Field of Science

Life with Termites

Termitusodes lativentris. Image from Seevers (1957).

The Staphylinidae are one of the largest (over 46,000 species) and most easily recognisable of the commonly accepted beetle 'families'. Among other things, they can be readily distinguished from other beetles by the reduction of the elytra so that they no longer cover the abdomen, similar to what is seen in the earwigs. The most familiar staphylinids are relatively large predatory species (known as rove beetles or devil's coach-horses) but the family includes a wide variety of other forms.

The animal rather poorly illustrated at the top of the post is a member of the subtribe Termitusina in the subfamily Aleocharinae. Termitusina are found in tropical Africa where they live within the colonies of certain species of termite. The image doesn't show the distinguishing features of this group very well, unfortunately, but if you look carefully (and perhaps use a bit of imagination) you may be able to make out the strongly deflexed head and ninth abdominal tergite divided between two elongate halves. Unlike many termite-associated staphylinids, Termitusina of the genera Termitusa and Thoracotusa do not generally exhibit physogastry (a swollen abdomen). Termitusodes, the third genus in the subtribe, differs from the other two genera in this respect, though it still comes nowhere near the level of physogastry found in some termite-associated staphylinids.

I haven't been able to find out whether the Termitusina are in some way social parasites of their host termites or whether they are finding their food in the nest some other way (such as by scavenging or feeding on fungi) but the association is undoubtedly a close one. Termitusina are, as a rule, host specific (Jacobson & Kistner, 1975). Termitusa are found only in nests of the termite genera Cubitermes and Noditermes, Thoracotusa is found with Thoracotermes, and Termitusodes has been found with Cubitermes and Pericapritermes magnificus. Pericapritermes often nest in association with Cubitermes (they may occupy abandoned Cubitermes nests or they may invade the nest while the Cubitermes are still living there) and it seems likely that this association is what allowed Termitusodes to change hosts at some point in its history.


Jacobson, H. R., & D. H. Kistner. 1975. Numeric analyses of relationships of genera and species of the subtribe Termitusina (Coleoptera: Staphylinidae). Systematic Zoology 24 (2): 191-198.

Seevers, C. H. 1957. A monograph on the termitophilous Staphylinidae (Coleoptera). Fieldiana: Zoology 40: 1-334.

Name the Bug # 39

Attribution to follow. And yes, I realise that this image is beyond awful, but would you believe that this was the best that I could find?

Also, you may notice that there's been a bit of a design change here. All credit for that goes to Edward, the benevolent overlord of Field of Science.

Update: Identity now available here. Image from Seevers (1957).

Life in Sand

Paramesochra mielkei, from Huys (1987).

Paramesochra is a genus of minute marine copepods found around the world. Over twenty species are currently assigned to the genus, but it is likely that many more await description. The extremely small size of paramesochrids (most are less than half a millimetre in length) reflects the interstitial habitat of most species described to date, i. e. they live among the grains of sand beneath the surface of their substrate. Also related to their choice of habitat is their vermiform (worm-like) shape and reduced setation compared to other copepods. These features also mean that they would be poor swimmers so they probably do not often emerge above the substrate surface. Most of the species described so far are from shallower waters, but this possibly reflects a lack of study of deep-sea species rather than reflecting true diversity. For instance, a survey of deep-sea Paramesochridae in the southern Atlantic and Antarctic Oceans by Gheerardyn & Veit-Köhler (2009) identified four species of Paramesochra, none of which corresponded to previously described species. These species probably do not have the same lifestyles as the shallow-water interstitial species due to the deep-sea substrate being fine mud rather than sand. Vasconcelos et al. (2009) suggested that another deep-sea paramesochrid, Kliopsyllus minor, might burrow in fluid mud or live in the 'organic fluff layer' (wonderful words) on top of the sediment. Deep-sea Paramesochra would probably be similar.

For the most part, genera of copepods have generally been distinguished mechanistically—different genera have different combinations of key features (usually related to the number of setae or segments on appendages)—without an explicit consideration of how those characters relate to phylogeny. However, Huys (1987) did propose a phylogenetic arrangement for the genera of Paramesochridae in which he suggested that Paramesochra formed a clade with the genera Kliopsyllus and Kunzia on the basis of their possessing single-segmented exopodites on the antennae and mandibles. However, while his tree shows Paramesochra as a monophyletic sister group to a clade of the other two genera, he did not identify any synapomorphies for Paramesochra. Instead, the features distinguishing it from the other two genera (two-segmented endopodites on the second to fourth legs, four setae on the distal exopodite segment of the first leg and two setae on the distal exopodite segment of the fourth leg) are resolved as plesiomorphies relative to the other clade. So if any of you feel inspired to spend your time dissecting and examining the legs of animals about 0.3 of a millimetre in total length, I know a potential research project going begging...


Gheerardyn, H., & G. Veit-Köhler. 2009. Diversity and large-scale biogeography of Paramesochridae (Copepoda, Harpacticoida) in South Atlantic Abyssal Plains and the deep Southern Ocean. Deep-Sea Research I 56: 1804-1815.

Huys, R. 1987. Paramesochra T. Scott, 1892 (Copepoda, Harpacticoida): a revised key, including a new species from the SW Dutch coast and some remarks on the phylogeny of the Paramesochridae. Hydrobiologia 144: 193-210.

Vasconcelos, D. M., G. Veit-Köhler, J. Drewes & P. J. Parreira dos Santos. 2009. First record of the genus Kliopsyllus Kunz, 1962 (Copepoda Harpacticoida, Paramesochridae) from Northeastern Brazil with description of the deep-sea species Kliopsyllus minor sp. nov. Zootaxa 2096: 327-337.

Name the Bug # 38

These particular animals seem to generally be illustrated in bits:

Attribution to follow.

Update: Identity now available here. Figure from here.

A Review: Tetrapod Zoology: Book One

Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: United Kingdom.

This review is probably a little redundant. I suspect that most of my regular readers are probably already familiar with Tetrapod Zoology, Darren Naish's leading blog on all things vertebrates-that-ain't-fish-y. Nevertheless.

The original version of Tet Zoo debuted in January 2006, almost five years ago. Tetrapod Zoology: Book One is a collection of most of the posts which appeared from January to July of that year. For the most part, the text remains the same as it first appeared; the main change is that some of the topics that appeared as multiple posts have beeen fused into single chapters. The included chapters cover topics from cave-dwelling salamanders to corpse-hunting turtles, from contentious monkeys to elusive warblers. As always, Darren has the rare knack of combining breezy readibility with an extraordinary wealth of detail. And as well as chapters on dramatic topics such as giant pterosaurs and British big cat sightings, the book contains examples of Darren's even more admirable ability to highlight the extraordinary sides of seemingly unremarkable animals such as rabbits, gulls and owls.

There are few bloggers who could pull off a simple transition from blog post to book like Darren has, and even in Darren's case I'm not sure the process has been completely successful. The original appearance of the book's chapters as independent units means that there is a noticeable lack of connection between most of them, nor is there any obvious pattern to the topics covered. Also, some of the chapters might have been expected to have received more editing had they originally gone through a more leisurely . A chapter on stem-whale diversity refers to 'whales with trunks' both in the title and in the introductory paragraphs, but fails to enlarge on this further. Also, the time that has elapsed since the original composition of the chapters and their publication in the book means that some of the chapters can seem a little dated; both in obvious ways (references to 'recent' events that are no longer recent) and not-so-obvious ways (would Darren have revised his comments on Michael Heads' biogeography of birds-of-paradise had he known that Heads would later use the same principles to argue for a Jurassic divergence of modern primates?)

