Rod Eastwood

When and where did troidine butterflies (Lepidoptera : Papilionidae) evolve? Phylogenetic and biogeographic evidence suggests an origin in remnant Gondwana in the Late Cretaceous

The age, geographic origin and time of major radiation of the butterflies (Hesperioidea + Papilionoidea + Hedyloidea) are largely unknown. The general modern view is that butterflies arose during the Late Jurassic/Cretaceous in the southern hemisphere (southern Pangea/Gondwana before continental breakup), but this is not universally accepted, and is a best guess based largely on circumstantial evidence. The extreme paucity of fossils and lack of modern, higher-level phylogenies of extant monophyletic groups have been major impediments towards determining reliable estimates of either their age or geographic origin. Here we present a phylogenetic and historical biogeographic analysis of a higher butterfly taxon, the swallowtail tribe Troidini. We analysed molecular data for three protein-encoding genes, mitochondrial ND5 and COI–COII, and nuclear EF–1α, both separately and in combination using maximum parsimony (with and without character weighting and transition/transversion weighting), maximum likelihood and Bayesian methods. Our sample included representatives of all 10 genera of Troidini and distant ingroup taxa (Baroniinae, Parnassiinae, Graphiini, Papilionini), with Pieridae as outgroup. Analysis of the combined dataset (4326 bp; 1012 parsimony informative characters) recovered the Troidini as a well supported monophyletic group and the monophyly of its two subtribes, Battina and Troidina. The most parsimonious biogeographic hypothesis suggests a southern origin of the tribe in remnant Gondwana (Madagascar–Greater India–Australia–Antarctica–South America) sometime after the rifting and final separation of Africa in the Late Cretaceous (<90 Mya). Although an ancient vicariance pattern is proposed, at least four relatively recent dispersal/extinction events are needed to reconcile anomalies in distribution, most of which can be explained by geological and climatic events in South-east Asia and Australia during the late Tertiary. Application of a molecular clock based on a rate smoothing programme to estimate various divergence times based on vicariance events, revealed two peculiarities in our biogeographic vicariance model that do not strictly accord with current understanding of the temporal breakup of Gondwana: (1) the troidine fauna of Greater India did not become isolated from Gondwana (Antarctica) until the end of the Cretaceous (c. 65 Mya), well after Madagascar separated from Greater India (84 Mya); and (2) the faunas of Greater India, Australia and South America diverged simultaneously, also at the K/T boundary. A recent published estimate of the time (31 Mya) of divergence between Cressida Swainson (Australia) and Euryades Felder & Felder (South America) is shown to be in error.

Do ants enhance diversification in lycaenid butterflies? Phylogeographic evidence from a model myrmecophile, Jalmenus evagoras.

Abstract The ant-tended Australian butterfly, Jalmenus evagoras, has been a model system for studying the ecology and evolution of mutualism. A phylogeographic analysis of mitochondrial DNA cytochrome oxidase I sequences from 242 butterflies (615 bp) and 66 attendant ants (585 bp) from 22 populations was carried out to explore the relationship between ant association and butterfly population structure. This analysis revealed 12 closely related butterfly haplotypes in three distinct clades roughly corresponding to three allopatric subpopulations of the butterflies. Minimal genetic diversity and widespread haplotypes within biogeographical regions suggest high levels of matrilineal gene flow. Attendant ants are significantly more diverse than was previously thought, with at least seven well-defined clades corresponding to independent morphological determinations, distributed throughout the range of the butterflies. Nested analysis of molecular variance showed that biogeography, host plant, and ant associate all contribute significantly in explaining variation in butterfly genetic diversity, but these variables are not independent of one another. Major influences appear to come from fragmentation due to large-scale biogeographical barriers, and diversification following a shift in habitat preference. A consequence of such a shift could be codiversification of the butterfly with habitat-adapted ants, resulting in apparent phylogenetic concordance between butterflies and ants. The implications of these results are discussed in terms of possible effects of ant attendance on the diversification of Lycaenidae as a whole.

