THE AUSTRALIAN ENTOMOLOGIST

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Cover: Our cover illustration for 2013 shows a species of the pebble-mimicking

genus Raniliella (Orthoptera: Acrididae). There are two species in the genus, both

restricted to desert environments of the northern parts of South Australia. They are

commonly seen in stony areas around the Flinders Ranges. Raniliella is one of

several genera of Australian grasshoppers which closely resemble the stones of our

extensive gibber deserts. The illustration is reproduced by permission from CSIRO’s

Insects of Australia and is by the late Frank Nanninga, a Dutch-born artist who was

the leading insect illustrator of the 1960s in Australia. His work reached its peak in

the eight magnificent colour plates which grace the pages of Insects of Australia.

Australian Entomologist, 2013, 40 (2): 57-64 57

NOTES ON THE FEMALE COLOUR FORMS OF SYNTHEMIS

MIRANDA SELYS, 1871 (ODONATA: SYNTHEMISTIDAE) IN NEW

CALEDONIA

M. MARINOV! and S. RICHARDS?

!School of Biological Sciences, University of Canterbury, Private Bag 4800, Christchurch 8140,

New Zealand (E-mail: milen.marinov@canterbury.ac.nz)

"Conservation International, Atherton, Qld. Current address: Department of Terrestrial

Vertebrates, Museum and Art Gallery of the Northern Territory, PO Box 4646, Darwin, NT 0801

Abstract

Two female Synthemis miranda Selys specimens with strikingly different morphological features

(mainly wing colouration) were collected in sympatry in Province Nord, New Caledonia. Both

appear to be mature adults. Specimens with these two colour patterns have previously been

reported in the literature but this paper presents the first record of their co-existence. We

formally recognise the two forms based on colour pattern and note other morphological features

that may also distinguish them. The validity of these additional characters requires confirmation

through examination of further material.

Introduction

Synthemis miranda Selys, 1871 has often been referred to as the finest, most

interesting and spectacular species of the genus Synthemis Selys (Tillyard

1910, Campion 1921, Lieftinck 1971, Davies 2002). It is also one of the most

controversial species of the genus. The type locality has been extensively

debated because, in the original description, Selys (1871) did not specify

precisely where the female holotype originated. His statement ‘Patrie:

Nouvelle-Calédonie, par feu le pére Montrouzier. (Coll. Selys.)' is also

misleading because it is unclear who should be credited with the discovery of

the species. The situation became even more complicated when Tillyard

(1910) presented an anecdote about the discovery of the species. According

to him, the holotype was spotted by Selys pinned on a lady's hat in a fashion

shop in Paris. The origin of this story is unclear, but most probably it was a

rumour swirling around at that time and Tillyard had heard it from René

Martin in Paris. However, in the two large works (Martin 1906, 1914)

preceding and following Tillyard (1910), no details were given about the

discovery of S. miranda.

Martin (1906) illustrated the wing venation and colouration of the holotype

and Martin (1914) provided only a short note about the general distribution of

the species (as ‘Nouvelle-Calédonie’). Ris (1915) and Schmidt (1938)

commented on the validity of Tillyard's (1910) anecdote, concluding that it

was erroneous and probably arose because of uncertainties about the origin of

the species in the original description. However, Davies (2002) repeated this

story, claiming that the species was named after its discovery in the shop in

Paris and *... it was many years before its actual home was discovered to be

Melanesia’. Such stories, although intriguing, do little to advance our

knowledge of this species. Although the type locality is not certain and

probably never will be, the general origin of the species (Nouvelle-

58 Australian Entomologist, 2013, 40 (2)

Calédonie) was given in the first description and its occurrence as an endemic

species there has been confirmed by other researchers.

The species’ wing colouration is the second most discussed point in the

literature. The combination of bright yellow and brown occupying a large

area of the wing gives S. miranda a striking appearance. Campion (1921) was

the first to report that the extent of this colour is variable. Among the insects

collected from New Caledonia by Mr Paul D. Montague, there were two

males and three females. This material formed the basis of the description of

the [met]allotype male and a brief introduction to variation in the extent of

wing colouration in females. These females were collected on the same date

and all had limited colour that did not extend beyond the arculus, unlike the

holotype which has substantially more extensive colour on the wings. It is

also important to note that one female was immature while the two others

were fully mature.

Lieftinck (1971) was the next author to report females of S. miranda with

reduced colour on the wings and the first who suggested that ‘... it would

appear that there are two main forms of female, which differ markedly from

one another as far as the wing colour is concerned, evidently quite

independent from the age of the individual, intermediates occurring at the

same time, though apparently more rarely'. However, no names for these

forms were proposed. He did not report the total number of specimens

investigated and, with the exception of seven specimens retained in the

Netherlands, the material was returned to the Bishop Museum in Hawaii.

Most of these seven females were collected from different locations; only two

were from the same site — Pouébo, in the northeastern part of the country.

One of those specimens had a wing colouration intermediate between that

reported for the holotype and the reduced colour, but it is unclear to which

form the second specimen from that location belongs. No further comments

on this issue were presented in two subsequent studies by the same author

dedicated to the Odonata of New Caledonia (Lieftinck 1975, 1976); he had

examined additional imagines without specifying the type of wing

colouration they possessed and regarded the wing venation (rather than

colouration) as a more important taxonomic feature (Lieftinck 1976).

Reduced extent of wing colouration was reported in all 12 females that

emerged during a larval rearing experiment (Winstanley 1983). No females

with extensive colouration similar to the holotype emerged during that study.

Davies' (2002) claim to priority of the discovery of females with reduced

wing colouration is therefore incorrect. Davies (2002) referred to these

specimens as ‘unusual style’ females but provided no specific description or a

name in his otherwise very detailed review of New Caledonian odonates.

Unfortunately, it is difficult to determine from Davies’ illustrations the full

extent of pigmentation on the wings. Apparently, he considered those females

to have an atypical colouration rather than being a distinct form, although on

Australian Entomologist, 2013, 40 (2) 59

one occasion he observed many ‘... properly mature ...' individuals with this

kind of colouration flying together with males. Formal description of the two

female types has never been suggested in the literature and the most likely

explanation is the fact that so far both types have never been collected from

the same site.

A recent biodiversity survey in New Caledonia documented females with

both types of wing colouration at the same location. This appears to be the

first record of sympatry reported for these types and we believe it will be

useful for future workers to describe and formally recognise them as distinct

forms. Our morphological analysis also hinted that additional features may

distinguish the ‘unusual’ form from the typical (i.e. holotype) form. The

names we suggest here are provided for convenience and to help avoid any

further misinterpretations pertaining to the different forms of this interesting

species.

Material and methods

The material analysed here was obtained during Conservation International’s

Rapid Assessment Program biodiversity survey carried out around Mt Panié,

New Caledonia. The study area was sampled on two occasions: 10-21

October 2010 (MM) and 01-15 November 2010 (SR). Females of the two

colour forms were encountered during the second period only. They were

compared with earlier descriptions in the literature, their affiliation verified

using the figures provided by previous researchers and names assigned to the

two forms.

Abbreviation of the wing venation follows Theischinger and Hawking

(2006).

Systematics

The two females reported here were collected on the same day from the same

locality (see below). Both appear to be mature.

Synthemis miranda f. typica (Figs 1-2)

Material examined. NEW CALEDONIA: 1 9, Dawenia Camp (20°32715”S,

164°40’50”E, 586 m a.s.1.), 15.xi.2010.

Etymology. The name is assigned according to priority of the description and

does not reflect the apparent abundances of the two forms.

This form is represented by the holotype described by Selys (1871). It has

been redescribed and illustrated on a number of occasions (Martin 1906,

Tillyard 1910, Campion 1921), which makes a new description unnecessary.

Figure 1 illustrates the extent of colour on the wings of the female reported

here. The pattern is similar to that of the holotype. In addition to the extent of

colouration the following features, which were not described by previous

researchers, may prove useful in future for distinguishing form typica from

the form described below.

60 Australian Entomologist, 2013, 40 (2)

Figs 1-4. Synthemis miranda females. (1-2) f. typica: (1) wings; (2) head. (3-4) f.

extenuata: (3) wings; (4) head.

Australian Entomologist, 2013, 40 (2) 61

Head (Fig. 2): Mandibles reddish brown with darker colour at the bases and a

white vertical bar at the level of the lateral edge of labrum. Clypeus is not

whitish as given in Campion (1921). A black margin borders almost the

entire postclypeus. It is broader at the anterolateral edges and central

posterior area and narrows towards and along the eyes and then continues

upwards along the borders with the anteclypeus, with the black bars not

meeting on the very top section of this border line. A weak, roughly

triangular black mark extends ventrally from the central part of posterior edge

of postclypeus, with its tip pointing towards anteclypeus. Slightly darker

areas are also present on the anterolateral corners of anteclypeus.

Abdomen with a pair of white ventrolateral spots present on both sides of S1

and a characteristic white ventral edge on each side of S2 to S8. Campion

(1921) reported supplementary basal spots on S5, a feature not included in

the original description of the holotype. They are, however, clearly visible on

the specimen reported here.

Measurements (mm). Total length (with appendages) 68.7; abdomen (with

appendages) 55.4; forewing length 49.4; hindwing length 48.5.

