Field of Science

Jewels among Beetles

There are many contenders for the title of most stunning-looking insect but there is no question that the jewel beetles have a place among the line-up. Some of these brilliantly coloured insects look as if they could have been sculpted from gleaming metal:

Buprestis niponica, copyright Kohichiro Yoshida.

Buprestis is a genus of jewel beetles found in the Holarctic region, with the greater diversity around the Mediterranean and North America. Somewhere between forty and eighty species are recognised, depending on whether the closely related genera Cypriacis and Yamina are regarded as distinct or not. Species of Buprestis come in a variety of colours, with green, blue or black backgrounds often patterned with yellow or red.

Female Buprestis octoguttata ovipositing, copyright Christian Fischer.

Despite their attractive appearance, jewel beetles are not always welcome. They spend the larval part of their life cycle burrowing into wood so some are known for damaging timbers. The preferred hosts of most Buprestis species, where known, appear to be conifers such as pine, spruce and larch. They primarily attack dead and dying wood, and females of some species are known for searching the trunks of trees following fires to find where protective bark has cracked open (some jewel beetle species in other genera are commonly known as 'fire beetles' in reference to this habit). Buprestis larvae have been claimed to live for extraordinarily long periods. Mature beetles have been observed emerging from furniture and the like multiple decades after the original tree was felled, leading to claims of larval life spans of up to 51 years! It almost goes without saying that such inferences have attracted their share of scepticism, with detractors suggesting the possibility of eggs being laid after the wood was already worked. It is true that the low nutritious value of dry wood might be expected to lead to slow development, but how slow are you willing to believe?

Hairy-Winged Barklice

Forewing and fore tibia of Siniamphipsocus fusconervosus, from Mockford (2003). Scale bar for the femur = 0.1 mm.

For my next semi-random post, I drew Siniamphipsocus, a genus of more than twenty species of barklice known from eastern Asia. Most of these species were described by China by the almost ludicrously prolific psocopterologist Li Fasheng who over the course of his career has described close to 1000 psocopteran species—nearly a fifth of the world's barklouse fauna. It should be noted, though, that this productivity has not entirely come without criticism: for instance, in the case of the Siniamphipsocus species, most if not all are known from a single sex with some described from males and others from females (Li 2002).

Siniamphipsocus is a genus of the Amphipsocidae, a family of barklice most easily recognised by their wings which have a double row of setae along each of the veins. Amphipsocids can be relatively large as barklice go: the largest Siniamphipsocus species, S. aureus, has a body length of four millimetres, with the forewings being up to 6.75 millimetres long. Features distinguishing Siniamphipsocus from other amphipsocids include the absence of the brush of hairs present at the base of the hind wing in many other species, the absence of a spur vein in the rear of the forewing pterostigma, and the presence of a row of minute spines along the fore femur (Li 2002). Distinguishing the individual species of the genus requires fine attention to details such the patterns of markings on the face, the proportions of the wing veins, and details of the genitalia.


Li F. 2002. Psocoptera of China (2 vols). Science Press: Beijing.

Mockford, E. L. 2003. New species and records of Psocoptera from the Kuril Islands. Deutsche Entomologische Zeitschrift 50 (2): 191–230.

Sordariomycetidae: Soil Fungi A-Plenty

I'm pretty sure I've commented before that, although most of us tend to associate the word 'fungi' with mushrooms and other eye-catching fruiting bodies, the vast majority of fungal diversity is minute and tends to go unnoticed. Nevertheless, despite their obscurity, many of these microfungi are crucial to our own continued existence. These are the decomposers, the organisms that break down fallen plant matter and animal wastes in their own search for nourishment and so contribute to the release of locked-up nutrients back into the environmental cycle.

Neurospora growing on sugar cane waste, from here.

The group of fungi that I drew for today's post, the Sordariomycetidae, is primarily made up of these minute decomposers. Sordariomycetids have already made an appearance here at Catalogue of Organisms, in a post from ten years ago on black mildews. Depending on how broadly the group is circumscribed, the Diaporthales could also be included. Due to a simple morphology that provides few distinct characters, the Sordariomycetidae are primarily defined on the basis of molecular phylogenies. The difficulty of classifying microfungi by morphology alone is underlined by cases where species previously classified within the same genus have proven to belong to entirely distinct fungal lineages.

In general, the vegetative body of most Sordariomycetidae consists of little more than disassociated hyphae embedded in their substrate, with the only distinct structures being the reproductive fruiting bodies. These are perithecia: that is, globular or flask-shaped fruiting bodies with a single small opening or ostiole at the top through which the mature spores are released. In some cases, the internal structure of the mature perithecium will simply dissolve, freeing the spores to escape through the ostiole in the manner of a miniature puffball. In others, the spores become entangled in a long strand or seta that is then extruded through the ostiole like toothpaste being squeezed out of a tube.

Perithecium of Chaetomium extruding spore-bearing setae, from here.

Sordariomycetids are found in almost every habitat imaginable: as well as soil- and dung-dwelling forms, they may also be found in aquatic and even marine habitats. Perhaps the best-known sordariomycetid is Neurospora crassa, red bread mould, which is widely used in laboratories as a model organism for genetic research. Indeed, it was investigations into N. crassa in the 1950s that first led to the proposal of the 'one gene, one enzyme' model that became a cornerstone of molecular genetics.

A Western Rockweed

Rockweed Silvetia compressa, from here.

Silvetia compressa is a species of brown alga found on shorelines on the western coast of North America, from British Columbia to Baja California. It is a member of the wrack family Fucaceae that I covered in an earlier post. Silvetia compressa is found in midtidal habitats, generally higher up on the shoreline than other large seaweeds. Individual thalli can reach a maximum length of about three feet (90 cm) but are often smaller. This is a slow-growing species, so patches of Silvetia are slow to recover from damage due to trampling and other disturbance. Please try to avoid walking on the rockweed!

Thalli of Silvetia compressa are composed of thin strands a few millinetres in width with irregular, dichotomous branching. Strands of the thalli lack a midrib (distinguishing them from some other Fucaceae species found in the same area). The width of the strands and regularity of the branching varies with environmental conditions: for instance, individuals growing in locations with stronger wave action have more robust strands that branch more frequently. As with other Fucaceae, the reproductive structures a produced on swollen branch tips called receptacles, but these receptacles do not become inflated with gases and buoyant like those of other species. The exact size and shape of the receptacles is, again, variable.

In many older references, Silvetia compressa may be referred to as Pelvetia fastigiata. The supposed species 'Fucodium compressum' and 'F. fastigiatum' were originally distinguished on the basis that the latter was smaller than the former with more fastigiate branches (that is, the branches remained subparallel). As indicated above, these characters represent the effects of environmental conditions, not fixed differences (Silva 1996). They were eventually included in the genus Pelvetia, together with the Atlantic species P. canaliculata, on the basis of the thalli without a midrib, and the production of just two eggs from each oogonium in the receptacles. However, later analyses supported the separation of the Atlantic and Pacific species of Pelvetia. Not only did they not form a clade in molecular analyses, the eggs in the oogonia were separated by a horizontal division in the Atlantic species but a longitudinal or oblique division in the Pacific species (Silva et al. 2004). As such, the Pacific Pelvetia were transferred into a new genus Silvetia.

