At The Feet Of The Giants

The Lords of the Grasslands, in Kaziranga National Park.

The children’s stories of the ant and the elephant have always made me wonder what their true relationship is – the versions I heard pinned the elephant, proud and powerful, against the ant, timid but sharp, the tale ending with the ant stinging the elephant in a place it cannot reach – literally and allegorically. In most stories, the ant symbolised the underdog who triumphed over the elephant, rarely did they both work together or become friends. Long after, I started working on the concept of ants to elephants, and not merely because they rhyme. Through the tales and their scale, they represent most of the animals I grew up watching and admiring and studying, but in this case, it was simply connecting two organisms I am passionate about, insects and elephants. Come to think about it, plants to elephants encompass a much larger scope under this philosophy.

On various occasions, I explored insects and elephants independently. Studying both the organisms has always been mutually exclusive, but I started to develop the concept along the way with some questions: what is it that connects elephants with ants? How are ants and elephants connected in an ecosystem? Simply put, the concept of ants to elephants recognizes the functions of insects (represented by ants) and large animals (represented by elephants), and how they both complement, cooperate, and even antagonise one another while interacting with the shared terrestrial habitat or ecosystem they are a part of. I continue to connect the dots or, in other words, observe the threads, one at a time, of the grand tapestry of nature. This is one step in that long journey.

The Giants of the World, Mount Everest (centre left), Lhotse to its right, and Makulu (centre right).

It wasn’t until an opportunity arose that I would start connecting the dots ecologically. Long before I could make this connection, the scale of things as they are was made clear through a window some nine kilometres up in the sky as I stared dumbstruck straight at Mount Everest, Lhotse, and Makulu, slowly but quickly moving past Jomolhari, past the gigantic Sleeping Buddha of Khangchendzonga, but the mere sight of the mighty Himalaya, skirting west-to-east in a straight line over the horizon, did not give a perspective until the clouds below cleared to reveal a vast stretch of draining rivers to form the most populous fertile plains on the planet – the terai and the duar, formed by many rivers big and small, emerging from the foothills of the Himalaya, trickling into being further up the tall glaciers along the shoulders of the mountains looming over the clouds.


Spraining my neck to take a good look at the mountains – three of the world’s tallest mountains (over 8 km straight up) – I have experienced only a few times over much of my adult life, I found myself fighting the urge between photographing through the tinted glass and absorbing all that I could with naked eyes – it felt so near yet it was so far. I whisked past them in no time to be greeted by a great haze over much of another giant that takes birth in the mountains, the Brahmaputra, as I touched ground.

The Guwahati air did not feel very different from Mumbai’s. In a perspective, I witnessed the mangroves to the snow-clad peaks within half-a-day, but I did this with the doom of car-brained development over intertidal zones to dams built upon mountain rivers bursting – all happening at the same time. Fully aware that my footprint on this planet is not worth the views, I descended back into myself thinking of the harm in invisible but not-so-congruent footprints I was leaving behind.

Big views helped me set my mind on big hopes – surely the Himalaya had to be followed by other entities larger than life. Few places offered this privilege, and I was headed towards one. Kaziranga. It appeared not over the horizon but as a low-lying plain more bowl like than a dish, the Assam Trunk Road leading through it built upon pile of rubble splitting the bowl into two halves, one up north towards the Brahmaputra and the rest down south leading up the Karbi-Anglong mountains. Traveling eastward, the living giants – the elephant, the rhino, and the wild buffalo, lived somewhere here, I thought to myself, between the great Himalaya to my left, I travelled through the vast grasslands and cane-riddled forests of the Brahmaputra, and the Karbi-Anglong mountains to my right. It was between these scales – not with these scales – that I was to work.

Heading into the forests of Kaziranga landscape.

One bright sunny morning at 4:45 AM, I was to wake up listening to birds chirruping and light gleaming in from the high window, an unusually bright early morning for a Konkan resident. Before 6 AM, I greeted six armed guards, sharp and military-like, intimidating me more than the talks of wild elephants, rhinos, and buffalos walking the same forest I was to. Luckily, they were here precisely because these big three walked this landscape, of course not to shoot them but to ring shots in the air should they charge at us. Not that they had to at any point in the course of my explorations, but it took me a while to get used to guns around guiding my way through the forests.

A landscape of woodlands transitioning into wetland-grassland complex of Kaziranga's beels.

