The annotated budak

Handsome buggers

Fungus beetles can be reliably found by trails where damp suffuses the air and ample deadwood litters the forest floor, providing nourishment to the hyphae of basidiomycetous fungi, whose fruiting bodies in turn feed new generations of mycophagous insects. Two groups of beetles share this broad label: the pleasing hordes of Erotylidae and the handsome ranks of Endomychidae. Both families tend to be dull black and hairless, with reddish, yellow or orange markings, but pleasing fungus beetles are generally more streamlined in profile, with a convex body bearing striking zig-zag lines on the elytra. The most commonly encountered Endomychids are rather less elongate, even squat, and display variable sprinklings of spots or specks on their sclerotised forewings, which may also have broad margins that partially conceal the hindlegs.

The beetles probably form a chain of aposematism, using their bold patterns, and in some, a foetid smell, to warn off predators before a fatal bite. Against intrusive fingers, however, they often resort to rolling off their perch to lose their pursuers in a tangle of low vegetation. Not a few endomychids in the tribe Amphisternini also sport elytral projections that may serve a defensive function, borne as they are by both sexes. In Spathomeles, the spines may assume the shape of high tubercles or resemble the dorsal fins of sharks and whales. Another genus, Amphisternus, sprouts thick, blunt excrescences, while Cacadaemon has tall, keel-like protuberances, and in one exuberant Bornean species, a battery of spikes arising from the pronotum to the tip of the wingcase.

The possession of such armature, along with potent chemical deterrents, may have encouraged a curious beetle to place itself, late one mildewy afternoon, on a low blade by a well-trodden path. Endomychidean in build, poise and colouration, though markedly smaller than the black and yellow species that inhabits the undersides of bracket fungi, the beetle has a pronotal flap that extends into a blunt shield around the head. The anterior fringe of each elytra forms a short ridge ending in a pigmented knob. Most eye-catching of all are two rearward-pointing processes with thickened bases, which simmer with a menace ill-befitting a beast of mouldy appetites. The creature matches, with some hazardous nips and tucks, the description of either Amphisternus vomeratus or A. malaccanus, both of which have been reported from Malacca. Little more can be said about the insect, for it was a quarry unsought for on a day of dragon-hunting, and, quite possibly, a prize unreckoned in the records of a land too mean to account for such small, scurrilous losses.

Cord in the net

Pseudobiceros bedfordi, Cyrene Reef.

In October 1898, two young men from Cambridge arrived in Singapore on a year-long mission to explore the marine zoology of its neighbourhood. Eschewing the heart of the colony, where commerce thrived and coolies dropped like flies in the hovels of Chinatown, Francis Perch Bedford and William Foster Lanchester established field stations at Labrador, Changi, Raffles Lighthouse as well as Cape Rachado (now Tanjong Tuan in Malacca), where they dredged, trawled and walked, sampling vials at hand, the seabed to draft an annotated inventory of its littoral wealth.

Soft corals, reef flat of Pulau Jong.

The fruit of their labour was shipped back to England to form the basis of a regular stream of scholarly reports by the collectors themselves and their associates. In 1900, Lanchester presented, in the Proceedings of the Zoological Society of London, what he intended to be the first of a series of papers describing crustaceans encountered in Singapore and Malacca. On his part, Bedford provided, in the same volume, an overview of the echinoderms found in these two possessions. Lanchester continued to pen treatises on crabs, mantis shrimp, peanut worms and soft corals in the following decade, but his erstwhile companion did not long survive his first formal publication, succumbing to a fatal bout of malarial poisoning on 7 October 1900, aged 26. The passing of this young academic was not overlooked by fellow biologists, however, as his name was soon honoured by way of appendage to an extravagant polyclad and a widespread arrow worm.

About a year before his death, while still in the halls of the Raffles Museum, Bedford had written another article, one more suggestive than scientific, in which he offered "stray impressions" of his surveys, marvelling at the diversity of taxa snared by his nets or hidden under stones, and remarking on the striking similarities in form, features and habits between the coastal invertebrates of Singapore and those of his native land, save in the corners where corals flourish. He also gives a topographical account of the latter, which convey a vision little different from the reefs that survive to this day at Pulau Semakau and other southern islands spared the imprisonment of breakwaters.