Further taxonomic complications involved subspecific variation in Silvetia compressa. A distinctive form of 'Pelvetia fastigiata' found at Pebble Beach in California's Monterey Bay, with smaller, finer thalli and more abundant, regular branching, was labelled as a separate forma gracilis. Similar individuals were also found on the islands off California's coast. However, when Silva (1996) examined the original type specimen of P. fastigiata, he discovered that it was an individual of this 'gracilis' form, not the more typical larger form. Later, Silva et al. (2004) examined genetic variation within the Silvetia compressa of California and Baja California. They found that the individuals of the offshore islands were indeed genetically distinct from continental individuals. As well as the differences in growth habit, there was also some difference in receptacle shape: the continental form had receptacles that tended to be linear and pointed whereas those of the island form were ellipsoidal and blunt. However, the island form still could not be labelled with either of the 'fastigiata' or 'gracilis' monikers, as individuals from the type locality of Pebble Beach did not align genetically with insular individuals but with other continental forms. As such, yet another name had to be coined for the insular form which now goes by the name of Silvetia compressa ssp. deliquescens. Let's see if it sticks this time.


Silva, P. C. 1996. California seaweeds collected by the Malaspina expedition, especially Pelvetia (Fucales, Phaeophyceae). Madroño 43 (3): 345–354.

Silva, P. C., F. F. Pedroche, M. E. Chacana, R. Aguilar-Rosa, L. E. Aguilar-Rosa & J. Raum. 2004. Geographic correlation of morphological and molecular variation in Silvetia compressa (Fucaceae, Fucales, Phaeophyceae). Phycologia 43 (2): 204–214.

The Millenium Post

Apparently, this is the 1000th post to appear at Catalogue of Organisms. When I first started this site, over ten years ago, I don't know if I had any idea when, if ever, I would reach this point and where I would be when it happened. I probably imagined I would be thinner.

I want to thank everyone that has followed Catalogue of Organisms over the years. I particularly want to thank those readers who have supported me on Patreon: Paul Selden, Sebastian Marquez, Rob Partington, William Holz. Your contributions have meant a lot to me. Apropos of that, some news: some of my readers may recall that my employment status has been a little up in the air for a large chunk of the last couple of years (though I was able to find casual positions for some of that time). A few months ago, though, an opportunity came up to work on a project looking at insect diversity in mangroves in Hong Kong. Though it means being away from my home, my partner and my dog for a couple of years, the opportunity was too good to pass up and for the next little while I'll be based in the city of the Fragrant Harbour (especially around the port district in high summer).

So on to the next 1000 posts, then? We'll just have to see. Certainly I'm not putting stuff up here as frequently as I did in the past, when I was a carefree post-graduate student. There have been times when I've wondered if I should keep going. People far more talented and perspicacious than I have had a great deal to say elsewhere about the apparent decline of science blogging as a format, and it certainly doesn't seem to attract the audience it once did. Nevertheless, I think I'll be going for a while yet. I've noted before that this blog functions as my own means and motivation for investigating things that I might find interesting, and there's certainly no shortage of things left to investigate. And as for the health of science blogging overall: a glance to the sidebar to the right of this page reminds me that there's still a lot out there worth following. There's Deep Sea News, there's Small Things Considered, Bug Eric, Tetrapod Zoology, Letters from Gondwana, Synapsida, Beetles in the Bush, and so many more. If you don't already know these sites, check them out!

For my part, the main indicator I see as to whether people are reading anything here is when people leave comments. A big thank you again to those who have contributed over the years. I'm needy, and need validation... And in that light, I'd like to specially ask my readers to comment on their general feelings (if any) about Catalogue of Organisms. Has there been anything you've particularly liked about the site over the years? Any favourite posts? Anything you'd like to see going forward? And once again...

The Shrinking World of Bandicoots

A bandicoot is a very disagreeable animal to clean, therefore it should be done as soon after killing as possible, and then the flesh can be left in strong vinegar and water for a few hours before dressing. Sweet potatoes and onion make a good stuffing for bandicoot, which is good either boiled or baked.--Mrs Lance Rawson, Australian Enquiry Book of Household and General Information.

Golden bandicoots Isoodon auratus barrowensis, copyright Kathie Atkinson.

Back when I used to work on Barrow Island in the north-west of Australia, one of the more noticeable animals to be seen around the place was the golden bandicoot Isoodon auratus. In the evenings, the place seemed to absolutely swarm with them. About the size of a guinea pig, with no tails to speak of (bandicoots are actually born with fairly long tails but tend to lose them in the course of their quite vicious fights with one another; few if any individuals reach maturity with their tails intact), there was no question about their qualifications when it came to cuteness.

Bandicoots are a group of twenty-odd species of marsupial found in Australia and New Guinea (one species, the Seram bandicoot Rhynchomeles prattorum, was described from montane forest on the Indonesian island of Seram to the west of New Guinea). Most are primarily insectivorous, but they also eat varying amounts of small vertebrates and plant matter such as bulbs and fruit. The largest bandicoot, the giant bandicoot Peroryctes broadbenti, has been recorded to reach close to five kilograms in weight. The smallest, the Papuan bandicoot Microperoryctes papuensis, weighs less than 200 grams. I suspect many people in Australia assume that the name 'bandicoot' comes from one of the the Aboriginal languages, but it is in fact Indian (specifically Telugu) in origin. The original bandicoot Bandicota indica is a large rat that is widespread in southern Asia and Australian bandicoots were named for their resemblance to this species. Personally, I have maintained in the past that Australian bandicoots look more like rats than rats do: with their relatively long snouts, bandicoots bear a distinct resemblance to the sort of cartoon figure that comes to most people's minds when they hear the word 'rat'.

New Guinea spiny bandicoot Echymipera kalubu, copyright Michael Pennay.

Bandicoots are highly distinctive from all other marsupials in appearance. Their hind legs are noticeably longer than their forelegs and more or less specialised for cursorial locomotion (especially so in one example that I'll get to shortly). The fourth and fifth toes of the hind foot are much larger than the other three; the first toe in particular is reduced to a non-functional stub. The second and third toes of the hind foot, as in diprotodontian marsupials such as kangaroos and possums, are externally joined together with the two claws at the end forming a comb that is used in grooming more than in locomotion. The fore feet, in contrast, are mostly functionally three-fingered (with the first and fifth fingers reduced) and adapted for digging with the claws large and flat.

Many bandicoots are rapid reproducers with their gestation periods among the shortest of any mammal, less than two weeks between fertilisation and birth. Bandicoots also have the most developed placentas of any marsupial group (yes, most marsupials do have a placenta, albeit a much simpler one than found in placental mammals); it is presumably because of this that, despite their short gestation, bandicoot young are born at a more advanced stage of development than those of some other marsupials. When the young are born, they initially remain attached to their mother via the umbilical cord; this latter does not become detached and the placenta ejected until after the joey is firmly attached to a teat in the rearward-opening pouch. The young remain in the pouch for about two months and grow rapidly; they may reach full sexual maturity at the age of only three months. As a result, bandicoot populations may increase rapidly if conditions permit.

Greater bilby Macrotis lagotis, copyright Bernard Dupont.

