THE AUSTRALIAN

ntomologist

published by

THE ENTOMOLOGICAL SOCIETY OF QUEENSLAND

Volume 34, Part 2, 25 May 2007

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ISSN 1320 6133

THE AUSTRALIAN ENTOMOLOGIST

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Cover: Parobia alipilus Seeman & Nahrung (Podapolipidae) is one of three sexually-

transmitted mites that infest the eucalyptus leaf beetle Paropsis atomaria. The adult

male (pictured) is unlike the female: his legs have spurs, the genital capsule is mid-

dorsal and the fourth pair of legs is modified. Nevertheless, Parobia species are

among the most mite-like of the Podapolipidae, where physogastry and loss of legs is

common and some males have their genitalia on prongs above their heads. Parobia

mites are often more common on female beetles and at least one species significantly

reduces survival of the host during overwintering. Illustration by Owen Seeman.

Australian Entomologist, 2007, 34 (2): 33-42 33

PHYLOGENY OF THE TROIDINE BUTTERFLIES (LEPIDOPTERA:

PAPILIONIDAE) REVISITED: ARE THE RED-BODIED

SWALLOWTAILS MONOPHYLETIC?

DAVID L. HANCOCK

PO Box 2464, Cairns, Qld 4870

Abstract

The phylogenetic relationships and biogeography of troidine butterflies are reassessed. The

subtribe Troidina is considered to comprise the following generic groups, representing vicariant

separation within western, central and eastern Gondwana respectively: Parides Hiibner +

Euryades C. & R. Felder; the ‘red-bodied swallowtails’ Atrophaneura Reakirt + (Pachliopta

Reakirt + (Losaria Moore, stat. rev. + Cressida Swainson)); and the ‘birdwings’ Trogonoptera

Rippon + (Troides Hübner + Ornithoptera Boisduval). Pharmacophagus Haase, with its sole

species P. antenor (Drury) from Madagascar, is retained as a subgenus of Pachliopta. The

suggestion that Pharmacophagus, Cressida and Euryades are relict taxa is rejected.

Introduction

Recent studies on the phylogeny and biogeography of troidine butterflies

have shown little agreement. For example, Hancock (1983, 1988), Miller

(1987) and Parsons (1996a, b) produced classifications which differed

markedly, particularly with regard to the placement of Atrophaneura Reakirt

and its allies. Until now, it has not been possible to reconcile these

differences. However, a recent study by Braby ef al. (2005) has strongly

suggested that almost all previous phylogenies were based on a

misconception - that Cressida Swainson and Euryades C. & R. Felder were

sister genera. The suggestion by Braby et al. (2005) that this is not the case

enables several morphological characters, previously considered to be

homoplasious (e.g. the distinctive red abdomen seen in several genera,

including Cressida), to be reassessed and seen as potential synapomorphies.

Morphological relationships

Monophyly of the tribe Troidini and subtribes Battina and Troidina has been

recognised by most recent authors (e.g. Hancock 1983, Miller 1987, Braby et

al. 2005) and appears well established. Subtribe Battina contains the sole

genus Battus Scopoli, known only from North and South America. It is of

either North American (Hancock 1983) or Gondwanan (Braby et al. 2005)

origin. Subtribe Troidina contains all the remaining genera in the tribe and is

of undoubted Gondwanan origin. Three distinct groups of genera are

recognisable on morphological grounds; these appear to be vicariant groups

originating in eastern, western and central Gondwana respectively. Genera in

this subtribe are united by numerous characters (Hancock 1983, Miller 1987),

including the presence of a basal seta on the subdorsal tubercles of the first

instar larva (Igarashi 1984) and red hairs on the head and thorax (and often

the abdomen) of the adult, a condition not seen elsewhere in the Papilionidae.

Larval food plants for the Troidini are all members of the family

Aristolochiaceae (Aristolochia, Pararistolochia, Thottea and Apama).

34 Australian Entomologist, 2007, 34 (2)

Trogonoptera Rippon, Troides Hiibner and Ornithoptera Boisduval

Although disputed by Parsons (1996a, b), monophyly of this group of genera,

the ‘birdwings’, is well established. The relationship Trogonoptera +

(Troides + Ornithoptera) was suggested by Hancock (1983, 1991) and

supported by Miller (1987, as subordinate groups within Troides), Hancock

and Orr (1997) and Braby et al. (2005). [Note that in Fig. 3 of Braby et al.

(2005), the names Troides and Troides (Ornithoptera) should be transposed].

The genera are restricted to the Indo-Australian region. They share the

following characters: abdomen without red hairs; juxta U-shaped;

pseuduncus generally long, apically pointed and with a basal suture

(secondarily reduced in Trogonoptera); aedeagus short; bursa large with

membranous ductus bursae and well developed appendix bursae; signum

comprised of concentric folds (all characters similar to those seen in Battus).

Defining characters were noted by Hancock (1983) and Miller (1987).

Parides Hiibner and Euryades C. & R. Felder

Although their evidence was not well supported, Braby et al. (2005)

suggested that these two genera were sister groups. Significantly, they found

no evidence of a relationship between Euryades and Cressida. The genera

occur in Central and/or South America and share the following characters:

abdomen with red hairs, especially posteroventrally; juxta Y-shaped with

narrow anterolateral expansions; pseuduncus long, apically pointed and with

a basal suture; aedeagus short; female bursa with membranous ductus bursae

and no appendix bursae. Larvae with spinose setae on subdorsal tubercles

retained beyond the first instar [absent in other groups and Battus] (Parsons

1996a). In Parides the valvae are entire, the aedeagus broad, the bursa large

and the signum broadly C-shaped and transverse. In Euryades the valvae are

reduced, the aedeagus narrow, the juxta with long apodemes, the ovipositor

lobes melanised, the bursa small and the signum absent.

Atrophaneura Reakirt and allies

Monophyly of this group has not been recognised previously. Four Indo-

Australian genera and one Madagascan subgenus are included. All have

extensive red hairs and scales on the abdomen, much better developed than in

Parides and Euryades. The female bursa has no appendix bursae and the

signum, when present, is neither broadly C-shaped nor formed of concentric

folds. The first instar larva has the sclerotised, setose apex of the tubercles

rounded and dome-like [pinnacle or pagoda-like in other groups] and the

pupa has the head short and truncate in lateral view [distinctly produced

anterolaterally in other groups and Battus] (Igarashi 1984, Parsons 1996a).

Atrophaneura

This is an eastern and southeast Asian genus of some 27 species referred to

the /atreillei and nox groups by Hancock (1980, 1988). Miller (1987)

regarded these two groups as subgenera included in an unresolved trichotomy

Australian Entomologist, 2007, 34 (2) 35

with Parides. It differs from other genera in the group by the retention of a

well developed scent organ on the male hind wing, a distinct, apically pointed

pseuduncus with a basal suture, a short, broad aedeagus, and a large bursa

with membranous ductus bursae. The juxta appears to be a modification of

the Y-shape, while the signum is longitudinal, expanded anteriorly and

narrowly tapered posteriorly. On the hind wing the tail is broadly spatulate

(latreillei group) or absent (nox group) and white areas, when present, at

most weakly enter the discal cell. The valvae are entire or weakly emarginate.

Larva with a distinct supraspiracular tubercle on first abdominal segment and

pupa with distinct, apically pointed subdorsal projections on abdominal

segments 2-10 and a broad lateral flange on segment 4 (Igarashi 1984).

Losaria Moore, Cressida Swainson and Pachliopta Reakirt

These genera differ from Atrophaneura in the reduction of the scent organ

and shortened vein 1A+2A on the hind wing, elongate and narrow aedeagus,

and sclerotised posterior part of the ductus bursae [not evident in Cressida].

They occur primarily in the Indo-Australian region, with a single species in

Madagascar. The tegumen and socii are frequently enlarged and the valvae

are usually reduced. Larva with supraspiracular tubercle on first abdominal

segment usually absent and pupa with apically rounded or truncate subdorsal

projections on abdominal segments 4-7 (and very weakly on segment 8) and

with lateral flange on segment 4 present or absent.

Losaria

This is a Southeast Asian genus of four species referred to the coon group by

Hancock (1983, 1988). An affinity with Pachliopta was recognised by

Igarashi (1984), Hancock (1988) and Miller (1987), who placed it either as a

subgenus of the latter (Miller 1987) or as a synonym of it. Because of its

apparent sister-group relationship with Cressida, it is here raised to generic

status (stat. rev.), characterised by the elongate forewing discal cell, deeply

indented margin to cell Cu of the male hind wing, strongly petiolate and club-

shaped tail, posteriorly downcurved abdomen, Y-shaped juxta with broad

anterolateral expansions, often reduced but apically acute pseuduncus with a

basal suture, reduced valvae, and large bursa with a small, round signum. A

sphragis is present in one species and in two the ‘red’ abdominal markings

are yellow. When the hind wing has a white area that crosses the discal cell,

the apex of the cell is black. Larva with supraspiracular tubercle on first

abdominal segment absent (Igarashi 1984) and subdorsal tubercles on

abdominal segments 2 and 3 conspicuously reduced (Weintraub 1995). Pupa

with subdorsal projections large and lateral flange present (Igarashi 1984).

Cressida

This is a monotypic Australian genus previously placed with Euryades. It

differs from related genera in the absence of a tail, large precostal hindwing

cell, vestigial bursa and lack of a signum. It shares with Losaria the elongate

forewing discal cell, deeply indented margin to cell Cu of the male hind

36 Australian Entomologist, 2007, 34 (2)

wing, black apex of the hindwing discal cell, posteriorly downcurved

abdomen, presence of a Y-shaped juxta with broad anterolateral expansions,

apically pointed pseuduncus with a basal suture, and similarly reduced

valvae. With both Euryades and L. palu (Martin) it shares a well-developed

sphragis but these appear to be independently acquired and not homologous

(Orr 1988, 1995). Larva with supraspiracular tubercle on first abdominal

segment absent and all other tubercles reduced; pupa with head truncated in

lateral view, subdorsal projections small and lateral flange absent (Igarashi

1984). The male valvae also resemble those of Euryades but other characters

of Euryades [including the socii, pinnacle-like apex of the first instar larval

tubercles and anterior projection of the pupal head in lateral view (Miller

1987, Parsons 1996a)] more closely resemble those of Parides than those of

Cressida. The distinctively red abdomen, dome-like apex of the first instar

larval tubercles and truncated pupa in Cressida provide the best evidence of

the affinity of this genus with others in the Atrophaneura complex.

Pachliopta

This genus contains 13 species in the Indo-Australian region and one in

Madagascar. They were referred to the polydorus, hector and antenor groups

by Hancock (1988). The genus is characterised by the spatulate tail [reduced

in P. polydorus (Linnaeus)], T-shaped juxta, short, blunt pseuduncus with

basal suture absent (at least medially), valvae either entire or vestigial, and

large bursa with an elongate signum of uniform width. When the hind wing

has a white area that crosses the discal cell, the apex of the cell is also white.

