Australian gall-inducing scale insects on Eucalyptus: revision of Opisthoscelis Schrader (Coccoidea, Eriococcidae) and descriptions of a new genus and nine new species

We revise the genus Opisthoscelis Schrader, and erect the genus Tanyscelisgen. n. with Opisthoscelis pisiformis Froggatt as its type species. Species of both genera induce sexually dimorphic galls on Eucalyptus (Myrtaceae) in Australia, with Opisthoscelis subrotunda Schrader also in Papua New Guinea. We synonymise the following taxa (junior synonym with senior synonym): Opisthoscelis fibularis Froggatt, syn. n. with Opisthoscelis spinosa Froggatt; Opisthoscelis recurva Froggatt, syn. n. with Opisthoscelis maculata Froggatt; Opisthoscelis globosa Froggatt, syn. n. (= Opisthoscelis ruebsaameni Lindinger) with Opisthoscelis convexa Froggatt; and Opisthoscelis mammularis Froggatt, syn. n. with Opisthoscelis verrucula Froggatt. We transfer seven Opisthoscelis species to Tanyscelis as Tanyscelis conica (Fuller), comb. n., Tanyscelis convexa (Froggatt), comb. n., Tanyscelis maculata (Froggatt), comb. n., Tanyscelis maskelli (Froggatt), comb. n., Tanyscelis pisiformis (Froggatt), comb. n., Tanyscelis spinosa (Froggatt), comb. n., and Tanyscelis verrucula (Froggatt), comb. n. We redescribe and illustrate the adult female of each named species of Opisthoscelis for which the type material is known, as well as the first-instar nymph of the type species of Opisthoscelis (Opisthoscelis subrotunda) and Tanyscelis (Opisthoscelis pisiformis). We describe four new species of Opisthoscelis: Opisthoscelis beardsleyi Hardy & Gullan, sp. n., Opisthoscelis thurgoona Hardy & Gullan, sp. n., Opisthoscelis tuberculataHardy & Gullan, sp. n., and Opisthoscelis ungulifinis Hardy & Gullan, sp. n., and five new species of Tanyscelis: Tanyscelis grallator Hardy & Gullan, sp. n., Tanuscelis megagibba Hardy & Gullan, sp. n., Tanyscelis mollicornuta Hardy & Gullan, sp. n., Tanyscelis tripocula Hardy & Gullan, sp. n., and Tanyscelis villosigibba Hardy & Gullan, sp. n. We designate lectotypes for Opisthoscelis convexa, Opisthoscelis fibularis, Opisthoscelis globosa Froggatt, Opisthoscelis maculata, Opisthoscelis mammularis, Opisthoscelis maskelli, Opisthoscelis pisiformis, Opisthoscelis recurva, Opisthoscelis serrata, Opisthoscelis spinosa, and Opisthoscelis verrucula. As a result of our taxonomic revision, Opisthoscelis has six species and Tanyscelis has 12 species. We describe the galls of females for all 18 species and galls of males for 10 species of Opisthoscelis and Tanyscelis, and provide photographs of the galls for most species. A key to the adult females of the species of both genera is included.     

Phylogeny of Australian Coptotermes (Isoptera: Rhinotermitidae) species inferred from mitochondrial COII sequences

Six Australian species of Coptotermes are traditionally recognized, but recent cuticular hydrocarbon studies suggest that some of these may represent more than one species. An understanding of the phylogenetic diversity of Australian Coptotermes, particularly the pest species, is likely to be important for the improvement of termite management strategies. A study of phylogenetic relationships among species of this genus was performed, based on the mitochondrial cytochrome oxidase (COII) gene, comparing the data with recent data from Asian species. Representatives of the species C. lacteus (Froggatt), C. frenchi Hill and C. michaelseni Silvestri were each found to form closely related monophyletic groups, however representatives of C. acinaciformis (Froggatt) were not. For C. acinaciformis, representatives from northern mound-building populations were found to form a distinct group to southern, tree-nesting forms. Among southern C. acinaciformis, two Western Australian representatives were found to be divergent from other populations. The results suggest that C. acinaciformis probably represents a complex of species rather than one, as has been suggested previously. One unidentified Coptotermes sp. taxon from Melbourne was found to be divergent from other taxa. Notably, some Australian species were more closely related to Asian species than other Australian species.     