Based on its natural vegetation, terrestrial Kaziranga can be simply divided into grasslands and woodlands. The more nuanced types are versions of these two broad categories, encompassing floodplains and marshes to riverine woodlands to semi-evergreens. The purpose of my visit was to explore insects and spiders that call the greater Kaziranga landscape (comprising the National Park and the adjoining Reserve Forests) home, but in a short time, we could only focus on a fraction of one, and we decided upon woodlands given various factors including the threat from the big three in grasses denser than forests and taller than elephants. We began in earnest after seeking approval for non-sampling exploration of insects and spiders – much like how one sees mammals and birds in their natural habitat – with a camera and a lens.


It began with the dazzling giants of the insect world flitting about wherever I roamed. Under a leaf hid a Nawab – there are at least five species of this butterfly here compared to only two where I come from, all looking more-or-less the same, leaving me amazed at the diversity that the place has to offer. The most common were larger-than-life, the aptly named Great Mormon; this large black-and-blue butterfly was everywhere, and there was the large Yellow Helen, also everywhere – butterflies that would cover more than my palm if they were to perch on my hand.

A Great Mormon basking along the edge of a forest.

During one of the walks along the Hathidondi – any and every forest trail made by elephants is called so –, an elderly guard made me aware of a large butterfly that flew like a bird. Like a bird? – I asked him. He said yes, black-and-yellow. Not really aware of the butterfly diversity of the region, I wondered if it could be a birdwing. And it surely was. On the way back, I saw one gliding on its large sail-like wings across the forest edge, disappearing into the tall canopy as I shouted for my companions to look for a butterfly as big as a bird.

Days later, I found myself in the middle of a sandy meshwork of streams originating from the Karbi-Anglong mountains, a land invaded by tall lantana thickets turning it into a maze. Somewhere here rested a male wild buffalo, we were informed by sand-diggers. Save for a loud grunt we heard at one point it remained unperturbed by our wanderings around the stream. By this time, I had spoken to everyone I met who led the way about Common Birdwing. It wasn’t difficult to capture attention and imaginations of the guards. I put it up with the famous Big Five of Kaziranga: the one-horned rhinoceros, Asiatic elephant, Asiatic water buffalo, the tiger, and the Eastern Swamp Deer. Then there was the largest venomous snake of the world, the King Cobra, the many-hued birds – from hornbills to storks to raptors – to the largest gecko of India (and second globally) the Tokay Gecko. The Common Birdwing, I proposed, is the sixth wonder – if not literally, definitely proportionally – largest wing to body-size ratio, the largest tongue (proboscis) of all animals, and of course, its free-spirited flight through the forest when the sun shines the brightest – even the cobra is weary of where it treads and the eagle where it perches.

The Common Birdwing, a graceful symbol of the insect diversity of the Kaziranga landscape.

After climbing down the mountain resonating with Hoolock Gibbon cacophony, Milan Deka, who patrols this woodland, spotted one puddling by the streambed. I ran to see it fly away and out of sight, leaving Yellow Helen alone, who didn’t mind our company. I requested we spend a few more minutes, which turned into an hour, and Deka ji’s sharp eyes helped us realise we were in the midst of a birdwing haven made up mostly of an invasive plant, lantana. Whenever the sun was out, they were everywhere, feeding on flowers, chasing one another, both male and female on their glorious wings, and when clouds rolled in, they settled on the tallest shrubs. This game of hide-and-seek took me back in time when I would chase butterflies where not one is seen anymore, in places where no one can visit again. Those black-and-yellow wings were mesmerising enough, they entranced Deka too, who memorised the name and became a companion butterfly-watcher in the buffalo marshes.

A female Common Birdwing, hover-feeding on Lantana camara covering a floodplain-marsh at the foot of the Karbi-Anglong mountains.

The birdwings fervently fed on lantana flowers, taking no breaks while doing so – typical of all Papilionids, and they chased one another whenever they fed close by, hinting at some form of territoriality, especially among males who could be told apart from females for having uniform black forewings. Indeed, they behaved more like birds than the timidity butterflies are known for. Birdwings are perhaps a better bridge connecting people with the concept of ants to elephants, that is no surprise, for butterflies are always a field biologist’s first insect.

Deep in the embalmed darkness of the semi-evergreen forests of Kaziranga landscape, along elephant-trampled pathways and elephant flattened mountain shoulders, beneath the shade of giant trees lay a world: surprisingly like the world our eyes are accustomed to, of hunting, fighting, of love and companionship – bound to the laws of nature. For ten days I peered into this niche of Kaziranga landscape, my eyes slowly adjusting to the shadowed understory and hidden forest floor. That view of the Himalaya and that expanse of Brahmaputra in the back of my mind, aware of the elephants, rhinos, and buffaloes always around, and assured by the gun-wielding guards who took as much interest in insects as they were watchful for sounds the large animals make, I dove in and lost myself in this world as mosquitoes, ticks, and leeches got the better of me – one just cannot escape them.