Such issues are not unique to blogs, of course—had this been a collection of magazine essays or newspaper columns, there would be cause for the same complaints. Still, I feel that the layout of this book should have embraced its history more, rather than simply presenting the sections as if they were written for the book. It would have been nice to have been given the original release date for each of the chapters (Darren does make it clear in the introduction that they were written in 2006, but it would have been good to have individual dates, ideally at the head or foot of each chapter). If nothing else, doing so would allow the inclusion of Darren's legendarily detailed April 1 posts*. Also, while Darren does include an update section in the book's introduction mentioning some of the significant events that have happened since relevant to the original chapters, it may have been better to present each chapter's updated separately as an addendum to the chapter itself. And I'm surprised that Darren failed to mention Roosmalen et al. (2007) in which the new peccary species discussed in the final chapter was named Pecari maximus (the validity of the species apparently remains contentious).

*I'm also personally disappointed by the absence of one of my favourite Tet Zoo posts from the period included—indeed, one of my favourites of all time—the story of Walter Rothschild and his all-consuming cassowary obsession.

None of these complaints, of course, are particularly serious. Most of them disappear if the book is read as its contents were originally intended: not as a unit, but as a series of fascinating essays to be indulged in, one chapter or two at a time, when looking for a half-hour or so's entertainment. And there would be few people indeed who would fail to come away from it without an increased sense of wonder about the animals it describes. Of course, you may be wondering why you should be paying for something that is available for free online. To which I can simply point out that, as well as the moral satisfaction of doing your part to help keep Darren and his family in stockings and fans, the success of this book can only encourage him to produce more. Hopefully, Tetrapod Zoology: Book Two will not be long in coming.


Roosmalen, M. G. M. van, L. Frenz, P. van Hooft, H. H. de Iongh & H. Leirs. 2007. A new species of living peccary (Mammalia: Tayassuidae) from the Brazilian Amazon. Bonner Zoologische Beiträge 55 (2): 105-112.

Learning to Like Lichen

The lichen Parmelia saxatilis. The red cups at the top of the photo are the lichen's fruiting bodies (apothecia) that produce fungal spores. Photo from here.

We all know what lichens are. They're the standard example of a mutualistic association that we were all presented with in high school, an association of fungus and unicellular alga allowing both to survive long-term in situations that would normally be fatal for them both. More than 15,000 species of lichen have been described—or, rather, species of lichenised fungi, as names applied to lichens technically apply to the fungal member of the association (only a relatively small number of algae form lichen associations). Though these species can all be attributed to the Ascomycetes among the main fungal subdivisions*, they do not form a single clade within the asomycetes. Instead, it appears that the lichen lifestyle has been gained and/or lost on numerous occasions.

*Lichen-like associations are sometimes formed by other fungi such as Basidiomycetes but they lack the integrity of the ascomycetous examples. Lab workers have even been able to induce lichen-like associations between unicellular algae and colonial or hyphal bacteria such as myxobacteria and streptomycetes (Ahmadjian 1965).

Parmelia is a genus of foliose lichens which is found worldwide but has its highest diversity in Asia (Molina et al. 2004). Well over 1000 species have been assigned to the genus over the years but many (though not all) recent authors have tended towards a much more restricted circumscription of about forty species. True Parmelia, in this sense, is distinguished from other genera in the lichen family Parmeliaceae by its linear pseudocyphellae (pore-like structures in the upper-surface of the lichen's cortex) and its particularly small spores and conidia (conidia are reproductive structures like spores but produced asexually rather than sexually) (Elix 1993). ITS rDNA phylogeny is mostly consistent with many of the proposed segregate genera, including the restricted Parmelia, though it provides little information on their higher relationships (Crespo & Cubero 1998).

Another view of Parmelia saxatilis. As well as the spore-producing apothecium, this photo also shows numerous isidia, the small finger-like protrusions covering the thallus. Containing both fungal and algal cells, the isidia can break off to form new lichens. Photo by Stephen Sharnoff.

Parmelia achieves its highest diversity in temperate or boreal regions. The type species, P. saxatilis, is one of the world's most widespread lichen species, found in both the Arctic and the Antarctic, as well as cooler localities in between (Molina et al. 2004). Lichens can reproduce in one of two ways: small pieces of the thallus containing both algal cells and fungal hyphae may break off to grow directly into a new thallus elsewhere, or the lichen can release spores and/or conidia in the manner of other fungi. A germinating lichen spore will grow extremely slowly: even in laboratory cultures on agar, some lichen fungi will only reach a diameter of 1 mm within the course of a year when grown without algal symbionts(Ahmadjian 1965). Formation of the lichen association is dependent on the fungus randomly coming into contact with an alga, and growing lichen fungi will form exploratory hyphae around anything (even grains of sand) that they touch that might turn out to be an alga (Ahmadjian 1960). The low variety of algal species occuring in lichens appears to be dependent not on any direct attraction of the alga for the fungus, but on the alga's ability to resist digestion by the fungus' hyphae. Lichens are famed for their slow growth even after an association is established, and may increase in diameter by only a millimetre a year*, but the limiting factor is probably not so much their inherent growth abilities as that their favoured environments such as exposed on rocks may only allow growth for a minute part of the year.

*If you're thinking that that doesn't sound any greater than the rate for symbiont-less fungi that I cited above, remember that the latter rate applies to growth in the laboratory under theoretically optimal conditions; growth in the natural environment would be much, much slower.


Ahmadjian, V. 1960. The lichen association. Bryologist 63 (4): 250-254.

Ahmadjian, V. 1965. Lichens. Annual Review of Microbiology 19: 1-20.

Crespo, A., & O. F. Cubero. 1998. A molecular approach to the circumscription and evaluation of some genera segregated from Parmelia s. lat. Lichenologist 30 (4-5): 369-380.

Elix, J. A. 1993. Progress in the generic delimitation of Parmelia sensu lato Lichens (Ascomycotina: Parmeliaceae) and a synoptic key to the Parmeliaceae. Bryologist 96 (3): 359-383.

Molina, M. del C., A. Crespo, O. Blanco, H. T. Lumbsch & D. L. Hawksworth. 2004. Phylogenetic relationships and species concepts in Parmelia s. str. (Parmeliaceae) inferred from nuclear ITS rDNA and β-tubulin sequences. Lichenologist 36 (1): 37-54.

Cunoniaceae and Friends

The Albany pitcher plant Cephalotus follicularis of south-western Australia. The streaked colours on the inside of the lid attract insects into the pitcher; the incurved teeth around the rim stop them from climbing back out. Photo by Holger Hennern.

The plant order Cunoniales was first established in 1926 to include plants with similar flowers to the Saxifragales but that were primarily woody rather than herbaceous (Dickison, 1975). Woodiness vs. herbaceousness is no longer considered that significant a feature in plant classification (sometimes you can find both in the same genus) and the content of the order has varied between classifications*. Today, the type family Cunoniaceae is included in the order Oxalidales and a taxon "Cunoniales" is no longer used as such. However, one of the two basal clades within the Oxalidales includes the Cunoniaceae and two ex-Cunoniales genera placed in their own families, Cephalotus and Brunellia, together with the family Elaeocarpaceae (previously in its own order) (Matthews & Endress, 2002). With the notable exception of Cephalotus, the members of this clade are mostly shrubs or trees. Economically, the clade is not overly significant: some species are used for wood; some have good reputations as honey sources for bees; a few produce edible fruits but do not appear to have been systematically cultivated for them. Members of all families bear their flowers clustered into (most often cymose) inflorescences; the size of individual flowers in the inflorescences varies between species. Brunellia and Cephalotus both produce flowers with thick sepals and no petals; in the other two families, petals may be present or absent (Matthews & Endress, 2002).

*As have most flowering plant "orders". There's a reason why order-level taxa don't get much day-to-day use among botanists compared to families.

Coachwood, Ceratopetalum apetalum, a member of the Cunoniaceae from eastern Australia. Photo by Melburnian.