Holothuria leucospilota with shrimp, Cyrene Reef.

"In places where the reef is present, its distance from the shore varies from a few yards to half a mile or more, and in many cases no part of the reef proper is dry at low spring-tides; the actual width of the reef itself is also very variable, but rarely exceeds about ten yards; on its outward edge it slopes somewhat abruptly to about five or six fathoms, and then more gradually seawards. Between the reef and the shore there is nearly always a flat covered with mud, and very often with an abundant growth of brown sea-weed which harbours a large fauna.

The mud flat is very nearly level, and at lowest spring-tides there is left about a foot of water in the deepest parts, the highest portions of the 'flat' being just dry. The mud sometimes extends nearly up to high-water mark, but as a rule it is separated from the land by a belt of sandy or rocky ground, or occasionally by projecting volcanic rocks excavated by the sea into hollows, which on the retreat of the water form tide-pools, and contain numerous nooks and crannies in which molluscs, crabs, and other animals find a hiding place.

Blue dragon, Pulau Jong.

Here at any rate at first sight the naturalist will readily admit that he might be on English ground. As he looked more cloely he would probably see large fleshy Alcyonarians abounding on the mudflat, and to some extent replacing our anemones, the latter being only locally common; he might also see large Holothurians basking in the sun, either stationary or crawling slowly over the mud... Hermit-crabs abound everywhere, and at night the shore would sometimes be almost covered with them; crabs and prawns shelter themselves in crevices or under stones or in the sand, and Spatangids, Chaetopods, and Gephyreans make their burrows in the sand or rocks; limpets, too, of a diminutive size it it true, but still obvious limpets, stick to the rocks with the same tenacious grip as elsewhere, and obviously fill the same place in the economy of nature; our common littoral Gasteropod genera, such as Nassa, Purpura, Littorina, Trochus, etc., are represented by forms closely similar both in form and habits, and many of the species seem to have extremely variable coloration as on our own coasts; in fact it would be difficult to name any characteristic difference.

Rock formations, Pulau Jong.

Polychaet tabes [sic] project from the surface on nearly every sand-flat, Lamellibranchs abound in the mud and bore into rocks and wooden landing-stages, Nudibranchs of brilliant colours, together with Polyclads, creep about on stones and sea-weed, and even the abundant Periophthalmus which forms so marked a feature of the littoral fauna as it bounds over the surface of the pools, or rests on some adjacent object just above the water, is after all only a goby, such as every boy-naturalist delights to hunt at home."

Bedford also managed, a few months before his demise, to dispatch a brief note, in Nature no less, on a group of animals few have since studied, much less seen, in Singapore. He was probably based on Pulau Satumu when his specimens were nabbed "just outside or over the edge of the reef surrounding the island", amid very strong tidal currents that swept plankton from the depths towards the surface. What he caught was described as larvae no more than 5 mm in length, in various life stages, including individuals with a full complement of fin-rays and ventral fin chambers. Later, he dug up a single adult from about "six fathoms of water on a bottom composed of somewhat coarse gravel-sand close to the west entrance to Singapore Harbour."

Earlier this week, just a little south of a terminal that is supplanting the new harbour of Keppel's day, a sample of similarly rough substratum from the seafloor between Semakau Landfill, Pulau Jong and Pulau Sebarok yielded another lancelet, which was initially thought to be a measly worm but demonstrated its affinities by dashing about the confines of a small container with the vigour of whitebait. Barely an inch long, the beast looks nothing more than a pale, undulating rod surrounded by chevrons of thin muscle and with unpaired flap-like fins running the length of the body. What was probably its last meal could be seen in a strip of colour on the ventral edge that ends shortly before the caudal tip.

