Field of Science

Anthaxia: More Modest Jewels

The jewel beetles of the Buprestidae are best known for their spectacularly patterned exemplars, a couple of which I've presented on this site before. But as with most animal groups renowned in this way, they also include their fair share of less immediately eye-catching members. The species of the genus Anthaxia are among these more modest jewels.

Anthaxia hungarica, photographed by Frayle.

Which is not to say they are unattractive. Anthaxia species still usually have the metallic gloss so widespread among the Buprestidae but they tend to be more uniform in colour, and those colours are often shades of bronze or blue-green rather than yellows or purples. They are also smaller than the species previously shown: a length of 6.5 millimetres would be relatively large for an Anthaxia. Some of the smallest species don't quite make it to three millimetres (Bílý & Kubáň 2010). Nevertheless, Anthaxia are incredibly diverse. Something in the range of 700 species are known from around the world (though they appear to be absent from Australia, with the single species described from Victoria now thought to have been based on a mis-labelled African specimen) and a quick Google Scholar search indicates new species continue to be described regularly. It should come as no surprise that many of these species would be difficult to distinguish without close examination.

Anthaxia scutellaris, a more colourful species of the genus, copyright Hectonichus.

Like other buprestids, Anthaxia species are wood-borers as larvae and flower-feeders as adults. The larvae seem to run the gamut of preferred tree hosts: Anthaxia have been found emerging from hosts ranging from pines to pears, from oleander to oaks. Some species appear to be quite catholic in their tastes: the recorded host list for the most polyphagous known species, A. millefolii, includes maples, chestnuts, carobs, oleanders, pistachios, plums, pears, oaks and rowans (Mifsud & Bílý 2002). Others are more discerning. Species of the subgenus Melanthaxia are only known to feed on conifers (Bílý & Kubáň 2010) and records for A. lucens indicate a dedication to stonefruit trees (Mifsud & Bílý 2002). Nevertheless, the larval hosts of many species remain unknown and there may be surprises. The North American species A. hatchi might be expected to be a conifer feeder like other Melanthaxia species but to date it has been collected in riparian habitats where conifers do not grow (Nelson et al. 1981). Could this member of an otherwise conifer-loving group have developed a taste for the willows and alders amongst which it lives? The question is yet to be answered.


Bílý, S., & V. Kubáň. 2010. A study on the Nearctic species of the genus Anthaxia (Coleoptera: Buprestidae: Buprestinae: Anthaxiini). Subgenus Melanthaxia. Part I. Acta Entomologica Musei Nationalis Pragae 50 (2): 535–546.

Mifsud, D., & S. Bílý. 2002. Jewel beetles (Coleoptera, Buprestidae) from the Maltese Islands (central Mediterranean). Central Mediterranean Naturalist 3 (4): 181–188.

Ferreting up a Bird's Nose

Mites, as I may have commented before, seem to have an almost fractal level of diversity: the closer you look, the more there is of it. This is nowhere more apparent than when it comes to parasitic mites which infest almost any host in any way that you can imagine. For the subject of this post, I drew one such mite: the honeyeater nasal mite Ptilonyssus myzanthae.

Venter (left) and dorsum of female Ptilonyssus myzanthae, from Domrow (1964). The scale bar equals 500 µm.

Bird nasal mites of the family Rhinonyssidae are, as their name indicates, inhabitants of the nasal passages of birds. General adaptations of the family for their parasitic lifestyle include tendencies towards reduction of the body sclerotisation and reduction in the length and number of setae. They use the claws on their front legs to tear openings in the host's mucous membranes and then feed on its blood. Transmission of nasal mites seems to happen during bill-to-bill contact such as when parents are feeding their young or during mating activities, or indirectly through water or on the surface of perches or the like. Rhinonyssid nasal mites are not known to transmit any actual diseases between hosts but they can cause the formation of lesions or inflammation or the like. All in all, probably not very pleasant for the bird (see here for some more details).

Whole-body illustration of a different rhinonyssid species, from Greg Spicer.

