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Friday, February 19, 2010

Of Nymphs, Naked Pupae And Callows.

Ants and termites generally live (have their nests) in trees (or plants) or underground (including in cracks of rocks). Some live in both, either at all times of they colony life (the lifespan of their colony) or during some part of their colony life. True with both ants and termites (for those species that applies), some species begin their colony (i.e. the founding) underground and then migrated up trees (or other wooden structures) once the nest reach a certain size.

Some species of army ants because of the highly nomadic live style and colony size, dispense with the laborious nest building from mostly to totally, and the ants just form bivouac nests using the living ant workers themselves against some structure as a backdrop and/or overhang.

Several genera of termites build nests made either wholly or partially from their excrement. Talk about being green and environment friendly. you just can't top that. In both dry wood termites and damp wood termites, the termites' excrement are their first line and main defense. They are used to cut off and stop up the relative safety of their tunnels and passage ways from exposure to their unfriendly surrounding. While damp wood termites generally employed the same mandible strategy of mandibled termites, the general dry wood termites (outside of the few incisor species) second line of defense is to block off the tunnels and passage ways with the specifically engineered head of the soldiers.

Of the larger mound building termites of the Macrotermitinae sub family, the nest mounds are made from a mixture of sand and clay soil (where the condition exists in the habitat). Much had been written about the architectural and engineering exploits of mound building termites but little is mentioned about why (i.e. the reason and purpose) these termites (as least those of this location) would use a mixture of sand and clay to build up their nest mounds. Sand and clay both have their attributes in the face of the weathering by heat (dryness) and rain (water erosion) but when combined the sand holds the mixture together in the face of rain and the clay does the same when it is dry with the sand also preventing the clay from cracking from the lost of moisture.

Globitermes and Dicuspiditermes on the other hand build nest mounds and structures respectively from a mixture of soil, cellulose and excrement. The excrement, wood and soil nest of Globitermes sulphureus rival those of the Macroterminitae termites in ability to withstand the elements those of Dicuspiditermes are slightly less robust. But which ever the case may be all these nest mounds need active maintenance by the termites and once vacated by the termites they would eventually be worn down by the elements (particularly the rain), some more quickly than others.

Generally forest ants and non forest ants are commonly differentiated by their humidity tolerance, as is also true with subterranean ants and arboreal ants. The latter in both cases has a lower humidity tolerance. This is the reason ants such as Paratrechina longicornis, Anoplolepis gracilipes, Odontomachus simillimus, Oecophylla smaragdina and Solenopsis geminata are not found in deep forest though they may be found on the fringes. Though occasionally these may be found in the forest during the dry season, once the rainy season commences any colony that managed to establish itself in these habitats will quickly collapse with the brood and even the queen dying from the excess humidity of the forest.



The Eggs Of Ants

Ants that lay more rounded or oval eggs generally have naked pupae. In some species the larva can successfully pupate without the aid of her queen mother or her adult siblings.

Eggs of Pheidologoeton ants on the underside of the abdomen of a new queen. 
Eggs of Pheidologeton ants

Pheidologeton ants' eggs.

Pheidole ants' eggs.
eggs of Pheidole ants


Ants that lay elongated sort of sausage shaped eggs are typified by cocooned pupae (but again there are exceptions). Personally I feel that ants with cocooned pupae are more "primitive" from an evolution chronological perspective than those that has naked pupae. Some species that has cocooned pupa such as most species of Camponotus (and includes many Formicinae ants), the larvae do not need any covering material to spin their ant silk to become a cocoon while for others such as Odontoponera (also generally true for Ponerinae) the covering material whether it is soil or debri are very essential. Without this covering to aid the pupation the larvae will fail the pupating process and eventually die.

Odontoponera eggs (next three images below).


eggs of Odontoponera transversa

This Diacamma egg is near to hatching indicative from the darken center.
eggs of Diacamma ant

Diacamma worker with a clutch of baton shaped eggs.

Anochetus worker holding a clutch of eggs.
eggs of Anochetus ants

eggs of Solenopsis geminata


The first clutch of eggs by a newly mated queen or gyne whether an ant or a termite tend to have this slightly honey colored tint. This seems to apply to those which lay sausage shaped eggs.

eggs of Camponotus or Carpenter ants

Iridomyrmex.


