Field of Science

Spotless Ladybirds

Ladybirds are one of the first types of insect most children learn to recognise. Most people recognise these beetles straight away with their bright, contrasting colour patterns and shiny elytra. Many people are also aware of the beneficial role they may play in gardens and horticulture, feeding as both adults and larvae on plant-sucking insects such as aphids. However, ladybirds are a lot more diverse than many people realise. The great majority of species in the ladybird family Coccinellidae are not brightly coloured, and many of them do not feed on aphids.

Rhyzobius cf. chrysomeloides, a typical member of the Coccidulini, copyright Mick Talbot.

The ladybirds of the cosmopolitan tribe Coccidulini are mostly small species; many are dully coloured, though some are more distinctly patterned. As recognised by Ślipiński (2007) in his revision of the Australian Coccinellidae, the group is a diverse one and difficult to define; common features of its members include pubescent rather than shiny elytra, and certain features of the genitalia. A phylogenetic analysis of coccinellids by Seago et al. (2011) did not resolve the Coccidulini as a monophyletic clade, suggesting the possibility that they may represent a basal grade of unspecialised taxa, but no reclassification of the group appears to have been proposed as yet.

Stethorus punctillum, copyright Gilles San Martin.

The most familiar ladybirds are more or less hemispherical, but coccidulin species may vary in shape from rounded to oblong, with the body convex to flattened. The Tasmanian species Nat vandenbergae* particularly stands out in this regard, being oval and flattened with relatively long legs and antennae, and at first glance might be mistaken for a chrysomelid leaf beetle rather than a coccinellid. The rounded species of the genera Stethorus and Parastethorus include some of the smallest of all ladybirds, some being only a millimetre or so in length.

*Yes, the genus name for this species is Nat. Other coccidulin genera named by Ślipiński (2007) include Roger and Robert.

A mealybug destroyer Cryptolaemus montrouzieri engaged in destroying a mealybug, copyright Cynthia Bingham Keiser.

Being so small and easily overlooked, the life habits of many coccidulins remain poorly known. Many feed on aphids or scale insects in the usual ladybird manner. The orange-and-black Cryptolaemus montrouzieri was one of the first ladybirds to be deliberately used for biological control of pests; originally native to eastern Australia, it has been introduced to other parts of the world such as the United States to help control mealybugs. The tiny Stethorus and Parastethorus species feed on spider mites (Tetranychidae), species of which may similarly cause damage to crops. But other coccidulins have more unexpected diets. The larva of another Australian species, Bucolus fourneti, is a specialised predator of ants, lying in wait under the bark of Eucalyptus trees to ambush its prey as it passes by.


Seago, A. E., J. A. Giorgi, J. Li & A. Ślipiński. 2011. Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetics and Evolution 60: 137–151.

Ślipiński, A. 2007. Australian Ladybird Beetles (Coleoptera: Coccinellidae): Their biology and classification. Australian Biological Resources Study: Canberra.

Perilampella acaciaediscoloris: An Australian Gall Wasp

The small but incredibly diverse chalcidoid wasps are mostly known as parasitoids, their larvae attacking the eggs and young of others insects. Some, however, have chosen the vegetarian option, inserting their eggs into plant rather than animal tissue. As the larva develops, it induces the host plant to develop an often bizarre-looking growth around it that provides both shelter and food; this growth is known as a gall.

Antenna, forewing venation and dorsum of Perilampella acaciaediscoloris, from Bouček (1988).

Perilampella acaciaediscoloris is an gall-forming wasp that was first described by Froggatt in 1892 from galls that he collected on the wattle Acacia discolor, a species now regarded as a synonym of the sunshine wattle A. terminalis of south-eastern Australia. Froggatt placed his species in the genus Cynips (which is not part of the Chalcidoidea but belongs to a different micro-wasp superfamily, the Cynipoidea) but it is now placed in the chalcid subfamily Ormocerinae in the (polyphyletic) Pteromalidae. Ormocerinae are fairly generalised 'pteromalids' that are non-metallic in colour and often finely sculpted. So far as is known, ormocerines are all associated with galls in one way or another, either as gall-causers themselves or as inquilines (species that lay their eggs in the galls caused by other insects). Another ormocerine species, Trichilogaster acaciaelongifoliae, has been introduced from Australia to South Africa to help control the Sydney golden wattle Acacia longifolia.

