Field of Science


Over the years, I've put up several posts about the diversity of oribatid mites. It's time for another one.

Scheloribates laevigatus, copyright R. Penttinen.

One of the largest genera of oribatids out there is the genus Scheloribates, for which well over 200 species have been described. Their distribution is pretty much worldwide; they are found in a range of microhabitats, such as in leaf litter, in pastures or marshes, or among rocks. Distinguishing features of the genus from other oribatids include well-developed, immobile pteromorphs, tridactylous (three-clawed) legs, and a notogaster with ten pairs of setae and three pairs of sacculi (little sac-shaped glandular openings) (Ermilov & Anichkin 2014).

Considering their abundance in soil habitats, Scheloribates probably have a significant role to play in decomposition and nutrient cycling. Studies on the diet of one of the better-known species, S. laevigatus, have found that it will eat almost any type of vegetable or fungal matter, though its preferred diet is microscopic algae (Hubert et al. 1999). Indeed, they are most abundant in damper habitats that would provide good conditions for the growth of such algae.

Scheloribates species may impact on human lives in other ways too. They are an intermediate host for the larvae of anoplocephalid tapeworms that infect livestock when the mites are accidentally ingested during grazing. S. laevigatus is a known host for at least eight tapeworm species in North America. Rates of tapeworm infestation in Scheloribates can be quite high; over 60% of the individuals of one species at a particular locality in Australia were infected (Lee & Pajak 1990). Scheloribates species are also noteworthy as a likely source of the toxic alkaloids found in the skin of arrow-poison frogs. The alkaloids are likely to be synthesised by the mites (as suggested by their presence in adults but not in juveniles, despite no known difference in diet between the two life stages) and then sequestered by the frogs after they eat the mites (Saporito et al. 2011). And if they eat enough mites, they end up becoming dangerous even to something the size of a human.


Ermilov, S. G., & A. E. Anichkin. 2014. A new species of Scheloribates (Scheloribates) from Vietnam, with notes on taxonomic status of some taxa in Scheloribatidae (Acari, Oribatida). International Journal of Acarology 40 (1): 109–116.

Robert, J., V. Šostr & J. Smrž. 1999. Feeding of the oribatid mite Scheloribates laevigatus (Acari: Oribatida) in laboratory experiments. Pedobiologia 43: 328–339.

Lee, D. C., & G. A. Pajak. 1990. Scheloribates Berlese and Megascheloribates gen. nov. from southeastern Australia, with comments on Scheloribatidae (Acarida: Cryptostigmata: Oriopodoidea). Invertebrate Taxonomy 4: 205–246.

Saporito, R. A., R. A. Norton, N. R. Andriamaharavo, H. M. Garraffo & T. F. Spande. 2011. Alkaloids in the mite Scheloribates laevigatus: further alkaloids common to oribatid mites and poison frogs. Journal of Chemical Ecology 37: 213–218.

A Parasitic Eel?

The following post was inspired by an e-mail that I was sent recently by Sebastian Marquez. He told me about a friend of his catching a trevally when fishing, then cutting it open to find a snake eel inside the body cavity (but outside the stomach), wrapped around the trevally's internal organs. According to Sebastian, the lead suspicion for what had happened was that the eel had somehow burst out of the trevally's stomach before it was caught, and he wanted to know if I'd ever heard of anything similar. I didn't have an explanation for him, but his story did get me thinking about the snub-nosed eel.

Snub-nosed eel Simenchelys parasitica, from Jordan (1907).

The snub-nose eel Simenchelys parasitica is a small deep-sea eel, about 20 to 35 centimetres long. It has attracted note by being found a number of times burrowed into the body cavity of larger fishes with perhaps the most renowned case being two juveniles that were found nested inside the heart of a mako shark. This lead to the description of S. parasitica as an endoparasite (hence the species name). However, acceptance of this tag has been far from universal. The snub-nosed eel has been caught free-living more regularly than it has been found in other fish and because of its deep-sea habitat it has never been observed in life. An alternative suggestion has been that Simenchelys is normally a scavenger; because many of its recorded 'hosts' have been collected through non-targeted methods such as trawls, it is not impossible that the snub-nosed eels may have burrowed into their body cavity after they were already deceased.

It was with this conundrum in mind that the cranial anatomy of the snub-nosed eel was described by Eagderi et al. (2016). The jaws of Simenchelys are relatively short and muscular (hence its 'snub nose'). It also has teeth arranged in such a way that they form an even cutting edge (in contrast to the more spaced and uneven teeth of other eels). Eadgeri et al. came to the conclusion that the snub-nosed eel probably feeds by biting out plugs of flesh, in a similar manner to a cookie-cutter shark. Simenchelys also resembles a cookie-cutter in having large, fleshy lips that are probably used to form a seal between jaws and food source. A large hyoid ('tongue') apparatus probably works to provide suction to maintain the seal. The snub-nosed eel may also rotate while biting, a behaviour known from both cookie-cutters and other eels.

So is Simenchelys a parasite? It is probably not a habitual endoparasite, lacking as it does any clear adaptations to the endoparasitic lifestyle. There are fish that could be described as ectoparasites, in that they habitually feed on live animals larger than themselves in a manner that does not normally lead to the host's death. The cookie-cutter is one such fish; another is the candiru Vandellia cirrhosa, a small freshwater catfish from the Amazon basin that feeds on blood from the gills of other fish. It is possible that the snub-nosed eel could have a similar lifestyle to one of these. However, recorded evidence of its habits is even more consistent with scavengers such as hagfish and the candiru-açu Cetopsis candiru (another South American catfish) that tear flesh from the submerged bodies of dead animals, and may often burrow their way into the corpse's body cavity as they do so.

Of course, the two modes of feeding are not mutually exclusive. The only difference between predator and parasite in this scenario is whether the attacked animal is alive or dead, and the thing about flesh-feeders is that they're not always picky. A habitual scavenger may easily choose the opportunity to take a nibble from a still-living host, especially is said host is in some way incapacited (as a result of being swept up by a trawl, for instance). The snub-nosed eel may not be a habitual parasite, but it may be an opportunistic one.


Eagderi, S., J. Christiaens, M. Boone, P. Jacobs & D. Adriaens. 2016. Functional morphology of the feeding apparatus in Simenchelys parasitica (Simenchelyinae: Synaphobranchidae), an alleged parasitic eel. Copeia 104 (2): 421–439.

Of Shrimp Plants and Bear's Breeches

For today's semi-random post, I drew the plant subfamily Acanthoideae. As recognised by Scotland & Vollesen (2000), the Acanthoideae is the largest of the subfamilies of the Acanthaceae by a considerable margin, including about 95% of the family's 2500+ species. Though perhaps not hugely familiar to readers in more temperate climes, the Acanthoideae are one of the dominant groups of herbs and shrubs in tropical parts of the world.

Golden shrimp plant Pachystachys lutea, copyright Dryas.

The Acanthoideae have been recognised as a morphological group since the late 1800s and their integrity has been confirmed by more recent molecular studies. They are distinguished from related plants (within the Lamiales, the order that also includes such plants as the mints and snapdragons) by having capsular fruits that dehisce explosively when mature to scatter their seeds. The seeds are attached within the capsule by hook-shaped stalks called retinacula that presumably play a role in determining how the seeds are released. A classification of Acanthaceae published in 1965 by Bremekamp restricted the family to species with explosive fruits and retinacula, dividing them between two subfamilies, the Acanthoideae and Ruellioideae, based on the absence or presence, respectively, of cystoliths. These are outgrowths of the epidermal cell walls that are impregnated with calcium carbonate. They are visible in the stems and leaves, at least in dried specimens, as hard white streaks. As phylogenetic studies have supported division of Acanthoideae in the broad sense between a cystolith-possessing and a cystolith-lacking clade, the decision whether to recognise 'Ruellioideae' as a separate subfamily comes down to a ranking choice only. At lower levels, the classification of Acanthoideae is less straightforward. Over two hundred genera of Acanthoideae are recognised but just three of those—Justicia, Strobilanthes and Ruellia—account for about half the total number of species. Each of these mega-genera is morphologically diverse and likely to be para- or polyphyletic with regard to related taxa, raising the distinct likelihood of future revisions.

Spiny bear's breeches Acanthus spinosus, copyright Magnus Manske.

Economically, few of the Acanthoideae are of great significance except for a number of species being grown ornamentally. One such species is Acanthus mollis, which goes by the vernacular name of 'bear's breeches' (why, I have absolutely no idea). Acanthus was a popular decorative motif in classical Greece and forms the basis for the design of Corinthian columns. Its use as an ornamental has lead to it becoming regarded as an invasive weed in some regions, largely because this is one of those garden plants that Just Will Not Die, spreading easily from seeds and tubers. We've got some in a pot outside that is currently flourishing despite having been burnt down to a nub by the searing Perth summer sun, metaphorically shouting its defiance at an uncaring world.


