Field of Science

Gender's Just a State of Gonads

It didn't take long for Adam Yates to recognise this animal:

Juvenile Pagrus auratus. Photo by Richard Ling.

This is the fish that goes by the name of 'snapper' in New Zealand, though that name is used for different kinds of fish elsewhere. In older references, you'll find this species under the name of Chrysophrys auratus, but the genera Chrysophrys has since been synonymised with Pagrus (Paulin, 1990). However, the molecular phylogenetic analysis of Chiba et al. (2009) failed to recover monophyly for Pagrus, so we may yet see Chrysophrys make a comeback some day.

Mature individuals of Pagrus major, a north-west Pacific species regarded by some authors as a synonym of P. auratus. These two are probably engaging in courtship behaviour. Photo from here.

As this young snapper gets older, its body will change in numerous ways. One is that the blue spots along its side will fade away and it'll become more evenly pink. Its head will become deeper, and if it may develop a large supraorbital boss on its forehead. And one other significant change that it may go through is a reassignment of gender. Members of the marine fish family Sparidae, to which Pagrus belongs, show a bewildering range of sexual development, including forms which show protandrous hermaphroditism (they start life as males before developing into females), protogynous hermaphroditism (starting as females, developing into males) and gonochorism (completely separate males and females, as we have ourselves). Other species start life with the rudiments of both male and female gonads but have only one or the other develop to maturity, without any subsequent sex changes, while a single species has been recorded as possessing simultaneously functional gonads of both sexes (Buxton & Garratt, 1990).

Different species of sparids feed on a variety of different diets, from predators of other fish such as the Dentex species to herbivores on algae such as Sarpa salpa. This variation in diet is reflected in a variety of dental morphologies. Predators such as Dentex possess pointed caniniform teeth while invertebrate feeders such as Pagrus auratus have a combination of pointed teeth in the front and round molariform teeth in the back. Algal feeders have flat-topped incisiform dentition, leading to occassional reports on fish with human teeth:

Teeth of sheepshead, Archosargus probatocephalus. Photo from Nathan Thurston.

In the past, dentition has been used as the basis for dividing sparids into a number of subfamilies, but both molecular (Chiba et al., 2009) and morphological (Day, 2002) analyses indicate multiple polyphyletic origins of the various dentition types. Contrast that to the situation in the possibly related* family Lethrinidae where trophic type and phylogeny show a much closer fit.

*A relationship between the two has been suggested on morphological grounds; molecular analyses have so far not supported such a relationship, but nor have they produced any strong relationships for either family.


Buxton, C. D., & P. A. Garratt. 1990. Alternative reproductive styles in seabreams (Pisces: Sparidae). Environmental Biology of Fishes 28: 113-124.

Chiba, S. N., Y. Iwatsuki, T. Yoshino & N. Hanzawa. 2009. Comprehensive phylogeny of the family Sparidae (Perciformes: Teleostei) inferred from mitochondrial gene analyses. Genes and Genetic Systems 84 (2): 153-170.

Day, J. J. 2002. Phylogenetic relationships of the Sparidae (Teleostei: Percoidei) and implications for convergent trophic evolution. Biological Journal of the Linnean Society 76 (2): 269-301.

Paulin, C. D. 1990. Pagrus auratus, a new combination for the species known as "snapper" in Australasian waters (Pisces: Sparidae). New Zealand Journal of Marine and Freshwater Research 24: 259-265.

Name the bug # 26

I haven't put up any posts earlier this week as I was at the Australian Entomological Society conference, but now it's back to work. Tomorrow's post will be related to this animal:

Attribution to follow.

Update: Identity now available here. Photo from here.

The Trouble with Coelosclerites

A couple of years ago, I posted a brief review of the chancelloriids, mysterious sessile animals from the Cambrian period. As explained in that post (which I'd recommend reading before this one), chancelloriids are remarkable for how much we know about them while still being unable to place them anywhere in the animal family tree. However, there are two main options that are currently supported: one is that chancelloriids are sponge-grade animals, probably in the stem-group of modern Epitheliozoa (the clade including coelenterates and bilaterians, which differ from sponges in having a differentiated external skin around their bodies); the other is that chancelloriids are related to other Cambrian animals such as halkieriids, which have themselves been shown to be closely related to molluscs (Vinther & Nielsen, 2005).

Diagram of coelosclerite microstructure from Porter (2008).

The sponge interpretation of chancelloriids has some strong points marshalling in its favour: chancelloriids lack any sign of bilateral symmetry and no sign has been recognised in them of differentiated organ systems. The main feature associating them with halkieriids is the microstructure of their sclerites. Chancelloriids and halkieriids (and a couple of other Cambrian families) possessed sclerites with a microstructure unknown for any other animal group. Known as coelosclerites, these structures were hollow and would have been secreted as a single unit without any subsequent growth*. The greater part of the sclerite was formed of aragonite fibres, arranged parallel to the axis of the sclerite. External protrusions on the sclerite were formed by aragonite bundles sitting at an angle to the main body. A thin layer, probably originally organic, covered the outer surface of the sclerite (Porter, 2008).

*But see Jakob Vinther's comment on the earlier post.

Porter (2008) felt that the similarity between chancelloriid and halkieriid sclerites was so great that it was unlikely that they had evolved independently. The coelosclerite was far from being the only way to develop such a structure: the Cambrian and subsequent periods have seen the evolution of many other sclerite-possessing animals, all of which exhibited different sclerite microstructures. Nor could any convergence be explained by selective pressures: the sessile chancelloriids and slug- or chiton-like halkieriids would have ecologically very different animals. If the coelosclerite structure arose independently in the two groups, the similarities would have to be accepted as pure coincidence.

However, if we accept that coelosclerites had a single origin, we have to explain the complete absence of apparent bilaterian traits in chancelloriids. Many groups of bilaterians have lost their ancestral bilateral symmetry: tunicates, entoprocts, echinoderms, for instance. None of them, however, have lost all trace of their ancestry to quite the same degree that chancelloriids would have had to. Porter (2008) proposed two options: (1) chancelloriids were indeed highly derived bilaterians forming a clade with halkieriids, or (2) chancelloriids were sponge-grade stem-epitheliozoans; coelosclerites arose in the common ancestor of chancelloriids and bilaterians but were subsequently lost by bilaterians other than halkieriids.