A typical break in woodland canopy showing forest floor dominated by tree saplings.

This survey revealed 190 species – comprising of moths and butterflies and dragonflies (most of these were wilfully ignored because they were very thoroughly studied in this landscape), including many beetles, bugs, flies, ants, bees, wasps, and spiders. In addition, we saw 94 species on light curtains at night. In other words, I saw some 28 species – most of them new to me – every day, from butterflies to ants. I wished I didn’t have to stop looking. Sitting in my dusty hometown now, I am trying to make sense of what it all means under the concept of ants to elephants.

A band of Raider Ants (Aenictus sp.) on the way up a Hal (Shorea robusta) tree.

Of particular interest to me were the social insects of the region. As I peered into the undergrowth, ants stood out to be the most abundant of all easily-distinguishable invertebrates (the real crown belongs to collembolans, diplurans, and mites – none of them insects or spiders – but since they are very difficult to find and count because of their small size, they are often omitted in field-based/non-lab studies). In nine plots with five replicates each, we found ants in all – from just one species to as many as six, in an area as small as five square metres. Overall, we found 29 species of ants, and two species of termites – both eusocial organisms. I focus only on the social insects because they resemble the larger animals like elephants as much as they resemble their invertebrate counterparts. This is where, I thought, I could lead the concept.

Titles like architects, engineers, farmers – the landscapers – of an ecosystem are generally associated with large mammals. The concept is so popular that it overshadows the functionality of ‘little things that run the world’. The social insects in particular are as close to the way megaherbivores and carnivores function as they are to their own kind. A way of addressing this bias is by framing a question: How will the both measure if compared side-by-side, weight-by-weight for their ecosystem functions? This is not about winners and losers, it is about who receives more attention and who remains the underdog as much as it is about addressing the bias in understanding how ecosystems really function – simply put, it takes more than an elephant to shape a forest – it takes a community, and that includes ants, but by what measure? It is relevant to talk about these eusocial insects as representing other invertebrates the way elephants represent most megaherbivores (I include wild buffalo in regional context) and tigers carnivores.


In Kaziranga, ants and termites are everywhere, high up on trees, in the understory, on the floor. Of 217 ant species known from Assam (as per a 2016 assessment) of the 830-odd species known from India, 29 were found within ten days of scouting, although Kaziranga hasn’t been thoroughly assessed yet. (Another large online citizen-science-based repository, iNaturalist, has photographic records of 290 species of India, 56 species of Assam, and 29 ant and two termite species from Kaziranga landscape, mostly uploaded by me and identified by the experts).

A typical worker Raider Ant, with long legs, long antennae, and no eyes.

They constitute a host of unique life-history strategies for survival. Very few are nomadic, they move from one place to another, sheltering in bivouacs and moving on, raiding termite and other ant colonies along the way. Ants in the genus Aenictus are called raider ants or army ants – they are blind and do all the ant-stuff using chemical cues and sense of touch using their long antennae. Their presence is an indication of an active eat-or-be-eaten dynamic in the undergrowth, their target, most often than not, are the lichen-eating termites.

A column of lichen-grazing termites, Hospitalitermes sp. not far from the Raider Ant migration.

Known for the formation of large moving columns, not unlike a herd of buffaloes, Hospitalitermes sp. are a highly organized group of termites that actively graze on lichen and moss in the forest as opposed to making their own fungus nurseries underground like most termites. Unlike the wood-eating termites, they are pigmented – usually dark brown, indicating an active ‘outside’ life – they move out in day and night through the forest and live inside tree hollows close to the ground in their thousands, made previously by other social dwellers like ants or crevasse-nesting bees. The workers pile out in large columns often five metres long, using human and elephant-made pathways looking for lichen pastures to feed on.

A nasute lichen-grazing termite, the one-horned soldier, guarding the column.

The soldiers, called nasutes, carry a weapon on their helmet-like head, a horn not very unlike the one-horned rhinoceros of Kaziranga, which they use to inject a terpene-based chemical which causes severe burning. The nasutes number less than workers; they stand guard along the outer margins of the procession, their horns raised up in the air and antennae moving to detect threat. Curiously, they are all blind, too.

