ANTS
Ants probably evolved from parasitoids [Malyshev]. during the Jurassic. It is probable that the development of the secretion of phenyl acetic acid from the metapleural glands is what made it possible for ants (Formicidae of the Hymenoptera) to take up a subterranean habit very successfully by acting as a fungicide and bactericide [Holldobler & Wilson (1990), p30]. The metapleural glands had evolved at least by mid Cretaceous [Wilson et al., 1967]. That the fossil which establishes this was very primitive does not remove the possibility that progenitors of legionary ants existed already in, say, South America, as well as probably Ponerine ants all over, any more than discovery of a duckbill platypus would deny the existence of rats today. A Ponerine fossil has been discovered [Agosti, et al]. It is plausible these glands existed by early Cretaceous or earlier. This statement is also suggested by molecular genetic analysis, which indicates that most of the major subfamilies had started to separate by mid Cretaceous [Moreau]. It is probable that progenitors of rain forest legionary ants (Dorylinae) were able to follow the rain forest termites across a Bering Sea bridge on the moist rain forest coastal regions sometime during the Cretaceous eventually, most likely near the close [Schneirla]. Molecular genetic analysis places their divergence many millions of years after the Cretaceous close [Moreau]. However such huge colonies with a single queen and implications of inbreeding implies very slow evolution, so their actual diversification must go back well before that, probably closer to Schneirla’s estimate. Their progress must have been very slow to migrate. This is because they do not send off female mating flights but reproduce by colony fission. Therefore they could not cross a swamp or river unless a raft happened to touch down on the opposite side, which would be a very rare event for a subterranean insect fielding large colonies, or an oxbow was cut off. Leaf cutting ants can bore under a river in the current world and thus presumably provide a path for Dorylines in South America [Branner (1910), p473], but it is doubtful that they had such an attribute in the Cretaceous, or leaf cutters even existed then. Dorylines could not inhabit a raft for more than a couple of days, for they starve to death in a short time without food [Holldobler & Wilson (1990), p586]. African above ground driver ants can form a ball and float the whole colony on water [Wheeler 1910]. I suspect that underground driver ants can not. That last trait may have helped them reach Africa but such a ball almost certainly could not have crossed an ocean and they never reached Madagascar, which last was no doubt additionally ruled out because of no rain forest on the opposite African shore. The absence of animals on Madagascar other than those which can fly, swim, or could reasonably be expected to occupy a floating tree, such as dinosaurs, birds, lemurs, a hippopotamus, and Ponerine ants, would imply that Madagascar has been cut off from Africa for a long time, probably always. Large mammals are very poor at crossing oceans [Carlquest p35-38] and there are no Triassic mammals at present, for instance, such as Australia's duckbill platypus and few in the distant past. Cretaceous primitive mammals are not known to have had much diversity [Flynn] on Madagascar and all four of the existing mammal groups are from a single species origin [Yoder]. Madagascar termites appear to be stray immigrants, recently arrived [Paulian R p293] and are primarily from semiarid regions [MacKay p118].
Army ant's progress across the world was probably very slow. Gotwald believes that they split up into two families in the new world, Dorylini tribe in Africa (but 4 species back to Asia), and Aenictini tribe [Holldobler & Wilson (1990), p584] in Asia. If so, their Ponerine, possibly Cerapachyini tribe [Gotwald (1979)], progenitors must go back a long way in moist tropical rain forest, at least to late Cretaceous, but probably much earlier. The underground species of both those legionary ants prey on termites and other ants, Aenictini specialized more on ants. None of the army ants attack their own species and seldom other army ant species [Gotwald 1995 p158]. That would be a logical development for they are nomadic and it would be ruinous to defend a nonexistent territory from such a dangerous foe. They have since both migrated widely through the old world moist tropics and Aenictini alone of the two to Australia. Ecitoninae and the primitive Cheliomyrmicini tribe remained in South America, and, maybe previously before the glaciers, in North America.
