Blogs updates.

The published pages on this blogs are not static. Aside from publishing new post pages, existing posts of this blog are periodically updated with photos of new species, additional photos of existing species and additional information. All materials published here are the property of the author. Reproduction of any material published here in part or in total without the expressed permission of this author is strictly forbidden.



Monday, September 07, 2009

Macrotermes Carbonarius

This black termite species of the Macrotermes genus forages in open columns usually during late evenings through the night to the early morning hours. Though Macrotermes carbonarius will feed on wood in fallen trees and branches it seems to have a preference for dried leaves, grass and twigs. Of all the termites from this locale Macrotermes carbonarius is the only free ranging foragers other than those of the Nasutitermitinae sub family. Even as Macrotermes gilvus also frequently forages above ground without any cover in periods of low light their coverage area is mostly small and their columns distance from tunnel opening to feeding area less than three or four meters. Other species of snapping termites mostly of the Macrotermitinae sub family almost invariably completely cover over (with earth/soil) any of their targeted food source that are exposed above ground.

These column marches of M. carbonarius usually take place at late evening hours, at night and early morning hours. On cloudy days as well as in areas of heavy vegetation cover they may occur at mid day as well as throughout the day too. Their columns may stretch for a few meters though they are not as long as that of Hospitalitermes, a lichen eating termite species that may march more than a hundred meters to their food sources up tall trees. Macrotermes carbonarius is the only documented black termite of the Macrotermes genus in this region.

In heavily wooded areas where little sunlight reaches the forest floor such as in Malaysian primary forest the Macrotermes carbonarius column foragings may well span the day.

Macrotermes carbonarious above ground forays typically starts with a few workers termites and soldiers emerging from an opened hole in the ground. From these first forays where no predators such as ants are encountered, a few columns of workers accompanied by major and minor soldiers will then fan out to cover an area of a few square meters. Once a food source is targeted, the workers set to work chewing out small pieces of vegetable matter to carry back to the nest while the soldiers stand guard on the perimeter of these forays.

Other Macrotermes species such as Macrotermes gilvus too comes out in the open but their open foraging are restricted to within two to three meters from the underground tunnels and almost invariable during hours of low light or in area of low light.

Nest defense behavior in Macrotermes carbonarius is apparently much more advance than in most of the termites species that I have encountered. In the M. carbonarius nest mound there is clearly an organised distribution of inhabitants. Found in the fungus combs in the upper level of the nest are the workers and last instar worker and soldier caste larvae (nymphs). The bulk of the larvae especially the younger larvae are found in fungus combs in the lower levels close to as well below ground level. At the center or core of the nest is a network of labyrinth of mixed soil (usually clay) and excrement.

Most of the nymphs and  eggs are only found from ground level and below. So despite the huge mound above the ground most of the inhabitants of a Macrotermes carbonarius nest reside at ground level and deeper down. The bulk of the nest is below ground level.

In a structured defense system this black termite, Macrotermes carbonarius, response using different defensive strategy when the nest mound is breached. In a typical breach to a nest tunnel or on the mound structure a few major soldiers will rush out, probably never to find their way back to the nest again. The numbers are usually small, less than twenty individuals accompanied occasionally by a few minor soldiers especially if the breach is deep reaching into the fungus growing area. There are however few termites found in these upper layer combs. I suspect that these combs are used by last instar nymphs and workers. These easily and swiftly flee into the fortress walls and tunnels going downwards to lower level chambers once any noticeable disturbance is felt by the termites. It is probably an 'evolved' strategy against pangolins and other animals that dig into their mounds. The mound walls of Macrotermes carbonarius are typically particularly thick, over a hundred millimeters.

But once the breach extends downwards towards the lower level chambers and fungus combs close to or below the ground level, the Macrotermes carbonarious major soldier adopt a different defensive strategy. Instead of rushing out of the nest, they gather together in large numbers a few rows deep directly in front of the breached chamber with their antennae waving and vibrating their bodies in a constant rhythmic vibration that is so precisely synchronized it may well put symphony orchestras to shame. The entire nest reverberate with this perfectly time rhythmic vibration which last for several minutes.


A major soldier nymph of Macrotermes carbonarius host to Verticia fasciventris larva.