The clade formed by these four families has a distinctly southern distribution in southern Africa, South and Central America, south-east Asia, Australia and New Zealand. Elaeocarpaceae are absent from continental Africa but are present in Madagascar. Also, while currently absent from India, they have been recorded from the fossil record there (Crayn et al., 2006). The Cunoniaceae include about 300 species, half in the genus Weinmannia, whereas the Elaeocarpaceae include about 600 species. Molecular studies have shown that an Australian radiation of dry-habitat shrubs previously regarded as the family Tremandraceae is in fact a subclade of the otherwise mostly rainforest-inhabiting Elaeocarpaceae (Crayn et al., 2006). Interestingly, the molecular dating study by Crayn et al. (2006) suggests that the 'Tremandraceae' developed scleromorphy (a suite of adaptations such as hardened leaves that are usually associated with arid habitats) some time before the Australian continent developed its current arid climate. While this may seem counter-intuitive, it is worth pointing out that the fossil record supports the same thing in the evolution of the genus Banksia (Mast & Givnish, 2002). It has been suggested that scleromorphy in these groups was therefore not originally an adaptation for arid living, but for growing in the poor soils of the Palaeogene Australian rainforest.

Prima donna, Elaeocarpus reticulatus. Fringed petals are characteristic of a number of species of Elaeocarpaceae. Elaeocarpaceae flowers are also adapted for buzz-pollination, where anthers do not release their pollen until vibrated by the wingbeats of their pollinating bees. Photo from Kate's Photo Diary.

The oddest member of this clade, however, has to be the Albany pitcher plant, Cephalotus follicularis. Restricted to the south-west corner of Western Australia, this plant bears a superficial resemblance to pitcher plants from other parts of the world, the mostly Indomalayan Nepenthaceae and the North American Sarraceniaceae. All three of these families have evolved the pitcher morphology independently, and can in fact be placed in separate subclasses: Cephalotus in the Rosidae, the Nepenthaceae in the Caryophyllidae and the Sarraceniaceae in the Asteridae. In the Nepenthaceae, the pitchers are developed from trendrils at the ends of the leaves; in the other two families, the entire leaves form the pitchers growing from a ground-level rhizome. In Cephalotus, only the outer leaves of an individual plant are developed into pitchers; the inner leaves remain flat and simple.


Crayn, D. M., M. Rossetto & D. J. Maynard. 2006. Molecular phylogeny and dating reveals an Oligo-Miocene radiation of dry-adapted shrubs (former Tremandraceae) from rainforest tree progenitors (Elaeocarpaceae) in Australia. American Journal of Botany 93 (9): 1328-1342.

Dickison, W. C. 1975. Studies on the floral anatomy of the Cunoniaceae. American Journal of Botany 62 (5): 433-447.

Mast, A. R., & T. J. Givnish. 2002. Historical biogeography and the origin of stomatal distributions in Banksia and Dryandra (Proteaceae) based on their cpDNA phylogeny. American Journal of Botany 89 (8): 1311-1323.

Matthews, M. L., & P. K. Endress. 2002. Comparative floral structure and systematics in Oxalidales (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae). Botanical Journal of the Linnean Society 140 (4): 321-381.

Name the Bug # 37

Attribution to follow.

Update: Identity now available here. Photo from here.

The Fall of Dryandra

Banksia nivea, previously Dryandra nivea. Photo from here.

Adam Yates has requested that I do a post on the relationship between Banksia and Dryandra, two Australian genera (but read on) of the plant family Proteaceae. Banksia has a generally coastal distribution around Australia, but is conspicuously absent through the Nullarbor region and northern Western Australia (i.e. where the coast is driest). Dryandra is restricted to the south-west corner of Australia. A close relationship between the two genera has long been accepted, supported as it is by features including the bearing of flowers in compact inflorescences (cone-shaped in Banksia, capitate in Dryandra) and production of seeds with a hard bony endocarp. These seeds are contained in bivalved woody follicles, only a relatively small number of which develop to maturity in any given inflorescence. In most cases, the follicles do not open at maturity but remain closed until heated by the passage of a bushfire; only after the fire do they open to release their seeds. The appearance of remnant Banksia cones with the protruding follicles resembling eyes or gaping mouths has long affected Australian folklore, with the most familiar example being May Gibbs' 'bad banksia men'.

The banksia men, villainous characters from May Gibbs' Snugglepot and Cuddlepie books, in their natural habitat.

However, recent years have seen the species of Banksia involved in a greater controversy than the mere kidnapping of gumnut babies: the kidnapping of an entire genus. Molecular studies have shown that Dryandra is phylogenetically nested within Banksia, rendering Banksia in its familiar sense paraphyletic (Mast & Givnish, 2002; Mast et al., 2005). This has lead to the formal synonymisation of the two genera by Mast & Thiele (2007), an action that has incited its fair share of grumbling among Australian botanists and horticulturalists. However, the alternative—dividing Banksia into multiple genera—would have required establishment of a number of new genera as the type species, Banksia serrata, is among the closer relatives of Dryandra. Also, these new genera would probably not have been easily distinguishable morphologically.

Banksia gardneri, showing both new and persistent cones. Photo by Brian Walters.

Though initially incited by molecular studies, the paraphyly of Banksia excluding Dryandra is also supported by morphological factors. Mast & Givnish (2002) supported a division of Banksia into two clades which Mast & Thiele (2007) recognised as the subgenera Banksia (including Dryandra) and Spathulatae. Members of Banksia subgenus Banksia have beaked follicles while the follicles of Banksia subgenus Spathulatae are unbeaked. Also, most members of subgenus Banksia have the stomata on their leaves recessed into deep pits, though this feature has appeared apparently independently in one small subclade of subgenus Spathulatae (most of which have more superficial stomata). The sinking of the stomata into pits (an adaptation for living in arid environments) is also supported as a derived feature by the fossil record of Banksia sensu lato, the earliest known representatives of which in the Late Palaeocene lack such arid adaptations (arid-adapted banksiines are not known until the Late Eocene).

Images of Banksia ilicifolia, a short-coned potential relative of Dryandra from south-western Australia. Photos by T. J. Alford & C. Hortin.

Within Banksia subgenus Banksia, the clade now known as Banksia series Dryandra falls within a clade also containing other south-west Australian species of what have previously been regarded as 'series Banksia' and 'subgenus Isostylis'. The latter are interesting in this regard as having particularly short cones, and had been compared to Dryandra even when the two were regarded as separate genera. But while the short-coned banksias are close to series Dryandra, they do not necessarily form an exclusive clade with them, so it is uncertain whether their short cones represent a transition between the normal Banksia cone and the capitate Dryandra inflorescence, or whether they have shortened convergently.


Mast, A. R., & T. J. Givnish. 2002. Historical biogeography and the origin of stomatal distributions in Banksia and Dryandra (Proteaceae) based on their cpDNA phylogeny. American Journal of Botany 89 (8): 1311-1323.

Mast, A. R., E. H. Jones & S. P. Havery. 2005. An assessment of old and new DNA sequence evidence for the paraphyly of Banksia with respect to Dryandra (Proteaceae). Australian Systematic Botany 18 (1): 75-88.

Mast, A. R., & K. Thiele. 2007. The transfer of Dryandra R.Br. to Banksia L.f. (Proteaceae). Australian Systematic Botany 20 (1): 63-71.

Name the Bug # 36

Adam Yates chose to request a post topic as his prize for Name the Bug. So what better way to preview his request than with a Name the Bug post?

Attribution, as always, to follow.

Update: Identity now available here. Photo from here.

More than Four and Twenty Blackbirds

The Tristan thrush or Starchy, Turdus eremita, an endemic bird of the remote Tristan da Cunha group of islands in the South Atlantic, scavenging on a dead penguin (starchies have decidedly more catholic tastes than other thrushes). Photo by Lex.