Nevertheless, infection rates in bird populations can be very high and most (if not all) bird species will be host to some nasal mite species. Most species of nasal mite are very host specific, known on only one or a few bird species (it must be noted, though, that the question of just how many researchers choose to look up a bird's schnozz in search of mites may not be irrelevant here). Ptilonyssus myzanthae was described by Domrow (1964) from two species of honeyeater in Queensland, Australia: the noisy miner Manorina melanocephala and the little wattlebird Anthochaera chrysoptera. Distinctive features of this species compared to others in the genus include a subhexagonal anterior dorsal shield on the body, a narrow genital shield, and a divided pygidial shield (the small pair of shields near the rear of the dorsum). Both of the known hosts are widespread and common in eastern Australia and it is likely that this mite is similarly ubiquitous. Studies of honeyeater phylogeny tend to place the genera Manorina and Anthochaera as close relatives, so it is possible that P. myzanthae has been infesting them since before their lineages diverged. It would be worth looking for the species in other related honeyeaters to see if we find any further clues.


Domrow, R. 1964. Fourteen species of Ptilonyssus from Australian birds (Acarina, Laelapidae). Acarologia 6 (4): 595–623.


The common perception of monkeys tends to be dominated by a relatively small number of species, generally those most commonly seen in zoos, such as capuchins, macaques, baboons or tamarins. But as is usual when it comes to biodiversity, there are a lot of varieties of monkey out there that may be less familiar to the general public. This post will look at one of those less familiar groups: the guenons of the genus Cercopithecus.

Moustached monkey Cercopithecus cephus, copyright Rufus46.

Cercopithecus is a genus of monkeys found in sub-Saharan Africa. The exact number of species has shifted around a bit (though it currently sits around twenty). Some authors have included almost all species of the monkey tribe Cercopithecini, characterised by self-sharpening lower incisors and four cusps on the lower third molars (Lo Bianco et al. 2017), in the single genus Cercopithecus. However, more recent authors have tended to favour dividing this tribe between a number of phylogenetically and ecologically distinct genera. Under this latter system, Cercopithecus would be restricted to a group of more arboreal species. A number of these species have been divided between multiple subspecies and there may be some back and forthing about what is recognised as which. One entirely new species, previously not even known as a subspecies, was described as recently as 2012 by Hart et al.: the lesula C. lomamiensis.

Young female lesula Cercopithecus lomamiensis, from Hart et al. (2012).

A large part of this uncertainty relates to the fact that Cercopithecus species are most diverse in dense forests of western and central Africa, in regions that may be both physically and politically difficult to access and which have received less attention from researchers than others. The aforementioned lesula was described from the Lomami River basin near the middle of the Democratic Republic of the Congo (the one that used to be called Zaire, though I think they prefer not to talk about it). Another Congolese species, the dryas monkey C. dryas, was long thought to be known from only a single juvenile specimen until it was realised that the adult form had been described as a separate species C. salongo. It's still only known from a handful of records and is thought to be critically endangered.

Diana monkey Cercopithecus diana, copyright Ikmo-ned.

Some species of guenon are notable for their striking colour patterns. Perhaps the species I've most commonly seen in zoos is the diana monkey C. diana, native to the region between Sierra Leone* and the Côte d'Ivoire (though it is possible that at least some of these 'diana monkeys' were actually roloway monkeys C. roloway, until recently treated as a subspecies of the diana monkey). This species has a bright white throat, chest and front of the fore arms that contrasts with the black face and dark grey back. It also has a white band across its brow which is where its name comes from, the band having been thought to resemble the crescent moon. De Brazza's monkey C. neglectus of central Africa has a crescent-shaped orange mark on its forehead and a white muzzle and beard, making it look reminiscent of a grumpy old man (Wikipedia claims that it has also been dubbed the 'Ayatollah monkey'). Male De Brazza's monkeys also have a bright blue scrotum. Large bright blue patches are also present around the scrotum and backside of males in the lesula and the owl-faced monkey C. hamlyni.

*Having grown up in New Zealand in the 1980s, I'm going to have that stuck in my head all day now. Nothing to do with the subject of this post, I just thought I'd mention it.

Male De Brazza's monkey Cercopithecus neglectus, copyright Heather Paul.