Larva with subdorsal tubercles not conspicuously reduced. Two subgenera

are recognised: Pachliopta Reakirt and Pharmacophagus Haase. In both the

pupa is strongly S-shaped in lateral view and has the lateral flange on

abdominal segment 4 present; the anal segments and cremaster, viewed

ventrally, are also very similar (Igarashi 1984, Parsons 1996c).

Subgenus Pachliopta

This subgenus contains all the Indo-Australian species. Braby et al. (2005)

noted that P. hector (Linnaeus) was more closely related to species in the

polydorus group than to P. antenor, with which it was associated by Hancock

(1988). This is supported by the wide separation between tergite 8 and the

hypertrophied tegumen and socii, and by the vestigial valvae (see Miller

1987); thus P. hector is here subsumed into the polydorus group. Larva with

the supraspiracular tubercle on the first abdominal segment reduced to a spot

in P. hector (Woodhouse and Henry 1942), absent in other species

(Woodhouse and Henry 1942, Igarashi 1984). Pupa with subdorsal

projections large and lateral flange well developed (Igarashi 1984).

Subgenus Pharmacophagus

The sole species [P. antenor (Drury)] occurs in Madagascar. It differs from

subgenus Pachliopta in the presence of normal (not vestigial) valvae, a well

developed supraspiracular tubercle on the first abdominal segment and

Australian Entomologist, 2007, 34 (2) 37

reduced subdorsal tubercles on abdominal segments 3 and 4 of the larva, and

small subdorsal projections and lateral flange on the pupa (Parsons 1996a, c).

The tail is relatively narrow, similar to that of P. hector. Often regarded as a

distinct genus, the form of the juxta, pseuduncus and signum, and the

presence of blue-green hindwing scales in a rare form of P. hector

(Woodhouse and Henry 1942), support a close relationship with Pachliopta.

It was regarded as a subgenus of the latter by Hancock (1993).

Discussion

Pharmacophagus was regarded as a distinct genus by Igarashi (1984), Miller

(1987), Parsons (1996a, b, c) and Braby et al. (2005). Igarashi (1984)

considered it to be ‘closely related to Pachliopta but highly specialised’.

Miller (1987) placed it at the base of the subtribe, i.e. as the most primitive

genus. However, he identified only one character that unambiguously

separated it from Pachliopta - the 3-segmented palpi with a ‘basal fleck’

present. All other genera in the Troidina have 2-segmented palpi [which

appear to have formed by fusion of segments 1 and 2: see illustrations in

Miller 1987] and no ‘basal fleck’. Hancock (1989) suggested that the 3-

segmented palpi of P. antenor resulted from a secondary loss of this fusion

(character reversal) and that the ‘basal fleck’ was also secondarily acquired,

with neither homologous to the condition seen in Battus. Its placement as

sister genus to Ornithoptera (Parsons 1996a, b) was disputed by both

Hancock and Orr (1997) and Braby et al. (2005) and is not sustainable.

In their molecular study, Braby et al. (2005) also regarded Pharmacophagus

as basal to all other genera in the subtribe. However, only one of the three

genes studied (EF-/a) supported that arrangement. The NDS gene produced a

trichotomy of (i) Pharmacophagus, (ii) Cressida, and (iii) all other genera,

whereas the CO/-COII gene produced an unresolved polytomy of all genera

plus Battus. Their combined analysis also produced a clade with

Pharmacophagus at its base, but the placement of Papilio Linnaeus as sister

to the Parnassiinae in the combined clade, and the placement of the

Baroniinae and Parnassiinae between Graphium Scopoli and the Troidini (or

Papilio + Troidini) in all of them, suggest that the use of these genes in

resolving higher-level groupings requires considerable caution. Unreliability

of the NDS gene at higher levels was also noted by Braby et al. (2005).

At lower taxonomic levels, recognition of the generic groupings Parides +

Euryades, [Trogonoptera +] Ornithoptera + Troides, and Atrophaneura +

Pachliopta (including Losaria) by Braby et al. (2005) appears to be well

founded, although they were unable to effectively place the seemingly

isolated genera Pharmacophagus and Cressida. Resolution in these cases is

best done using morphological characters that, although subject to various

interpretations, strongly suggest a sister-group relationship between

Pharmacophagus and Pachliopta in the first case and between Cressida and

Losaria in the other.

38 Australian Entomologist, 2007, 34 (2)

An Indo-Australian plus Madagascan pattern of distribution, as shown by

subgenera Pachliopta and Pharmacophagus, also occurs in pitcher plants

(Nepenthes: Nepenthaceae) (Heywood 1978) and the fruit fly genus Dacus

Fabricius (Diptera: Tephritidae), where the largely Indo-Australian subgenus

Neodacus Perkins is represented in Madagascar by the relatively primitive

xanthaspis group (Hancock and Drew 2006, White 2006). Furthermore, the

largely Indo-Australian butterfly genus Euploea Fabricius (Nymphalidae)

[and which, like Pachliopta hector, is known to be migratory] is represented

in the Malagasy region by apparently unrelated endemic species in the

Mascarenes and Seychelles, but not in Africa or Madagascar (Holloway and

Nielsen 1999). Under the above classification, there is no need to invoke an

ancient (pre break-up Gondwana) origin for P. antenor, which appears to

have differentiated much more recently. Its only confirmed food plant,

Aristolochia acuminata (= A. indica; = A. tagala) [also utilised by P. hector,

P. polydorus and P. aristolochiae (Fabricius)], is widespread throughout the

Indo-Australian region (Sands 2002) and is the only Aristolochia species

known from Madagascar (Parsons 1996c). Ar. acuminata is also likely to be a

relatively recent arrival from India and there is no evidence that a suitable

host existed in Madagascar [or in Africa] prior to its arrival.

The association of Cressida with Losaria suggests it is not the Gondwanan

relict previously believed but a relatively recent arrival to Australia from a

Southeast Asian, Losaria-like ancestor. As in the case of Pharmacophagus,

its dispersal as a small founder population into a new environment might

have accentuated its genetic differentiation. In both cases, the modified pupa

is likely to reflect a need for crypsis rather than an ancestral state. The

various specialisations seen in Euryades suggest that it, too, is of relatively

recent derivation. Sphragis development in Cressida, Euryades and Losaria

palu appears to reflect their dispersal into open habitats, as has been

demonstrated in the nymphalid genus Acraea Fabricius (Orr 1988).

Phylogenetic relationships of the genera and subgenera, as proposed here, are

shown in Fig. 1.

Biogeography

Based on the above morphological evidence, the following biogeographical

scenario for the subtribe Troidina is proposed. [For dating see Braby et al.

2005 and references therein]. During the continental drifting phase following

the break-up of Gondwana, the three main lineages within the subtribe

Troidina (Fig. 1) appear to have been confined to the Southeast Asian (or

Australian), South American and Greater Indian plates respectively.

Assuming Africa had already separated, this dates the fragmentation of the

ancestral population at ca 90-80 Mya [84-80 Mya if Madagascar is also

excluded]. The present restriction of both Parides and Euryades to Central

and/or South America strongly supports the suggestion that these two genera

now represent the western lineage of the original Gondwanan ancestor.

Australian Entomologist, 2007, 34 (2) 39

Subtribe BATTINA

Battus

Subtribe TROIDINA

(Eastern lineage)

Trogonoptera

Troides

Ornithoptera

Western lineage

( ge) Parides

Euryades

(Central lineage)

Atrophaneura

Losaria

Cressida

Pachliopta

(Pachliopta)

Pachliopta

(Pharmacophagus)

Fig. 1. Suggested phylogenetic relationships of the genera and subgenera of tribe

Troidini. See text for discussion of defining character states.

40 Australian Entomologist, 2007, 34 (2)

Similarly, the present distributions of Trogonoptera, Ornithoptera and

Troides support the suggestion that they represent the eastern lineage of the

Gondwanan ancestor. It remains unclear whether the lineage radiated directly

from the Southeast Asian (Sundaland) block (see Hancock 1988) into the

Papuan subregion via Wallacea [twice, first as Ornithoptera via a Sulawesian

Troides ancestor similar to T. hypolitus (Cramer) then, more recently, as

Troides oblongomaculatus (Goeze)], or (see Braby et al. 2005) from

Australia into southeast Asia [T7rogonoptera and Troides, leaving

Ornithoptera in Australia] and, in the case of 7. oblongomaculatus, back

again [with subsequent extinction of the Trogonoptera and Troides ancestors

in Australia]. The former scenario does not require extinctions and seems the

more likely. Based on molecular dating, Braby et al. (2005) suggested that

the Ornithoptera-Troides split occurred around 42 + 8 Mya but this is

doubtful and the split is likely to be significantly younger. The Southeast

Asian and Australian plates did not make contact until 25 Mya (Braby et al.

2005) and whether suitable host plants reached Australia prior to the break-up

of Gondwana, or only after contact with Southeast Asia, is uncertain.

Biogeography of the central lineage, the red-bodied Atrophaneura and its

allies, is more complex. Due to the reassessment of genera such as

Pharmacophagus, Losaria and Cressida, the scenario proposed here differs

in some details from those suggested by Hancock (1988) and Braby et al.

(2005). This lineage appears to have been associated with the Indian plate as

it broke free from the rest of Gondwana (ca 80 Mya), dispersing into Asia as

Atrophaneura [with increased pupal projections] once contact with it was

possible (50-45 Mya). An ability to utilise Thottea and Apama as host plants

might have assisted this dispersal. The suggestion of an Indian origin is

supported by the presence of the relatively unspecialised /atreillei group in

eastern Asia, which subsequently appears to have dispersed into Southeast

Asia as the more specialised nox group (Hancock 1988).

A second dispersal of the Indian ancestor, now differentiated into a

Pachliopta-like taxon following major climatic, tectonic and environmental

changes within the subcontinent, appears to have brought a Losaria-like

ancestor into Southeast Asia - Sundaland. This dispersal, and that of the nox

group, possibly accompanied the southward spread of Thottea and Apama

following contact between the Sundaland block and the rest of Asia.

Subsequent dispersal and differentiation has brought Cressida to Australia,

Timor to Tanimbar Is and southern Papua New Guinea, and Pachliopta

(Pharmacophagus) to Madagascar, leaving Losaria in southeast Asia and

subgenus Pachliopta (as P. hector) in India. Dispersal to Madagascar

possibly took place via the Laccadive-Maldive-Chagos archipelago during a

period of lowered sea level. Unlike the condition in Losaria and Cressida,

none of the distinguishing characters of Pharmacophagus appear to be

distinctive enough to recognise it at generic level; most are homoplasious.