The related ormocerine Trichilogaster acaciaelongifoliae, copyright Simon van Noort. Perilampella acaciaediscoloris most obviously differs from this species in its hairier and darker wings, and more shiny mesosoma.

The genus Perilampella differs from other ormocerines in being particularly shiny, with little clear setation. Bouček (1988) listed four Australian species in the genus, noting that P. acaciaediscoloris could be recognised by its very dark, long wings and shiny orange-yellow mesosoma. Froggatt (1892) described the galls of P. acaciaediscoloris as formed at the inception of a leaf bud or new shoot. Sometimes, they would be little more than swellings at the base of the shoot. More often, they would be oval with three irregular horns formed from aborted leaf buds. Sometimes, P. acaciaediscoloris galls would be the target of inquilines of their own that caused the gall to degrade to a shapeless mass.


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

Froggatt, W. W. 1892. Notes on Australian Cynipidae, with descriptions of several new species. Proceedings of the Linnean Society of New South Wales, second series 7: 152–156.

The Adaeines: South Africa's Cryptic Micro-Giants

Adaeulum sp., copyright Charles Haddad.

The Triaenonychidae are the family of Gondwanan harvestmen. While there are other families of harvestmen with a Gondwanan distribution (such as my own favoured family, the Neopilionidae), none of them are nearly as widespread and diverse as the triaenonychids. Despite their diversity, however, our understanding of triaenonychid relationships remains uncertain, and the family's classification poorly defined.

Within their range in Africa, Australasia and South America, triaenonychids can easily be distinguished from most other families of short-legged harvestmen by the structure of the claws on the hind two pairs of legs. Whereas members of other families bear a pair of simple claws on these legs, triaenonychids have a single claw with side branches on each leg. Branched claws are also found in the New Zealand genus Synthetonychia, which occupies its own distinct family, but that genus is easily recognised by its unusual body shape without a distinct eyemound. The Gondwanan Triaenonychidae were divided by Roewer into three subfamilies (Triaenonychinae, Triaenobuninae and Adaeinae) based on the shape of the sternum (the plate running along the underside of the body between the leg coxae). The significance of this feature was later questioned by Forster (1954) who recognised two subfamilies Triaenonychinae and Soerensenellinae on the basis of claw morphology (soerensenellines having longer side branches on the claws than triaenonychines) and reduced Roewer's subfamilies to tribes of Triaenonychinae. No large-scale analysis of triaenonychid phylogeny has been done so far, so it remains unestablished whether we should prefer one classification or the other (or possibly neither).

Typical triaenonychine (left) and adaeine (right) sternal shapes, from Forster (1954).

The Adaeinae or Adaeini may be one of the better defined of Roewer's original subgroups and recent authors have expressed the opinion that this may indeed turn out to be a natural clade. Whereas members of the Triaenonychinae sensu stricto and Triaenobuninae have a sternum that has a spearhead-shaped expansion at the front end and a broadened base at the back, members of the Adaeinae have a sternum that is a triangular or wedge shape without a posterior expansion. The adaeines are likely to be endemic to southern Africa; Kury et al. (2014) did list a single Australian species, Dingupa glauerti, in the Adaeinae but I would hazard a guess that future study proves this species to be misplaced (as has been found with other Australasian 'adaeines').

About forty species of adaeines are currently recognised, all from South Africa, but it is entirely likely that more remain to be described. The hard, granular body surface of adaeines inevitably picks up a covering of dirt and grit, making them exceedingly difficult to spot when not moving. Nevertheless, adaeines can be quite large as harvestmen go, with some being up to a centimetre in body length. Conversely, Micradaeum rugosum, a species found in the vicinity of Cape Town, is only about three-and-a-half millimetres in body length (Lawrence 1929). As with other species of Triaenonychidae, the large, raptorial pedipalps are larger and more robust in male adaeines than in females, and often have more pronounced spines. In some species of the genus Larifuga, nowever, spines or denticles may be more prominent on the female's pedipalps than on the male's, though the male's pedipalps are still larger and stronger overall (Lawrence 1937).


