Field of Science

Sympiesis

Female Sympiesis, copyright Lyle J. Buss.


We often imagine that parasites select their hosts largely on the basis of type: one parasite prefers caterpillars, for instance, while another prefers flies. However, sometimes what is important is not so much what type of host a parasite attacks, as where they find it. The wasp in the photo above represents Sympiesis, a sizeable genus (the Universal Chalcidoidea Database lists over 130 species) of microscopic parasitoid wasps found worldwide. The majority of Sympiesis larvae attack the larvae of Lepidoptera, but others feed on the larvae of Diptera. A few have been recorded as hyperparasitoids, attacking the larvae of other parasitic wasps. The main thing that all hosts of Sympiesis have in common, though, is that they are all found in secluded, vegetative habitats: either mining in leaves, or in retreats formed by rolling or tying leaves (sometimes boring in stems). Depending on species, Sympiesis larvae may be either ectoparasites or endoparasites: those species feeding on leaf-rolling hosts tend to be ectoparasites, while those targeting leaf-miners are endoparasites (Miller 1970).

Sympiesis is a genus of the chalcid family Eulophidae. Eulophids used to be the subject of some disagreement between myself and a colleague of mine about their ease of recognition. Eulophids are a diverse group in appearance, coming in a bewildering array of shapes and colours. However, I have always maintained that they are nevertheless readily recognisable. Whatever their appearance, eulophids seem to always a distinctive stamp of 'eulophid-ness'. They tend to be slender, relatively soft-bodied wasps, often with a flat top to the gaster. Most identification guides will tell you to look out for their four-segmented tarsi (as opposed to the five-segmented tarsi of most other chalcid wasps); eulophid tarsi are rendered even more recognisable by the point that, though they have less segments than the tarsi of other chalcids, they are not any shorter so the individual tarsal segments are all relatively long. The features distinguishing Sympiesis from other eulophid genera are, of course, finer and require fairly close examination: notably, they have relatively few dorsal setae (only four on the scutellum) (Bouček 1988). As far as I know, they are mostly metallic green in coloration.

Male Sympiesis, showing branched antennae, from here.


As with many eulophids, males of Sympiesis usually differ from females in having long branches on the antennae. However, the first species of the genus to be described, the European Sympiesis sericeicornis, is distinctive in that these antennal branches are much reduced so that the males' antennae look little different from the females' (if you look very closely, they still have just a bit of a finger on each of the middle antennal segments). This led historically to a fair bit of confusion in the recognition of Sympiesis, with many species originally being placed in segregate genera (often with tongue-twistery compound names such as Asympiesiella or Sympiesonecremnus; thank you again, Alexandre Girault). Even now, the status of Sympiesis with regard to some related smaller genera could do with further investigation; we may yet see it grow again.

REFERENCES

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Miller, C. D. 1970. The Nearctic species of Pnigalio and Sympiesis (Hymenoptera: Eulophidae). Memoirs of the Entomological Society of Canada 102 (Suppl. S68): 5–121.

The Velvet Spiders: High Society

Communal web of Stegodyphus, copyright V. B. Whitehead.


In John Wyndham's novel Web (published in 1979, some ten years after Wyndham's own death), a group of settlers attempting to establish a utopian society on a remote Pacific island find themselves besieged by spiders. Contrary to the usual solitary habits of their kind, the spiders of Web have evolved a social structure like that of wasps or ants, and form roving packs that can overwhelm and devour animals many times their size. Fortunately for us, no such rapacious beasts exist in real life. But there are social spiders, even if they do not present a threat to anything much larger than a big insect.

The social habit has evolved in spiders on a number of occasions, but is perhaps best developed in some species of the genus Stegodyphus. This is a genus of the family Eresidae, commonly known as the velvet spiders. Eresids are small spiders, distinguished from most others by their subrectangular carapace with the front edge produced into a hood above the chelicerae (Miller et al. 2012). They have the full spider complement of eight eyes, with the posterior median eyes generally enlarged and directed forwards. Together with their covering of plush fur (hence the name 'velvet' spiders), this gives them an appearance distinctly reminiscent of some sort of carnivorous muppet.