Australian Entomologist, 2007, 34 (2) 4l

A second major radiation within subgenus Pachliopta then appears to have

occurred, dispersing again (as the polydorus group minus P. hector) to

Southeast Asia - Sundaland and leaving P. hector in India. In these species

the wing pattern appears to mimic that of Losaria, supporting the suggestion

that they radiated first to southeast Asia and from there back to India and Sri

Lanka (twice, first as P. pandiyana (Moore) + P. jophon (Gray), then more

recently as P. aristolochiae). Subsequent radiations within southeast Asia

appear to have resulted in the differentiation of: (1), P. mariae (Semper), P.

polyphontes (Boisduval) and P. oreon (Doherty) + P. liris (Godart) in the

Philippines, Sulawesi and northern Moluccas, and the Lesser Sunda Islands +

Timor and neighbouring islands, respectively; (2), the elongate-tailed and

largely allopatric P. schadenbergi (Semper), P. leytensis Muruyama [=

phegeus (Hopffer)], P. phlegon (C. & R. Felder) [= annae (C. & R. Felder)]

and P. atropos (Staudinger) in the Philippines; and (3), P. polydorus from the

Moluccas to Australia and the Solomon Islands, leaving the widespread P.

aristolochiae in much of south and southeast Asia, including Sundaland,

Sulawesi and the Philippines.

The centre of origin of the tribe Troidini remains unresolved. Braby et al.

(2005) objected to the suggestion of a North American origin (Hancock

1983) on the grounds that it ‘does not explain the absence of the Troidini in

Europe and Africa’. However, such absence might merely reflect a lack of

suitable host plants or habitat, either now or in the past, or the tribe’s failure

to disperse there. Within the Troidina the red abdominal hairs, shape of the

juxta and loss of the appendix bursae suggest that the Atrophaneura group is

more closely related to Parides + Euryades than to the Troides group and that

the latter is the most primitive. If so, then Battus (subtribe Battina) likely is of

South American origin, with subtribe Troidina evolving in eastern Gondwana

and subsequently dispersing back to South America as the ancestor of

Parides + Euryades. Any original North American ancestor, if such existed,

presumably become extinct due to subsequent climatic changes (ice ages)

and/or competition from reinvading Battus.

Acknowledgement

I thank Bert Orr for helpful comments on the manuscript.

References

BRABY, M.F., TRUEMAN, J.W.H. and EASTWOOD, R. 2005. When and where did troidine

butterflies (Lepidoptera: Papilionidae) evolve? Phylogenetic and biogeographical evidence

suggests an origin in remnant Gondwana in the Late Cretaceous. /nvertebrate Systematics 19:

113-143.

HANCOCK, D.L. 1980. The status of the genera Atrophaneura Reakirt and Pachliopta Reakirt

(Lepidoptera: Papilionidae). Australian Entomological Magazine 7(2): 27-32.

HANCOCK, D.L. 1983. Classification of the Papilionidae (Lepidoptera): a phylogenetic

approach. Smithersia 2: 1-48.

HANCOCK, D.L. 1988. A revised classification of the genus Atrophaneura Reakirt

(Lepidoptera: Papilionidae). Australian Entomological Magazine 15(1): 7-16.

42 Australian Entomologist, 2007, 34 (2)

HANCOCK, D.L. 1989, The relationships of Meandrusa Moore. Papilio International 6: 460-

470.

HANCOCK, D.L. 1991. Notes on the phylogeny and biogeography of Ornithoptera Boisduval

(Lepidoptera: Papilionidae). Tyô to Ga 42(1): 17-36.

HANCOCK, D.L. 1993. Origins and evolution of the Afrotropical Papilionidae (Lepidoptera).

Arnoldia Zimbabwe 9(40): 557-583.

HANCOCK, D.L. and DREW, R.A.I. 2006. A revised classification of subgenera and species

groups in Dacus Fabricius (Diptera, Tephritidae). Pp 167-205, in: Merz, B. (ed.), Phylogeny,

taxonomy, and biology of tephritoid flies (Diptera, Tephritoidea). Instrumenta Biodiversitatis

Vol. VII, Natural History Museum, Geneva; 274 pp.

HANCOCK, D.L. and ORR, A.G. 1997. Ornithoptera euphorion (Gray) (Lepidoptera:

Papilionidae): species or subspecies? Australian Entomologist 24(4): 165-168.

HEYWOOD, V.H. (ed.). 1978. Flowering plants of the world. Oxford University Press, Oxford;

336 pp.

HOLLOWAY, J.D. and NIELSEN, E.S. 1999. Biogeography of the Lepidoptera. Pp 423-462,

in: Kristensen, N.P. (ed.), Lepidoptera, moths and butterflies. Vol. 1: Evolution, systematics, and

biogeography. Handbook of Zoology, Vol. IV, Part 35. De Gruyter, Berlin; 491 pp.

IGARASHI, S. 1984. The classification of the Papilionidae mainly based on the morphology of

their immature stages. Tyô to Ga 34(2): 41-96.

MILLER, J.S. 1987. Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae).

Bulletin of the American Museum of Natural History 186(4): 365-512.

ORR, A.G. 1988. Mate conflict and the evolution of the sphragis in butterflies. PhD thesis,

Griffith University, Brisbane.

ORR, A.G. 1995. The evolution of the sphragis in the Papilionidae and other butterflies. Pp 155-

164, in: Scriber, J.M., Tsubaki, Y. and Lederhouse, R.C. (eds), Swallowtail butterflies: their

ecology and evolutionary biology. Scientific Publishers, Gainesville.

PARSONS, M.J. 1996a. Gondwanan evolution of the troidine swallowtails (Lepidoptera:

Papilionidae: Troidini): cladistic reappraisals using mainly immature stage characters, with focus

on the birdwings Ornithoptera Boisduval. Bulletin of the Kitakyushu Museum of Natural History

15: 43-118.

PARSONS, M.J. 1996b. A phylogenetic reappraisal of the birdwing genus Ornithoptera

(Lepidoptera: Papilionidae: Troidini) and a new theory of its evolution in relation to Gondwanan

vicariance biogeography. Journal of Natural History 30: 1707-1736.

PARSONS, M.J. 1996c. The immature stages of Pharmacophagus antenor (Drury)

(Papilionidae: Troidini) from Madagascar. Journal of the Lepidopterists’ Society 50(4): 337-344.

SANDS, D. 2002. The food plants of the birdwing larvae. Pp 14-17, in Sands, D. and Scott, S.

(eds), Conservation of birdwing butterflies. SciComEd and THECA, Brisbane; 48 pp.

WEINTRAUB, J.D. 1995. Host plant association patterns and phylogeny in the tribe Troidini

(Lepidoptera: Papilionidae). Pp 251-252 [figs], 307-316, in: Scriber, J.M., Tsubaki, Y. and

Lederhouse, R.C. (eds), Swallowtail butterflies: their ecology and evolutionary biology.

Scientific Publishers, Gainesville.

WHITE, I.M. 2006, Taxonomy of the Dacina (Diptera: Tephritidae) of Africa and the Middle

East. African Entomology Memoir 2: [i-v], 1-156, cd-rom.

WOODHOUSE, L.G.O. and HENRY, G.M.R. 1942. The butterfly fauna of Ceylon. Ceylon

Journal of Science, Colombo; 172 pp.

Australian Entomologist, 2007, 34 (2): 43-48 43

REVIEW OF THE GENUS DUNGOORUS CARNE (COLEOPTERA:

SCARABAEIDAE: RUTELINAE: ANOPLOGNATHINI)

ANDREW B.T. SMITH

Canadian Museum of Nature, Research Division, PO Box 3443, Station D, Ottawa, Ontario,

KIP 6P4, Canada. Email: asmith@mus-nature.ca

Abstract

The genus Dungoorus Carne is reviewed and now contains two species: D. murrumbullus Carne

and D. frater sp. n. An identification guide is given for the genus and both species, including

descriptions, diagnoses and a key to species. Distributional data and a description of the female

are presented for the first time.

Introduction

The ruteline beetle tribe Anoplognathini occurs only in Australia and the

Neotropics. Phillip Carne last reviewed the Australian taxa in a series of

papers published in the 1950s. In his main work on this group, Carne (1958)

described many new taxa, including the monotypic genus Dungoorus Carne.

He placed it in the subtribe Schizognathina and commented that it was

similar to the genus Saulostomus Waterhouse. The description of Dungoorus

and the new species D. murrumbullus Carne were both based on a single

specimen without locality data [although Carne (1958) hypothesized that the

specimen was collected in Queensland, based on other specimens he was

familiar with bearing similar labels]. Since the original description, no new

information has been published on this genus. Recently, I discovered further

specimens in three collections. This paper reviews the genus Dungoorus

based on these additional specimens and provides previously unknown

information on this genus, including a description of females, reliable

distributional data and the discovery of a new species.

Specimens were borrowed from and deposited in the following institutions

(collections managers and/or curators listed in parenthesis). A total of 23

specimens formed the basis of this review. All specimens examined were

labelled with a determination label or one of my red or yellow type labels.

ABTS - Andrew B.T. Smith Collection, Ottawa, ON, Canada; ANIC -

Australian National Insect Collection, CSIRO, Canberra, ACT (Tom Weir);

CASC - California Academy of Sciences, San Francisco, CA, USA (David

Kavanaugh, Roberta Brett); CMNC - Canadian Museum of Nature, Ottawa,

ON, Canada (Robert Anderson, François Génier); QMBA - Queensland

Museum, Brisbane, Qld (Geoff Monteith).

Genus DUNGOORUS Carne, 1958

(Figs 1-3)

Dungoorus Carne, 1958: 198, 218 [key to genera of Australian Rutelinae, original

description]; Machatschke, 1965: 13, 47 [distribution, catalogue listing];

Machatschke, 1972: 297 [catalogue listing]; Cassis and Weir, 1992: 375 [catalogue

listing]; Smith, 2003: 199 [checklist].

44 Australian Entomologist, 2007, 34 (2)

Type species. Dungoorus murrumbullus Carne, 1958, by original designation. Gender

of genus: masculine.

Description. Length 11.7-14.8 mm, width 6.8-8.2 mm. Colour yellowish-

brown. Body ovate, convex. Head (Fig 1-2): Dorsal surface moderately

punctate on frons, densely punctate on clypeus; clypeus with erect, golden-

brown setae. Clypeal apex strongly reflexed. Labrum reduced, apex without

medial tooth or projection. Mandible plate-like, projecting well past apex of

clypeus in dorsal view; apex and lateral margin strongly reflexed. Terminal

maxillary palpomere in males greatly enlarged with elongate sulcus,

approximately as long as antennal club. Terminal maxillary palpomere in

females not greatly enlarged, much shorter than antennal club. Mentum

constricted apically but not curved into oral cavity, without apical notch or

tooth. Antenna with 9 antennomeres. Pronotum: Disc glabrous, moderately

punctate. Marginal bead present laterally, absent medially on base and apex.