In terms of classification, there is a general consensus that Recent bandicoots can be divided between four groups though there has been some disagreement about exactly these groups are interrelated (and hence exactly how they should be ranked). The most diverse, but probably also the least studied, group of modern bandicoots are the rainforest bandicoots of the Peroryctidae or Peroryctinae. These are about a dozen species found mostly in New Guinea with the aforementioned Rhynchomeles prattorum on Seram and the the long-nosed spiny bandicoot Echymipera rufescens extending its range to the northern tip of Queensland. Most of continental Australia is home to the dry-country bandicoots of the Peramelidae sensu stricto or Peramelinae, of which there are six Recent species (one of these, the northern brown bandicoot Isoodon macrourus, is also found in southern New Guinea). Peramelids tend to have shorter snouts and flatter skulls than peroryctids. The other two groups are both very small and also native to arid regions of Australia. Two Recent species are known of the genus Macrotis, the bilbies, though one of these is extinct and the other is endangered. Bilbies are larger than most other bandicoots, with long ears (hence their alternative vernacular name of 'rabbit-bandicoots') and a long, silky-haired tail.

Gerard Krefft's 1857 illustration of the pig-footed bandicoot Chaeropus ecaudatus, from here.

The final representative of the Recent bandicoots is unquestionably the weirdest of them all. Unfortunately, it is also now extinct, last recorded some time about the middle of the 20th Century, a fact that cannot be called anything less than a fucking tragedy. The pig-footed bandicoot Chaeropus ecaudatus was the most cursorial of all bandicoots. Its forelegs, rather than being adapted for digging as in other bandicoots, had only two functional toes on which the claws were modified into hooves. The hind legs went a step further and had only a single functional toe (raising the question of how this animal groomed itself without the aforementioned claw-comb of other bandicoots Edit: That was a bit of a blonde moment; a second look at the Krefft illustration above shows that the comb is definitely there). The most extensive observations of its habits seem to have been made by Gerard Krefft (1866) who kept a pair alive for about six weeks in 1857 on a trip to the Murray-Darling region before killing them to provide specimens because, you know, 19th-Century naturalist. Krefft recorded that his bandicoots subsisted primarily on plant foods such as lettuce, grass and roots, refusing all meat offered to them (Krefft also refers to providing grasshoppers for them but his account is unclear about whether they were ever eaten). A herbivorous diet was also indicated by the animals' droppings, which where dry and similar to a sheep's. The bandicoots constructed a covered nest from grass and leaves in the tin enclosure in which Krefft kept them in which they sheltered during the day, only becoming active after nightfall. Krefft notes that he acquired "about eight" specimens of pig-footed bandicoot during his six-month camp, admitting that some met a stickier end than others: "They are very good eating, and I am sorry to confess that my appetite more than once over-ruled my love for science; but 24 hours upon "pig-face" (mesembryanthemum) will dampen the ardour of any naturalist". Krefft also noted that several of the specimens found were female, and that despite being provided with eight teats the females never carried more than two joeys. A particularly interesting detail was that the fourth toe of the joeys' fore foot, rather than being reduced as in the adults, remained large so that the feet resembled those of other bandicoots. Presumably this was so that the fore-claws could still be used to allow the newborn joeys to climb from the birth canal to the pouch.

Krefft also noted that the pig-footed bandicoot was already declining in abundance, blaming its increased rarity on competition with introduced grazing livestock. Sadly, changing habitats and introduced predators have caused other bandicoot species to also become endangered since Krefft's time. Please, don't let them go the way of the pig-footed bandicoot.


Gordon, G., & a. J. Hulbert. 1989. Peramelidae. In: Fauna of Australia vol. 1B. Mammalia. Australian Biological Resources Study: Canberra.

Krefft, G. 1866. On the vertebrated animals of the lower Murray and Darling, their habits, economy, and geographical distribution. Transactions of the Philosophical Society of New South Wales 1862–1865: 1–33.

A Second Look at Scallops

In a post that appeared at this site over eight years ago, I described some of the distinctive features of the Pectinoidea, the group of bivalves commonly known as scallops. It's time to look in a bit more detail at some of the points mentioned in that post.

Fossil of Pernopecten, the earliest scallop genus, from

Pectinoidea, in the sense recognised by Waller (2006), first appear in the fossil record way back in the late Devonian. They were probably derived from earlier members of the Aviculopectinoidea, an extinct group of bivalves that closely resemble scallops in their overall appearance and were included in the Pectinoidea by many earlier authors (such as in the 1969 Treatise on Invertebrate Paleontology volume on bivalves). However, the shell ligament of aviculopectinoids was reinforced by aragonite fibres (a primitive feature for bivalves) rather than having the specialised rubbery core found in pectinoids. As such, aviculopectinoids would have lacked the swimming abilities of true scallops. The Palaeozoic pectinoids belong to a single genus, Pernopecten, that possesses a number of features such as details of the shell crystalline structure that indicate a position outside the pectinoid crown group. In the early Triassic, Pernopecten begat the family Entolioididae that includes the ancestors of living pectinoids.

As mentioned in the previous post, four pectinoid families survive to the present day: the Pectinidae, Propeamussiidae, Entoliidae and Spondylidae. The first three families diverged in the early Triassic. Spondylids (usually classified in a single genus, Spondylus) were not to appear until the mid-Jurassic and Waller (2006) argued for their derivation from within the Pectinidae. The Pectinidae are otherwise distinguished from other pectinoids by a structure called the ctenolium. This is a row of teeth that develops on the shell in the gap between the disc and one of the auricles (the triangular 'wings' at the top of the shell). During the earlier part of the scallop's life, when it lives attached to the ocean bottom by a byssus (what in mussels we call the 'beard'), the ctenolium functions to hold the byssus threads in place and help stop the shell from twisting. In those pectinid species that lack a byssus in the latter part of their life, the ctenolium may end up getting overgrown by the expanding shell and disappearing, but all pectinids (ignoring the aforementioned Spondylus question) have a ctenolium for at least part of their life.

The propeamussiid Cyclopecten secundus, copyright Museum of New Zealand Te Papa Tongarewa.

The Pectinidae is the largest scallop family in the present day, followed by the Propeamussiidae. The Entoliidae were diverse during the Mesozoic but declined dramatically after the end of the Cretaceous (I'm not clear whether or not their decline was a direct part of the end-Cretaceous mass extinction). Indeed, entoliids are completely unknown from the fossil record between the Palaeocene and the late Pleistocene; like the tuatara, it might be that the post-Mesozoic survival of entoliids could have gone completely unrecognised were it not for the single surviving relictual genus.

In the earlier post, I implied that propeamussiids lack the eyes and guard tentacles of other pectinids; it turns out that this was a mistake on my part. Many propeamussiids found in the deep sea do indeed lack these features but they are present in shallow-water propeamussiids. It appears that these features are ancestrally common to all crown-group pectinoids but have been lost as an adaptation to life below the photic zone. The anatomy and lifestyle of many propeamussiids remains poorly known but those species that have been investigated have simplified gills compared to pectinids. The filaments of the gills are free rather than being connected by ciliary junctions. The lips of the mantle are also simplified, lacking the complex lobes found in pectinids. These features may be related to the carnivorous diet of many propeamussiids that feed on zooplankton rather than smaller phytoplankton and organic particles.


Waller, T. R. 2006. Phylogeny of families in the Pectinoidea (Mollusca: Bivalvia): importance of the fossil record. Zoological Journal of the Linnean Society 148 (3): 313–342.


Trichosternus vigorsi, copyright Udo Schmidt.

Many of the carabid ground beetles tend to attract a lot of attention from amateur entomologists due to their size and striking appearance, but it must be admitted that they are often not the easiest of animals to work with from a taxonomic perspective. The larger species tend to fall into the category of 'big, black, massive sharp mandibles' and it can require a lot of practice to reliably identify which genus a specimen belongs to, let alone species.

