked out with DNA sequencing. But even as the biogeography of these beautiful and striking creatures is more and more understood,  and as their distribution is being mapped by collectors and cartographers like myself, there is still little that can substitute for the shear joy and astonishment, much like that experienced by Godman in the nineteenth century, of seeing these insects "on the wing." An experience that unfortunately in today's rapidly developing world, is getting less and less possible.
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For a modern survey of the butterflies of Tikal and surrounding region see George Austin et. al., "Annotated Checklist of the Butterflies of the Tikal National Park Area of Guatemala," Tropical Lepidotera 7 (1996) 21-37. A total of 535 species are currently known to exist in the park.