Synthemis miranda f. extenuata (Figs 3-4)

Material examined. NEW CALEDONIA: 1 9, Dawenia Camp (20?32'15"S,

164°40’50”E, 586 m a.s.1.), 15.xi.2010.

Etymology. The name translates from the Latin for alleviated, reduced or

thinned out (H. Fliedner pers. comm.) and refers to the reduced colouration

on the wing area compared with the holotype.

The most notable difference of this form is the extent of colouration on the

wings. Other potentially important characters involve colouration of the

clypeus, frons and abdomen but they are weakly expressed and it is not clear

if they are features typical of the form or are just morphological variation.

Those characters are described following the description of the wings. Other

morphological features that have already been introduced in the literature

were found to be nearly the same as the holotype.

Wings (Fig. 3): Tinged with brown at the bases of both pairs. The colour is

deeper along the main longitudinal veins and fades on the wing membrane to

opaque yellow. It is much darker in the area between Sc and R1 in both pairs

of wings. In the forewings the brown area occupies the total length of the

Ax6 with some patchy signs of colour in subcostal space between Ax9-10.

The colouration extends posteriorly to the anal margin with the wing

membrane becoming more diffuse yellow with light brownish touches along

the transverse veins. The colouration occupies the entire hypertriangle and

cubital space. Subtriangle is free of colour. Weak diffuse brown marks are

developed along the nodus, more intensively at the proximal end of the Sc;

they do not continue on the subnodus. In the hind wings the colouration is

developed as follows: the subcostal space is the darkest area; opaque yellow

62 Australian Entomologist, 2013, 40 (2)

becomes a diffuse brownish around Ax4-5 and continues until Ax7-8; very

weak patches are still present on the cells between Ax9-10 also; the

colouration completely covers the sector of arculus, hypertriangle, triangle

and cubital space and continues posteriorly to the level of the top rows of

cells of the three divisions of the anal loop (middle to the posterior end of

second row in proximal and middle divisions and only first row in the distal

division); the brown continues along the anal vein up to the point where it

joins CuP; weak diffused brown follows the CuP for a very short distance

after this joins at about the level of the first cell formed at the division of CuP

and 1A and also goes one more cell distance along 1A; nodal area is marked

on the same manner as the forewings, with the brown forming an almost

elliptical spot. Scattered diffused brown spots occur at several other places on

the wing membrane.

Head (Fig. 4): Black colouration of clypeus generally resembles the pattern

described above for f. typica but with projections into the whitish area much

more marked. Two additional white spots (one divided by a transverse dark

bar) are developed on the top area of the frons.

Abdomen with no supplementary basal spots on S5. The base of S5 on the

single investigated female is slightly retracted into S4, which obscures clear

vision of this part of the abdomen and this feature needs further verification.

Measurements (mm). Total length (with appendages) 69.3; abdomen (with

appendages) 59.4; forewing length 48.3; hindwing length 47.5.

Discussion

This paper presents the first evidence for co-existence of female S. miranda

with strikingly different wing colouration pattern and we formally refer them

to two separate forms to aid future discussion of this species. Although the

present descriptions are based on two females only, there are multiple data in

the literature indicating that wing colouration is not related to the state of

maturity and this is the first record of sympatry of the same aged (mature)

females collected on the same day. Variation in the extent of wing colour was

also found by Campion (1921) and Davies (2002), who reported it extending

only as far as the level of arculus, which is shorter than in f. extenuata

described here. Also, Lieftinck (1971) reported a specimen with colouration

extent intermediate between f. typica and f. extenuata. Variations at the

infraspecific level are common and were, for example, encountered in a very

extensive survey on female polymorphism in Odonata (Fincke et al. 2005).

That study reported difficulty in assigning some females to a particular form

because of phenotypic variations in otherwise unequivocally andromorph and

heteromorph females. They proposed that female polymorphism must be

heritable in order to be considered as genetic and designated the term

‘functional polymorphism’ to describe genetically monomorphic females

with colour states which are age dependant. It is evident, from previous

Australian Entomologist, 2013, 40 (2) 63

literature records and present observations, that the wing colour in S. miranda

is not influenced by state of maturation and thus is probably inherited.

More evidence is needed to determine whether the other morphological

features mentioned here as potentially diagnostic are useful because our

observations are based on single specimens. Further morphological research

is also necessary to determine whether the additional differences described

here are age dependant, inheritable or vary between populations.

Future identification keys should consider both forms. Lieftinck (1971)

provided the only key to New Caledonian species of Synthemis which, if

followed precisely, may not identify the holotype female. It gives the wing

colouration extending only as far as the arculus, which is typical of f.

extenuata, whereas the dark area in f. typica extends beyond the nodus.

Synthemis miranda is a common species (Davies 2002) and future

investigations will hopefully provide more data about the abundance of and

ratio between the two female colour forms. The names suggested here are

proposed for convenience in subsequent interpretations of the results from

odonatological studies in New Caledonia. The names may not reflect the true

ratio between both female forms and have no formal standing under the

International Code of Zoological Nomenclature. So far it is not clear if the so

called f. typica is really the typical (dominant, most common or often

encountered) of the two. The name is assigned by priority as this form was

the first to be described for the species. It is evident that females with both

types of wing colouration are commonly encountered among the insect

samples from the country. A quick scan through the literature revealed five

records of f. typica and 18 records of f. extenuata. This is simply an account

of the published data and in no case proves the dominance of one form over

the other. Researchers do not always report on the type of females examined

during their studies, nor on the number of specimens examined. For example,

Lieftinck (1971) examined a large collection but reported only on a subset

(six males and six females) of this material and neither Lieftinck (1975) nor

Davies (2002) gave any indication of the total number of females that they

examined. À

Acknowledgements

We thank our local guides for providing us with the best possible assistance

in the field and for kindly hosting us on their land. Our special gratitude goes

to the organisers of the 2010 New Caledonia RAP, particularly to François

Tron and Romain Franquet but also to the whole Conservation International-

New Caledonia staff, their partner organisation Dayu Biik, and J.J. Cassan of

Province Nord. We also thank Vincent Kalkman for the important

suggestions and comments made on early drafts of the paper, Heinrich

Fliedner for consultation on the scientific name of the newly described

female form and Matt Walter for providing photos of the wing colouration.

64 Australian Entomologist, 2013, 40 (2)

References

CAMPION, H. 1921. Odonata collected in New Caledonia by the late Mr. Paul D. Montague.

Annals and Magazine of Natural History 8(9): 33-67.

DAVIES, D. 2002. The odonate fauna of New Caledonia, including the descriptions of a new

species and a new subspecies. Odonatologica 31(3): 229-251.

FINCKE, O., JODICKE, R., PAULSON, D. and SCHULTZ, T. 2005. The evolution and

frequency of female colour morph in Holarctic Odonata: why are male-like females typically the

minority? International Journal of Odonatology 8(2): 183-212.

LIEFTINCK, M. 1971. Studies in oriental Corduliidae (Odonata) I. Tijdschrift voor Entomologie

114(1): 1-63.

LIEFTINCK, M. 1975. The dragonflies (Odonata) of New Caledonia and the Loyalty Islands —

Part 1. Imagines. Cahiers O.R.S.T.O.M., Série Hydrobiologie 9: 127-166.

LIEFTINCK, M. 1976. The dragonflies (Odonata) of New Caledonia and the Loyalty Islands —

Part 2. Immature stages. Cahiers O.R.S.T.O.M., Série Hydrobiologie 10: 165-200.

MARTIN, R. 1906. Collections Zoologiques du Baron Edm. de Selys Longchamps, Catalogue

Systématique et descriptif. Cordulines. Fascicule XVII: 94 pp.

MARTIN, R. 1914. Odonata. Fam. Libellulidae. Subfam. Cordulinae. Genera Insectorum,

Fascicule 155: 1-32.

RIS, F. 1915. Libellen (Odonata) von Neu-Caledonien und den Loyalty-Inseln. Pp 55-72, in:

Sarasin, F. and Roux, J. (eds.), Nova Caledonia. Forsdiungen in Neu-Caledonien und auf den

Loyalty-Inseln, A. Zoologie, Vol. II, L. I.

SCHMIDT, Er. 1938. Libellen als Objekte der angewandten Entomologie. Eine Literaturstudie.

Verhandlungen VII Internationaler Kongress für Entomologie, Berlin: 1494-1505.

SELYS-LONGCHAMPS, E. de. 1871. Synopsis des Cordulines (1), 2"* Légion. — Macromia.

Bulletins de l'Académie Royale des sciences, des lettres et des Beaux-Arts de Belgique (2) 31:

519-565.

TILLYARD, R. 1910. Monograph of the genus Synthemis. Proceedings of the Linnean Society of

New South Wales 35: 312-377.

WINSTANLEY, W. 1983. Terrestrial larvae of Odonata from New Caledonia (Zygoptera:

Megapodagrionidae; Anisoptera: Synthemistidae). Odonatologica 12(4): 389-395.

Australian Entomologist, 2013, 40 (2): 65-66 65

OPETIOPALPUS SCUTELLARIS PANZER (COLEOPTERA:

CLERIDAE: KORYNETINAE) ESTABLISHED IN THE WESTERN

AUSTRALIAN WHEATBELT

JUSTIN S. BARTLETT! and ANDRAS SZITO?