Option 2 might appear tempting if halkieriids were close to the base of bilaterians, but it is well-established that they are not. If halkieriids are interpreted as stem-trochozoans (a fairly conservative interpretation) then coelosclerites would have had to have been lost at least six times, in the ancestors of ctenophores, cnidarians, deuterostomes, ecdysozoans, bryozoans and platyzoans (and that is ignoring more phylogenetically contentious groups such as acoelomorphs and chaetognaths that could potentially increase the number even further). If, as seems more likely, halkieriids are stem-molluscs, we have to factor in another two losses for brachiozoans and annelids (and, again, I'm ignoring phylogenetic renegades such as entoprocts). And in the case of brachiozoans, the greater part of the brachiozoan stem group appear to have themselves possessed sclerites; Porter's hypothesis 2 would require the stem-brachiozoans to have lost coelosclerites, only to re-evolve a distinct new sclerite form shortly afterwards.

So, in my opinion, the only really viable options are coelosclerites evolved convergently in two entirely separate lineages, or coelosclerite-possessing animals formed a single monophyletic clade. My personal inclination would be to favour the latter; the examples of ascidians and others demonstrate that significant re-organisations of the bilaterian body plan are not a priori impossible. Of course, the supporting evidence either way remains shaky, and the whole structure could still come tumbling down tomorrow.


Porter, S. M. 2008. Skeletal microstructure indicates chancelloriids and halkieriids are closely related. Palaeontology 51 (4): 865-879.

Vinther, J., & C. Nielsen. 2005. The Early Cambrian Halkieria is a mollusc. Zoologica Scripta 34: 81-89.

Freak of the Week: Wingless, Legless Flies

I found this while looking up identification info for phorid flies, of which we currently seem to be receiving something of an influx in our samples.

Photoes from here.

Most of what you see in the lower of the two photoes above are larvae of army ants of the genus Aenictus. The odd one out is the whiter 'larva' in the centre—which is not a larva at all, but a fully adult female of the phorid fly Vestigipoda longiseta! (The upper photo shows the same animal in close-up.) This bizarre animal makes its living by imitating its host larvae and being fed by the larvae's deluded carers. Five species of Vestigipoda have been described to date from Malaysia (Disney et al., 1998; Murayama et al., 2008).

Close-up of head of Vestigipoda maschwitzi, from Disney et al. (1998).

Cases of neoteny, where insects develop full sexual maturity while still 'larvae', are not unknown among holometabolous insects (I earlier described a case involving the beetle Micromalthus). However, Vestigipoda cannot be regarded as neotenous because the female has a fully developed adult head.

So far, Vestigipoda seems to only be known from females. It is possible that males, when found, may turn out to be much more normal phorid flies. The challenge would be recognising them as related to their bizarre females.


Disney, R. H. L., A. Weissflog & U. Maschwitz. 1998. A second species of legless scuttle fly (Diptera: Phoridae) associated with ants (Hymenoptera: Formicidae). Journal of Zoology 246 (3): 269-274.

Maruyama, M., R. H. L. Disney & R. Hashim. 2008. Three new species of legless, wingless scuttle flies (Diptera: Phoridae) associated with army ants (Hymenoptera: Formicidae) in Malaysia. Sociobiology 52 (3): 485-496.

A Little Bit of Gastrocopta

Gastrocopta armigerella, from Okinawa in Japan. Photo by Hiroshi Ishikawa.

Gastrocopta is a genus of terrestrial snail found pretty much throughout the world, mostly in drier habitats. Despite its abundance, it is not hugely familiar to most people by virtue of the fact that most species are only a couple of millimetres in size. Shells of Gastrocopta species are whitish and somewhat translucent, and have a number of distinct projections around the shell aperture. These projections may function for protection or they may help to reduce water loss: both important functions for a tiny snail.

Gastrocopta is a member of the superfamily Pupilloidea. All pupilloids are small, cylindrical snails, and authors have differed significantly as to how the group is divided up: some authors have recognised only a single family Pupillidae, others have recognised a number of families. The classification of Bouchet et al. (2005) (which is as good a baseline as any) includes the subfamily Gastrocoptinae in the family Vertiginidae. As well as the cosmopolitan Gastrocopta, Gastrocoptinae includes a number of genera with more restricted distributions (Pokryszko, 1996). The relationships between these genera do not appear to have been studied in detail; while I haven't found an explicit statement of such, there seems to be something of an implied suspicion that Gastrocopta in its current sense may represent a plesiomorphic paraphylum to at least some of the other gastrocoptines.

Live specimen of Gastrocopta contracta. Photo by Aydin Örstan.

There is certainly no shortage of described species for Gastrocopta, with probably at least as many species waiting to be described. However, the taxonomy of this genus is not on the firmest of grounds. Soft-anatomy characters such as genitalic features have been little studied within Gastrocopta, and have generally not shown much variation when they have been studied (Pokryszko, 1996). As such, species are distinguished by features of the shell, particularly the arrangement of teeth around the aperture. Unfortunately, at least some species have been shown to vary between individuals in these features, making species identification potentially difficult without access to multiple specimens (of course, Gastrocopta is hardly exceptional in this regard). Also, while the genus has been divided between a number of subgenera, the boundaries between these subgroups tend to be somewhat blurry and not all authors have accepted their validity.


Bouchet, P., J.-P. Rocroi, J. Frýda, B. Hausdorf, W. Ponder, Á. Valdés & A. Warén. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1-2): 1-397.

Pokryszko, B. M. 1996. The Gastrocoptinae of Australia (Gastropoda: Pulmonata: Pupilloidea): systematics, distribution and origin. Invertebrate Taxonomy 10: 1085-1150.

Name the Bug # 25

A couple of my readers should find this a fairly easy one, at least for the genus:

Attribution to follow.