Turdus, the thrushes, is a large cosmopolitan genus of birds found throughout the world except for Australia. The extent of the genus' circumscription has varied between authorities, though most recent authors exclude the ground-thrushes of the genus Zoothera. Conversely, phylogenetic studies have indicated that the previously monotypic genera Cichlherminia lherminieri of the Caribbean and Nesocichla eremita from Tristan da Cunha should be subsumed within Turdus (Voelker et al., 2007). At present, it seems unlikely that Turdus will be further subdivided; as the basalmost species in the genus is likely to be the mistle thrush Turdus viscivorus, which happens to also be the type of the genus, any subdivision would require that Turdus be reduced to a single species and all other species placed in new genera.

The blackbird Turdus merula, a species found throughout northern Eurasia (and introduced to New Zealand). Only the males are black; the females are dark mottled brown and have grey rather than yellow beaks. Photo by Bence Mate.

Of the 60+ species remaining in Turdus, many are widespread and divided into a number of subspecies that may or may not be promoted to separate species by future researchers. As an extreme example, a study on variation between geographically separated populations of the island thrush Turdus poliocephalus, whose distribution extends from Sumatra and the Philippines east to Norfolk Island* and Vanuatu, suggested that there may be grounds for dividing them between nearly forty diagnostic taxonomic units (Peterson, 2007).

*At least, it did. The Tasman Sea populations of T. poliocephalus have, unfortunately, since shuffled off this mortal coil.

The St Lucia forest thrush, Turdus lherminieri sanctaeluciae. Like T. eremita, this is a distinctive species that was previously placed in its own genus. Photo by Jean-Michel Fenerole.

The base coloration of most species of Turdus can be described as 'mottled brown', though notable exceptions (at least as males) include the grey and red American robin T. migratorius and the blackbird T. merula. Most members of the genus are more highly regarded for their voices rather than their looks, an attribute honoured in both the vernacular and scientific names of the song thrush Turdus philomelos* ("lover of song"). As with other speciose songbird clades, variation in song has turned out to be significant in separating closely related species. Both the Príncipe thrush T. xanthorhynchus (Melo et al., 2010) and the black-throated thrush T. atrogularis (Sangster et al., 2009) differ in their songs (among other things) from species with which they were previously considered conspecific.

*Older references may one of the names Turdus musicus or Turdus ericetorum for this species. Both these names have since been suppressed by the ICZN. The history of Turdus musicus is particularly turgid, as authorities had disagreed over whether the name should be applied to the song thrush or to the redwing (now Turdus iliacus) (Mayr & Vaurie, 1957). Both T. musicus and T. iliacus appeared in Linnaeus' 1758 Systema Naturae. Unfortunately (whether because he was unclear on the distinction between the species, or by a simple composition error), Linnaeus confused the two species' descriptions: under T. musicus, he gave a description of the redwing but provided sources referring to the song thrush, while the entry for T. iliacus attached a description of the song thrush to references referring to the redwing! (The significance of Linnaeus' sources to his descriptions has previously been discussed in the sperm whale nomenclature post.) Mayr & Vaurie's (1957) application buried the name Turdus musicus and designated a neotype to fix Turdus iliacus firmly to the redwing.


Mayr, E., & C. Vaurie. 1957. Proposed use of the plenary powers to suppress the specific name "musicus" Linnaeus, 1758, as published in the combination "Turdus musicus" and to approve a neotype for "Turdus iliacus" Linnaeus, 1758, the Eurasian redwing (class Aves). Bulletin of Zoological Nomenclature 13 (6): 177-181.

Melo, M., R. C. K. Bowie, G. Voelker, M. Dallimer, N. J. Collar & P. J. Jones. 2010. Multiple lines of evidence support the recognition of a very rare bird species: the Príncipe thrush. Journal of Zoology 282 (2): 120-129.

Peterson, A. T. 2007. Geographic variation in size and coloration in the Turdus poliocephalus complex: a first review of species limits. Scientific Papers, Natural History Museum, The University of Kansas 40: 1-17.

Sangster, G., A. B. van den Berg, A. J. van Loon & C. S. Roselaar. 2009. Dutch avifaunal list: taxonomic changes in 2004–2008. Ardea 97 (3): 373–381.

Voelker, G., S. Rohwer, R. C. K. Bowie & D. C. Outlaw. 2007. Molecular systematics of a speciose, cosmopolitan songbird genus: defining the limits of, and relationships among, the Turdus thrushes. Molecular Phylogenetics and Evolution 42: 422-434.

Name the Bug # 35

I promise I'll get the answer to this one up faster than last time:

And for the record, I'm interested in the animal on the left.

Attribution, as always, to follow.

Update: Identity now available here. Photo from here.

A Halfway House, Halfway Down Honshu Island

Epiophlebia superstes from Saitama, Japan. Photo by chochoensis.

The insect order Odonata contains two types of insect, the dragonflies and the damselflies. All living odonates can be placed in one or the other of these groups except for just two species. The two species of the genus Epiophlebia, E. superstes in Japan and E. laidlawi in the Himalayas, are inhabitants of fast-flowing streams in montane rainforests. In overall appearance, they resemble a dragonfly (and are more closely related to dragonflies than damselflies) but they retain a number of primitive features shared with damselflies. Their wings are more like a damselfly's, and like a damselfly they are able to raise their wings directly above their body so that the wings are together at rest. In dragonflies, the wings have a much smaller arc of movement, so when the insect is resting the wings are still held extending outwards. Like all other living odonates, Epiophlebia nymphs are aquatic. Like dragonflies, they have their gills internalised within the rectum* rather than external as in damselflies. However, unlike dragonflies, Epiophlebia nymphs do not jet-propel themselves through the water by firing water out of their rectum, but move about by the slower but more dignified method of crawling. Growth of Epiophlebia nymphs is slow and they can take up to eight years to reach maturity (Tabaru, 1984).

*Yes, that's right, dragonfly nymphs breathe through their arses.

Despite lacking a fossil record of their own (probably due to their montane habitat), Epiophlebia seem to have diverged from the dragonfly lineage very early on, at least as early as the Triassic, as indicated by the presence in that time of Odonata more closely related to dragonflies than Epiophlebia (Grimaldi & Engel, 2005). The Japanese name for Epiophlebia superstes refers to their presumed age: they are mukashitombo, the "dragonfly from long ago".


Grimldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press.

Tabaru, N. 1984. Larval development of Epiophlebia superstes in Kyushu. Tombo 27: 27-31.

Name the Bug # 34

Okay, let's have another one, probably much easier than the last couple have been. Anyone recognise this very distinctive insect from Japan?

Attribution to follow.

The leader board currently has Adam Yates in front with five points, followed by tf with three, intercostal with two and Reprobus with one. Remember, oneupmanship is not only permitted, it is encouraged.

Update: Identity now available here. Photo from here.

Conical Problematica

Scattered throughout the fossil record are little mysteries, organisms whose remains have been preserved but which are not obviously relatable to any more familiar group. Either their remains are too simple to preserve much evidence of their affinities (as with the 'tubular problematica' I've discussed before), or they are too distinct from other organisms for their affinities to be clear, or both, or some other reason. Unless they are particularly common or otherwise significant, most of these problematica are probably doomed to remain so. Case in point:

The figures above show Asymmetroconus splendidus, described by Korde in 1975 from the Albian (early Cretaceous) of the Crimea. The photos are of thin sections of the fossils; the complete skeleton would have probably been shaped rather like a wine goblet. The largest specimens of Asymmetroconus were just under 8 mm in height. In the same paper, Korde described a number of similar fossils aged from the Albian to the Danian (earliest Palaeocene), assigning them all to the new order Asymmetroconida. Korde attributed the asymmetroconidans to the Hydroconozoa, a group of similar fossils he had himself described previously from the early Cambrian. Hydroconozoa have generally been assigned to the Cnidaria, though their exact position therein remains obscure. Asymmetroconida resembled hydroconozoans in being small and goblet-shaped, with a conical interior to the cup and a basal globular hollow below the point of the cone. However, they differed from Cambrian hydroconozoans in their skeletal microstructure and in the asymmetry of the cup, with one side much thicker than the other. Rozanov & Zhuravlev (1992) later dismissed the idea of Mesozoic hydroconozoans, stating simply that structures described as such had 'little in common with this group'. No alternative identification of the Asymmetroconida has ever been proposed, and they do not appear to have been properly studied since Korde's original description.