"Biosecurity Queensland, Department of Agriculture, Fisheries and Forestry (Qld), Ecosciences

Precinct, GPO Box 267, Brisbane, Qld 4001 (Email: justin. bartlett@daff.qld.gov.au)

?Plant Biosecurity Entomology, Department of Agriculture and Food Western Australia,

3 Baron-Hay Court, South Perth, WA 6151 (Email: andras.szito@agric.wa.gov.au)

Abstract

The status of the exotic clerid beetle Opetiopalpus scutellaris Panzer has been unclear due to the

ambiguous nature of the single previous Australian record. Recent pheromone trapping at grain

stores in Western Australia indicate that O. scutellaris is locally naturalised within the Western

Australian wheatbelt. It is considered likely that the trapped O. scutellaris specimens originated

from surrounding areas rather than being directly associated with grain.

Introduction

Corporaal (1950) listed 28 species of Opetiopalpus Spinola, distributed

widely throughout Africa and the Palaearctic. References to the biology of

this genus indicate associations with bovid manure, fungi and various plants,

especially legumes, in meadows and grasslands (Bahillo de la Puebla and

López-Colón 2006, Valcarcel et al. 2009). Opetiopalpus scutellaris Panzer

(Fig. 1), from Africa, central and southern Europe and western Asia, has been

associated with old timber (Gerstmeier 1998) and bird nests (Hicks 1959).

This species was first reported in Australia from a single specimen (in

Australian National Insect Collection, Canberra, ACT) from Western

Australia, labelled ‘Acropolis, Geraldton, G22. 22/9/76’ (Kolibaé 2003).

Fig. 1. Opetiopalpus scutellaris, specimen from Wagin, WA (in JSBC).

Discussion

The collection data of the Geraldton specimen give no clues as to the nature

of its capture or whether Acropolis is a place or a freight vessel; however, as

66 Australian Entomologist, 2013, 40 (2)

no location ‘Acropolis’ has been gazetted (Geoscience Australia 2012), as a

general cargo ship of that name was found to have been in service in 1976

(Cameron 2005) and as Geraldton is a port city, the latter seems most likely.

In isolation, this record suggests an interception event and implies adventive

status for O. scutellaris in Australia.

Since January 2011, numerous O. scutellaris specimens have been collected

in traps using aggregation pheromone lures to monitor the stored product

pests Tribolium castaneum (Herbst) (Tenebrionidae) and Rhyzopertha

dominica (Fabricius) (Bostrichidae) near grain stores at Mingenew, Three

Springs, Coorow, Koorda and Wagin in Western Australia. These recent

collections provide evidence that O. scutellaris is locally naturalised within,

and just north of, the wheatbelt region of Western Australia. It is not known

whether the trapped specimens were directly associated with the grain stores,

or were inhabiting surrounding areas in association with dung, timber or bird

nests. Since various non grain-associated insect species were collected in the

same traps and as O. scutellaris has not previously been reported in

association with grain, the latter seems more likely. Specimens are lodged in

the Dept of Agriculture & Food Collection, South Perth, WA (DAFWA), the

Queensland Primary Industries Insect Collection, Dutton Park, Qld (QDPC),

and in the collection of the first author (JSBC).

Acknowledgements

We thank Dave Cousins and Michelle Chami (Dept of Agriculture & Food, Western

Australia) for making recently collected Opetiopalpus scutellaris material available

and Greg Daglish (Agri-Science Queensland, Dept of Agriculture, Fisheries &

Forestry) for reviewing the manuscript.

References

BAHILLO de la PUEBLA, P. and LÓPEZ-COLÓN, J.I. 2006. Los cléridos de la comunidad de

Madrid (Coleoptera, Cleridae). Graellsia 62(número extraordinario): 403-418.

CAMERON, J. 2005. Acropolis - Piraeus, Greece. Great Lakes and International Ship

Photograph Archives [website]. [Accessed 9 October 2012.] Available: http://www.

wellandcanal.ca/salties/a/acropolis/acropolis.htm

CORPORAAL, J.B. 1950. Pars 23 (Editio secunda), Cleridae. In: Hinks, W.D. (ed.),

Coleopterorum Catalogus Supplimenta. W. Junk, The Hague.

GEOSCIENCE AUSTRALIA. 2012. Gazetteer of Australia Place Name Search [website].

[Accessed 9 October 2012.] Available: Attp://www.ga.gov.au/place-names/

GERSTMEIER, R. 1998. Checkered beetles. Illustrated key to the Cleridae of the Western

Palaearctic. Margraf Verlag, Weikersheim.

HICKS, E. A. 1959. Check-list and bibliography on the occurrence of insects in birds' nests.

Iowa State College Press, Ames.

KOLIBÁC, J. 2003. A review of Australian genera of Korynetinae (Coleoptera, Cleridae).

Entomologica Basiliensia 25: 41-97.

VALCÁRCEL, J.P., PILONA, F.P. and RUIZ-TAPIADOR, I. 2009. Sobre la presencia de

Opetiopalpus bicolor (Laporte de Castelnau, 1836) (Coleoptera, Cleridae) en lahunas de Espana

Central. Arquivos Entomolóxicos 1: 17-21.

Australian Entomologist, 2013, 40 (2): 67-78 67

AERIAL BROOD CELLS CONSTRUCTED BY SOME AUSTRALIAN

RESIN BEES (HYMENOPTERA: MEGACHILIDAE) AND A CASE

OF GREGARIOUS NESTING

TERRY F. HOUSTON! and DAVID T. PIKE”

"Department of Terrestrial Zoology (Entomology), Western Australian Museum, Locked Bag 49,

Welshpool Delivery Centre, WA 6986 (Email: Terry.Houston@museum.wa.gov.au)

?37 Everingham St, Carine, WA 6020

Abstract

This paper brings together a number of observations and photographs of ‘aerial’ (i.e. free-

standing and exposed) resin brood cells constructed by Australian Megachile Latreille species.

The cells were attached either singly or in groups to dead twigs on shrubs, standing grass flower

stalks and the underside of a rock. In southwestern Australia, at least two species are known to

build cells attached to dead woody twigs. Three females of one of these species were observed

constructing and provisioning cells at the same time in close proximity on the same twig, thus

suggesting gregarious nesting. The selection pressures which may have led to resin bees

constructing such exposed and seemingly vulnerable brood cells are discussed briefly.

Introduction

The world-wide family Megachilidae is comprised of essentially solitary bees

and its largest genus Megachile (sensu Michener 2007) is notable for

transporting material to its nesting sites for use in nest construction. This

material includes resin, leaf mastic, leaf or petal pieces, plant hairs, mud and

pebbles. Megachiline bees lack the basitibial and pygidial plates

characteristic of burrowing bees and many species (but certainly not all

(Eickwort et al. 1981)) use preformed cavities such as beetle borer holes in

dead wood or the vacated nests of mud wasps in which to build their brood

cells. Such species are often referred to as ‘lodger bees’. Female resin bees

usually apply their loads of resin to the walls of their nesting cavities so that

the cavity, to a large extent, determines the shape of the cells. Only partitions

built across the lumens of cavities to close off individual cells or to seal nest

entrances could be termed ‘free-standing’. Here, we report a very different

mode of nest construction, where cells are built in exposed situations and are

almost completely free-standing. Our observations and photographs were

recorded over many years at various localities in Western Australia and South

Australia. Only in some cases were bees taken with these cells but probably

all were the work of Megachile species.

Taxonomy and identification

The identification of many Australian resin bees, such as those discussed in

this paper, is fraught with difficulty. First, their generic-level classification

has changed several times over the years and remains in a state of flux. Many

species were established in the genus Megachile Latreille (e.g. Cockerell

1930) but Michener (1965) transferred some of them to the genus

Chalicodoma Lepeletier (leaving Megachile for those megachilids that cut

leaves) and he recognized several subgenera. Much later, in his major work

on the bees of the world, Michener (2000, 2007) returned the resin bees to the

68 Australian Entomologist, 2013, 40 (2)

genus Megachile because of the presence of taxa with features intermediate

between those of Megachile and Chalicodoma. More recently, Gonzalez

(2008) made a case for separating some resin bees from both Megachile and

Chalicodoma and proposed that they be placed under the oldest available

generic name, Thaumatosoma Smith. As we are uncertain as to whether this

move will gain general acceptance, we have preferred here to follow

Michener’s (2007) scheme of classification and nomenclature. Second, the

Australian resin bees have not been completely revised. King (1994)

commenced a revision of the Australian Chalicodoma (sensu Michener 1965)

but has not extended it to the subgenera most relevant to the bees discussed

here. Further difficulties are outlined below under ‘Identification of

associated bees’.

All bee specimens and nests collected in the course of this study are lodged in

the entomology collection of the Western Australian Museum.

Observations

Cases ## 1-3

Three clusters of aerial resin brood cells were found on different occasions by

one of us (DTP) in Star Swamp Reserve, North Beach (a suburb of Perth),

Western Australia. This is an area of remnant bushland comprising mainly

Banksia woodland with emergent Tuart (Eucalyptus gomphocephala) trees

and a floristically diverse understorey of shrubs and herbs. One of the clusters

(case #2) was found in an established native garden in the grounds of the

Henderson Environment Centre on the western side of the Reserve.