Update: Identity available here. Photo from here.

Contents Complete

Just a note to let you all know that the Table of Contents for this site that I first uploaded last week now contains links to all the major posts on this site to date. I hope you find it useful!

Ants on the Move (Taxon of the Week: Dorylidae)

Dorylus male. Photo by A. Buschinger.

As expected, it did not take long for this animal to be identified as a male driver ant of the genus Dorylus, commonly known as a sausage fly due to the appearance of the elongate abdomen. African Dorylus species include the largest of all ants: the male in the photo is about 3 cm long while females can be up to 5 cm.

Army ants and driver ants are a clade of tropical ants characterised by a number of distinct behavioural and morphological characteristics, including the construction of temporary nests with at least occasional colony migrations to new nest sites and obligatory foraging in large groups. Queens are permanently wingless; both queens and workers are eyeless or possess very rudimentary eyes (the weirdness of army ant eyes was previously referred to in this post) and often exhibit a large variation in size within a colony. Males possess fully-developed eyes and wings and appear so different to the queens that Linnaeus originally described one in 1764 as a species of wasp, and the relationship between males and females was not established until Savage collected them in association in 1849 (earlier authors, noting the common proximity of males to workers, had suggested that the former might be parasitoids of the latter!) During the 1800s, army ants were commonly recognised as a separate family Dorylidae, but since then they have been included in the family Formicidae with all other ants and 'Dorylidae' has not been used as a valid taxon. The former Dorylidae is now referred to as the 'doryline' or 'dorylomorph' group (also including the related subfamily Cerapachyinae, the members of which do not show the full range of army ant characteristics) with the army ants divided between four subfamilies. This subdivision was established when it was thought for a large part of the 1900s that the subfamilies might be convergent rather than related, but in recent years both morphological and molecular studies have supported a single origin for the army ant lifestyle (Brady, 2003; Brady & Ward, 2005). Within the army ant clade, the basal division has been consistently identified as between the South American Ecitoninae and an Old World (African and Asian) clade including the genera Dorylus, Aenictus and Aenictogiton*, each currently placed in a separate monogeneric subfamily. The biology of army ants was reviewed by Gotwald (1982); I've drawn heavily on that publication for this post.

*Aenictogiton is a rather enigmatic genus, known only from the rarely collected males. Whether the species of this genus exhibit army ant lifestyles, as suggested by its apparent phylogenetic position, requires the identification of females to establish.

Workers of the South American ecitonine Labidus praedator, showing the huge variation in size between large and small workers. Photo by Alex Wild.

Army ants can be roughly divided between species that forage above ground (epigaeic forms) and those that forage subterraneously or beneath leaf litter (hypogaeic forms), though some hypogaeic species will forage epigaeically if conditions permit (usually at night). Similarly, epigaeic foragers may in turn be either epigaeic or hypogaeic nesters. For obvious reasons, the epigaeic species are the best studied with little being known about the sociobiology of most hypogaeic species; however, it is the hypogaeic forms that account for the majority of army ant species. In all army ants studied to date, foraging is done by advancing columns or networks of groups of workers, without individual exploratory scouts searching ahead for food sources. Location of food sources is therefore largely fortuitous, though advancing columns are attracted to movement (potential prey animals have been observed avoiding predation by the simple expedient of not moving). In the subterranean forager Dorylus laevigatus, smaller workers produce small exploratory tunnels, often by muscling into pre-existing cracks in the ground; once a suitable food source is discovered, the tunnels are enlarged to allow the passage of larger workers (Berghoff et al., 2002). Digging is done by workers wither carrying away soil pieces or passing them beneath their body to on-following workers (Berghoff et al., 2003). In epigaeic forms, all sizes of worker participate equally in exploration, though the largest of workers (often referred to as 'soldiers'; however, different worker sizes exist as points on a spectrum rather than as discrete classes) usually only perform defensive rather than food-gathering roles for the simple reason that their hypertrophied mandibles (which increase allometrically with body size) are largely incapable of handling food items. Smaller workers will also handle smaller food particles or may transport liquid rather than solid food sources. Most food items are small animals such as earthworms or arthropods though some species will also take plant foods such as oily fruits; despite the somewhat macabre stories sometimes told about army ants, larger animals are rarely at risk unless injury or confinement restricts their ability to move away from the foraging column. Army ants have functional stings that most use in addition to their mandibles for attacking prey or other animals disturbing the column; however, Dorylus species appear to use their mandibles only. Many African tribes have supposedly exploited the aggressiveness of driver ant soldiers for suturing cuts; the ant was held to the open wound and its head broken off after its mandibles had clamped into the patient's skin.

Queen of Eciton burchelli during the reproductive phase. When the colony becomes nomadic, her expanded abdomen will shrink down and she will become difficult to distinguish from a large worker. Photo from Arkive.

In many species, a regular transition occurs between nomadic and stationary phases in the life cycle; in others, the transition is more irregular and less defined. In those species with separate phases, the queen of a colony lays eggs during the stationary phase, her abdomen becoming massively swollen while laying. An army ant queen may produce hundreds of thousands of eggs over the course of a year (which will of course include a number of nomadic phases in those species that have them); the queen of Dorylus wilverthi has been estimated to produce from 3 to 4 million eggs per month. When the young reach the larval stage, the ants become nomadic and the queen's abdomen telescopes down in size so that she is able to move along with the rest of the colony (Dorylus queens, which do not have such clearly separated life phases, do not undergo this regular expansion and contraction). When the larvae moult into pupae, the colony returns to the stationary phase and produces a new nest. Hypogaeic nesters dig a nest into the ground; epigaeic nesters produce a bivouac between plant branches by means of larger workers linking themselves together into a living net with the queen safely held in the centre.

Bivouac of the South American Eciton burchelli. Photo from here.