Scotland, R. W., & K. Vollesen. 2000. Classification of Acanthaceae. Kew Bulletin 55 (3): 513–589.

Magnificent Eurhins

Eurhinus festivus(?), copyright Andreas Kay.

Weevils are one of the most incredibly diverse of beetle groups, coming in an incredible array of shapes and structures, but they are not usually renowned for their bright colours. Nevertheless, in a group of this size, there is always scope for surprise: witness the image above. Eurhinus is a genus of absolutely stunning metallic-coloured weevils native to Central and South America; one species, E. magnificus, was first recorded in Florida in 2002 and has since become established there. The photo above was identified on Flickr as E. magnificus but looking over the descriptions in Casey (1922) I suspect it is more likely to be the closely related E. festivus. Eurhinus magnificus differs in having patches of red on the pronotum and elytral humeri (the 'shoulders'); see photos here, for instance.

Eurhinus species feed on vines of the Vitaceae, the grape family. Eggs are inserted into young stems where the larvae cause distinct galls as they develop. It does not look like they are known to cause significant damage to economically important species though studies on whether it can successfully attack grapes are inconclusive.


Casey, T. L. 1922. Studies in the rhynchophorous subfamily Barinae of the Brazilian fauna. Memoirs on the Coleoptera 10: 1-520.

Finches in Drag

Green-headed tanager Tangara seledon, copyright Dario Sanches.

In many parts of tropical South America, it is common to see small flocks of brightly coloured small birds foraging among vegetation, plucking off berries or hunting for insects. In many cases, these flocks may contain individuals of multiple or even several species. These are the tanagers, one of the Neotropical region's most characteristic bird families.

Tanagers are members of the bird clade known as the nine-primaried songbirds (so-called because their wings have nine functional primary feathers rather than the ten of other songbirds) that also includes the finches, buntings and cardinals. The largest genus of tanagers, and indeed one of the larger genera of birds in general, is Tangara. This genus includes about fifty species found in various parts of the neotropics. In their overall structure, they are fairly uniform: small, sturdy birds with a stout, moderate-length bill and an average-length tail (Hilty 2011). In other words, they have a fairly unremarkable, finchy-type appearance. In colour and patterning, however, they are considerably more varied, to the extent that I am at a loss to know where to begin. There are species of a rich, deep blue and of a bright, emerald green. There are species with bold, contrasting patterns of blues, blacks, greens or golds; there are species of a solid, uniform brilliance. There are species with caps or chests of orange or black. There are even a few, such as the plain-coloured tanager Tangara inornata, that eschew the gaudy pigments of their congeners entirely in favour of more restrained patterns of greys and beiges. In many species, males and females show little or no difference in appearance; however, in the black-capped group (including species such as the black-capped tanager T. heinei), the males have contrasting patterns of black and blue or yellow whereas the females are largely green and grey.

Golden tanager Tangara arthus, copyright Alejandro Bayer Tamayo.

As noted above, tanagers feed on a mixed diet of fruit and insects. The fruit part is dominated by small berries that they can either swallow whole or mash with their bills before swallowing them piecemeal. Studies on the mixed-species flocks formed by Tangara species have found that while different species show very little variation in how they obtain the fruit component of their diet, they usually show very distinct specialisations in how they forage for insects. Some hunt for insects along branches, others prefer to look on leaves. Branch-hunting species may differ in the thickness and density of branches preferred, or in the mode of searching employed. For instance, the golden tanager T. arthus and flame-faced tanager T. parzudakii can both be found foraging on moss-covered branches, but the flame-faced tanager usually catches insects by probing directly into the moss whereas the golden tanager usually either focuses on the moss-free sections or catches insects sitting on the moss surface without probing. A few species catch insects aerially, making short sallies from a perch.

Blue-grey tanager Thraupis episcopus, indicated by phylogenetic analysis as a species of Tangara, copyright Mdf.

Somewhat unexpectedly for a genus of this size and diversity in a group as taxonomically challenging as the tanagers, molecular phylogenetic studies have largely corroborated Tangara's monophyly. They have also supported the monophyly of most of the species groups recognised within the genus of the basis of similarities in plumage patterns (Sedano & Burns 2010). The only exception has been the discovery that many of the species previously included in the genus Thraupis form a clade nested within Tangara, leading to the suggestion that these two genera should be synonymised (apart from in informal discussions online, I'm not aware of anyone suggesting the alternative that Tangara be split). The 'Thraupis' species are larger and plainer in coloration than most other Tangara species. A few taxonomists have also suggested that the colourful green tanagers of the genus Chlorochrysa should be included in Tangara, but this relationship has not been supported by molecular data. Chlorochrysa species are glossier than the often more matt-coloured Tangara, and they have an acrobatic mode of foraging involving postures such as regularly hanging upside-down that differ from any Tangara species.


Hilty, S. L. 2011. Family Thraupidae (tanagers). In: del Hoyo, J., A. Elliott & D. Christie. Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 46–329. Lynx Edicions: Barcelona.

Sedano, R. E., & K. J. Burns. 2010. Are the northern Andes a species pump for Neotropical birds? Phylogenetics and biogeography of a clade of Neotropical tanagers (Aves: Thraupini). Journal of Biogeography 37: 325–343.

The Wingless Penguin

A couple of weeks ago, I put up a page on the 'terrestrial penguin' Cladornis pachypus, described from the Oligocene of Patagonia by the Argentine palaeontologist Florentino Ameghino. As it happens, Cladornis wasn't the only unusual penguin recognised from the Patagonian fossil record by Ameghino nor was it even necessarily the most unusual. That title should probably go to another species, the wingless Palaeoapterodytes ictus.

Anterior (left) and posterior view of humerus of Palaeoapterodytes ictus, from Acosta Hospitaleche (2010). Scale bar = 10 mm.

Like Cladornis, Palaeoapterodytes was based on only a single bone, in this case a humerus (upper wing bone) from the Early Miocene. And also like Cladornis, Ameghino's description of this bone indicated a truly remarkable bird. The distal part of the humerus lacked any sign of the facets that would normally articulate with the succeeding wing bones and, as a result, Ameghino concluded that the wing skeleton had been reduced to the humerus only. The crest and pits on the humerus marking the attachment of the wing muscles were also reduced. Ameghino's Palaeoapterodytes presumably had wings reduced to the merest nubs, effectively functionless and probably of little mobility. Nevertheless, the humerus of Palaeoapterodytes remained relatively robust, its breadth little less than that of other penguins.

I am not aware of any other bird with a wing structure anything like this. In other birds without functional wings, the entire wing skeleton becomes reduced, not simply truncated. Perhaps the closest approximation I have found is the wing of Hesperornis, which also lacks known wing bones beyond the humerus. However, the Hesperornis humerus is slender and gracile, and even without direct indication of the presence of more distal bones, it still looks to retain some remnant of the ancestral articulation. Also, the whole concept of a wingless penguin is decidedly problematic. Hesperornis derived its main propulsion in swimming from its feet and so its wings became reduced because they served little function. Penguins, on the other hand, get most of their propulsion from their wings, swimming in a manner that has been compared to flying underwater. Despite being flightless, penguins retain a wing skeleton that is, if anything, even more well developed than that of their flying relatives. For Palaeoapterodytes to have lost functional wings, it would have somehow had to change its mode of propulsion.

Reconstruction of the Palaeoapterodytes humerus with missing sections restored, from Acosta Hospitaleche (2010).

As a result, even while authors were cautiously considering Ameghino's interpretation of Cladornis, they treated Palaeoapterodytes with more scepticism. This scepticism was eventually concerned when the humerus was re-examined by Acosta Hospitaleche (2010). The reason for the lack of structure at its distal end was very simple: the original distal end had been broken off. The apparent lack of development of the muscle attachment structures was the result of erosion, not any indication of the bone's original appearance. When alive, Palaeoapterodytes had probably been very similar to, if not identical with, one of the several other penguin species known from around the same time and place. Unfortunately, the state of preservation of the type humerus is so poor that its exact identity cannot be determined, and Palaeoapterodytes ictus has been cast into the taxonomic limbo of nomen dubium. Ameghino's Cladornis may remain an intriguing mystery, but his Palaeoapterodytes is just a red herring.