Guenons tend to be found living in small troops consisting of one adult male and a harem of females with their offspring; unmated adult males will be found living solitary lives. Males are usually larger than females, up to about 1.5 times the size of their mates. Multiple guenon species may be found in a single location though closely related species tend not to overlap. Famously, hybrids have been described from the Kibale forest in Uganda between the blue monkey C. mitis and the red-tailed monkey C. ascanius, two species that are quite distinct in external appearance. Larger species such as the spot-nosed monkey C. nictitans and the blue monkey tend to eat a higher proportion of leaves in their diet. Smaller species such as the mona monkey C. mona may be more insectivorous (Macdonald 1984).

Blue monkeys Cercopithecus mitis stuhlmanni, copyright Charles J. Sharp.

The origins of the Cercopithecus radiation are relatively recent with the tribe Cercopithecini as a whole probably originating in the late Miocene (Lo Bianco et al. 2017). Karyological studies of the group show a wide variation in chromosome number from 58 in the diana monkey to 72 in the blue monkey. In contrast, the sister group of the Cercopithecini, the Papionini (which includes baboons and macaques) always has 42 chromosomes. Polymorphism in chromosome arrangements has also been described within Cercopithecus species. The possibility that this gene variability is related to their rate of speciation remains a worthwhile line of study.


Hart, J. A., K. M. Detwiler, C. C. Gilbert, A. S. Burrell, J. L. Fuller, M. Emetshu, T. B. Hart, A. Vosper, E. J. Sargis & A. J. Tosi. 2012. Lesula: a new species of Cercopithecus monkey endemic to the Democratic Republic of Congo and implications for conservation of Congo's central basin. PLoS One 7 (9): e44271.

Lo Bianco, S., J. C. Masters & L. Sineo. 2017. The evolution of the Cercopithecini: a (post)modern synthesis. Evolutionary Anthropology 26: 336–349.

Macdonald, D. (ed.) 1984. All the World's Animals: Primates. Torstar Books: New York.

Matoniaceae: Ferns with a Heritage

Ferns are one of those groups of organisms, like sharks and cockroaches, that are not really as ancient as most people imagine. For all that ferns are indelibly associated in the public conscience with antediluvian imagery of steamy coal swamps and great lumbering reptiles, the dominant fern groups that can be seen today did not arise until the Cretaceous and diversified as part of a flora that would have been largely modern in appearance (Schneider et al. 2004). Nevertheless, there are some fern lineages around today that might be said to have a genuine claim to a more venerable pedigree. One such group is the Matoniaceae.

Matonia pectinata, copyright Ahmad Fuad Morad.

In the modern flora, the Matoniaceae are a small family, including only three or four species in two genera, Matonia and Phanerosorus, found in south-east Asia (Lindsay et al. 2003). The two genera are distinct in appearance and habits. Matonia is found on more or less exposed montane summits and ridges and has pedate fronds with pectinate pinnae radiating from an erect central stipe that may grow well over a metre in height. Phanerosorus is found on vertical limestone walls and has pendulous, branching fronds whose pinnae are simple or more weakly pectinate (Kato & Setoguchi 1999). Both genera have the fronds arising from a long, hairy, creeping rhizome. Lateral veins in the pinnules show one or more bifurcations and in Matonia these branching forks may anastomose with each other to form a reticulate vein pattern. The genera also share features of the reproductive anatomy such as massive, deciduous sporangia.

Phanerosorus major, copyright Wally Suarez.

The fossil record of Matoniaceae indicates that they were far more widespread in the past; indeed, Matonia was illustrated from preserved compression fossils before it was described as a living genus (Klavins et al. 2004). Leaf fossils of Matoniaceae go back to the Late Triassic, and the Middle Triassic stem taxon Soloropteris rupex has been more tentatively assigned to the family (van Konijnenburg-van Cittert 1993). Fossil forms are more similar to Matonia in overall appearance and this is presumed to be the plesiomorphic morphology for the family. A certain resemblance exists between Phanerosorus and younger fronds of Matonia and it seems likely that the former genus evolved from Matonia-like forms by a process of paedomorphosis (Kato & Setoguchi 1998). The family was most widespread during the Jurassic and Early Cretaceous but became extinct in temperate regions of the Northern Hemisphere during the Late Cretaceous. It persisted longer in the Southern Hemisphere, with the stem taxon Heweria kempii known from the Early Tertiary of Australia, but at some point following that it became restricted to its modern localised range.


