Barrow's Scaly Bark-louse

Male of Lithoseopsis humphreysi, from Taylor (2013). As this specimen has been preserved in ethanol, most of the wings' scales have been washed off.

In yesterday's post, I told you about our project's new book on the terrestrial invertebrates of Barrow Island. In this post, I want to tell you about my own main contribution: a description of Barrow Island's resident species of Amphientomidae.

Amphientomidae is a family of the bark-lice, the Psocodea (or Psocoptera, perhaps). They differ from most other bark-lice in having the wings densely covered in scales, like the wings of a moth; someone on BugGuide once referred to an amphientomid as a "moth-hopper-louse-bug thingy". These scales are often arranged into striking patterns of contrasting colours (take a look at the individual below). When the late Courtenay Smithers initially identified our collections of bark-lice from Barrow Island (which we briefly reviewed last year, though most are still not identified to formal species), he highlighted the presence of an amphientomid in the collection as particularly interesting. Amphientomids are incredibly little known in Australia. Only three Australian species had previously been described, and all three were only known from a single specimen. The description in my paper is drawn from six specimens, so it represents a tripling of Australia's published tally!

A North American species of Lithoseopsis, L. hellmani, photographed by Diane Young.

With this in mind, a description of the Barrow Island amphientomid seemed in order. This, of course, required comparing it to the already-described Australian species, which had thankfully been given detailed descriptions (Smithers 1989; New 1994). Two of these were from Western Australia: one from the Kimberley region in the far north, the other from the Cape Range which is on the mainland close to Barrow Island. Both of these had been placed in the genus Seopsis, which is otherwise known from Africa and Asia. However, when I compared the Barrow Island specimens to descriptions of Seopsis and other Asian genera, things didn't quite add up. For instance, take a look at the face of the Barrow Island species:

In particular, note the position of the ocelli (the three simple eyespots) at the front of the face. In all the Australian species, these are widely spaced, and the two lateral ocelli are right alongside the compound eyes. In contrast, here is the face of a Japanese amphientomid:

The individual in this photo, from here, isn't identified but may belong to Paramphientomum yumyum (yes, really, that's its name). Note how in this species, all three ocelli form a close triangle in the centre of the face; Seopsis also has this arrangement of ocelli. Other features such as genital morphology were also inconsistent between the Western Australian species and Seopsis.

Instead, the Western Australian species are better placed in a genus called Lithoseopsis*. The Barrow Island specimens I assigned to the same species that had earlier been described from the Cape Range, Lithoseopsis humphreysi. There are some minor differences between the Barrow specimens and the Cape Range holotype, but with only one specimen available from the latter locality I couldn't really assess whether these were indicative of more than one species. The recognition of this species as Lithoseopsis is interesting, as this genus is otherwise known only from southern North America. How such an oddly disjunct distribution may have come about I have no idea; my first guess would be that it's somehow relictual. However, it's also worth pointing out that there are three other genera very similar to Lithoseopsis that share its broadly-spaced ocelli: the African genus Hemiseopsis and the circum-Mediterranean genera Marcenendius and Nephax. The relationship between these genera really deserves a proper study.

*And here's a bit of a cautionary tale for you all. When I initially submitted the manuscript of this paper for review, one of the reviewers pointed out that the Western Australian species shouldn't be placed in Lithoseopsis due to the absence of one of that genus' primary features, a sclerotised plate on the back of the abdomen. Therefore, the manuscript was revised and accepted to establish the WA species as a new genus. At this point, I should note, the only amphientomids that had been available from Barrow Island were males. However, shortly before the book was due to be published, I finally received a female specimen. This specimen possessed the sclerotised plate that had been absent in the males! The paper was quickly revised at the last minute to return the WA species to Lithoseopsis. The ability to examine a female from Barrow also lead me to change my mind about whether the Barrow species was distinct from the Cape Range species, represented only by a female holotype. Unfortunately, once the book had been published and I could see the final product, I found that I had missed correcting the genus and species name in the figure captions! Much wailing and gnashing of teeth immediately commenced, I can bloody well tell you. After the whole pureora/pureroa thing, 2013 has not made me look good as a proof-reader.

The Barrow Island specimens also tell us something interesting about intra-specific variation in amphientomids. A common feature of bark-lice is variation within species of wing development: individuals of a single species may have the wings fully developed, reduced or absent. In amphientomids, however, such variation has been rarely recorded. A couple of species are known in which males and females differed in wing development, but so far as was known all males and all females had wings the same size. In the Barrow Island specimens, however, some males had both pairs of wings large and fully developed, but others had the forewings slightly shortened and the hind wings reduced to minute flaps. As the reduced-wing specimens were otherwise little different from the fully-winged specimens, they seem unlikely to represent different species. Instead, Lithoseopsis humphreysi is the first recorded example for amphientomids of wing polymorphism within a single sex.

REFERENCES

New, T. R. 1994. A second species of Amphientomidae (Insecta: Psocoptera) from Western Australia. Proceedings of the Linnean Society of New South Wales 114 (4): 233–236.

Smithers, C. N. 1989. Two new species of Amphientomidae (Insecta: Psocoptera), the first record of the family for Australia. Proceedings of the Linnean Society of New South Wales 111 (1): 31–35.

