Field of Science


The concept of ranks in taxonomy is ultimately an arbitrary one. There is no real definition of what constitutes an 'order', a 'family' or a 'subfamily'. What determines the rank that a given taxon is recognised at is a combination of tradition, convenience, and the taxon's relationships to other recognised taxa. As such, the question of whether a given classification is overly 'split' or 'lumped' is a meaningless one and arguing the point is a complete waste of time. That said, the classification of the 'higher' oribatid mites is massively oversplit.

A big part of the reason why oribatid classification seems such a mess, with large numbers of small families containing only a handful of genera and/or species apiece, can be attributed to simple ignorance. We simply do not have a good handle on how many oribatid taxa are related to each other and as a result we find ourselves with a great many orphan taxa still hunting for a good home. The Caloppiidae may be regarded as one such taxon.

Dorsal view of Luissubiasia microporosa, from Ermilov (2016). Scale bar = 100 µm; labels with 'A' indicate areae porosae.

Caloppiids are a pantropical group of about thirty species of poronotic oribatids (the group of oribatids exhibiting the octotaxic system, an arrangement of glandular openings on the notogaster), with three genera recognised in the family by Ermilov (2016): Zetorchella, Brassiella and Luissubiasia. Zetorchella, which includes the majority of the family's species, is also pantropical in distribution. Brassiella is known from the Indo-Pacific region and Liussubiasia is known from a single species from Cuba. Past authors have often referred to Zetorchella and the Caloppiidae by the names Chaunoproctus and Chaunoproctidae, respectively, but as the name Chaunoproctus had already had dibs called on it before the mite was named (by a bird, the now-extinct Bonin grosbeak Chaunoproctus ferreorostris), their respective most senior synonyms have to take over. Caloppiids are more or less egg-shaped in dorsal view. They lack the distinct pteromorphs of most other poronotics though they may have quadrangular projections in the humeral region (the 'shoulders'). The integument is usually heavily sculpted and foveate. The legs end in three claws apiece. The most characteristic feature of the group is that the openings of the octotaxic system on the notogaster, of which five pairs are present, are extremely small. The octotaxic system can take two forms, recessed saccules or porose patches. Those of caloppiids have usually been described as saccules but Ermilov (2016) states that, at least in some species, they are very small porose areas.

Going by their overall appearance, caloppiids are classified within the superfamily Oripodoidea. However, one of the most characteristic features of the Oripodoidea as an evolutionary group is that their nymphs have notogastral setae borne on individual off-centred sclerites (oribatid nymphs often look very different from their adults and are often more soft-bodied). At this point in time, we simply do not know what the nymphs of caloppiids look like so we cannot say whether they possess this crucial feature. Conversely, with their lack of pteromorphs, caloppiids bear a distinct similarity to the more diverse oripodoid family Oribatulidae. The two families have mostly been separated on the basis of caloppiids supposedly having an octotaxic system of saccules rather than porose areas, a distinction that I've already noted may not hold up. There's also something of an open question whether the distinction between saccules and porose areas is really as significant as it has been thought in the past. So, at present, we can't say with confidence whether caloppiids are true oripodoids... or whether they are not only oripodoids but don't even warrant recognition as a distinct family from oribatulids.


Ermilov, S. G. 2016. Luissubiasia microporosa gen. nov., sp. nov. (Acari, Oribatida, Caloppiidae) from Cuba. International Journal of Acarology 42 (2): 127–134.

The Grisons

Spend a bit of time following discussions of nature documentaries and other popular representations of biodiversity, and one topic you're likely to see come up is the biases that tend to exist in what gets represented. Images from eastern and southern Africa predominate while the west and north of that continent get overlooked. Europe and North America receive much more attention than the temperate regions of Asia. Another region whose diversity tends to go underrepresented is South America. The casual observer might think this continent is all monkeys and jaguars but South America is also home to notable radiations of dogs, deer, rodents, and other animals that many people would associate more with other parts of the world. Among these overlooked elements of the South American fauna are the local species of mustelid, including the grisons of the genus Galictis.

Greater grison Galictis vittata, copyright Tony Hisgett.