Acosta Hospitaleche, C. 2010. Taxonomic status of Apterodytes ictus Ameghino, 1901 (Aves; Sphenisciformes) from the Early Miocene of Patagonia, Argentina. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 255 (3): 371–375.

Brittle Stars, Brittle Taxa

Amphiura arcystata brittle stars extending their arms above the sediment, copyright James Watanabe.

The brittle stars are something of the poor cousin among echinoderm classes. Their tendency to relatively small size and cryptic habitats means that they do not attract the level of attention given to starfish, sea urchins or sea cucumbers. Despite this, they are perhaps the most diverse of the living echinoderm classes, with more recognised species around today than any other.

It should therefore come as no surprise that the internal classification of brittle stars remains decidedly up in the air. The basic framework of the surrent system was established over a hundred years ago by Matsumoto (1915) and changes to this arrangement since have been fairly cosmetic. However, a significant challenge to Matsumoto's system has been arisen following the input of molecular data to the mix: many of Matsumoto's higher groupings have not been supported by moleculat analyses. Perhaps the nail in the Matsumoto system's coffin has come from a recent publication by Thuy & Stöhr (2016) who found that a formal analysis of morphological data also failed to support the pre-existing classification. At this point in time, we know that a new classification of brittle stars is needed but we don't yet know what form it will take.

Excavated specimen of Amphiuridae, copyright Arthur Anker. The radial plates are visible as a pair of bars alongside the base of each arm; I don't think that the genital plates are visible externally.

Perhaps one of Matsumoto's groupings that will survive the transition is the Gnathophiurina. Notable features of this group include a ball-and-socket articulation between the radial shields (large plates that sit on the aboral side of the central body on either side of the insertion of each arm) and the genital plates (sitting below and alongside the radial shields), with the socket in the radial shield and the ball on the genital plate. The genital plates are also firmly fixed to the basal vertebra of each arm. I haven't been able to find what the functional significance of this arrangement is, such as whether it renders the body more flexible that in other groups where the radial-genital plate articulation is more fixed. At least one of the families of Gnathophiurina, the Amphiuridae, includes species that commonly live in burrows with the tips of their arms extended into the water column, using their tube feet to capture food particles (Stöhr et al. 2012). In contrast, some Ophiotrichidae are epizoic, living entwined around black corals and the like. The Gnathophiurina as a whole seem to be most diverse in relatively shallow waters.

Matsumoto's (1915) original concept of the Gnathophiurida included species that are now classified into four families, the Amphiuridae, Ophiotrichidae, Amphilepididae and Ophiactidae, and recent analyses have returned results not inconsistent with this association. In Thuy & Stöhr's (2016) morphological analysis, Gnathophiurina species all belong to, and make up the bulk of, their clade IIIc. In the molecular analysis presented by Hunter et al. (2016), the families belong to two separate clades but the branch separating them is very weakly supported. Further research is needed, of course, but it may turn out that Matsumoto was on to something when he focused on that ball-and-socket joint.


Hunter, R. L., L. M. Brown, C. A. Hill, Z. A. Kroeger & S. E. Rose. 2016. Additional insights into phylogenetic relationships of the Class Ophiuroidea (Echinodermata) from rRNA gene sequences. Journal of Zoological Systematics and Evolutionary Research 54 (4): 269–275.

Matsumoto, H. 1915. A new classification of the Ophiuroidea: with descriptions of new genera and species. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (1): 43–92.

Stöhr, S. T. D. O'Hara & B. Thuy. 2012. Global diversity of brittle stars (Echinodermata: Ophiuroidea). PLoS One 7 (3): e31940.

Thuy, B., & S. Stöhr. 2016. A new morphological phylogeny of the Ophiuroidea (Echinodermata) accords with molecular evidence and renders microfossils accessible for cladistics. PLoS One 11 (5): e0156140.

Public Service Announcement: Page priority is Not A Thing

I'll admit it, the rules governing taxonomy and nomenclature can seem horribly complicated when you don't spend a lot of time dealing with them directly. This isn't because taxonomy itself is inherently complicated: in fact, the underlying principles are really quite simple. The primary rationale behind each of the various codes of nomenclature can be distilled down to two points: (a) each single taxon should have a single name that differs from that of any other taxon, and (b) if more than two possible names can be assigned to a single taxon then the name given to that taxon first should be the one used (this latter point is called the principle of priority). Where things get complicated is that taxonomy is a process run by and for human investigators. And if there's one thing that can be said about all human endeavours, from science to politics to the selection of sports teams, it's that any application of simple principles is going to run afoul of complex practicalities. So questions arise that any code of nomenclature has to deal with: is it always ideal to simply use the oldest name? How do we determine which name is 'oldest', and what do we do if it's not clear? Each of the codes has developed its own methods of dealing with these questions and others, but it is not uncommon for these methods to be overlooked or misunderstood, sometimes even by people who might be expected to know better. One particularly pernicious misunderstanding that I've often come across (and which I was reminded of recently by one paper in particular that will go unnamed) is 'page priority'.

As mentioned above, it is not uncommon for two or more names to turn out to be synonymous without one particular name being obviously 'older'. Perhaps the papers naming each species were published at the same time, or they were named within a single paper. In these situations, many authors will invoke the principle of page priority in determining which name should be used: the name which appeared in an earlier place in the publication (say, on p. 23 rather than p. 25) should be the one used. The problem is that, in the case of the International Code of Zoological Nomenclature at least, no such principle is mandated (I'm not so familiar with the other codes, but I don't think they have page priority either). Instead, the code resolves indeterminate priority through the principle of the 'first reviser'. In the code's own words:
24.2.1. When the precedence between names or nomenclatural acts cannot be objectively determined, the precedence is fixed by the action of the first author citing in a published work those names or acts and selecting from them; this author is termed the "First Reviser".

In other words, the question of which name takes priority is determined by the choice of the first person who treats them as synonyms. Any subsequent authors are required to abide by the decision made by the first reviser. There is no restriction placed on how the first reviser should make their decision; if they want to follow page priority, they're perfectly free to do so. The problem comes when a first reviser doesn't follow page priority, only to have later authors claim they made the "wrong" decision. Maybe the name that appeared on a later page in the original publication was better described, or had been more commonly referred to by later authors. Whatever the situation, the decision of the first reviser is final. No correspondence shall be entered into.

"Page priority" is a bit of a case of what's been called a hypercorrection, when someone 'corrects' something that was already right. Someone who's familiar with a principle (in this case the principle of priority) but doesn't fully understand the reasons for the principle may try to apply it more widely than they should. So, for instance, someone who knows the plural of 'hippopotamus' is 'hippopotami' may assume that the plural of 'octopus' is 'octopi'. Hypercorrections persist because, on a superficial level, they 'make sense'. Sadly, being sensible is no barrier to being wrong.

Interestingly enough, there was a brief time in zoological nomenclature when page priority was a mandated rule (Dubois 2010). Between 1948 and 1953, a page priority clause was inserted into the Règles Internationales de la Nomenclature Zoologique, the earlier code of zoological nomenclature that was used before 1961. In 1953, this clause was suppressed and invalidated, and the first reviser principle now applies almost universally. Anyone who argues that a taxonomic decision violates 'page priority' can be safely ignored.


Dubois, A. 2010. Retroactive changes should be introduced in the Code only with great care: problems related to the spellings of nomina. Zootaxa 2426: 1–42.


Parastenocaris brevipes, copyright A. Hobaek.

It's time for another consideration of the overwhelming diversity of stygofaunal copepods. Parastenocaris is a genus of copepods found on almost all the landmasses of the world except, presumably, Antarctica (New Zealand also stands out as an intriguing void in the genus' distribution). The majority of species in this genus are insterstitial, mostly found in soils saturated with fresh water; a small number of species are found in brackish habitats such as estuaries. A few species have also been found above ground, particularly in the tropics (Galasi & Laurentiis 2004). The type species of the genus, P. brevipes, has been found in sphagnum bogs (Karanovic 2005).

As commonly recognised, Parastenocaris is a pretty huge genus, with well over 200 species having been assigned to it over the years. However, the genus has been poorly defined and many authors have questioned its integrity. Galasi & Laurentiis (2004) suggested that Parastenocaris should be restricted to those species most closely related to the type species, P. brevipes. Such a group would still be pretty cosmopolitan; indeed, P. brevipes itself has a Holarctic distribution and is known from both Europe and North America (this stands in pretty stark contrast to the super-short ranges of some stygofaunal copepods). Distinctive features of this restricted P. brevipes group include a characteristic endopodal complex on leg 4 of the male, with the endopod hyaline and with one or two large claws. In contrast, the leg IV endopod in females is long and distally serrate.

