Field of Science

Into the Labyrinth

Climbing perch Anabas testudineus emerging from water, as illustrated by Richard Lydekker.

Amongst the unholy mess that is the Percomorpha, one group that has long been recognised is the labyrinth fishes of the Anabantoidei. The anabantoids are a group of freshwater fishes found in southern Asia and Africa (but not Madagascar) that get their vernacular name from their possession of a distinctive respiratory organ called the labyrinth. This organ, found in a cavity above the gills, is derived from part of the first gill arch; the bone has become expanded and much-folded, and is covered with a layer of respiratory epithelium. So long as the gills do not actually dry out, the labyrinth allows these fish to take in oxygen directly from the air, and they can survive in warm, low-oxygen waters. They can even survive for limited periods entirely out of water (a feature that has helped make some of the larger species popular food fish, due to the greater ease of keeping them fresh in a tropical environment). Recent phylogenetic studies (e.g. Li et al. 2009) have agreed in placing labyrinth fishes as related to a number of other freshwater Indo-Australian fishes, such as the snakeheads of the Channidae and the swamp eels of the Synbranchidae, many of which are also tolerant of air-breathing.

Kissing gouramis Helostoma temminckii, from Peter Bus.

Labyrinth fishes can be divided between three families (Rüber et al. 2006). One of these contains a single species, the kissing gourami Helostoma temminckii of south-east Asia. Kissing gouramis are primarily specialised filter feeders, though they may also graze on algae or insects. The vernacular name refers to their enlarged lips, making them look permanently puckered up. Kissing gouramis even 'kiss', pressing their smackers against one another, though this is regarded as an act not of affection but of aggression (kind of like a 1930s Hollywood melodrama) as the fish push against one another.

A rather unfortunate Cape kurper Sandelia capensis, photographed by Darryl Lampert.

The climbing perches of the Anabantidae include the south Asian Anabas and the African Ctenopominae. These short-bodied carnivores have serrated edges to their gill covers that the Asian species use to pull themselves over land when travelling between water bodies (imagine lying on your stomach and pulling yourself along with your chin). You can see video of some climbing perch Anabas testudineus emerging from water here.

Giant gourami Osphronemus goramy, photographed by E. Naus.

The most diverse subgroup of the Anabantoidei is the gouramis of the Osphronemidae, another south Asian group. The largest of the Osphronemidae, the giant gourami Osphronemus goramy, grows up to 70 cm, but most species are quite a bit smaller. A number of gourami species (as well as the kissing gourami) are popular aquarium fishes; the most popular by far is the Siamese fighting fish Betta splendens, males of which have been bred to exhibit much longer and more ornamental fins than found in the wild. The gouramis are generally omnivorous, with species varying in the extent to which they prefer plant or animal food. The most specialised carnivore of the Osphronemidae is the pikehead Luciocephalus pulcher, a small but elongate species that has been described as having the most protrusible mouth of any fish (and that, by the way, is no small claim). You can see the pikehead in action below:

The pikehead is so divergent from other labyrinth fishes that past authors have regarded it as its own family, possibly the sister taxon to all other anabantoids, or even questioned whether it was a labyrinth fish at all. However, as confirmed by Rüber et al. (2006), Luciocephalus is not only a true anabantoid but nested well within the Osphronemidae as sister to the chocolate gouramis of the genus Sphaerichthys. These and two other genera, Ctenops and Parasphaerichthys, form what is known as the 'spiral egg' clade, named after the presence of spiraling ridges on the egg leading to the micropyle, that have been suggested to act as guides for the sperm.

Siamese fighting fish Betta splendens mating below a bubble-nest, photographed by Stephen & John Downer.

The anabantoids are also known for the bubble-nests constructed by a number of species, in which the eggs are contained within a floating nest of bubbles that is guarded by the male parent (both parents in the Ceylonese combtail Belontia signata). Bubble-nesting has evolved at least twice among the anabantoids: once in the Osphronemidae, and once in the ctenopomine genus Microctenopoma (other anabantids and Helostoma are free spawners that do not construct nests or guard their eggs; the ctenopomine Sandelia capensis digs a nest in the bottom substrate) (Rüber et al. 2006). Though bubble-nesting is probably the ancestral behaviour for Osphronemidae, it has been modified in a number of sublineages. Osphronemus species build submerged nests from vegetation, while members of the 'spiral egg' clade (except Parasphaerichthys) and a number of Betta species are mouthbrooders. Usually the male broods the fry in these species, but the female is the brooder in a couple of Sphaerichthys species.


Li, B. A. Dettaï, C. Cruaud, A. Couloux, M. Desoutter-Meniger & G. Lecointre. 2009. RNF213, a new nuclear marker for acanthomorph phylogeny. Molecular Phylogenetics and Evolution 50: 345-363.