Kato, M., & H. Setoguchi. 1999. An rbcL-based phylogeny and heteroblastic leaf morphology of Matoniaceae. Systematic Botany 23 (4): 391–400.

Klavins, S. D., T. N. Taylor & E. L. Taylor. 2004. Matoniaceous ferns (Gleicheniales) from the Middle Triassic of Antactica. Journal of Paleontology 78 (1): 211-217.

Konijnenburg-van Cittert, J. H. A. van. 1993. A review of the Matoniaceae based on in situ spores. Review of Palaeobotany and Palynology 78: 235–267.

Lindsay, S., S. Suddee, D. J. Middleton & R. Pooma. 2003. Matoniaceae (Pteridophyta)—a new family record for Thailand. Thai Forestry Bulletin 31: 47–52.

Schneider, H., E. Schuettpelz, K. M. Pryer, R. Cranfill, S. Magallón & R. Lupia. 2004. Ferns diversified in the shadow of angiosperms. Nature 428: 553–557.

Radiolarians of the Globe

Radiolarians are one of the primary groups of micro-organisms to be found among the marine plankton. These unicellular greeblies are justly famed for their intricate mineralised skeletons, leading to their comparison to living works of art. Today's post is covering one particular group of radiolarians, the Spumellaria.

Haeckel's (1899–1904) figure of Hexancistra quadricuspis from Kunstformen der Natur.

Spumellaria are one of the major subdivisions of radiolarians, containing species characterised by a generally spherical skeletal form. Many authors have also included the colonial radiolarians, which often lack a coherent skeleton and may form colonies up to several metres long, in the Spumellaria but these have more recently been treated as a distinct group. The skeleton of radiolarians is entirely enclosed by cytoplasm in life, though in those species in which the skeleton bears radiating spines, those spines may extend beyond the main body of the cell and be covered by only a thin cytoplasmic layer distally. In Spumellaria and anothre major radiolarian group, the Nassellaria, the skeleton is composed of opal, making these living jewels in more ways than one (another radiolarian group, the Acantharea, composes its skeleton of a mineral by the somewhat ethereal-sounding name of celestite). The cytoplasm of radiolarians is internally divided by a fibrous capsule into two structurally distinct sections, the internal endoplasm and external ectoplasm. The denser endoplasm contains most of the cell's primary organelles, such as the nucleus and large mitochondria. Linear microtubular structures called axonemes extend outwards from the endoplasm, passing through pores in the internal capsule and through the ectoplasm. The ectoplasm is often frothy in texture, containing an extensive assemblage of cellular vacuoles. In many of these radiolarians, some of these ectoplasmic vacuoles will house symbiotic algae that contribute much of the radiolarian's nutrition. Otherwise, radiolarians may feed on other small organisms that are captured on axopodia supported by the axonemes, which in spumellarians radiate outwards from the cell body in all directions. Extension and contraction of the axopodia may also help maintain the radiolarian's position in the water column (Cachon et al. 1990).

Schematic diagram of organisation of Didymocyrtis tetrathalamus from Sugiyama & Anderson (1998).

In many spumellarians, the basic skeletal architecture is one of nested spheres and/or globules. Sugiyama & Anderson's (1998) description of Didymocyrtis tetrathalamus stands as a fairly typical example. The central part of the skeleton is a double sphere well within the cytoplasmic capsule with the lobate nucleus contained in the spaces between the spheres. Radiating axes connect the inner shell with an outer shell mostly just outside the capsule (the capsular wall crosses the skeleton at some points). In Didymocyrtis, this outer shell is not spherical but a sort of peanut shape. At each end of the 'peanut', a further cap is added beyond the main shell. In many spumellarians, the outer shell appears spongy in texture, being constructed of densely criss-crossing fine opal fibres. There may be further extensions of the outer shell such as polar spines or funnels.