Because army ant queens do not possess wings at any point in their lives, they do not perform dispersive nuptial flights in the way of many other ants. The winged males do still function to disperse genes between colonies; mature males leave their parent colony to find a suitable mate, dying after a period of a few days to a few weeks. New colonies are formed by colonial fission; if a brood contains developing queens, some workers in the colony become attached to the new queens instead of the old. Usually only the first one or two young queens to emerge from the pupa establish new colonies; once they reach maturity, they will leave their parent colony, taking those workers that have transferred their allegiances with them. The workers are unable to survive long without a queen, but if a colony should happen to lose its matriarch all may not be lost for the individual workers. If they should find another colony before they perish, they may be adopted into that colony and transfer their loyalty to a new queen. Of course, if they are carrying any larvae or eggs produced by their old queen, those will soon be consumed by the new colony.


Berghoff, S. M., J. Gadau, T. Winter, K. E. Linsenmair & U. Maschwitz. 2003. Sociobiology of hypogaeic army ants: characterization of two sympatric Dorylus species on Borneo and their colony conflicts. Insectes Sociaux 50 (2): 139-147.

Berghoff, S. M., A. Weissflog, K. E. Linsenmair, R. Hashim & U. Maschwitz. 2002. Foraging of a hypogaeic army ant: a long neglected majority. Insectes Sociaux 49 (2): 133-141.

Brady, S. G. 2003. Evolution of the army ant syndrome: The origin and long-term evolutionary stasis of a complex of behavioral and reproductive adaptations. Proceedings of the National Academy of Sciences 100 (11): 6575-6579.

Brady, S. G., & P. S. Ward. 2005. Morphological phylogeny of army ants and other dorylomorphs (Hymenoptera: Formicidae). Systematic Entomology 30: 593-618.

Gotwald, W. H., Jr. 1982. Army ants. In: Hermann, H. R. (ed) Social Insects, vol. 4 pp. 157–254. Academic Press: New York.

Name the Bug # 24

Tomorrow's post will be related to this animal:

I'm pretty sure a couple of my readers will find this an easy one to identify. Attribution, as always, to follow.

Update: Identity now available here. Photo from here.

The Attack of Mega-matrix

Lienau, E. K., R. DeSalle, M. Allard, E. W. Brown, D. Swofford, J. A. Rosenfeld, I. N. Sarkar & P. J. Planet. 2010. The mega-matrix tree of life: using genome-scale horizontal gene transfer and sequence evolution data as information about the vertical history of life. Cladistics 26: 1-11.

The last two decades have seen a great deal of discussion about the role of horizontal gene transfer (HGT) in phylogenetic reconstruction of prokaryotes. That HGT occurs among prokaryotes, occassionally between members of far distant lineage, is undeniable; the question is whether HGT is a common event in bacterial evolution or whether it is mere occasional noise. Some researchers have gone so far as to argue that HGT is so rampant among prokaryotes that the reconstruction of a reliable tree of life for bacteria is an impossibility. As I've noted elsewhere, I have to admit to a certain degree of hostility towards this idea, but I immediately have to confess that this hostility is entirely due to personal prejudice (I really want there to be a tree of life for prokaryotes) and not supported by anything rational.

Attempts to reconstruct the prokaryote tree of life have usually attempted to circumvent the issue of HGT by focusing on a small subset of genes that are believed to be resistant to this problem, such as ribosomal RNA genes. However, this method carries two major issues: (1) the assumption that horizontal transfer of these genes is not possible may not be as robust as believed (some have suggested that there may be no such thing as a truly HGT-free gene), and (2) the smaller the data set used, the greater the chance that other complicating factors may interfere with results. For instance, it is now generally accepted that high-level phylogenetic reconstructions of eukaryotes using rRNA are very vulnerable to the effects of inequal evolutionary rates, with many supposedly 'basal' branches being shown to in fact be highly derived. There is no a priori reason to assume that the same problem would not apply to rRNA phylogenies of prokaryotes.

A paper just published in Cladistics takes the opposite approach to the problem: it uses an absolutely enormous amount of data to see whether a coherent tree can still be recovered. Two main data sets were used analysing 166 genomes from taxa throughout the tree of life (mostly prokaryotes). One concatenated direct amino acid sequences from 12,381 genes to provide 846,999 phylogeny-informative characters (out of a potential 4,540,579 characters). The other compared presence vs absence of genes from the 166 genomes. Analysis was done using parsimony, which is potentially problematic for sequence data but probably necessary to simply work with this amount of data. One analysis was run on the sequence data alone; another was run using the combined sequence and gene presence/absence data (the gene presence/absence data alone had been analysed by an earlier study).

The heartening result of this analysis is that a coherent phylogeny was recovered, particularly using the combined data set (shown above from the paper; a few anomalies were present using the sequence data alone). Most previously recognised major bacterial groups analysed were recovered by the combined data as monophyletic* (the only exception being the spirochaetes, with Leptospira failing to associate with the two Spirochaetaceae). Many of the higher-level relationships were also congruent with earlier proposals: α-proteobacteria as sister to the clade of β- and γ-proteobacteria, with δ-proteobacteria the next group out; a clade of the sphingolipid-producing bacteria (Chlorobium + Bacteroidales; and a clade uniting ε-proteobacteria with Aquifex + Thermotoga, which would then include all known hydrogen-oxidising Eubacteria. It appears unlikely that HGT fatally compromises large-scale analyses.

*Or perhaps I should say 'congruent'. As far as I can see, the study glosses over the question of the rooting of the tree of life; the tree shown is rooted between Neomura (Archaea + eukaryotes) and Eubacteria but no discussion is given on that position.

Of course, the tree is not without warning signs. The aforementioned polyphyletic spirochaetes are a bit worrying in light of the distinctive spirochaete ultrastructure. Some of the relationships within the major clades are a bit off: Gloeobacter is nested well within other cyanobacteria rather than being the most divergent (Gloeobacter is the only known cyanobacterium to lack thylakoids), and the arrangement of eukaryotes is all wrong. However, it must be stressed that, as large as this study was, the taxa analysed still represent only a small proportion of the world's total diversity. What is more, the choice of organisms to have their whole genome sequenced (a necessary pre-requisite for this study) has not been evenly distributed through prokaryote diversity. Many little-studied but potentially phylogenetically significant taxa (such as many of the low-diversity bacterial 'divisions') are significant by their absence, as are many significant subgroups of those divisions that are represented. This story is not yet over.