Elytron: Surface glabrous; longitudinal striae poorly defined, densely

punctate; without membranous border. Suture apically rounded, without

acute spine or projection. Pygidium: Surface convex, smooth, setose apically;

setae erect, golden brown. Venter: Thorax setose. Mesothoracic process not

developed. Abdominal sternites sparsely setose. Legs: Protibia with 3 sharp,

subequal teeth; apical spur absent. Tarsomeres 1-4 in males as wide or wider

than long, cup-shaped. Protarsomere 1 in females long, approximately as long

as protarsomeres 2-4. Protarsomeres 3-4 in males with weak internoapical

stridulatory ridges: Protarsomere 5 in males with weak internomedial,

stridulatory surface. Unguitractor plate laterally flattened with 2 setae.

Modified tarsal claws in males thickened, elongate when compared with

other claw, without teeth or bifurcations. Modified tarsal claws in females

simple, not thickened or toothed.

Male genitalia. Phallobase not fused with parameres. Parameres weakly

fused, longitudinally contiguous.

Diagnosis. Dungoorus is distinguished from all other genera of

Anoplognathini by the following combination of characters: labrum reduced,

apex without medial tooth or projection; mandible plate-like, projecting well

past apex of clypeus in dorsal view and with apex and lateral margin strongly

reflexed; terminal maxillary palpomere in males greatly enlarged,

approximately as long as antennal club; mentum flat, constricted apically but

not curved into oral cavity; antenna with 9 antennomeres; elytron without

membranous border, glabrous; mesothoracic process absent; protibia without

spur; tarsomere 5 with simple claws; unguitractor plate with 2 setae; male

genitalia with phallobase and parameres not fused; parameres with apices

close together (not widely separated), not fused.

Distribution. Central and southern Northern Territory to southwestern

Queensland, Australia (Fig. 3).

Australian Entomologist, 2007, 34 (2) 45

Key to species of Dungoorus

1 Frontoclypeal suture a well-defined, elevated ridge (Fig. 1); clypeal apex

in males evenly rounded; mesotarsomere and metatarsomere 1-4 in males

compact, as wide as long. Queensland .............. D. murrumbullus Carne

- Frontoclypeal suture absent except at lateral margins (Fig. 2); clypeal

apex in males bidentate, teeth at lateral edges; mesotarsomere and

metatarsomere 1-4 in males strongly compact, wider than long. Northern

Territory: anA te sete cir ee tee rea a ey as ie atest D. frater sp. n.

Figs 1-2. Dungoorus spp. (1) Dungoorus murrumbullus head and pronotum; (2)

Dungoorus frater head and pronotum.

46 Australian Entomologist, 2007, 34 (2)

Fig. 3. Known distribution of D. frater (white circles) and D. murrumbullus (black

circle).

Dungoorus murrumbullus Carne, 1958

(Figs 1, 3)

Dungoorus murrumbullus Carne, 1958: 219 [original description]; Machatschke,

1965: 47 [catalogue listing]; Machatschke, 1972: 297 [catalogue listing]; Cassis

and Weir, 1992: 375 [catalogue listing]; Smith, 2003: 199 [checklist].

Type material examined. Holotype ©, labelled: (a) ‘Type’ (round label with red

border, typeface); (b) ‘Lea has not 13-12-18’ (typeface and handwritten); (c) ‘Not in

coll. of S. Aus. Museum’ (typeface); (d) ‘QUEENSLAND MUSEUM’ (pink label,

typeface); (e) ‘Prob. n. g. near Saulostomus Claws uneven Curious palpi’

(handwritten); (f) “Holotype of Dungoorus murrumbullus, sp.nov. P.B. Carne det.,

1957’ (handwritten and typeface); (g) ‘DUNGOORUS MURRUMBULLUS CARNE,

1958 HOLOTYPE ©” (red label, handwritten and typeface); (h) ‘QM Reg. No.

T.5533’ (handwritten) [in QMBA]. Carne (1958) indicated that this species was

described using a single specimen. He speculated that the specimen was collected in

Queensland, based on the labels and his experience dealing with Queensland

specimens examined by A.M. Lea. Type locality: Queensland, Australia.

Other material examined. QUEENSLAND: 1 0%, Milroy (30 km N Quilpie; 26° 03'S,

144° 21'E) [in QMBA].

Australian Entomologist, 2007, 34 (2) 47

Diagnosis. Male (n = 2). Length 11.7-12.2 mm, width 6.8-7.5 mm. Head:

Frons flat apically. Frontoclypeal suture complete, weakly elevated ridge.

Clypeus concave, apex evenly rounded without lateral teeth. Legs:

Tarsomeres 1-4 thickened, width approximately equal to length, cup-shaped.

Female unknown.

Distribution (Fig. 3). Southwestern Queensland, based on the holotype and

one additional specimen in QMBA from Milroy, near Quilpie.

Temporal data. November (1).

Dungoorus frater sp. n.

(Figs 2-3)

Types. Holotype &', NORTHERN TERRITORY: labelled ‘Ti-Tree N.T. 30/12/1982

D.P. Carne At light’ (handwritten) and ‘Dungoorus murrumbullus Carne m det. T.A.

Weir 1991’ (handwritten and typeface) [in ANIC]. Paratypes: 2 99 [including

allotype], same data as holotype except with ‘fP instead of ‘m’ on the second label [in

ANIC]; 12 00%, 5 29, labelled ‘AUSTRALIA: Northern Territory, 11 mi ne Yambah,

nr Alice Springs, 625 m. 29 October 1962’ (typeface) and ‘Collectors: E.S. Ross D.Q.

Cavagnaro’ (typeface) [9 00", 2 9° in CASC; 1 07, 1 2 each in QMBA, CMNC and

ABTS]; 1 0%, labelled ‘burrowing in soil below cattle dung CADNEY BORE 40 km

NW ALICE SPRINGS 6-xii-1975 G. Griffin.’ (handwritten), ‘Dungoorus sp. DET. T.

WEIR 1977’ (handwritten), and ‘Dungoorus murrumbullus Carne P.B. Carne det.,

1977’ (handwritten and typeface) [in ANIC]. Type locality: Ti Tree, Northern

Territory, Australia (22° 08'S, 133° 16'E).

Description. Male [holotype]. Length 14.8 mm, width 8.1 mm. Colour

yellowish-brown. Body ovate, convex. Head: Dorsal surface with depression

from clypeus to apex of frons; depression setose with erect, golden-brown

setae. Frontoclypeal suture absent except laterally. Clypeal apex strongly

reflexed, bidentate with lateral teeth. Terminal maxillary palpomere greatly

enlarged with elongate sulcus, approximately as long as antennal club.

Terminal labial palpomere enlarged with elongate sulcus, less than half as

long as terminal maxillary palpomere. Pronotum: Widest near apex; disc

glabrous, moderately punctate. Elytron: Surface glabrous; longitudinal striae

poorly defined, densely punctate; without membranous border. Suture

apically rounded, without acute spine or projection. Pygidium: Surface

convex, smooth, setose apically; setae erect, golden brown. Legs: Protibia

with 3 subequal teeth. Mesotibia and metatibia robust, thickest medially and

apically. Tarsomeres 2-4 greatly thickened, wider than long, cup-shaped.

Protarsomeres 3-4 with weak internoapical stridulatory ridge. Protarsomere 5

with weak internomedial stridulatory surface. Modified tarsal claws

thickened and elongate when compared with other claw, without teeth or

bifurcations. Male genitalia: Phallobase slightly longer than length of

paramere. Paramere with elevated apical and apicolateral margins; apex

strongly deflexed.

48 Australian Entomologist, 2007, 34 (2)

Female [allotype]. Length 14.8 mm, width 8.2 mm. As holotype except in the

following respects. Head: Clypeal apex strongly reflexed, evenly rounded

without lateral teeth. Terminal maxillary palpomere not greatly enlarged,

without sulcus. Terminal labial palpomere not enlarged, without sulcus. Legs:

Tarsomeres 2-4 thickened, length approximately equal to width.

Protarsomeres 3-4 and protarsomere 5 without stridulatory surfaces. Tarsal

claws not modified or thickened; paired claws weakly asymmetrical.

Variation. Male (n = 13). Length 12.4-14.8 mm, width 7.4-8.2 mm. Female

(n= 6). Length 12.8-14.8 mm, width 6.9-8.2 mm. The paratypes do not differ

significantly from the holotype and allotype.

Etymology. Frater is the Latin word for brother and this species has a very

similar gestalt to D. murrumbullus. According to Carne (1958), Dungoorus is

derived from the aboriginal word meaning ‘a stranger.’ If the first species

described in this genus is a stranger, then this second species must be that

stranger’s brother!

Distribution (Fig. 3). Known from the following localities in southern

Northern Territory, Australia: Cadney Bore (40 km NW Alice Springs; 23°

29'S, 133° 31'E), Ti Tree (22° 08'S, 133° 16'E) and Yambah (17.6 km NE; 23°

07'S, 133° 49'E).

Temporal data. October (17), December (4).

Acknowledgements

I thank David Kavanaugh (CASC) and Tom Weir (ANIC) for the loan of

specimens. I also greatly appreciate the encouragement and assistance of

Geoff Monteith and Peter Allsopp in studying Australian scarabs. This

publication was supported, in part, by an NSF/BS&I grant (DEB-0342189)

and an NSERC-PDF award to the author.

References

CARNE, P.B. 1958. A review of the Australian Rutelinae (Coleoptera: Scarabaeidae). Australian

Journal of Zoology 6(2): 162-240.

CASSIS, G. and WEIR, T.A. 1992. Rutelinae, pp 359-382, in: Houston, W.W.K. (ed.),

Zoological Catalogue of Australia. Vol. 9, Coleoptera: Scarabaeoidea. Australian Government

Printing Services Press, Canberra; 544 pp.

MACHATSCHKE, J.W. 1965. Coleoptera Lamellicornia. fam. Scarabaeidae, subfam. Rutelinae,

section Rutelinae Orthochilidae. Genera Insectorum 199°: 1-145,

MACHATSCHKE, J.W. 1972. Scarabaeoidea: Melolonthidae, Rutelinae. Coleopterum

Catalogus Supplementa 66(1): 1-361.

SMITH, A.B.T. 2003. A monographic revision of the genus Platycoelia Dejean (Coleoptera:

Scarabaeidae: Rutelinae: Anoplognathini). Bulletin of the University of Nebraska State Museum

15: 1-202.