THESIS ABSTRACT

Geographical distribution, seasonal abundance and habitat preference of fruit fly species (Diptera: Tephritidae) in South-East Queensland, with special reference to Bactrocera tryoni (Froggatt) and Bactrocera neohumeralis (Hardy) . S. Raghu.     

Taxonomy of Heterotermes (Isoptera: Rhinotermitidae) in south-eastern Australia: cuticular hydrocarbons of workers, and soldier and alate morphology

Abstract Heterotermes ferox (Froggatt) auctt. is shown to be a complex of species, each characterized by the composition of cuticular hydrocarbons in the worker, by canonical variate scores calculated from eight measurements of the soldier, and by small differences in the size and shape of the alate. H.ferox is redefined and two species are described, H.brevicatena sp.n. and H.longicatena sp.n.     

First record of the termite family Kalotermitidae (Blattodea: Termitoidae) in Korea

The drywood termite Kalotermitidae Froggatt, represented by Glyptotermes nakajimai Morimoto, is identified for the first time in Korea. They were collected in Yeoseo-do Island, the southern part of the Korean peninsula. A key to the termite species in Korea, habitus photographs of all castes, ecological information, and illustrations of diagnostic characters are provided.     

Insect, not plant, determines gall morphology in the Apiomorpha pharetrata species-group (Hemiptera: Coccoidea)

Abstract  Scale insects of the genus Apiomorpha Rübsaamen (Hemiptera: Coccoidea: Eriococcidae) induce sexually dimorphic galls on Eucalyptus, but the Apiomorpha pharetrata species-group is unusual in that nymphal males aggregate on the surface of the maternal gall where they induce a compound structure. Although originally described as distinct species on the basis of differences in gall morphology, A. pharetrata (Schrader) and A. thorntoni (Froggatt) were subsequently synonymised, primarily because the adult females of the two are morphologically indistinguishable. The two gall types of A. pharetrata sensu lato are allopatric and found on different host eucalypt species. To test the hypothesis that gall morphology may be determined by the host species on which the female feeds, we reared crawlers of field-collected females from both gall types on eucalypts in a glasshouse. Both gall types were induced on the same eucalypt species, with the gall type matching that of the maternal gall from which the crawlers had emerged. This indicates that it is the insect, not the host, that determines the gall morphology in these taxa. In addition, insects from the two gall types were different chromosomally and could be distinguished by one fixed allozyme difference. Thus, Brachyscelis (=  Apiomorpha ) thorntoni (Froggatt) revised status is removed from synonymy with Brachyscelis (=  Apiomorpha ) pharetrata (Schrader), and recognised as a distinct biological species. However, the extent of chromosomal variation among the populations of both A. pharetrata and A. thorntoni suggests that there may be further cryptic species present.     

REDUCTION IN TRAP CAPTURES OF FEMALE FRUIT FLIES (DIPTERA: TEPHRITIDAE) WHEN SYNTHETIC MALE LURES ARE ADDED

Abstract
Captures of females of 4 species of fruit flies on or in both coloured sticky traps and protein baited traps were reduced by synthetic male attractants. Captures of female Dacus tryoni (Froggatt), Dacus neohumeralis Hardy and Dacus cacuminatus (Hering) were reduced by both cuelure and methyl eugenol, and female Ceratitis capitata (Weidemann) by Capilure®. These effects were sufficient to overcome the normally attractive stimuli of colour and odours associated with yeast.     