Molecular phylogeny and evolutionary biology of Acrodipsas (Lepidoptera: Lycaenidae)

Most butterflies in the family Lycaenidae associate with ants but fewer than 3% are myrmecophagous. Larvae of the Australian endemic butterfly genus Acrodipsas parasitise the nests of ants from two subfamilies and, thus, constitute an interesting and uncommon myrmecophagous radiation within the Lepidoptera. Phylogenetic relationships among Acrodipsas species were inferred from fragments of mitochondrial cytochrome oxidase subunits I and II totalling 1155 bp using maximum parsimony and a neighbour joining method. Monophyly of the genus was confirmed, as was the sister genus status of Lucia. Acrodipsas myrmecophila was established as the plesiotypic Acrodipsas species, which together with Acrodipsas brisbanensis parasitises the ancestral dolichoderine host-ants. A speciation event associated with a radical host-ant shift to Myrmicinae occurred in an ancestor of Acrodipsas cuprea but subsequent speciation events may have been driven by climatic fluctuations during the Pleistocene. Modifications to leg morphology in several species were found to be a synapomorphic state, which arose subsequent to the host-shift to Myrmicinae. Minimal genetic variation detected in allopatric species exhibiting divergent morphology suggests that phenotypic variability has been driven by strong environmental selective pressures. As a result, morphological differences between closely related allopatric species have evolved faster than genetic differences, most notably between Acrodipsas arcana and Acrodipsas illidgei. In contrast, sympatric lineages of A. brisbanensis across similar habitats show considerable genetic differentiation, yet have remained phenotypically indistinguishable. Successful amplification of short overlapping fragments of DNA from museum specimens confirms their utility for phylogenetic analysis when the availability of fresh tissue is limited.

The Provenance of Old World Swallowtail Butterflies, Papilio demoleus (Lepidoptera: Papilionidae), Recently Discovered in the New World

Abstract A potential citrus pest, the Old World swallowtail butterfly Papilio demoleus L. was recently discovered in the Dominican Republic; however, there are many subspecies of this butterfly and not all are considered pests of citrus. We used genetic markers to determine the source population (subspecies), by comparing sequences from butterflies collected in the Dominican Republic with individuals whose provenance was known, and to indicate whether single or multiple introductions had taken place. Sequences of up to 1,400 nucleotides from the mitochondrial cytochrome oxidase subunit I gene were characterized from 28 specimens collected in the Dominican Republic and compared with known P. demoleus sequences available from GenBank. We confirmed earlier estimates, based on morphology, that the butterflies had originated in Southeast Asia, where the species is known to be a citrus pest. Lack of sequence variation among specimens from the Dominican Republic suggests that a single introduction in early 2004 was most likely. It is not known how the butterflies reached the Dominican Republic.

Taxonomy, ecology, genetics and conservation status of the pale imperial hairstreak ( Jalmenus eubulus ) (Lepidoptera:Lycaenidae): a threatened butterfly from the Brigalow Belt, Australia

The taxonomic status of Jalmenus eubulus Miskin stat. rev. is revised and considered to be specifically distinct from J. evagoras (Donovan) based on fundamental differences in morphology, ecology and genetics. Miskin’s holotype is fixed by monotypy and illustrated, with type locality Rockhampton, Queensland. Fixed differences in the mitochondrial genomes of J. eubulus and J. evagoras in which the mean pairwise divergence is only 0.85% indicate absence of matrilineal gene flow, whereas allozyme data show significant structure within and between populations of both species consistent with recent diversification. Underlying causes for the observed genetic patterns are investigated. The two species are parapatric, with a narrow range of overlap along the Great Escarpment in south-eastern Queensland. Jalmenus eubulus is restricted to vegetation communities comprising brigalow-dominated old-growth open-forests and woodlands in the Brigalow Belt (with larvae monophagous on Acacia harpophylla F. Muell. Ex Benth), whereas J. evagoras occurs in a range of disturbed eucalypt woodlands/open-forests predominantly in montane and coastal areas east of this bioregion (with larvae polyphagous on Acacia species other than A. harpophylla). The conservation status of J. eubulus is considered to be vulnerable nationally and critically endangered in New South Wales according to International Union for Conservation of Nature (IUCN) criteria. Nationally, the geographic range has an estimated area of occupancy of less than 2000 km2, is severely fragmented, and the extent or quality of its habitat, which is poorly conserved, continues to decline. It is recommended that the taxon be used as an indicator for identification of remnant old-growth forest for conservation planning, as well as a flagship for the conservation of invertebrate biodiversity associated with this threatened ecological community.