Reconstruction of the hydroconozoan Hydroconus mirabilis, from Rozanov & Zhuravlev (1992). Whether actually related or not, the Asymmetroconida would have probably looked superficially similar.


Korde, K. B. 1975. [Hydroconozoa from Cretaceous and Palaeocene deposits of the Crimea]. In: Shimansky, V. N., & A. N. Soloviev (eds) Razvitie i smena organičeskogo mira na rubeže Mezozoâ i Kajnozoâ. Novye dankye o razvitii fauny pp. 32-38. Nauka: Moscow. [in Russian]

Rozanov, A. Yu., & A. Yu. Zhuravlev. 1992. The lower Cambrian fossil record of the Soviet Union. In: Lipps, J. H., & P. W. Signor (eds) Origin and Early Evolution of the Metazoa pp. 205-282. Plenum Press: New York.

Name the Bug # 33

I apologise that this is not the best possible reproduction of an image. In my defense, it is quite possible that even in the original publication the image quality was poor.

The organism shown grew to just under 8 mm in height, and dates to the Albian epoch of the early Cretaceous.

Oh, and it's only fair to give you all a warning: in the past, I've put up what turned out to be some pretty evil ID challenges. I think I can safely say, however, that this is my most evil challenge yet. Any other challenge I have put up, no matter how evil, was but a room full of fluffy white kittens paddling their paws in pink marshmallow compared to this challenge. If this challenge came across a fluffy white kitten and a bowl of pink marshmallow, it would use the marshmallow to drown the kitten. This challenge feasts on the flesh of virgins while bathing in their blood and idly drawing pentagrams alongside its pool. Rumour even has it that this challenge played a significant role in the chart success of Hear'Say. Seriously, though, if you enter the name of this challenge into Google, you will get one result, and that's the original description from whence these figures are taken.

Attribution, as always, to follow.

Update: Identity now available here. Figure from Korde (1975).

A little Linguipolygnathus

Variants of Linguipolygnathus linguiformis over time, from Bardashev et al. (2002).

Three points for this ID challenge go to Adam Yates who recognised the objects in the figure as P-elements of an ozarkodinid conodont (the first person to identify them as a conodont looses out on points because they didn't supply any supporting comments). Linguipolygnathus linguiformis is the type species of Linguipolygnathus, one of the genera carved by Bardashev et al. (2002) out of the large older genus Polygnathus. I've commented on the taxonomic insanity of Bardashev et al. in a previous post, though the idea of subdividing Polygnathus is not in itself a bad one (and note that if Linguipolygnathus were synonymised with its supposed polyphyletically-ancestral genus Eolinguipolygnathus we'd be left with a single monophyletic genus).

Many discussions of conodonts make reference to their minuteness (I've done it myself in the past) and the preserved conodont fossils are certainly minute. However, I must confess to only realising fairly recently that, just because the preserved fossils are minute, doesn't necessarily mean that the (largely soft-bodied and hence rarely preserved) animals themselves were. Of the two best-preserved body fossils of conodonts available to us, the remains of Promissum are those of an animal about 20 cm long. Even the more modestly sized Clydagnathus, which is apparently more like the usual run of conodonts, would have been about 6 cm long in life: not huge, but still comparable in size to a modern anchovy.


Bardashev, I. A., K. Weddige & W. Ziegler. 2002. The phylomorphogenesis of some Early Devonian platform conodonts. Senckenbergiana Lethaea 82 (2): 375-451.

Name the Bug # 32

Does this mean anything to anyone?

As before, you can get three points for the best answer, two points for second-best and one point for third best. And remember, the best answer will not necessarily be the first with the most accurate identification.

Update: Identity now available here. Figure from Bardashev et al. (2002).

Snails that Never See the Light of Day

Diagrams of Hauffenia tellinii from Bodon et al. (2001). Figure 67 is the shell, figures 68-71 are opercula, 72-75 are male anatomy, 76-80 are female anatomy.

Hauffenia is a genus of freshwater snails of the family Hydrobiidae found in south-eastern Europe (Slovenia, Croatia, adjoining parts of Italy and Austria, etc.) Hydrobiids are a very diverse but very minute group of gastropods: Hauffenia species, for instance, are less than three millimetres in diameter and less than 1.5 millimetres in height. Hauffenia species differ from many other hydrobiid genera in having much flatter shells with only the slightest of turrets. This shape, which commenter 'tf' described as "almost but not quite planiform", is known as 'valvatiform', after Valvata, another freshwater snail with a similar shell. Though the shells of Hauffenia and Valvata are similar enough that the two genera have been confused in the past, the internal anatomy of Valvata shows that it is not a hydrobiid or even closely related. Hydrobiids belong to the major gastropod clade known as caenogastropods but Valvata is a heterobranch, more closely related to garden snails or sea slugs than to hydrobiids (Dayrat & Tillier, 2002). Valvata species are also hermaphroditic while hydrobiids such as Hauffenia have separate males and females.

Distinguishing Hauffenia from other valvatiform hydrobiids is difficult and requires examination of the internal anatomy. Bodon et al. (2001) characterised Hauffenia as possessing a penis with a stylet in the male, while females possessed proximal seminal receptacles only (no distal receptacles) and a reduced bursa copulatrix. Identifying these characters can be difficult because Hauffenia species are subterranean, mostly living in caves and springs in limestone karsts though the Hungarian Hauffenia kissdalmae was recently described from a spring in andesite (Erőss & Petró, 2008), making collecting fresh material difficult. Most early studies on Hauffenia were based on shell morphology only, and species previously assigned to the genus from western Europe or North America were regarded by Bodon et al. as belonging to other genera. Even more doubtful is the assignation of Miocene marine fossils to this genus, refuted by Iljina (2010) on the basis that the Hauffenia-like opercula attributed to the fossils were probably not validly associated, and possibly not even gastropod opercula. Some Hauffenia species, such as H. tellinii in the figure at the top of this post, possess a distinctive knob on the inside of the operculum that distinguishes them from other valvatiform hydrobiids. Earlier authors distinguished separate subgenera in Hauffenia based on whether or not a species possessed such a knob, but Bodon et al. (2010) did not use such a formal distinction.

And if you've just been reading this post to see who won the ID challenge, three points go to 'tf' who recognised the animal as a hydrobiid and provided the diagnostic features; two points go to 'intercostal' who mistook it for a valvatid but pointed out that the presence of an operculum meant that it couldn't be a pulmonate.


Bodon, M., G. Manganelli & F. Giusti. 2001. A survey of the European valvatiform hydrobiid genera, with special reference to Hauffenia Pollonera, 1898 (Gastropoda: Hydrobiidae). Malacologia 43 (1-2): 103-215.

Dayrat, B., & S. Tillier. 2002. Evolutionary relationships of euthyneuran gastropods (Mollusca): a cladistic re-evaluation of morphological characters. Zoological Journal of the Linnean Society 135 (4): 403-470.

Erőss, Z. P., & E. Petró. 2008. A new species of the valvatiform hydrobiid genus Hauffenia from Hungary (Mollusca: Caenogastropoda: Hydrobiidae). Acta Zoologica Academiae Scientiarum Hugaricae 54 (2): 159-167.