Nothing is known directly of their evolution since they have left no early fossils [Holldobler & Wilson (1990), p585]. It is possible that their Ponerine progenitors go back a long time (probably before 100 million years ago based on genetic analysis) in the past in some isolated area, most likely in South America. They may be considerable of the reason why huge forest herbivores remained in South America until late in the Cretaceous. The early species were underground and very specialized [Gotwald (1978)]. Very little is known of the habits of present day under ground species because of the extreme difficulty of digging in the hot tropics for a nomadic, widely dispersed animal armed with a poisonous sting. There are 248 species known, 144 in South and North America alone [Holldobler 7 Wilson (1990), p587] and all are nomadic mass hunters. This is a high number for an insect with so low a density of reproductive units, such a long queen life and such a circumscribed life style, with its implication of inbreeding. Of course, on the other hand, the difficulty in emigrating must have increased species formation somewhat by creating many cul-de-sacs. Also they tend to specialize. Some prey mainly on ants, for instance. This would tend to permit species separation even in the same area. If a species that preyed on ant colonies entered an area where there were no Dorylinae ants that specialized on termites, one would suspect that termites there would receive an immediate and dramatic boost because non army ants are half as abundant where Eciton occurs [Gotwald 1995 p225]. The termites would then enjoy this advantage for a long time because of the difficulty Dorylines must have in emigrating. Where legionary Dorylinae are present, legionary Ponerines and other similar families do poorly or are eliminated completely [Holldobler & Wilson (1990), p590]. Since the underground Aenictini tribe tend to specialize in preying on ants [Holldobler & Wilson (1990), p595], it may be that the Asian rain forests were especially hard hit by termites before those few African Dorylini tribe species had a chance to migrate back to Asia. It is possible that this is still the present day situation in northern Australia.
Dorylinae South American progenitors must go back at least before the time when North and South America separated. This is not absolutely certain because lower Miocene or Oligocene fossils are found in the Dominican Republic [Holldobler & Wilson (1990), p585]. The Neivamyrmex involved has 117 species, ten times as many as other Ecitoninae genera, so it is possible that they have a fairly modern attribute that we do not know about which permits them to cross short ocean barriers. Neivamyrmex have a very wide distribution. Dorylines must all have been derived from totally underground species originally because they are all blind. This, along with the absence of flying female reproductives and an instinct to rescue trapped nest mates, would help explain why there are so few fossils. There is a good chance that they reached western North America by a million and a half years after the close of the Cretaceous because there was a rain forest growing on the western slope of the Rocky Mountains in Colorado then and this forest had much more diversity than forests growing elsewhere [Johnson]. Also coal appeared immediately after the start of the Paleocene [Archibald p 249].
Other ants radiated out enormously in almost every herbivorous and carnivorous direction during the Tertiary, with several families started by the Eocene. This would make one suspect that this radiation started even before the Cretaceous closed. However it is not possible to tell for sure from distribution because ants other than Dorylinae can cross fairly wide ocean barriers. However, the likely circumstance is that ants which could hunt in packs were absent from Cretaceous savannas because that type are not cosmopolitan even today. Primitive ants make up only 1% of the Cretaceous insect fossils [Holldobler & Wilson 1990, p23]. There were only two individuals out of thousands in the Alberta, Canadian Amber, while in the Tertiary ants were among the most abundant fossils [Wilson 1987]. However, there were very few fossil insects in the Cretaceous anyway [Carpenter 1953 p268], especially in the tropics. What few were trapped in amber tended to be tiny, perhaps because larger insects were able to escape. Therefore it is very likely the lack of ants that hunted in packs made Amitermitinae and other termites still virtually unopposed in savanna areas during the Cretaceous.
Soil borne termites in general undoubtedly would have been able to spread and/or regenerate immediately after a Yucatan meteorite impact [Alvarez] and/or the Decca traps formation [McLean 1985], the last probably meteorite induced by antipode (opposite side of a sphere) disruption from a meteorite impact, for it is unlikely that an underground creature would be much affected by the wild fires hypothesized at the close [Melosh et al. 1990,]. It is even less likely that a saprophytic organism would find any scarcity of food with enormous amounts of dead vegetation all over and there was surely plenty of new small plant growth for plant smothering Amitermitinae (now designated inside Termitinae). There was not even much change in other families of insects across the Cretaceous-Eocene boundary [Whalley (1987)], never mind termites. Neither would a brief two week period of meteorite impact cold [Wolfe 1991] have affected Amitermitinae much because they withstand such a situation every year in Texas. Thus the Paleocene was primed for a deluge of termites.