A new family of Strepsiptera parasitizing fruit flies (Tephritidae) in Australia

ABSTRACT. A new family of Strepsiptera (Dipterophagidae) from Australia containing a new genus and species ( Dipterophagus daci ) is described and figured. This is the first species of Strepsiptera to be described as a parasite of Diptera. D.daci is recorded from ten fruit fly species, viz Dacus aquilonis (May), D.bellulus Drew, D.cacuminatus (Hering), D.decurtans (May), D.mayi Hardy, D.neohumeralis Hardy, D.peninsularis Drew, D.tenuifascia (May), D.tryoni (Froggatt) and Dacus sp. n. ( near musae ). Gregarious parasitism (two to six adult parasites per fly) is common.     

The development of «workers» and reproductives inMastotermes darwiniensis Froggatt (Isoptera)

Summary The five larval stages ofMastodermes darwiniensis Froggatt lead to the pseudergate «worker» stages, which can undergo stationary moults. The reproductives and soldiers develop from the pseudergates.

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Zusammenfassung Die fünf Larven-Entwicklungsstufen vonMastotermes darwiniensis Froggatt leiten zu pseudergaten «Arbeiter» Formen, welche feststehende Häutungen untergehen können. Die Sexuellen- und Soldatenformen entwickeln sich von den Pseudergaten.

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Résumé Les cinq états larvaires deMastotermes darwiniensis Froggatt aboutissent aux états ouvriers pseudergate, qui peuvent subir des mues «stationnaires». Les reproducteurs et les soldats se développent à partir des pseudergates.

     

Insect pests of eucalypts in California: implications of managing invasive species

For the first 150 years following their introduction, eucalypts planted in the California landscape were free of both insect pests and diseases. In the last 15 years, numerous herbivorous insect species have been introduced accidentally into the State and have caused significant damage to the trees. Several of these species, e.g. Phoracantha semipunctata (Fabricius), Phoracantha recurva Newman (Coleoptera: Cerambycidae) and Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae), have also been introduced into other parts of the world where eucalypts are grown, whereas others, e.g. Glycaspsis brimblecombei Moore (Hemiptera: Spondyliaspidae) and Eucalyptolyma maideni Froggatt (Hemiptera: Spondyliaspidae), are currently restricted to California and Australia. Research programmes have provided management solutions to individual pest problems, but as more pest species are introduced, these solutions must be integrated across broad geographic, horticultural, and economic scales, in a systems approach.     

The likely fate of hybrids of Bactrocera tryoni and Bactrocera neohumeralis

Bactrocera tryoni (Froggatt) and B. neohumeralis (Hardy) (Diptera: Tephritidae) are sympatric species which hybridise readily in the laboratory yet remain distinct in the field. B. tryoni mates only at dusk and B. neohumeralis mates only during the day, but hybrids can mate at both times. We investigated the inheritance of mating time in successively backcrossed hybrid stocks to establish whether mating with either species is more likely. The progeny of all backcrosses to B. tryoni mated only at dusk. The majority of the progeny of the first and a minority of the progeny of the second backcross to B. neohumeralis also mated at dusk, but the third successive B. neohumeralis backcross produced flies that mated only during the day. This trend towards dominance of the B. tryoni trait was also reflected in a diagnostic morphological character. We discuss the possible genetic background for these phenomena and propose that unidirectional gene flow might explain how the two species remain distinct in the face of natural hybridisation.     

The structure of calling songs in the cicada Pauropsalta annulata Goding and Froggatt (Hemiptera: Cicadidae): evidence of diverging populations?

Variation recorded within species is often taken to represent evidence for local and ongoing adaptation, but often without the interpopulation variation being subject to analysis across the geographic distribution occupied by the taxon. Here we investigate the rhythmic song structure across the range of three known song types in a variable cicada, Pauropsalta annulata Goding and Froggatt. Statistical analysis of the structure of songs across individuals reveals four discrete clusters that are demonstrated to be independent and stable across extensive geographic space in areas of allopatry and, generally, also into areas of sympatry. This suggests that P. annulata is a cryptic species complex. Unique combinations of plant species are linked with each of the clusters, suggesting that the different populations have independent plant associations. These findings are discussed in relation to similar case studies on cicadas and other organisms, with particular emphasis on the most appropriate approach to testing variation across populations, especially when it is thought to represent populations in the initial stages of evolutionary divergence.     