Grisons are somewhat ferret- or skunk-like animals found across almost the entirety of South America, and north into southern Mexico. They are greyish in colour dorsally (the name 'grison' itself means 'grey') with a black face and underparts. A pale stripe separates the upper and lower parts across the top of the face and continues diagonally back to the shoulders. They feed on small vertebrates and tend to be solitary hunters though they may sometimes form small family groups. They are primarily terrestrial and diurnal in habits. They have a reputation for ferocity; residents of Chile apparently have a history of using comparisons to grisons to describe unchecked rage (Yensen & Tarifa 2003b), in a similar manner to references to wolverines and honey badgers in other parts of the world. Contrasting colour patterns like those of the grisons are associated in other musteloids (such as skunks) with the production of offensive odours for defence, and grisons also produce strong-smelling secretions from their anal glands. Though some sources have claimed the odour produced by the lesser grison to be worse than a skunk's, it appears that these reports are exaggerated (Yensen & Tarifa 2003b).

Lesser grison Galictis cuja, copyright Ken Erickson.

Most authors have recognised two species of grison, the greater grison Galictis vittata and the lesser grison G. cuja*, as corroborated by a recent taxonomic study of the genus by Bornholdt et al. (2013). As their names indicate, the greater grison is generally larger and more robust than the lesser, being about 60 to 76 cm in total length versus 44 to 68 cm for the lesser grison (Yensen & Tarifa 2003b). The tail is also proportionately shorter in the greater grison (30% of the total length for the greater, 40% for the shorter). Fur is relatively longer and denser in the lesser grison, giving it more of a fluffy look. Whereas the dorsal fur is always a plain grey in the greater grison, it may often have a yellowish tinge in the lesser (not always, though). The two are generally distinct in range and habitat, as well. The greater grison is an animal of tropical forests and inhabits the northern part of the genus' range in Central America and northern and western South America. The lesser grison inhabits drier habitats, in arid or temperate regions, and so occupies the southern and eastern parts of the continent. The ranges of the species are known to overlap in Bolivian and Paraguay where their respective biomes approach each other.

*Some sources have listed a third species G. allamandi but this seems have been something of a 'ghost' taxon born from confusion whether the name 'G. vittata' applied to the greater or lesser species.

The genus Galictis arrived in South America as part of the Great American Biotic Interchange, about three million years ago. The general consensus is that it is derived from the genus Trigonictis of the North American Pliocene. Indeed, it has even been suggested that the two North American species of Trigonictis might represent independent ancestors of Galictis, with the larger T. macrodon giving rise to the greater grison and the smaller T. cookii birthing the lesser grison (Yensen & Tarifa 2003a). This certainly would seem overly complicated, though, and molecular data are more in line with a more recent separation of the species.


Bornholdt, R., K. Helgen, K.-P. Koepfli, L. Oliveira, M. Lucherini & E. Eizirik. 2013. Taxonomic revision of the genus Galictis (Carnivora: Mustelidae): species delimitation, morphological diagnosis, and refined mapping of geographical distribution. Zoological Journal of the Linnean Society 167: 449–472.

Yensen, E., & T. Tarifa. 2003a. Galictis vittata. Mammalian Species 727: 1–8.

Yensen, E., & T. Tarifa. 2003b. Galictis cuja. Mammalian Species 728: 1–8.

Nocardia pseudovaccinii

As noted on this site before, the Actinobacteria are one of the most significant groups of bacteria in the terrestrial environment. Among the more diverse genera of Actinobacteria is Nocardia, members of which produce fine, branching mycelia that often fragment into individual rod-shaped or coccoid segments, each of which is capable of developing into a new mycelium (Goodfellow et al. 2012). As is the way of things, Nocardia species are usually soil dwellers but are more commonly studied as facultative pathogens. Nevertheless, recent years have seen the recognition of an increasing number of species isolated from soil samples with one such species being Nocardia pseudovaccinii.