Forster, R. R. 1954. The New Zealand harvestmen (sub-order Laniatores). Canterbury Museum Bulletin 2: 1–329.

Kury, A., A. Mendes & D. Souza. 2014. World checklist of Opiliones species (Arachnida). Part 1: Laniatores—Travunioidea and Triaenonychoidea. Biodiversity Data Journal 2: e4094. doi: 10.3897/BDJ.2.e4094

Lawrence, R. F. 1929. The harvest-spiders (Opiliones) of South Africa. Annals of the South African Museum 29 (2): 341–508.

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, pls 14–15.

Pines for the Sea

Black pine Neorhodomela larix growing alongside lighter-coloured Analipus japonicus on a beach in California, copyright Peter D. Tillman.

There aren't many types of algae that receive their own vernacular name. Despite the fact that thousands of different species of macroscopic algae exist in the world, the majority tend to get lumped under the catch-all term of 'seaweed'. Among those seaweeds that are visible enough to stand out from the general crowd is Neorhodomela larix, the 'black pine'.

Neorhodomela larix is a common seaweed species on the west coast of North America. It has also been recorded from the western side of the Pacific Ocean, but it should be noted that it might not always be readily distinguished from related species (indeed, N. larix was not distinguished at species level from the primarily western Pacific N. aculeata until the revision of this genus by Masuda, 1982). The name 'black pine' that has been bestowed on it reflects its overall appearance. Black pine grows as erect axes radiating from a small basal holdfast. These primary axes give rise to regular branches arranged in a spiral fashion; most of these side branches, at least in young individuals, only grow to a fixed, short length and remain unbranched or only bifurcate (in more mature individuals, some of the side branches become indeterminate in length and grow in a similar manner to the primary axes). The overall effect is to make the seaweed stems look like a bottlebrush or, with a bit of imagination, like a pine-tree branch (or at least like a plastic-Christmas-tree branch). Black pine is particularly common on horizontal, wave-exposed beaches about a foot above the mean low-tide mark; in some places it may form large, continuous stands. It is most abundant where said shorelines are swept by sand, probably because the action of the sand's movement keeps away grazers such as urchins (D'Antonio 1986).

Neorhodomela larix was previously included in the closely related genus Rhodomela, and some sources still refer to it as 'Rhodomela larix'. Masuda (1982) established Neorhodomela as a separate genus from Rhodomela due to differences in vegetative structure. Both genera possess hair-like branchlets known as trichoblasts, but in Rhodomela these arise from the main branches spirally whereas in Neorhodomela they are positioned dorsally and arise in a zig-zag arrangement. Spermatangia are produced from trichoblasts in Neorhodomela, whereas in Rhodomela the spermatangia are produced directly on the unspecialised branches and the trichoblasts are only vegetative structures. In N. larix, few vegetative trichoblasts are produced until shortly before the production of reproductive trichoblasts bearing spermatangia; other Neorhodomela species (such as N. aculeata) may start producing vegetative trichoblasts soon after germination.

Apart from the ecological role it presumably plays in providing food and shelter to other coastal lifeforms, Neorhodomela larix does not have much of a direct impact economically. Like many macroalgae, black pine produces chemical compounds known as bromophenols that have been subject to some investigation due to their potentially beneficial (such as anti-microbial or anti-oxidant) activities. However, to date no practical pharmaceutical products have been developed from algal bromophenols, in part because the amount of these compounds produced by the algae in vivo is fairly low (Liu et al. 2011). For now, there is little to disturb these little pine forests by the sea.


D'Antonio, C. M. 1986. Role of sand in the domination of hard substrata by the intertidal alga Rhodomela larix. Marine Ecology—Progress Series 27: 263–275.

Liu, M., P. E. Hansen & X. Lin. 2011. Bromophenols in marine algae and their bioactivities. Marine Drugs 9: 1273–1292.

Masuda, M. 1982. A systematic study of the tribe Rhodomeleae (Rhodomelaceae,Rhodophyta). Journal of the Faculty of Science, Hokkaido University, Series 5: Botany 12 (4): 209–400.