Taylor, C. K. 2013. The genus Lithoseopsis (Psocodea: Amphientomidae) in the Western Australian fauna, with description of the male of Lithoseopsis humphreysi from Barrow Island. Records of the Western Australian Museum Supplement 83: 245-252.

The Terrestrial Fauna of Barrow Island

Nihara R. Gunawardene, Jonathan D. Majer, Christopher K. Taylor & Mark S. Harvey (eds) 2013. The Terrestrial Invertebrate Fauna of Barrow Island, Western Australia. Records of the Western Australian Museum, Supplement 83. 406 pp.

For several years now, my colleagues and I have been monitoring terrestrial invertebrates on Barrow Island here in Western Australia. Some of you will have already heard of Barrow Island; for anyone that hasn't, Barrow is the second-largest island off the coast of WA (it's about 25 km long and 12 km wide). It has two main claims to fame: (a) it has been a recognised nature reserve for over 100 years, with thriving populations of a number of animals that are rare or extinct elsewhere, and (b) for the last 50 years, it has also been a working oil field, most recently managed by the oil company Chevron. It also lies close to large offshore natural gas deposits, and in 2003 Chevron and its associates were given permission to build a processing plant on Barrow Island for extraction of the gas. This permit, however, carried strong caveats: development of the plant is not to compromise the value of Barrow as a nature reserve. That's where we come in: on a regular basis, we travel to the island to look for any undesirables that may have managed to slip through the stringent quarantine requirements that have been placed on transport to Barrow (nothing so far, touch wood). Before plant development was begun, a large-scale survey was also conducted to identify the pre-existing invertebrate fauna of Barrow Island: before you can say whether something isn't there, you need to be able to say what is.

Over the course of these surveys, a sizeable collection of material has been accumulated from an area that had previously been only sporadically sampled. Over two dozen taxonomic experts were consulted in the process of identifying this material, a lot of which represented species potentially new to science. And so, some time in 2012, we asked the people who had been involved with the project if they would like to contribute to a collection of papers on Barrow Island invertebrates. The response was mostly positive, and The Terrestrial Invertebrate Fauna of Barrow Island, Western Australia was released to the world a couple of weeks ago.

We're very pleased with how it turned out. Some of the contributors provided overviews of their taxon of interest; others provided descriptions of new species. Authors came from both the academic and private sectors, and we're grateful to everyone who put time and effort into answering our calls. In the end, we had 22 chapters on hand, including material on animals from arachnids to isopods to ants, and 25 new species: one snail, two spiders, a silverfish and 21 flies. Not all of these new species were from Barrow Island alone: the chapter on Dolichopodidae (long-legged flies) by Dan Bickel represents a review of the fauna of the entire Pilbara region.

The book is available for purchase from the Western Australian Museum, but I've noticed that their site doesn't provide an article listing. Therefore, I'm including one below, with the abstracts for each article. Contact details for the corresponding authors have been included as hyperlinks, if you want to ask them about their articles. And again, thank you to everyone involved.

The camaenid snail Rhagada barrowensis. The identity of Barrow Island's common Rhagada species has been subject to a bit of confusion over the years; Johnson et al. describe it as a new species in this book.

Dorian Moro and Russell Lagdon, pp. 1-8.
History and environment of Barrow Island

Barrow Island represents a unique island ecosystem off north-western Australia. It has ecological affinities to the Cape Range region of the Australian mainland, and it also supports an oil and gas resource industry. The island hosts a long-unburnt vegetation complex, and a diverse community of vertebrate and invertebrate fauna occupy the disturbed and undisturbed habitats of the island. In the absence of non-indigenous predators or herbivores, without extensive land clearing, and with an instituted level of island quarantine, these environmental values have persisted to make Barrow Island an important environmental asset for Australia, and an example where island ecology functions in the presence of resource extraction. To date, almost 2,800 species of terrestrial and subterranean species have been consistently recorded from Barrow Island. These include 378 native plant species, 13 mammal species (including two species of bats), at least 119 species of terrestrial and migratory birds, 43 species of terrestrial reptiles, one species of frog, three subterranean vertebrates, at least 34 species of subterranean invertebrates, and the most speciose of all, over 2,200 terrestrial invertebrates.

Russell Lagdon and Dorian Moro, pp. 9-11.
The Gorgon gas development and its environmental commitments

Chevron has made an important contribution to our knowledge and understanding of the Barrow Island flora and fauna, and to the Australian economy. This knowledge has been primarily founded from the investigations and commitments of joint venture partners associated with the environmental impact assessment for the Gorgon Gas Development. The Gorgon Gas Development is one of the world’s largest natural gas projects and the largest single natural gas project in Australia’s history. Development has been balanced between energy needs and environmental management. Through plans, procedures, programs and research, Chevron Australia and its joint venture participants have established a benchmark for environmental management of this important island reserve. Furthermore, the Gorgon Joint Ventures have contributed to one of the largest biodiversity offset and Net Conservation Benefit programs in Western Australia.