Nocardia pseudovaccinii was described as a new species by Kim et al. (2002). In culture, N. pseudovaccinii grows a beige-red substrate mycelium supported a scarce, white aerial mycelium. Kim et al. (2002) identified the species as able to utilise a wide range of organic substrates such as ribose and glucosaminic acid though it could not break down others such as sucrose or citrate. Molecular analyses of Nocardia in Kim et al. (2002) and Goodfellow et al. (2012) do not really indicate a clear association of N. pseudovaccinii with any other species. Bacterial systematists apparently still maintain that neighbour-joining analyses are something more than a complete waste of time. I do not support this view.

The strains assigned to N. pseudovaccinii by Kim et al. (2002) had previously been identified as another species, N. vaccinii, hence the new species' name ('vaccinii', in case you were wondering, has no direct connection to vaccines but refers to Vaccinium, the plant genus including blueberries and from which N. vaccinii was first isolated). Nocardia vaccinii has been known to act as a facultative plant pathogen but I am not aware of this role being identified for N. pseudovaccinii. The original isolates were cultured from soil (though Kim et al. say nothing about what kind of soil or even where it was sampled). Nocardia pseudovaccinii has also been found forming part of the microbiome of wireworms of the genus Agriotes, beetle larvae that feed on plant roots. It may form a symbiotic association with these grubs that provides the latter with antibiotic protection from the pathogenic fungus Metarhizium brunneum (Kabaluk et al. 2017). A good thing for the wireworms but perhaps not so good for agriculturists who would like to keep them under control.

Goodfellow, M., P. Kämpfer, H.-J. Busse, M. E. Trujillo, K. Suzuki, W. Ludwig & W. B. Whitman (eds) 2012. Bergey's Manual of Systematic Bacteriology 2nd ed. vol. 5. The Actinobacteria, Part A and B. Springer.

Kabaluk, T., E. Li-Leger & S. Nam. 2017. Metarhizium brunneum—an enzootic wireworm disease and evidence for its suppression by bacterial symbionts. Journal of Invertebrate Pathology 150: 82–87.

Kim, K. K., A. Roth, S. Andrees, S. T. Lee & R. M. Kroppenstedt. 2002. Nocardia pseudovaccinii sp. nov. International Journal of Systematic and Evolutionary Microbiology 52: 1825–1829.


I have referred in the past to there being something of a divide in approaches to the classification of the Foraminifera. This divide arises from disagreements such as the relative significance of various character complexes. One taxon that stands as an example of such disagreements is the subject of this post, the family Pyrgoidae as recognised by Mikhalevich (2005).

Pyrgo williamsoni, copyright Michael.

Pyrgoids are members of the group of forams generally recognised as the Miliolida, the porcelaneous forams. In this group, the wall of the test is composed of calcite but the calcite crystals are not regularly lined up with each other so the wall is not transparent. As a result, the wall of the test resembles porcelain in appearance. Most miliolidans have the chambers of the test coiling in a single plane. The Pyrgoidae were distinguished from other miliolidans by Mikhalevich (2005) by the overall structure of the test which is primarily biloculine (with the whorls of the test composed of two chambers). The family was divided into subfamilies by the nature of the test aperture: single with an inner tooth in Pyrgoinae, single with a flap in Biloculinellinae, and multiple (at least when mature) in Cribropyrgoinae and Idalininae. Idalininae also differed from other subfamilies in that the very last chamber was further enlarged to envelop the entire test. Members of the Pyrgoidae are known from the fossil record going back to the Jurassic period.

In the system of Loeblich & Tappan (1964), however, the pyrgoids were not recognised as a single group. Instead, they were dispersed among separate subfamilies of the family Miliolidae. Part of the reason was simply that Loeblich & Tappan did not divide the miliolidan families as finely as Mikhalevich later would but a bigger difference was one of priority. Loeblich & Tappan regarded the nature as an aperture as a more important feature taxonomically than the arrangement of chambers. Both classifications seem to have been constructed more from a diagnostic viewpoint than necessarily intended to reflect phylogenetic relationships.

Cribropyrgo aspergillum, from the National Museum of Natural History.