All cells were oriented with their long axes more or less vertical and their

mouth ends uppermost. They attached to horizontal dead twigs at or very

close to their proximal (mouth) ends and hung mostly below them. All

clusters would have been exposed to the sun for most of the day. Female bees

(apparently of the same species) were observed working on two clusters but

not the third. However, the similarities of the clusters and their occurrence in

the small bushland reserve suggest that all were the work of the same species.

Case # 1

A cluster of three cells (Figs 1-2) was found on 25 November 2006. They

were attached near the end of a dead twig of Melaleuca systena almost 1 m

above ground. Two sealed cells were on one side of the twig and an open cell

was on the other, each cell being in contact with its neighbours. A female bee

was observed working in the open cell (Fig. 2).

Case # 2

A group of eight cells (Figs 3-4) was found on 28 February 2009 in the

Henderson Environment Centre garden. The cells were arranged in a linear

series on a dead horizontal twig protruding from an otherwise healthy

Olearia axillaris and were about 1 m above ground. All cells were attached

Australian Entomologist, 2013, 40 (2) 69

to the same side of the twig at their upper (mouth) ends and were in contact

except for a gap between the sixth and seventh (counting from the left in Fig.

3). Five cells were sealed and three were open (fourth, seventh and eighth).

Three females were active at the cluster. One, seen carrying pollen in Fig. 3,

was provisioning cell # 8 while another was busy capping cell # 7 (Fig. 4).

Figs 1- 2. Case #1: du of diee cells from Star r Samy Reserve, Noni Beach,

WA, viewed from two sides. The abdomen of a female is visible in the open cell in

Fig. 2. Photos: David Pike.

Figs 3-4. Case # 2: group of eight cells found in a native garden on the edge of Star

Swamp Reserve, North Beach, WA. (3) five cells were sealed and three (4 , 7" and

8" from left) were open; two females, one loaded with | pollen, hover over the cluster;

(4) closer view showing female constructing cap on 7" cell and another female in 8"

cell. Notice pale inclusions in resin. Photos: David Pike.

Case #3

A cluster of eleven cells attached to a dead horizontal twig (Fig. 5) was found

on 16 November 2009. The cells occupied both sides of the twig and each

was in contact with one or more neighbouring cells. The pale material coating

the cells was not identified but may have been soil. No bees were observed

with this cluster.

70 Australian Entomologist, 2013, 40 (2)

Fig. 5. Case # 3: a cluster of 11 cells photographed at Star Swamp Reserve, North

Beach, WA. Photo: David Pike.

Case #4

A cluster of five cells (Figs 6-7) was found by DTP in his home garden in the

Perth suburb of Carine on 13 February 2012. The cells were attached to one

side of a dead, horizontal twig of a Tecoma plant and would have been

exposed to the sun for most of the day. The twig was 1.0-1.5 mm in diameter

and about 20 cm above ground. Each cell was attached at its upper (mouth)

end and hung below the twig and each was fused to one or two of its

neighbours. The cells were 9-10 mm long and c. 5 mm in diameter (the group

measuring 24 mm in width). Externally, they were smooth and shiny, the

resin walls containing only a few small, pale inclusions. Three cells (first to

third from left in Fig. 6) were open and, because of the presence of cocoons,

must have yielded adult progeny. The remaining two cells were sealed and,

when opened two days after discovery, each was found to contain a newly

eclosed adult female within a cocoon.

9 ; = 7

Figs 6-7. Case # 4: a five-cell cluster found in a suburban garden in Carine (Perth),

WA. (6) three cells on the left were vacated (cocoons were present) but two others

were sealed and contained newly eclosed females (scale bar = 1 cm); (7) enlarged

view of the three vacated cells showing their relatively smooth, shining surfaces with

few inclusions.

Australian Entomologist, 2013, 40 (2) 71

The cap of each cocoon was comprised of multiple layers of coarse, criss-

crossing, pale brown, silk threads. The remainder of the cocoon was

membranous, colourless and transparent and closely applied to the inner

walls of the cell. Clearly visible beneath this membrane was a thin layer of

faecal material, laid down as longitudinal ribbons. These obscured the

underlying resin walls except at the base of the cell. Microscopic examination

of the faeces revealed that they were composed of a single kind of pollen

grain consistent in morphology with those of Fabaceae.

Case #5

This single cell (Fig. 8), collected by Geoff Allen at Cape Naturaliste, WA in

December 1993, was donated to the Western Australian Museum along with

a female Megachile (registered WAM #17621). The cell and its maker had

been collected at night. The incomplete cell, measuring 9 mm in length and

5.5 mm in maximum diameter, was attached to a dead twig c. 1.5 mm in

diameter. The cell was attached at its mid section (in contrast to cells in cases

##1-4) but its original orientation is unknown. It was constructed from pale,

yellow-brown, translucent resin and revealed distinct annulations at its

proximal (mouth) end. It had been partly provisioned with pollen. The pollen

consisted of one kind of grain: spherical, tricolpate and finely sculptured.

Fig. 8. Case # 5: a cell from Cape Naturaliste, WA, collected in December 1993 by G.

Allen.

72 Australian Entomologist, 2013, 40 (2)

Case # 6

A series of ten cells attached to a dead horizontal twig (Fig. 9) was

photographed by Jean Hort, 5.7 km north of Calingiri, c. 120 km NE of Perth,

on 8 April 2010. The twig, extending from a shrub of Daviesia angulata, was

c. 80 cm above ground in an exposed situation. The habitat was degraded,

regrown native vegetation between a road and a railway. Each cell was

attached by its upper (mouth) end and well separated from its neighbours. AII

cells were sealed and no bees were observed on or about them.

- —m—— ~

Fig. 9. Case # 6: a series of ten cells found near Calingiri, c. 120 km NE of Perth,

WA. Photo: Jean Hort.

Case #7

Two cells attached to a vertical, standing flower stalk of ‘spinifex’ (Triodia

sp.) (Fig. 10) were found by TFH c. 30 km S of Mount Bruce, Karijini

National Park, Hamersley Ranges, WA, on 10 May 1980. The lower cell was

opened, revealing a pharate adult within. This specimen was clearly a

Megachile but could not be identified to subgenus.

Case # 8

Two series of seven and nine cells, respectively, were found attached to the

underside of a rock at Peake Homestead ruins, c. 73 km SE of Oodnadatta,

South Australia, by TFH in April 1977. The cells had been vacated and the

maker was not determined. All cells were attached to the rock by one side,

were in contact with one or two neighbouring cells and faced in the same

direction (Fig. 11). Cell length was estimated to be c. 8.5-9.0 mm.

Australian Entomologist, 2013, 40 (2) 73

Fig. 10. Case # 7: two resin cells attached to a flower stalk of ‘spinifex’ (Triodia sp.)

in Karijini National Park, northern Western Australia. (Match length c. 43 mm). Part

of lower cell was cut away revealing a pharate adult within.

Identification of associated bees

Females were photographed working on some of the Star Swamp cells (cases

## 1-2), taken as unemerged progeny from two Carine cells (case # 4), and a

female was collected with the Cape Naturaliste cell (case # 5). Only a pharate

adult was observed within one of the cells in case # 7. No bees were observed

in the remaining cases.

74 Australian Entomologist, 2013, 40 (2)

UA D uL d UEM di. d LUCAN 1

Fig. 11. Case # 8: two series of cells attached to the underside of a rock at Peake

Homestead ruins, c. 73 km SE of Oodnadatta, SA. Scale is provided by the match

(length c. 43 mm) and a bushfly (upper left).

The females from the Carine cells (case # 4: Figs 12-14) appeared to be

conspecific with the females appearing in Figs 2-4 (cases ## 1-2). The diffuse

orange tomentum over the dorsum of the metasoma and the prominent white

*hair' spot on each side of the first metasomal tergum are distinctive. Other

features which set this species apart from most other Western Australian resin

bees are the following: dense white pubescence forming seven spots on

thorax (pronotum with one median and two lateral spots; mesoscutum with an

anterior paramedian pair and a posterolateral, preaxillary pair); a line of beige

tomentum on posterior margin of mesoscutum; clypeus unmodified except

for a small median notch and tubercle in ventral margin; mandibles with short

tufts of orange setae on outer surfaces subapically (Fig. 14). A specimen in

the Western Australian Museum (WAM 22533), possessing all of these

character states and identified as an unnamed species of Austrochile

Michener (det. J. King), appears to be conspecific. It was collected 103 km

west of Neale Junction, Western Australia, on 21 September 1982, by T. F.

Houston and B. P. Hanich on flowers of Swainsona (Fabaceae). It was

assigned WAM bee species code F437.

Australian Entomologist, 2013, 40 (2) 75

Figs 12-14. Female Megachile reared from one of the Carine cells (case # 4). (12)

dorsal view; (13) lateral view; (14) facial view. Arrows indicate characteristic setal

tufts on mandibles.

The female collected with the cell from Cape Naturaliste (case # 5) is almost

identical to those from North Beach and Carine (cases ## 1-4) but differs as

follows: metasomal terga lacking a general covering of orange tomentum;

T2-4 with complete apical bands of white tomentum; T5 with scattered white

76 Australian Entomologist, 2013, 40 (2)

plumose setae; T6 sparsely covered with ochreous setae. These differences

suggest that it is a distinct species and the specimen (WAM 17621) has been

assigned species code F473.