Rüber, L., R. Britz & R. Zardoya. 2006. Molecular phylogenetics and evolutionary diversification of labyrinth fishes (Perciformes: Anabantoidei). Systematic Biology 55 (3): 374-397.

The Kellyclams

Ruddy lasaeas Lasaea adansoni photographed in a rock crevice by David Fenwick.

Members of the family Lasaeidae, commonly known as kellyclams, are small thin-shelled bivalves that often live in close association with larger invertebrates such as crustaceans, worms or cnidarians. The clam may be directly attached to its host or share a burrow with it; one genus, Entovalva, includes associates of sea cucumbers that live within their host's esophagus (you can find some more details of this particular relationship here). Though the clam's presence may not be entirely without physical effect on its host, such effects are usually minor and kellyclams are generally regarded as commensals rather than parasites. Other species live independently and may live nestled among rocks or buried in sediment; these free-living forms may possess a relatively large muscular foot for mobility. Like most bivalves, kellyclams are filter feeders; the invertebrate-commensal species are believed to take advantage of water currents created by the host to increase the effectiveness of their own feeding currents.

Commensal Pseudopythina rugifera attached to a ghost shrimp Upogebia pugettensis, photographed by Arthur Anker.

The family-level classification of the kellyclams has been fairly turbulent. The family has variously been known as the Lasaeidae, Erycinidae or Leptonidae, owing to confusion over which of these names has priority, while some authors have regarded them as separate families and/or recognised further segregate families Kelliidae or Montacutidae. With their small size, kellyclams have simplified a number of the characters used in classifying other bivalves, and a number of the commensal species have become modified in order to co-exist with their host. The Peregrinamor species, for instance, live attached longitudinally underneath the thorax of the ghost shrimp Upogebia; they have accordingly become low and elongate, and were until recently misclassified as mussels of the Mytilidae (note that the Lasaeidae and Mytilidae represent evolutionary lineages that first diverged some time in the Ordovician). In the broad sense, the Lasaeidae have been separated from the closely related family Galeommatidae by the fact that members of the latter have the soft body enlarged so that the shell becomes internal. However, Goto et al. (2012) established that even this distinction is not reliable, with the Galeommatidae nested and probably polyphyletic within the Lasaeidae. Also, while Goto et al. did not support recognition of the Lasaeidae as a holophyletic group, nor did they support any of the segregate families. Commensal species are scattered phylogenetically among free-living species, and host-switching has apparently happened numerous times within commensal lineages.

The Lasaeidae are hermaphrodites, and often brooders. Rather than being released as eggs, young are retained within their parent until they are released as veligers (later-stage larvae that have begun to develop a shell) or as miniature versions of the adults.


Goto, R., A. Kawakita, H. Ishikawa, Y. Hamamura & M. Kato. 2012. Molecular phylogeny of the bivalve superfamily Galeommatoidea (Heterodonta, Veneroida) reveals dynamic evolution of symbiotic lifestyle and interphylum host switching. BMC Evolutionary Biology 12: 172.

"Anatomy of Mollusca": A Case of Plagiarism

Whilst researching material for an upcoming post, I came across this book on Google Books, published in 2010 by the International Scientific Publishing Agency, New Delhi:
Anatomy of Mollusca, by Rita Rawat

Looking at the section previewed through Google Books, I couldn't help feeling that it seemed a little... familiar. Take a look at this screenshot from Google Books:

Now take a look at this screenshot, taking from a page written by my erstwhile associates at, last modified in 2006:

I can only go by what was available in the Google Books preview, of course, without direct access to the actual book, but I can confirm that most if not all of the material in pages 15 to 72 at least of the "Anatomy of Mollusca" appears to have been copied directly from (unfortunately, a large part of is currently offline as the site gets revamped). A quick Google search failed to uncover anything indicating whether the International Scientific Publishing Agency has a reputation for publishing lifted material, so I can't say if this is part of a broader issue.

Amazon currently has the "Anatomy of Mollusca" on sale for just over $75. Not that expensive by the standards of technical publications, but a fair chunk of change by the standards of books in general. Certainly a lot more than the cost of reading the same stuff on, which carries no extra cost beyond that of the ISP charges.

Dung Beetles

Flat-headed dung beetles Pachylomerus femoralis with a ball of the good stuff, photographed by Guido Coza.

The dung beetles of the Scarabaeini include 146 species found in Africa and Asia, classified by Forgie et al. (2006) into three genera: Pachysoma, Pachylomerus and Scarabaeus, with the last including the vast majority of species. The Scarabaeus species are perhaps the most famous of all dung beetles, renowned since ancient history when Egyptians saw a dung beetle rolling a ball of dung along the ground as a metaphor for the movement of the sun through the heavens*. Dung beetles collect their turd balls to use as food for themselves or for their larvae. Ball-rolling is not unique to the Scarabaeini as a method of transporting dung, however (it is also done by members of other dung beetle tribes), nor do all Scarabaeini species engage in ball-rolling.