Where Do You Put Your Camels?

A dromedary Camelus dromedarius dares you to say what you make of it. Copyright John O'Neill.

In any discussion of the conflicts that may exist between morphological and molecular data in phylogenetic analysis, hippos and whales are bound to come up sooner or later. The claim in the late 1990s that these are each other's closest living relatives (and hence, that whales are nested fairly deeply within the artiodactyls, or even-toed hoofed mammals) was greeted with amazement, incredulity and more than a little skepticism. The story even caught the interest of the general public through news stories like this one, meaning that many people who are otherwise unfamiliar with the trivia of mammalian phylogeny may have picked up this detail. Since then, the whale-hippo relationship has been tested, re-tested and examined again, using every data source available. But the insertion of the whales was not the only way that molecular data mixed up the artiodactyl family tree. There was also the question of where one put the camels.

Based on anatomical data, it had previously been generally agreed that camelids (including camels and llamas) were most closely related to the ruminants, the group including such artiodactyls such as cattle, deer or giraffes. Both camelids and ruminants regurgitate cud pellets from the stomach back to the mouth in order to break their food more efficiently*, and both camelids and ruminants have a stomach divided into chambers with food only travelling to the rear section of the stomach after it has been re-chewed. They do differ in that whereas ruminants have the stomach divided into four distinct chambers, camelids only possess three; the rear two chambers (the abomasum and omasum) are not clearly differentiated in camelids. They were also united by features of the dentition, such as the presence of distinctly crescent-shaped cusps on the rear teeth. This latter feature lead the camelid+ruminant grouping to commonly be referred to as the Selenodontia (the 'moon-teeth').

*Yes, giraffes do chew cud. Yes, the cud does travel all the way between the stomach and the mouth each time.

However, the advent of molecular analyses cast doubt on this long-accepted arrangement. Instead of supporting the expected Selenodontia clade, molecular analyses placed camelids as the sister group to all other artiodactyls, with the ruminants instead being sister to the whales+hippos clade (with a pigs+peccaries clade the next clade out). This implied that the shared features of camelids and ruminants had arisen convergently (or else all other artiodactyls had reverted to a state considered more plesiomorphic for the group as a whole). In support for such a proposition, one might point to the ecological similarities in play. Camelids and ruminants are more specialist browsers than the non-selenodont artiodactyls, which are commonly more omnivorous (pigs and peccaries) or even carnivorous (whales).

Alternative phylogenies of artiodactyls, based on morphological (left) and molecular (right) data, from Spaulding et al (2009).

However, even if one is willing to credit that the 'selenodont' characters may have been the result of similar dietary pressures, one must also consider the issue that there are a number of fossil artiodactyl groups with selenodont or quasi-selenodont features. Examples of these include the Protoceratidae, a North American group that has commonly made some sort of appearance in popular books on fossil animals due to the weird home arrangements of some species, and the semi-bipedal Anoplotherium. In an influential morphological study of artiodactyl relationships, Gentry & Hooker (1988) referred to the possibility that some of these groups might be members of the selenodont stem, sitting outside the exclusive camelid+ruminant clade. Obviously, if selenodonts were not monophyletic, fossil 'selenodonts' might be aligned to either camelids or ruminants, but they couldn't be connected to both. Most studies that posited selenodont polyphyly, however, looked at living taxa only and did not consider extinct groups.

The most detailed study that I've found so far that considers the relationship between data from fossil taxa and from molecular sources in artiodactyl phylogeny is that published by Spaulding et al. (2009). This combined analysis of both morphological and molecular data produced results that were largely concordant with the latter, generally supporting placement of camelids as the sister group to all other artiodactyls (it's worth noting, mind you, that the size of the molecular data set used was considerably larger than that for the morphology, and an analysis of their morphological data only resulted in selenodont monophyly). The various 'proto-selenodonts' were scattered to the stems of various Recent clades. Protoceratids, for instance, were associated with ruminants rather than with camelids*. There are still a number of groups that remain yet to be analysed, but they've made a start.