Jonathan D. Majer, Shae K. Callan, Karl Edwards, Nihara R. Gunawardene and Christopher K. Taylor, pp. 13-112.
Baseline survey of the terrestrial invertebrate fauna of Barrow Island

Barrow Island is Western Australia’s second largest offshore island and its flora and fauna have been able to evolve without major human disturbances. Chevron Australia Pty Ltd and its Joint Venture Participants made an application to construct a plant to liquefy natural gas on the island in 2001. One of the conditions under which approval was granted was the implementation of a rigorous biosecurity effort to ensure that no non-indigenous species (NIS) are introduced or allowed to establish on the island. To fulfil this condition it was first necessary to characterise what was already present on the island. A series of surveys have been performed using a purpose-designed sampling protocol in order to provide baseline data on the existing terrestrial invertebrates on Barrow Island. A total of 1,873 morphospecies were sampled but subsequent surveys and taxonomic developments have increased the count to 2,397. This compares with an estimated species richness of 2,481 terrestrial invertebrate species on the island. Composition of the fauna varied considerably between the wet and dry seasons and between years, even when samples were taken during the same month. Composition also varied with distance from the coast, which may be associated with soil type and vegetation association. Twenty five non-indigenous species and seven putative non-indigenous species have been found, all of which are believed to have been present prior to commencement of the Gorgon Gas Development project.

Peter Whittle, Frith Jarrad and Kerrie Mengersen, pp. 113-130.
Design of the quarantine surveillance for non-indigenous species of invertebrates on Barrow Island

The Ministerial conditions for regulatory approval for the Gorgon gas project on Barrow Island included a quarantine surveillance program having detection power of 0.8 for non-indigenous species of terrestrial invertebrates, vertebrates and plants. No method was available for design of such a program, so we developed a new method and designed surveillance systems that were implemented successfully in 2010−11 for the first of four years over the construction period. Here we describe the method and outline the invertebrate surveillance system, after the experience of the first year. We discuss a set of issues that characterised the design problem, which we consider typical of many surveillance applications. We suggest that the method is broadly applicable for objective design of surveillance, for biosecurity and other settings.

Ken Walker, pp. 131-134.
Providing web based diagnostics for the Barrow Island baseline survey

During the years of 2005 to 2007, an extensive baseline study of the Barrow Island invertebrate fauna was conducted. This survey included more than 50 sample sites across the island and multiple collecting techniques were used at each site. Over 14,000 specimens were collected during this survey. Taxonomic specialist who examined this material nominated over 2,000 morphospecies of which about 300 could be placed to species rank. Having done all of this collecting and identification, the question then was how best to access and use this valuable resource. All of the specimens were stored in two institutions in Perth – several thousand kilometres south of Barrow Island. Manual access to these specimens was slow which hindered the decision making processes needed when a suspected non-indigenous species was found on the island. The decision was made to digitise the diagnostic characters for representative of each morphospecies. These images were to be made available through a website called PaDIL (Pests and Diseases Image Library). Each species was to have its own webpage containing at least 4 diagnostic images of each species and all of the species collection points to be displayed on an interactive Google Map. Species, as well as higher ranks, could be queried alone or against sample localities or against Indigenous or Non-Indigenous status. Individual species pages could be opened and comparative images tables could be pre-defined and presented or users could build their own comparative image tables in real time. The development of the Barrow Island PaDIL website made the results of the entire Baseline Study accessible to anyone with a web browser from anywhere with an internet connection. The Barrow Island PaDIL website is a major part of the Quarantine efforts of Chevron on Barrow Island.

Christopher K. Taylor, pp. 135-144.
Annotated bibliography for Barrow Island terrestrial invertebrates

A bibliography is provided of publications treating terrestrial invertebrates on Barrow Island. A brief overview is also given of natural history and invertebrate collections on Barrow Island.

Garth Humphreys, Jason Alexander, Mark S. Harvey and William F. Humphreys, pp. 145-158.
The subterranean fauna of Barrow Island, north-western Australia: 10 years on

Barrow Island, situated off the north-west Australian coast, is well recognised for its subterranean fauna values. Sampling for both stygobitic and troglobitic fauna has taken place on the island since 1991, and Humphreys (2001) summarised the then current state of knowledge of the island’s subterranean fauna. Sampling for impact assessment purposes on the island over the past decade has substantially increased the recorded species richness of Barrow Island. The number of documented stygal taxa has more than doubled since 2001, from 25 to 63 species now known. Troglobitic diversity has also substantially increased, with six species known in 2001 and 19 troglobitic taxa known today. The total recorded subterranean species richness for Barrow Island at this time stands at 82 species. It is likely that considerably more species remain to be recorded, as even the additional surveys of the past decade leave many areas of the island unsampled.
The distributions and minimum area of occupancy for many species known from Barrow Island in 2001 have also been significantly expanded by the sampling efforts of the last decade. This includes specially protected species listed under State and Commonwealth Government legislation. The available data suggest the fauna of the island may number in the hundreds of species, many of which are endemic, confirming its status as internationally significant for subterranean biota.

Michael S. Johnson, Sean Stankowski, Corey S. Whisson, Roy J. Teale and Zoë R. Hamilton, pp. 159-171.
Camaenid land snails on Barrow Island: distributions, molecular phylogenetics and taxonomic revision

Three species of camaenid land snails occur on Barrow Island: Quistrachia barrowensis and two previously unassigned species of Rhagada. Based on morphological re-evaluation and analysis of sequences of the mitochondrial gene COI, we have revised the taxonomy of these species, providing a clearer understanding of their geographic distributions and origins. The supposed Barrow Island endemic Q. barrowensis is synonymous with Q. montebelloensis from the Montebello and Lowendal Islands. The small species of Rhagada, confined to the northern tip of Barrow Island, is conspecific with R. plicata, whose distribution also includes the Montebellos and the Lowendals. The large species of Rhagada is described here as R. barrowensis sp. nov., known only from Barrow Island and adjacent Pascoe Island. The three camaenids represent deeply divergent lineages with different geographic origins, indicating that the local diversity on Barrow Island has come about through a complex history. With maximum geographic spans of only 22 to 70 km, the short-range endemism of these species highlights the conservation significance of Barrow Island.