Male Eresus cinnaberinus, copyright Ferenc Samu.


There are nine currently recognised genera of eresids, though only Stegodyphus includes social species. The family is mostly restricted to the Old World, with a single species Stegodyphus manaus known from Amazonian Brazil. A second species, S. annulipes, was originally described as Brazilian, but has since been collected from Israel and appears to have been mislabelled (Miller et al. 2010). Members of the temperate Eurasian genus Eresus are commonly known as 'ladybird spiders' as males often have a striking abdominal colour pattern of black spots on a red background. Most eresids live in silken tubes under objects such as stones or underground, whereas Stegodyphus species construct their webs in vegetation (Miller et al. 2012).

Mature Stegodyphus lineatus feeding hatchlings, copyright jorgemotalmeida.


The communal webs of social Stegodyphus species may extend for several metres. When an animal becomes trapped in the web, as many spiders as are able to reach it swarm over, all biting and salivating as they can. As a result, the members of the colony are able to kill and digest much larger prey than they could otherwise handle alone. Sociality in Stegodyphus appears to have arisen as an extension of the parental care found in other eresids. Females of both Stegodyphus and Eresus will regurgitate food for newly hatched young (Kullmann 1972). In social Stegodyphus, young are cared for communally and females will feed the young of their nest-mates as well as their own. Eventually, the young begin feeding directly on the caring female herself, draining her haemolymph to the point of rapid death. Again, in social Stegodyphus, this fate awaits all mature adults in the colony, and there is no overlap between generations (Schneider 2002). After the death of their mother, juvenile Eresus and non-social Stegodyphus remain in a group until they are closer to maturity; social behaviour could have arisen through a simple delay in the time of dispersal.

REFERENCES

Kullmann, E. J. 1972. Evolution of social behavior in spiders (Araneae; Eresidae and Theridiidae). American Zoologist 12: 419–426.

Miller, J. A., A. Carmichael, M. J. Ramírez, J. C. Spagna, C. R. Haddad, M. Řezáč, J. Johannesen, J. Král, X.-P. Wang & C. E. Griswold. 2010. Phylogeny of entelegyne spiders: Affinities of the family Penestomidae (NEW RANK), generic phylogeny of Eresidae, and asymmetric rates of change in spinning organ evolution (Araneae, Araneoidea, Entelegynae). Molecular Phylogenetics and Evolution 55: 786–804.

Miller, J. A., C. E. Griswold, N. Scharff, M. Řezáč, T. Szűts & M. Marhabaie. 2012. The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae). ZooKeys 195: 1–144.

Schneider, J. M. 2002. Reproductive state and care giving in Stegodyphus (Araneae: Eresidae) and the implications for the evolution of sociality. Animal Behaviour 63: 649–658.

The Problem with Sacesphorus

Probably not the subject of today's post: an unidentified assamiid from Thailand, from here.


One of the most frustrating things about many older taxonomic resources can be the shortage of illustrations. Up until the early part of the twentieth century, at least, it was something of a rarity for a publication to include extensive figures of their subject(s). So when I was presented for my semi-random subject of the week with the Burmese assamiid Sacesphorus maculatus, described by T. Thorell in 1889, I was not entirely surprised to discover that this Asian harvestman has never actually been illustrated.