Common in East Asia, where they have not escaped the appetites of hungry tribes, amphioxiforms have been described as "a vertebrate with almost everything subtracted, a half-sketched blueprint" (Fortey, 2011). Boneless but not entirely spineless, amphioxi resemble the young of lampreys, and at some point in deep time, a similar creature cultivated and conserved traits that would allow it to brave open waters in adulthood and break into the pelagic zone. Those Bedford bagged he suggested to be Branchiostoma cultellum or B. belcheri, two species then known to occur in the Indo-Pacific. Larvae of the latter species were later observed in Singapore to briefly join the planktonic hordes just after sunset. After the wars, one new amphioxus, B. malayana, was named from this locality, specifically the "extreme eastern end of the island" and large populations of Branchiostoma lanceolatum are said to live in the Bay of Bengal. 

Living in loose substrates in maturity, lancelets writhe through sand and mud, exposing only what sections are required to take in suspended matter and unleashing bursts of speed that foil the plans of more advanced chordates. Ed Ricketts, who stamped on packed sand to flush out the animals during expeditions in southern California, recommended that a collector "turn up a spadeful of sand, trusting to one's quickness and skill to snatch some of the animals before all of them disappear." He continued, "Until one has actually seen them in the act, it is hard to believe that any animal can burrow as rapidly as amphioxus, for the tiny, eel-like creatures are as quick as the proverbial greased lightning. Seemingly, they can burrow through packed sand as rapidly as most fish can swim." Swift in body, slim in build and subject to seasons of unfathomability, these stripped-down survivors of prehistory have somehow retained a foothold in the Singapore Strait, where life has progressed so its heirs have all but forgotten their roots and live in disharmony with the land, the sea and the undigested remains of a disquiet city .

A strand of blue

Some dragonflies, though widely distributed and tolerant of disturbed habitats, are neither particularly abundant nor uncommonly rare. Their presence, or no-show, from what would appear to be suitable localities could simply be a phenological quirk or the result of competitive forces in pools that favour naiads with fiercer appetites or faster rites. One could also miss an ode by showing up at odd hours or looking in the wrong places. Overcast days can trigger an exodus of all but the most insouciant species. Torrid seasons may drive libellulids into the shadows or sweep the insects to breezier perches high in the canopy. There are also perpetual flyers, such as banded skimmers and water monarchs, who never settle down, or do so only to tease, and aeshnids that dart out to eyeball intruders before dashing off into the deep.

Conversely, insects with a reputation for skittishness can forget themselves in the heat of the moment and line up for the taking, as a small group of common chasers did one afternoon at Bollywood Veggies after a meal of fragrant portions. Near the rear of the farm, close to a lotus pond and artificial swamp dominated by a small Sonneratia tree, the proprietors have fenced some plots with robust cords strung from sturdy posts. The ropes have in the past supported the hives of paper wasps, but what caught the eye that day was the sight of mature Potamarcha congener displaying little interest in courtship, with both sexes alighting on the twine and raising their rear ends in homage to the sun. Granted, the males seemed a little past their prime, with ragged wings and faded colours, but they were otherwise quick on the uptake, readily leapfrogging their companions to land on slightly more private stretches. Tethered to a line of their choosing, there was little one could do but stand still, squint at stray rays and be held close and captive by these stiff bodies of dull gold and blue.

Soft bottom lines

Paguroids are among the most abundant and amusing decapods in the littoral zone, where they flourish on the bleached remains of gastropod mortality. Shorn of their periostraca, the shells lose what colours they had in life, fading into a dull coat of nacre which soon acquires a fresh body of settlers: algae and sea anemones land on the whorls, while slipper limpets and anomurans with soft bottoms squirrel themselves into coils that can still support life from without. On soft expanses, tiny diogenids trundle in the shells of Umbonium, moon snails, cerithiids, batillariids, whelks and frog snails. On rocky shores, windrows of columbeliids, drills, periwinkles, planaxids and trochids are fought over by crabs who gather in ungainly piles on slippery slopes to partake of scarce resources or indulge in common appetites.