The subgeneric placement of the two species associated with cases ## 1-4

and # 5 must remain in doubt until a revision of Austrochile and/or

Hackeriapis Cockerell sens. lat. is undertaken. None of the females exhibits a

clear spine on the first metasomal tergum (a diagnostic feature of

Austrochile) and, unfortunately, we are unaware of any conspecific male

specimens that might help decide on the species’ subgeneric placement.

Subsequent to acceptance of this paper, one of us (TFH) reared adults of both

sexes from a series of 11 cells found in the Perth suburb of Hamersley.

Females matched those reared from the Carine cells (case # 4). Males possess

diagnostic features of Megachile (Austrochile) but, at this time, do not assist

species identification (J. King pers. comm.).

A potential maker of the kind of cells reported in case # 8 was found in the

WAM collection: a female (WAM 22535) labelled as an undescribed species

of Austrochile by J. King and collected at Woomera, South Australia, 15

September 1968 by H. Mincham, carries a label stating that it was

“constructing resin cell in cliff face’. With it is a conspecific female (WAM

22532) taken 70-75 km ENE of Norseman, WA. While similar in size and

several features to species F437 from Carine and North Beach, these

specimens lack white spots of tomentum on the pronotum and mesoscutum

and almost certainly represent a distinct species (coded F436).

Discussion

Some different methods of construction can be deduced among the different

cases reported here. In cases ## 1-4, construction must have begun with the

‘mouth’ of the cell and, in case # 5, with the middle section, given the

different points of attachment to the twig. Also, in cases # 1 and #5 the cells

have distinct annulations towards the mouth end but no annulations were

evident in other cells. External finishes of cells in the various cases varied

from smooth and glossy to rough and dull. Some variation among the finishes

of the Star Swamp and Carine cells (cases ## 1-4), believed to have been

made by the one species, could be explained, perhaps, by use of resin from

different sources.

While adult bees were collected or observed with brood cells in only four of

the eight cases described above and appeared to represent just two species,

the variety of cell forms and arrangements suggest that the eight cases may

represent the work of at least five species: (Sp. 1) the Star Swamp and Carine

cells (cases ## 1-4); (Sp. 2) the Cape Naturaliste cell (case # 5); (Sp. 3) the

Calingiri cells (case # 6); (Sp. 4) the Karijini cells (case # 7); (Sp. 5) the

Peake Homestead cells (case # 8).

Australian Entomologist, 2013, 40 (2) 77

Aerial brood cells are constructed by numerous kinds of apocritan wasps,

including some Sphecidae, Crabronidae, Pompilidae and Vespidae (Evans

and Eberhard 1970, Gess 1996), and by various bees (many Megachilidae

and some Apidae in the tribes Euglossini, Bombini and Apini (Michener

2000, 2007)). Mud is used by the majority of the aerial nesting wasps and by

species of Chalicodoma sensu stricto. Resin is used only rarely by wasps

(e.g. Rayment 1935, Mudd and Corbet 1975) and bees other than megachilids

(mainly Apidae: Euglossini, Meliponini and Apini, and in combination with

secreted wax in the last two tribes (Michener 2000, 2007)). The habit of

building aerial cells or nests appears to have arisen independently in these

various hymenopteran families and multiple times in some of them. Aerial

nesting is generally believed to have evolved from cavity nesting which, in

turn, evolved from ground nesting. In Megachilidae, aerial nesting has been

reported in some genera of Anthidiini and Osmiini and in one subgenus

(Megachile (Chalicodoma)) in Megachilini (Eickwort et al. 1981, Michener

2007). It should be noted that Anthidiini are represented in Australia by only

two species known only from Queensland and New South Wales [including

the recently established Afranthidium repetitum Schultz], while Osmiini and

Chalicodoma sensu stricto are not represented in Australia (Michener 1965,

2007). Aerial nesting has not previously been reported for any of the

Australian Megachilini. Doubtless, the cases we describe here represent a

further instance (or instances) of independent evolution of aerial nesting

habits.

To account for multiple origins of aerial nesting among wasps and bees there

must be one or more selection pressures favouring this style of nesting.

Perhaps the most likely advantage would be that the bees are less constrained

in where they can nest. There could be considerable competition among

cavity nesting wasps and bees (and other insects) when few suitable cavities

are available. Aerial nesting females could therefore expend less time and

energy searching for a suitable nesting site and more on building brood cells.

Whatever the benefits of aerial nesting, they must be considerable to out-

weigh new risks from exposure to physical damage from storms, bird attack,

extremes of temperature, bush-fires and parasitoids. When cells are open

during provisioning, they would be especially vulnerable to water entry

during rain and attack by depredators and parasitoids.

Case # 2 was especially interesting in that three females were observed

building and provisioning cells concurrently on the same twig. While

gregarious nesting is common among many solitary, ground nesting bees, it

occurs only rarely among cavity nesting or ‘lodger’ bees. The latter are, of

course, confined to nesting in the available cavities in their environments.

Gregarious nesting, therefore, is a rarity among the Megachilidae and has

< been reported previously for very few species. Females of the American

Dianthidium sayi Cockerell (Anthidiini) build cells of resin and other

materials attached to roots in cavities in the soil and colonies of from 8 to 50

78 Australian Entomologist, 2013, 40 (2)

or more nesting females have been reported (Custer and Hicks 1927). The

females excavate their own nesting cavities and so are not constrained by

availability of pre-existing hollows. Females of Afranthidium repetitum are

known to have constructed a mass of c. 1750 cells from plant hairs in a man-

made box (Michener 2000, 2007). We can see here that when lodger bees

change to aerial nesting they gain another benefit — the potential to nest

gregariously, when defence of the brood cells can be shared among a number

of females. With gregarious nesting, too, comes the potential for development

of higher levels of sociality.

Acknowledgements

We thank Jean Hort of Stratton, Perth, for permission to use one of her

images (Fig. 9) and for associated information. Victor Gonzalez (Natural

History Museum, University of Kansas) kindly read an early draft of this

paper and offered constructive suggestions for its improvement.

References

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6(3): 205-236.

CUSTER, C.P. and HICKS, C.H. 1927. Nesting habits of some anthidiine bees. Biological

Bulletin 52(4): 258-277.

EICKWORT, G.C., MATTHEWS, R.W. and CARPENTER, J. 1981. Observations on the

nesting behavior of Megachile rubi and M. texana with a discussion of the significance of soil

nesting in the evolution of megachilid bees (Hymenoptera: Megachilidae). Journal of the Kansas

Entomological Society 54(3): 557-570.

EVANS, H.E. and EBERHARD, M.J.W. 1970. The wasps. David & Charles (Holdings) Ltd,

Newton Abbot, England.

GESS, S.K. 1996. The pollen wasps, ecology and natural history of the Masarinae. Harvard

University Press, Cambridge, Massachusetts and London, England.

GONZALEZ, V.H. 2008. Phylogeny and classification of the bee tribe Megachilini

(Hymenoptera: Apoidea, Megachilidae), with emphasis on the genus Megachile. ProQuest UMI

Dissertation Publishing 1-274.

KING, J. 1994. The bee family Megachilidae (Hymenoptera: Apoidea) in Australia. I.

Morphology of the genus Chalicodoma Lepeletier, and a revision of the subgenus Hackeriapis

Cockerell. Invertebrate Taxonomy 8(6): 1373-1419.

MICHENER, C.D. 1965. A classification of the bees of the Australian and South Pacific

Regions. Bulletin of the American Museum of Natural History 130: 1-362.

MICHENER, C.D. 2000. The bees of the World. The Johns Hopkins University Press, Baltimore.

MICHENER, C.D. 2007. The bees of the World. 2™ edition. The Johns Hopkins University

Press, Baltimore.

MUDD, A. and CORBET, S.A. 1975. Use of pine resin in nests of pemphredonine wasps.

Transactions of the Royal Entomological Society of London 127(3): 255-257.

RAYMENT, T. 1935. A cluster of bees. Endeavour Press, Sydney.

Australian Entomologist, 2013, 40 (2): 79-88 79

KANALIELLA GEN. N., A NEW GENUS OF CLERIDAE FROM THE

MELANESIAN REGION (COLEOPTERA: CLERIDAE: CLERINAE)

ROLAND GERSTMEIER and MARTIN SEITNER

Technische Universität München, Department Ökologie und Okosystemmanagement, Lehrstuhl

für Tierékologie, Hans-Carl-von-Carlowitz-Platz 2, 85350 Freising, Germany

(E-mail: r.gerstmeier@googlemail.com; martin.seitner@mytum.de)

Abstract

Kanaliella dregei (Perroud, 1864), comb. n. and Kanaliella triangularis (Perroud, 1864), comb.

n., from the Melanesian territories of New Caledonia and Vanuatu, are transferred from

Eunatalis Schenkling, 1909 to the new genus Kanaliella Gerstmeier, gen. n. and lectotypes

designated. Colour photographs, figures of genitalia and distribution maps are provided.