*Another result of their analysis that is not directly relevant to the selenodont question, but cannot go unremarked upon, is that their tree indicates that Andrewsarchus, a lead contender for the title of largest terrestrial mammalian carnivore ever, might some sort of giant entelodont. I don't know how much I should read into this—not all of Andrewsarchus' potential relatives were included in Spaulding et al.'s analysis—but that's the sort of result that one just wants to be true.


Gentry, A. W., & J. J. Hooker. 1988. The phylogeny of the Artiodactyla. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol 2. Mammals pp. 235–272. Clarendon Press: Oxford.

Spaulding, M., M. A. O’Leary & J. Gatesy. 2009. Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretations of key fossils and character evolution. PLoS ONE 4(9): e7062. doi:10.1371/journal.pone.0007062

What We Have Here is a Failure to Communicate

To those without much of a background in taxonomy, the various rules governing the naming of organisms can seen frustratingly byzantine and laborious. "Surely," they think to themselves as they despairingly attempt to come to grips with concepts of holotypes and lectotypes, synonyms and homonyms, "there must be an easier way of doing this". Nevertheless, the easiest way to develop an appreciation for just how valuable it is to have a set of rules governing nomenclature is to attempt to deal with anything dating back to the days before such rules were established. Settle back, readers, while I tell you a tale. Pour yourself a drink. You're going to hate this.

Collonista glareola, a species that just might be related to the subject of this post, copyright Huang, Fu & Poppe.

These days, it is generally accepted that before a new name can enter general use, it should be clearly established in some form of formal, widely-accessible publication just to what it is that the name is supposed to refer. Back in the day, however, this was not always the case. A century or two ago, the communities of researchers working on a particular group of organisms were often small, and it was not uncommon for names to effectively spread through personal correspondence or word of mouth alone. One naturalist might refer to a new genus he had come to recognise in a letter to another, and the latter naturalist may then assign his own species to that genus without the first naturalist ever publishing a formal description. At the time, this might not be seen as much of an issue: after all, if there was ever any question as to the first naturalist's original intent, surely it could be clarified by simply writing to him personally?

The name Leptothyra seems to have been established in this kind of way in the mid-1800s by the American naturalist James Graham Cooper for a genus of small marine gastropods (belonging to the vetigastropods, related to the top shells and cat's-eyes) found on the coast of California. In 1871, W. H. Dall attributed the name to an unpublished manuscript of Cooper's and cited the type species as Linnaeus' Turbo sanguineus, a Mediterranean species to which Cooper had also attributed specimens from the Pacific. As it happens, Turbo sanguineus was already the type species for an earlier genus name, Homalopoma, so Dall's 'Leptothyra' would be considered invalid and give precedence to Homalopoma. However, in 1869 the name Leptothyra had been used by W. H. Pease for L. costata, a species from Hawaii, without direct reference to any other species (thus making L. costata the effective type species of Leptothyra). Subsequent authors often considered 'Leptothyra Pease' to be a separate genus from Homalopoma/'Leptothyra Dall'. At least one author who did not, Henry A. Pilsbry (1888), nevertheless used the name Leptothyra under the mistaken belief that the name Homalopoma was preoccupied. Over time, numerous species both living and fossil from around the Pacific were assigned to Leptothyra in one way or another.

It was not until over a century later that Coan (1986) pointed out that the name Leptothyra had appeared in print even earlier than Pease's usage. Cooper himself had used the name in a list of Californian molluscs in 1867. Even though Cooper's list lacked any descriptive details, this counts as enough to validate the name because he included species for which descriptions had already been published under other genera. One of these was Turbo sanguineus, which Coan officially designated as type species and fixed Leptothyra's status as an invalid later name for Homalopoma.

Which leaves open the question of what one should call the genus formerly known as 'Leptothyra Pease'. It doesn't help matters that Pease's 'Leptothyra costata' has apparently never been illustrated and its identity has been open to question. Iredale (1918) proposed the name Collonista for use with species previously included in Leptothyra, stating that the latter "proves to have been first published by Pease in connexion with a juvenile shell of a different genus", but gave no further elaboration or explanation how he reached that conclusion. The online resource WoRMS lists L. costata as a junior synonym of the widespread Pacific species Collonista verruca, but I have been unable to find where that synonymy was published. Nevertheless, any sort of replacement name for 'Leptothyra Pease' seems like it would be misguided at best. It is unlikely that L. costata represents any genus otherwise unknown and, even without any explicit statement to the effect, it is quite possible that Pease only intended to assign his species to Cooper's manuscript genus rather than establish a new genus of his own. Any concept of a genus Leptothyra is best left to sink into the annals of history.