Australian Entomologist, 2007, 34 (2): 49-50 49

THE IDENTITY OF ‘TRYPETA’ NIGRICANS WIEDEMANN

(DIPTERA: TEPHRITIDAE: TEPHRITINAE)

D.L. HANCOCK

PO Box 2464, Cairns, Qld 4870

Abstract

Trypeta nigricans Wiedemann, described from an unknown locality, is confirmed as a species of

Metasphenisca Hendel and the Indian species Metasphenisca bifaria (Munro) is placed as a new

synonym of it.

Introduction

Trypeta nigricans Wiedemann was described from an unknown locality

(Wiedemann 1830). Bezzi (1913) suggested it might belong in Acidia

Robineau-Desvoidy or Aciura Robineau-Desvoidy. Hardy (1968) examined

the types (in Naturhistorisches Museum, Vienna [NHMV]), provisionally

referred it to the genus Metasphenisca Hendel and suggested a possible

Afrotropical origin. This was followed by Norrbom ef al. (1999).

Metasphenisca is an Afrotropical-Oriental genus currently included in the

tephritine tribe Tephrellini (Hancock 1990).

Through the kindness of Peter Sehnal (NHMV), I have examined

photographs of the lectotype female. Although both wings are damaged,

sufficient remains to leave no doubt that T. nigricans is the same species as

Metasphenisca bifaria (Munro), described from southern India (Munro 1947)

and only known from there.

Metasphenisca nigricans (Wiedemann)

Trypeta nigricans Wiedemann, 1830: 509. (Unknown locality).

Isoconia bifaria Munro, 1947: 111. (Coimbatore, India). Syn. n.

Metaspheniscus ? nigricans: Hardy, 1968: 147.

Metasphenisca bifaria: Hancock, 1990: 45.

Comments. Since its original description, the name Trypeta [or

Metasphenisca] nigricans appears to have been used only in a list of names

(Bezzi 1913), a catalogue of types (Hardy 1968) and a catalogue of World

names (Norrbom et al. 1999), No additional specimens have been referred to

it and it was not mentioned in any recent regional study or catalogue for

either the Indian (e.g. Kapoor 1993, Agarwal and Sueyoshi 2005) or

Afrotropical (e.g. Cogan and Munro 1980, Hancock 1990, 1991) faunas,

unlike the currently used name M. bifaria. However, the Rules of the

International Commission on Zoological Nomenclature (ICZN 1999),

regarding automatic departure from the Principle of Priority for unused

names proposed prior to 1900, do not appear to be fully applicable, with

insufficient useage of the name M. bifaria. Accordingly, M. nigricans

(Wiedemann) is accepted here as the senior and valid name, with M. bifaria

(Munro) placed as a new synonym.

50 Australian Entomologist, 2007, 34 (2)

The wing pattern of this species is distinctive (Munro 1947), particularly the

size and orientation of the two elongate, posterior hyaline indentations.

Metasphenisca species normally have 3 pairs of frontal setae but Hardy

(1968) recorded 5 pairs in the types of M. nigricans and Munro (1947: fig.

12) showed 4 pairs in his illustration of the head of M. bifaria, suggesting

that the number of frontal setae in this species is variable.

Distribution. Known only from southern India (Tamil Nadu).

Host plant. Pods of Barleria sp. (Acanthaceae) (Munro 1947).

Acknowledgements

I thank Peter Sehnal (Naturhistorisches Museum, Vienna) and Bernhard Merz

(Muséum d’Histoire Naturelle, Geneva) for their help in obtaining

information on the type of 7. nigricans.

References

AGARWAL, M.L. and SUEYOSHI, M. 2005. Catalogue of Indian fruit flies (Diptera:

Tephritidae). Oriental Insects 39: 371-433.

BEZZI, M. 1913. Indian trypaneids (fruit flies) in the collection of the Indian Museum, Calcutta.

Memoirs of the Indian Museum 3: 53-175.

COGAN, B.H. and MUNRO, H.K. 1980. Family Tephritidae. Pp 518-554, in: Crosskey, R.W.

(ed.), Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History),

London; 1437 pp.

HANCOCK, D.L. 1990. Notes on the Tephrellini-Aciurini (Diptera: Tephritidae), with a

checklist of the Zimbabwe species. Transactions of the Zimbabwe Scientific Association 64(5):

41-48.

HANCOCK, D.L. 1991. Tephrellini (Diptera: Tephritidae: Tephritinae) from Madagascar.

Journal of the Entomological Society of Southern Africa 54(2): 173-184.

HARDY, D.E. 1968. The fruit fly types in the Naturhistorisches Museum, Wien (Tephritidae-

Diptera). Annalen des Naturhistorisches Museums in Wien 72: 107-155.

ICZN (International Commission on Zoological Nomenclature). 1999. International Code of

Zoological Nomenclature. Fourth edition adopted by the International Union of Biological

Sciences. The International Trust for Zoological Nomenclature, London.

KAPOOR, V.C. 1993. Indian fruit flies (Insecta: Diptera: Tephritidae). International Science

Publisher, New York; viii + 228 pp.

MUNRO, H.K. 1947. African Trypetidae (Diptera). A review of the transition genera between

Tephritinae and Trypetinae, with a preliminary study of the male terminalia. Memoirs of the

Entomological Society of Southern Africa 1: i-viii, 1-284.

NORRBOM, A.L., CARROLL, L.E., THOMPSON, F.C., WHITE, I.M. and FREIDBERG, A.

1999. Systematic database of names. Pp 65-251, in: Thompson, F.C. (ed.), Fruit fly expert

identification system and systematic information database. Myia 9, Backhuys Publishers, Leiden;

ix + 524 pp.

WIEDEMANN, C.R.W. 1830. Aussereuropaische zweiflugelige Insekten. Vol. 2. Schulz,

Hamm; xii + 684 pp.

Australian Entomologist, 2007, 34 (2): 51-55 51

THE IMMATURE STAGES OF CEPHRENES MOSELEYI (BUTLER)

(LEPIDOPTERA: HESPERIIDAE) FROM TORRES STRAIT,

QUEENSLAND

TREVOR A. LAMBKIN

Queensland Department of Primary Industries and Fisheries, 665 Fairfield Road, Yeerongpilly,

Qld 4105. Email: Trevor.Lambkin@dpi.qld.gov.au

Abstract

The final instar larva and pupa of Cephrenes moseleyi (Butler) are described and illustrated from

two specimens collected on Dauan Island, Torres Strait, Queensland and compared with the

immature stages of C. trichopepla with which it occurs. Overall, on Dauan Island, larvae of C.

moseleyi were encountered much less frequently than those of C. trichopepla (2:17). The host

plant is Cocos nucifera Linnaeus (coconut palm), with juvenile palms found to be preferred by

larvae of both Cephrenes species. The rearing of these two specimens from Dauan Island,

together with another reared from Saibai Island in 2001, confirms the species’ establishment in

Australia.

Introduction

Cephrenes Waterhouse & Lyell is an Indo-Australian genus of skipper

butterflies, with three species known from Australia (Braby 2000, Lambkin

and Knight 2004). The larvae of the Australian species all feed on palms

(Arecaceae) (Braby 2000, Lambkin and Knight 2004). Although

predominately tropical, C. trichopepla (Lower) and C. augiades (C. Felder)

are widespread along coastal Australia (Braby 2000), with C. trichopepla

(Lower) also known from central Australia (Braby 2000). The invasion of

these two species into new areas in recent years has been significantly aided

by their accidental introductions on cultivated palms (Braby 2000). The third

species, C. moseleyi (Butler) is known in Australia from only two islands in

the northern sector of Torres Strait, Queensland (Lambkin and Knight 2004).

Elsewhere, it occurs from the Moluccas, Aru and Kai Islands in Indonesia to

New Britain and Bougainville, including mainland New Guinea and

surrounding islands.

In Torres Strait, Cephrenes trichopepla occurs commonly, where it is known

from almost all inhabited islands (unpublished data). In this region, larvae of

C. trichopepla almost exclusively occur on Cocos nucifera Linnaeus

(coconut palm). Oddly, despite its frequency in Torres Strait, Parsons (1998)

found it uncommon on mainland New Guinea, but assumed that it was

possibly widespread throughout much of the lowlands. Conversely, C.

augiades, which in Torres Strait is only occasional locally and is much more

rarely observed than C. trichopepla, is probably restricted to the islands in the

lower half of the strait, namely Thursday, Prince of Wales, Moa and Badu

(Mathew 1885, Waterhouse and Lyell 1914, Valentine and Johnson 1993,

Braby 2000, collection records of A.I. Knight, T.A. Lambkin, C.G. Miller

and G.B. Monteith).

52 Australian Entomologist, 2007, 34 (2)

Figs 1-7. Cephrenes moseleyi (Butler). (1-3) larval head capsule: (1) frontal view,

width = 3.0 mm; (2) lateral view; (3) dorsal view. (4-5) final instar larva: (4) lateral

view, length = 45 mm; (5) dorsal view, length = 43 mm. (6-7) pupa, length = 25 mm:

(6) lateral view; (7) ventral view.

Australian Entomologist, 2007, 34 (2) 53

The life histories and larval hosts of C. trichopepla and C. augiades on the

Australian mainland are well known (Dunn 1993, 1994, 1995, Lyons 1999,

Braby 2000), including the final instar larval head capsule patterns, which are

particularly diagnostic in identifying larvae of both species (Dunn 1993,

Braby 2000). The mature larva and pupa of C. moseleyi were illustrated and

briefly described by Parsons (1998), from material originating from Bulolo,

Papua New Guinea.

During two field trips to Dauan Island, Torres Strait in 2004 and 2006, a

number of Cephrenes larvae were collected from coconut and golden cane

palms (Dypsis lutescens [H. Wendl.] H. Beentje & J. Dransfield) and reared

to adults. The predominant species was found to be C. trichopepla (n = 17),

with only two larvae of C. moseleyi collected. These two, together with a

specimen reared from Saibai Island (Lambkin and Knight 2004), confirm its

establishment in Australia. In this work, the final instar larva, head capsule

and pupa of C. moseleyi are illustrated and described in detail for the first

time from Australian material and compared with C. trichopepla.

Immature stages of C. moseleyi

Final instar larva (Figs 1-5). 43-45 mm long. Head capsule fawn in colour

with a granulated surface and four conspicuous, reddish-brown to black

vertical, almost parallel stripes consisting of a submedial and lateral stripe on

each side of the head capsule joining at either side of the mandibles; frons

with a short, centrally placed, thin brown vertical stripe immediately above

mandibles. Body elongate, translucent, pale green to yellowish-green with a

darker middorsal line extending from 4th abdominal segment to the anal

plate; spiracles white, joined by a dull translucent white line; legs and

prolegs, including bases, the same colour as body; anal plate granulated with

prominent setae along its outer perimeter.