Not surprisingly, spumellarian classification has most commonly been based on skeletal architecture. Some attempts have been made to construct alternative classifications incorporating cytoplasmic features such as the relationship between the axopods and the nucleus (Cachon et al. 1990) but, as these systems require access to live specimens to place taxa, they have been less popular (especially as most people studying radiolarians are primarily working with fossil material). A phylogenetic study of recent spumellarians by Ishitani et al. (2012) found evidence for two main lineages within the class that differ in ecology. One, including the families Pyloniidae and Sponguridae, contained species found in temperate and cold waters. The other, including the families Astrosphaeridae, Hexalonchidae and Coccodiscidae, was found in tropical waters. Species assigned to the family Spongodiscidae were divided between both lineages, suggesting the need for some further tinkering with the morphological classification.


Cachon, J., M. Cachon & K. W. Estep. 1990. Phylum Actinopoda. Classes Polycystina (=Radiolaria) and Phaeodaria. In: Margulis, L., J. O. Corliss, M. Melkonian & D. J. Chapman (eds) Handbook of Protoctista. The structure, cultivation, habitats and life histories of the eukaryotic microorganisms and their descendants exclusive of animals, plants and fungi. A guide to the algae, ciliates, foraminifera, sporozoa, water molds, slime molds and the other protoctists pp. 334–346. Jones & Bartlett Publishers: Boston.

Ishitani, Y., Y. Ujiié, C. de Vargas, F. Not & K. Takahashi. 2012. Two distinct lineages in the radiolarian order Spumellaria having different ecological preferences. Deep-Sea Research II 61–64: 172–178.

Sugiyama, K., & O. R. Anderson. 1998. Cytoplasmic organization and symbiotic associations of Didymocyrtis tetrathalamus (Haeckel) (Spumellaria, Radiolaria). Micropaleontology 44 (3): 277–289.


The photo above (copyright Dave), may or may not show Dalmanella, a brachiopod originally described from the later Ordovician of Sweden. Dalmanella belongs to the Orthida, one of the earliest groups of articulate brachiopods to appear in the fossil record ('articulate' meaning that the two valves of the shell are hinged together, not that they are particularly well spoken). The Dalmanellidae, the family to which Dalmanella belongs, are known from the lower Ordovician to the lower Carboniferous (Williams & Wright 1965).

Over the years, numerous fossil brachiopods from Europe and North America have been assigned to Dalmanella, leading Jin & Bergström (2010) to describe it as "perhaps one of the most commonly reported orthide brachiopods". However, if truth be told, the main reason Dalmanella is so widely recognised is because of how perfectly unremarkable it is. It is small and unspecialised, and the genera within Dalmanellidae have mostly been separated by somewhat vague characters such as shell shape and ribbing pattern. Some studies of variation in dalmanellid populations have questioned whether characters used to separate genera can even be used to separate species or whether they may vary within a single population.

This uncertainty lead Jin & Bergström (2010) to restudy the original type species of Dalmanella, D. testudinaria. Their conclusion was that D. testudinaria was morphologically distinct from North American species attributed to the genus: for instance, the midline of the dorsal valve bore an interspace (the furrow between two costae) in D. testudinaria but a raised costa in the American species. The myophore, a process associated with the hinge to which the muscles responsible for opening the shell would have attached in life, is much narrower in D. testudinaria than in the American species. Not only were the morphologically distinct, they were ecologically distinct as well: D. testudinaria being found in cooler, deeper waters while the American species basked in tropical shallows. Not for the first time, it appears that an external sameyness masks an internal divergence.


Jin, J., & J. Bergström. 2010. True Dalmanella and taxonomic implications for some Late Ordovician dalmanellid brachiopods from North America. GFF 132 (1): 13–24.

Williams, A., & A. D. Wright. 1965. Orthida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H299–H359. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

Ghost Moths and Other Obscurities

During my early years in rural New Zealand, I would often take note of the variety of insect life that could be seen coming to the screen doors at night, attracted by the light from inside the house. Among the most spectacular animals that would sometimes turn up was a gigantic pale green moth, about three inches long as it crawled across the screen. This was the puriri moth Aenetus virescens, perhaps New Zealand's best known member of the moth clade Exoporia.

Puriri moth Aenetus virescens, copyright Nga Manu Images NZ.