Introduction

Schenkling (1909) erected Eunatalis Schenkling for Australian clerid species

formerly included in the South American genus Natalis Laporte. Kolibaé

(1998) synonymised Metademius Schenkling, Eunatalis Schenkling and

Cormodes Pascoe with Natalis, although Solervicens (2007) disputed this and

reinstated Metademius and Eunatalis as valid genera without commenting on

the status of Cormodes. In his cladogram, Solervicens (2007) showed the

relationships between the South American genera Eurymetomorphon Pic,

Neogyponyx Schenkling and Natalis with the Australian genera Metademius

and Eunatalis. Further assessment of these genera is required in order to

clarify their taxonomic status.

When revising Eunatalis (Gerstmeier and Seitner in press), it became evident

that a new genus was required for the Melanesian species Eunatalis dregei

(Perroud) and Eunatalis triangularis (Perroud). Kanaliella Gerstmeier gen. n.

is herein established, based on differences of elytral punctation/striation,

anterior mesoventral process, protibiae and hindwing venation (see genus

diagnosis below).

Material and methods

This study is based on 37 specimens from 8 museums and institutional

collections. Lectotypes are designated in order to fix the taxonomic identities

of the species involved. Museum acronyms and curators are as follows:

BMNH - Natural History Museum, London, UK (Beulah Garner, Max

Barclay); HNHM — Hungarian Natural History Museum, Budapest, Hungary

(Otto Merkl); IAC — Institut Agronomique néo-Calédonien, La Foa, New

Caledonia (Christian Mille); ISNB — Institut Royal des Sciences Naturelles

de Belgique, Brussels, Belgium (Pol Limbourg); MNHN — Museum National

d'Histoire Naturelle, Paris, France (Thierry Deuve, Antoine Mantilleri);

RGCM - Roland Gerstmeier Collection, Munich (deposited in the collection

of the Technical University Munich); SDEI — Senckenberg Entomologisches

Institut, Müncheberg, Germany (Lutz Behne); QMB — Queensland Museum,

Brisbane, Australia (Susan Wright).

80 Australian Entomologist, 2013, 40 (2)

From all examined specimens the label data were transcribed and presented at

the beginning of each species description. The depository of the specimen is

given in parentheses.

The taxonomic changes herein are based only on morphological studies,

including preparations of the male genitalia, the pygidium and the sixth

ventrite. For this purpose the specimens were heated in distilled water and the

genitalia removed with fine forceps through small incisions made along the

sides of the terminal abdominal segments. Aedeagi were cleaned of soft

tissue in 10% KOH, cleared in 70% ethyl alcohol and submerged in

glycerine, then stored in genitalia vials pinned below each specimen.

Measurements were made under a stereo microscope using an ocular

micrometer. Total body length is the distance measured from the apical

clypeal margin to the elytral apices. Elytral length was measured alongside

the elytral suture and pronotal length was taken from the dorsal middle line.

Pronotal width was measured at the broadest extreme. Measurements of the

elytral width were made across the middle. The scale bar is 1 mm (except as

noted otherwise).

Habitus photographs of adult beetles were taken using a DFC 490 digital

camera fitted to a Leica Z6 stereo microscope, then composited with Leica

LAS 3.8 automontage software.

Plot points of distribution maps were made with QuantumGIS (1.8.0) and are

based on locality data sourced from specimen labels. These data included

exact locality information based on longitude/latitude coordinates. Red labels

were applied to newly designated lectotypes and paralectotypes.

Kanaliella Gerstmeier, gen. n.

Type species: Natalis dregei Perroud, 1864.

Diagnosis. In Kanaliella gen. n. the elytral punctation in the basal part (up to

middle) is arranged into ten striae (in Eunatalis the 9" stria is represented by

distinct punctation near the apex in some species only), the punctation is

without nodules (Eunatalis with two or three nodules), the anterior

mesoventral process is lacking the central excavated area (in Eunatalis

slightly open), the distal margin of the broadly dilated pro-intercoxal process

is thick and bulging (not bulging in Eunatalis), the protibial spine (short

apical apophysis) is absent (Eunatalis with spine), the hind wings are missing

the MP3+4 (Eunatalis with short MP3+4), and the apical field is about one-

third of the elytra (in Eunatalis less than one-third).

Head: Labrum broadly emarginate, V-shaped; terminal maxillary palpomeres

with sides parallel to very weakly divergent and apex obliquely truncate,

terminal labial palpomeres securiform; eyes coarsely facetted, protruding

laterally, margined, strongly ovate; interocular space about 2-2.5 eye widths

(as viewed from above); head including eyes as broad as or broader than

Australian Entomologist, 2013, 40 (2) 81

anterior width of pronotum; gular sutures converging, gular process

conspicuous, with two papillae. Epistomal suture distinct. Antennae with 11

antennomeres, filiform; antennomeres with dense, fine-punctation, terminal

three antennomeres densely tomentose, forming a loose club.

Thorax: Pronotum longer than wide, sides more or less sinuate, with

wrinkles, middle with an elongated groove (sometimes missing or weak);

sub-basal collar distinct, grooves of sub-basal collar fused or separated by an

indistinct bar. Pro-intercoxal process narrow, broadly dilated distally, distal

margin thick and bulging (Fig. 1); procoxal cavities posteriorly closed;

anterior mesoventral process distinct, sinuate, closed apically (Fig. 2);

metaventrite with a conspicuous discriminal line. Metendosternite with

conspicuous furcal arms; laminae acute, triangular (Fig. 3).

Elytra: Elytral base not margined, lateral margins parallel or broadest behind

middle, apices broadly rounded, elytral punctation arranged into ten striae,

interstices with a conspicuous microsculpture (wrinkled); punctation without

nodules. Hind wing with closed wedge cell, margin of radial cell strongly

sclerotized, MP3, MP4, AA3+4 and CuA+AA reaching margin, MP3+4 and

RP2 absent (Fig. 4).

Legs: Inner edge of protibiae and sometimes mesotibiae with a row of short,

thick bristles, with more or less conspicuous punctation and fine wrinkles;

profemora conspicuously swollen, protibiae slightly bent, mesotibiae slightly

bent or straight, metatibiae straight, all tibiae without longitudinal carinae,

protibiae terminating in a single spur (without a spine), tibial spur formula 1-

2-2; tarsal pulvillar formula 3-3-3, tarsal pulvilli broadly lobed, lobes not

emarginate; claws simple.

Abdomen: Six abdominal ventrites.

Etymology. The name Kanaliella refers to Kanala, the New Caledonian type

locality of the two species.

Redescription of the species

Kanaliella dregei (Perroud, 1864), comb. n.

(Figs 5, 7, 9)

Natalis dregei Perroud, 1864: 108; Fauvel 1904: 144.

Type material examined. Lectotype ĝ (designated here): NEW CALEDONIA. dregei

Perroud, Kanala, nov. cal.; ex coll. Perroud; type (MNHN). Paralectotype (designated

here): Coll. R. I. Sc. N. B., Nouvelle Calédonie, Kanala, Fevrier rec. Montrouzier, ex

coll. Fauvel; Coll. A. Fauvel Natalis triangularis Perr. 1864; Syntype (ISNB).

Additional specimens. NEW CALEDONIA: 2 ex., Nova Caled. (MNHN). Bourail

(MNHN). Nov. Caledon. Fairmaire; Natalis dregei Perr.; Eunatalis dregei Perroud,

1985 Ginter Ekis det. (MNHN). Noumea, N. Caledonie; Natalis dregei Perr.; desire

(MNHN). La Foa (MNHN). Yahoué; dregei Perroud (RGCM). Nouvelle Caledonie

(RGCM). New Caledonia, Maré, sifted, 25.v.-6.viii.1987, leg. Janos Balogh (RCGM).

82 Australian Entomologist, 2013, 40 (2)

5 6

Figs 1-6. Kanaliella gen. n. (1) pro-intercoxal process (Scale bar 0.5 mm); (2) anterior

mesoventral process (Scale bar 0.5 mm); (3) metendosternite (Scale bar 1 mm); (4)

wing venation (Scale bar 5 mm); (5) habitus K. dregei; (6) habitus K. triangularis

Lectotype.

Australian Entomologist, 2013, 40 (2) 83

2 ex. New Caledonia, Maré, Araucaria forest, beaten, 25.v.-6.viii.1987, leg. Janos

Balogh (HNHM). Nouvelle-Calédonie, Farino/Barbou L. Face 4 la scierie Alt. 305 m;

Collecte sur: Codiaeum peltatum abattu Le 22/03/05 Par S. Cazéres & C. Mille (IAC).

Nouvelle-Calédonie, Farino/Barbou L. Ensemble de la propriété; Collecte sur: Divers

par battage Le 22/03/05 Par S. Cazéres & C. Mille (IAC). Nouvelle-Calédonie,

Sarraméa, Col d’Amieu; Collecte sur: Divers Le 21 et 23/11/05 Cazéres & Nugues

(IAC). Sarramea: N-Caledonie, Reserve du Col d’Amieu le 17/03/07 par T. Salesne.

Amp. Vap. Mercure (IAC). VANUATU: Brazier; New Heb; Schenkling det;

Brit.Mus. 1907; Eunatalis dregei, Ginter Ekis det. 1995; Col-02942 (SDEI). 46302;

Brazier; New Heb; Fry Coll. 1905-100; Natalis dregei Montrz.; Natalis dregei Montr.