*It probably does not say much for the standard of ancient Egyptian public sanitation that they were apparently so willing to believe that the ultimate source of all life on the planet was a giant mass of burning poop.

Flightless orange dung beetle Pachysoma denticolle, photographed by Alex Dreyer.

The flightless dung beetles of the genus Pachysoma, for instance, transport their food by dragging it along between their hind legs. Pachysoma species are also less choosy than other Scarabaeini, feeding not just on dung but all manner of organic detritus. They have specialisations allowing them to feed on drier food particles than other Scarabaeini, suitable for their arid habitats in southern Africa. In contrast, species of the subgenus Sceliages within Scarabaeus are the epicures of the scarabaein world: they feed entirely on dead millipedes, which they push along in front of themselves bulldozer-style (Forgie et al. 2005). Relatively few species of Scarabaeini feed by burrowing directly alongside piles of dung where they lay, but this may be done by Pachylomerus and Scarabaeus galenus (both of which may also transport food).

Individual of Sceliages transporting a millipede, photographed by Shaun Forgie.

Most Scarabaeini are active during the day, but a small number such as Scarabaeus satyrus are nocturnal in habit. In the phylogenetic analyses conducted by Forgie et al. (2005), these nocturnal species usually formed a single clade. Had the ancient Egyptians observed the nocturnal dung beetles as well, they could have presented us with a sky full of poo at all hours.


Forgie, S. A., U. Kryger, P. Bloomer & C. H. Scholtz. 2006. Evolutionary relationships among the Scarabaeini (Coleoptera: Scarabaeidae) based on combined molecular and morphological data. Molecular Phylogenetics and Evolution 40: 662-678.

Forgie, S. A., T. K. Philips & C. H. Scholtz. 2005. Evolution of the Scarabaeini (Scarabaeidae: Scarabaeinae). Systematic Entomology 30: 60-96.

Snap! goes the Termite

The snapping termite Cavitermes tuberosus, from Wiki Termes.

For the subject of today's post, I drew the termite subfamily Termitinae. Termites are extraordinary animals: socially complex, ecologically vital, dietically remarkable. Personally, I'm rather found of these communal cockroaches.

Termites of the family Termitidae (commonly referred to as the 'higher termites') differ from other, 'lower' termites in the nature of their gut biota (without which they would not be able to digest their cellulose diets): instead of having flagellated protozoa in their gut, termitids carry symbiotic bacteria. This difference in symbionts is reflected by a difference in diet. Higher termites feed on more decayed wood or plant matter than lower termites; some higher termites feed directly on organic-rich soil that contains little or no plant material (Inward et al. 2007). Subfamilies within the Termitidae are also distinguished on the basis of their gut anatomy: members of the Termitinae have what is called a 'mixed segment' on the outer edge of their intestine (Lo & Eggleton 2011). In the mixed segment, instead of the division between the mesenteron (the middle section of the intestine) and the proctodaeum (the posterior section) being simple and straight across, the mesenteron wall extends backwards along one side of the gut only; it has been suggested that the mixed segment functions to pump alkaline fluids into the gut, maintaining appropriate pH and fluid levels for the symbiotic bacteria in the hindgut (Bignell et al. 1983).

Workers of Amitermes dentatus repairing a damaged nest, from here.

The Termitinae have also been distinguished on the basis of the morphology of their soldiers, with most genera having soldiers with elongate mandibles that have relatively few large teeth. These are used to bite and slash at threats to the colony. However, phylogenetic analyses have contradicted this distinction (Inward et al. 2007). The Termitinae are paraphyletic with regard to the Nasutitermitinae, who have developed a very different method of defense: the mandibles are reduced, and instead the front of the head is drawn out into an elongate 'nose'. At the end of the 'nose' is a glandular opening from which the soldiers squirt a sticky glue at their opponents. Also nested within the Termitinae are the Syntermitinae whose soldiers combine both methods of defense: they retain sickle-shaped mandibles that are used to pierce the cuticle of attackers while the protruded glandular opening is used to apply toxic secretions. Chemical defenses are also not unknown among more standard termitines: soldiers of Globitermes sulphureus were dubbed 'walking bombs' by E. O. Wilson due to their explosive (and often self-destructive) discharge of toxic chemicals from hypertrophied labial gland reservoirs in the abdomen. It should also be noted that a small number of termitines do not produce soldiers at all: they may live in association with other soldier-producing termites, like the Australian Invasitermes, or they may feed on low-nutrient soils (presumably making the maintenance of a soldier caste too nutritionally expensive), like the Indomalayan genera Protohamitermes and Orientotermes.