A major soldier nymph of Macrotermes carbonarius host to Verticia fasciventris, a solitary endoparasitoid of Macrotermes soldiers. The eggs are laid by the parasitic fly on the major and minor soldier nymphs. The parasite causes the nymph to grow extra large, twice its normal size but the resulting major soldier is smaller than the normal major soldier. 

A major soldier of Macrotermes carbonarius host to Verticia fasciventris. The final molt of the soldier result in a much smaller major soldier of Macrotermes carbonarius. The parasitoid larva inhabit the head of the soldier.

A major soldier of Macrotermes carbonarius host to Verticia fasciventris. The final molt of the soldier result in a much smaller major soldier of Macrotermes carbonarius with smaller head and mandibles.


Verticia fasciventris larva.

Major soldiers of Macrotermes carbonarius guards the foraging columns
Major soldiers of Macrotermes carbonarius guards the foraging columns.

Workers of Macrotermes carbonarius termites dismantling a branch
Workers of Macrotermes carbonarius dismantling a dried branch.

Trails of Macrotermes carbonarius are 'paved' with soil to soak up water after a downpour
Macrotermes carbonarius' 'paved' road. Trails of Macrotermes carbonarius are 'paved' with soil to soak up residual water after a downpour.


The paving works of Macrotermes carbonarius workers
The paving works of Macrotermes carbonarius workers. Where commonly termites build tubes or fully covered corridors for their above ground or exposed passageways, Macrotermes carbonarius paved their trails smoothing out the water logged gaps making for a smoother run for their forays.

Workers of Macrotermes carbonarius attacking a dried Oil Palm branch and leaves
Workers of Macrotermes carbonarius attacking a dried Oil Palm branch and leaves.

Minor soldiers of Macrotermes carbonarius confront a gang of Oecophylla smaragdina workers
Confrontation. Minor soldiers of Macrotermes carbonarius confront a gang of Oecophylla smaragdina workers. In the middle of this photo is a casualty, the worker of Oecophylla smaragdina.

Closer look at the Oecophylla casualty
Closer look at the Oecophylla smaragdina casualty.

Workers of Macrotermes carbonarous that strayed a bit too far from the protection of the soldiers are attacked by waiting Oecophylla smaragdina workers.

A major soldier of Macrotermes carbonarius guarding the foragers.

Major soldiers of Macrotermes carbonarius guarding the foragers.

Major and minor soldiers of Macrotermes carbonarius guarding the foragering columns.

Soldiers of Macrotermes carbonarius guarding the foraging columns.

Soldiers of Macrotermes carbonarius guarding the foraging columns.

Soldiers of Macrotermes carbonarius guarding the foraging columns
Soldiers of Macrotermes carbonarius guarding the foraging columns.

Soldiers of Macrotermes carbonarius guarding the foraging columns
Soldiers of Macrotermes carbonarius guarding the foraging columns.

The tunnel opening guarded by a major and a minor soldier of Macrotermes carbonarius
The tunnel opening guarded by a major and a minor soldier of Macrotermes carbonarius.

Macrotermes carbonarius termite fungus
A Macrotermes carbonarius fungus comb. The fungus comb of this species is not as advance as that of Macrotermes gilvus. This comb below is even less defined of most that I have come across.

A typical fungus comb in the upper chambers.

imago of Macrotermes carbonarius
A Macrotermes carbonarius imago or adult.

Wings pairs of Macrotermes carbonarius (left) and Macrotermes gilvus (right).

The Macrotermes carbonarius alate queen and a minor worker. The male or king is usually nominally smaller and slimmer.

View from top the reproductives (delated queen and king) of this black termites species "almost" look like cockroaches.

View from top the reproductives (delated queen and king) of this black termites species "almost" look like cockroaches.

Some minor workers with a major worker (left) of Macrotermes carbonarius. Minor workers are stationed mostly in the nest and are rarely found with the foraging columns.

 Head to head comparision of the Macrotermes carbonarius minor and major worker.

 Major workers of Macrotermes carbonarius.

The strong powerful jaws of the workers are meant to chew through woody substances and are not equipped for battling ants, their most common enemy. Nevertheless if an ant should falls into this set of power jaws it will be easily cut in two.