Pupa (Figs 6-7). 23-25 mm long; elongate and slender, variable in colour,

pale yellow to yellowish-green; thorax and abdomen covered in short setae;

wing cases pale with a distinctive brown enlarged thoracic spiracle at the

base of each; a darker greenish-brown, trifid, projecting operculum covered

with prominent setae; haustellum extending to 5th abdominal segment; tip of

haustellum and antennae pale brown; cremaster rounded, with crenulate

flange covered with posteriorly pointed spines.

Life history

The habits of the larva of C. moseleyi are similar to those of other Cephrenes

spp. The larva constructs a silk-lined shelter on the host plant, in which it

finally pupates. The surface of the pupa is covered in a light dusting of white,

waxy powder. The three larvae of C. moseleyi encountered thus far in

Australia, from Dauan and Saibai Islands (Lambkin and Knight 2004), were

all collected from juvenile coconut palms growing in sand not far from the

water’s edge. In contrast, larvae of C. trichopepla, although found

54 Australian Entomologist, 2007, 34 (2)

exclusively on juvenile palms on Dauan, also occurred on coconut and

golden cane palms away from the beach. Despite larvae of both species

occurring on the same host species near the water’s edge, it is not known

whether they occur together on individual host plants. In general, adults of C.

moseleyi were not often observed and, when observed, preferred to rest for

long periods on or near their host plants (Lambkin and Knight 2004), as is the

habit of the other two Australian Cephrenes spp (Dunn 1993, Lyons 1999).

On Dauan, adults have also been collected from Melaleuca blossom.

Discussion

Final instar larvae of C. moseleyi and C. trichopepla closely resemble each

other, but can be separated primarily by the different patterns on the head

capsules. The pattern of vertical striping on the head capsule of C.

trichopepla consists of two lateral black stripes, with two frontal black stripes

running both sides of the frons, these stripes becoming laterally thicker just

above the frons and thicker again closer to the mandibles. There is an orange-

yellow patch on the cheeks either side of the mandibles at the base of the

frontal stripes. Dunn (1993) provided very useful line drawings of the typical

head pattern of both this species and C. augiades. The pattern of striping on

the head capsule of C. moseleyi (Figs 1-3) is distinctly different from that of

C. trichopepla and consists of two lateral and two frontal, almost parallel

stripes which tend to be reddish-brown to black instead of the typical black

stripes of C. trichopepla. There are no orange-yellow patches on the cheeks

either side of the mandibles in C. moseleyi, with the two stripes either side of

the head capsule joining at the base of the head. In addition, the ground

colour of the head capsule of C. moseleyi is fawn while that of C. trichopepla

is white. Larvae of C. moseleyi (Figs 4-5) are more elongate than those of C.

trichopepla, are more yellow in colour and have the spiracles white rather

than brown.

Pupae of the two species also differ. That of C. moseleyi (Figs 6-7) tends to

be pale yellow to yellowish-green in colour, as opposed to the dirty-green to

brown colour of C. trichopepla; it is also smoother in appearance than that of

C. trichopepla and has a characteristic trifid, projecting operculum as

opposed to the very dark, blunt and rounded operculum of C. trichopepla.

The lengths of the haustella are also different (Parsons 1998), that of C.

moseleyi extending to and just reaching the Sth abdominal segment while in

C. trichopepla the haustellum reaches the 6th abdominal segment.

On Dauan, C. moseleyi is generally encountered less frequently than C.

trichopepla. The collection of only two larvae of C. moseleyi during two field

trips to Dauan, together with the paucity of adult Australian material known

in collections (now about 10 specimens) attests to its overall rarity on Dauan

and Saibai Islands. Parsons (1998) also reported the wide-ranging rarity of

this species in Papua New Guinea but did indicate that it can be occasional

locally.

Australian Entomologist, 2007, 34 (2) 55

Acknowledgements

I thank the Dauan Island community council for its cooperation during the

periods of fieldwork spent on Dauan Island, and C.G. Miller and G.B.

Monteith for providing collection records. J.S. Bartlett gave valuable

assistance by preparing the colour plate.

References

BRABY, M.F. 2000. Butterflies of Australia; their identification, biology and distribution.

CSIRO Publishing, Collingwood, Victoria; xx + 976 pp.

DUNN, K.L. 1993. Notes on the biology and larval hosts of Cephrenes (Lepidoptera:

Hesperiidae: Hesperiinae). Victorian Entomologist 23(5): 97-110.

DUNN, K.L. 1994. Oviposition and territorial behaviour in Cephrenes trichopepla (Lower)

(Lepidoptera: Hesperiidae: Hesperiinae), and a new distribution record. Victorian Entomologist

24(1): 21-25.

DUNN, K.L. 1995. Notes on the biology and larval hosts of Cephrenes (Lepidoptera:

Hesperiidae) — Part II. Victorian Entomologist 25(1): 3-12.

LAMBKIN, T.A. and KNIGHT, A.I. 2004. The first Australian record of Cephrenes moseleyi

(Butler) (Lepidoptera: Hesperiidae) from Torres Strait, Queensland. Australian Entomologist

31(3): 107-109.

LYONS, K.A. 1999. The palm darts, Cephrenes augiades and C. trichopepla (Hesperiidae). Pp

105-114, in: Kitching, R.L., Scheermeyer, E., Jones, R.E. and Pierce, N.E. (eds), Biology of

Australian butterflies. Monographs on Australian Lepidoptera, Volume 6. CSIRO Publishing,

Collingwood, Victoria; xvi + 395 pp.

MATHEW, G.F. 1885. An afternoon among the butterflies of Thursday Island. Proceedings of

the Linnean Society of New South Wales 10(2): 259-266.

PARSONS, M.J. 1998. The butterflies of Papua New Guinea: their systematics and biology.

Academic Press, London; xvi + 736 pp, xxvi + 136 pls.

VALENTINE, P.S. and JOHNSON, S.J. 1993. The butterflies of Moa Island, Torres Strait.

Victorian Entomologist 23(6): 116-121.

WATERHOUSE, G.A. and LYELL, G. 1914. The butterflies of Australia. Angus and Robertson,

Sydney; vi + 239 pp.

56 Australian Entomologist, 2007, 34 (2)

BOOK REVIEW

Dragonflies of Peninsular Malaysia and Singapore. A Pocket Guide, by A.G.

Orr. Natural History Publications (Borneo), Kota Kinabalu. 2005; vi + 127

pp; soft covers; illustrated throughout. ISBN 983-812-103-7.

This pocket guide to the odonates of Peninsular Malaysia and Singapore is

the first to cover this region and is ideally suited to its purpose. Attractively

presented with colour paintings on every page, it will be useful to beginner

and expert alike. Of the 230 species recorded, 98.7% are discussed and their

adult males illustrated. Small differences between similar species are

depicted clearly by additional illustrations, as well as by details of the male

appendages where needed. A representative selection of larvae is shown in

line drawings. Anyone familiar with the author’s previous award-winning

publication on the dragonflies of Borneo [see Australian Entomologist 31: 4

(2004)] will recognise the quality of his artwork, which in this instance

renders the use of complex keys in the field superfluous.

The species accounts are necessarily brief but include details of size,

distinguishing characters, habitat, general distribution and notes on behaviour

and larvae. The book is a model of conciseness, including a general

introduction, a checklist and a guide to further reading, in addition to the

species accounts - all of which is encompassed within 127 pages.

It is interesting to note that there are only 88 species of Zygoptera recorded,

against 141 of Anisoptera. This ratio is very different from the typical ratio in

other countries, which is closer to 1:1. Could this be due to the large number

(75 species) of libellulids — which include strong fliers and often migrants, or

to the possibility that there are still a lot of Zygoptera out there awaiting

discovery?

As a stimulus to the study of Malaysian and Singaporean species, this little

book will surely encourage the discovery of more species and will be

welcomed by all dragonfly enthusiasts. It could serve well as a model for

similar treatments in other geographical regions and I, for one, would look

forward to such an advance.

J.N. Yates

Imbil

Australian Entomologist, 2007, 34 (2): 57 57

A NEW SYNONYM AND A NEW COMBINATION IN THE FRUIT

FLY TRIBE PLIOMELAENINI (DIPTERA: TEPHRITIDAE:

TEPHRITINAE)

D.L. HANCOCK

PO Box 2464, Cairns, Qld 4870

Abstract

Pliomelaena udhampurensis Agarwal & Kapoor is placed as a new synonym of P.

spathuliniforma (Dirlbek & Dirlbek), comb. n. [transferred from Metasphenisca Hendel]. The

species is known from NW India and NE Afghanistan.

Introduction

The fruit fly tribe Pliomelaenini contains several Indo-Australian species

associated with the flowers of Acanthaceae. They were reviewed recently by

Hancock (2004) but continued investigation has revealed a new synonym and

a new combination within the group.

In addition, a second male of the little known species Quadrimelaena

quadrimaculata (Agarwal & Kapoor) has been located in The Natural

History Museum, London. It is labelled ‘India, W.W. Saunders’ and ‘prisca

Walker’, an apparent manuscript name. As expected, vein R, has a gap in the

row of setae opposite the end of vein Sc.

Pliomelaena spathuliniforma (Dirlbek & Dirlbek), comb. n.

Metasphenisca spathuliniforme Dirlbek & Dirlbek, 1968: 175. (Darunta & Laghman,

Nengrahar Province, NE Afghanistan).

Pliomelaena udhampurensis Agarwal & Kapoor, 1988: 119. (Udhampur, Jammu and

Kashmir, NW India). Syn. n.

Comments. Although described in genus Metasphenisca Hendel (tribe

Tephrellini), P. spathuliniforma clearly belongs in genus Pliomelaena Bezzi

in tribe Pliomelaenini. Descriptions and illustrations of both taxa listed above

(Dirlbek and Dirlbek 1968, Agarwal and Kapoor 1988) leave no doubt that

they are conspecific. This species is known from NE Afghanistan and NW

India. For further discussion see Hancock (2004),

Acknowledgement

I thank Nigel Wyatt (NHM, London) for access to material in his care.

References

AGARWAL, M.L. and KAPOOR, V.C. 1988. Four new species of fruit flies (Diptera:

Tephritidae: Tephritini) together with redescription of Trupanea inaequabilis Hering and their

distribution in India. Journal of Entomological Research (New Delhi) 12: 117-128.

DIRLBEK, J. and DIRLBEK, K. 1968. Beiträge zur Kenntnis der Fauna Afghanistans.

Trypetidae, Diptera. Casopis Moravskeho Musea, Vedy Prirodni (Acta Musei Moraviae,

Scientiae Naturales) 53 (Supplement): 173-180.

HANCOCK, D.L. 2004. A review of the fruit fly tribe Pliomelaenini (Diptera: Tephritidae:

Tephritinae) in the Indo-Australian Region. Australian Entomologist 31(3): 133-136.