Iljina, L. B. 2010. On the taxonomic position of Miocene valvatiform gastropods and their ecological features. Paleontological Journal 44 (4): 391-394.

Name the Bug # 31

Last week, I put up an ID challenge that I thought would be relatively simple but which was apparently more difficult. Does that mean that today's challenge, which I think is damn near impossible, is really quite simple?

As a clue, the animal shown lives in fresh water. The scale bar represents 1 mm for all figures except fig. 73, for which it represents 0.5 mm. Attribution to follow.

The last few ID challenges have been getting good responses, so I think it may be time to up the stakes a little. Like Alex Wild does for his challenges, I'm going to award points for correct answers. But unlike Alex, I'm not going to have any set rules about how those points are divvied up. Basically, three points will be awarded to the person who gets the closest to the correct identification. Bonus points (one or two points) may be awarded to up to two less precise answers that contribute to the correct identification (say, if someone identifies the family but not the species, or draws attention to a significant diagnostic feature).

But that's not all there is to it, because I'm also going to take a certain degree of inspiration in awarding points from QI. If you want to win points, but find that someone else has already correctly identified the species, don't despair: if you can provide a better answer than the one they gave, you may be able to steal their three-point spot off them! Examples of better answers would be if you provide a better explanation of the diagnostic features, or if you tell me something really interesting about the organism in question. If you do bump the previous leader from the top spot, they'll be knocked down to the two-point position. Even if you don't successfully knock down the leader, you may be able to get the two-point or one-point position yourself. Unless, of course, someone else pushes you out in turn. Also, I should note that all three point-spots are awarded entirely at my discretion: if there are not suitable answers, I may not award one or more of the point slots.

The first person to get a total of ten or more points from successive Name the Bug posts will win the first round. I'm still deciding what exactly you'll win; my current inclination is to give you a choice between requesting a post on the topic of your choice, or of being given a guest post on Catalogue of Organisms.

As this is the first round, I'll be generous in my judgement of which comments are worthy of the points (so please have a go—after all, I'd look a right twit if I didn't get to award any points in the first challenge!) Let battle commence!

Wigs and Wings and Other Things

Congratulations go to Adam Yates for successfully identifying this animal:

Arixenia esau, photographed in Deer Cave in Sarawak by Alan Cressler.

This "very interesting, though repulsive, insect" (to use the words of Hebard in 1927) is a member of today's Taxon of the Week, the Neodermaptera. Neodermaptera is the clade containing all living members of the Dermaptera, the earwigs, distinguished from various stem groups of the Dermaptera by features such as three-segmented tarsi and the lack of veins in the forewings (Engel 2003). Earwigs are one of the few groups of insects other than beetles to have the forewings hardened into elytriform cases, which in earwigs have also been greatly reduced in size (in earwigs, the hardened forewings are referred to as 'tegmina' rather than 'elytra', but these seem to be just different words for much the same sort of thing). The rarely-seen hindwings remain folded under the tegmina unless the earwig is flying (which they do not often do) and are simply bizarre. One of the characteristics of polyneopterans, the group of insects including crickets, cockroaches, earwigs and various others, is a tendency towards enlargement of the anal fan, the posterior part of the wing; in earwigs, the anal fan of the hindwing has become enlarged to the point that the wing is almost entirely anal fan with the anterior parts of the wing greatly reduced and crammed into a small toughened section towards the base. One of the stories floating about to supposedly explain the origin of the name 'earwig' claims that it is a corruption of 'ear-wing'. While the wings are indeed ear-shaped, the story rather loses credibility in face of the detail that the average person would probably never see them.

Earwig (probably a female Doru using the key in Engel 2003) with its wings spread, showing the semicircular shape and radiate anal veins. Photo by Sean McCann.

The other distinctive feature of most living earwigs is the development of the cerci at the end of the abdomen into a pair of large, hard forceps. The forceps are used for defense as well as capturing prey in those species that eat animal matter (most earwigs are omnivorous); they may also be used to help fold the wings under the tegmina. In most species, the males have heavier forceps than the females. The only earwigs to have filamentous cerci rather than forceps are the Arixeniidae and Hemimeridae, two families that live in association with mammalian hosts. Arixeniids (such as Arixenia in the top picture) are about 2 cm long and live on bats in south-east Asia; hemimerids are about half that size and live on African giant rats. Not are they distinctive among earwigs, they are the only known quasi-parasitic polyneopterans—I say 'quasi'-parasitic because they probably feed more on dead skin and host secretions than the actual living host itself. The arixeniids probably feed mostly on the rich deposits of bat poo in host roosting sites. Because of the lack of forceps and other features, these two families have often been placed in separate suborders from the remaining earwigs; at least one author argued that hemimerids should be removed from Dermaptera entirely and treated as a separate order. However, the current consensus is that the two families are probably derived from more normal earwigs, with their distinctive features being adaptations to their symbiotic lifestyles.

Forceps of the recently extinct Labidura herculeana of St Helena, the largest known earwig, alongside a 22 mm specimen of its more average close relative L. riparia. Photo by Philip Ashmole.

Another distinctive feature of the two mammal-associated families is that they are live-bearers. In all other families, the female lays a batch of eggs, usually in a burrow, that she watches over until the young hatch out. She continues to protect her young for their first one or two instars; after that they must fend for themselves. In fact, if the young do not move out quickly enough, their mother will eat them (Rentz & Kevan, 1991). Something, perhaps, to be kept in mind by all those parents who feel their adult offspring are taking too long to get their own place.


Engel, M. S. 2003. The earwigs of Kansas, with a key to genera north of Mexico (Insecta: Dermaptera). Transactions of the Kansas Academy of Science 106 (3-4): 115-123.

Hebard, M. 1927. Studies in Sumatran Dermaptera. Proceedings of the Academy of Natural Sciences of Philadelphia 79: 23-48.

Rentz, D. C. F., & D. K. McE. Kevan. 1991. Dermaptera (earwigs). In: CSIRO. The Insects of Australia, 2nd ed., vol. 1, pp. 360-368. Melbourne University Press.

Name the Bug # 30

If I'm continuing with the supposed pattern of these challenges, it's time for an easy one:

But because it is relatively simple, I'm going to want at least a genus ID (and preferably some supporting info).

Attribution to follow.

Update: Identity available here. Photo from here.

Archechiniscus: Distinctively Indifferent

Archechiniscus marci. Figure from Pollock (1976).

Archechiniscus is a genus of three species of marine tardigrade found in littoral habitats. They can be readily distinguished from other marine tardigrades by their unique arrangement of claws: two pairs, with the internal pair on the end of a long pair of toes but the external pair set directly onto the foot. Most of the other distinguishing features of Archechiniscus are more negative: they lack conspicuous segmentation or ornamentation. The presence of cephalic appendages marks Archechiniscus as belonging to the heterotardigrades rather than the eutardigrades; within the Heterotardigrada, it belongs to the paraphyletic 'arthrotardigrade' group. Opinions have differed as to whether it should be placed in the family Halechiniscidae or in its own separate family; Jørgensen et al. (2010) identified the broad Halechiniscidae as polyphyletic and plumped for placing Archechiniscus in its own family (though potentially as the sister group of their more restricted Halechiniscidae).

As for most marine tardigrades, there doesn't appear to be a great deal of info about the lifestyle of Archechiniscus. Archechiniscus symbalanus got its name due to being collected in association with barnacles (Chang & Rho, 1998) but I don't know what it was doing there. As littoral inhabitants, Archechiniscus are resistant to a higher degree of desiccation than other marine tardigrades (Jönsson & Järemo, 2003) but do not show the extremes of resistance found in some other tardigrades (remember, not all tardigrades are resistant to adverse conditions, and not all tardigrades are resistant to the same adverse conditions).