Outnumbered than the minute hordes, though less likely to be mistaken, and thus overlooked, for sluggish molluscs by casual beachcombers is a hermit crab with orange-brown longitudinal stripes on its limbs and eyestalks. Clibanarius infraspinatus serves as the archetype of its superfamily on local shores, shuffling as it does on Changi's seagrass flats, Chek Jawa's sand bars and the sinking core of Cyrene Reef. Smaller animals probably compete with their punier cousins for modest lodgings, but adults have a knack for finding the shells of noble volutes, spiral melongenas, conches, bonnets and balers, which often seem several sizes too large for the crab. Mature crustaceans expend more energy on gonads than growth, though, so there is merit to hoarding ill-fitting drags, especially when they'd withstand a good few moults and are scattered to boot. The main drawback to domestic tightwadry is probably an accumulation of dead weight: porcelain crabs, amphipods, polychaetes and slipper limpets worm their way into the lumen to hitch a ride and share the spoils of messy feeders. 

At least eight other congenerics occur in Singapore's waters, though few attain the bulk of the buff bruiser. One species of similar length but lighter build is Clibanarius longitarsus, which forages in the mangrove creeks of Pulau Semakau as well as silty habitats on St John's Island, Pasir Ris and Pulau Ubin. In life, this hermit is rather variable in colour, being "mottled brown and blue or light brown with several blue patches" and having brown claws covered with blue or blue-green tubercles and spines.

Another, rather uncommon, paguroid encountered on occasion at Beting Bronok and muddier parts of Changi Beach is Paguristes longirostris, which, despite its generic designation, is a member of the same 'left-handed' family as Clibanarius. The carapace of this hermit is, however, "nearly naked" and the outer portions of the wrist, hand and fingers are distinguished by flat, overlapping tubercles which lend the claws a "squamiform appearance". The subequal chelipeds also sport bright blue patches on the inner surface of the carpi, which may serve as signals for intraspecific recognition or signs of reproductive health. There is little to indicate if these crabs, which have been recorded from Singapore and the Bay of Bengal, enjoy a comparative advantage in areas of a certain depth or substratum, but the genus is known to produce few, relatively large eggs that hatch into larvae exhibiting abbreviated development. Such a strategy, unlike the scattershot approach of those who gamble on short bursts of fertility, hinges on the persistence of stable states and a climate of certainty, vestiges of a kinder, gentler time that no longer exists in this age of miracles in which life is chained to a perpetual horizon with neither space for exploration nor room for thought.

Trash talking

Land hermit crabs are more easily heard than seen, for these supralittoral wanderers seldom bother to hide their tracks, shuffling as they do and clicking their heels against the debris of the high shore with little regard for aural sensitivities. They abhor seafronts that have been pick over and rendered plain for the sake of visitors who regard every speck and spot on the sand as well as unruly grasses as blights on a landscape awash in green delusions. It's a shame, for these anomurans gladly labour without recompense to cleanse the waste of coastal life, including perhaps the droppings of unretentive beach apes, leaving the inorganic portions to the mercies of gloved crews.

A stern breakwater now links Lazarus Island to St John's, stifling a passage of fringing reefs abutting a rocky strand. Overlooking the shore is a ridge of Callophyllum, sea almonds, pandans and figs, whose terminal buds and tough husks litter the gravel below. Pied songbirds and sea eagles erupt from the canopy long before sunrise. Where the embankment meets the hill, there is a nook into which the tides have shoved a midden of jetsam. The debris is a magnet to hermit crabs of rare bulk, which clamber over foam and foliage and even brave the lower reaches of the seawall in their nocturnal scurry. Some lug the remains of nerites and turban snails, while not a few individuals are at home in lightweight imports. When picked up, some stretch out in a frantic grasp at damp air or even bail out, ditching their shelters for a flight of naked panic. But most just scoot into their coils, sealing the aperture with a stout claw and betraying their impatience by peering out within a minute and resuming, if the coast appears clear, their seedy patrols.

Such armies now thrive only on islands with little afoot, where food abounds and the sea laps at the base of lowland woods. Where these conditions are met, hermit crabs can rival their landlocked cousins in numbers and vie with earthbound arthropods for choice spots on low trees. Large concentrations of small coenobites can be found thus in the littoral forest of Tangkoko Duasaudara Nature Reserve in North Sulawesi, where a thin corner of the peninsula has been preserved for the natives of a fiery region. Fuelled perhaps by the ash of nearby volcanos, Terminalia, Barringtonia and Morinda attain monumental proportions in a strip of land between a stained beach and volatile slopes.