MARINE PHOSPHORUS
Marine carbonate maximums correlated with phosphate minimums in the Cretaceous [Follimi (1994)]. This is corroborated by an experiment in the present day on the Great Barrier Reef. Phosphorus but not nitrogen inhibited calcification 50% when the phosphorus was raised 10 times. At the same time productivity rose 25% during the 8 month duration [Burke (1994)]. This gives additional support to the concept that phosphorus increased in the ocean at that time inversely to the calcium carbonate's rise and fall. This in turn could have been tied to the relative rise and fall of the fortunes of Amitermitinae termites in their evolutionary battles with ants. In addition, there is the possibility of the complications of migrations of evolutionary changes back and forth across land bridges and narrow ocean barriers. While South America probably did not participate in the early migrations of Amitermitinae, North America and Eurasia make up the bulk of the tropical land area now and probably did then, so these areas alone could easily have accounted for the marine phosphorus. There could have been some migrations of primitive Termitidae termites, which made nests in logs, early on in the Jurassic from Africa to South America via floating logs carried by lenses of fresh water in the equatorial ocean current. This is because fresh water fish were sporadically present on both continents later than any possible reasonable connection even given the unlikely circumstance of a closed Triassic ocean, with South American species descended from few fish ancestors [Darlington 1965, p168]. Indeed, I doubt if there ever was a land connection in the last 3 billion years because of shallow ridge earthquakes and continental rocks in an Atlantic ocean rise among other anomalies. There was also extensive interchange of plants worldwide in mid Cretaceous, probably because of bird evolution, which may have permitted carrying angiosperm seeds from the Ontong-Java Plateau and considerable interchange in late Cretaceous and Paleocene. Even in the Eocene there were common mangroves across the Atlantic [van der Hammen p308,309]. There must have been an occasional lens of fresh or brackish water from a very heavy ancient million year Congo River flood that was able to ride across the Atlantic on the south equatorial current. Such a lens could have kept primitive Termitidae alive in floating logs on rare occasions, as well as large dinosaurs capable of swimming. Obviously primitive Termitidae did arrive early on (I suspect the late Jurassic) in South America from some source to permit the evolution of Nasuti termites there, but Amitermitinae were very late.
Those Amitermitinae probably came down to North America from east Asia across the Bering straits. The spike of temperature in mid Cretaceous [Huber], the Arctic Ocean going up to 27 degrees centigrade in the Coniacian (93.5 million years ago) [Jenkyns] possibly from trees in the high Arctic [Otto-Bleiser] but even more likely from removal of vegetation by soil smothering termites because leaves in warm climates have adaptations that lower leaf temperatures toward their optimum photosynthetic temperature [Helliker], which should lower climate temperature a little. Also that removal of vegetation probably allowed the soil to become very hot in the subtropics at least, and may have permitted migration of Amitermitinae across a Bering Sea land bridge to North America. You can verify such a rise in temperature by touching a stone walk or bare soil and then adjacent grass and noting the dramatic difference in temperature.
From early Valanginian [Follimi 1994] to early Hauterivian, early to the middle of the late Aptian, latest Aptian and earliest Albian of the Cretaceous there was widespread phosphogenesis both on the shelves and in the deep ocean [Follmi (1994), p742,743]. I have a very strong suspicion that this ocean phosphorus was made largely possible by Amitermitinae runways.
It may be that the decline of ammonites near the end of the Cretaceous [Paul] was because the rise in phosphorus gave their competitors, the fish, a relative advantage given that the ammonites used a calcium carbonate shell while the fish require phosphate for theirs. The ammonites could not escape the huge air breathing vertebrates also made possible by the added phosphate while the agile fish could. Also anoxic conditions on the sea floor which tends to follow an excess of phosphorus may have been hard on a bottom dwelling creature [Harries 1993]. Ammonites almost completely disappeared by the Paleocene, although this may also have been related to a considerable extent to the pH and/or calcium status arising from enormous loss of calcium in Foramina, mollusk shells, and etc.