Ants that lay more oval shaped eggs are typically characterized by a less mobile larvae and naked pupae. The development cycle from eggs to eclosion is also considerably shorter than those that lay sausage shaped eggs. Typically the time from egg to callow adult is about a month, the eggs taking around 6 days to hatch. This is typical of most ants from this region.

Sausage shaped eggs from most ants that lay them take around 11 or more days to hatch. Larvae growth rate varies with food intake. Pupa stage is around 9 to 11 days. The pupa forms cocoon which in some species required some covering over the pupating larva for this to successfully happen. Typically the time from eggs to eclosion is around 45 days. Some sources has claimed that the ants bury the larva for the pupation to take place. This may be true in some ants but for this region I have yet to encounter a species that actually buried the larva. The covering aid do not bury the larva such that it had to be dug up after successfully pupating. However not all ants of cocoon pupa needs this debris or soil covering.

Not all eggs hatched. In some species the unfertilized eggs produce male ants but in others these unfertilized eggs doesn't hatch. These are with reference to those laid by the dealated females. In quite a few species unmated gyne will break off their wings and burrow into their nesting 'media' and seal themselves in.  While in others they never do - the unmated female alates that is.

In most species of ants, the mating in nuptial flights most commonly takes places within the first half hour. As time wears on after the first hour of the nuptials, of those female alate still unmated, the bulk will remain unmated evern if there are still numerous male alates around. This is common among those species that will only mate in mid flight, all the unmated females will like the males eventually die off or become food of their predators.

Ants nuptials varies with the species and even among those of the same species there is a variation depending on the location of the nests. Some species files only at night while others only during the day. Of those that release nuptial in the day most do so during the early morning hours from the first light onward until mid morning. The night fliers flies at dusk, after dark, late into the night or towards morning before first light depending on the species with variation also within the same species.

Peak times of ants nuptials are dusk and dawn hours. All these generally applies to ants in this region (South Asia).


Eggs Of Termites.
Termites eggs are sort of bean shaped curving from the centre towards both ends. The eggs enlarged slightly as they gets closer to hatching.

Macrotermes termite's eggs. The image above showed the egg from a mature nest. Note the egg color. Below some eggs from a new queen of Macrotermes gilvus. The larger eggs are those closer to hatching.
eggs of Macrotermes termites

Eggs of a newly nested Macrotermes gilvus termite.
eggs of Pericapritermes termites

Pericapritermes termites' eggs.


Eggs of drywood termites are much larger than the typical termites' eggs, larger even than those of much larger species. they are also much longer. Typically most termites eggs are roughly around the same size, the difference between the eggs of Macrotermes from those of Labritermes are not that significant.


The Larvae of Ants.

Pheidologeton workers tending to a clusters of larvae.

Pheidologeton ants' larvae.

Typically Pheidologeton diversus suspend their larvae from the walls and ceiling of the nest chambers. The larvae are able to adhere to one another. This is most likely done to make it easier to feed these hungry larvae (see the Pheidologeton page for photos of this). Pupae on the other hand are usually left on the floor of the nusery chambers. Several other ants species does this too including some Pheidole species.

Odontoponera larvae and eggs.

Polyrhachis ants' larva.


Oecophylla ants' larva.

Odontomachus ants larvae.

Anochetus ants and larvae.

Larvae of Nylanderia with some pupae. One larva has stopped feeding and is excreting the residue of all its feeds since hatching from an egg. Larvae of ants  excrete only once after they have stop feeding.


Termites Nymphs

Newly hatched nymphs of the termite Macrotermes gilvus beside some eggs.

Newly hatched nymphs of Macrotermes gilvus with her second instar older sibling.

A molting worker nymph. Termites nymph that develop into soldiers typically are apparent only in the last two instars. Termites larvae or nymphs need to be helped by their adult siblings to successfully molt.

A Macrotermes carbonarius minor worker beside a nymph.


Macrotermes carbonarius nymphs of major, minor soldiers and workers

The last instar soldier nymph of a Nasutitermes termite species.

A soldier nymph of a Odontermes termite species in its fourth instar. Soldier mandibles characteristically only showed up in the last two instar.

An Odontermes termite soldier nymph in her preadult instar.