Parastenocaris lacustris, from here

Members of the Parastenocaris brevipes groups are found closer to the soil surface than many other members of their family (Karanovic 2005). They are also relatively large, reaching the absolutely monstrous size (I'm sure) of half a millimetre or more. Karanovic (2005) suggested that this larger size could reflect the larger size of the sand grains they live among closer to the surface, or it could simply reflect their access to more reliable food sources that are available to their more deeply buried relatives.


Galasi, D. M. P., & P. de Laurentiis. 2004. Towards a revision of the genus Parastenocaris Kessler, 1913: establishment of Simplicaris gen. nov. from groundwaters in central Italy and review of the P. brevipes-group (Copepoda, Harpacticoida, Parastenocarididae). Zoological Journal of the Linnean Society 140: 417–436.

Karanovic, T. 2005. Two new subterranean Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from Western Australia. Records of the Western Australian Museum 22: 353–374.

The Patagonian Land Penguin

Take a good look at the figure above, which comes from Mayr (2009). It shows the fossilised tarsometatarsus (the fused long bone of the foot) of a bird from the late Oligocene of Patagonia. This may be one of the single most mysterious specimens in the fossil record. It represents all we know to date of Cladornis pachypus, described by Argentinean palaeontologist Florentino Ameghino in 1895. The appearance of the bone, being very broad and flat relative to its length, is quite bizarre and does not much resemble the tarsometatarsus of any other known bird.

The first thing that should be pointed out is that, whatever it was, Cladornis was a large bird. The specimen is not completely preserved (part of the proximal end of the bone has been lost) but its overall shape suggests that its original length was probably not too much longer than what we have. As such, the tarsometatarsus was probably comparable in length to that of a large pelican. However, it was much wider relative to length than that of a pelican, suggesting the possibility of a more robust bird. The shape of the bone's end indicates that the toes would have been widely spaced, and it may have even approached a zygodactyl arrangement (with two toes pointed rearwards and two forwards, like a modern parrot*) (Mayr 2009).

*When explaining this to my partner, I suggested that he imagine a parrot the size of a pelican. He shuddered and declared that he would rather not.

When Ameghino (1895) first described Cladornis, he interpreted it as an aquatic bird and suggested a relationship to the penguins, albeit in an extinct family Cladornidae (later authors would correct this to Cladornithidae). Later, noticing that it was preserved in association with terrestrial mammals, he declared that it was not marine and was possibly even terrestrial (he also included another species from the same formation, Cruschedula revola, in the Cladornithidae; this species is based on part of a scapula and there is no telling if it was related to Cladornis or not). He still maintained its relationship to the penguins (Ameghino 1906). Ameghino had a bit of a thing for trying to find the origins of all major modern vertebrate groups in his native South America (one of his other works was a book arguing for an Argentinean origin of humans) and it is possible that this was in play here. Nevertheless, the idea of a 'Patagonian land penguin' held sway until Simpson's (1946) review of the fossil penguins, in which he declared that Cladornis was "so very unlike any other penguin, recent or fossil, that I can only consider its reference to that group as erroneous".

This left Cladornis' taxonomic position completely up in the air (the question of whether Cladornis itself could get up in the air is, of course, currently completely unswerable). Wetmore (1951) included Cladornis in the Pelecaniformes, because...reasons. The closest he gave to an explanation was, "The only suggestion that has come to me is that possibly they may belong in the order Pelecaniformes, in which I have placed the family tentatively in the suborder Odontopteryges, where it is located with two others of almost equally uncertain status. This allocation is wholly tentative and is no indication of belief in close relationship in the three diverse groups there assembled". He would later move Cladornis into its own suborder, Cladornithes, and no close relationship to the 'Odontopteryges' (now the Pelagornithidae) has been suggested since. Our current understanding of bird phylogeny finds Wetmore's remaining 'Pelecaniformes' to correspond to three or four independent clades (the Pelecanidae, Suliformes, Phaethontidae and probably Pelagornithidae) so his assignment of Cladornis to this group becomes almost completely uninformative.

Which is pretty much where we're forced to leave things. Mayr (2009) included Cladornis in his chapter on 'land birds', with other taxa discussed in this chapter belonging to the clade Telluraves. However, this was motivated more by a lack of any idea what to do with it otherwise than anything else (it is possible that Cladornis' sub-zygodactyly played a role, but not all zygodactylous birds belong to the Telluraves). I did notice a similarity in proportions between the Cladornis tarsometatarsus and the corresponding bone in the large phorusrhacid Brontornis, making me wonder if anyone had ever compared the two, but this may well be only superficial. Most recent authors have assumed that the Cladornis tarsometatarsus is simply too weird, too unique, for any resolution of its affinities to be reached without first finding more complete remains of the animal.


Ameghino, F. 1895. Sur les oiseaux fossiles de Patagonie et la aune mammalogique des couches a Pyrotherium. Boletín del Instituto Geográfico Argentino 15 (11–12): 501–602.

Ameghino, F. 1906. Enumeración de los Impennes fósiles de Patagonia y de la Isla Seymour. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 97–167.

Mayr, G. 2009. Paleogene Fossil Birds. Springer.

Simpson, G. G. 1946. Fossil penguins. Bulletin of the American Museum of Natural History 87 (1): 1–99.

Wetmore, A. 1951. A revised classification for the birds of the world. Smithsonian Miscellaneous Collections 117 (4): 1–22.

Camponotus: A Sugary High

I think I may have said before that Australia is the land of ants. When travelling in Australia's arid regions (i.e. most of the continent), ants are often the most visible animals about. Perhaps the most visible of all Australia's ants are the meat ants (Iridomyrmex), but not too far behind them are the sugar ants of the genus Camponotus.

Workers and emerging queens of banded sugar ants Camponotus consobrinus around the nest opening, copyright Steve Shattuck.

Camponotus is a genus of the ant subfamily Formicinae found pretty much everywhere around the world that ants are to be found. It is massively diverse: well over 1000 species have been assigned to this genus over the years, with probably more to be described. They are correspondingly diverse in habits and appearance. Some are among the giants of the ant world, others are much smaller. Some form massive colonies that are difficult to miss and forage during the day, others are more retiring and emerge only at night. Some construct their nests in holes under the grounds, others hollow out wood or use the holes left by other wood-boring insects. Most (but not all) Camponotus species exhibit some form of worker polymorphism: rather than having just a single worker caste, a colony will often include large major workers and much smaller minor workers, with the two forms superficially looking quite different. Sometimes the distinction between majors and minors will be quite clear, other times there will also be workers of intermediate sizes. In some Australian species, known as honeypot ants, there are specialised workers called 'repletes' who spend their lives hanging in one spot inside the nest, being fed by the other active workers until their gasters swell into engorged round balls. These repletes serve the colony as a living larder, able to regurgitate their stored excess of food when needed by their nestmates. Despite all this diversity, most Camponotus species are readily recognisable as Camponotus: they usually lack spines on the mesosoma (the 'thorax'), the back end of which is narrow and often arched. This smoothness and slimness gives Camponotus a distinctive look that kind of puts me in mind of the ant version of a greyhound. The majority of Camponotus species also differ from other ants in lacking the metapleural gland, a gland producing an antibiotic chemical whose opening is usually visible near the rear of the mesosoma.

Camponotus aurocinctus, copyright Steve Shattuck.

Camponotus species have been referred to in Australia as 'sugar ants' in reference to their diet, which is commonly dominated by the sugary excretions of plant-sucking bugs that they attend. In other parts of the world, they have sometimes been referred to as 'carpenter ants' in reference to the wood-tunneling habits of their most notorious representatives. Bug-derived honeydew is high in sugar but low in other essential nutrients, so the ants also feed on things such as the scavenged bodies of the bugs themselves after death. They are also probably assisted in meeting their nutritive needs by Blochmannia, an endosymbiotic bacterium that infests specialised cells in the gut of Camponotus and closely related genera (Wernegreen et al. 2009). Genetic data from the endosymbiont indicates that it probably synthesises nutrients the ant does not otherwise ingest. It may also play some role in compensating for an absence of metapleural gland secretions. As well as the gut, Blochmannia infest the ovaries of reproductive females and are passed to the next generation via the developing oocytes. Phylogenetic analysis of Blochmannia indicates that it is closely related to other endosymbiotic bacteria found in mealybugs, and it is possible that the ancestors of Camponotus picked it up in the course of feeding on honeydew.

Honeypot ant Camponotus inflatus repletes hanging in the nest, copyright Mike Gillam.