Chang, C.-Y., & H.-S. Rho. 1998. Three new tardigrade species associated with barnacles from the Thai coast of Andaman Sea. Korean Journal of Biological Sciences 2: 323-331.

Jönsson, K. I., & J. Järemo. 2003. A model on the evolution of cryptobiosis. Annales Zoologici Fennici 40: 331-340.

Jørgensen, A., S. Faurby, J. G. Hansen, N. Møbjerg & R. M. Kristensen. 2010. Molecular phylogeny of Arthrotardigrada (Tardigrada). Molecular Phylogenetics and Evolution 54 (3): 1006-1015.

Pollock, L. W. 1976. Marine Flora and Fauna of the Northeastern United States. Tardigrada. NOAA: Seattle.

Name the Bug #29

I've been accused of swinging wildly between the easy and the impossible for these ID challenges. The last one was fairly easy, so I suppose it's time for the impossible:

To give a bit of a hand, the animal shown is a marine species of its phylum.

Attribution to follow.

Update: Identity available here. Figure from Pollock (1976).

Borage and Comfrey and Bugloss

Anchusa undulata ssp. granatensis. Photo by James Gaither.

The tribe Boragineae includes about 170 species of herbaceous flowering plants, mostly found in the Palaearctic region with only a couple of species extending into southern Africa. The group is well-distinguished by the presence of what are called fornices, the whitish lobes at the base of each petal that you can see in the photo above, as well as features of their seeds. Many Boragineae seeds have an elaiosome, a fatty plug at one end that attracts foraging ants (Hilger et al., 2004). The ants carry the seed back to their nest as food, but the plant produces enough seeds that at least some will not be eaten but will be able to germinate after being carried under the ground and away from anything else that might eat them.

Abraham-Isaac-Jacob, Trachystemon orientalis, a native of forests around the Black Sea and one of the more unusual species of Boragineae. Apparently the unusual name refers to the flowers changing colour as they age. Photo by Daniel Mosquin.

The species are divided between about fifteen genera (the exact number varies depending on whom you ask). The largest generally-recognised genus, Anchusa (the buglosses), was identified by Hilger et al. (2004) as para-/polyphyletic with a number of smaller genera also nested within the Anchusa clade, suggesting that the currently recognised constituent subgenera may need to be recognised as separate subgenera (or else the genera Lycopsis and Cynoglottis submerged into Anchusa). Other relationships within the tribe recognised by this and other studies include a close relationship between the genera Borago (borage) and Symphytum (comfrey), and between Nonea and Pulmonaria (lungwort). The basalmost member of the tribe is Pentaglottis sempervirens, which is also the only member of the tribe found in the Atlantic region of southwest Europe. The relict distribution of this species, as well as the concentration of diversity for the tribe overall, have been cited as supporting a Mediterranean origin for the Boragineae.

Green alkanet, Pentaglottis sempervirens, the sister species to all other Boragineae. Photo by Carl Farmer.

A number of members of the tribe have long been cultivated and many are even labelled by their botanical names as officinal (the Medieval Latin term 'officinalis' refers to a plant or substance that is kept in an apothecary; not surprisingly, many plants with supposed medicinal values are also eaten for their nutritional values). Borago officinalis, borage, is used as a salad or pot herb in Europe. Symphytum officinale, comfrey, has also been widely used medicinally, mainly for external uses such as soothing bruises (some of the properties attributed to comfrey verge on the ridiculous: a bath steeped in comfrey was supposedly able to restore a woman's virginity). Pulmonaria officinalis, lungwort, received its name because of the supposed resemblance of its blotchy leaves to lung tissue. Under the unabashedly loopy herbalist principle known as the Doctrine of Signatures, this outward resemblance indicated its suitability in treating lung diseases such as tuberculosis (in fact, lungwort contains toxic alkaloids that make it dangerous to take internally).


Hilger, H. H., F. Selvi, A. Papini & M. Bigazzi. 2004. Molecular systematics of Boraginaceae tribe Boragineae based on ITS1 and trnL sequences, with special reference to Anchusa s.l. Annals of Botany 94 (2): 201-212.

Name the Bug # 28

Tomorrow's Taxon of the Week will certainly be bringing the pretty:

Anyone recognise it?

Attribution to follow.

Update: Identity now available here. Photo from here.

Thoughts Inspired by a Private Publication

A number of sources, including this message on the DML archive, drew my attention yesterday to the existence of a recent online publication proposing a new North American sauropod species 'Amphicoelias brontodiplodocus', as well as synonymising a whole slew of other familiar dinosaur taxa (such as Apatosaurus and Diplodocus) under the relatively unfamiliar name Amphicoelias. Quite apart from 'Amphicoelias brontodiplodocus' perhaps being the most intensely unaesthetic name ever proposed for a dinosaur (and this in a field including such wince-inducing monikers as Raptorex and Tyrannotitan!), the pdf has once again lifted the lid on a number of arguments that have been simmering away for some time now. I recommend reading the responses to the DML message linked to above, as well as Mike Taylor's discussion at SV-POW. I'm not in a position to discuss the technical details of the 'brontodiplodocus' pdf itself but I would like to discuss some of the broader issues raised by it, the questions of regulating publication and of publications based on privately owned specimens.

It should be noted that the 'brontodiplodocus' pdf currently appears to be an online document only and hence not validly published in the view of the ICZN (and whatever its publication status, there is no obligation on anyone else to accept the proposed synonymies). However, Mike Taylor has pointed out that it would be a simple matter for the publishers of the document to produce it in printed form and thereby validate it. Also, it would be easy for a non-expert in taxonomic procedure (and even a few supposed experts in taxonomic procedure) to mistake the current pdf for a valid publication as it is. As I've noted before, end-users of taxonomy should not be required to consider ICZN esoterica every time they are presented with a new name, just as non-specialist computer users should not always have to familiarise themselves with thousands of lines of computer code before using a new word-processing programme.

While self-publication of poor-quality works has always been a potential issue for taxonomy and the ICZN, modern technology has made self-publication much easier than before. It can now pontentially be done by anyone with access to a word processor of some form: i.e. pretty much everyone in the developed world and a large number of people in the developing world. Some have suggested that, to counteract this issue, only names published in peer-reviewed journals should be accepted as valid (I've pointed out why I disagree with this proposal in an earlier post). Others have suggested taking this even further and establishing a single journal for the publication of all zoological taxonomy. This is already the method used by the Prokaryote Code of Nomenclature, which requires that all new bacterial taxa be published or validated in the International Journal of Systematic and Evolutionary Microbiology. However, it is debatable whether such a model could be applied to the ICZN. Firstly, the sheer number of new zoological names published each year is much higher than the number of bacterial names, perhaps by a few orders of magnitude. A single zoological taxonomic journal would be a major undertaking, especially for an organisation such as the ICZN which lacks any major sources of funding on which to draw. Secondly, the Prokaryote Code of Nomenclature has very high basic requirements for a taxon to be considered published (such as deposition of cultures of the type strain in two separate collections in separate countries). It is unlikely that the ICZN would be able to implement such across-the-board requirements because it deals with a much broader range of organism types (such as fossils and protozoans) than the bacterial code, each of which may have their own specialised requirements for appropriate typification.

In addition, one of the reasons that prokaryote nomenclature functions so well under these restrictions is the usage of a wide range of what might be called 'grey taxa', taxa widely regarded as recognisable but which, for various reasons (most commonly their inability to date to be cultured in the laboratory), cannot be 'validly' published (one such taxon that I've previously discussed is the minute 'Nanoarchaeum equitans'). Bacterial nomenclature therefore has two distinct nomenclatural classes of taxa. This is not necessarily a problem: 'grey taxa' cannot compete for priority with 'valid taxa', for instance, which reduces the chance of an old poorly-characterised name supplanting a newer familiar one.