The column leading up a boulder and further deep into the woodlands of the Karbi-Anglong mountains.

In the undergrowth, the two societies are almost always at odds, ants fight ants and ants and termites fight among themselves. It is as competitive as it is among mammalian carnivores and herbivores, if not more. I found 16 species of ants to be ground-nesting, either among rocky crevasses, base of a tree, or having subterranean colonies, 13 of them hunters and eight of them also scavengers and five opportunists. 12 were tree-dwelling, ten largely arboreal and two alternating between ground and trees. Not that they don’t visit the forest floor at all – they all spread across the nest in search of food.

A pack of Weaver Ants (Oecophylla smaragdina) overpower a solitary forager Diacamma.

Under a large Ficus tree, its mountain-like buttresses dappled in sunlight, I was witness to a clash between two ants with distinct behaviours. From the looks of it, a solitary forager Diacamma rugosum (now curiously called the Asian Bullet Ant) was out and about looking for small invertebrates to hunt or scavenge, and carry it back to its nest. On reaching the base of the tree it might have met someone unexpected – a watchful guard of the Weaver Ant, Oecophylla smaragdina. The Weaver Ant guards often posture themselves with open mandibles so that they can latch onto things in defense or to hunt – and the lone Diacamma worker walked right into them.

Among the tall shoulders of the Ficus, a fight ensues.

As the troop of Weaver Ants pulled the hapless solitary worker up the buttresses of the Ficus, I imagined the hunt as happening along the edge of mountains of gigantic proportions, bound by the same laws of nature – survival of the fittest. Even as the Diacamma is one of the most common ants in the undergrowth, indicating its successful survival in the Kaziranga landscape, this solitary worker fell prey to an ant known to work as a team. Once long ago, I had compared Weaver Ants to a pack of dhole (Indian wild dog), then Diacamma is a solitary hunter, a leopard, if you will.

The Karbi-Anglong mixed forest overlooking vast expanse of the Brahmaputra, a habitat of ants and elephants.

It is hard to escape allegorizing smaller lifeforms with their larger, not-so-different, vertebrate counterparts. This is why the philosophy of ants to elephants, and the inconvenient comparisons between the functional roles of the two in an ecosystem, is relevant. One of the reasons is that these 29 species were found in just 45 sq m area sampled randomly across the woodlands of Kaziranga landscape – an indication of their sheer abundance and diversity. Yet, in order to talk about them and their conservation, we don’t know much about them in regional context. We don’t know how exactly raider ants (Aenictus sp.) lead their nomadic lives, how much area they cover, when and how colonies split if they do at all, or why are they blind – is it nature’s way to square-off the rivals – the raider ants and lichen grazing termites?

Very small but unique woodland ant species in the genus Philidris, from the mountains.

There was one small species of ant I stumbled upon in the woods – Philidris, considered to be one of the most common ants of evergreen and rainforests of the Indo-Australian Region. I didn’t find it to be very common here. Very few plants are known for their deep-tied bond with ants (plants to ants also rhymes), aptly labelled myrmecophytes – plants which have a symbiotic relationship with ants, developing specialised modifications to attract and even house ants. Philidris is one of those species, yet where, how, and with which plants they maintain such a relationship, remains unknown in the South Asian context. One of the plants Philidris in North East India are associated with may be Dischidia bengalensis, an epiphytic milkweed with ‘pouches’ that is not uncommon in mountainous woodlands – this is likely to be this ant’s home, but it needs further exploration. Curiously, I found a small trail of Philidris on the forest floor, with about 44% canopy cover, 34% litter, and 62% ground vegetation – a mixed forest of the Karbi-Anglong mountnains. On the other side of the mountain lay the vast Brahmaputra, stretching over the horizon, framed by tall trees and a clear blue sky. The thing about many species of ants is, we don’t know what we don’t know, and that is one of the reasons why it is difficult to fully grasp the concept of ants to elephants.


I was able to mix my back-breaking view of the ants in their natural environment with that of frolicking rhinos, buffalos, and elephants. I was told that I would see all the big five, but saw only three up-close, while the tiger eluded me and the swamp deer chose to stay farther along the beel shallows. It was a landscape I had not seen before: in perfect weather conditions, usually during winters, one is witness to the Himalaya as rhinos graze along the many beels. It was too hazy for me to witness this.

The greater one-horned rhinoceros in a playful battle.