As with most other forams, pyrgoids exist in what are called megalosphaeric and microsphaeric forms. These forms represent alternate generations in the foram life cycle: microsphaeric forams are the sexually reproducing generation whereas megalosphaeric forams reproduce asexually. The names refer not to the overall size of the individuals but to the size of the proloculus, the very first embryonic chamber that sits at the center of the test. In megalosphaeric pyrgoids, the developing test is biloculine from the very start. In microsphaeric individuals, the earliest stages of the test are quinqueloculine (with five chambers per whorl) then become triloculine then finally biloculine (with a further progression for the idalinines, of course). The significance of the differences between the two forms has historically been the subject of discussion with some authors arguing that the microsphaeric forms represented a retention and overwriting of ancestral forms, or an expression of the trajectory the lineage might evolve along in the future (Loeblich & Tappan 1964). The most likely explanation, though, seems to me to be the simplest. The size of the proloculus correlates with the amount of cytoplasm in the young foram. Megalosphaeric pyrgoids start with fewer chambers per volution from the start for the simple reason that they don't have the space to pack in more.


Loeblich, A. R., Jr, & H. Tappan. 1964. Treatise on Invertebrate Paleontology pt C. Protista 2. Sarcodina: chiefly "thecamoebians" and Foraminiferida vol. 1. The Geological Society of America, and The University of Kansas Press.

Mikhalevich, V. 2005. The new system of the superfamily Quinqueloculinoidea Cushman, 1917 (Foraminifera). Acta Palaeontologica Romaniae 5: 303–310.

Donaldina: Palaeozoic Turrets

Within the last few decades, we've developed a reasonably good idea of what are the primary subdivisions of gastropods alive today. One such generally accepted lineage is the Heterobranchia (or, depending on the author, the Heterostropha), a group that includes (among others) the air-breathing pulmonates as well as the marine sea slugs and bubble shells. In the fossil record, the roots of this lineage extend well back into the Palaeozoic with early members recognisable by their distinctive mode of shell development. The larval shell, the protoconch, of these forms spirals in the opposite direction from the mature teleoconch, so the animal will start its life sinistral (spiralling left) and end it dextral (spiralling right; if you're having difficulty imagining how this works, the protoconch often ends up sitting upside down relative to the teleoconch). Among the earliest heterobranchs in the fossil record is the genus Donaldina.

Specimen of Donaldina (1.2 mm in height), with close-up of protoconch, from Bandel et al. (2002).

Fossils of Donaldina have been found around the world and the genus persisted for a long time. The earliest potential Donaldina have been described from the Early Devonian but their inclusion in the genus is uncertain (Bandel et al. 2002). The protoconch on these early forms is poorly preserved and it is uncertain whether they truly showed a heterobranch development. The genus was definitely present by the early Carboniferous and persisted into the lower Permian. This is an impressive length of time: the Carboniferous alone last for around sixty million years.

Donaldina was a genus of small, high-spired gastropods, less than a centimetre in height. Many early members of the Caenogastropoda, the likely sister group of the Heterobranchia, also had shells of this kind and it may have represented the ancestral form for the two lineages. The sinistral protoconch of Donaldina was almost planispiral (spiralling in a flat plane) and completed between one and two whorls. The multi-whorled, dextral teleoconch was characterised by an ornament of spiral cords, usually only on the lower half of the whorl.

So what were Donaldina doing with their time when alive? Modern high-spired gastropods occupy a range of lifestyles, including free-living grazers, burrowers, or sedentary forms that live as filter feeders or parasites of other animals (Signor 1982). The morphology of Donaldina suggests that it is unlikely to be a burrower. The whorls are individually rounded whereas those of habitual burrowers tend to be flattened so the shell moves more smoothly through the sediment. The ornamentation on the underside of the whorl would presumably also have presented resistance to burrowing. The shape of the aperture in Donaldina is more suggestive of a free roamer, as a sinus in the upper part of the outer margin would have allowed the animal to pull back into its shell while the plane of the aperture was held as flat as possible against the substrate to protect against predators. Overall, the lifestyle of Donaldina may not have been dissimilar to that of the modern mudsnails of Cerithium and similar genera, crawling about in search of algae and other tasty morsels.


Bandel, K., A. Nützel & T. E. Yancey. 2002. Larval shells and shell microstructures of exceptionally well-preserved Late Carboniferous gastropods from the Buckhorn Asphalt Deposit (Oklahoma, USA). Senckenbergiana Lethaea 82 (2): 639–689.