The sheer size of Camponotus as a genus has been a challenge to understanding relationships within the genus. Over thirty subgenera have been proposed at one time or another, but many of these are poorly defined and many authors eschew using them in favour of informal species groups. It does not help matters that, since the early 20th century, most reviews of Camponotus have been conducted at a local rather than a global level. Those studies that have touched on Camponotus phylogeny in recent years suggest the need for a large-scale revision, with few of the subgenera supported as monophyletic.


Wernegreen, J. J., S. N. Kauppinen, S. G. Brady & P. S. Ward. 2009. One nutritional symbiosis begat another: phylogenetic evidence that the ant tribe Camponotini acquired Blochmannia by tending sap-feeding insects. BMC Evolutionary Biology 9: 292. doi:10.1186/1471-2148-9-292.


Slide-mounted male of Zercon gurensis, copyright Holger Müller.

The animal depicted above is a mite of the Zerconidae, one of the numerous families in the major mite clade known as the Mesostigmata. This family is mostly found in soil habitats such as leaf litter, mosses, decaying vegetation, or occasionally in animal nests (Lindquist et al. 2009). The zerconids are restricted to the Northern Hemisphere and are most diverse in temperate to Arctic regions; those species found in tropical parts of the world are restricted to high altitudes away from the hot lowlands (Ujvári 2012). Like many other Mesostigmata, they have the dorsal surface of the body mostly covered by shields of hardened cuticle. In most zerconids, separate shields cover the front (podonotal) and rear (opisthonotal) sections of the dorsum; the opisthonotal shield wraps around the rear margin of the mite and forms a continuous unit with the ventrianal shield that usually protects most of the underside of the mite behind the legs. Among the most noticeable features of the zerconids are two pairs of large openings near the rear of the opisthonotal shield (the four orange-segment-like structures in the photo above). These represent the openings of secretory glands, but I don't know if it has been established just what they're secreting; comparable structures in other mites may secrete pheromones, or defensive chemicals, or oils that prevent debris from sticking to the body. Other features of the zerconids include slender, relatively simple chelicerae that lack the modifications seen in the males of some other Mesostigmata, and peritremes (grooves on the underside of the body that channel air to the openings of the respiratory stigmata) that are relatively short. These peritremes are longer in zerconid nymphs, but become shortened when the mite moults to maturity.

Zerconids are another of those mite groups where the vast majority of what has been written about them relates to their basic taxonomy, with little yet known about their natural history. Several genera of zerconids are recognised, distinguished by features such as the shape of the body's various shields and the appearance of various setae. The form of their chelicerae indicates that zerconids are predatory like many other Mesostigmata. Because they are mostly found at ground level rather than on vegetation they have not attracted the economic interest of other predatory mites, but those few species that have been observed feeding were chowing down on nematodes. Mating does not appear to have been directly observed in zerconids, but again their anatomy and comparison with other mesostigs allows us to infer that the male fertilises the female by using his chelicerae to pass a spermatophore from his own genital opening on the underside of the body between the legs to hers. Where she then lays her eggs, and how her offspring spend their time to maturity, seem to be questions still awaiting an answer.


Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009. Order Mesostigmata. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 124–232. Texas Tech University Press.

Ujvári, Z. 2012. Draconizercon punctatus gen. et sp. nov., a peculiar zerconid mite (Acari: Mesostigmata: Zerconidae) from Taiwan. Opusc. Zool. Budapest 43 (1): 79–87.

Midges of the Macabre

In a recent post, I commented that derived members of an insect 'order' could sometimes be all but unrecognisable as belonging to that order. Take a look at this:

Miastor sp., copyright Charles Olsen.

Believe it or not, sitting in the middle of that photo is a fully reproductively mature insect (I'm not so sure about the maturity of the other individuals). In fact, it's a fully mature fly. Miastor is a genus of midges found living in rotting wood or fungal fruiting bodies. Members of this genus are found worldwide; I believe that several species are recognised, but distinguishing the individual species is extremely difficult. Miastor exhibit what is called paedogenesis: they can become reproductively mature while still in the larval state. They are not the only genus of the family Cecidomyiidae to exhibit this process; another such genus, Oligarces, is found in similar habitats. Miastor is probably the best known such genus, in part because it has been cultured for study in the laboratory, and in part because of the decidedly macabre way in which its paedogenesis plays out.

Each paedogenetic Miastor larva develops several eggs within its ovaries (generally four to ten or more—Harris 1923). No males are involved in this process: the larvae are parthenogenetic as well as paedogenetic. These eggs hatch while still inside their mother, after which the daughter larvae are nourished by the mother's own tissues. Eventually, the daughters are not born so much as they escape. In the words of Quammen (1985), "While the food lasts, while opportunity endures, no Miastor female can live to adulthood without dying of motherhood". But karma still seems to have its place, because by the time the daughter larvae escape they carry their own fate within them: they emerge with their own eggs already developing inside them.

Metamorphosed male Miastor metraloas, copyright John Plakidas.

In this way, a Miastor colony can go through an entire generation in as little time as two weeks, until changing conditions (such as exhaustion of food supplies, or a change in season) induce a change in tack. Larvae are produced that do not reproduce paedogenetically in the way that their mothers did, but instead pupate in the usual way to emerge as more ordinarily formed midges, both males and females. As with another paedogenetic insect that has already been featured on this site, the beetle Micromalthus debilis, these metamorphosing larvae can be readily distinguished from paedogenetic larvae. Not only do they not produce the precocious reproductive organs of their fellows, but they have visible imaginal discs (the clumps of tissue that develop into adult structures during metamorphosis), and have eyes that are clearly separated on top rather than touching as in paedogenetic individuals (Harris 1923). After the mature adult midges emerge from their pupae, they can disperse in search of new habitats, seeking food for their offspring and mates for themselves in order to begin the cycle again.


Harris, R. G. 1923. Occurrence, life-cycle, and maintenance under artificial conditions, of Miastor. Psyche 30 (3–4): 95–101.

Quammen, D. 1985. Natural Acts. Schocken Books.

The Mongolian Death Worm

This would have been a comment on a recent post by Darren Naish at Tetrapod Zoology on the behaviour of amphisbaenians, but the commenting system they have at Scientific American these days means that any comment on a post more than a couple of days old will never be seen by anyone. As such, I'm posting it up here:

PhilJTerry's comment in response to Darren's post: "As I love introducing cryptozoology into the conversation wherever possible - are Amphisbaenians a likely influence for the Mongolian Death Worm? Can they live in desert environments?"

Image from National Geographic.

For those not already aware, the "Mongolian death worm" or "olgoi-khorkhoi" is a supposedly incredibly dangerous animal found in the deserts of Mongolia. It's first mention in Western literature came in Roy Chapman Andrew's (1926) On the Trail of Ancient Man. Andrews heard about the animal in a meeting with Mongolian officials:

Then the Premier asked that, if it were possible, I should capture for the Mongolian government a specimen of the allergorhai-horhai. I doubt whether any of my scientific readers can identify this animal. I could, because I had heard of it often. None of those present ever had seen the creature, but they all firmly believed in its existence and described it minutely. It is shaped like a sausage about two feet long, has no head nor legs and is so poisonous that merely to touch it means instant death. It lives in the most desolate parts of the Gobi Desert, whither we were going. To the Mongols it seems to be what the dragon is to the Chinese. The Premier said that, although he had never seen it himself, he knew a man who had and had lived to tell the tale. Then a Cabinet Minister stated that "the cousin of his late wife's sister" had also seen it. I promised to produce the allergorhai-horhai if we chanced to cross its path, and explained how it could be seized by means of long steel collecting forceps; moreover, I could wear dark glasses, so that the disastrous effects of even looking at so poisonous a creature would be neutralized. The meeting adjourned with the best of feeling; for we had a common interest in capturing the allergorhai-horhai.

Since then, there have been a number of expeditions have been conducted in search of the Premier's "allergorhai-horhai"; all have come up fruitless. Various opinions have been expressed as to what the stories may have been based on, with the most popular suggestions being some sort of reptile (Darren says in his response to the above comment on the original post that he "could buy that the stories are based on exaggerated tales of erycine boas or something"). For my part, I suspect that the question of the 'original identity' of the Mongolian death worm may be a futile one. When I first heard Andrews' account, I was not reminded of an amphisbaenian or a boa; I was immediately put in mind of a drop bear.

I feel almost certain that Andrews was being told a local tall tale, a popular joke at the expense of visiting travellers. The nature of Andrews' response to the officials suggests that he was in on the joke and more than happy to play his part in communicating it. Admittedly, other accounts of the Mongolian death worm have been recorded at more recent dates. And in the same way, I've never seen a drop bear myself, but I can assure you that my cousin did once and got the fright of his life. Be careful. They're out there.