These phosphorus trends may not have been absolute, especially at the close of the Cretaceous. Sands deposited on the banks of an ancient river delta in New Jersey was low in phosphorus at the close of the Cretaceous [Weber (1993), p114]. It is conceivable that termites migrating from several regions in both directions created such a diversity in them that they were able to strip upland savannas of much of their tree, shrub, herb, and mulch cover and thus allow large amounts of infertile upland subsoil to erode down near the close in New Jersey, for on a short grass prairie in the south west of North America from which termites were killed, runoff and erosion were much less [Spears]. Also it has been shown that where vegetation and woody debris was removed from southern Australia river banks, the river channel was carved much deeper and the meandering of the river was dramatically increased [Brooks 2003) ]. Removing forests from a valley in Nepal caused earth to be eroded away one hundred times as fast as when it was forested [Hanson]. Alternately, the development of Mastotermitidae in eastern North America, especially their ring barking capability, could have been one of the phenomenon involved for eastern North America was cut off from the west by an inland sea until just before the close of the Cretaceous (also see an extensive discussion at this site.) Mastotermes in Australia is not very numerous nor are the colonies large. Its diet is very varied. It will eat introduced plants, damage ivory and leather, and eat wood and debris. It becomes a major agricultural pest, to the extent that vegetable farming has been virtually abandoned in Northern Australia [Hill 1942] wherever this termite is numerous, which it is outside of the rain forest or bauxite soils [Brittan]. It has developed the ability to bore up into a living tree and ring bark it such that it dies and becomes the center of a colony. Late Cretaceous rain forest had an open canopy lasting into the Paleocene [Collinson]. There was open tree growth with rotted interiors in Texas also, with few shrubs [Lehman]. It would not take long. Four years after termites were removed from an Arizona desert soil, the mulch had tripled and was still rising [Bodine and Veeckert (1975)]. I assume it would be even more rapid in the tropics. Even today in Africa, even though there are potent savanna ant predators, the soil is much lower in organic content where termites are present [Jones1989 (this article also contains phosphorus data)][Jones1990]. It is possible that this phosphorus dearth on land may have paradoxically caused significant increase of plant diversity [Wassen ] and maybe even small animals.
A low productivity of the oceans lasted for half a million years into the Paleocene [Zachos et al. (1989)]. It is possible that this low productivity was a function of a temporary dearth of phosphorus because there was a downward spike of phosphorus in ocean bottom sediments in the Indian Ocean at the beginning of the Paleocene [Cook (1984), p263]. Phosphorus is what determines the productivity of oceans, not nitrogen or iron [Tyrrell]. If this productivity decline was a function of phosphorus, the phosphorus famine must have also ended after the one half million years because the marine phosphorus sedimentation rose to even greater heights in the Paleocene than it did in late Cretaceous and lasted until the end of the Eocene. Those deposits were clustered around Africa and India [Cook (1984), p255], So it may be that these areas previously had termite faunas which conserved the soil's phosphorus in addition to being inefficient, and then were afflicted with Amitermitinae in an area which was still rich with soil phosphorus on the savannas. This is plausible because the dominant termite in Africa even today is the Basidiomycetes fungus [Gillott (1995), p168] cultivating termite, Macrotermes [Harris (1949)] or Termitomyces fungus [Howse p 19]. ]Its prototypes would seem to go back at least to the Miocene [Jarzembowski (1980)] for that is when they are thought to have reached the Orient [Bouillon 1970]. They must go back in Africa to when primitive African Termitidae split from South American Termitidae [Emerson 1955 p486] in late Jurassic or early Cretaceous because Macrotermitinae use aliphatic ant repellent while Nasutitermitinae use terpenes [Prestwick 1983]. Macrotermes uses earth from the subsoil at a 60-150 cm depth and does not enrich the constructions with phosphorus [Malaka (1977)]. The fact that mound soil is high in sodium while topsoil is not [Watson 1977] gives additional support to the concept that subsoil is used. Therefore it would be expected that the soil would be covered with a protective layer of phosphorus deficient material from mounds and runways, since subsoils are usually low in phosphorus and in any case was not further enriched by the termites. Present day Macrotermes bring up 2,000 kilograms of earth per year per hectare in Senegal of which 950 kg are runways and sheets [Wood 1978, p255]. It is plausible that the Macrotermes progenitors were also dominant in late Cretaceous. African savannas are among the most fertile of the present day world's tropical savannas. Even so, its savanna soils are lower in phosphorus than the parent materials, unlike the opposite in the forest soils [Nye and Bertheux (1957)] and normally elsewhere plants concentrate nutrients in the top soil [Jobbagy]. If it did happen this way it would be necessary to explain why Macrotermes never reached North and South America during the Tertiary.