Volker W. Framenau and Anna E. Leung, pp. 173-184.
Costacosa, a new genus of wolf spider (Araneae, Lycosidae) from coastal north-west Western Australia

A new genus of wolf spider (family Lycosidae Sundevall, 1833), Costacosa gen. nov. is described from north-west Western Australia to include C. torbjorni sp. nov. (type species) and C. dondalei sp. nov. The genus belongs to the subfamily Lycosinae Sundevall, 1833 and differs from all other Australian genera in this subfamily with similar somatic morphology, in particular Venator Hogg, 1900 and Knoelle Framenau, 2006, mainly in genitalic characters. The tegular apophysis of the male pedipalp has a pronounced ventral spur, a distinct ventral edge of species-specific shape and serrations along its apical edge. The female epigyne has an elongated triangular atrium and the medium septum is longer than the posterior transverse part. Costacosa are medium-sized wolf spiders of overall brown colouration and with broad light median and sublateral bands on the carapace and a black patch in the frontal two-thirds of the venter. Costacosa torbjorni is the most commonly recorded wolf spider on Barrow Island, from where currently seven species of Lycosidae are known.

Simon Judd and Giulia Perina, pp. 185-207.
An illustrated key to the morphospecies of terrestrial isopods (Crustacea: Oniscidea) of Barrow Island, Western Australia

This paper presents an illustrated key to eighteen morphospecies of terrestrial isopods from Barrow Island with a brief summary regarding their currently known distribution and potential endemicity to the island. Six described species are recorded, Ligia exotica (family Ligiidae), Alloniscus pallidulus (Alloniscidae), Laevophiloscia yalgooensis (Philosciidae), Porcellionides pruinosus (Porcellionidae), Barrowdillo pseudopyrgoniscus, Buddelundia hirsuta (both Armadillidae), but the identifications of most need to be confirmed following genus-level revisions and examination of type- or topotypical material. The key includes twelve undescribed species and at least two undescribed genera from the family Armadillidae, one of which is apparently restricted to Barrow Island. Although there is still considerable taxonomic work required to evaluate distributions, it appears that at least six of the eighteen species are potential short-range endemics (SRE).

Catherine A. Car, Megan Short, Cuong Huynh and Mark S. Harvey, pp. 209-219.
The millipedes of Barrow Island, Western Australia (Diplopoda)

Six species of millipedes are recorded from Barrow Island, including three species of pin-cushion millipedes of the order Polyxenida, Lophoturus madecassus (Marquet and Condé, 1950) (Lophoproctidae), Unixenus mjoebergi (Verhoeff, 1924) (Polyxenidae) and Phryssonotus novaehollandiae (Silvestri, 1923) (Synxenidae), a single species of the order Spirobolida, Speleostrophus nesiotes Hoffman, 1994 (Trigoniulidae), and two species of the order Polydesmida, Boreohesperus dubitalis Car and Harvey, 2013 (Paradoxosomatidae) and one species of the family Haplodesmidae (genus and species indet.). Lophoturus madecassus is circum-tropical in distribution, Unixenus mjoebergi and Phryssonotus novaehollandiae are found also on mainland Australia, but the other three species are endemic to the island. Speleostrophus nesiotes is a highly modified troglobiotic species, currently listed as threatened by the Western Australian government. It is unclear at present whether the haplodesmid specimen is a troglobite.

Penelope Greenslade, pp. 221-228.
Composition of Barrow Island collembolan fauna: analysis of genera

Collembola have been collected from Barrow Island for the first time; a maximum of seventy one species were detected, of which a high proportion are undescribed. Only four non-indigenous species (NIS) species have been collected, three in very small numbers but one was a large population introduced to the island in lengths of timber which were subsequently sent off the island. Despite few of the species being described, most have been collected before and endemism is low. One new genus record for Australia, Calx, was found. The presence of a species of Temeritas is unusual in that the males showed strong sexual dimorphism, and a species of Acanthocyrtus that lacked any pigment was collected in reasonable numbers. Collections from bore holes were rich in species. Five species were recorded only from bore holes and may be island endemics. The intertidal fauna was also rich in species with 14 found, all restricted to this habitat. Soil fauna density of Collembola was found to be high, with a mean average potential density of nearly 47,000/m². A proportion of the terrestrial Collembola fauna is active under all weather conditions but other species are only active after rain. In general, the terrestrial fauna shows a dominance of the families Isotomidae and Bourletiellidae, which is typical for the wet/dry tropics where trees are absent.

Graeme Smith, pp. 229-240.
A new species of Heterolepisma from Barrow Island (Zygentoma: Lepismatidae)

The silverfish fauna of Barrow Island is discussed and Heterolepisma parva sp.nov. is described from extensive material collected mostly in pitfall traps or Winkler sac leaf litter samples.