Macrotermes carbonarius major worker.

Macrotermes carbonarius major worker.

Macrotermes carbonarius major soldier
Macrotermes carbonarius major soldier.

Macrotermes carbonarius major soldier
Macrotermes carbonarius major soldier.

Macrotermes carbonarius major soldier


Macrotermes carbonarius minor soldier
Macrotermes carbonarius minor soldier. 

Long and powerful mandible of these Mactotermes carbonarius major soldiers will make short work of any ants within striking distance.

Below a major soldier and a minor soldier of Macrotermes carbonarius.

Two major soldiers of Macrotermes carbonarius. These huge and agile soldiers ferociously rushed headlong out of the nest or foraging columns whenever they sense approaching enemies. Estate and plantation workers of the early days must have suffered numerous painful bites walking in their bare feet in the early morning hours. That could possibly explain why this particular species of the Macrotermes genus is not as common as those of Macrotermes gilvus and even Macrotermes malaccensis, their open foraging activities singled them out for extermination.

The inward curving of the pointed ends of the mandibles suggest that they might be targeting more than just ants. Their targets probably include small forest creatures such as rats, lizards, snakes and frogs.

At close to 20 millimeter in total length, the major soldier of Macrotermes carbonarius is the largest soldier termite of this region.


Worker nymph (larva) of Macrotermes carbonarius.
Worker nymph (larva) of Macrotermes carbonarius


Nymphs of the major soldier of Macrotermes carbonarius
Nymphs of the major soldier of Macrotermes carbonarius.

A major and minor soldier nymph of Macrotermes carbonarius
A major and minor soldier nymph of Macrotermes carbonarius.

Nymphs and minor workers of Marcotermes carbonarius
Nymphs and minor workers of Marcotermes carbonarius.

Nymphs and minor workers of Marcotermes carbonarius
Nymphs and minor workers of Marcotermes carbonarius.

Nymphs and minor workers of Marcotermes carbonarius
Nymphs and minor workers of Marcotermes carbonarius.

A major soldier of Macrotermes carbonarius beside a major soldier nymph
A major soldier of Macrotermes carbonarius beside a major soldier nymph.

Mound of Macrotermes carbonarius.

Mound of Macrotermes carbonarius contain less than a quarter of the total nest, the bulk of which are below the ground level. Their nest can easily be differentiated from those of Macrotermes gilvus in that Macrotermes carbonarius mounds are more angular whereas those of M. gilvus more rounded from rain erosion.

Macrotermes carbonarius extending their nest
Macrotermes carbonarius extending their nest.

Macrotermes carbonarius extending their nest
Macrotermes carbonarius extending their nest.



Termitomyces termite mushroom growing from a Macrotermes carbonarius nest
Mushroom growing from a Macrotermes carbonarius nest.

Termitomyces termite mushroom growing from a Macrotermes carbonarius nest
Mushroom growing from a Macrotermes carbonarius nest.

Like the Macrotermes gilvus species there are a few subspecies of Macrotermes carbonarius. The most notable differences among Macrotermes carbonarius in different location are the size differences especially of the major soldiers. This is possibly also true of Macrotermes malaccensis though I have not personally confirm this.


Taxonomy
(no rank) Root 1
(Top Node) Cellular organism 131567
Superkingdom: Eukaryota 2759 [Domain]
(no rank) Opisthokonta 33154
Kingdom: Metazoa 33208
(no rank) Eumetazoa 6072
(no rank) Bilateria 33213
(no rank) Coelomata 33316
(no rank) Protosomia 33317
(no rank) Panarthropoda 88770
Phylum: Arthropoda 6656
(no rank) Mandibulata 197563
(no rank) Pancrustacea 197562
Superclass: Hexapoda 6960
Class: Insecta 50557
(no rank) Dicondylia 85512
(no rank) Pterygota 7496
Subclass: Neoptera 33340
Superorder:Dictyoptera 6970
Order: Blattodea 85823
Suborder: Isoptera 7499
Family - Termitidae 46569
Subfamily - Macrotermitinae 62955
Genus - Macrotermes 46572
Species - carbonarius 144742


Last Updated: 2014 07 30
First Posted: 2009 09 07
© 2009 - 2014 Quah. All rights reserved.