The current version of the ICZN does allow in Article 79 for the potential publication of Lists of Available Names for selected taxa. Under this article, a list could be published of (for instance) all published bird names that would become the definitive listing for that group. Any names published prior to the list's end-date that were not included are regarded as unavailable (and hence not competing for priority with names included in the list), even if they normally would be under the general rules of the code (the Prokaryote Code followed this path with the publication of the Approved Lists that were the foundation of current bacterial nomenclature). To date, I am not aware of any such list being successfully ratified by the ICZN (it would not be a simple process), but the facility is potentially there. At present, the Lists of Available Names are only intended to clarify the status of previously published names rather than regulate any published subsequent to the relevant list. However, even if it is not currently feasible to introduce a single journal model for all zoological nomenclature, perhaps it could be done for specific groups? Imagine if a List of Available Names could be published for (say) dinosaurs, with a requirement that all subsequent names be validated by a prominent journal such as the Journal of Vertebrate Paleontology? (On the flipside, there would have to be a requirement that the journal must validate all names that meet certain pre-existing requirements [as currently exists for the IJSEM] in order to prevent names being refused for reasons unrelated to the diagnosability of the taxa concerned, such as personal disputes between taxonomists.) If done on a group-by-group basis, such a process might also allow for workers in each group to determine the appropriate requirements for that group.

The second major issue raised by the 'Amphicoelias brontodiplodocus' pdf is that its publishers are commercial fossil dealers and at some point the brontodiplocus 'holotype' may (probably will) be privately sold. The ICZN currently recommends, but does not require, that the type material for new taxa be deposited in publicly accessible collections. If a specimen is privately held, it is less likely to be available for study by future researchers (however, it is worth noting that a privately held specimen is not always unavailable, nor is a specimen in a public collection necessarily available). In the case of fossils, some people are of the opinion that their commercial sale should be banned completely in order to prevent scientifically significant material from entering private hands. However, not all fossil material is scientifically significant—some fossils are extremely abundant (to the extent that some localities have profitable fossil mines) and there would seem to be little sense in banning the sale of a fossil species for which thousands are already held in public collections. If the possibility is floated of only banning the sale of 'scientifically significant' specimens, we run afoul of the problem of specifying how to determine which specimens are scientifically significant. For instance, articulated Tyrannosaurus skeletons are very rare and of great scientific interest, but isolated Tyrannosaurus teeth are very common (or so I've heard) and of little scientific interest. Where exactly does one draw the line?

Type specimens, however, are by definition of scientific interest, and it is usually clear whether a specimen is a type. So while it would be somewhat ridiculous to ban the commercial sale of all fossils, perhaps it would be reasonable to preclude the commercial sale of type material? Note that I am not suggesting banning the use of privately held material in describing taxa, only that such material could not be subsequently traded*. I can think of two potential issues that might have to be considered in such a scenario. One is that commercial collectors may refuse to allow any examination of material that they hold by researchers to prevent its effective devaluation (of course, some may not see this as a problem). Another potential problem would be if a privately owned specimen is made part of the type material of a new species without the informed consent and/or involvement of the specimen's owner.

*The distinction between privately and publicly owned specimens is a lot fuzzier than those working outside the taxonomic field might expect. Many new species are described by authors on the basis of specimens they have themselves collected; while the authors may intend on eventually depositing the specimens in a public collection, this may not have yet happened at the time of publication. Depending on circumstances, it may be some time before the specimen(s) are finally passed on to the collection**.

**To give an example from my own experience, two years ago I published the new harvestman species Templar incongruens based on two specimens previously held at the Western Australian Museum that had been collected in Canterbury, New Zealand. It was decided that the new types should be transferred to the Canterbury Museum as it seemed more appropriate for them to be held in a collection in their home country, and this was the depository recorded in the publication. However, because T. incongruens has also formed part of my phylogenetic analysis of Monoscutidae currently in preparation, the specimens have not yet actually made it to Canterbury: they're still sitting in my lab here in Perth.

Small lizards of South America

Three readers made comments about the apparent identity of yesterday's ID challenge; none of them, I'm afraid, even came close.

The gymnophthalmid lizard Leposoma hexalepis, photographed in Venezuela by Carl Franklin.

Gymnophthalmids are a family of nearly 200 species (new ones continue to be described at a steady rate) of small (4-15 cm excluding the tail) insectivorous lizards from South America. So little regarded is this family that no really good vernacular name exists for it and its members are generally referred to by what they are not: they are referred to as 'microteiids' in contrast to the related but physically larger family Teiidae. Today's Taxon of the Week is a clade within the Gymnophthalmidae known as the Ecpleopini or Ecpleopinae*, depending on whom you ask.

*Technically, that should be 'Ecpleopodini', but all recent publications have used the 'incorrect' spelling (so far I've only seen one publication from 1887 use the correct spelling). The online page for one recent article includes a footnote mentioning the correct spelling but the note does not appear to be present in the printed article.

The Ecpleopini include (at present) about thirty species, about half of which are placed in the genus Leposoma. The clade is currently supported by molecular analyses without any identified morphological synapomorphies (Pellegrino et al., 2001; Rodrigues et al., 2005). Different analyses recover different relationships between the constituent genera except for a small clade of the genera Colobosauroides, Dryadosaura and Anotosaura (Rodrigues et al., 2005). This clade represents one of a number of lineages within Gymnophthalmidae to develop an elongate body form and reduced limbs together with a fossorial lifestyle. Anotosaura has also lost its external ear openings.

Another ecpleopin, Arthrosaura reticulata, photographed in Peru by Thomas Stromberg.

The species of the genus Leposoma are more generalised in their overall appearance but still not without their intrigues. Leposoma species can be divided between two groups distinguished by their chromosome number and arrangement. The L. scincoides group possess 52 chromosomes of a range of sizes while species of the L. parietale group ancestrally possess 44 chromosomes with a clear size distinction between 20 major and 24 minor chromosomes. The only exception to this pattern is L. percarinatum, a parthenogenetic species* from Mato Grosso in Brazil with 66 chromosomes: one of the few known examples of a triploid genome in vertebrates. When the triploid nature of L. percarinatum was identified, it was suggested that it might be derived from a hybridisation event between two diploid parents. Since then, a diploid form of L. percarinatum has also been identified that may represent one of the parents of the triploid form; perhaps the other is the sympatric bisexual** diploid L. ferreirai (Laguna et al., 2010). Which does still leave the question of how the usual parthenogenesis of the diploid L. percarinatum came to be in the first place.

*Or, as it seems to be called, a 'parthenoform' (presumably to avoid having to refer to a parthenogenetic taxon as a 'species').

**In the sense of possessing two sexes, not the other sense.


Laguna, M. M., M. T. Rodrigues, R. M. L. dos Santos, Y. Yonenaga-Yassuda, T. C. S. Ávila-Pires, M. S. Hoogmoed & K. C. M. Pellegrino. 2010. Karyotypes of a cryptic diploid form of the unisexual Leposoma percarinatum (Squamata, Gymnophthalmidae) and the bisexual Leposoma ferreirai from the lower Rio Negro, Amazonian Brazil. Journal of Herpetology 44 (1): 153-157.

Pellegrino, K. C. M., M. T. Rodrigues, Y. Yonenaga-Yassuda & J. W. Sites Jr. 2001. A molecular perspective on the evolution of microteiid lizards (Squamata, Gymnophthalmidae), and a new classification for the family. Biological Journal of the Linnean Society 74: 315-338.

Rodrigues, M. T., E. M. X. Freire, K. C. M. Pellegrino & J. W. Sites Jr. 2005. Phylogenetic relationships of a new genus and species of microteiid lizard from the Atlantic forest of north-eastern Brazil (Squamata, Gymnophthalmidae). Zoological Journal of the Linnean Society 144 (4): 543-557.