Field of Science


Metabiantes leighi, from Schönhofer (2008).

The handsome fellow in the photo above represents a species of the genus Metabiantes, currently the largest recognised African genus of harvestmen in the family Biantidae. Metabiantes species are currently recognised from the greater part of sub-Saharan Africa, with species known from as far north as Kenya on the east coast and the Ivory Coast in the west (Staręga 1992). The bulk of study on this genus, however, has focused on the South African species. Kauri (1961) divided the southern African species between two groups based on genital morphology. Schönhofer (2008) recorded that the widespread Transvaal species M. leighi inhabits leaf litter in evergreen forests. It seems to inhabit slightly drier microhabitats than other harvestmen in the area.

Compared to many other harvestmen, Metabiantes species do not show a high degree of sexual dimorphism. The males' chelicerae tend to be a bit larger, and consequently the front of the carapace tends to be a bit broader (Schönhofer 2008). In some species, the metatarsus of the males' second leg bears a series of teeth (Lawrence 1937). While I haven't found any explicit investigation of the role that such modifications play, the fact that the second legs in harvestmen fill a sensory function (being used much like the antennae in insects) provides a likely suggestion.


Kauri, H. 1961. Opiliones. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 9–197. Almqvist & Wiksell: Uppsala.

Lawrence, R. F. 1937. The external sexual characters of South African harvest-spiders. Transactions of the Royal Society of South Africa 24 (4): 331–337.

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.

Staręga, W. 1992. An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33 (2): 271–336.

I Said Primrose-Willows, Darling

The plant shown in the photo above (copyright Forest and Kim Starr) is Ludwigia octovalvis, commonly known (along with other species in the same genus) as primrose-willow. This is a very common plant in tropical and subtropical regions around the world; indeed, it is so widespread that we have little idea where in the world it originated*. The name 'primrose-willow' derives, of course, from its combination of primrose-like flowers with willow-like leaves, but it is no close relation to either. Primrose-willows belong to the Onagraceae, the same plant family as evening primrose or fuchsias. Ludwigia octovalvis is a shrubby plant, sometimes growing up to four metres in height. Lower parts of the stem may become woody with age, but the greater part of the plant is herbaceous. It prefers to grow in damp habitats, in swampy soil or alongside streams, even rooted in ponds. The seeds are minute and easily spread by water or mixed in with other materials. They are also durable: Raven (1977) refers to the possibility of propagating Ludwigia from seeds preserved in herbarium specimens.

*Which, if one were of a panbiogeographical bent, might be taken to indicate that it has survived unchanged since the Triassic at least.

Ludwigia octovalvis may even grow as floating mats upon the surface of water. The floating roots then produce spongy, upright branches called aerophores. These have been interpreted as floatation devices, but the plant is apparently perfectly buoyant without them. It is more likely that they allow oxygen to reach the waterlogged roots. The lower part of the stem may also become covered in aerenchyma, porous tissue that also aids in the diffusion of gases. If conditions dry up and the plant becomes rooted in the ground, the aerophores disappear and the roots resume their normal rootly business.

Close-up of flower of Ludwigia octovalvis, copyright Bob Peterson.

Primrose-willows are generally toxic to humans. In the usual way, I have come across references to Ludwigia octovalvis being used folk-medicinally, mostly to help with ailments of the digestive tract such as diarrhoea and worms. A quick look through Google Scholar indicates that this has lead to a certain degree of pharmaceutical research, but so far this doesn't seem to have lead to much major commercial application. At the present point in time, the main economic impact of Ludwigia octovalvis is as a weed. It can grow mixed in with fields of crops such as rice and taro, or particularly lush patches of primrose-willow may clog up waterways. On the flipside, I did find this summary of the species that notes that, "Its yellow flowers add a splash of color to areas often devoid of colorfully flowering plants".


Raven, P. H. 1977. Onagraceae. Flora Malesiana, ser. I, 8 (2): 98–113.

The Diprotodontids: Marsupials Go Large

Reconstruction of Diprotodon optatum by Anne Musser, from Long et al. (2002). Offhand, running a search for Diprotodon through Google Image brings up some true horrors of digital imagery.