Trichosternus is a genus of ground beetles found in far eastern Australia, from the base of Cape York in Queensland to a bit north of Sydney in New South Wales, in the band of land between the coast and the Great Dividing Range. There is also a single isolated species T. relictus in the southwest corner of Western Australia, and apparently another in New Caledonia (Darlington 1961). However, considering the difficulty that many authors have had in the past in providing an exact definition for Trichosternus relative to other closely related genera, it would be interesting to see if future studies corroborate the inclusion of these outlying species. By way of contrast, a reasonable number of New Zealand species assigned at one time or another to Trichosternus have all long since been moved elsewhere.

Trichosternus species are all flightless and in most the elytra are fused and cannot open (the exception is T. relictus). Most species have a distinctive male genital morphology, with the genital opening deflected to the right and the right paramere (the parameres are two sclerotised 'arms' on either side of the genitalia) modified into a specialised falcate shape, the exact functional significance of which seems to remain unknown. Again, the outlier in this regard is T. relictus in which said paramere retains a primitive styloid shape. Similar falcate parameres are also known from members of related genera such as Megadromus and Nurus; the latter is particularly similar to Trichosternus with the only real difference between the two being that Nurus is more robust with longer mandibles. Trichosternus relictus also has a distinctive female genital morphology, in which the internal passage between the median oviduct (where emerging eggs are fertilised by sperm stored in the spermatheca) and the vagina is remarkably extended and concertina-like. Again, the function of this structure is unknown though Moore (1965) suggested that it might be related to viviparity.

Northern Trichosternus species found in tropical Queensland are all inhabitants of rainforest (hence the restriction of the genus to east of the Great Dividing Range: on the western side of the range, rainforests are absent and the arid zone begins). Southern species are found in upland temperate rainforests or in savannah woodland (Darlington 1961). Some species have very restricted ranges: T. montorum, for instance, is known from two mountains on the Spec Plateau, Mts Bartle Frere and Bellenden Ker.


Darlington, P. J., Jr. 1961. Australian carabid beetles VII. Trichosternus, especially the tropical species. Psyche 68 (4): 113–130.

Moore, B. P. 1965. Studies on Australian Carabidae (Coleoptera). 4.—The Pterostichinae. Transactions of the Royal Entomological Society of London 117 (1): 1–32.

The Lonely Life of the Cave Collembolan

For a few weeks last year, I had the job of sorting and identifying a collection of Collembola, springtails. Prior to doing this work, I had only the vaguest of understandings of springtail diversity: I knew that there were the round blobby ones, the long thin ones, and the ones that look a bit like sausages, but that was about as far as it went. Needless to say, there's a bit more to it than that.

Pseudosinella immaculata, copyright Andy Murray.

Pseudosinella is the largest genus of Collembola currently recognised, with over 280 described species. The greater number of those species are in Europe and North America, but various Pseudosinella have also been described from other regions of the world (there don't appear to be any from South America, but then I don't know how thoroughly anyone's looked). Pseudosinella species are mostly associated with subterranean habitats, from soil and litter to deep caves, with the highest diversity in the latter. According to a key at, Pseudosinella are distinguished from related genera by having reduced eyes (with six or fewer ommatidia, as opposed to the eight ommatidia of other genera), and a bidentate mucro lacking a projecting lamella (the mucro is the claw-like structure at the end of the furcula, the posteroventral prong that forms a springtail's 'spring'). The key also distinguishes Pseudosinella from the similar genus Rambutsinella by it's not having the fourth antennal segment swollen as in the latter, but Bernard et al. (2015) described the species Pseudosinella hahoteana as also having the fourth antennal segment swollen so I'm not sure how reliable that feature is. Pseudosinella is very similar to another genus Lepidocyrtus, the main difference between the two being Pseudosinella's reduced eyes, and more than one author has raised the possibility that Pseudosinella may be a polyphyletic assemblage derived from Lepidocyrtus adapted for life underground.

As well as the reduced eyes, Pseudosinella tend to show a number of other features commonly associated with a subterranean lifestyle, such as a pale coloration and relatively elongate appendages. The claws of the feet also tend to become modified, with the larger of the two becoming longer and progressively narrower (Christiansen 1988). This latter feature is probably an adaptation to movement on the wet surfaces that predominate in caves. At a moderate length, the claws dig into the substrate surface more than those of surface-dwelling forms, allowing greater grip. At longer lengths, the claws are suited to allow the springtail to walk over the surface of the water itself (most springtails float on water surfaces due to their small size and low density, but not all can move with purpose in this position).

Pseudosinella hahoteana, from Bernard et al. (2015). Scale bar = 200 µm.

The aforementioned Pseudosinella hahoteana is worthy of extra attention, as it is one of a half-dozen springtail species endemic to caves on Rapa Nui, the landmass previously known as Easter Island. Many of you will be aware of the ecological catastrophe that beset Rapa Nui following human settlement, as its entire forest covering was cleared away. As a result of this clearing, the native fauna was also all but wiped out; no vertebrates survive, and of about 400 arthropods known from the island only about twenty are indigenous (Bernard et al. 2015). As such, the handful of minute animals clinging to survival in patches of ferns and moss at the entrance to caves represent a significant proportion of Rapa Nui's surviving native fauna.


Bernard, E. C., F. N. Soto-Adames & J. J. Wynne. 2015. Collembola of Rapa Nui (Easter Island) with descriptions of five endemic cave-restricted species. Zootaxa 3949 (2): 239–267.

Christiansen, K. 1988. Pseudosinella revisited (Collembola, Entomobryinae). Int. J. Speleol. 17: 1–29.

Define 'Trichostomum'

The moss in the above photo Icopyright Hermann Schachner) generally goes by the name of Trichostomum crispulum. Trichostomum is a cosmopolitan genus in the Pottiaceae, the largest recognised family of mosses with about 1500 species overall. But with great diversity comes great difficulty of identification. Pottiaceae tend to be small mosses that are common in harsh habitats. Features of pottiaceous mosses are often hard to distinguish and may be quite variable, making it difficult to confidently define taxa. As a result, Pottiaceae is a prime example of what I like to call 'taxonomic blancmange': something that tends to just get prodded nervously then backed away from when it wobbles ominously.

Characteristic features of Trichostomum as it is commonly recognised tend to include symmetric leaves with more or less plane margins, and with the basal cells of the leaf differentiated straight across the blade or in a U-shape. The peristome of the capsule also tends to be short and straight, and the sexual system is usually dioicous (with separate male and female plants) (Flora of North America). However, none of these features are entirely reliable, and some species have been the subject of extensive disagreement about whether they should be placed in Trichostomum, or in a related genus such as Weissia or Tortella.

To date, only a selection of Pottiaceae species have been subject to molecular analysis, but these analyses have confirmed the unsatisfactory nature of the current system. A molecular phylogenetic analysis of the pottiaceous subfamily Trichostomoideae by Werner et al. (2005) did not identify Trichostomum species as a monophyletic clade; instead, various representatives of the 'genus' were scattered throughout the subfamily. The type species of Trichostomum, T. brachydontium, was associated with a few close relatives such as T. crispulum in a broader clade containing numerous species of the genus Weissia. As a result, it has been suggested that the two genera should perhaps be synonymised, in which case the name Trichostomum would be absorbed by the older Weissia. But first, someone would need to work out just how such a genus could be recognised...


Werner, O., R. M. Ros & M. Grundmann. 2005. Molecular phylogeny of Trichostomoideae (Pottiaceae, Bryophyta) based on nrITS sequence data. Taxon 54 (2): 361–368.