While the rhinos mostly just browsed, one evening in Baghori, my colleagues and I witnessed two high-spirited rhinos, probably a young bull and a cow, playfully fighting and softly rubbing faces. At times their horn-to-horn battles took a ferocious form, and the next moment they suddenly resumed rubbing against one another, leaving us confused whether they were courting or sizing each other up. Eventually, they moved farther over the treed horizon, where we think they finally mingled.

Two elephants greeting one another while feeding on aquatic vegetation.

At Agoratoli, I was witness to a slender-tusked elephant cozy-up to another as they fed on aquatic vegetation along one of the beels. They nuzzled and continued to feed on the vegetation as the other left for a soil-bath. All this while, frozen still, I became aware to the many biting midges (Ceratopogonid flies) that left bitemarks all over my hands that itched for over a week – the same flies that were troubling the elephants who had the privilege of a soil-bath while I was left to scratch myself through the rest of the journey.

Taking care of an itch with a dusting of soil.

The elephant may have been bitten by something pesky, for he proceeded to what looked like their regular path along the beel, grabbed a trunk-full of soil, and applied over the back of foreleg before proceeding to a continuous shower of soil till the flies were driven away.

It is important that I write about these seemingly mundane events in the wild, for to write only about ants while discussing the concept is also a bias. This was the closest I was to wild rhinos and elephants, and the most enigmatic of large mammals I am fascinated with presented itself along the banks of the Sohola beel. Even as the migratory waterfowls like the bar-headed geese and pintails were only just arriving this unusually warm early November, the wild water buffaloes were loitering and grazing on natural pastures.

A wild water buffalo grazing on the fresh grass along one of the large beels.

A cow buffalo was closest to me as the jeep stopped one bright, crisp morning. She grazed oblivious to my presence, about a hundred metres from where I sat, rotating as her lips got hold of new shoots. At one point she turned just enough for me to truly grasp the spread of her curved horns framing her entire body within its expanse, a marvel of nature – each horn at least a metre in length.

A mother and a calf water buffalo enjoying what water buffaloes do best - remunating in the cool waters.

She moved slowly toward the herd where her calf joined her, and they all grazed peacefully, just like the lichen-eating termites somewhere up the mountains. After a while they all moved to the yonder shore, where a bull sat half-submerged in the shallows. Eventually, when returning the same way, I found the mother and the calf sitting in the water, only the calf’s head and the cow’s horns visible at times, regurgitating the bank grass over much of the afternoon. Out of curiosity, I asked the guide if the cow would charge at me if I stood on our side of the bank – he said she would be upon me within a blink of an eye, enraged and ready to flick me over the trees.

A herd of wild water buffaloes at home at Sohala beel.

Indeed, my view of these megaherbivores was peaceful – playful even – as they loitered and grazed and nuzzled one another, compared to the stories I have been hearing, and the reason for walking with guards and guns. Between talks of ants and spiders, I asked them which one of the megaherbivores was the most dangerous. The consensus remains undetermined, some rank rhinos before buffaloes, the elephants rank third, probably because elephants are more predictable. When I asked who poses the biggest risk, the answer is usually elephants, buffaloes a close second, but rhinos aren’t far behind. I’m told that buffalo population has increased significantly, making them a major threat to humans, with population increase also linked to increased interactions with rhinos.

A small channel leading to another beel, showing the typical wetland habitat of Kaziranga landscape.

In the scheme of nature, human-nature interactions take many shapes and hues. It is a fine line to tread between celebrating megaherbivores wandering the grasslands and grieving for the death of a person in an elephant attack, or a rhino or elephant being shot for wildlife trade. In my brief association with elephants, it is the negative interactions that have had my attention. To acknowledge both the sides of the interactions is important when it comes to conservation of landscapes. Not all that glitters is gold, some of it is also blood.


From tall elephant grass, I espied a head peep out, and slowly a large, graceful cow elephant. She looked alert, raising her trunk half-way up like a periscope to sense the situation about her. Everything looked calm to me, but elephants hear and smell better than humans.

A mother and her baby, along the edge of elephant grass, contemplating crossing a water hyacinth marsh to other side.

As soon as she lowered her trunk a calf emerged from the thickets, following along the edge. The mother was eyeing the other end of elephant grass, where, as we came to know in the evening, her family grazed, but between them lay a vast marsh overridden by water hyacinth (Eichhornia crassipes), one of the biggest scourges of the wetlands of Kaziranga-Brahmaputra ecosystem complex.

As my attention turned to a juvenile Crested Serpent Eagle that swooped down right next to our jeep, my guide pointed to a moving bump in the marsh – an eye and a periscopic trunk barely visible. The marsh had consumed the whole of the elephant as she tried to cross to the other side.