The Great Catalogue Table of Contents

Welcome to the Catalogue of Organisms Table of Contents. Here you can find links to all the major posts on this site, arranged by their subject. Enjoy!

Methods and Concepts:
        Wot? No inverts?
        Blog Action Day—what have we lost?
        A dragonfly in amber: how it got there
        Indian entomologists cut off
        More than one way to skin a cat (or fertilise a female)
        Implications of Aetogate: who owns the data?
        Inevitable moles in a lonely universe
        Is taxonomy a science
        Inevitable Spandrels on a biology blog
        The overwhelming diversity of life
        Why do we bother?
        What would the ICZN do?
        The gender of a table
        Phylogenetic nomenclature—oui ou non?
        Getting the hang of compromisation
        Why use phylogeny?
        What's in a name?
        Hey, old taxo! My genus is better than yours!
        A minor complaint about Google, and a major complaint about ranked taxonomy
        The claim-jumpers and grave-robbers of taxonomy
        A new stem-bird and publication in the digital age
        Electronic publication in the ICZN—new proposals
        Define "published"
        The perils of peer review
        Some thoughts on how to make electronic publication work
        The problem of publication again
        Thoughts inspired by a private publication
        It's the end of the world as we know it...
        The ICZN and electronic publication: where did it go wrong?
        O ZooBank, where art thou?
    Species Recognition
        The Phylogenetic Species Concept: is there such a thing as too much?
        What are the bare necessities?
        The significance of type specimens, and more on Utrecht
        The importance of vouchers: even molecular workers need herbaria
        Poor taxonomic practice takes some f***ing liberties!
        Keeping an eye on inflation
        How to recognise a species
        How to write a key

Holy careening continents, Batman!
Of serpentine soils

Maison Verreaux—animals of all varieties
Alexandre Girault: a man against the world
The worst of Girault

Taxonomy Quizzes:
Completely frivolous taxonomy quiz
42 [answers to above]
Taxonomy trivia quiz #2: you've come a long way, baby
Secret identities [answers to above]

Specific Taxa:

        Salinella—what the crap was it?
        Is it a sponge, or is it a plant?
        Naming the monad
        Giants of the Silurian
        Why animals are not plants
        More things in heaven and ocean
        Fungus and the individual
        Prototaxites: a giant that never was?
        Life on Mars: the Cambrian terrestrial environment
        Prototaxites revisited
        The diversity of slime moulds
            Crossing the algal divide
            Little discs of doom
            A parasite in the family
            Coral—it's not just an animal thing
            Carpospores in chains (Taxon of the Week: Schizoserideae)
            The alga of uncertainty
                    Giving plants the glove
                    Some like it cold (Taxon of the Week: Saccogynidium vasculosum
                    Southern moss (Taxon of the Week: Ptychomitrium muelleri)
                    Mosses: not as simple as you think (Taxon of the Week: Ectropothecium)
                    The trials and tribulations of tree moss
                    Mosses have a place for reproduction
                    Brachythecium salebrosum: some like it temperate
                            Before the word for world was forest
                                    The origins of flowers
                                    Most unbelievable organisms evah!
                                                    Flowers from two to five
                                                    The Dilleniaceae: tropical enigmas
                                                    Taxon of the Week: Misplaced Hawaiians
                                                        Are you sucking on a lemon or a lime?
                                                        The fall of Rafflesiales
                                                        Taxon of this Week: not all violets are violet
                                                        Reference review: Brooms of New Zealand
                                                        Taxon of the Week: Cotoneaster
                                                        Nettle, where is thy sting?
                                                        In a bunch, in a bunch!
                                                        A South American paradox (Taxon of the Week: Sellocharis)
                                                        A simple stream life
                                                        Cunoniaceae and friends
                                                        Milk-vetches, liquorice and locoweeds
                                                        Lecantheae and/or Elatostemateae
                                                    Christmas is coming
                                                    Stars in the pasture
                                                    Prickly pears
                                                        Taxon of the Week: Misplaced Hawaiians
                                                        Name the bug: Fouquieria columnaris
                                                            Hebe or Veronica?
                                                            My flower is a trumpet (Taxon of the Week: Solanales)
                                                            Borage and comfrey and bugloss
                                                            Sending forget-me-nots
                                                            Thistle be the one (Taxon of the Week: Carduoideae)
                                                            Ginseng and ivy
                                                The fall of Dryandra
                                            Reference review: The monocot tree
                                                Flowers in the water (Taxon of the Week: Hydrocharitaceae)
                                                Taxon of the Week: Rhaphidophora
                                                Name the Bug: Pistia stratiotes
                                                The thalli that are green (Taxon of the Week: Lemnoideae)
                                                    Peeling the lily
                                                    Strangers from parts unknown (Taxa of the Week: Juncus section Juncotypus, Juncus amabilis)
                                                    Scattering the sheaves (Taxon of the Week: Elymus)
                                                    Stacks of barley (Taxon of the Week: Hordeum)
                                                    From giant reeds to tiny leaves
                                                    The resurrection of grass
                                                    There's treasure everywhere
                                                    Most unbelievable organisms evah!
                                                    Patterns on a squill
                                            Taxon of the Week: Spices of Gondwana
                            When ferns don't look like ferns
                            Focus on a fern (Taxon of the Week: Polystichum vestitum)
                            It's not what you think
                            From tree moss to tree ferns
            Little whirling photosynthetic (and not so photosynthetic) thingies
            Pseudo-worms and such
            Taxon of the Week: A selection of ciliates
            Another non-missing not-quite-link
            Of macros and micros
            Of gregarines
            Like, wow. Just... wow
            The schizosphere (Taxon of the Week: Schizosphaerella)
            Taxon of the Week: Protoperidinium grande
            The state of Peridinium
            Glenodinium and the horseshoe of light
            Filling in the gaps
            Parcelling plastids
            Giant cannibal algae from the watery ditch
            Slime nets: another group of not-fungi
            Return of the slime-nets
            Algal threads (Taxon of the Week: Myrionemataceae)
            The wracks
            Name the Bug: Sticholonche zanclea
                    If a komokiacean turns up in a phylogeny, will anybody notice?
                    Living with poo—a new xenophyophore
                    A small bag of grains (Taxon of the Week: Saccamminidae)
                    Three random foram genera (Taxon of the Week: Pelosininae)
                    Star sands (Taxon of the Week: Calcarinidae)
                    Floating forams (Taxon of the Week: Globorotaliidae)
                    The Rotaliida: building a wall
                    The Osangulariidae: deep-water trochospires
        The diversity of slime moulds
        TAFKAMI walks
        Amoeba: much weirder than you think
        Amoebozoan classification: putting the formless in formation
        Tubulinea: the paragons of amoeboids
        Discosea: keeping a low profile
        Amoebozoan oddments
        Archamoebae: the apogee (or nadir) of amoebozoan evolution
                    Taxon of the Week: A barely pronounceable yeast
                        Saddling the truffles
                        Learning to like lichen
                        Reference review: Messing about with mildews
                        Reference review: The trials of anamorphic fungi
                        If they only wood (Taxon of the Week: Diaporthales)
                A relict fungus on a relict host
                Most unbelievable organisms evah!
                Ending life in a puddle of ichor (Taxon of the Week: Coprinopsis herbivora)
                The mushroom tree
                Tons of little tubes
                More little tubes—not just tons but tonnes
                Scleritome Week: The cactus animals
                Scleritome Week: A mystery ending
                The trouble with coelosclerites
                The return of Buddenbrockia
                Buddenbrockia: the gift that keeps on giving
                E pluribus unum
                Coral love
                Thought-crime: I have slandered the gelatinous
                Conical problematica
                From three to two
                A brain explosion
                        Scleritome Week: Worm buttons
                        It's nematodes all the way down
                        Oh #$%^& me—it's the Taxon of the (last) Week
                        My first tardigrades
                        Return of the water bears (Taxon of the Week: Tardigrada)
                        Archechiniscus: distinctively indifferent
                        Welcome to Scleritome Week: The little nets
                            Chain, chain, chain
                            Blinding me with Science
                            A quick primer on arthropod growth
                                        Another word on arachnid phylogeny
                                            Cyphophthalmids wait for the mountain to come to them
                                            More tales of the crunchy
                                                Saintly harvestmen (Taxon of the Week: Equitius)
                                                Taxon of the Week: Cynortula, Cynortula
                                                Taxon of the Week: Stygnoplus
                                                Taxon of the Week: Collonychium
                                                Gonyleptids are just so cool
                                                Taxon of the Week: Metarhaucus
                                                Biantidae: the importance of titillators
                                            What is a daddy-longlegs?
                                            Gnah! Gagrella! Headdesk!
                                            Remarkable things
                                            More Gagrellinae (Taxon of the Week: Harmanda)
                                            Score one for biogeography
                                            Possibly the coolest thing I had published this year
                                            How to wipe out a family
                                            Disco opilioni
                                            The Gagrella problem cranked up to eleven
                                            The saga of Forsteropsalis fabulosa
                                            The hard way to be a bloodsucker (Taxon of the Week: Ixodidae)
                                            Brown ticks
                                            Your little friends that are with you always
                                            Most unbelievable organisms evah!
                                            The Pygmephoroidea: lives of phoresy and fungi
                                            The Prostigmata: endless forms
                                                        On a wing and a mite
                                                        South American mites
                                                    Life in the fast lane (Taxon of the Week: Astigmata)
                                                    An introduction to Malaconothrus
                                            Tarantulas sans tarantella
                                                What is a daddy-longlegs?
                                                Spiders losing their lungs
                                                    The strangest of spiders
                                                    Taxon of the Week: Eye of the spider
                                                    Amaurobioidea: rummaging through a wastebasket
                                                    Salticid spider bollocks
                                                    Taxon of the Week: Amphinectidae
                                                    Big bad wolfies (Taxon of the Week: Lycosidae)
                                                    More wolfies (Taxon of the Week: Artoriinae)
                                                    Lace web weavers
                                                    The one about sexual cannibalism
                                                    Araneidae—with web and with scent
                                    Brine fairies
                                    Taxon of the Week: Some copepods for your reading pleasure
                                    Another case of mistaken identity
                                    Life in sand
                                        Reference Review: The secret of Bubba-Gump's success
                                        Getting crabs
                                        Kneel before the shrimp queen
                                        Crabs that cannot scratch their heads (Taxon of the Week: Parthenopidae)
                                        The Grapsidae: from sea to shore
                                        Taxon of the (this) Week—Holarctic subterranean amphipods, Batman!
                                        Southern crustacean relicts
                                        Another case of mistaken identity
                                        Snail mimics and marine symbionts (Taxon of the Week: Pleustidae)
                                        Burrowing beaky amphipods
                                        Life among a shrimp's gills
                                    Reference review: Barnacles among the coral
                                    The secret of y-larvae
                                    Forcing out the secret
                                    Most unbelievable organisms evah!
                                    (Possibly) The World's Smallest Tetrapods
                                        Bouncing bristletails
                                        Textbook death match: Insect palaeontology
                                            The origin of insect wings
                                            A Devonian pterygote?
                                            What is the sound of one mayfly fossilising?
                                            Everything you knew about mayflies is wrong (Taxon of the Week: Pisciforma)
                                            Mayflies in their spring
                                            A halfway house, halfway down Honshu island
                                            Big suckers
                                            Because it's Friday...
                                            A seclusion of Embioptera
                                            A choir of Zoraptera
                                            When parsimony goes wrong: the wings of stick insects
                                            All about Gerarus
                                            Name that Bug: Ponopterix axelrodi
                                            Wigs and wings and other things
                                            The stoneflies: old or new?
      ;                                      Psocoptera
                                                Barklice and booklice and such
                                                The Psocoptera of Barrow Island
                                                Love Hurts
                                                Soft waxy scales
                                                Soft yet scaly (Taxon of the Week: Coccidae)
                                                The overall scale
                                                A vision of thrips
                                                The really abominable mystery
                                                            A bunch of apocrites
                                                            Eureka! It's an ant!
                                                            Most unbelievable organisms evah!
                                                            Ants on the move (Taxon of the Week: Dorylidae)
                                                            Ants go out in the noonday sun
                                                            What to do with a dead hummingbird
                                                            Multifarities most horrid (Taxon of the Week: Braconidae)
                                                                Taxon of the Week: To give Lovecraft nightmares
                                                                    Ormyrids: attacking the gall
                                                                    A new short-horned Elasmus
                                                                    Brachymeria perflavipes and beyond
                                                        Diversity and distribution of tropical Lepidoptera: a bit of cross-purposes
                                                                    Taxon of the Week: butterflies on parasites
                                                                    The sphinxes that aren't like the others (Taxon of the Week: Smerinthini)
                                                                    Caterpillars and their capers (Taxon of the Week: Belenois)
                                                                    Blues (not all of them blue) (Taxon of the Week: Polyommatus)
                                                                    Colour vs crypsis
                                                                    Deceptive and poisonous sisters
                                                                    The butterflies get all the glory (Taxon of the Week: Gelechioidea
                                                    Insects never fail to amaze
                                                    What is a daddy-longlegs?
                                                    A queenage of Strepsiptera
                                                    Drosophila forever?
                                                    More on Drosophila and Sophophora
                                                    "Electronic publication of nomenclatural acts is inevitable"
                                                    Name the Bug: Boreus
                                                    More in the bloodsucking vein (Taxon of the Week: Simulium)
                                                    In which, despite not being the crowd favourite, Drosophila funebris holds D. melanogaster down and kicks it repeatedly in the teeth
                                                    More than just Sophophora (Taxon of the Week: Drosophilidae)
                                                    Freak of the Week: wingless, legless flies
                                                    What is the sound of one mayfly fossilising?
                                                    Eating mum from the inside out
                                                    The diversity of ground beetles
                                                    Ground beetles for today
                                                        Life with termites
                                                        The Corotocini in their gut-swollen glory
                                                        Bryaxis on the prowl
                                                        There's something on your back (Taxon of the Week: Prioninae)
                                                        Weevil ball (Taxon of the Week: Diorymerina)
                                                Of lions and lace
                                                Butterflies before there were butterflies
                                                Name that Bug: Meioneurites spectabilis
                                                Name the Bug: Psectra diptera
                                                The red-lined wings of South America
                                                Ant-lions of Australia
                                The august history of filter-feeding ostracods
                            The trouble with coelosclerites
                                Tons of little tubes
                                So nice when people agree with you
                                    Snails letting it all hang out
                                    Nerites old and new
                                    A different kind of shell
                                    Name the Bug: Alaskiella medfraensis
                                    Marginal limpets
                                            In which I am defeated by shells
                                            The life of an ostrich foot
                                            Stop giggling (Taxon of the Week: Fartulum)
                                            Careful with that spelling (Taxon of the Week: Barleeiidae)
                                            Snails that never see the light of day
                                            Taxon of the Week: Toxic sea snails
                                            A whole new twist on things, or just shifting back and forth?
                                            Fifteen seconds of mediocrity
                                            Asperdaphne, I don't know who you are any more
                                            (Belated) Taxon of the Week: The bishop's Mitra
                                            The coral-lovers
                                    I's been ejucated, now I can haz snails pleez? Kthnx
                                    Food that puts more than just hairs on your chest
                                    The beautiful angel of death
                                    Taxon of the Week: Clausilioidea
                                    Wild slug chases (Taxon of the Week: Gastrodontoidea)
                                    Name that Bug: Cornu aspersum
                                    Re-opening the door (Taxon of the Week: Clausilioidea)
                                    A little bit of Gastrocopta
                                    Limpets of the north-east Atlantic
                                The calcareous heart
                                Pig's toes and water nymphs
                                Bivalves born free (Taxon of the Week: Pectinoidea)
                                Triassic, glorious Triassic
                                With plate and girdle (Taxon of the Week: Ischnochitonidae)
                                The beak-shells's legacy
                                Name the Bug: Yochelcionella daleki
                                Day of the Tentacle
                                More giant cephalopods
                                The floating egg
                                Open query—what are cephalopod shells for?
                                When is a cephalopod like a snake?
                                How to be straight
                                Nectocaris: largely irrelevant to cephalopods?
                                Ammonites of the Arctic (Taxon of the Week: Arctocephalitinae)
                                The horns of Ammon
                            Taxon of the Week: The lamp (shell) post
                            Back to the scleritome—tommotiids revealed!
                            More giant larvae
                            More crunchy scleritome goodness
                            The Athyrididae: spiralia and lamellae
                            Yay, machaeridians!
                            Separating segments
                            More crunchy scleritome goodness
                            My genitals just grew eyes and swam away: the life of a syllid worm
                            Building a home of your own (Taxon of the Week: Hydroides)
                            Of interstitial annelids (Taxon of the Week: Pisionidae)
                            Earthworms of the Amazon (Taxon of the Week: Urobenus buritis)
                            Exogone sexoculata, a worm of the interstitial
                        Tiny flowers of the sea
                        Colonies on the move
                        Taxon of the Week: Lacy Lepraliellidae
                        Gunky lace
                        The long-whipped bryozoan
                        Taxon of last Week: I can't think of a clever title involving gastrotrichs
                        Sex and the rotifer
                        I can has mutant larvae?
                        The writing in the rocks
                        Further readings from the rocks (Taxon of the Week: Graptolithina
                        Name the Bug: Acanthastus luniewskii
                        Rastrites: stringing out thecae
                            Conversations with Cothurnocystis
                            Name the Bug: Polyplacus kilmeri
                            A beginner's guide to blastoids
                            Callocystitids: ambulacra advancement and rhomb reduction
                            Taxon of the Week: So many arms
                            Return to the crinoids
                            Clutching crinoids
                            Forgotten feather stars
                            Crinoids of the open seas
                            Mystery animal for today
                            Hemiaster: an echinoid with heart
                                Time for teeth (Taxon of the Week: Polygnathus)
                                Conodonts: they just got scarier
                                A little Linguipolygnathus
                                Linguipolygnathus redux
                                        Scleritome Week: Not just an invert thing
                                        Just when you thought it was safe
                                        A bizarre new shark
                                                Who left all this fish lying around? (Taxon of the Week: Neopterygii)
                                                        Taxon of the Week #1: Gonostomatidae
                                                        Living larvae and fossil fish
                                                        Hunters in the deep sea
                                                            The bush at the top of the tree
                                                            Perciformes go bye-bye?
                                                                Taxon of the Week #2: Trachinoidei
                                                                The live-bearing brotulas
                                                                Pomfrets of the high seas
                                                                The ugly stick in action
                                                                Taxon of the Week: Give us a kiss!
                                                                Gender's just a state of gonads
                                                                Sculpins go wild
                                                                Knocked off the perch (Taxon of the Week: Percidae)
                                                    More Really Ugly Fish
                                                    It's a moray Friday
                                                    The surprisingly mysterious eels
                                                Some history of the history of tetrapods
                                                Taxon of the Week #3: Rana
                                                Sooglossidae: Deja vu all over again
                                                Relict frog sex
                                                Most unbelievable organisms evah!
                                                        The tuna-lizards
                                                                Because it's Friday...
                                                                Tortoise resurrection
                                                                Small lizards of South America
                                                                Southern snakes at sea
                                                                Anole, anole, anole, anole
                                                                Obama's lizard? Not so fast
                    Our faceless cousins?
                    Gutless wonders
                    Name that Bug: Stoecharthrum giardi
    The nature of 'Nanoarchaeum'
    Taxon of the Week: Pick from a wide range of pathogens
    Reference review: Cutting up the excess
    Taxon of the Week: Life in mycolates
    Standing the heat
    Epsilon of the deeps—coming to an organ system near you
    Life before it had facial features
    Hyphae without nuclei: filamentous bacteria
    The diversity of slime moulds
    Most unbelievable organisms evah!
    Pathogens, or more than pathogens? (Taxon of the Week: Aeromonas)
    The endosymbiotic hammer strikes again
    Bacteria, too, grow old and die
    The attack of mega-matrix
    The Nostocaceae: tangled filaments
    The Rhodospirillales: it's photosynthesis, but not as you know it