Cryptophytes: Four Genomes for the Price of One

Sometimes, the little things really do make a difference. Cryptophytes (or cryptomonads) are one of the many groups of minute flagellate protists to be found around the world whose role in our lives tends to get dismissed because of their microscopic size. Nevertheless, cryptophytes make up a large part of the photosynthetic phytoplankton in both freshwater and marine habitats and so ultimately are a starting point for many of the food chains that we depend on. They also had an important role to play in our developing understanding of how modern eukaryote cells have evolved.
Structure of a typical cryptophyte, from here.

As well as occurring in the phytoplankton, cryptophytes have also been found in damp soil and snow. They have a distinctive, slightly lop-sided cell morphology with two haired flagella of unequal length inserted in an invaginated gullet towards the right side of the front of the cell. This invagination is also lined on the ventral side by organelles called ejectosomes (sometimes spelled 'ejectisome'). When the organism is threatened, these ejectosomes shoot out a proteinaceous ribbon that propels the cell rapidly away from the source of irritation. Some of the references to ejectosome function that I've found seem to imply that the expelled ribbon is itself toxic, but I'm not sure if I've understood correctly. Smaller ejectosomes may also play a role in capturing bacteria and the like for the cryptophyte to feed on. Cryptophytes have a distinctive way of moving through the water column, resulting from the uneven lengths of their two propellent flagella, that has been reffered to as 'recoiling'. Essentially, they move in a series of circular tumbles while the cell itself corkscrews around its axis. This movement is distinctive enough that cryptophytes have been dubbed with the Dutch vernacular name of 'rekylalger', 'recoiling algae' (Novarino 2003).

Diagram of typical cryptophyte movement, from Novarino (2003).

The majority of cryptophytes are heterotrophic: one or more large chloroplasts provide much of the cell's energy, but they are also capable of ingesting particulate matter through the gullet. As alluded above, the cryptophyte chloroplast has been significant in the study of how chloroplasts evolved. The 1960s and 1970s saw an increasing acceptance of the concept that some organelles, most notably mitochondria and chloroplasts, had originally appeared through a process of endosymbiosis: bacteria had become intimately associated with eukaryote cells, becoming embedded in the host cell and eventually ceding enough of their vital functions to the host to be unable to function as independent organisms. The chloroplasts of the ancestors of land plants arose in this manner from cyanobacteria, as indicated by the presence of a remnant but reduced bacterial genome within the chloroplast itself, and the presence of a double membrane around each chloroplast (corresponding the cyanobacterium's original cell membrane, plus the vacuoule membrane in which it had been enclosed by the host eukaryote). In the early 1970s, however, it was found that cryptophyte chloroplasts have not two but four surrounding membranes. What is more, wedged between two of those membranes was a tiny remnant cell nucleus, dubbed the nucleomorph. The nucleomorph was a crucial piece of evidence in demonstrating that cryptophyte chloroplasts had arisen by a process of secondary endosymbiosis. A eukaryote cell containing a chloroplast that had arisen in the manner described above was itself engulfed and converted to a chloroplast by another eukaryote. The four membranes around the cryptophyte membrane were therefore, from the inside out, the original cyanobacterium cell membrane, the vacuole membrane containing the cyanobacterium, the cell membrane of the primary host cell (with the nucleomorph between this and the last), and the vacuole membrane in which that had been contained in turn. Other groups of eukaryotes also have chloroplasts that arose in this way, such as brown algae and dinoflagellates, but in these the nucleus of the captured eukaryote cell has entirely disappeared.

Another cryptophyte structural diagram of the species Guillardia theta, showing the arrangement of the chloroplast, from here. This also shows the sites of the four genomes contained in the typical cryptophyte cell.

Exactly when the cryptophyte chloroplast arose remains a contentious subject. Various lines of evidence point to the captured chloroplast donor being a red alga, as is also the case with the aforementioned brown algae and dinoflagellates. As such, some have argued for the chloroplasts of all such algae being descended from a single capture event. However, there are also a number of protists related to such taxa that lack chloroplasts. In the case of cryptophytes, there is strong evidence that the sister clade to the the photosynthetic cryptophytes is the chloroplast-less genus Goniomonas. The subsequent sister to these two clades together is less certain but a number of recent studies have pulled forward another chloroplast-less group, the katablepharids. If the cryptophyte chloroplast shares an origin with that of brown algae, then it must have somehow been lost in the ancestors of both Goniomonas and katablepharids. So far, an author's preference for a single or multiple origins of red alga-derived chloroplasts tends to come down to whether they think it is easier for chloroplasts to be lost or gained, a question whose answer is still unclear.

The diversity within cryptophytes is still not that well understood, largely due to difficulties in observing significant characters. Prior to the advent of scanning electron microscopy, some authors had gone so far as to dismiss cryptophytes as essentially unclassifiable. Nevertheless, not everything was as bleak as the pessimists would have it. Cryptophyte taxa may differ from each other in overall size and shape. They may also differ in cell colour, due to the presence of various accessory pigments in addition to chlorophyll. The primary accessory pigments found in cryptophytes are known as phycocyanin and phycoerythrin; species containing the former are a blue-green colour whereas those containing the latter are reddish, golden or a greenish yellow. The use of scanning electron microscopy has led to the discovery of other useful features such as those relating to the periplast, a protein envelope that covers the inside and outside of the cryptophyte cell membrane. Electron microscopy has shown that the outer periplast layer is often ornamented, such as by being divided into scales. And even more recently, of course, researchers have recognised the value that molecular tools may have to offer cryptophyte taxonomy, though said tools have also complicated matters by, for instance, giving hints that previously recognised 'taxa' may represent different life cycle stages of a single organism. Whatever the eventual result, there is no question that we still have a lot to learn about cryptophytes.


Novarino, G. 2003. A companion to the identification of cryptomonad flagellates (Cryptophyceae = Cryptomonadea). Hydrobiologia 502: 225–270.


It's time to meet the sweepers.

Smallscale bullseyes Pempheris compressa, copyright John Turnbull.

Sweepers, Pempheridae, are a group of moderately sized marine fish (usually about fifteen to twenty centimetres in length) found around tropical reefs in the Indo-Pacific and western Atlantic. I don't know why they're called sweepers, but in some areas they may be among the most abundant fish on the reef. Distinctive features of the group include a short, high dorsal fin and a long anal fin. The lateral line is also distinctively long, extending past the end of the tail right onto the caudal fin. Perhaps the feature that most stands out about sweepers is their large eyes. The eyes are so big because sweepers are nocturnal; during the day they retreat into protected crevices and caves, emerging at night to feed on minute crustaceans and other small animals (Mooi 2001).

Pygmy sweeper Parapriacanthus ransonneti, from here.

Sweepers are divided between two quite distinct genera. Members of the genus Parapriacanthus have a more 'average fish-like' elongate profile with the body less deep than the head is long. The other genus, Pempheris, has a distinctively deep profile, deeper than the head is long. The exact number of species of pempherid appears to still be uncertain. Pempherids lack the striking markings of other tropical fish and species can appear very similar to each other. What is more, they have two layers of scales on the body, with the outer scales being larger than the inner and deciduous (easily shed), and loss of the outer scales has the potential to change an individual's superficial appearance. Early descriptions of pempherid species are often inadequate for their reliable identification, and new species continue to be described at a quite rapid pace. A recent publication by Randall & Victor (2015), for instance, described no less than thirty-four new species of Pempheris from various locations in the Indian Ocean, close to doubling the number of species in the genus at a stroke. The genus Parapriacanthus is much less diverse, with only about five recognised species.