At present, Sacesphorus maculatus is the only recognised species in its genus, known only from the Bago region in southern Burma. Unfortunately, that in itself doesn't necessarily indicate much. The Assamiidae are perhaps the most diverse group of Laniatores (short-legged harvestmen) in the tropics of the Old World, but they are also some of the least studied. The last extensive revision of the family was by our old friend Carl-Friedrich Roewer in 1935, and like many of Roewer's classifications its accuracy is suspect. Roewer divided the assamiids between seventeen subfamilies, but the characters separating most of these subfamilies are fairly superficial and probably do not reflect actual relationships (Staręga implicitly synonymised some of the African subfamilies in 1992 when he synonymised genera from different 'subfamilies' together). Roewer placed Sacesphorus in the Erecinae, which he characterised by features such as the absence of a pseudonychium (a claw-like process between the two true claws at the end of each leg), smooth leg claws, a two-segmented telotarsus on the front legs, and the absence of a median spine along the front edge of the carapace. All of these are fairly generalised characters, and some (such as tarsal segment number) are probably more variable than Roewer realised. Many short-legged harvestmen possess a pseudonychium as nymphs but lose it as they grow into adulthood, and the distribution of this feature in the assamiids may require more investigation.

Similar issues attend the identification of assamiid genera. Thorell (1889) originally distinguished Sacesphorus from the genus Pygoplus, also found in Burma and eastern India, by the presence in the former of a small spine in the middle of the eyemound. Roewer recognised a number of 'erecine' genera in Burma and eastern India, largely on the basis of tarsal segment numbers and armature of the dorsum. The relationship between all these genera deserves a second look. Many other groups of harvestmen have been successfully raised from the Roewerian quagmire in recent years; the assamiids are still waiting.

REFERENCES

Roewer, C.-F. 1935. Alte und neue Assamiidae. Weitere Weberknechte VIII. (8. Ergänzung der “Weberknechte der Erde” 1923). Veröffentlichungen aus dem Deutschen Kolonial- und Uebersee-Museum in Bremen 1: 1–168.

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

Thorell, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XXI.—Aracnidi Artrogastri Birmani raccolti da L. Fea nel 1885–1887. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 521–729.

Let's You and Me Enter Syzygy

Finally, you and your beloved are together. For the two of you, there are no others; all the world is yours alone. You gaze into each other's eyes, and then you pull your beloved into an embrace. Your lips touch in a passionate kiss. Your arms and legs intertwine in a firm hold. As you press so close to one another, it almost feels like you can no longer tell where the dividing line is between you. The excitement builds, and then... the two of you explode, each dissolving into a cascading avalanche of twitching gobbets of flesh.

Life cycle of Lecudina, from Clopton (2002).


This, roughly, is syzygy, a key event in the life cycle of many of the invertebrate-gut-parasitising protists known as eugregarines. Originally, the term 'syzygy' referred to the conjunction of two heavenly bodies, and provides a very poetic label for the process by which two of these unicellular organisms conjoin, rotating around one another as they produce and outer membrane to contain themselves within a single gametocyst. Once within the gametocyst, each divides into numerous gametes (which are produced through straight mitotic division, as eugregarines are haploid at maturity rather than diploid like ourselves). The resulting gametes will then be released from the gametocyst to fuse with one another in the production of diploid zygotes. Each zygote encloses itself in a resistant oocyst, in which state it may be passed out of the host's digestive system and be swallowed by a new host. While within the oocyst, the zygote divides to produce a number of new haploid individuals. Once the oocyst is in a suitable host, the new eugregarines are released, ready to feed and hopefully to eventually find a syzygy of their own.

Mature individuals of Blabericola in association, copyright R. E. Clopton.


Eugregarines have been referred to at this site before. As described in that post, they are part of the group of protists known as gregarines. Eugregarines differ from the other two major subgroups of gregarines, the archigregarines and neogregarines, in that they do not include an extensive asexually reproducing phase in their life cycle in addition to the sexual phase. All known eugregarines are parasites of invertebrates: their hosts include arthropods, molluscs, annelids and tunicates. Most eugregarines parasitise only a single host species over the course of their life cycle. The only known exception is members of the family Porosporidae, which are believed to spend part of their life cycle in a crustacean, and part in a mollusc. However, the porosporid life cycle has only been observed in its entirety once in 1940, when H. F. Prytherch fed infective spores from an oyster to crabs. It has been suggested that Prytherch may have conflated two separate parasites, with the eugregarine infection observed in the crabs after feeding them the spores actually representing a pre-existing infection that they had been carrying before the start of the experiment (Clopton 2002).