Signor, P. W., III. 1982. Resolution of life habits using multiple morphologic criteria: shell form and life-mode in turritelliform gastropods. Paleobiology 8 (4): 378–388.

Mooching Off the Relatives

Something I've referred to before but only (I think) in passing is that, among the enormous diversity of bees that inhabit this world, there are a large number of species that act as cleptoparasites*. That is, instead of constructing and provisioning their own nests, they lay their eggs in the nests of other bee species. When the eggs hatch, the emerging larvae feed on the provisions that the constructing bee intended for her own offspring. One lineage of these cleptoparasites is the megachilid genus Coelioxys.

*Depending on the source, you may see this term spelt as either 'cleptoparasite' or 'kleptoparasite'. Personally, I've never been able to decide just which I should be using.

Coelioxys sodalis, copyright jgibbs.

Coelioxys is a diverse, cosmopolitan genus with nearly 500 known species, closely related to the even more diverse leafcutter bees and resin bees of the genus Megachile. Species vary in size from half a centimetre to nearly an inch in length. They are fairly similar to species of Megachile in overall appearance, the most obvious difference being that (as with most cleptoparasitic bees) their covering of hair is greatly reduced. In particular, the dense scopa of hairs that covers the underside of the metasoma in female Megachile is absent. The primary function of the hairs in bees is to carry pollen; with no nest of their own to worry about, cleptoparasitic bees have no need for such dense hairs. Coelioxys females also differ from Megachile in the shape of the metasoma which is tapering and ends in a narrow tip. More on that in a moment.

As might be expected for such a large genus, Coelioxys has been divided between a number of subgenera. Until recently, the definitions of a number of these subgenera was somewhat uncertain. The biggest problem was that most revisions of the genus had been done on a regional level so (for instance) North American taxa were more finely subdivided than in the Old World. However, a recent phylogenetic analysis of the genus by da Rocha Filho & Packer (2017) redefined a number of subgenera and adjusted their definitions. For instance, the type subgenus Coelioxys, recognised as subcosmopolitan by Michener (2007), became restricted to just two species, the European C. quadridentata and the North American C. sodalis. Whereas Michener's concept of Coelioxys was essentially recognised by lacking the specialised features of other subgenera, the restricted Coelioxys sensu stricto can be recognised by having the outer margin of the pronotal lobe conspicuously rounded, as well as having the pilosity on the mesosoma suberect, long and thin, without spots of appressed hairs (da Rocha Filho & Packer 2017).

For the most part, Coelioxys species are cleptoparasites of Megachile though some have also been found mooching off Apidae species. Coelioxys quadridentata, for instance, has been found in association with nests of both Megachile and Anthophora. In most cases, a female Coelioxys will lay into a host nest before it is closed, while the constructor is away foraging for supplies. The narrow metasoma allows the Coelioxys to reach into the cavity containing the nest and insert her eggs into the nest wall where the host will not notice it. Often, multiple eggs will be laid in a single nest. After the nest is closed, the eggs hatch into larvae that look fairly unremarkable for their first one or two instars: like other bee larvae, not doing much more than sit there and eat. But upon reaching the second or third instar, the Coelioxys larva develops greatly enlarged mandibles that it uses to stir through the nest's food mass and execute any other larvae and eggs contained therein. Both the original host larva and any other Coelioxys larvae the nest may contain are dealt with in this manner (presumably the process of finding an appropriate host nest is difficult enough that the waste of eggs is still worth it for the parent Coelioxys to increase the chance that at least one reaches maturity). Its competitors thus removed, the larva them moults back to a more average form with nothing more agin to do but eat until the time to mature is reached.


Michener, C. D. 2007. The Bees of the World 2nd ed. John Hopkins University Press: Baltimore.

Rocha Filho, L. C. da, & L. Packer. 2017. Phylogeny of the cleptoparasitic Megachilini genera Coelioxys and Radoszkowskiana, with the description of six new subgenera in Coelioxys (Hymenoptera: Megachilidae). Zoological Journal of the Linnean Society 180: 354–413.