The Exoporia is one of the more basal moth groups alive today. The name of the clade refers to one of its most distinctive features: a female genital system with separate external openings for the seminal receptacle and the oviduct, meaning that the male's sperm has to travel along an external groove between the two if it is to fertilise the egg (in other Lepidoptera, there is a single cloacal opening, or there are separate openings but the cavities are connected by an internal duct). Other important features of the clade include dicondylic antennae, with two instead of just one articulations between the antenna and the head, and a male reproductive system without a sclerotised tubular intromittent organ (Kristensen 1978; the males instead have the gonopore opening on a shorter protuberance). Six families are generally recognised within the clade but the majority of species (including the puriri moth) belong to just one of these families, the Hepialidae, commonly known as the ghost moths.

Bentwing ghost moth Zelotypia stacyi, copyright CSIRO.

Hepialids definitely buck the phylogenetic trend among moths. Lepidopterists commonly divide the moth and butterfly order between two main groupings, somewhat self-explanatorily referred to as Micro- and Macrolepidoptera. To some extent, this is merely a division of convenience (the practicalities of working with smaller and larger moths can be quite different) but Macrolepidoptera is also used as the name of a major clade within the order with micro-Lepidoptera then indicated for any lepidopteran not belonging to this clade. By this measure, hepialids are by far the largest micro-Lepidoptera out there (most other examples are unquestionably micro). I've already alluded to the fifteen centimetre wingspan of the puriri moth but this isn't even close to being the largest hepialid out there. The honour perhaps goes to the bentwing ghost moth Zelotypia stacyi of eastern Australia which reaches a wingspan of 25 centimetres, a full ten inches. The larvae of hepialids are commonly borers in live trees; the puriri moth, for instance, gets its name because it burrows into puriri trees Vitex lucens. Other species live as larvae in burrows in soil, emerging at night to feed on pasture or leaf litter, or feeding externally on tree roots. Adult hepialids are short-lived and do not feed, and as such their proboscis is reduced or absent. They may emerge en masse at particular times of year. Following mating, females may scatter their eggs at random during flight or lay them in loose masses on the ground, with larvae finding a suitable food source after hatching. Because of the high mortality rates associated with this scatter-shot method, laying rates can be exceedingly large: females of some genera may produce around 18,000 eggs apiece (Nielsen & Common 1991).

Mnesarchaea acuta, copyright George Gibbs.

The other exoporian families are all much less diverse and more localised. They are also all small moths, far more typical 'micro-Lepidoptera'. The genus Mnesarchaea, endemic to New Zealand, retains functional mouthparts and is believed to be the sister group to all other exoporians. Larvae of Mnesarchaea live in silken galleries among mosses and liverworts, feeding on moss and liverwort leaves, algae, fungal spores and the like. The remaining families all lack functioning mouthparts as adults but their habits are otherwise all but unknown. Anomoses hylecoetes is placed in its own family known from rainforests in eastern Australia. The family Neotheoridae was until recently known from only a single female specimen collected in Brazil, but a few further species of this family were described recently by Simonsen & Kristensen (2017). Prototheora, another genus held worthy of its own family, is found in southern Africa. Finally, the family Palaeosetidae is known from a small number of genera with disjunct distributions in Colombia, south-east Asia and Australia. Because of its scattered distribution, some authors have questioned whether this last family is monophyletic, but an analysis of exoporian phylogeny by Simonsen & Kristensen (2017) continued to support it as such. It is not impossible that this family is more widespread, its apparent rarity due to the overlooking of small moths emerging for only very short periods, living just long enough to breed and deposit their eggs in as-yet-unknown locales.


Kristensen, N. P. 1978. A new familia of Hepialoidea from South America, with remarks on the phylogeny of the subordo Exoporia (Lepidoptera). Entomologica Germanica 4 (3–4): 272–294.

Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

Simonsen, T. J., & N. P. Kristensen. 2017. Revision of the endemic Brazilian 'neotheorid' hepialids, with morphological evidence for the phylogenetic relationships of the basal lineages of Hepialidae (Lepidoptera: Hepialoidea). Arthropod Systematics and Phylogeny 75 (2): 281–301.