David T. Jones, pp. 241-244.
The termites of Barrow Island, Western Australia

Forty years ago D. H. Perry, the renowned termite expert, published a checklist of 18 species that he had collected on Barrow Island. That checklist is now updated with the results of a recent invertebrate survey of the island, and a literature search for additional records. The updated list now runs to 27 species, all of which appear to be indigenous to the island.

Christopher K. Taylor, pp. 245-252.
The genus Lithoseopsis (Psocodea: Amphientomidae) in the Western Australian fauna, with description of the male of Lithoseopsis humphreysi from Barrow Island

The Australian Amphientomidae species Seopsis incisa Smithers, 1989 and S. humphreysi New, 1994 are transferred to the genus Lithoseopsis Mockford, 1993 as L. incisa new combination and L. humphreysi new combination, as a result of the discovery of speciens of L. humphreysi from Barrow Island, Western Australia. The male of L. humphreysi is described for the first time, and both macropterous and brachypterous individuals are described. The genus Lithoseopsis was previously known from North America only, and the addition of the Western Australian species significantly increases its range. A key is provided to the genera of Amphientomidae.

David Gopurenko, Murray Fletcher, Holger Löcker and Andrew Mitchell, pp. 253-285.
Morphological and DNA barcode species identifications of leafhoppers, planthoppers and treehoppers (Hemiptera: Auchenorrhyncha) at Barrow Island

The hemipteran suborder Auchenorrhyncha comprises a rich assemblage of plant feeding species, many of which are widespread in distribution and act as vectors of viral and fungal diseases affecting plants. Species level identifications in this group generally are possible only by examination of male specimens; prior DNA barcode analyses of a limited range of Auchenorrhyncha indicate that this approach may provide an expedient means to identify species within this diverse group. In this study we explored the utility of DNA barcoding for identification of a wider range of Auchenorrhyncha species than has been examined previously. Diverse fulgoroid (planthopper) and membracoid (leafhopper and allies) Auchenorrhyncha were sampled from Barrow Island, Western Australia, and identified to the least inclusive taxonomic units using morphology. DNA barcodes from 546 adult specimens were obtained and analysed using a General mixed Yule – Coalescent (GMYC) modelling approach to genetically delimit putative species, as a comparison to the morphospecies identifications. Additional DNA barcodes (N = 106) were obtained from nymphs and these were compared to adult DNA barcodes to identify species present among immature specimens.
Among adult specimens, 73 species were congruently delimited by morphology and genetic analyses when modelled using a single threshold GMYC. Congruence between morphological and molecular species assignments was greatly reduced when the Yule – Coalescent transition was allowed to vary across genetic lineages. In a separate DNA barcode analysis of all specimens using neighbour joining distance metrics, nymphs and physically degraded specimens were in most cases genetically linked to adult conspecifics. Ten genetic clades detected among the nymphs were not observed among adults and did not match pre-existing sequence accessions in GenBank or DNA barcode records in BOLD.
Of the 73 adult Auchenorrhyncha species congruently identified by DNA barcoding and morphology, most were Cicadellidae (N = 53 morphospecies), the remaining 20 morphospecies were sparsely representative of ten other families. Formal identifications to species level were available for only 36% of these 73 morphospecies, owing mainly to an absence of diagnostic male specimens within many of the delimited species. Indeterminate species detected among adults and nymphs are designated with interim species codes.
The work presented here demonstrates that DNA barcoding is likely to be a powerful investigative tool for identifying and understanding species limits in the Auchenorrhyncha, particularly if it is used within an integrative taxonomic framework.

Laurence A. Mound, pp. 287-290.
Thysanoptera (Insecta) of Barrow Island, Western Australia

Almost 50 species of the insect order Thysanoptera are here listed from Barrow Island, Western Australia, of which several are known only from this island. This cannot be interpreted as indicating that any species is endemic to the island, because almost nothing is known of the Thysanoptera fauna of the nearby mainland.

Daniel J. Bickel, pp. 291-348.
The family Dolichopodidae (Diptera) of the Pilbara region, Western Australia in its Australasian biogeographic context, with the description of 19 new species

The Dolichopodidae (Diptera) of the Pilbara Region (here also including Barrow Island and Cape Range), Western Australia are described, keyed and illustrated. The fauna comprises 41 species, including three with generic names only, being represented by females or badly damaged males. The following 19 species are newly described: Pseudoparentia canalicula sp. nov., Pseudoparentia niharae sp. nov., Paraclius manglar sp. nov., Medetera junensis sp. nov., Corindia gascoynensis sp. nov., Thinophilus eboricoxa sp. nov., Thinophilus yarraloola sp. nov., Chaetogonopteron capricorne sp. nov., Chaetogonopteron vexillum sp. nov., Sympycnus colliepa sp. nov., Sympycnus lacrimulus sp. nov., Sympycnus pistillus sp. nov., Sympycnus weano sp. nov., Sympycnus ephydroides sp. nov., Sympycnus hamulitarsus sp. nov., Diaphorus karijini sp. nov., Diaphorus garnetensis sp. nov., Chrysotus austrotropicus sp. nov. and Chrysotus pilbarensis sp. nov. Paraclius obtusus Hardy, 1939 is regarded as a new senior synonym of Paraclius albodivisus Parent, 1941, syn. nov. The Pilbara fauna is treated in the context of the wider Australian fauna, and many extralimital records are included. Many Pilbara species are found across tropical northern Australia, and sometimes into adjacent Melanesia. However, some species have a trans-continental distribution south of the monsoonal belt and also occur in central Northern Territory and subtropical interior Queensland suggesting a biogeographic track that now comprises favorable relictual habitats in a largely arid region. The Millstream site along the Fortescue River is particularly rich in species, and it is the only known locality of the isolated monotypic genus Pilbara Bickel.