58 Australian Entomologist, 2007, 34 (2): 58-60

BOOK REVIEW

The Smaller Majority: The Hidden World of the Animals that Dominate the

Tropics, by Piotr Naskrecki. Harvard University Press, Cambridge,

Massachussets & London, UK. 2005; x + 278 pp; hardback. ISBN 0-674-

01915-6. Price US$35.

"Invertebrates and other small animals play crucial roles in ecosystem health, including

soil production, water filtration, pollination and provision of food for the other, larger

vertebrates. In short, invertebrates make the Earth a liveable planet, yet insects and

other invertebrates are disappearing from the globe at a rate faster than their larger

vertebrate cousins, particularly in tropical regions. We do not know the precise rate of

loss (nor for sure which species have gone), due to uncertainties in taxonomic

composition and number and the general lack of baseline data on their distributions,

but we can be fairly certain that the current extinction rate — the sixth (human-

induced) mass extinction event — is faster than anything the planet has experienced in

the past. The reason is simple: most tropical invertebrates have narrow, specialised

niches, but the vast tropical forests of central West Africa, SE Asia and South

America in which they live are disappearing at a phenomenal rate. Each species lost is

another chapter of the genetic diversity and evolutionary history destroyed forever.

Moreover, most animals are small and live in tropical latitudes, yet most people do

not notice small animals and live in temperate latitudes, compounding the problem.

So what is the solution? A key first step is the need for better education and

conservation advocacy, promoting the importance and popularity of invertebrates and

the need to conserve the habitats in which they live. The Smaller Majority fills this

critical need. The author, Piotr Naskrecki, is Director of the Invertebrate Discovery

Initiative of Conservation International, and has provided a compelling tool to

promote invertebrates and their urgent need for conservation. His message is simple.

If the general public can understand and appreciate the beauty and ecological

importance of invertebrates, they are more likely to care for them; and caring is the

key to their long-term preservation.

The purpose of the book is therefore to celebrate everything that is small and

misunderstood in the natural biological world — to understand and notice them, and to

highlight the enormous diversity of life found right under our very feet. This

ambitious goal is admirably achieved with the presentation of a comprehensive

collection of more than 400 stunning photographs from the tropical areas of the world,

particularly Central America and Africa but also Australia and, to a lesser extent, the

Solomon Islands and Madagascar, augmented with an informative text. It thus covers

insects and other terrestrial invertebrates, together with small amphibians and reptiles,

which Naskrecki collectively refers to as the ‘smaller majority’. Giver that nearly all

animals on Earth are small and largely ‘unseen’, there is clearly a limit to how much

attention can be allocated to each group. As Naskrecki notes [on page 3] ‘It does not

pretend to be an exhaustive overview of tropical biota, and its taxonomic coverage is

fragmentary. Each page provides only a glimpse into an animal’s world rather than a

comprehensive account of its life cycle’.

The book focuses on three major terrestrial ecosystems: humid forests, savannas and

deserts. These three biomes are not treated equally, with 174 pages (72%) devoted to

humid forests compared with 46 pages (19%) to savannas and 22 pages (9%) to

Australian Entomologist, 2007, 34 (2) 59

deserts: the bias in coverage reflects differences in species richness between each

biome rather than their spatial representation on the globe. A prologue sets out how

the author discovered the smaller majority, and why it is important to understand and

conserve invertebrates. The three major biomes follow, which make up the bulk of the

book, with attention given to threatening processes. Then follows an epilogue in

which Naskrecki highlights the taxonomic impediment of invertebrates against the

current biodiversity crises and the urgent need to document the Earth’s biological

heritage more effectively. The epilogue includes a series of images depicting several

undescribed taxa but, more importantly, we are reminded of the thrill and adventure of

discovering species new to Science. A short chapter on photographing the smaller

majority provides useful practical tips on working in the rainforest and the basic

equipment needed. A list of international organisations (almost all in USA) devoted to

conservation, acknowledgements and a species index (including both common and

scientific names) complete the work.

The photographs are masterpieces in their own right and many fill an entire page!

Particularly stunning are the pictures of Cholus cinctus (a weevil from Costa Rica: p.

15), in flight, Gasteracantha metallica (a jewel spider from the Solomon Islands: p.

108), Pseudatteria leopardina (a diurnal leafroller moth from Costa Rica: p. 147),

Uroplatus phantasticus (a Malagasy leaf-tailed gecko: p. 159), Tympanophora

uvarovi (an Australian balloon-winged katydid: p. 193) and Polyspilota aeroginosa

(an African savanna mantid: p. 222) against the sunset, to name just a few. There is

also an evocative image of two workers of Oecophylla smaragdina (green tree-ant or

weaver ant from Australia: p. 132) attending the larva of a lycaenid butterfly

(scientific name not given but clearly Hypolycaena phorbas). Most, but not all

terrestrial invertebrates mentioned are illustrated. There is a strong bias towards

katydids and allied insects, although this is probably understandable given that the

author is a world authority of the group.

The Smaller Majority is more than a compilation of first class natural history

photographs. The superb images are interwoven with text that is lucid, detailed,

scientifically accurate and easy to read. Naskrecki is one of those rare authors who

can communicate effectively to both the scientific audience and the wider general

public. He also writes with flair and passion. Thus, the illustrative material is

augmented with a considerable amount of information on natural history, ecology and

evolutionary principles, as well as personal anecdotes. For example, we learn that the

evolutionary success of weevils, the largest family of all living organisms, is probably

due to their mouthparts being able to exploit seeds and nuts of flowering plants and

the co-diversification of these plants in the Late Cretaceous. We also learn about the

unique and spectacular radiation of the Australian spur-throated grasshoppers

(Cantatopinae), and that the pointillist art style of Aboriginal paintings may have had

its origins based on the colour patterns displayed by Australian spotted pyrgomorph

grasshoppers. On pages 86-87, Naskrecki recalls his discovery of cockroaches and

moths feeding at night on the honeydew of plant-feeding fulgorids (Auchenorhyncha)

— a most unusual association in which the benefit to both parties is not entirely clear. |

was delighted to see a section devoted to the ‘heelwalkers’ — Mantophasmatodea — the

newly recognised insect order from Africa that previously were thought to have died

out in the Miocene.

I found few weaknesses with this book. A glossary would have been a useful addition,

as some of the terms used (e.g. co-existence, cryptic, diurnal, mutualism, mimicry,

60 Australian Entomologist, 2007, 34 (2)

parasite) require some basic knowledge or understanding of tertiary-level biology,

although the author has attempted to qualify many of these terms in the text; for

example: ‘A butterfly pupa, also known as the chrysalis, looks like a beautiful

sculpture’ [p. 144]. The index is unworkable; for instance, all ant species are listed

under ‘ants’ but not under Hymenoptera or Formicidae, but all moths and butterflies

are listed under ‘Lepidoptera’. As a result, it is difficult to trace entries; for example,

there is no separate entry for Oecophylla smaragdina under ‘O° or ‘S’, nor under ‘H’

for Hymenoptera, ‘F’ for Formicidae or ‘W?’ for weaver ant, and there is no separate

entry for Eurema hecabe under ‘E’ or ‘H’, or under ‘S’ for sulphur butterfly. As it

stands, the index assumes the reader knows the correct systematic placement of these

and all other taxa. This is an unreasonable assumption given the audience for which

this book id intended.

Despite these very minor shortcomings, The Smaller Majority is a landmark

publication bringing together superb natural history, macro-photography, biological

science and conservation concerns: it serves to promote both the importance of insects

and the conservation of tropical habitats in which they live. The wide scope and

general knowledge that Naskrecki has brought together in a single work are truly

breathtaking. If you love insects or close-up photography, or desire to learn more

about those ‘far off tropical places, then this book is for you. It will especially appeal

to youngsters or anyone with a fascination of our natural world. I am confident that

Naskrecki’s dream ‘that the images in this book will reinforce a child’s interest in the

natural life of caterpillars or frogs, or perhaps they will awaken a long-forgotten

fascination with small creatures in an older reader’ will be fulfilled. | am also

confident that it will inspire the next generation of invertebrate conservation

biologists. In summary, this work is a must buy!

M.F. Braby

Invertebrate Conservation Biologist at the Biodiversity Conservation Division,

Department of Natural Resources, Environment and the Arts, PO Box 496,

Palmerston, NT 0831 and Visiting Fellow at the School of Botany and Zoology, The

Australian National University, Canberra, ACT 0200.

Australian Entomologist, 2007, 34 (2): 61-62 61

BOOK REVIEW

Of Peaches and Maggots: The Story of Queensland Fruit Fly, by A.C.

Courtice. Hillside Books, Marengo, NSW. 2006; 336 pp; softback. ISBN 0-

9588239-0-1. Available from Hillside Books, 1187 Marengo Road, Marengo

via Dorrigo, NSW 2453. Price A$34.50 (including postage).

This is an entertaining book, easy to read and well researched. Intended for non-

professionals, it nevertheless has a much wider appeal. It entwines a history of fruit

fly outbreaks in Australia and elsewhere, from the 1880s onwards (in particular the

Toowoomba outbreak of 1885), with the author’s own efforts to understand why they

happened. It breathes life (in a romanticised way) into the minds and actions of many

of the early entomologists involved with fruit fly research or control, including the

familiar names of Henry Tryon, Walter Froggatt and Hubert Jarvis. Some are

criticised and others praised, but the difficulties all these early workers faced are ably

documented. Along the way some myths are debunked — for example, the history of

the 1853 ‘Kiama’ fly, long held to be one of the earliest records of Queensland fruit

fly (Bactrocera tryoni) south of northern New South Wales but with no actual

specimens in existence, has more than one intriguing twist. Some of the underlying

causes attributed to many of the early outbreaks [and to some even earlier plant

diseases, such as that resulting in the Irish ‘potato famine’] are entirely unexpected.

I found particularly enjoyable, and enlightening, the tales of Dagobert Daldorf in India

and Walter Froggatt in Australia — both of which help shed light on the vexed and

enigmatic question [see below] of Mediterranean fruit fly, or Medfly (Ceratitis

capitata) in India. Equally enjoyable are the discussions of Medfly’s first detection in

London in 1822 and the Queensland-New South Wales ‘banana wars’. Noteworthy,

too, is the well founded assertion that Medfly was present in Sydney several years

before the 1898 date normally quoted and well before the arrival of Queensland fruit

fly, which subsequently displaced it. In fact there is much of interest in this book, not

just for those interested or involved in the study of fruit fly outbreaks, past or present.