Above and below the nymphal form of a reproductive of the termite Odontotermes.

Above the nymphal stage of the reproductives caste of Odontotermes, one in the final instar.

Nymphs of Macrotermes gilvus in their various pre-adult stages.
Nymphs of Macrotermes gilvus in their various pre-adult stages


The Naked Pupae.

Pheidole pupae and larvae that are about to pupate.

Minor worker pupa and some larvae of Pheidologeton.

Pupa of Pheidologeton super major.



Oecophylla larva pupating. Naked pupae (i.e. ants that have naked pupae) usually need the aid of the adult workers to pupate successfully.

The Cocooned Pupae.



Camponotus pupae. One in the process of eclosion (left centre). Some larvae in the centre. Ants larva do not discharge excrement until they stop feeding and begin to pupate. In cocooned pupa the dark spot at one end is the excretment whereas in naked pupa these are remove by the workers once it is ejected as the pupating metamorphosis occurs. The cocoon typically darkens as it mature towards eclosion.


Odontoponera. As a pupa gets closer to the eclosion it typically darkens.


The Callows.
Callows are ants that have newly eclosed (emerge as adult ants) completing their metamorphosis. They exoskeleton are not fully hardened and they are typically lighter in colour than their fully 'matured' nest siblings.


Small Pheidole speices. This callow worker is about 2.5mm.

Callow of a minor worker of Pheidole ant.

Pheidole major worker viewed from bottom.

A newly eclosed callow of a Pheidologeton major worker surrounded by her siblings.

The Minor.

Pheidole worker.

Pheidologeton minor workers.

Camponotus gigas minor worker.



Monomorphic workers
Anochetus workers.
Anochetus ant workers

Amblyopone worker.
Amblyopone worker

Gnamptogenys ant worker.

Echinopla ant worker.
Echinopla worker

Cataulacus ant worker.
Cataulacus ant worker


Termites Workers

Macrotermes carbonarius. Major termite worker.

Macrotermes gilvus major termite worker.

M. carbonarius minor termite worker.

Macrotermes gilvus minor termite worker.

Globitermes worker termites.

Worker termites of Termes rostratus.

Worker termite of Nasutitermes.



Worker termite of Pericapritermes.

Termites workers are usually the first to flee into the safety of the nests or tunnels once these are breached and later working furiously to seal any breached once the soldiers have secure that from enemy ants. But the workers can be extremely tenacious and ferocious when they encounter termite workers of other nests or of other species. Below a worker of the Termes genus without even the slightest hesitation ferociously attack a worker from another nest.
Termite workers battling one another to the death

Below a worker of the Termes genus attacking a worker from another nest. The ferocity of attack rival those of the most ferocious ants. 
Termite workers battling one another to the death

Two termite workers ferocious biting one another until both are dead.
Termite workers battling one another to the death


The Major Workers.




Carebara (Oligomyrmex) major and minor workers

Pheidole major and minor workers.

Camponotus gigas major worker.

Solenopsis super major with a normal worker (right).

Major worker of Mystrium ant.

Dorylus ants major, median and minor workers


Termites Soldiers

Major soldiers of Macrotermes carbonarius

Major soldier of the termite Macrotermes malaccensis

Macrotermes gilvus major soldier
.

Coptotermes gestroi soldier.

Coptotermes soldiers

Termites Autothysis
Autothysis. A Coptotermes soldiers 'exploded' as it fought against the larger Odontotermes soldier. However this is not particularly helpful against termites as this defensive act is mainly targeted at ants.

Globitermes soldiers.


Amitermes soldier.
Amitermes soldier

Odontotermes soldiers.


Schedorhinotermes major and minor soldiers.



Parrhinotermes.
soldier of Parrhinotermes

The Gyne or queen.
Pheidole queen.

Pheidole queen.


Pheidole queen.

Anoplolepis gracilipes ants' queen or gyne.


Camponotus queen.

Camponotus queen semi claustral


The gyne of Odontomachus ant.

Queen of Anochetus ant.
Queen of Anochetus ant

Queen of Strumigenys

Queen of the Polyrhachis ants.

Queen of the Polyrhachis ants.