Individual of Schneideria quadrinotatus, from Clopton (2002); scale bar = 100 µm. Offhand, I can't be the only one who can't help seeing the nuclei in these sort of drawings as eyes. And for some reason, they always seem to look a bit wistful.


The eugregarines are usually divided between three suborders. Two of these, the Septatorina and Aseptatorina, include the great majority of species and are distinguished (as their names suggest) by the presence or absence of septae dividing the cell into sections. The third suborder includes the single small genus Siedlickia, parasites of marine annelids, which differs from other eugregarines in that it does not go through syzygy; instead, reproductive cells are budded directly off the mature feeding cells. The relationships between the three suborders are largely unknown; the Aseptatorina in particular seems to be defined largely by plesiomorphies. Clopton (2009) argued for a marine ancestry of eugregarines as a whole, and that the radiation of the septate eugregarines had been driven by adaptations of the gametocyst allowing their transmission in freshwater and terrestrial habitats. However, both the aseptate and septate eugregarines include parasites of marine, freshwater and terrestrial hosts. The fact that Clopton did not refer in 2009 to the marine members of the Septatorina (in the Porosporidae and various families of the Gregarinoidea) is somewhat bemusing as he himself had reviewed them some years earlier in his 2002 chapter on the eugregarines for The Illustrated Guide to the Protozoa. It is possible that he simply assumed the marine species to sit outside the terrestrial-freshwater clade, but it would have been nice for hime to say so.

Multiple syzygy in Hyalospora roscoviana, from Clopton (2002). The one in front doesn't look like it was quite expecting this.


Ignorance of the marine eugregarines does seem to be a theme, though: they're definitely less well-studied than the parasites of terrestrial species. Not that the latter can claim to have been exhaustively studied either: as noted by Clopton (2002), while over 1600 species of eugregarine have been described, only a fraction (much less than one percent) of potential hosts have been investigated for their presence. As almost every investigation of a new host results in the description of new parasite species, it is possible that the total number of eugregarine species out there ranks in the millions. Eugregarines are morphologically and behaviourally diverse. Attachment to the cells of the host's intestinal lining is via a structure called the epimerite, which may be a simple nubbin or may be a complex branching, fingered, collared or dart-like structure. When not attached to the host cell, most eugregarines move by gliding, but the worm-like Selenidiidae move by nondirectional swinging or thrashing. Many taxa are all distinguished by the characteristics of their syzygy. They may connect end to end, or they may lie top-to-tail. Members of the septate superfamily Gregarinoidea form associations some time before entering actual syzygy, so they are often found connected (whereas other taxa that do not become conjoined until the point of syzygy are more often found as isolated cells). Syzygy is most often between two individuals, but some Gregarinoidea regularly form associations of three or more. At least one species, Hirmocystis polymorpha, has been found in head-to-tail chains of up to twelve individuals. Whether such polygamous associations lead to all the individuals involved combining to form one gametocyst, or whether some form of competition occurs to whittle them down to a single victorious pair, is something I haven't yet discovered.

REFERENCES

Clopton, R. E. 2002. Order Eugregarinorida Léger, 1900. In: Lee, J. J., G. Leedale, D. Patterson & P. C. Bradbury (eds) Illustrated Guide to the Protozoa, 2nd ed., vol. 1 pp. 205–288. Society of Protozoologists: Lawrence (Kansas).

Clopton, R. E. 2009. Phylogenetic relationships, evolution, and systematic revision of the septate gregarines (Apicomplexa: Eugregarinorida: Septatorina). Comp. Parasitol. 76 (2): 167–190.

Melastomes and Pals

Princess flower Tibouchina heteromalla, copyright João Medeiros.


For most botanists currently working on flowering plants, the default taxonomic framework for their studies is the classification published by the Angiosperm Phylogeny Group. This is not the only classification for angiosperms proposed in recent years, but it is the most widely recognised, and it is the one that all its competitors are compared to. If there is one major deficiency of the Angiosperm Phylogeny Group classification, it is that it eschews the use of formal categories between 'orders' (which it tends to define broadly) and 'families'. As such, there are a number of well-supported clades that require one to turn to alternative classifications for suitable names.