S. Schenkling det. (BMNH).

Description. Length: 10-13.7 mm (19 specimens measured). Head: Black

with isolated pilosity, dense punctation and wrinkles and a more or less

conspicuous longitudinal fovea between the eyes; eyes separated by about

2.1-2.5 eye widths.

Thorax: Pronotum black with light brown to black pilosity; sides rounded

then divergent before the anterior margin, widest behind middle, constricted

basally; surface with large and small punctation, wrinkles on basal half

laterally and dorsally and around central longitudinal fovea; grooves of sub-

basal collar fused, flattened; pronotal length to width ratio 1.06-1.15:1.

Scutellum triangular, dark brown to black with isolated pilosity and dense

punctation. Thoracic ventrites black with isolated pilosity; anterior

mesoventral process lacking deeply sculptured process; mesoventrite finely

wrinkled; posterior part of mesepisternum with isolated punctation.

Elytra: Elytral length to width ratio 2.35-2.45:1; elytra dark brown to black,

with isolated pilosity, with a sharply S- to Z-shaped fascia in the middle of

most specimens which spans from the suture to the lateral margin and is often

broken before the lateral margin, the fascia formed of thick golden setae;

elytral base slightly broader than pronotum, sides broadening towards apex;

punctation very large, deep and irregularly-shaped (sub-aereolate) to apical

two-fifths internally and apical quarter externally; apical area smooth in part

with irregular, shallow, elongate impressions; punctation conspicuously

wider than interstices, becoming flattened and slightly decreasing towards

apex; punctation of ninth and tenth striae conspicuous towards apex;

interstices with fine wrinkles, not ridged.

Legs: Black with short dense pilosity, dense punctation and fine wrinkles;

protibiae slightly bent, mesotibiae straight.

Abdomen: Abdominal ventrites brown to black, some specimens with a

yellowish-coloured posterior margin; tegmen opened ventrally, phallic tip Z-

shaped (in side view), with a hook-like structure; length of phallobase in

relation to whole tegmen about 50%, lateral sclerotization of phallobase

distinct, but not strong, apical lateral sclerotization of parameres weak, V-

shaped branches of ventral sclerotization distinct; spicular fork fused less

84 Australian Entomologist, 2013, 40 (2)

than one-fifth its length; sixth ventrite long, slightly and broadly emarginated;

pygidium compact, straight, processes short, stout (Fig. 7).

Distribution. New Caledonia (including Isle of Pines and Mare, after Fauvel

1904) and Vanuatu (Fig. 9).

Fig. 7. Kanaliella dregei. (A) aedeagus ventral; (B) phallus ventral; (C) tegmen

ventral; (D) spicular fork; (E) pygidium; (F) sixth ventrite (Scale bar 1 mm).

Kanaliella triangularis (Perroud, 1864), comb. n.

(Figs 6, 8, 10)

Natalis triangularis Perroud, 1864: 104; Fauvel 1904: 144.

Type material examined. Lectotype 9 (designated here), NEW CALEDONIA:

triangularis Perroud, Kanala, N. Caled.; ex coll. Perroud; type (MNHN).

Paralectotype (designated here): Coll. R. I. Sc. N. B., Nouvelle Calédonie, Kanala,

Janvier rec. Montrouzier, ex coll. Fauvel; det. Perroud 1864, Natalis triangularis

Perr., ex coll. A. Fauvel; Syntype (ISNB).

Australian Entomologist, 2013, 40 (2) 85

Additional specimens. NEW CALEDONIA: Coll. R. I. Sc. N. B., Nouvelle Calédonie,

Ile des Pins, rec. Deplanche, ex coll. Fauvel; coll. et det. A. Fauvel, Natalis

triangularis Perr. R.LSc.N.B. 17.479 (ISNB). New Caledonia 12263, 21?34'S x

165?50'E Col d'Amieu, N waterfall 26Apr2005, GB Monteith beating, 270 m;

Eunatalis triangularis (Perroud, 1864) Det. JS Bartlett, 2005 (QMB). 1 d, 1 9,

Nouvelle-Calédonie, Farino/M. Barbou, Lat. S 21.61431 Long. E 165.70065 Alt. 408

m; Collecte sur: S. gabriellae Le 27/8/04 Par battage du piége Par S. Cazéres (IAC). 2

ex. Nouvelle-Calédonie, Farino 3/Barbou L. Lat. S 21.63084 Long. E 165.70576 Alt.

315 m; Collecte sur: S. gabriellae Ralia Le 27/10/04 Par S. Cazéres (IAC). 2 3d,

Nouvelle-Calédonie, Farino 7/Barbou L. Lat. S 21.63266 Long. E 165.70684 Alt. 263

m; Collecte sur: S. gabriellae Ralia Le 27/10/04 Par S. Cazéres (IAC). 2 ex. Nouvelle-

Calédonie, Farino/M. Barbou L. Lat. S 21.63084 Long. E 165.70576 Alt. 314 m;

Collecte sur: Schefflera gabriellae Le 27/5/04 Par S. Cazéres (IAC). 1 9, Nouvelle-

Calédonie, Farino/M. Barbou L. Lat. S 21.63084 Long. E 165.70576 Alt. 314 m;

Collecte sur: Schefflera gabriellae Le 09/6/04 Par S. Cazéres (IAC). Nouvelle-

Calédonie, Farino/M. Barbou L. Lat. S 21.63266 Long. E 165.70684 Alt. 263 m;

Collecte sur: Schefflera gabriellae Le 16/7/04 Par S. Cazéres (IAC). Nouvelle-

Calédonie, Farino/M. Barbou, Lat. S 21.61431 Long. E 165.70065 Alt. 408 m;

Collecte sur: S. gabriellae Le 09/9/04 Par battage du piége Par S. Cazéres (IAC). 1 3,

Nouvelle-Calédonie, Farino/Barbou sur S. gabriellae 23/06/04 S. Cazéres coll. (IAC).

Nouvelle-Calédonie, Poya/Beaupré, chez M. Dalstain 26/04/2007 S/C & J/P Coll.

Battage B/cutting 4, S 21?28.634' E 165?11.237, Alt 12 (IAC). Nouvelle-Calédonie,

SRFP 28 ‘Perroquet? Lat S 21.73393 Long. E 165.89605, alt. 23 m; Collecte sur:

Ficus sp. ‘Figuier’ Piège par battage Le 05/01/05 Par S. Cazéres (IAC).

Description. Length: 8.5-11.7 mm (18 specimens measured). Head: Black

with light brown pilosity; with very dense large punctation and dense

wrinkles; frons deeply sunken and bordered posteriorly by a triangular

formation of golden setae, in the centre of the frons are further tuft-like

formations of golden setae; eyes separated by about 2.1-2.33 eye widths.

Thorax: Pronotum dark reddish brown with dense light brown pilosity;

conspicuously constricted behind apex and at base, widest behind middle;

dense punctation and fine wrinkles; dorsal longitudinal fovea distinct;

pronotum with four gibbosities; grooves of sub-basal collar separated by an

indistinct bar; pronotal length to width ratio 1.12-1.22:1; scutellum triangular

to circular, black with dense pilosity and punctation; thoracic ventrites light

brown to dark brown with isolated whitish pilosity and isolated punctation;

anterior mesoventral process not deepened.

Elytra: Elytral length to width ratio 2.59-2.69:1; elytra light brown to dark

brown in basal half, in apical part yellow to light brown, with dense

yellowish pilosity; elytra parallel to sub-parallel, elytral base slightly broader

than pronotum; punctation irregular, punctation in basal half distinct and

deep, becoming smaller towards apex, apical half with isolated punctation,

diameter of punctures wider than interstices, punctation of ninth and tenth

striae conspicuous towards apex; interstices with distinct microsculpture

(wrinkles), interstices 3 and 5 ridged in basal half.

86 Australian Entomologist, 2013, 40 (2)

Legs: Brown to black-brown with dense, light brown pilosity, dense

punctation and wrinkles; pro- and mesotibiae slightly bent.

Abdomen: Abdominal ventrites brown to black with a yellowish-coloured

posterior margin; tegmen opened ventrally; phallic tip not Z-shaped; length

of phallobase in relation to whole tegmen about 50%, lateral sclerotization of

phallobase distinct, but not very strong, apical lateral sclerotization of

parameres very weak, V-shaped branches of ventral sclerotization distinct,

but not very strong; spicular fork fused less than one-fifth its length; sixth

ventrite long more or less straight; pygidium compact, rounded, processes

stout (Fig. 8).

Distribution. New Caledonia (including Isle of Pines and Lifou, after Fauvel

1904; Fig. 10).

Fig. 8. Kanaliella triangularis. (A) aedeagus ventral; (B) phallus ventral; (C) tegmen

ventral; (D) spicular fork; (E) pygidium; (F) sixth ventrite (Scale bar 1 mm).

Australian Entomologist, 2013, 40 (2)

Vanuatu i)

Å, Fü 7 pl

b vs

' S EN e

` S

-29.000| S -20.000

New Caledonia »

160.000 170,000 180.000.

Solomon Islands

PaaS :

Fig. 10. Distribution map of Kanaliella triangularis.

Australian Entomologist, 2013, 40 (2)

Differential diagnosis

Kanaliella dregei is unmistakable by the slightly curved fascia of the elytra

(which is densely vested with golden-yellow setae), the longitudinal fovea

between the eyes, the longitudinal punctation behind the middle of elytra,

fused grooves of the pronotal sub-basal collar and the straight mesotibiae.