Wading through head-deep water hyacinth marsh, the little head of the calf barely visible behind the mother.

Worried how the mother and the calf would traverse this seemingly invisible wetland choked by a network of thick leaves and roots, I sat at the edge of my seat, the eagle taking off with something I didn’t notice. After a while, a small head peeped from right behind the mother, and after many more biting minutes, the little bump moved ahead of the mother, now closer to the other end of the marsh.

It was overwhelming to see how an aquatic, floating herb has occupied this part of the world, choking small beels and straddling along the banks of steady-flowing waters, ruining this unique ecotone. In a world that appears peaceful on the surface, invasives like Eichhornia present a very pressing challenge – a survival of the fittest that it is no doubt winning.

The Yellow Crazy Ant, Anoplolepis gracilipes, waiting for a sugary treat from the Urena lobata leaf-base.

On the other end of the highly specialised blind hunters and grazers is the Yellow Crazy Ant (Anoplolepis gracilipes) – an ant known for its industrious nature. It is the only species of ant I dedicated one whole article to a decade ago, where I cover a lot of things these ants that form multiple satellite-colonies are capable of. (For more). Among many things, they are fully capable of putting an entire island’s faunal and floral biodiversity in peril. I was a bit concerned to see them in the woodlands. A species that loves humid, semi-evergreen to evergreen habitats, in India, they are mostly associated with humans – the more the disturbance caused by humans, the more widespread they are in the region. They mostly occur in the Western Ghats and the East Himalaya foothills and plains.

In the Kaziranga landscape, I found them in four plots of the 45, all of them in mixed or dry forests, all the sample sites far from direct human disturbance. It might just be a natural population, for they were not ‘everywhere’ the way they are in certain pockets of the northern Western Ghats, where trees and the forest floor were full of them. For one, it is considered native to South Asia and South East Asia, which might explain their presence here. So how can a native species be invasive? The Yellow Crazy Ants poses a unique case. Their invasiveness is due to their nature – they are opportunists, they will take over a hapless invertebrate prey, fight other ants and mostly win by sheer numbers, scavenge on dead animals, and tend to aphids and mealybugs for honeydew. Doing so, they outcompete and deplete invertebrate biodiversity of the region – if their spread is vast, they do so over a very large area. Yet, what determines their invasiveness, and why they are invasive in some but not in all areas, like in the woodlands of Kaziranga, remains to be seen. It is important to monitor invasives and potential invaders that are, more often than not, spread by humans – and that applies not just to things that we can see standing up, but also things at the feet of the giants.

Elephants making a meal out of water hyacinth Eichhornia crassipes.

Interestingly, megaherbivores love to eat Eichhornia, they consume it in vast quantities. On our return, we saw two sister elephants munching on the juicy Eichhornia stems as the sun set. It was a delightful view – rhinos too enjoy eating this weed, but it is just too much for the megaherbivores to eat it all up. In the scheme of nature, having an edible invasive plant loved by megaherbivores is a curious fit – I will go on a limb and say that Eichhornia is one of the many reasons, even if less significant, for all the megaherbivores to cohabit this shore of Brahmaputra, but who am I to theorize?


Like in the macro-ecosystem where animals interact with one another and the ecosystem – say by feeding on Outenga (Dillenia indica) and aiding in its dispersal, insects too play an important role within that same thread. Outenga became my favourite tree during my stay, mostly because it was fairly easy to identify and it was rather common. At this time of the year, the fruits were ripe for the picking. No wonder the elephants love them, for which they are called Elephant Apple. For an elephant, they are quite a mouthful, and an entire meal if a human is to eat it – raw or cooked.

Outenga, elephant apple, Dillenia indica, whole fruit, cross-section, and in wild elephant dung.

My fascination with Outenga increased when I learnt from the guards that it is essential for its seed to pass through the elephant’s long digestive system to increase its chances of germination. It was only natural then to seek Outenga remains in elephant dung, but my interest wasn’t solely elephants, of course. Over a long period of peering through elephant dungs along roads, we found a fresh pile, wet and warm.

A burrow dug by the large dung beetle right next to fresh wild elephant dung.

Azim ji, a guard with a wealth of knowledge about trees, helped me set to work. I wanted to see dung beetles, and to our surprise, there were very few to be seen – most had immediately buried themselves right next or underneath the pile of dung, more than a foot deep. Save one small dung beetle, the giant ones retreated. In this pile we found a piece of Outenga, but no seeds.