Crypteronia paniculata, copyright Tony Rodd.


The Melastomatineae, as recognised by Reveal (2012), for instance, is a clade of mostly tropical and subtropical plants within the Myrtales commonly recovered by molecular phylogenetic analyses. Most of its members are included in the pantropical family Melastomataceae, but it also includes three much smaller and more localised families: the southeast Asian Crypteroniaceae, the western Neotropical Alzatea and the African Penaeaceae. Some authors also divide the Melastomataceae into two families Melastomataceae and Memecylaceae, but as the two groups are universally accepted as sister clades this is purely a matter of taste. Morphological characters uniting the families are few (see the Angiosperm Phylogeny Website). Many accumulate aluminium in the leaves, to the extent that the leaves of the small tree Memecylon edule were used in India as a mordant for fixing dyes to cloth. The flowers lack nectaries, and when nectar is produced in the Melastomataceae it exudes from locations such as the anthers or the corolla. Some Olisbeoideae (the Memecylaceae of other classifications) produce oil from glands on the anthers that is collected by pollinators in lieu of nectar. In the South American genus Axinaea, the anthers have a sugary appendage that is eaten by tanagers; as they attack it, a puff of pollen dusts their head. The anthers of Melastomataceae are often distinctly coloured from the rest of the flower, and arranged in a distinctive seried row on one side of the flower. The Melastomatoideae (i.e. Melastomataceae sensu stricto) also have distinctive leaves, with three or more strong longitudinal veins arising from the leaf base and connected by cross-veins.

Mountain hard pear Olinia emarginata, copyright H. Robertson. Olinia leaves smell of almonds when crushed, due to the presence of a cyanogenic compound.


Though Melastomataceae are often significant members of the tropical forest understorey, they tend not to have much direct economic significance for humans. Some, such as the princess flowers or glory trees of the genus Tibouchina, are grown as ornamentals. The hard pear Olinia ventosa of the Penaeaceae (no, I don't know why it's called that) is grown as a shade tree in South Africa. A few species of Melastomataceae are significant invasive weeds in warmer regions, including such luminaries as the Straits rhododendron Melastoma malabathricum and the evocatively-named Koster's curse Clidemia hirta. The velvet tree Miconia calvescens has earned itself the label of the 'purple plague' in Hawaii, where it over-runs native forest.

REFERENCE

Reveal, J. L. 2012. An outline of a classification scheme for extant flowering plants. Phytoneuron 37: 1–221.

Orioles: The Genuine Article

Eurasian golden oriole Oriolus oriolus, copyright Crusier.


It is widely appreciated that the British during the age of exploration were probably not the most imaginative of baptisers. Thanks to their tendency to label the fauna of foreign lands with the names of familiar animals back home, we are regularly confronted with warblers that aren't warblers, cod that aren't cod, monkeys that aren't monkeys. And for years, many an American has laboured under the mistaken impression that they know what an oriole is. This post is about the real orioles.

The Oriolidae are a family of birds found mostly in the tropics of the Old World, from Africa to Australia. Only a few species in the family are known from temperate climes. One of these is the original oriole, the European Oriolus oriolus, which migrates between sub-Saharan Africa and its breeding range in Europe and central Asia. The name 'oriole' is derived from the Latin word for 'golden', and there is no question of this being an appropriate name for the European bird. The male's plumage is almost entirely golden yellow, with the wings being black. As is commonly the way with birds, the females are less dramatic, being predominantly green. Despite the males' bright coloration, though, orioles are by all accounts fairly retiring birds, usually remaining secluded in the tree canopy, where they seek out fruit and small insects.

Black-and-crimson oriole Oriolus cruentus malayanus, copyright Christopher Hill.