        Insectivores: possibility of puggles (Taxon of the Week: Australosphenida)
                A little bit mole-ish in the Miocene
                The mysterious name of Queen Lestoros
                    Life in the Palaeocene—we don't need no Placentalia?
                    More mysterious Palaeogene eutherians
                            The swimming sloth
                            Meet the Shrews (Taxon of the Week: Soricidae)
                            Little yellow bats
                            Taxon of the Week: Phocidae
                            Taxon of the Week: Capra—the goats
                            The camel that walked on two legs
                            The whale that looked like a walrus
                            Horns and guts
                            West Indian raccoons: from endangered endemics to invasive introductions
                            Moustache whales
                            Three-quarters of a century, and we still care about a dead horse
                            Support your local taxonomy
                            Fantastic Mr Fox
                            Dog's life
                            The uglier side of the family (Taxon of the Week: Ceratomorpha)
                            Wolf and wolf and wolf and wolf and cub
                            "Creodonts": carnivores by association
                            How the badger became (Taxon of the Week: Meles thorali)
                            More than just Moby (Taxon of the Week: Physeteridae)
                            The sad, sad story of Physeter
                            The wolf in time
                            Taxon of the Week: Anthropoidea
                            Taxon of the Week: Marmotini
                            Taxon of the Week: Apodemus (no pliers)
                            Taxon of the Week: Nothing to do with teapots
                            Sacred monkeys
                            Most unbelievable organisms evah!
                            Name the Bug: Anomalurus pelii auzembergeri
                            The parrot of King Charles I
                            Beginning to grasp things (Taxon of the Week: Euprimateformes)
                            There he goes! (Taxon of the Week: Indriidae)
                            Squirrels and cedar-mice (Taxon of the Week: Sciuridae)
                            Old men of the woods
                            Origins - a day in the Broom Room
                            Groundhogs, woodchucks and other big squirrels
                            Beaver fever
        The rise and fall of the multi-cusped