Prior to the arrival of humans, the Australian fauna included many strange, and often dramatic, animals that are sadly no longer with us. Enormous python-like snakes, monitors that would have made a Komodo dragon look underwhelming, drop bears, and of course the notorious demon duck of doom. But among the most iconic of Australia's extinct fauna were the Diprotodontidae, heavyset herbivores that included the largest of all marsupials. Diprotodontids are sometimes referred to in the popular press as giant wombats, but this is a bit misleading: though more closely related to wombats than any other living marsupials, they were a quite distinct group of animals (besides, they shared their world with actual giant wombats that reached the size of a cow). A potentially more appropriate descriptor that has been suggested is 'marsupial rhinos', though at least some diprotodontids were decidedly not like rhinos either.

Skull of Zygomaturus trilobus in Museum Victoria, photographed by Nigel Waring.

The most famous of the diprotodontids was also the first to be described, and indeed the first fossil mammal of any kind described from Australia. Diprotodon optatum, named by Richard Owen in 1838, was the largest of the diprotodontids, sometimes standing more than six feet tall at the shoulder, and reaching estimated weights of around two and a half tonnes. At the time of human arrival, Diprotodon would have been one of the dominant herbivores in the arid central region of Australia. A number of species of Diprotodon have been named over the years, but a review of the genus by Price (2008) recognised only a single species, with the two different size classes present probably representing the different sexes. In the less arid coastal regions, Diprotodon was replaced by various species of the slightly smaller (but still formidably sized) genus Zygomaturus (Long et al. 2002). The best known species in this genus, Z. trilobus, bore a distinctive large bony boss on the snout, giving its skull a profile reminiscent of a cartoon bear. Two other diprotodontid species that would have come into contact with humans are known from the Pleistocene of montane New Guinea, Hulitherium tomasettii and Maokopia ronaldi. Both these species were smaller than the mainland Australians, being about the 100 kg mark. Maokopia has been interpreted as a grazer, while Hulitherium has been seen as a browser, and suggested as a direct analogue of the Asian giant panda (Long et al. 2002).

Reconstruction of Hulitherium tomassettii as a panda analogue, by Peter Schouten.

The broader record of diprotodontids goes back to the Oligocene, with two main lineages being recognised, the Diprotodontinae and Zygomaturinae. Of the species referred to above, all but Diprotodon optatum are zygomaturines. The two groups are primarily distinguished by their dentition, with the premolars being generally more complex in zygomaturines than diprotodontines. In both lineages, the earlier members were smaller: Long et al. (2002) describe a number of genera as 'sheep-sized'. The smallest known diprotodontid, the late Oligocene Raemeotherium yatkolai, they describe as 'lamb-sized'. Black et al. (2012) estimated the weight of the middle Miocene Nimbadon lavarackorum as abut 70 kg. They also suggested that it was an adept climber, in a similar manner to the modern koala, making it the largest known arboreal mammal from Australia. It might seem odd to picture an animal of this size up in a tree, even allowing for the higher density of the canopy in Australia's Miocene rainforests. However, there are larger arboreal mammals alive even today: male orangutans, for instance, may weigh over 100 kg.

Reconstruction of a climbing pair of Nimbadon lavarackorum (adult and juvenile) by Peter Schouten, from Black et al. (2012).

Interestingly, Nimbadon is not placed as a particular basal diprotodontid in the phylogeny of zygomaturines presented by Mackness (2010). As other related marsupial families, such as koalas or thylacoleonids (marsupial lions), also include climbers, it would not be unreasonable to consider such habits plesiomorphic for diprotodontids as a whole. The 'rhino-like' appearance of the later giants would then be something of a novelty, an adaptation to the drier conditions and more open woodlands that arose at the end of the Miocene. If we are to regard the diprotodontids as marsupial rhinos, then we must consider the possibility of rhinos in trees.


Black, K. H., A. B. Camens, M. Archer & S. J. Hand. 2012. Herds overhead: Nimbadon lavarackorum (Diprotodontidae), heavyweight marsupial herbivores in the Miocene forests of Australia. PLoS ONE 7 (11): e48213. doi:10.1371/journal.pone.0048213.

Long, J., M. Archer, T. Flannery & S. Hand. 2002. Prehistoric Mammals of Australia and New Guinea: One hundred million years of evolution. University of New South Wales Press: Sydney.