Queen of Leptogenys ant.
Queen of Leptogenys ant

Queen of Mystrium ant.
Queen of Mystrium ant

Queen of Gnamptogenys.
Queen of Gnamptogenys


Secondary Queens

Secondary queens of polygynous Monomorium sp. I prefer the term 'secondary' queen to 'neotenic' as these are not necessarily babies (or having baby features). Basically these queens dispense with nuptial flight (or even mating) all together and so as is the fact with evolution whatever you don't use will soon (in a matter of a few generations) be evolved (done) away with (not always completely). 


Pristomyrmex pungen's queen is not a worker  queen as is common referred to but a secondary queen

In termites secondary queens are fairly common, but in ants there are currently (as far as I know) none documented except here. In this instance these Monomorium ant queens do not ever carry wings (meaning they never took part in the nuptial swarms) of their more typical imago queens and therefore they have a correspondingly narrower thorax, especially the second (mesonotum). Please note that there is a difference between secondary queens from worker 'queens' (i.e. ants where the workers can also lay eggs and take on the role as the reproductives in the colony).

The compound eyes too are smaller than in typical imago queens of similar Monomorium genus such as M. pharaonis. The head shape is similar to that of the workers (rather than that of the imago queens) just larger. Size (i.e. length especially) these secondary queens are nominally smaller.

Ocelli are present but noticeably smaller, the presence of ocelli make these queens stands out from other non swarming queens (of army ants). Ocelli are found in drones and flying queens (i.e. alates) and in some ants also among worker ants.

All not all termites produce secondary reproductives (i.e. those with neotenic features). Some termites don't produce any at all while others the replacement king and queens are primary reproductives undergoing the full development phrases that nuptial reproductives undergo before shedding their wings to take up the role as primary reproductives.

A true neotenic queen of Prorhinotermes flavus. Note the full developed worker (left) the head capsule is far larger than that of the neotenic queen (right) which transformed from a nymph or larva (not a fully developed worker).
True neotenic (i.e. baby) reproductives of Prorhinotermes flavus, here (photo above) we have three different larva stages (instar) of neotenic reproductives (center is the largest, left the second largest and top the smallest)
The closest thing to polygyny in termites, the termite Prorhinotermes (possibly all dampwood termites and also other termites) tend to produce a large number of reproductives (i.e. secondary queens) in a colony. In this colony of around sixty (60) individuals, eight are reproductives (one male and seven female, one of which is nanitic a solder)


The Male or Drone.

Ants are like bees where all the contributing members of the nest are female except for the seasonal male whose sole existence in the order is to mate with the female alate during the nuptial flight.

In Termites the workers and soldiers can be male or female. Researchers have determined that the larger of these workers and soldiers tend to be females while the smaller (minor) are males. Generally also in the reproductives, the female alates are noticeable larger than their males. 

There may also be a job separation between the reproductive pair as they found new colonies. Unlike ants where the gyne or queen will feed the larvae with her eggs, in termites generally it is the males which feeds the nymphs usually from his internal reserves while the female lay the eggs until the first brood of workers starts tending to the nest.

In some termite species, those that feeds directly on wood, it is not uncommon to find males with truncated antennae or a missing limb or an amputated one. These appeared to have ended up as nourishment for the female during the nest incubation period. In this case it is literally true that the male have to give up a limb or two to raise his family.

Solenopsis geminata.

Odontomachus rixosus.

Odontomachus simillimus
male of Odontomachus simillimus


Proceratium

Camponotus gigas


The Female Alate (unmated queen).




Tetraponera

 Camponotus (Colobopsis) female alate

 Camponotus (Colobopsis) female alate

Proceratium

Termites Alates.




Termite Queens or Gyne.

Macrotermes gilvus queen and king (partially hidden behind the queen). For fungus growing termites physogastry of the queen normally takes place after the first year of mating.

  A young queen of a small termite species

Queen from a Nasutitermitidae termite

Neotenic (secondary) queen of Prorhinotermes flavus from a nymph or larva (not a fully developed grown worker).