The majority of the about thirty remaining species of Oriolus are also some combination of gold, green and/or black, but there are notable exceptions. A clade of Australo-Papuan and Moluccan species, identified by Jønsson et al. (2010) as the sister group to the other Oriolus species, contains relatively dull brown or greenish species. The Moluccan species in this clade bear a strong resemblance to friarbirds, a group of honeyeaters found in the same region, to the extent that the black-eared oriole Oriolus bouroensis was first described as a friarbird. It has been suggested that this represents a case of mimicry with the retiring orioles gaining a degree of protection from their resemblance to the aggressive friarbirds (Dickinson 2004). Another Asian clade identified by Jønsson et al. (2010) includes mostly red and black species. It also includes the silver oriole Oriolus mellianus in which the red coloration has been mostly lost, so that it is mostly silver-white with a black head and wings.

Male and female Australasian figbirds Sphecotheres vieilloti, copyright Jim Bendon.


Also included in the Oriolidae are the three species of figbird in the genus Sphecotheres, found in the Australo-Papuan region. The figbirds, as their name suggests, have a higher proportion of fruit in their diet than orioles. They are also more sociable, living in small flocks. Figbirds are distinguished from orioles by the presence of patches of bright red bare skin around their eyes; they are otherwise a dull greenish colour. Recent studies have also indicated oriolid affinities for Pitohui, a genus of two red and black birds, the hooded pitohui P. dichrous and variable pitohui P. kirhocephalus, found in New Guinea. Previous authors have included six species in Pitohui, but phylogenetic studies have revealed that the genus in the broad sense is widely polyphyletic, with the remaining species belonging to different bird families. The red and black markings of the 'pitohuis' are a case of aposematic coloration, advertising that its bearer is toxic. The pitohuis contain batrachotoxins in their skin and feathers, a similar substance to that found in the poison-arrow frogs of South America. Contrary to what you may read elsewhere, the pitohuis were not the first known case of toxicity in birds, though it was one of the most definite ones. It has been known since ancient times that migratory quail Coturnix coturnix are toxic at certain points on their migratory route: the biblical book of Numbers describes a case of mass poisoning suffered by the Israelites during the exodus. Other examples of birds that are at least seasonally toxic include the spur-winged goose Plectropterus gambensis and the bronzewing pigeons of the genus Phaps (a brief review of bird toxicity is provided by Bartram & Boland, 2001). As far as is known, all cases of toxicity in birds result from feeding on something containing the relevant toxic substance (probably beetles, in the case of pitohuis) which is then sequestered by the bird.

Mounted North Island piopio Turnagra tanagra, copyright Te Papa.


The Australo-Papuan distribution of these two genera, together with the basal position of the Australo-Papuan species in the genus Oriolus, suggests that the family originated in this area before crossing the Wallace Line to diversify in Eurasia and Africa (Jønsson et al. 2010). An Australo-Papuan origin for the orioles also correlates with the presence of a fossil oriolid, Longmornis robustirostrata, in the early Miocene Riversleigh deposit of Australia (Boles 1999). It also correlates with the recent identification as oriolids of the now extinct New Zealand piopios of the genus Turnagra (Zuccon & Ericson 2012). The piopios were two species (the South Island piopio Turnagra capensis and the North Island T. tanagra) of mostly brown songbirds, also commonly known as the New Zealand thrushes. Their song was described as being amongst the most beautiful of any New Zealand bird, both complex and with a propensity towards mimicking other birds. Though seemingly common at the time of European settlement, they declined rapidly and probably became extinct around the start of the 20th Century. The affinities of the piopios were long contentious, with leading suggestions including a relationship with the whistlers of the Pachycephalidae, or with the bowerbirds of the Ptilonorhynchidae. Zuccon & Ericson (2012) marshalled an array of molecular, morphological and behavioural evidence in favour of a relationship with the orioles, though this stands in contrast with an earlier molecular study that supported the bowerbird hypothesis (Zuccon & Ericson noted that the cytochrome b sequence reported in the earlier study did not correspond with the one they found themselves, and suggested that it may have been the result of contamination). The dull coloration of the piopios compared to other orioles was explained by Zuccon & Ericson as a loss of sexual dimorphism, but this may have been unnecessary: they seem to have overlooked the similarly dull coloration of a number of other basal oriolids. The fact that the piopios were described as more terrestrial than the other oriolids is also not unusual in the New Zealand context. After all, the New Zealand bird fauna is famed for its tendency towards terrestrialisation (it even included a terrestrial owlet-nightjar!) In an environment where the main threat came from above in the form of birds of prey, the ground must have seemed like a welcoming place to be.