Mackness, B. S. 2010. On the identity of Euowenia robusta De Vis, 1891 with a description of a new zygomaturine genus. Alcheringa 34 (4): 455–469.

Price, G. J. 2008. Taxonomy and palaeobiology of the largest-ever marsupial, Diprotodon Owen, 1838 (Diprotodontidae, Marsupialia). Zoological Journal of the Linnean Society 153: 389–417.

The Cancellothyridids: A Modern Success Story

Northern lamp shels Terebratulina septentrionalis, from Oceana.

As has been noted on this site more than once before, brachiopods are a group of animals probably more familiar to the student of palaeontology than of zoology. From the brief gloss that tends to be their only coverage in textbooks, one might be forgiven for thinking them all but inconsequential in the modern fauna. But where conditions suit them (usually sheltered locations where low levels of light and water flow favour their slow metabolisms over the higher energy requirements of bivalves), brachiopods can still be abundant, and even dominant.

One of the most diverse families of brachiopods in the modern fauna is the Cancellothyrididae. Cancellothyridids first make their appearance in the Jurassic, becoming widespread in the Cretaceous (Cooper 1973). Members of this family have shells with a large foramen (the opening at the rear of the shell through which passes the pedicel or stalk by which the brachiopod is attached to its substrate), usually with the deltidial plates surrounding the foramen greatly reduced. The main defining feature of the Cancellothyrididae is the structure of the brachidium, the skeletal structure that provides the support for the base of the lophophore, the tentacle-like feeding structures. In cancellothyridids, the two sides of the brachidium coalesce in the middle to form a tube.

Dorsal valve of the Cretaceous cancellothyridid Cricosia filosa in (A) lateral, (B) ventral and (C) posterior views, from Cooper (1973), showing the tubular brachidium.

The brachidium does not extend into the arms of the lophophore, which are instead strengthened by unattached spicules. The tubular shape of the brachidium distinguishes the Cancellothyrididae from the closely related family Chlidonophoridae, whose members share the large posterior foramen but have the two sides of the brachidium open in back. Cooper (1973) recognised two subfamilies of cancellothyridids, the living Cancellothyridinae and the Cretaceous Cricosiinae; the cricosiines have the tubular section of the brachidium longer and narrower than the cancellothyridines.

Modern cancellothyridids are found in the Indo-Pacific and the North Atlantic, but seem to be absent from the South Atlantic. The majority of living species are included in the widespread genus Terebratulina, with the other living genera all having restricted distributions in the Indo-Pacific. However, a molecular phylogenetic analysis of species of Terebratulina and the Australian genus Cancellothyris by Lüter & Cohen (2002) indicated that both Atlantic Terebratulina and Cancellothyris were nested within Pacific Terebratulina. Paraphyly of the widespread genus would also correlate with its palaeontological distribution: while the other genera are known only from the Recent fauna, Terebratulina has a fossil record dating right back to the origins of the cancellothyridids in the Jurassic (Muir-Wood 1965). Lüter & Cohen (2002) tentatively suggested the possibility of a North Pacific origin for Terebratulina (and, by implication, for Cancellothyrididae as a whole), with dispersal to the North Atlantic occurring through the gap between North and South America before formation of Central America. Their preference for this option rather than the alternative of dispersal through the Tethys (the seaway that once separated Africa from Eurasia) was based on their estimate via molecular clock of a separation of about 100 million years between the Atlantic and Pacific species, supposedly too early for the Tethys option. However, it must be stressed that their sampling of even modern cancellothyridid diversity was not comprehensive. A trans-Tethys dispersal of cancellothyridids may also be indicated by the presence of the fossil genus Rhynchonellopsis in the lower Oligocene of northern Europe (Muir-Wood 1965). Of course, there is no inherent reason why cancellothyridids could not have travelled in both directions!


Cooper, G. A. 1973. Fossil and recent Cancellothyridacea (Brachiopoda). Tohoku Univ., Sci. Rep., 2nd Ser. (Geol.), Special Volume 6: 371–390.

Lüter, C., & B. L. Cohen. 2002. DNA sequence evidence for speciation, paraphyly and a Mesozoic dispersal of cancellothyridid articulate brachiopods. Marine Biology 141: 65–74.

Muir-Wood, H. M. 1965. Mesozoic and Cenozoic Terebratulidina. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H762–H816.