Orange-striped bullseyes Pempheris ornata in hiding during the day, copyright Peter Southwood.

Because of their relatively small size and retiring habits, sweepers are mostly not that significant economically. At least one species, Pempheris xanthoptera, is fished off the coast of Japan and mostly eaten as fish paste; it is supposed to be quite tasty. Some have appeared in aquaria.

When foraging at night, sweepers communicate with each other by producing popping noises through muscular flexing of the swim bladder wall. Noise production increases in the presence of potential threats, perhaps to warn other members of the school. At least some pempherid species also have bioluminescent glands associated with the posterior part of the gut. The bioluminescent compound is not directly produced by the fish itself but obtained by consuming bioluminescent ostracods. I haven't found whether the function of this bioluminescence is specifically known for pempherids, but similar ventral glows in other fish provide camouflage by breaking up the fish's silhouette when seen from below.


Mooi, R. D. 2001. Pempheridae. Sweepers (bullseyes). FAO Species Identification Guide for Fishery Purposes. The Living Resources of the Western Central Pacific vol. 5. Bony fishes part 3 (Menidae to Pomacentridae) pp. 3201–3204. Food and Agriculture Organization of the United Nations: Rome.

Randall, J. E., & B. C. Victor. 2015. Descriptions of thirty-four new species of the fish genus Pempheris (Perciformes: Pempheridae), with a key to the species of the western Indian Ocean. Journal of the Ocean Science Foundation 18: 77 pp.

Long-legged Harvestmen of Southern Africa

New paper time!

Rhampsinitus conjunctidens, a new species of harvestmen from north-east South Africa, from Taylor (2017).

Taylor, C. K. 2017. Notes on Phalangiidae (Arachnida: Opiliones) of southern Africa with description of new species and comments on within-species variation. Zootaxa 4272 (2): 236–250.

When I first started research for my PhD thesis, *cough* years ago, I asked a number of museums if they could loan me their collections of monoscutid (now neopilionid) harvestmen. The species that I was interested in are found in Australia and New Zealand but when I opened a package of specimens sent to me from the California Academy of Sciences, I found a number of specimens from Africa in the mix. I immediately recognised what they were: not neopilionids, but representatives of another harvestment family, the Phalangiidae.

It seemed an easy enough error to make. Many species of southern African Phalangiidae resemble a lot of neopilionids in that the males have over-sized, elongate chelicerae. I referred to some of these species in the genus Rhampsinitus in an earlier post. To those not familiar with harvestmen diversity (which, let's face it, is the majority of people out there), the two groups can look very similar. True, the phalangiids are all distinctly much spikier than the neopilionids, but that doesn't seem that major a difference. To really see where they diverge from each other, you need to reach underneath the males' genital opercula and pull out their todgers.

Anywho, the specimens sat in storage for much longer than they should have, until I finally got around to looking them over in the latter part of last year. I then decided that it was worth writing them up into a short paper. Not only was there at least one new species among the specimens, they told me some very interesting things about variation within the species. Not only do Rhampsinitus species resemble Australasian neopilionids in their enlarged chelicerae, they resemble them in that individuals of a species vary in how enlarged the chelicerae are.

Major (left) and minor males of Rhampsinitus nubicolus, from Taylor (2013).

Now, I was not the first person to observe this point. Axel Schönhofer (2008) had already provided some detailed examples of variation in males of Rhampsinitus cf. leighi. I did, however, observe that the variation was even greater than Axel seemed to have recognised. Some of the least developed males of the species I was looking at had chelicerae that were pretty much no more developed than those of females. In some ways, the variation was even more remarkable than what I was familiar with in neopilionids. In most of the latter, major and minor males tend to be pretty similar to each other in features other than cheliceral development. In Rhampsinitus, we can see variation in almost all the features related to sexual dimorphism. In the species pictured immediately above, R. nubicolus, major males have massively long pedipalps as well as the long chelicerae; minor males have short, stubby pedipalps like those of a female. We can tell that they are the same species because they are found in the same location and have matching genitalia, but on the outside you would be hard pressed to pick them as such. Just to confuse matters even more, major males of two species may look very different to each other whereas minor males are externally almost identical. Without looking at the genitalia, it is all but impossible to identify which species a minor male belongs to.

As with the neopilionids, we can't yet say for sure what this variation means for the species' behaviour. In many other animal species with comparably varying males, large males will fight to protect and contain females while small males adopt a sneaking behaviour and try to spot females that are not being watched by large males. It seems quite possible that a similar thing is going on with Rhampsinitus. If you're a keen natural historian or behavioralist, there's something here that is crying to be looked into.


Schönhofer, A. L. 2008. On harvestmen from the Soutpansberg, South Africa, with description of a new species of Monomontia (Arachnida: Opiliones). African Invertebrates 49 (2): 109–126.

A Mystery Ammonoid

Münster's (1834) figure of Goniatites hybridus.

Looks like I drew another dud. For today;s semi-random post, I ended up tasking myself to write something about the Devonian ammonoid genus Heminautilinus. But as it turns out, there simply isn't that much to say about this genus, and what there is isn't really worth saying.

Heminautilinus was established as a genus by A. Hyatt in 1884. He diagnosed it as including "species with whorls similar to those of Anarcestes, but with angular lateral lobes in the adults", and designated George de Münster's (1834) Goniatites hybridus as type species on the basis of that author's original figure. The problem is that Münster's figure is apparently not very reliable; the original specimen was only fragmentary and Münster himself expressed uncertainty as to just what section of the ammonoid conch he had on hand. So Hyatt's assumption that Münster's species retained some juvenile features to maturity should not be considered reliable.

As a result, Hyatt's genus seems to have been pretty roundly ignored. Those authors who have made some speculation as to its identity have suggested that it is probably synonymous with some better known genus such as Cheiloceras or Imitoceras. This might present something of an issue because either one of these genera was published more recently than 1884, meaning that Heminautilinus should be considered the senior name. Because there would be little to be gained from replacing a familiar name with one that is all but forgotten, it seems most likely that, even if Heminautilinus' identity could be reliably established, it would be somehow suppressed. As such, Heminautilinus seems doomed to remain in obscurity.


Hyatt, A. 1883–1884. Genera of fossil cephalopods. Boston Soc. Nat. History, Proc. 22: 253–338.

Münster, G. de. 1834. Mémoire sur les clymènes et les goniatites du calcaire de transition du Fichtelgebirge Annales des Sciences Naturelles, seconde série, Zoologie 1: 65–99, pls 1–6.

Blue Moon

Male blue moon butterfly Hypolimnas bolina, photographed by Comacontrol.

My native country of New Zealand is not home to a large diversity of butterflies. Only a couple of dozen or so species are known from the entire country. It would not be unreasonable for a keen butterfly spotter to attempt to track down them all. But one particular species of butterfly generally included in New Zealand lists would a touch of luck: the aptly named blue moon Hypolimnas bolina.

This is because the blue moon is not a regular resident of New Zealand (I've personally never spotted one). It is native to a wide region stretching from Madagascar and India to Japan and northern Australasia where it is usually referred to by the more prosaic name of common or greater eggfly. The only examples found in New Zealand are vagrants who lost their way on southwards migrations. Nevertheless, such vagrants are regular enough for its local appellation to be thought worth coining. Not only does it reflect their rarity, it also describes the appearance of the male, with the wings bearing blue-ringed white spots on a black background.