There are some minor factual errors and some of a more serious and unnecessary

nature, the latter possibly related to the author’s apparent disdain for professional

entomologists working in government departments. Bactrocera tryoni and B.

neohumeralis are certainly not members of the B. dorsalis complex, which is

represented in Australia by B. cacuminata, B. opiliae and B. endiandrae. Despite its

primary use of cultivated fruits in southern Queensland and New South Wales, B.

tryoni has a very large number of native hosts in northern Queensland. The geneticist

referred to on page 153 was from Hawaii, not New Zealand and B. occipitalis is not

the only pest member of the B. dorsalis complex in the Philippines — it occurs

alongside the very similar, but distinct, B. philippinensis. The suggestion that the

eradication of B. papayae from northern Queensland was ‘staged for the benefit of

trading partners ... but could not possibly have succeeded’ is little more than a

conspiracy theory which, like others of its ilk, does not withstand scientific scrutiny.

B. papayae was eradicated due to the determined efforts, in exceptionally difficult

circumstances, of those actively involved and the effectiveness of the attractant

methyl eugenol. B. papayae has also invaded Papua New Guinea, where it remains

well established. Cue-lure is less effective and this might in part explain why B. tryoni

has not been eradicated successfully from Lord Howe Island. Another of these

62 Australian Entomologist, 2007, 34 (2)

seeming ‘conspiracy theories’ concerns the status of B. aquilonis in Darwin, implying

ulterior motives which did not, and do not, exist. ‘Ceratitis’ dentipes [now Lenophila

dentipes] belongs in family Platystomatidae and does occur in New South Wales.

The question of Mediterranean fruit fly in India is problematical. It does not occur

there now and almost certainly never did. The frequent assertion that the type

specimen of C. capitata was collected by Daldorf personally ‘along or off the coast of

Bengal’ is at odds with the specimen’s label, which states ‘in mari indico’ [in Indian

Ocean]. It is more likely that the specimen was presented to Daldorf by one of the

Danish sea-captains plying the trade routes between Copenhagen [or East Africa] and

Calcutta, many of whom, as a Royal representative, he would have known. However,

the suggested collection date (between 1798 and 1802) is likely to be correct. Since

then, the only specimens actually recorded from India are 4 females and a male from

Pusa [an Agricultural Research Station in Bihar] dated 20.viii.[19]07 and 23.ix.[19]08

but with no indication of who collected them (Munro 1938). However, there are no

reports of a Medfly outbreak in India in 1907-08 [or at any other time] and the ‘Pusa’

record is more likely to be of mislabelled reference specimens obtained from

overseas. [This does happen: specimens of Medfly in the Suriname Department of

Agriculture labelled ‘Paramaribo’ actually came from California (DLH, pers. obs.

1989)]. Bezzi (1913) noted that the only material in the Calcutta Museum at that time

were 2 specimens from Australia presented by W.W. Froggatt [presumably in 1908,

when he visited both Pusa and Calcutta]. This congruence of dates is telling and there

can be little doubt that the ‘Pusa’ specimens were also gifts from Froggatt. No one

took any notice of them until Munro received them from New Delhi. Bezzi (1909) did

not refer to them, merely stating that C. capitata was ‘recorded from India [based on

the type], where apparently comparatively rare’. Neither Froggatt himself (1909) nor

Senior-White (1924) made any mention of them and both doubted the occurrence of

Medfly in the Oriental Region. The 3 females recorded by Munro (1938) as ‘bred

from peach’ in August 1907 were most likely bred by Froggatt in Sydney where, at

that time, Medfly was abundant.

Despite the above comments and criticisms, the overall conclusions reached regarding

the history and causes of past fruit fly outbreaks and the species actually involved are

both persuasive and plausible. Whether they withstand the test of time or not remains

to be determined, but the journey there is nonetheless a fascinating read.

References

BEZZI, M. 1909. Le specie dei generi Ceratitis, Anastrepha e Dacus. Bolletino del Laboratorio

di Zoologia Generale e Agraria della Regia Scuola Superiore d'Agricoltura, Portici 3: 273-313.

BEZZI, M. 1913. Indian trypaneids (fruit flies) in the collection of the Indian Museum, Calcutta.

Memoirs of the Indian Museum 3: 53-175.

FROGGATT, W.W. 1909. Fruit flies: a general account of the flies belonging to the family

Trypetidae, that damage sound fruit. Government printer, Sydney; 56 pp, 8 pls.

MUNRO, H.K. 1938. Studies on Indian Trypetidae (Diptera). Records of the Indian Museum 40:

21-37.

SENIOR-WHITE, R. 1924. Catalogue of Indian insects. Part 4 — Trypetidae (Trypaneidae).

Government of India, Calcutta; iii + 33 pp.

D.L. Hancock `

PO Box 2464, Cairns, Qld 4870

Australian Entomologist, 2007, 34 (2) 63

RECENT ENTOMOLOGICAL LITERATURE

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2007 Systematics, biogeography and diversification of the Indo-Australian genus Delias

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2006 Jewel bugs of Australia (Insecta, Heteroptera, Scutelleridae). Pp 275-398, in: Rabitsch,

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2006 A new genus and species of Chironominae (Diptera: Chironomidae) with wood-mining

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2006 A revised classification of subgenera and species groups in Dacus Fabricius (Diptera,

Tephritidae). Pp 167-205, in Merz, B. (ed.), Phylogeny, taxonomy, and biology of

tephritoid flies (Diptera, Tephritoidea). Instrumenta Biodiversitatis Volume VII. Natural

History Museum, Geneva; 274 pp.

HUXHAM, K.A., FAY, H.A.C. and HANCOCK, D.L.

2006 Two new species and a new Australian record of Bactrocera Macquart (Diptera:

Tephritidae: Dacinae) from northern Queensland, Torres Strait and Papua New Guinea.

Australian Journal of Entomology 45(1): 34-37.

KOLESIK, P., ADAIR, R.J. and EICK, G.

2005 Nine new species of Dasineura (Diptera: Cecidomyiidae) from flowers of Australian

Acacia (Mimosaceae). Systematic Entomology 30: 454-479.

64 Australian Entomologist, 2007, 34 (2)

KOLESIK, P., GEYER, C. and MORAWETZ, W.

2006 First known gall midge (Diptera: Cecidomyiidae) from Arecaceae: Normanbyomyia

Jrructivora gen. & sp. n. damaging fruit of the black palm, Normanbya normanbyi, in

tropical Australia. Australian Journal of Entomology 45(1): 38-43.

MEATS, A.

2006 Attributes pertinent to overwintering potential do not explain why Bactrocera

neohumeralis (Hardy) (Diptera: Tephritidae) does not spread further south within the

geographical range of B. tryoni (Froggatt). Australian Journal of Entomology 45(1): 20-

25.

MONTEITH, G.B.

2006 Maternal care in Australian oncomerine shield bugs (Insecta, Heteroptera,

Tessaratomidae). Pp 1135-1152, in: Rabitsch, W. (ed.), Hug the bug — for love of true

bugs. Fetschrift zum 70. Geburtstag von Ernst Heiss. Denisia 19, zugleich Kataloge der

OO. Landesmuseen Neue Serie 50.

[Includes notes on Bromocoris soefi (Distant) (Pentatomidae: Halyinae)]

MOULDS, M.S.

2005 An appraisal of the higher classification of cicadas (Hemiptera; Cicadoidea) with special

reference to the Australian fauna. Records of the Australian Museum 57: 321-446.

PARETAS-MARTINEZ, J. and PUJADE-VILLAR, J.

2006 Charipinae (Hymenoptera: Figitidae) from Australia: revision of the genus Thoreauana

Girault, 1930 and description of Dilapothor n. gen. Australian Journal of Entomology

45(3): 219-226.

PAULL, C. and AUSTIN, A.D.

2006 The hymenopteran parasitoids of light brown apple moth, Epiphyas postvittana (Walker)

(Lepidoptera: Tortricidae) in Australia. Australian Journal of Entomology 45(2): 142-156.

SINCLAIR, D.P.

2002 A generic revision of the Oncomerinae (Hemiptera: Heteroptera: Tessaratomidae).

Memoirs of the Queensland Museum 46(1): 307-329.

SMITH, G.B.

2006 New species of Metrinura Mendes (Zygentoma: Nicoletiidae) from Queensland,

Australia. Australian Journal of Entomology 45(2): 163-167.

SOTA, T., TAKAMI, Y., MONTEITH, G.B. and MOORE, B.P.

2005 Phylogeny and character evolution of endemic Australian carabid beetles of the genus

Pamborus based on mitochondrial and nuclear gene sequences. Molecular Phylogenetics

and Evolution 36: 391-404.

STEBNICKA, Z.T.

2006 Revision of the Indo-Australian genus Cnematoplatys Schmidt (Coleoptera:

Scarabaeidae: Aphodiinae: Eupariini). Australian Journal of Entomology 45(2): 168-175.

TAKAML, Y. and SOTA, T.

2006 Four new species of the Australian Pamborus Latreille (Coleoptera: Carabidae) carabid

beetles. Australian Journal of Entomology 45(1): 44-54.

WEBSTER, K.W., COOPER, C. and MOUND, L.A.

2006 Studies on Kelly’s citrus thrips, Pezothrips kellyanus (Bagnall) (Thysanoptera:

Thripidae): sex attractants, host associations and country of origin. Australian Journal of

Entomology 45(1): 67-74.

WINTERTON, S. and de FREITAS, S.

2006 Molecular phylogeny of the green lacewings (Neuroptera: Chrysopidae). Australian

Journal of Entomology 45(3): 235-243.

ENTOMOLOGICAL NOTICES

Items for insertion should be sent to the editor who reserves the right to alter,

reject or charge for notices.

NOTES FOR AUTHORS

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The Australian Entomologist

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Authors should refer to recent issues for layout and style. All papers will be

forwarded to two referees and the editor reserves the right to reject any paper

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comply with the mandatory provisions of the International Code of

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THE AUSTRALIAN

Entomologist

Volume 34, Part 2,25 May 2007

CONTENTS

HANCOCK, D.L.

Phylogeny of the troidine butterflies (Lepidoptera: Papilionidae)

revisited: are the red-bodied swallowtails monophyletic?

HANCOCK, D.L.

The identity of Trypeta’ nigricans Wiedemann (Diptera: Tephritidae:

Tephritinae).

HANCOCK, D.L.

A new synonym and a new combination in the fruit fly tribe

Pliomelaenini (Diptera: Tephritidae: Tephritinae).

LAMBKIN, T.A.

The immature stages of Cepbrenes moseleyi (Butler) (Lepidoptera:

Hesperiidae) from Torres Strait, Queensland.

SMITH, A.B.T.

Review of the genus Dungoorus Carne (Coleoptera: Scarabaeidae:

Rutelinae: Anoplognathini).

RECENT ENTOMOLOGICAL LITERATURE

BOOK REVIEWS:

Dragonflies of Peninsular Malaysia and Singapore. A pocket guide.

A.G. Orr.

The smaller majority: the hidden world of the animals that dominate

the tropics. P. Naskrecki.

Of peaches and maggots: the story of Queensland fruit fly.

A.C. Courtice.

ISSN 1320 6133