David K. Yeates and Stefanie K. Oberprieler, pp. 349-354.
Two new species of the Australian bee fly genus Comptosia (Diptera: Bombyliidae) from Barrow Island, Western Australia

Two new species of the bee fly genus Comptosia Macquart from Western Australia, C. barrowensis and C. karijinii, are described.

Nicholas B. Stevens, Syngeon M. Rodman, Tamara C. O’Keeffe and David A. Jasper, pp. 355-374.
The use of the biodiverse parasitoid Hymenoptera (Insecta) to assess arthropod diversity associated with topsoil stockpiled for future rehabilitation purposes on Barrow Island, Western Australia

This paper examines the species richness and abundance of the Hymenoptera parasitoid assemblage and assesses their potential to provide an indication of the arthropod diversity present in topsoil stockpiles as part of the Topsoil Management Program for Chevron Australia Pty Ltd Barrow Island Gorgon Project. Fifty six emergence trap samples were collected over a two year period (2011 and 2012) from six topsoil stockpiles and neighbouring undisturbed reference sites. An additional reference site that was close to the original source of the topsoil on Barrow Island was also sampled. A total of 14,538 arthropod specimens, representing 22 orders, were collected. A rich and diverse hymenopteran parasitoid assemblage was collected with 579 individuals, representing 155 species from 22 families. The abundance and species richness of parasitoid wasps had a strong positive linear relationship with the abundance of potential host arthropod orders which were found to be higher in stockpile sites compared to their respective neighbouring reference site. The species richness and abundance of new parasitoid wasp species yielded from the relatively small sample area indicates that there are many species on Barrow Island that still remain to be discovered. This study has provided an initial assessment of whether the hymenoptera parasitoid assemblage can give an indication of arthropod diversity. However, further work would still be required to more robustly establish the use of the hymenoptera parasitoid assemblage as indicators of arthropod diversity.

B. E. Heterick, pp. 375-404.
A taxonomic overview and key to the ants of Barrow Island, Western Australia

This work characterises the ant (Hymenoptera: Formicidae) fauna of Barrow Island, Western Australia, and provides a key to the workers and several unique reproductives of the 117 species recorded from the island thus far. In all, 11 of the 13 subfamilies of Western Australian ants have been recorded from Barrow Island, but Myrmeciinae and Heteroponerinae are absent. At a generic level, the fauna of the island is less rich, holding 36 of the 71 genera currently known from Western Australia. The ant fauna is characteristic of the Eremaean Botanical Province of the Pilbara, rather than that of the Carnarvon Basin from which Barrow Island is geologically derived. Ninety-three ant species (79.5% of the total on Barrow Island) are shared with the ant fauna of the Pilbara region on the adjoining mainland, but only 52 species (44.4% of the total) are shared with the ant fauna of the Carnarvon Basin. The island is very rich in unspecialised and thermophilic ant species. Five such genera, i.e., Iridomyrmex (14 spp.), Monomorium (13 spp.), Polyrhachis (12 spp.), Melophorus (10 spp.), and Camponotus (nine spp.) make up almost 50% (i.e., 49.6%) of the island’s ant fauna. Very few ants appear to be endemic to Barrow Island. The relative proportions of the two major subfamilies (Formicinae and Myrmicinae, together comprising 61.5% of the total ant richness) are similar to the proportions found in the South-west Botanical Division for these two subfamilies (i.e., 65.9%), with Barrow Island having a slightly lower ratio of formicines to myrmicines than is found in the south-west of the state. An estimate of the total number of ant species likely to occur on Barrow Island, using the Estimate-S program (Colwell 2009), suggests that a maximum of fourteen additional species may be as yet unrecorded.

Jonathan D. Majer, Nihara R. Gunawardene, Christopher K. Taylor and Mark S. Harvey, pp. 405-406.
A last word

The work reported on in this volume is the culmination of nine years of data gathering stemming from the original baseline surveys on Barrow Island. Not surprisingly, this has resulted in one of the most comprehensive terrestrial invertebrate surveys ever performed on an offshore island on this continent. There are other substantial surveys, but these have generally focussed on specific taxonomic groups, rather than the whole spread reported here.

Trichadenotecnum: Six Spots and Spiny Terminalia

Trichadenotecnum sexpunctatum, photographed by Brian Valentine.

The animal in the photo above is a typical representative of Trichadenotecnum, a diverse genus of the barklice. About 200 species have been assigned to this genus from almost all the major biogeographic regions of the world except Australia (Yoshizawa et al. 2007); of the two species recorded from Australia and assigned to this genus, one (Ptycta enderleini) has recently been excluded from Trichadenotecnum, and the other (Trichadenotecnum circularoides) is probably a recent introduction from the Americas (Yoshizawa & Smithers 2006). Though long regarded as suspectly heterogenous, the genus has been extensively reviewed in recent years, particularly by Kazunori Yoshizawa of Hokkaido University and associates (Yoshizawa 2001, 2004; Yoshizawa et al. 2008). Members of Trichadenotecnum are characterised by a distinctive array of wing markings, visible in the above photo. Note, in particular, the series of six submarginal spots forming a U-shape towards the end of each wing (though, confusingly, these characteristic markings can become difficult to distinguish in species in which the wings are more heavily spotted overall). The genus is also distinguished by certain features of the male terminalia (or, in layman's terms, the bum) with a number of processes developed on the hypandrium, the posteriormost segment of the underside of the abdomen that covers the phallosome, the intromittent organ in Psocoptera. These processes vary in development between species, and often themselves bear arrays of small spines or teeth. A number of species of Trichadenotecnum also have the terminalia assymmetrically developed, with the left and right lobes of the hypandrium (for instance) differently sized and/or shaped, though the functional significance of this arrangement (if any) remains unknown.