Two females of Hypolimnas bolina. On the left, a mimetic individual, copyright Greg Hume; on the right, a non-mimetic individual, copyright W. A. Djatmiko.

The appearance of the female is a bit harder to explain because it can vary between individuals. Females of the eggfly genus Hypolimnas are commonly mimics of other, poisonous butterflies of the subfamily Danainae, to which eggflies are only distantly related (both groups belong to the family Nymphalidae but eggflies are placed in the subfamily Nymphalinae). For instance, the diadem or danaid eggfly H. misippus of Africa and Asia (and also introduced into parts of the Americas adjoining the Caribbean) is a mimic of the plain tiger Danaus chrysippus. The chosen model of H. bolina in the western part of its range is the common crow Euploea core and in the region of India almost all females are a remarkably good copy of that species (above left). But as one moves east, one starts seeing females of H. bolina that are not mimics like the individual shown above right; by the time one reaches Australia these make up the greater part of the population. Mimetic females may also vary to resemble different Euploea species, depending on which model is locally present.

Female danaid eggfly Hypolimnas misippus, copyright Raju Kasambe. Males of this species are similar to those of H. bolina but lack the blue rings around the white spots on the wings.

There are about two dozen species of Hypolimnas eggflies found in various parts of the Old World tropics. Hypolimnas misippus is also found in parts of the Americas around the Caribbean where its presence is usually explained as the result of an early introduction (possibly, and somewhat poignantly, in connection to the slave trade). Their vernacular name is probably derived from the unique behaviour (for butterflies) of a number of species whose females stand guard over their eggs, beating their wings over them to protect them from predators until hatching. About two-thirds of Hypolimnas species are mimics. In some of these species, both sexes are mimetic; others resemble H. bolina and H. misippus in that only the females are mimics (Vane-Wright et al. 1977). One might be tempted to ask why this variation exists. One point to be considered is that there are limits on when mimicry is likely to be effective. The mimic needs to be much less abundant than its model, otherwise potential predators may not learn to associate the distinctive coloration with the toxic original. Swinhoe (1896) noted that males of Hypolimnas misippus were very active, aggressively defending their territories from other butterflies, and suggested that this agility might provide males with alternative defences to mimicry. The more sedentary females (especially when egg-guarding) might be expected to benefit more from the passive protection mimicry provides, but mimesis might be expected to disappear in areas where their model is less abundant.


Swinhoe, C. 1896. On mimicry in butterflies of the genus Hypolimnas. Journal of the Linnean Society, Zoology 25: 339–348.

Vane-Wright, R. I., P. R. Ackery & R. L. Smiles. 1977. The polymorphism, mimicry, and host plant relationships of Hypolimnas butterflies. Zoological Journal of the Linnean Society 9: 285–297.

False Spider Mites

Among the enormous diversity of the world's mites, some families are particularly notorious for the damage that they inflict on commercial plant crops. Among such Acari non grata are the spider mites of the family Tetranychidae or the gall mites of the Eriophyidae. But a third, equally notorious group is the false spider mites or flat mites of the Tenuipalpidae.

Red and black flat mite Brevipalpus phoenicis, false-colour SEM by Christopher Pooley.

False spider mites include about 800 known species of more or less flattened, slow-moving mites. They are closely related to the true spider mites and both families have the chelicerae modified into a pair of long, whip-like retractable stylets that are used to pierce and suck fluids from plant tissues. In the case of the false spider mites, though, their commercial infamy comes not only from the direct damage caused by the feeding mites themselves but also from the effects of transmitted viruses. Viruses transmitted by false spider mites include the causative agents of diseases such as citrus leprosis and coffee ring spot and may cause significant reductions in the yield and lifespan of infected plants.

Morphologically, false spider mites differ from true spider mites in the absence of what is called the 'thumb-claw' process, an arrangement of the tarsus of the pedipalp alongside a claw on the seta (hence the family name which means 'slender palp'). The palps are often reduced, with some species having only the barest remnant. Some species also show reduction in the fourth pair of legs, and females of a number of species are six-legged as adults. This merely stands as another example of how mite morphology functions purely to play silly buggers with anything one might learn in basic animal biology.

Hebe stem gall mites Dolichotetranychus ancistrus inside an open gall, copyright Plant and Food Research.

Parthenogenesis is also common in false spider mites. Species found in cooler climes will often overwinter as females, with a new generation of males not appearing until the next spring. In some species, eggs produced parthenogenetically will hatch into males; in others, they will produce females. A few species almost entirely lack functional males. A small group of these species in the genus Brevipalpus is unique among animals in being both parthenogenetic and genetically haploid.

Almost all forms of seed plant seem to be vulnerable to some form of flat mite or another; some mite species are very catholic in their tastes and will latch onto almost anything green and photosynthesising. Others are more discerning. How false spider mites make their way from one host plant to another is little known but they may be passively carried through the air on the wind. Alternatively, they may be inadvertently carried from place to place by feeding herbivores, or by the very human horticulturalists that suffer so much from their presence.


Walter, D. E., E. E. Lindquist, I. M. Smith, D. R. Cook & G. W. Krantz. 2009. Order Trombidiformes. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 233-420. Texas Tech University Press.

Oily and Salty Trees

The Annonaceae is another one of those plant families like Acanthaceae that, despite containing a high diversity of speceis, tend to be overlooked because that diversity is mostly tropical. A number of species in the type genus Annona produce commercially significant fruits: custard apples, cherimoyas, soursops and the like. However, these are just a few of the 2400+ species of trees and lianes assigned to this family.

Ylang-ylang flowers Cananga odorata, from here.

Taxonomically, the Annonaceae is well established as distinct, readily recognised by a number of distinctive features. Among these is a characteristic 'cobweb' appearance to the wood structure when seen in cross-section, resulting from prominent rays of xylem connected by narrow cross-bands of parenchyma (Chatrou et al. 2012). Relationships within the family have been much harder to work out, not becoming well established until the advent of the molecular era. Recently, Chatrou et al. (2012) have recognised four subfamilies within the Annonaceae. The majority of species are placed in the subfamilies Annonoideae and Malmeoideae (which together form a clade), but a handful of species are placed in two basal subfamilies: one for the single genus Anaxagorea, and the Ambavioideae. Anaxagorea and the ambavioids differ from the annonoid-malmeoid clade in the structure of their seeds. Seeds of Annonaceae have what is called ruminate endosperm: that is, the surface of the endosperm is not smooth, but divided by wrinkles and grooves (the term 'ruminate' literally means 'chewed'). In Annonoideae and Malmeoideae, the ruminations of the endosperm are shaped like spines or lamellae. In Anaxagorea and the Ambavioideae, the ruminations are irregular in appearance. Molecular analyses place Anaxagorea as the sister taxon to all other Annonaceae.

View into the canopy of a salt-and-oil tree Cleistopholis patens, copyright Marco Schmidt.>

The Ambavioideae, despite not being very diverse, are widespread, with species found in the tropics of Africa, Asia and the Americas. Perhaps the best known ambavioid is the ylang-ylang tree Cananga odorata, native to south-east Asia, whose flowers are used as a source of perfume. Other south-east Asian ambavioids belong to the genera Cyathocalyx, Drepananthus and Mezzettia. The type genus, Ambavia, is native to Madagascar; other ambavioids in the genera Meiocarpidium, Cleistopholis and Lettowianthus are found in continental Africa. Finally, a single genus Tetrameranthus is found in South America. Most species of ambavioid are not systematically economically exploited but a number are locally used as sources of wood. The wood is light and not suitable for structural uses, but can be shaped and finished for utensils and other small items. The West African species Cleistopholis patens, whose Ghanaian name has been translated as 'salt and oil tree' (in reference to the taste of the bark when chewed), provides a fibrous bark that is readily stripped from the tree and is used for such purposes as matting and carrying straps (see here).