Coan, E. 1986. Some additional taxonomic unites that first appear in publications by J. G. Cooper. Nautilus 100 (1): 30–32.

Cooper, J. G. 1867. Geographical catalogue of the Mollusca found west of the Rocky Mountains, between latitudes 33° and 49° north. Geological Survey of California: San Francisco.

Dall, W. H. 1871. Descriptions of sixty new forms of mollusks from the west coast of North America and the North Pacific Ocean, with notes on others already described. American Journal of Conchology 7 (2): 93–160, pls 13–16.

Iredale, T. 1918. Molluscan nomenclatural problems and solutions.—No. 1. Proceedings of the Malacological Society of London 13 (1–2): 28–40.

Pease, W. H. 1869. Descriptions of new species of marine Gasteropodae inhabiting Polynesia. American Journal of Conchology 5 (2): 64–79.

Philonthus: Too Many Staphylinids

Philonthus marginatus, copyright James K. Lindsey.

Working with staphylinids, it has to be said, can be horrible. They are treated as one of the most diverse of the beetle families—perhaps the most diverse of all—but compared to other diverse families they attract relatively little study. The majority of staphylinids are usually either very small or soft-bodied, not uncommonly both together, making them difficult to prepare and maintain as dry specimens. For the soft-bodied species, with their reduced elytra, many of the easily visible features that can be so useful for other beetle groups are obscure or unavailable. They also tend to be drab in coloration, without much in the way of striking patterning. As a result, it is often impossible to identify staphylinid species without examining minute features of the appendages or the genitalia. Something to keep in mind as you read the following.

Species of the genus Philonthus are relatively large as staphylinids go, often about half a centimetre in length, but they are certainly not free of the problems affecting other members of the family taxonomy-wise. The genus is massively diverse—over 1200 species have been described from around the world. Attempts have been made to break them down into more manageable chunks, such as through the recognition of subgenera, but these have mostly failed to gain much traction. Most recent authors have only recognised informal species groups within the greater mass.

Philonthus carbonarius, copyright James K. Lindsey.

In general, species of Philonthus are smooth, without excessive hairs, and have labial palps with the last segment fusiform and about as wide as the penultimate segment (Tottenham 1955; Stan 2012). Males have the aedeagus (the intromittent organ of the genitalia) rotated in the abdomen so its paramere (off-branch) is located on the left side rather than ventrally as in other genera (Tottenham 1955). Some species may have a metallic sheen to their coloration; others are a plainer black or reddish. Species may also differ in the number and arrangement of setae on the pronotum.

Where their lifestyles are known, most Philonthus are associated with decomposing organic matter such as animal dung, compost or leaf litter. Some are predators of other insects and insect larvae found in such habitats (such as fly larvae); these species have highly developed senses to locate decaying matter, and are strong fliers to disperse to suitable habitats (Majka et al. 2009). Some species of Philonthus may act as predators of other pest insects, helping to keep their numbers down.


Majka, C. G., J.-P. Michaud, G. Moreau & A. Smetana. 2009. Philonthus hepaticus (Coleoptera, Staphylinidae) in eastern Canada: are distribution gaps distinctive features or collecting artifacts? ZooKeys 22: 347–354.

Stan, M. 2012. On the species of Philonthus Stephens (Coleoptera: Staphylinidae: Staphylininae: Staphylinini: Philonthina) in the collections of Romanian natural history museums. Travaux du Muséum National d'Histoire Naturelle "Grigore Antipa" 55 (2): 233–276.

Tottenham, C. E. 1955. Studies in the genus Philonthus Stephens (Coleoptera: Staphylinidae). Parts II, III, and IV. Transactions of the Royal Entomology Society of London 106 (3): 153–195.