Various views of the terminalia of Trichadenotecnum alexanderae, from Yoshizawa (2001). In life, the phallosome is contained within the underside of the terminalia.

The variability of the terminalia between species of Trichadenotecnum makes them a rich source of characters for use in taxonomy. The problem with this, of course, is that you need adult males, and that isn't always easy. Particularly in a group like Psocoptera, which seem to show a particular tendency for parthenogenesis. A number of Trichadenotecnum species are not, as yet, known to produce males, including the aforementioned T. circularoides, necessitating identifiers to fall back largely on wing markings. Trichadenotecnum circularoides has been recorded from Angola, east Asia, Australia, North America and Brazil; the distribution of closely related species suggests that the last locality represents its original homeland with human dispersal carrying it elsewhere (Yoshizawa et al. 2008). The New World species of Trichadenotecnum appear to fall within a small number of clades: one including T. circularoides is the sister group to other members of the genus, while the T. alexanderae species group is Holarctic in distribution. The majority of New World species, however, form a single lineage referred to as the 'bulky clade' by Yoshizawa et al. (2008). Members of the bulky clade have a movable median tongue on the hypandrium that bears a dorsal covering of denticles or spines. Yoshizawa et al. (2008) suggested that, as the bulky clade was nested amongst a number of Old World lineages, this group may represent a relatively recent invasion of the Americas, probably by way of the Bering Strait.

Nymphs of Trichadenotecnum possess glandular hairs to which frass and pieces of lichen become attached, providing them with camouflage. This individual was photographed by Charley Eiseman.
REFERENCES

Yoshizawa, K. 2001. A systematic revision of Japanese Trichadenotecnum Enderlein (Psocodea: ‘Psocoptera’: Psocidae: Ptyctini), with redefinition and subdivision of the genus. Invertebrate Taxonomy 15: 159-204.

Yoshizawa, K. 2004. Molecular phylogeny of major lineages of Trichadenotecnum and a review of diagnostic morphological characters (Psocoptera: Psocidae). Systematic Entomology 29: 383-394.

Yoshizawa, K., A. N. García Aldrete & E. L. Mockford. 2008. Systematics and biogeography of the New World species of Trichadenotecnum Enderlein (Insecta: Psocodea: ‘Psocoptera’: Psocidae). Zoological Journal of the Linnean Society 153: 651-723.

Yoshizawa, K., C. Lienhard & V. K. Thapa. 2007. Systematic study of the genus Trichadenotecnum in Nepal. Insecta Matsumurana, new series 63: 1-33.

Yoshizawa, K., & C. N. Smithers. 2006. Systematic position of Trichadenotecnum enderleini (Roesler) (Psocodea: “Psocoptera”: Psocidae). Records of the Australian Museum 58: 411-415.

The Psocoptera of Barrow Island

Courtenay Smithers, courtesy of the Sydney Morning Herald.

Gunawardene, N. R., C. K. Taylor & J. D. Majer. 2012. Revisiting the Psocoptera (Insecta) of Barrow Island, Western Australia. Australian Entomologist 39 (4): 253-260.

Our lab has just recently added to its publication list with the above title, which is part of a special issue of the Australian Entomologist printed in memory of the late Courtenay Smithers, who passed away last year. For many years, Courtenay was one of Australia's leading entomologists, particularly for those unfairly overlooked animals the bark-lice (non-parasitic Psocodea). An obituary for him can be found here.

We wanted to include this paper as a tribute to Courtenay, as it basically presents some identification work that he had done for us in the last few years. As some of you already know, we've been working for the last few years on surveying the terrestrial invertebrates of Barrow Island, off the north-west coast of Australia. Courtenay had first surveyed the bark-lice of Barrow back in 1982, when he collected only five species of Psocodea all up, including the cosmopolitan synanthrope Liposcelis entomophila (Smithers 1984). Because Barrow Island is a very arid habitat, with little to no standing fresh water, Courtenay felt that "The small size of the fauna is probably a reality not an illusion".

A cute litte critter from our collection that still only goes by the name of 'Pteroxanium sp. A'.

As it turns out, he was wrong. At least 26 species of Psocodea have been found on Barrow so far (the paper says 25, but we've had at least one more turn up since it was accepted for publication). Most of these have currently only been identified as morphospecies: identification of bark-lice is often a difficult task, and many Australian species probably remain undescribed (as an example, Courtenay's 1996 tally of the total described Australian Psocodea for the Zoological Catalogue of Australia includes less species of Liposcelididae than have been collected on Barrow Island alone). Three of the recorded species are synanthropes collected in buildings on the island; as far as we know, these species are not found in unmodified habitat. One of these, Dorypteryx domestica, was particularly interesting to me as it had not been recorded previously from Australia (and was my first real success at identifying a psocodean right down to species level, hurrah!), though Tim New (Australia's remaining bark-louse expert) informed us that its presence has always been expected. I have to say, while bark-lice in general are among the cutest of all insects, but the little jumping Dorypteryx really amps the cuteness right up there.