Scurvy and Cress

Without the subject of today's post, it's just possible that my home country of New Zealand could have had quite a different history. Sometimes, one shouldn't overlook the importance of cress.

Pepperwort Lepidium heterophyllum, copyright Anne Burgess.

Lepidium is a genus of herbs and subshrubs belonging to the Brassicaceae, the same family as cabbages, radishes and cauliflowers. The genus is found worldwide, and more than 150 species have been recognised to date. The fruit is a type of dry capsule called a silicle which is usually dehiscent (one subgroup of Lepidium, previously separated as the genus Cardaria, has indehiscent fruit), with strongly keeled or winged valves, and contains a single pendulous seed in each locule. The seeds are usually copiously covered in mucilage (Mummenhoff et al. 2001). Like other members of the Brassicaceae, Lepidium has not been overlooked for culinary uses. Leaves and stems of number of species in the genus, such as garden cress Lepidium sativum and dittander Lepidium latifolium, are used as pot or salad herbs. A South American species, maca Lepidium meyenii, is grown as a root vegetable.

Because of its wide distribution, some early authors suggested that Lepidium was a very ancient genus whose members had diverged with the break-up of the Mesozoic supercontinents. However, more recent phylogenetic analyses (Mummenhoff et al. 2001) have suggested just the opposite: the crown group of Lepidium may have originated in the Mediterranean-Central Asian region little more than two million years ago. The mucilaginous seeds of many species become sticky when damp, and can easily be carried long distances adhered to birds' feet and other such dispersal agents. Perhaps the most dramatic suggestion of intercontinental dispersal in the genus involves a clade of species found in Australia and New Zealand that phylogenetic analysis suggests originated via hybridisation between two divergent species—with one parent being native to South Africa and the other to California (Mummenhoff et al. 2004).

Cook's scurvy grass Lepidium oleraceum, copyright Andrea Brandon.

It was one of the members of the latter clade that played a small but significant role in New Zealand history. Lepidium oleraceum is an endemic New Zealand species that was once found growing over much of the country. It is commonly known as 'Cook's scurvy grass', because Captain James Cook was able to collect it while surveying New Zealand to provide vitamin C to stave off the scurvy that could have otherwise devastated his crew. Sadly, this once common plant is now extremely rare: the disappearance of mainland-nesting seabirds means that they are no longer around to provide the guano-enriched soils on which this plant thrived. It also proved extremely palatable to introduced herbivores. As a result, Cook's scurvy grass is now almost exclusively found on small offshore islets.


Mummenhoff, K., H. Brüggemann & J. L. Bowman. 2001. Chloroplast DNA phylogeny and biogeography of Lepidium (Brassicaceae). American Journal of Botany 88 (11): 2051–2063.

Mummenhoff, K., P. Linder, N. Friesen, J. L. Bowman, J.-Y. Lee & A. Franzke. 2004. Molecular evidence for bicontinental hybridogenous genomic constitution in Lepidium sensu stricto (Brassicaceae) species from Australia and New Zealand. American Journal of Botany 91 (2): 254–261.

Linnaeus' Infernal Fury

The starting point of modern zoological nomenclature (Clerck notwithstanding) has been established as the tenth edition of Linnaeus' Systema Naturae, published in 1758. Linnaeus divided the animal kingdom between six classes, with vertebrates making up four (Mammalia, Aves, Amphibia and Pisces) and invertebrates assigned to just two. One of these, Insecta, essentially corresponded to modern arthropods, and all other invertebrates were included in the class Vermes, 'worms'. Linnaeus' concept and arrangement of Vermes bears little resemblance to anything that exists in modern zoological classifications; with the study of invertebrate anatomy still in its absolute infancy, he was largely classifying animals based on their overall external appearance alone. One of Linnaeus' orders of Vermes, the 'Intestina', defined as 'simple, shell-less and limb-less', included animals now classified as annelids, nematodes, molluscs and even a chordate (the hagfish Myxine glutinosa). It also included a species whose identity would be debated for the next several decades: the 'infernal fury', Furia infernalis.

A reconstruction of Furia infernalis, from Piter Kehoma Boll.

Furia infernalis was described by Linnaeus as "Corpus filiforme, continuum, aequale, utrinque ciliatum: aculeis reflexis corpori appressis" ('body thread-like, continuous, uniform, ciliated on both sides with reflexed spinules appressed to the body'). It was found in marshes of southern Sweden and Finland. Linnaeus went on to record that F. infernalis was, "Pessima omnium, ex aethere decidua in corpora animalium, ea momento citius penetrat, intra horae quadrantem dolore atrocissimo occidit": the 'worst of all, falling from the sky onto the bodies of animals, into which it rapidly penetrates within a moment, striking [the victim] down with the most atrocious pain within quarter of an hour'. Linnaeus had good reason to highlight this animal's unpleasantness: he had been attacked by one himself when collecting botanical specimens in 1728, and barely escaped the resulting ailment with his life. A more detailed description of "der Höllenwurm" was compiled by Jördens (1802): it was a very slender worm, about the length of a nail, of a pale yellow or fleshy colour (other authors described it as greyish), with one end black. It climbed up standing vegetation, from whence it was carried by the breeze onto the exposed skin of humans and animals into which it rapidly burrowed. For victims, the first sign of its presence was usually a sudden pain in the afflicted spot, like the stab of a needle, and a small black spot marking the worm's entry point. A violent itching followed that developed into severe and extensive inflammation, often accompanied by fever; in the majority of cases, the affliction was so violent that the victim was dead within a matter of days if immediate action was not taken. If applied quickly enough, the worm could sometimes be drawn out with a poultice of fresh cheese curds. Otherwise, treatment required the careful dissection of the worms from between the muscle tissue into which they had entered, a process that (considering the surgical facilities available at the time) must have nearly as hazardous as the original infection.

As can be imagined, the attacks of this animal were greatly feared. In 1823–1824, an epidemic of Furia attacks spread through herds of livestock in Swedish and Finnish Lapland; thousands of head of reindeer perished, as well as countless cattle and sheep. Scavengers such as wolves feeding on the carcasses themselves sickened and died. One account from the time involves a young woman who was shearing wool from a recently deceased sheep (on a waste not, want not principle, I suppose) when she felt the tell-tale sting on a knuckle. Her life was saved by her master who was working nearby, when he quickly chopped off the affected finger with an axe. So great was the devastation that Norway, which had hitherto been free of the worm, passed an edict banning the import of animal furs from affected areas (Brooke 1827).

There were some, however, who greeted the description of Furia infernalis with skepticism. The idea of a tiny worm that somehow flew through the air and caused almost instantaneous mortality seemed fantastic. Even more problematic was the dearth of specimens. Many had seen the wounds caused by the worm and observed its effects; very few had seen the worm itself. Linnaeus himself had only seen a single, very poorly preserved specimen submitted to him by a church pastor. Most of the details about the worm's supposed appearance came from a single source, an article written by Solander, a student of Linnaeus'. The Academy of Sciences at Stockholm, naturally keen to discover all they could about such a scourge afflicting their country, offered generous rewards to anyone who could procure them a genuine specimen; no such specimen was forthcoming. Eventually, a consensus was reached: the worm Furia infernalis was an entirely fabulous animal, with no place in the annals of physical zoology. By 1827, notwithstanding the epidemic of only a few years previously, Brooke was able to comment that one could quite easily accept that something had affected the supposed victims of Furia without presuming that that something had to be the Furia itself. Even Linnaeus eventually came to accept that his inclusion of Furia in the Systema Naturae had been an error.

That Furia infernalis never existed outside the realms of fantasy remains the accepted wisdom to this day. But in that case, what did afflict Linnaeus and other unfortunates wandering the marshes of Sweden in the early 1700s? One thing that struck me was how much I was reminded of the more recent phenomenon here in Australia of 'white-tailed spider bites'. In recent decades, many people (including many medical professionals) have attributed serious ulcerative skin lesions, sometimes so serious that treatments such as skin grafts are required, to the bite of white-tailed spiders Lampona spp., common ground-running spiders often encountered near human dwellings. The actual evidence linking white-tailed spiders to such injuries is minimal; indeed, a clinical survey of 130 confirmed white-tail bites by Isbister & Gray (2003) found not a single incidence of one leading to ulceration. In both the 'Furia attacks' and the 'white-tailed spider bites', it seems likely that the primary culprit is bacterial infections resulting from opportunistic pathogens such as Streptococcus and Staphylococcus species. The initial wound may indeed have been caused by something like an animal bite or sting, or for that matter a splinter or pin-prick. Germ theory would not become widely accepted until the mid- to late 1800s; when Linnaeus compiled the Systema Naturae, flying worms probably seemed as good an explanation as any. The first 'attack' recorded by Furia victims may have simply been the first moment they noticed the infection's symptoms. And the 'worms' dissected out of advanced victims? Personally, I'm inclined to suspect that they may have been small pieces of tissue from the unfortunate sufferers themselves.