Secondary reproductives (female) and eggs of Prorhinotermes flavus. Secondary reproductives of termites are usually metamorphized from workers but those of dampwood termites can be from either soldiers and workers.
In some termites they are more commonly from nymphic stage larvae but in others (such as dampwood termites) from the pseudergates or non imago termite workers or even soldiers which transform (or metamorphize) with or without molting to become sexually mature with some even developing the rudimentary compound eyes found only in imago termites (i.e. matured reproductives) after their transformation. 
 Photos above and below, a pair of primary reproductives (king and queen) with a pair of secondary reproductives (male and female) of Prorhinotermes flavus. The (primary) king and queen are the ones with eyes while the secondary king and queen are without eyes. Those with fatter (or more bloated) abdomen are females (i.e. queens). In photo above, (top left) male secondary reproductives and (center) queen while the two with heads hidden (top) female secondary reproductive and (bottom) the king.

Photo below, top are the two females (i.e. queens) one a full fledged queen the other a secondary queen. The kings are below them in the center of the photo. These are from new colonies. Colony development and growth (in numbers) are pain takingly slow.


Secondary queens from termite soldiers (one full exposed the other partially hidden except for the abdomen). Not just workers (aka pseudergates) of this species (probably includes the rest in this genus) can transform into secondary kings and queens but the soldiers (also are pseudergates) also do the same though generally not as common as the case with the workers.

Some neotenic (secondary) reproductives (three hiding in the crevice on the left and two moving about on the right - note the two on the right the fatter is the queen) of Prorhinotermes flavus 
(click on video then click play to watch)

Termites mating takes place after the founding pair has successfully sealed themselves inside their nest. Mating position is similar to cockroaches with the male and female end to end. Mating is a frequent event (a few times a week during the founding stage and possibly the same thereafter, meaning that for termites sex is not just strictly for procreation) triggered by currently undiscovered (meaning I haven't got a clue except maybe they enjoy having sex) reasons.


The Nuptial Swarms

Post swarming frenzy of Odontotermes sp.
(click on video then click play to watch)

Odontotermes sp5 swarming in the morning
after a night of rain. This particular species has unusually fragile wings that disintegrate when in contact with water. So in them the nuptial flight has become the nuptial crawl.

Pericapritermes swarming immediately after the rain while it is still drizzling as a way to minimize predator on the colony while they are "opened" for swarming.

Macrotermes gilvus female alate fanning her pheromone to attract males. In many termites the female alate in the nuptial swarm once she landed will lift up the end of her abdomen to release her pheromone to attract the male alates. Not all termites species does this but many do. Nuptial swarms are perilous times for the participants and even more so for termites where large (not all termites release large swarms) swarming is basically a mass offering to nature to have a feast at their expense.

A female Cryptotermes cyanocephalus fanning her pheromone to attract males.

In some termite species the alates will check out one another to confirm that they have paired up correctly (both male/female pairing as well as same species pairing) before proceeding with their pairing tandem run. But not all species do this, some are in literally a mad rush to do so in order to quickly run for cover from the deluge of predators descending to feast on them.

 Acistrotermes alates begins their tandem pairing run to find a suitable nesting site. 
They soon break off their wings as they prepare to dig into the ground excavating their incubation chamber where they will remained until the first brood of workers and soldiers began to forage for food to build up the colony.

Tetramorium ants releasing nuptial at dusk
(click on video then click play to view)

Solenopsis geminata swarming at dusk
(around sunset time) after a day of rain. Male to female alates are more or less even in ratio.

 Camponotus parius nuptial begins soon after dark after a wet (rainy) day
 Typical swarms are very much larger than those of Solenopsis geminata which swarm more frequently, with males (drones) outnumbering female alates more than ten (my estimate around thirty to fifty to one female) to one. A typical swarm of a large mature nest releasing around thirty to forty females (yes I counted).


Fungus Cultivation

Here side by side is a fungus block of fungus growing termites (left) and (right) a piece of basically dry form excrement nest (of a species of Nasutitermes) that many wood eating termites nest are either made of (some are combined with wood, clay or fine sand) or are lined with. The progression from the dry form excrement to a fungus 'garden' is a basically natural progression.

The dry excrement nest of a Nasutitermes species.

With fungus growing leaf cutter ants, fungus growing in all likelihood began in a fairly similar way. Here is a relative of the leaf cutter ants with the makings of a potential beginning for fungus growing. These ants (Proatta butteli) line their nest bottom with dead and dried vegetation (among other things).



Updated: 2022 06 27
First Published: 2010 02 19
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