Chatrou, L W., M. D. Pirie, R. H. J. Erkens, T. L. P. Couvreur, K. M. Neubig, J. R. Abbott, J. B. Mols, J. W. Maas, R. M. K. Saunders & M. W. Chase. 2012. A new subfamilial and tribal classification of the pantropical flowering plant family Annonaceae informed by molecular phylogenetics. Botanical Journal of the Linnean Society 169: 5–40.

Hydromantes: Salamanders in Different Places

There are times when biogeography is able to throw us some real puzzlers: organisms whose distribution seems to defy expectations. Among these mysteries, special mention must be made of the salamanders of the genus Hydromantes.

Gene's cave salamanders Hydromantes genei courting, copyright Salvatore Spano.

Hydromantes is a genus containing a dozen species from among the lungless salamanders of the family Plethodontidae. Plethodontids are the most diverse of the generally recognised families of salamanders, with over 450 known species found mostly in Central and South America. Hydromantes, however, is a geographically isolated genus in this family with its species found in two widely separated regions: California in western North America, and mainland Italy and Sardinia in Europe. Though some authors have advocated treating the species found on each continent as separate genera, both morphological and molecular studies have left little doubt that the group represents a discrete clade.

Distinctive features of Hydromantes compared to other plethodontids include feet with five, partially webbed toes and a weakly ossified, flattened skull (Wake 2013). Members of this genus capture prey with a projectile tongue which is the most extensive of any amphibian, extending up to 80% of the animal's total body length (Deban & Dicke 2004). There are some differences between North American and European species notable enough for the recognition of separate subgenera (there is something of a gigantic clusterfuck surrounding the names of said subgenera but the details are far too tedious to relate here). The three North American species of the subgenus Hydromantes have bluntly tipped tails that they use as a 'fifth leg' when navigating smooth and/or slippery surfaces, whereas the European species have unremarkable pointed tails. Historically, the North American Hydromantes species have been poorly known, being isolated to restricted ranges. Hydromantes shastae is found in limestone around Lake Shasta whereas H. brunus is found in a small area of mossy talus habitat along the Merced River in the foothills of the Sierra Nevada (Rovito 2010). The third species, H. platycephalus, is found at higher altitudes in the Sierra Nevada, well over 1000 m above sea level. Individuals found living on steep slopes are known to escape predators by tightly coiling their bodies and simply rolling down the slope (García-París & Deban 1995). A molecular analysis of H. platycephalus and H. brunus by Rovito (2010) identified the former species as derived from within the latter, and Rovito suggested that H. brunus may have originated in a remnant population from when H. platycephalus moved into lower altitudes during the Ice Age.

Mt Lyell salamander Hydromantes platycephalus, copyright Gary Nafis.

The seven or eight European species are mostly placed in the subgenus Speleomantes; a single species, Hydromantes genei, is divergent enough to be placed in its own subgenus Atylodes (though most recent studies have indicated that the European Hydromantes overall form a discrete clade). Hydromantes genei and three species of Speleomantes are found in caves on the island of Sardinia; the remaining Speleomantes species on mountains of mainland Italy. Molecular analysis suggests that H. genei became isolated on Sardinia about nine million years ago, with the ancestors of the Sardinian Speleomantes arriving later about 5.6 million years ago when the Mediterranean dried out during what is known as the Messinian Salinity Crisis (Carranza et al. 2008). The absence of any Hydromantes on neighbouring Corsica is something of a mystery, and it has been suggested that they may have been present there in the past before going extinct.

Extinction also seems the most likely explanation for Hydromantes' unusual distribution. The fossil record for the genus is minimal, and provides little information not already available from living species, but molecular dating attempts agree that the division between European and North American Hydromantes happened too recently to be related to the tectonic separation of the two continents. Such a scenario would also leave open the Hydromantes' absence in eastern North America. The description in 2005 of the Korean lungless salamander Karsenia koreana demonstrated the presence of plethodontids in eastern as well as far western Eurasia, and it seems possible that Hydromantes dispersed into Eurasia via the Bering Strait landbridge, becoming widespread across the continent before extinction reduced it to the isolated relicts it is today.


Carranza, S., A. Romano, E. N. Arnold & G. Sotgiu. 2008. Biogeography and evolution of European cave salamanders, Hydromantes (Urodela: Plethodontidae), inferred from mtDNA sequences. Journal of Biogeography 35: 724–738.

Deban, S. M., & U. Dicke. 2004. Activation patterns of the tongue-projector muscle during feeding in the imperial cave salamander Hydromantes imperialis. Journal of Experimental Biology 207: 2071–2081.

García-París, M., & S. M. Deban. 1995. A novel antipredator mechanism in salamanders: rolling escape in Hydromantes platycephalus. Journal of Herpetology 29 (1): 149–151.

Rovito, S. M. 2010. Lineage divergence and speciation in the web-toed salamanders (Plethodontidae: Hydromantes) of the Sierra Nevada, California. Molecular Ecology 19: 4554–4571.

Wake, D. B. 2013. The enigmatic history of the European, Asian and American plethodontid salamanders. Amphibia-Reptilia 34: 323–336.

Leucicorus: FAKE EYES!

In an earlier post, I told you about the fishes known as brotulas. These are one of the most prominent groups of fish in the deep sea. They tend not to be attractive fish: their lack of outstanding dorsal and tail fins makes them look like something between an eel and a cod, and like many deep-sea fishes they look somewhat flabby and lumpish. There are numerous genera of brotulas out there; the individual in the photo below represents the genus Leucicorus.

Leucicorus atlanticus, from Okeanos Explorer.

Leucicorus belongs to the brotula family Ophidiidae, commonly known as the egg-laying brotulas though Leucicorus' own reproduction has (so far as I have found) not been directly observed. The feature that most immediately sets Leucicorus apart from other brotulas is the eyes: Leucicorus species have very large eyes but the actual lenses are rudimentary or absent (Cohen & Nielsen 1978). It almost looks like they grew bigger and bigger to cope with the low light of the deep sea before they just kind of gave up at some point.

Two species of Leucicorus are currently recognised, each known from separate parts of the world. Leucicorus lusciosus is found in the eastern Pacific, whereas L. atlanticus is known from around the Caribbean. The two species differ in meristic characters and proportions: for instance, L. lusciosus has more dorsal and anal fin rays, but fewer vertebrae and gill rakers, and has a deeper body (Nielsen & Møller 2007). Leucicorus has also been found in the vicinity of the Solomon Islands, but interestingly enough Nielsen & Møller (2007) identified the specimen found as L. atlanticus rather than L. lusciosus, despite the latter species' more proximate distribution. One wonders if perhaps a third species is involved, yet to be recognised.


Cohen, D. M., & J. G. Nielsen. 1978. Guide to identification of genera of the fish order Ophidiiformes with a tentative classification of the order. NOAA Technical Report NMFS Circular 417.

Nielsen, J. G., & P. R. Møller. 2007. New and rare deep-sea ophidiiform fishes from the Solomon Sea caught by the Danish Galathea 3 Expedition. Steenstrupia 30 (1): 21–46.