The mushroom-like mound of Cubitermes, a major soil-feeding genus in Africa, photographed by Marco Schmidt.

Another mode of defense that is found only among the termitines (though phylogenetic analysis indicates that it has evolved multiple times) is the production of soldiers with elongate snapping mandibles. In these termites, soldiers store kinetic energy through muscular deformation of the mandibles, allowing them to be suddenly closed with great force (Prestwich 1984). So great is the force involved, in fact, that it seems to be not uncommon for the jaws to become completely crossed over as has happened to the individual at the top of this post. Snapping termites generally live in subterranean colonies, and even after the soldier has been 'spent' on the discharge of its mandibles, its body acts as a physical barrier in the confined tunnel. In some snapping termites, the mandibles are strongly asymmetrical, so the force of the closure is channelled through the left mandible only with doubled force. Asymmetrical snappers of the genus Neocapritermes, in fact, are able to knock out fairly large ants with a single blow. The video below shows a soldier of Planicapritermes attacking an ant: Or you can see Neocapritermes in action in this video. Keep a close eye on the screen around the 20-second mark...


Bignell, D. E., H. Oskarsson, J. M. Anderson & P. Ineson. 1983. Structure, microbial associations and function of the so-called "mixed segment" of the gut in two soil-feeding termites, Procubitermes aburiensis and Cubitermes severus (Termitidae, Termitinae). Journal of Zoology 201: 445-480.

Inward, D. J. G., A. P. Vogler & P. Eggleton. 2007. A comprehensive phylogenetic analysis of termites (Isoptera) illuminates key aspects of their evolutionary biology. Molecular Phylogenetics and Evolution 44: 953-967.

Lo, N., & P. Eggleton. 2011. Termite phylogenetics and co-cladogenesis with symbionts. In: Bignell, D. E., et al. (eds) Biology of Termites: a modern synthesis pp. 27-50. Springer.

Prestwich, G. D. 1984. Defense mechanisms of termites. Annual Review of Entomology 29: 201-232.

Nosybelba: A Uniquely Madagascan Mite

Dorsal and ventral views of the main body of Nosybelba oppiana, from Mahunka (1994).

Why yes, it's another random oribatid! Nosybelba oppiana was described from Madagascar by Sándor Mahunka in 1994; Mahunka regarded it as distinct enough from other oribatids that he placed it in its own monospecific family. To date, the original description appears to be the sum total of our knowledge of Nosybelba oppiana. Subías et al. (2012) transferred it to a separate subfamily within the larger family Oppiidae, and transferred a second Madagascan species 'Oppia spinipes' Balogh 1964 to Nosybelba, but this was in the context of a species checklist only without supporting discussion (also, the name Oppia spinipes was used for an oribatid species by Banks in 1906, so whatever the status of Balogh's species it needs a new name).

Leg I of Nosybelba oppiana, from Mahunka (1994). Femur and genu of the right, tibia and tarsus on the left.

So what can we tell about Nosybelba from its description? One of the first things that attracts attention is that it has rather weird legs. The tarsi (the terminal segments) of the legs are really short, shorter on all legs than the adjoining tibia. On the first pair of legs, the tarsus is also compressed longitudinally, and a dorsal process on the tibia (that bears a large sensory seta) overhangs the tarsus. To my admittedly uneducated eyes, the overall structure does not give an impression of mobility. I'm guessing that Nosybelba is not the most agile of oribatids. At the end of each leg is a single large claw; as mentioned in a previous post, the number of claws on an oribatid's legs tends to correlate with habitat, with single claws suggesting a terrestrial lifestyle.

Lateral view of front end of Nosybelba oppiana (minus legs), from Mahunka (1994).

Another noteworthy feature of Nosybelba can be found in its mouthparts. The mentum, the 'under-head' shelf that underlies the chelicerae, does not have a basal articulation, so the chelicerae are limited in their range of movement. The chelicerae themselves do not have any teeth, so Nosybelba is not feeding on anything that requires a great deal of processing before swallowing. In another oribatid family, the Suctobelbidae, similar chelicerae are related to a diet of plant matter that is in an advanced state of decay; Nosybelba is presumably also a connoiseur of the rotten and the liquefied.


Banks, N. 1906. New Oribatidae from the United States. Proceedings of the Academy of Natural Sciences of Philadelphia 58 (3): 490-500.

Mahunka, S. 1994. Oribatids from Madagascar II. (Acari: Oribatida). Revue Suisse de Zoologie 101 (1): 47-88.

Subías, L S., U. Ya. Shtanchaeva & A. Arillo. 2012. Listado de los ácaros oribátidos (Acariformes, Oribatida) de las diferentes regiones biogeográficas del mundo. Monografías electrónicas S.E.A. 4.