And here it is! (Photo from Gunawardene er al. 2012.) Dorypteryx domestica is probably found worldwide, but records are scattered because of its unassuming nature.

Unfortunately, Courtenay's passing highlights that a large proportion of taxonomic expertise currently resides in the minds of retired individuals (of the experts who have made identifications of material from the Barrow Island project, nearly a third were either retired or amateur taxonomists working in their spare time). There is no shortage of material out there, but we still need the people to tell us what it is.

Suspiciously posed-looking photo, used in Gunawardene et al. (2012), of yours truly supposedly demonstrating an insect collection method.

REFERENCES

Smithers, C. N. 1984. The Psocoptera of Barrow and Boodie Islands, Western Australia. Entomologica Scandinavica 15: 215-226.

Smithers, C. N. 1996. Psocoptera. In: Wells, A. (ed.) Zoological Catalogue of Australia. Psocoptera, Phthiraptera, Thysanoptera pp. 1–79. CSIRO Publishing: Melbourne.

Barklice and Booklice and Such

Psocoptera is arguably the least deservedly obscure of the obscure insect orders. They're not uncommon - there's a reasonable chance that you'll have seen one in your life. You probably squashed it without giving much thought to what it was. And yet, so obscure is this order of insects that there isn't even a good vernacular name for the group. Psocoptera are minute insects (usually only a couple of millimetres long) that generally live among bark and litter, feeding on fungi. Some wingless species can be found in houses (where you might have seen one) and feed on such delicacies as dust or the glue used in book bindings, leading to their being known as booklice. The tree-living forms are sometimes referred to as barklice in comparison to booklice. Most entomologists that I know simply refer to the group as psocids, and that's exactly what I'm going to do.

Technically speaking, 'Psocoptera' is a paraphyletic group. The Phthiraptera, the true lice*, are derived from within the psocids. At the moment, things seem to be going through a transitional phase, with many authors dropping the paraphyletic 'Psocoptera' for the name Psocodea, which refers to the total group of psocids and lice. The 'Psocoptera' are divided into three suborders, the Trogiomorpha, Psocomorpha and Troctomorpha, the Phthiraptera being properly speaking a subgroup of the last. A representative of the second group, the psocomorph Blaste (photo from TOLWeb), can be seen at the top of this post, and it's the Psocomorpha that I'm looking at today.

*And holders of what is probably the worst insect order name of all to pronounce.



With over 3500 species, the Psocomorpha are generally regarded as the largest of the psocid suborders, though the Troctomorpha could give them a run for their money once the Phthiraptera are taken into account. We should probably be careful about making definite statements about this - because of their neglected nature, new species and sometimes even families of psocids continue to appear in the literature at a respectable rate. At the moment, though, it is a psocomorph that holds the honour of being probably the only invertebrate to get its picture plastered over Tetrapod Zoology, due to the nomenclatural issues that have arisen from the similarity in names of the psocid Caecilius and the amphibian Caecilia. The photo above is the one featured in Tet Zoo, and shows an identified member of the Caeciliusidae.



Most Psocomorpha are dwellers on bark or rocks. One group, the Caeciliusoidea, inhabits living foliage. Adults may be winged or wingless - many species have both forms. Many psocids cluster as nymphs - the photo above (from here*) shows one such congregation - and spin protective webs, but this is taken to the extreme in the genus Archipsocus. Archipsocus species form large colonies, and may build webs large enough to obscure tree-trunks, as can be seen in the picture below (from here). As with Embioptera, these colonies appear to be conglomerations of convenience, and there is no real social behaviour. Like aphids, Archipsocus may go through multiple generations in a summer, and the colony will contain individuals at all stages of development, both winged and wingless forms (Mockford, 1957). Once winter arrives, the colony breaks down and disperses, the survivors diapausing until the spring when they will start new colonies.

*This page also records a fantastic common name for psocids - "bark cattle", apparently because the nymphs move like a herd when disturbed.



Molecular and morphological data are mostly in agreement that the Psocomorpha can mostly be divided between four infraorders, the Psocetae, Homilopsocidea, Epipsocetae and Caeciliusetae (Johnson & Mockford, 2003; Yoshizawa, 2002). Both studies also agreed in placing Archipsocus outside these groups, as the basalmost member of the Psocomorpha. Unfortunately, beyond the bare morphology, information about most psocid groups seems to be few and far between, and there is a great deal about the order that we have yet to know.

REFERENCES

Johnson, K. P., & E. L. Mockford. 2003. Molecular systematics of Psocomorpha (Psocoptera). Systematic Entomology 28: 409-416.

Mockford, E. L. 1957. Life history studies on some Florida insects of the genus Archipsocus (Psocoptera). Bulletin of the Florida State Museum - Biological Sciences 1 (5): 254-274.

Yoshizawa, K. 2002. Phylogeny and higher classification of suborder Psocomorpha (Insecta: Psocodea: ‘Psocoptera’). Zoological Journal of the Linnean Society 136: 371-400.