CONCLUSIONS
It is known beyond a reasonable doubt that amphibious insect and vertebrate predators dominated the early half of the Permian near water from extensive fossil evidence. The middle of the Permian was a time of very extensive phosphorus deposits. That these two circumstances were connected is quite plausible. Because the bulk of the herbivorous habit was by insects in the Carboniferous, it is logical to assume that the bulk of the transfer of phosphorus was via the very competent dragonflies. When the marine phosphorus declined as the Triassic approached, it was not necessarily because of a decline in dragonflies alone. The rise of very numerous mammal like terrestrial reptiles could conceivably have neutralized that flow by preying on amphibians as well as by preempting dragonfly prey. This could have contributed to the decline in trilobites toward the Permian's close since their shells are high in phosphorus, radiation of the Ammonites in the Triassic when marine phosphorus may have declined, and the decline of armor and size of amphibious vertebrates during the upper Permian.
Phosphorus started to rise in marine deposits again in the late Jurassic with its implications of anoxic bottom waters and reduced shelf and reef calcium carbonate deposition. There is no fossil evidence to date that would refute the concept that primitive social Amitermitinae (now classified as merged with the Termitinae) subfamily of the Termitidae family of termites had appeared and were contributing significant amounts of phosphorus rich sediments from their runways. The scarcity of Cretaceous insect fossils prevents us from ruling out for sure whether an absence of ants that can hunt in packs as contributed to success of Amitermitinae in savanna regions. Present day distribution of ants tends to supports this proposition though. The acquisition of a very primitive fossil ant in mid Cretaceous does not prove that highly competent pack hunting ants were not present any more than discovery of a fossil duckbill platypus would rule out the presence of prior marsupials. It does hint at it though.
Marine phosphate deposits which were laid down in the late Jurassic and early Cretaceous tended to cluster around Australia [Cook 1984, p251], so it may be that it was this area on which Amitermitinae formed its initial plant smothering evolution and then spread to Southeast Asia from there. Since the now very numerous Termitidae family had differentiated on such widely separated areas as South America, Africa, and Australia during the Cretaceous, their progenitors must surely go back into the Jurassic at least. Amitermitinae would have had to spread to North America from Asia and thus help account for the drastic decline in savanna animal size in Asia up to the Paleocene [Archibald p XII] and the fact that phosphorus marine deposits rose to double what they had been in late Cretaceous during the Paleocene [Zhow 1992] after a brief decline. It is possible that the evolution of the blind, underground legionary ants of the Dorylinae family in South America helps explain why the largest of the Dinosaurs lasted much later in that region than elsewhere. In addition, arrival of Amiteritinae was probably late there. Determination of the timeline for phosphorite deposits off South America would throw light on this last.
Savanna vertebrates showed a trend toward increase in size and bony structure after the Eocene in Eurasia, such as horses. This corresponded to a decline in marine phosphorus deposits. It is plausible that the increased ability of Ponerine and Myrmicine, especially Ponerine, ants to hunt in packs on the savannas was the means by which both of these events happened.
Continue to affects of termites on soil phosphorus.
or to Paleocene phosphorus famine and Tertiary recovery
REFERENCES for "Permian Phosphorus", "Jurassic Termites", and "Cretaceous Ants (this article)"
Back to Permian wood roaches and the coal hiatus
Back to Jurassic termites and phosphorus flow to the ocean.
Back to phosphorus flow to Permian oceans by dragonflies and amphibians
For a geological time scale. You may see a chart of all the dinosaur families with respect to time and an extensive geographical discussion at this site. This is a discussion of dinosaurs’ probable posture.
This site shows very good photographs of all the termite families.
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A proposal as to why the human species seems to be derived from a single couple: The Eve Controversy.
There is a free browser called Firefox, which is said to be less susceptible to viruses or crashes, has many interesting features, imports information from Iexplore while leaving Iexplore intact. You can also install their emailer. A feature that lists all the URLs on a viewed site can be useful when working on your own site. .
Charles Weber Email = --- isoptera at att.net or 1 828 692 5816>