The exact causes of the 1823 epidemic are probably lost to history. Brooke (1827) stated that faculty at the Stockholm academy "had been led to consider the disorder by which [the reindeer] were attacked as a particular variety of hydrophobia". He also mentioned another possibility: reindeer were known to be vulnerable to inflammation of the brain, and dissections of the brains of deer killed by this condition sometimes revealed the presence of "a small vesicular worm". We can now recognise these vesicles as the cysts of hydatid tapeworms, which can hatch to cause tapeworm infections in any predator that eats the flesh of their host. So perhaps the 1823 epidemic was caused by a worm after all—just not the worm that was blamed.


Brooke, A. de C. 1827. A Winter in Lapland and Sweden, with various observations relating to Finmark and its inhabitants; made during a residence at Hammerfest, near the North Cape. John Murray: London.

Isbister, G. K., & M. R. Gray. 2003. White-tail spider bite: a prospective study of 130 definite bites by Lampona species. Medical Journal of Australia 179: 199–202.

Jördens, J. H. 1802. Entomologie und Helminthologie des Menschlichen Körpers, oder Beschreibung und Abbildung der Bewohner und Feinde desselben unter den Insekten und Würmern vol. 2. Gottfried Adolph Grau: Hof.

Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis 10th ed., revised, vol 1. Laurentius Salvius: Copehagen.

Two New Insect Orders?

When a new species of insect is described as being distinct enough to represent a new order, it's kind of a big deal. So it certainly caught my attention over the past year when, not one, but two species from Cretaceous Burmese amber were considered worthy of the honour. Now, I'm going to be up front here and say that, while both are very interesting specimens, in both cases I think that the 'new order' label may be a trifle overblown. What's interesting to me is that my reasons for thinking so are different for both. Let's take a look, shall we?

Lateral and dorsal views of holotype of Alienopterus brachyelytrus, from Bai et al. (2016). Pink scale bar = 1 mm.

The first was published in March of last year by Bai et al. (2016) under the name of Alienopterus brachyelytrus. In overall appearance, Alienopterus resembled a long-legged cockroach, but with the head clearly visible instead of hidden by the pronotum in the cockroach manner. The head would have been mobile and capable of being turned in the manner of a modern cockroach or mantis. The forewings were hardened and reduced to small pads covering only the base of the hind wings, which retained their full length. The femora of the front legs bore a pair of dense rows of setae on their underside, and Bai et al. suggested that Alienopterus may have used these setae to help it grab prey.

A phylogenetic analysis of Alienopterus placed it together with the modern cockroaches and mantids, specifically as the sister group to the latter. Because Alienopterus lacked the primary distinguishing features of a mantis (such as the spined raptorial forelegs), and because of its distinctive wing morphology, Bai et al. made it the type and only species of a new order, the Alienoptera. But there are a number of reasons why I find this designation problematic. It is generally agreed these days that cockroaches and mantids (and termites) together form a clade known as the Dictyoptera. Many people have an idea that cockroaches are one of the oldest living groups of insects, having supposedly been around for hundreds of millions of years. But modern cockroaches and mantids only diverged sometime during the Jurassic and Cretaceous; earlier members of the Dictyoptera were cockroach-like, certainly, but they were just as close to mantids as to cockroaches, and also had features very distinct from either. If we are to recognise a distinct 'order' for Alienopterus purely on phylogenetic grounds, then we would also have accept several separate 'orders' for each of the various lineages of stem-dictyopterans. And as distinctive as Alienopterus is morphologically, it is not the only (or even the most) unusual member of the Dictyoptera. This is, after all, the lineage that has given the termites with their wood-chomping biology and baroque caste system, beetle-like taxa with full-on elytra, and active leapers like the Jurassic Skok svaba or the modern Saltoblattella montistabularis.

There is a definite paradox at play here. On the one hand, the question of which lineages get designated 'orders' is completely arbitrary because there is no formal definition for an 'order' except that it is a taxon that is somehow more significant than a 'family' (itself a completely arbitrary level). From that perspective, there is no inherent reason why the Dictyoptera should not get divided between any number of orders. But on the other hand, the concept of 'order' has a certain cultural cachet. 'Orders' are kind of the base units of entomology: the first thing that any student of entomology is likely to do is learn to distinguish between the various insect orders. Labelling a particular taxon an 'order' is a statement of value; it says that that taxon is somehow fundamentally important in a way that other taxa are not. And while, again, Alienopterus is a very interesting animal in terms of what it can potentially tell us about cockroach-mantis relationships, it is hard to see how it can be called 'fundamental'. There have been extinct 'orders' recognised from the fossil record, such as the Palaeodictyoptera, but such taxa represent notable radiations. With only a single known species, referring to Bai et al.'s taxon as 'Alienoptera' tells us little more than calling it an unplaced species within the Dictyoptera.

Various views of Aethiocarenus burmanicus from Poinar & Brown (2016).

The other new 'order' made its appearance in December, when Poinar & Brown (2017) published Aethiocarenus burmanicus (if you're confused about the date, it reflects the difference between the online and print publication). This was a very odd little insect: a flattened and wingless yet long-legged animal with long antennae. The most distinctive feature of Aethiocarenus is its head, which is globular with great bulging eyes and placed on a narrow neck. Poinar & Brown suggest that it may have made its living hunting in confined spaces, such as crevices in bark or among epiphytes. Because of its highly distinctive appearance from any other known insect, Poinar & Brown placed it in its own new order, the Aethiocarenodea.

In this case, my issue with the establishment of a new 'order' is that it is essentially a statement of ignorance. As distinctive as Aethiocarenus is, there are many equally unusual-looking insects that are not placed in their own 'order'—particularly among wingless forms that can get up to all sorts of freakiness. The overall 'jizz' of Aethiocarenus, particularly the distinct cerci, suggest that its affinities probably lie somewhere within the Polyneoptera, the group of insects including such forms as cockroaches, grasshoppers and stoneflies. Within other polyneopteran orders, a novice entomologist would be hard-pressed to recognise a sandgroper as a grasshopper, or the Javan cave-dweller Arixenia esau as an earwig. Similarly, without a formal analysis it is difficult to exclude the possibility that Aethiocarenus represents a kooky member of some already recognised order. And again, with only one known species, recognition of an 'order' Aethiocarenodea tells us little more than recognition of an unplaced Aethiocarenus.


Bai, M., R. G. Beutel, K.-D. Klass, W. Zhang, X. Yang & B. Wipfler. 2016. Alienoptera—a new insect order in the roach-mantodean twilight zone. Gondwana Research 39: 317–326.

Poinar, G., Jr & A. E. Brown. 2017. An exotic insect Aethiocarenus burmanicus gen. et sp. nov. (Aethiocarenodea ord. nov., Aethiocarenidae fam. nov.) from mid-Cretaceous Myanmar amber. Cretaceous Research 72: 100–104.

Screech Owls

Eastern screech owl Megascops asio emerging from a nest-hole, copyright Zach.

For many people, owls are a group of birds known more by reputation than by encounter. Their nocturnal lifestyles and often cryptic habits make them rarely seen, and their diversity is often little appreciated. But despite being commonly referred to as a single homogeneity, owls actually come in a whole range of shapes and forms.

Megascops, the screech owls, is a genus of more than twenty species found over most of the Americas, though they are less diverse in North America than in the remainder of their range. They are also notably absent from the Caribbean; a single species from that region assigned to this genus, the Puerto Rican screech owl Megascops nudipes, was found in a recent phylogenetic study (Dantas et al. 2016) to be closer to the flammulated owl Psiloscops flammeolus of North America and may require reclassification. Despite what one might presume from the genus name alone, species of Megascops are small owls, around 20–25 cm in length. They have prominent grey facial discs and distinctly developed ear tufts. When disturbed, they will freeze upright in place with the ear tufts raised and the eyes almost closed; this, together with their broken brown or grey coloration, allows them to pass as an unremarkable piece of the tree they are sitting on. The most distinctive characteristic of the genus is their song, produced by members of both sexes, which comprises a rapid trill of several closely placed notes. The exact pattern of the song varies from species to species, and is commonly the most reliable method of telling each species apart (though, just for the sake of perversity, individuals that live in sympatry with members of another species may engage in mimicry). The species are otherwise often difficult to distinguish by external features alone. Just to confuse matters further, many (but not all) species of screech owl exhibit distinct colour morphs, with one morph being predominantly grey and the other rufous, that might be mistaken by the unwary for distinct species.