The darkened soil is coated by layers of bark, leaves and fruit, which support troops of black crested macaques as well as shy terrestrial crabs that forage for vegetation and carrion. Coenobita occurs here in disturbing densities, forcing those who wander off the path to watch their step lest heavy boots deliver a crack of doom. A few are so well-built they tread on the ground with little fear, but their undersized compatriots often cling en masse to sturdy saplings, creating the impression of a house plant infected by festive baubles. Others grip the trunks of more mature trees or dawdle near recesses that may serve as daytime retreats. A partial diet of scale insects is said to drive this rhythm, but others have caught no rhyme to this restless spirit, which has led to accusations of truancy that bounce off the hermits as they laze on the stems of tropical lianas or wing it on dangling vines. Encouraged by the onset of indiscreet hours, a few engage in conversations that waver between conflict and resolution, their antennules twitching with trepidation as fingers touch and feelings are exchanged via olfactory clues. But the signals are mixed and any semblance of pillow talk in this cinq à sept is soon quashed by the awkwardness of a tryst that spells surrender to the moon and spawns a rush to the sea for a moment of release in the dip end.

Looking with luck

Big bad things with a reputation for pointy ends, like sharks, stingrays and sea urchins, are hard to miss on the reef. A menacing fin, a splash of panic and black spineless orbs: these hazards of low water draw attention to themselves with furious dashes that rouse the surf or displays of armature that grip the mind and disarm the senses. The search for smaller majorities, those creatures that shun the limelight and hog the shadows, who retreat with pride or wield frames that fool their foes, on the other hand, demands a willingness to stop without cause, stoop at length and stifle the urge to move on to fresher fields. 

There are slugs that are barely distinguishable from their perch and nearly impossible to detect as they prowl about their prey, clad in a cloak of cerata that ape the colours and curvature of brown seaweed, green algae, firm sponges and soft polyps. There are crabs that cling to floating debris, lug hollow rubble, or co-opt sessile reefmates into service as shields, as well as snails that rock, fish that lie flat, and horses that hide on spongy trees. Luck, or the learned ability to tease apart living forms from the lay of the land below the waves, are the only guides to survival on dark shores, where there is no telling what lies a few steps ahead and time is running out and losing wind.  

It is often the case that a modest encounter – a hurtling carpet eel blenny, a tumbling xanthid, a hunting octopus – leads to punier discoveries. The animal plunges into a stumbling block and leaves its pursuer on the haunches or in thrall to fatigue, with no choice but to scan the immediate perimeter for consolation prizes. It is during such moments of exigency that resolution strikes and the sea reveals the secrets of its barest bottoms. In the cornea of one eye, a sliver of green paddles about a shallow basin of seagrass, scooting from blade to blade, settling just long enough to change its mind before another whirl of pleopods in search of right angles. Boasting a snout to be proud of and shaded to match the vascular halophytes of Cyrene Reef, the shrimp would have been invisible if it had not dissolved into a ruckus of motion, as if it had imbibed the witching hour that infects domestic felines every night, causing victims to bolt for imagined glories and show caution a digit of frippery. The caridean is probably a species of Latreutes, though one much larger than a squat, brown thing found close-by over a year ago. These hippolytids are usually associated with specific habitats, with some riding on jellyfish and others huddling amid gorgonians or kelp. Not a few, such as Latreutes pymoeus and L. mucronatus, dwell in seagrass meadows or algae beds, but information on their habits, density and sensitivities is as slim as their tapered flanks. Our surveys are alas too brief and easily distracted by other little beasts to determine if these decapods occur in dispersed enclaves or as disguised hordes, and there have been few efforts to break the back of this duck and prise open the spark of their capers before the fever ebbs and the riddler slips, in the bite of a blink, behind a fold of mysteries in a bid to lose interest and fade to blank.