        The ornithocheirids: misunderstood giants
            The Top Ten dinosaurs—Triceratops beats Tyrannosaurus
            Top Ten follow-up
            Dinosaurs all over the place
            A question of availability
            Most unbelievable organisms evah!
            The anchisaurs: near-lizards or near-sauropods?
                                    A new stem-bird and publication in the digital age
                                                A frustrating giant bird
                                                How irritable is this bird?
                                                Bird evolution—problems with Science
                                                More from the "They don't write papers like they used to" files
                                                    The voice of the turtle (Taxon of the Week: Streptopelia)
                                                    Jonathon Livingstone's cousin (Taxon of the Week: Sterna)
                                                    Name the bug #8: Prosobonia cancellata
                                                        Reference review: Parrots in the early days of molecular analysis
                                                        A King among parrots
                                                            Taxon of the Week: Cotinginae—Neotropical and fabulous!
                                                                    Products of kinky inter-species sex
                                                                    Birds of the sun
                                                                    The tomb of the unknown honeyeater
                                                                    Name the Bug #7—Apalopteron familiare hahasima
                                                                    More than four and twenty blackbirds
                                                                Yet another passerine 'family'
                                                                On hybrid birds
                                                                The shrikes of the south
                                                                Whistling for whistlers
                                                    Oh crake (Taxon of the Week: Amaurornis)
                                                    Cranes off the rails (Taxon of the Week: Grues)
                                                    Banana-eating birds that don't eat bananas
                                                    The stately herons
                                                The Hawaiian superducks
                                                A pathetic plea for recognition, and a platypus-billed duck
                                                Taxon of the Week: Leg or breast?
            Big horned lizards
            Ceratopsids: a Cretaceous flash in the pan

    Viruses upon viruses