Genetic delineation of sibling species of the pest fruit fly Bactocera (Diptera: Tephritidae) using microsatellites

Using a large set of microsatellites, the genetic relationships between three closely related Australian fruit fly species, Bactrocera tryoni (Froggatt), B. neohumeralis (Hardy) and B. aquilonis(May) were investigated. Bactrocera tryoni and B. neohumeralis are sympatric, while B. aquilonisis allopatric to both. The sympatric species, B. tryoni and B. neohumeralis, were found to be genetically distinct. It is likely that despite differences in mating time between these two species, some gene flow still occurs. In contrast, the sibling species B. tryoni and B. aquilonis were found to be closely related, despite allopatry. The level of genetic divergence was similar to that found within eastern Australian populations of B. tryoni. Consideration of all available genetic data suggests that this similarity is not due to recent (i.e. within the last 30 years) displacement of B. aquilonis by B. tryoni from the B. aquilonis region (north-western Australia). Instead the data suggests that, at least in the areas sampled, asymmetrical hybridization may have occurred over a longer timescale.     

Impact of introduction of Bactrocera dorsalis (Diptera: Tephritidae) and classical biological control releases of Fopius arisanus (Hymenoptera: Braconidae) on economically important fruit flies in French Polynesia

Oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), was discovered on Tahiti Island in July 1996. Eradication programs were conducted from 1997 to 2001, but failed. From 1998 to 2006, B. dorsalis was recovered from 29 different host fruit from the five Society Islands: Tahiti, Moorea, Raiatea, Tahaa, and Huahine. Analysis of coinfestation patterns by B. dorsalis, Bactrocera tryoni (Froggatt), and Bactrocera kirki (Froggatt) suggested B. dorsalis had displaced these two species and become the most abundant fruit fly in coastal areas. To suppress B. dorsalis populations, a classical biological control program was initiated to introduce the natural enemy Fopius arisanus (Sonan) (Hymenoptera: Braconidae) into French Polynesia from Hawaii. Wasps were released and established on Tahiti, Moorea, Raiatea, Tahaa, and Huahine Islands. In guava, Psidium guajava L., collections for Tahiti, F. arisanus parasitism of fruit flies was 2.1, 31.8, 37.5, and 51.9% for fruit collected for 2003, 2004, 2005 and 2006, respectively. Based on guava collections in 2002 (before releases) and 2006 (after releases), there was a subsequent decrease in numbers of B. dorsalis, B. tryoni, and B. kirki fruit flies emerging (per kilogram of fruit) by 75.6, 79.3, and 97.9%, respectively. These increases in F. arisanus parasitism and decreases in infestation were similar for other host fruit. Establishment of F. arisanus is the most successful example of classical biological control of fruit flies in the Pacific area outside of Hawaii and serves as a model for introduction into South America, Africa, and China where species of the B. dorsalis complex are established.     

DNA microsatellite analysis of naturally occurring colour intermediates between Bactrocera tryoni (Froggatt) and Bactrocera neohumeralis (Hardy) (Diptera: Tephritidae)

Abstract  The sympatric tephritid fruit flies Bactrocera tryoni (Froggatt) (Queensland fruit fly) and B. neohumeralis (Hardy) differ in time of mating and for the colour of the humeral callus ('shoulder pad'), which is typically entirely yellow in B. tryoni and typically entirely brown in B. neohumeralis . Field collections in sympatric regions usually include at least 1% of individuals whose humeral calli show mixed patches of yellow and brown ('intermediates'). Over 40 years, a number of studies have debated the possibility that these intermediates are interspecific hybrids. In the present study, we have used microsatellites to show that few if any of these intermediates are hybrids. Instead, most variation humeral callus appears to be confined to one species, B. tryoni . We discuss these results in the context of geneflow between the two species and suggest directions for future research.     