REFERENCES

Bartram, S., & W. Boland. 2001. Chemistry and ecology of toxic birds. ChemBioChem 2: 809–811.

Boles, W. E. 1999. A new songbird (Aves: Passeriformes: Oriolidae) from the Miocene of Riversleigh, northwestern Queensland, Australia. Alcheringa 23: 51-56.

Dickinson, E. C. 2004. Systematic notes on Asian birds. 42. A preliminary review of the Oriolidae. Zool. Verh. Leiden 350: 47-63.

Jønsson, K. A., R. C. K. Bowie, R. G. Moyle, M. Irestedt, L. Christidis, J. A. Norman & J. Fjeldsa. 2010. Phylogeny and biogeography of Oriolidae (Aves: Passeriformes). Ecography 33: 232–241.

Zuccon, D., & P. G. P. Ericson. 2012. Molecular and morphological evidences place the extinct New Zealand endemic Turnagra capensis in the Oriolidae. Molecular Phylogenetics and Evolution 62: 414–426.

Barrallier's Monkey

"Gogy told me that they had brought portions of a monkey (in the native language "colo"), but they had cut it in pieces, and the head, which I should have liked to secure, had disappeared. I could only get two feet through an exchange which Gogy made for two spears and one tomahawk. I sent these two feet to the Governor in a bottle of spirits."

In November 1802, Governor Philip King sent an exploratory expedition west of Sydney under the command of Ensign Francis Barrallier, a French ex-pat who had taken service with the British after fleeing France with his parents following the French revolution. As well as finding a passage across the mountains that barred Sydney from the interior, Barrallier was trying to find the seat of a figure that Governor King later referred to in letters as the 'King of the Mountains'. Who exactly this King of the Mountains was supposed to be is unclear. Many have thought he was supposed to be some sort of overlord of the local Aboriginals. David Levell, in his 2008 book Tour to Hell, argues the King of the Mountains to have been the head of a secret inland settlement that many of the convicts imprisoned in Sydney believed would offer sanctuary to any who escaped there. Barrallier returned to Sydney in late December, having failed to locate either passage or king (the one would be discovered later, the other would prove to be mythical under any interpretation). Barralier's journal of his expedition languished in relative obscurity until an English translation was published in 1897.

The main interest for later readers of Barrallier's account has been in his dealings with the indigenous people he encountered and worked with. Barrallier had an interest in developing a rapport with the local people he met that was not shared by most of his British associates and his notes, sparse as they may be, provide one of the few direct records available of pre-colonial life in the Sydney region. I've brought Barrallier into this post, however, because of an incident he describes briefly in his journal where the game procured by some of Barrallier's aboriginal associates included an animal that Barrallier refers to as a 'monkey'. Barrallier did not see the animal's remains before it had already been butchered, but he is still the first European known to have acquired a specimen of one of Australia's most iconic animals: the koala.

Koalas Phascolarctos cinereus, photographed by Dinkum.


Koalas are widespread in the east of Australia, though loss of habitat has rendered their distribution localised in some areas. To most people outside Australia, the koala seems like a plush toy come to life, the essence of cuteness manifest in a single animal. The Australians themselves often have a more ambivalent attitude: while the koala is certainly a high-ranking member of the pantheon of the Australian fauna, together with such luminaries as the kangaroo, the platypus, the kookaburra and the gumnut baby, Australians also tend to look upon it as indolent, bad-tempered, and steeped in the kind of aroma that only an exclusive diet of eucalyptus leaves can give an animal (many Australians look more affectionately on the koala's closest living cousin, the wombat). To zoologists, Phascolarctos cinereus is the only surviving species of a lineage that goes back at least to the late Oligocene. Three subspecies of koala have been recognised, but these probably represent clinal variations rather than geographically discrete units (Houlden et al. 1999).