Pair of tropical screech owls Megascops choliba exhibiting distinct colour morphs, from Nucleo de Fauna.

As is not uncommon for owls, the majority of Megascops species are poorly known from a natural history perspective. Gehlbach & Stoleson (2010) provided a review of one of the North American species, the western screech-owl Megascops kenniscottii, that is probably typical of the genus. The diet of screech owls (like other small owls) is dominated by insects, with small vertebrates making up less than a fifth of their regular daily intake. Nesting takes place in holes in trees; nesting holes are claimed and defended by the male, who advertises its (and his) availability to females via the medium of song. Females respond to the males' advertisements by singing in reply, and pairs of owls will also respond defensively to songs from other owls nearby. The series of calls and responses in densely populated regions may develop into a full chorus of acknowledgements and challenges. Incubation of the clutch of about four eggs (which in the western screech owl is usually laid around late March–early April) is done solely by the female with the eggs taking about a month to hatch. The male collects food for her and the newly hatched chicks; after the chicks begin to fledge, both parents hunt for food for them.

The majority of screech owl species are not considered threatened conservation-wise though some of the more localised species may be vulnerable to habitat loss. Screech owls are reasonably tolerant of human activity and will even live and nest in suburban regions. They usually only disappear from a region once it becomes completely urbanised.


Dantas, S. M., J. D. Weckstein, J. M. Bates, N. K. Krabbe, C. D. Cadena, M. B. Robbins, E. Valderrama & A. Aleixo. 2016. Molecular systematics of the new world screech-owls (Megascops: Aves, Strigidae): biogeographic and taxonomic implications. Molecular Phylogenetics and Evolution 94: 626–634.

Gehlbach, F. R., & S. H. Stoleson. 2010. Western screech-owl (Megascops kennicottii). In: Cartron, J.-L. (ed.) Raptors of New Mexico pp. 511–523. University of New Mexico Press: Albuquerque.

Hastocularis: A Fossil Harvestmen Allows Us to See

Sometimes the fossil record just gives us a gift, something that moves our understanding to an all-new level. One such gift saw publication a couple of years ago, but unfortunately I didn't have time to write about it then. I think it's about time I corrected that lacuna.

Reconstruction of Hastocularis argus, from Garwood et al. (2014).

By this point in time, we have a pretty good understanding of the basal framework of harvestmen evolution. The mite-like harvestmen of the Cyphophthalmi are well established as the sister group to all other Opiliones (which form a clade called the Phalangida). Unique features of the Phalangida include an intromittent penis in the males (phalangids are one of the few groups of arachnids to possess such a feature) and a central eyemound with a single pair of eyes. The Cyphophthalmi are more heavily armoured than most phalangids, and have a characteristic pair of raised cones (the ozophores) on either side of the carapace near the front that support the openings of odour-producing repugnatorial glands. Until recently, it was thought that most Cyphophthalmi lack eyes, but tiny, lens-less remnant eyes are now known to be present at the base of the ozophores in many cyphophthalmid subgroups.

There had long been questions about the nature of the cyphophthalmid eyes. The original arachnids possessed multiple pairs of eyes, and there is a good case to be made that the basal arrangement for arachnids as a whole is a single pair of larger median eyes in the middle of the carapace, and a number of pairs (up to three) of smaller lateral eyes at the margin. In some arachnid groups the median eyes have been lost; in others, the lateral eyes have become reduced in number or lost. In spiders, the lateral eyes have become enlarged and shifted about so the lateral/median distinction is less applicable (for the record, the posterior median eyes in spiders correspond to the original median eyes). Mites, of course, being mites, mess the whole system up entirely. Most mite eyes correspond to the original lateral eyes, but some mites possess a single median eye whose relation to the original arachnid median eye pair is up for grabs.

Phalangids, with their single central eyemound and single pair of eyes, had obviously kept the original median eyes and lost the lateral eyes. But what had happened with the Cyphophthalmi? Did their single pair of eyes near the edge of the carapace represent a single remnant pair of lateral eyes, or did they correspond to the median eyes of other Opiliones? It should be noted that some derived groups of undoubted Phalangida have lost the eyemound and have their eyes sitting directly on the carapace, and in some cases these unraised eyes may be widely separated. Arguments for both interpretations of cyphophthalmid eyes had been put forward by different authors, but the matter had certainly not been decided.

A representative member of Phalangida, Platybunus pinetorum, showing the central eyemound, from Opiliophilia.

That was until the description by Garwood et al. (2014) of Hastocularis argus, a remarkably preserved fossl harvestman from the Carboniferous of France. The appearance of this animal was established in some detail by the use of microtomography, allowing a number of details about it to be established. It was a heavily armoured animal with long legs, and like modern Phalangida it possessed a central eyemound on which there had been a pair of eyes (the eyes themselves were not preserved, but the sockets that had originally contained them were). The use of microtomography also allowed the identification of an intromittent penis like a phalangid. But Hastocularis also possessed a pair of raised ozophores like modern Cyphophthalmi, and at the base of those was preserved another socket indicating the presence of a second pair of eyes. There really could not be a more perfect answer to the cyphophthalmid eye question: the immediate ancestor of the Opiliones possessed two pairs of eyes, and the eyes of Cyphophthalmi do indeed correspond to the lateral eyes of other non-harvestmen arachnids and not to the median eyes of phalangids*.

*Pedantically speaking, Hastocularis is not the first four-eyed taxon assigned to the Opiliones. In 1875, an Austrian biologist by the name of Stecker described a remarkable animal from the Sudeten Mountains of Bohemia under the name of Gibbocellum sudeticum. Gibbocellum bore an overall resemblance to the Cyphophthalmi, except for possessing two pairs of eyes on raised cones, as well as two pairs of spiracles (other Opiliones possess a single pair). Despite enthusiastic searches, no other naturalist was ever able to find further specimens of Stecker's species, and at least one author suggested that it might be a poorly interpreted pseudoscorpion. However, a close criticism of various irregularities in Stecker's publications on Gibbocellum eventually lead Hansen & Sørensen (1904) to the conclusion that it had not merely been misrepresented, but was in fact a complete fabrication on that author's part.

A phylogenetic analysis of Hastocularis lead Garwood et al. (2014) to believe that it was more closely related to Cyphophthalmi than to Phalangida; together with another Carboniferous fossil species, Eophalangium sheari, they placed it within a new taxon Tetrophthalmi (meaning, of course, 'four eyes'). The main features cited in support of this relationship were the complete fusion of the dorsal surface (the only other harvestmen to show this feature are a southeast Asian family, the Oncopodidae, who are too deeply nested within the Phalangida to be a likely direct relative of Hastocularis) and the genital opening being a broadly open gonostome (in Phalangida, the genital opening is covered by an operculum). This implies that the immediate ancestor of all Opiliones was relatively long-legged, with the short legs of Cyphophthalmi a derived feature. However, I personally find the presence of an intromittent penis in Tetrophthalmi (it is also known to be present in Eophalangium) somewhat problematic in this regard. As noted above, the phalangid intromittent penis that directly injects sperm into the female ovipositor is highly unusual among arachnids. Cyphophthalmi do possess a penis-like structure (called the spermatopositor) but it is much shorter than in any phalangid and does not function as an intromittent organ. Instead, Cyphophthalmi males produce an encapsulated spermatophore that is attached by the spermatopositor to the female's underside, a more typical sort of arrangement for arachnids as a whole. An intromittent penis in the cyphophthalmid stem group would imply that Cyphophthalmi somehow reverted towards a more primitive-seeming reproductive arrangement at some point in the past. One possibility is that the penis of Tetrophthalmi did not function in exactly the same manner as that of Phalangida: perhaps tetrophthalmids still produced a spermatophore but were able to insert it more deeply in the female than Cyphophthalmi? Another possibility may be that Tetrophthalmi are stem-phalangids rather than stem-cyphophthalmids; only further analyses can possibly tell us more.


Garwood, R. J., P. P. Sharma, J. A. Dunlop & G. Giribet. 2014. A Paleozoic stem group to mite harvestmen revealed through integration of phylogenetics and development. Current Biology 24: 1017–1023.

Hansen, H. J., & W. Sørensen. 1904. On Two Orders of Arachnida: Opiliones, especially the suborder Cyphophthalmi, and Ricinulei, namely the family Cryptostemmatoidae. University Press: Cambridge.