This made me wonder, then, is it only elephant’s intestines that help Outenga germinate? What if insects like dung beetles help accelerate the process? Dung beetles are known to roll fresh dung into balls, and then bury them after laying an egg on them, the grub then consuming the mulch. What if, among the many balls, Outenga seeds are rolled, and then buried in the soil, much like how humans sow seeds, which help them germinate better? Like I said, we don’t know what we don’t know.

Oniticellus cinctus, a very shy and nervous (that's poop) dung beetle associated with wild elephants in Kaziranga landscape.

The elephant dung attracts a range of animals to it. It almost acts as a magnet. Humans, such as I, too am fond of it. I once wrote about all the mammals that are curious about elephant dung in Central India. (For more). Here, I observed a wild boar dig through the pile of elephant dung looking for morsels – undigested seeds, like that of Outenga, but also rice and other seeds the elephants may have razed from someone’s farm – to munch on.

Two dung flies on fresh elephant dung - the very cryptic Muscid fly (top) and the ant-mimicking Sepsiid fly.

The biggest attractants are no doubt the insects. Other than the dung beetles, a host of flies home-in on fresh dung, feeding on it and laying eggs in it (It is worth noting that most of these flies, in the family Sepsidae, mimic ants). Following them come spiders that want to hunt flies. Over a short span of time, elephant dung is consumed and disintegrated and spread on the forest floor, distributing the nutrients. It will be interesting to see how this process happens and how long it takes, and how, in the larger scheme of nature, vegetative communities are formed.


In the undergrowth, the inter-species relationships are equally, if not more, complex. Just as many animals depend on megaherbivores for survival (scavenging the leftovers, whether fresh or as dung), many, many others depend on ants for survival. Indeed, one way to weigh how important insects like ants are to an ecosystem, just count the ant-associates. These are animals, mostly invertebrates, who rely on ants for survival over much of their lifecycle.

Observing ants revealed twelve types of critters that are associated with ants, either as myrmecophiles – animals that live within ant colonies, myrmecophagous – animals that prey on ants (also includes birds and mammals), and many of them are myrmecomorphs – animals that mimic ants, with some animals doing so in general because mere appearance like an ant is enough to deter predators.

A myrmecophilous (ant-loving) Atelurine silverfish - one of the common inquilinens in ant nests, such as these Technomyrmex elatior.

Of these twelve, seven were true ant-associates, they lived either as inquilines – animals that live in ant colonies commensally as tenants (without harming their landlords), such as Atelurinae silverfish, or stealers like the tiny black fly that is conveniently riding on the petiole – or saddle – of the Diacamma forager (photo further below); when the ant finds food, she will either feed on it as the ant chews it or ride the ant to the nest, this Milichiid fly is a kleptoparasite, they steal food caught by others. Then there are Batesian mimics of ants like the Dung Flies (Sepsiidae) and the Aggressive mimics where the hunter mimics the prey – here, it is usually the spiders that mimic ants, like the Synagelides jumping spider, Mallinella ant-eating ground spider (that’s literally its common name), and Orthobula sp. (tentative identification, they are too small) of a rather uncommon Trachelidae family. These are some that I could find in the ten days, there are many others, especially beetles in the family Staphylinidae, which mimic ants not just morphologically but also chemically, becoming nest-dwelling predators of ant larvae, and ant-parasitoids known to decapitate ants.

Pheidole asperata and its associates - an Anthicid beetle and a ground spider, cf. Mallinella sp.

Every time I rose from beneath the feet of the giants, I said to myself, what a world within a world. I was quite fascinated to find two ant-mimics, an ant beetle (Anthicidae) and Mallinella the ant-eating ground spider living close to Pheidole asperata ants – all of them smaller than 0.5 cm – some living off by hunting the ants, some pretending to be them to escape other ant-fearing predators.

A Trachelid spider, cf. Orthobula sp., showing unique behaviour of an adult male piggy-backing a female - this tandem-carrying behaviour may be associated with their ecology.

On another occasion, I found the Trachelid near a ruckus of Lophomyrmex ambuguus ants. When I picked the leaf it was walking on, I found it oddly shaped. It fell on my notepad, and I saw something I had not seen spiders do before, it was a spider carrying another spider – not spiderlings, mind you –another adult spider on its back. I realised it was a female with a male riding on her back, holding onto her firmly as she tried to escape my prying eyes. After sufficient observation, I let them be where they were. An arachnologist from Japan told me this behaviour is not uncommon among Trachelids, especially Orthobula, and is a part of their mating ritual where the male rides the female to ensure they mate successfully. Is it because it is difficult to find a partner at the feet of the giants or is there fierce competition, but this was the only time I found them, leaving the mystery unsolved.