At just what point koalas became eucalyptus specialists is something we don't know for sure. The late Oligocene Perikoala palankarinnica possesses an ankylosed lower jaw (i.e. one that has the two sides fused together at the front) that may indicate a diet of tough leaves (Long et al 2002). Eucalyptus would be at least one candidate for such a diet. However, Perikoala's rough contemporary, Madakoala, lacked such a fused jaw and may have taken softer browse. Nor is a fused lower jaw present in the Miocene genera Litokoala or Nimiokoala (Louys et al. 2009). It seems likely that specialisation on Euclayptus may only have developed with the modern genus Phascolarctos, corresponding with the rise of eucalypt dominance in the Australian flora in the late Miocene. As well as being potentially less specialised, the fossil genera of koalas were also distinctly smaller than the living species. Koala evolution reached an apogee of sorts in the Pliocene and Pleistocene with the fossil species Phascolarctos yorkensis, which tipped the scales at nearly twice the size of P. cinereus (Long et al. 2002) (somewhat disappointingly, no-one seems to seen fit to present a fossil koala with the name of Katastaxarctos).


Koalas can be very vocal animals, using bellows and grunts as their main method of communicating. This video of a vocalising bull comes from here.


The specialisation of the modern koala is truly a remarkable thing. True exclusivity of diet seems to be a rarity among large terrestrial vertebrates (and as it can reach sizes of 20 kg, there is no denying that the koala is a large vertebrate). Many have their preferred delicacies but remain far from averse to the occasional variation (something that I really wish the ABC had been more aware of with that lorikeet article). Thus we have cattle gnawing on bones, cats eating grass, or deer killing and eating birds. Even the giant panda, perhaps the other specialist mammal most familiar to the general public, has been known to supplement its bamboo diet with roots and small animals. But the koala turns up its nose at almost anything other than Eucalyptus leaves—and usually only a small number of Eucalyptus species at that. The toughness of Eucalyptus leaves mean they require a great deal of digestive processing, and the small nutritive return is responsible for the extended periods of inactivity that koalas are known for. Early British naturalists often compared the koala to the South American sloth, which functions under similar constraints. The low nutrition of their diet is also reflected in the notoriously small brains of koalas, which have one of the smallest brains relative to body size of any mammal. So noxious is the eucalypt diet that koala joeys have to be weaned onto it through stages. When a joey is about six months old, its mother starts producing a faecal pap of half-digested leaves that the joey eats direct from her cloaca before moving to a more direct leaf diet about a month later.

Nevertheless, by specialising on Eucalyptus leaves, koalas have access to an abundant food source that few other mammals can handle. Even after the arrival of Europeans, koalas have handled the incursion of foreign predators better than many other Australian natives. The main threat to their continued existence is clearing of the forests on which they depend for food. The koala deserves its position as an icon, and an icon is worthy of respect.

ARKive video - Koala joey eating pap
Video of a koala joey feeding on pap, from Arkive.


REFERENCES

Houlden, B. A., B. H. Costello, D. Sharkey, E. V. Fowler, A. Melzer, W. Ellis, F. Carrick, P. R. Baverstock & M. S. Elphinstone. 1999. Phylogeographic differentiation in the mitochondrial control region in the koala, Phascolarctos cinereus (Goldfuss 1817). Molecular Ecology 8 (6): 999–1011.

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.

Louys, J., K. Aplin, R. M. D. Beck & M. Archer. 2009. Cranial anatomy of Oligo-Miocene koalas (Diprotodontia: Phascolarctidae): stages in the evolution of an extreme leaf-eating specialization. Journal of Vertebrate Paleontology 29 (4): 981–992.