The fig wax scale Ceroplastes rusci (Linnaeus) (Homoptera: Coccidae) in south-east Vietnam: pest status, life history and biocontrol trials with Eublemma amabilis Moore (Lepidoptera: Noctuidae).

The fig wax scale Ceroplastes rusci (Linnaeus) is a serious pest of fruit trees in many countries. In the present study we investigated the extent of C. rusci infestation and the range of host species, particularly in fruit orchards, in south‐east Vietnam. Captive populations of C. rusci were established to record life history parameters and to investigate the efficacy of an endemic predacious moth, Eublemma amabilis Moore, as a potential biocontrol agent. Heavy infestation of C. rusci (up to 100%) was recorded in soursop and other fruit orchards, and the culturally important Hoa Mai flower was also affected. Captive breeding trials found E. amabilis to be an efficient predator of C. rusci, but an unusual hyperactive trait in early instars of E. amabilis resulted in lower than expected survival rates. The implications of this trait in terms of the laboratory environment, augmentative release protocols and as a survival strategy are discussed.

New Australian butterfly genus Jameela gen. nov. (Lepidoptera: Lycaenidae: Polyommatinae: Polyommatini) revealed by morphological, ecological and molecular data

This paper proposes to split the Australian Polyommatini genus Erysichton Fruhstorfer, 1916 into two genera on the basis of molecular, morphological and early stage behavioral attributes. The utility of reflective eye pattern as a taxonomic character is discussed and a neotype of Lycaena lineata Murray, 1874 is designated.

Life history, damage assessment and control of Trioza hopeae (Hemiptera: Psylloidea), a serious pest on Hopea odorata (Malvales: Dipterocarpaceae) in Vietnam

Young plants of the commercially important timber species Hopea odorata Roxb. are seriously damaged by the psyllid Trioza hopeae Burckhardt & Vu in nurseries in South Central and Southern Vietnam. Farmers routinely use Class 1 toxic insecticides to control the psyllids but these pose serious health risks in the urban environment. As a precursor to developing a pest management strategy, we studied the life cycle of T. hopeae and trialed several control measures. Trioza hopeae is polyvoltine with overlapping generations peaking in mid wet season. Females laid approximately 135 eggs; however, there was a natural attrition rate of about 80% in first‐instar nymphs due to the nymphs inability to find a suitable feeding site. Feeding nymphs induced pit galls on young H. odorata leaves and were present on all trees examined during the study. The pre‐adult life cycle lasted approximately 13 days and adult life span approximately seven days. Several toxic insecticides were found to be effective for controlling T. hopeae, but a novel non‐toxic alternative was equally as effective. We discuss these methods and potential biological control measures.

Graphium agamemnon Linnaeus (Lepidoptera: Papilionidae), a pest of soursop (Annona muricata Linnaeus), in Vietnam: Biology and a novel method of control

Few butterflies are pests of economic significance, but some may be locally destructive, such as the papilionid Graphium agamemnon Linnaeus, which is known to feed on the commercially important soursop (Annona muricata Linnaeus) in Vietnam. This paper documents the life history and ecology of G. agamemnon and investigates commonly used control measures in south‐east Vietnam. A novel method of controlling G. agamemnon infestation is described. If soursop basal rootstock (Annona glabra Linnaeus) is encouraged to shoot during times of peak butterfly activity, ovipositing G. agamemnon females are preferentially attracted to the new growth where the resulting early stages may be easily detected and removed by hand.