If we connect the dots between ants and elephants, we end up creating a maze of connections and interconnections and the threads would extend well beyond where they started. Such is the complexity of ecosystems, symbolised by the concept of ants to elephants. That’s obvious, but it begs the question: is it possible to talk about insects, say ants, the way we talk about animals, like elephants? Yes, it is, but in terms of outreach, whether in writing or audio-visual, scientific or popular, the scale is so far biased towards elephants.

A Milichiidae fly, commonly called free-loader fly, piggy-backing a Diacamma worker, the moment the ant finds food, the fly will feed off it.

After counting insects, I was pressed with a question that haunted me during much of my stay. It is uncomfortable to address, but in the context of the concept, it is worth mentioning. The survey, temporarily and spatially small though it was, made one thing clear, that ants and termites are abundant in Kaziranga landscape. I’m confident in saying they’re everywhere (34 of the 45 had ants and/or termites, with an average of 3.6 species of ants per survey effort of nine). While this is too small a sample to make such a judgement, it is to be stated that in Kaziranga landscape ants and termites outweigh the biomass of all wild elephants, rhinos, and water buffaloes, combined by a large margin.

In such a context, the invisibility of those at the feet of the giants becomes tilted towards these social insects – and the argument of weight-per-weight functionality in an ecosystem becomes more comparable. This, in essence, is the concept of ants to elephants. Don’t fight this thought, I’m not the only one to make such an argument, even though mine is baseless at regional scale. Scientists have long pondered over this very simple question.

“The astounding ubiquity of ants has prompted many naturalists to contemplate their exact numbers on Earth,” starts a new publication of 2022 that estimates the abundance, biomass, and distribution of ants on Earth. (For more). It presents a conservative estimate of 20 x 1015 (20 quadrillion) ants on the planet, by weight, they exceed the combined biomass of all wild birds and mammals on the planet, or about 20% of human biomass.

Two ant species showing two different food procuring strategies, Stictoponera binghamii (top) actively hunting and carrying a subterranean termite and Lophomyrmex ambiguus group-foraging on what may have been a piece of fruit dropped by a vertebrate from up a tree.

This astounding abundance has significant impacts on ecosystems which yet remain unquantified. In the context of the concept, however, it is important to note that it takes both, the invertebrates and vertebrates, for an ecosystem to function. Addressing at the opening of the invertebrate exhibit at the National Zoological Park (now Smithsonian National Zoological Park) on May 7, 1987 – 98 years after the park opened – E. O. Wilson said, “It is a common misconception that vertebrates are the movers and shakers of the world, tearing the vegetation down, cutting paths through the forest, and consuming most of the energy. That may be true in a few ecosystems such as the grasslands of Africa with their great herds of herbivorous animals … But it is otherwise more nearly true in most parts of the world of the invertebrates rather than the nonhuman vertebrates.” (For more).

The heirs of the lord of the grassland practicing combat as the sun sets over Kaziranga.
Lord of the Grassland is the title of the book based in Kaziranga, written by Nirmal Ghosh, for more details, see.

There are many moments in the wild a naturalist admires – a herd of wildebeest on their annual migrations, a pack of wolves along the Rockies, a flock of flamingos in the mudflats, a school of fish in the Great Barrier Reef, a playful fight between teenage elephants as the sun sets over the mountains, what have you – rarely a few are dedicated to invertebrates.

A pair of Technomyrmex elaitor pass each other on the way to forage.

Amidst ticks and leeches, both annoying little invertebrates, I took pleasure in getting to know the most common ant of Kaziranga landscape, the pale-footed ant, Technomyrmex elaitor – as common in the woodlands as they are around kitchens, tying the remote wilderness with local, nature-conscious human habitations.

The nasutes stand guard as a column of foragers emerge from their abode.

Or the brave nasutes standing guard as a column of workers emerge from their tree-cavity abode to forage as the sun shines the brightest, the termites themselves blind to all the beautiful colourful world they are a part of. I cannot put a finger on any one moment, but that is the essence of the concept of ants to elephants, it is about acknowledging the privilege of being human, to experience everything in one breath.

One may say that my agenda was to get megaherbivores in an essay on ants, that is indeed true, I had to do so to make this argument: at the feet of the giants live the little things that run the world.


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