Microsatellite analysis of the Queensland fruit fly Bactrocera tryoni (Diptera: Tephritidae) indicates spatial structuring: implications for population control

The population structure of a tephritid pest species, the Queensland fruit fly Bactrocera tryoni (Froggatt), has been analysed over a five year period (1994-1998), using six microsatellites. Adult fly samples were collected to cover most regions of eastern and central Australia where the flies are regularly found. Tests for heterogeneity indicated that flies within geographically defined regions were homogeneous. The samples were allocated into five regions, including one very large region, Queensland, which encompasses that portion of the fly's range where breeding can occur year-round. With one exception, the collections from different regions were homogeneous between years, showing a fairly static distribution of the species. However, differences between regions were highly significant. The one case of a change in frequency between years indicated a gradual replacement of flies in a marginal region by flies from the main part of the range. The finding of stability in the distribution of a highly mobile insect is of interest, potentially also for other species which have expanded beyond their native range. It is argued that a contributing reason for this stability may be adaptation to different climatic regimes, and that strategies for control based on this hypothesis afford a reasonable chance of success.     

The termite (Isoptera) fauna of a monsoonal rainforest near Darwin, northern Australia

Abstract  Termite species richness, frequency of occurrence and functional diversity at Holmes Jungle Nature Reserve, a monsoon rainforest patch in northern Australia, was investigated at the end of the wet season in 2003. A sampling protocol that employed direct search, soil pits and baiting techniques was used to sample litter, wood, mound, soil and arboreal nest microhabitats for termites. Five species from five genera and three families (Mastotermitidae, Rhinotermitidae and Termitidae) were recorded. This included the first record of Ephelotermes taylori (Hill) from monsoon forest in Australia. The family Termitidae was dominant and represented 70% of termite occurrences. Termites were most frequently encountered in carton runways on tree trunks, followed by lying dead wood and baits. Four nesting habits were represented: arboreal, epigeal, hypogeal and within wood. The arboreal nest-builder Nasutitermes graveolus (Hill) accounted for 61% of termite encounters. Epigeal mound-building species were rare. Wood-feeders were the only trophic group represented. Relatively high activity of Mastotermes darwiniensis Froggatt was recorded at baits within 2.5 months. The low species richness recorded at Holmes Jungle is consistent with the low diversity reported for Australian rainforests generally, but remains relatively depauperate compared with other monsoon forest and savanna habitats of the Northern Territory.     

MORPHOLOGY OF THE IMMATURE STAGES OF DACUS TRYONI (FROGGATT) (DIPTERA: TEPHRITIDAE)

Abstract
External features of the puparium, larvae and egg of the Queensland fruit fly, Dacus (Bactrocera) tryoni (Froggatt) were studied by scanning electron microscopy and significant morphological characters are described and illustrated. Cephalopharyngeal skeletons are described from examination by light microscopy.     

Establishment of <Emphasis Type="Italic">Psyllaephagus parvus</Emphasis> and <Emphasis Type="Italic">P. perplexans</Emphasis> as serendipitous biological control agents of Eucalyptus psyllids in southern California

The free-living lemon gum psyllid, Cryptoneossa triangula Taylor, and the lerp-forming spotted gum psyllid, Eucalyptolyma maideni Froggatt (Hemiptera: Psyllidae) are invasive pests of eucalypts in California. In 2007, Psyllaephagus parvus Riek (Hymenoptera: Encyrtidae) was discovered parasitizing spotted gum psyllids and Psyllaephagus perplexans Cockerell was collected from lemon gum psyllids. While neither parasitoid species was purposely introduced, presence of the parasitoids was significantly associated with reduced intensity and duration of population peaks for both psyllid species. Spring peaks were reduced more than fall peaks. We estimated minimum rates of parasitism from the ratio of mummies to live nymphs. Higher parasitism was recorded in coastal than inland locations during the spring, while parasitism was similar for coastal and inland populations in the fall. Logistic regression models suggest parasitoids were the determining factor of psyllid population densities, although physical parameters, such as irrigation, may affect psyllid or parasitoid populations.