Kanaliella triangularis has typical coloration and pilosity of the elytra, the

pronotum is much longer than broad, the grooves of the pronotal sub-basal

collar are separated by an indistinct bar and the mesotibiae are slightly bent.

Acknowledgements

We thank Thierry Deuve, Antoine Mantilleri (MNHN), Beulah Garner, Max

Barclay (BMNH), Pol Limbourg (ISNB), Christian Mille (IAC), Susan

Wright (QMSB), Lutz Behne, Stephan Blank (SDEI) and Otto Merkl

(HNHM) for the loan of specimens, Marianne Miiller (the senior author’s

wife) for her help with the photos and John M. Leavengood Jr. (Lexington,

USA) and Justin S. Bartlett (Brisbane, Australia) for their profound revision

of the manuscript. Sincere thanks to Justin Bartlett for providing us with the

genitalia photos of Kanaliella triangularis.

References

FAUVEL, A. 1904. Faune analytique des coléoptéres de la Nouvelle-Calédonie. Revue d'

Entomologie 23: 113-207.

GERSTMEIER, R. and SEITNER, M. 2013. Revision of the checkered beetle genus Eunatalis

Schenkling, 1909 (Coleoptera: Cleridae, Clerinae). Zootaxa, in press.

KOLIBÁC, J. 1998. Notes on the classification of Natalis Laporte de Castelnau and

Notocymatodera Schenkling, with a description of Notocymatodera disjuncta sp. n. (Coleoptera:

Cleridae). Acta Musei Moraviae, Scientiae biologicae 82(1997): 191-198.

PERROUD, B.-P. 1864. Essai sur la faune entomologique de Kanala (Nouvelle-Calédonie) et

description de quelques espéces nouvelles et peu connues. Annales de la Societé Linnéenne de

Lyon (n.s.) 11: 46-256.

SCHENKLING, S. 1909. Aus den Sitzungen. Deutsche Entomologische Zeitschrift 1909: 162-

163.

SOLERVICENS, J.A. 2007. Cladistic analysis of species of Natalis Laporte (1836) and related

genera Eunatalis Schenkling (1909), Metademius Schenkling (1899) and Eurymetomorphon Pic

(1950) (Coleoptera: Cleridae: Clerinae) with redescription of a restored Clerinae genus. Zootaxa

1398: 1-14.

Australian Entomologist, 2013, 40 (2): 89-92 89

THE COURTSHIP BEHAVIOR OF THE BEE FLY MEOMYIA

VETUSTA WALKER (DIPTERA: BOMBYLIIDAE)

D.J. FERGUSON and D.K. YEATES

Australian National Insect Collection, CSIRO Ecosystem Sciences, PO Box 1700, Canberra,

ACT 2601

Abstract

Observations are provided on the courtship behaviour of Meomyia vetusta Walker in

southeastern New South Wales, Australia.

Introduction

The Bombyliidae are a species rich, globally distributed group of brachyceran

flies (Yeates 1994). Little is known of the life histories and behavioural

biology of the Australian species, although there are many records of

parasitism by non-Australian bombyliid larvae (Hull 1973, Yeates and

Greathead 1997). Dodson and Yeates (1990) and Yeates and Dodson (1990)

studied the territorial, hill-topping behaviour of male Comptosia tutela

Yeates in southeastern Queensland. During these studies, a male was seen to

intercept a female within a male territory, in mid-flight, achieving tail-to-tail

copulation without displaying any overt courtship behaviour (Yeates and

Dodson 1990). Toft (1989a, b) studied male Lordotus pulchrissimus

Williston swarming behaviour and interactions with females in Californian

sand dune environments; again courtship behaviour was not observed

although the species copulates in mid air. Thus, bombyliid courtship

behaviour, in territorial and swarming species at least, seems to consist of a

simple mid-air intercept, grapple and copulation. This behavioural pattern is

also common in many species of Lower Diptera such as Chironomidae,

where the males form swarms and the females visit the swarms and copulate

immediately with males in mid air (Downes 1969).

Colless (1977) observed a female Anastoechus sp. apparently displaying

ovipositing behaviour on the floor of a sandstone overhang in the Bukalara

Plateau, Northern Territory; however, this was most likely sand gathering

rather than actual oviposition (Yeates 1994). One of us (DJF) observed the

courtship behaviour of Meomyia vetusta Walker (Fig. 1) in sand dunes in

southeastern New South Wales and this is reported here. To our knowledge,

this species does not form hilltop aggregations, swarms or leks and the series

of behaviours leading up to copulation is much more complex than

previously reported for Comptosia or Lordotus.

The geographic distribution of Meomyia vetusta, determined from specimens

in the Australian National Insect Collection (ANIC), is from southeastern

Queensland to the Victorian border, mostly along the coast, with several

specimens taken at higher altitudes along the Great Dividing Range. Adults

fly from August to December in the northern part of the range and from

October to December towards the southern end of the range.

90 Australian Entomologist, 2013, 40 (2)

Fig. 1. Meomyia vetusta (ANIC_29:029405) female, collected on the same day at the

same location as the observations reported here. Scale = 5 mm.

Observations

Observations were made in a sand dune environment adjacent to Broulee

Island, southeastern New South Wales (35°51’S 150°10’E), on the morning

of 1 October 2012. Weather conditions were calm, 15°C at 0900h and under a

cloudless sky.

The first courtship observation was at approximately 0930h and was detected

through a loud buzzing noise produced by the male, which was hovering just

in front of a motionless female resting on the tip of a leaf of Lomandra

longifolia (Xanthorrhoeaceae). The male was slowly hovering in an arc of

approximately 45° directly in front of the female and in the same horizontal

plane. The male was also actively touching the female’s proboscis with his

throughout this hovering flight behaviour. After approximately 90 seconds

the female took flight, when the male intercepted and grappled with her in

mid air, achieving a tail-to-tail copulating position before the pair flew away.

Both male and female were netted and vouchered in ANIC.

The second observation occurred at around 1000h, when a female, flying

slowly approximately 30 cm above the ground, passed over a resting male.

The male immediately flew rapidly to intercept the female, bumped into her,

then dropped below and behind her, keeping a distance of about 15 cm. The

male followed the female for about 20 m until she landed on the end of an L.

longifolia leaf. Consistent with the previous observation, the male then

produced a loud buzzing sound, which was louder and of a different pitch to

the sound produced when flying normally. This loud buzzing sound was

produced while the male hovered left and right of the motionless female in an

Australian Entomologist, 2013, 40 (2) 91

arc of approximately 45°, and while the male actively touched his proboscis

against the female’s. This continued for approximately a minute, until the

female took flight, when the male then intercepted and grappled with the

female, achieving a tail-to-tail copulation in mid air. The pair then flew a

short distance away before landing and remaining in copula.

Discussion

The two observed courtships, involving different pairs, were consistent and

suggest the following series of six behaviours, by both male and female,

leading to copulation in Meomyia vetusta:

1. The female flies slowly near the ground in the vicinity of the resting male.

2. Once within range, the male intercepts the female in flight and grapples

with her.

3. The female continues flying while the male drops to a position behind and

below the female, separated by a distance of approximately 15 cm.

4. The pair fly in unison for a few metres, then the female lands while the

male hovers directly in front of her.

5. The male touches the female’s proboscis with his proboscis, while slowly

hovering in an arc of approximately 45° directly in front of the female in the

same horizontal plane. The female remains motionless during this stage.

While hovering, the male also produces a distinct, loud, buzzing sound with

his wings. This stage lasts 60-90 seconds.

6. The pair fly upwards, the male intercepts the female and grapples with her

before quickly achieving a tail-to-tail copulation position, the pair then flying

briefly together before landing to remain in copula.

Stage 5 is clearly the most complex and unusual and not previously described

in Diptera. No fluid transfer or nuptial gifting was observed between the

sexes during this stage, but remains a possibility and does occur in other flies

(Marshall 2012). The mechanism of sound production during stage 5 is

unknown, but presumably involves a change in the attitude of the wings as

the insect hovers. The series of behaviours elicited by both sexes leading up

to copulation in Meomyia vetusta shows that the courtship behaviour of bee

fly species that do not swarm or lek may be very complex.

Acknowledgement

We thank Cate Lemann, CSIRO Ecosystem Sciences for the photograph.

References

COLLESS, D.H. 1977. A note on apparent oviposition in a bombyliid fly (Diptera). Australian

Entomological Magazine 3(5): 90-91.

DODSON, G.N. and YEATES, D.K. 1990. The mating system of a beefly (Diptera,

Bombyliidae) II. Factors affecting male territorial and mating success. Journal of Insect

Behaviour 3: 619-636.

92 Australian Entomologist, 2013, 40 (2)

DOWNES, J.A. 1969. Swarming and mating flight of Diptera. Annual Review of Entomology 14:

271-293.

HULL, F.M. 1973. Bee flies of the World. Smithsonian Institution Press; 687 pp.

MARSHALL, S.A. 2012. Flies: the natural history and diversity of Diptera. Firefly Press,

Ontario; 616 pp.

ROBERTS, F.H.S. 1928-29. A revision of the Australian Bombyliidae (Diptera). Parts 1-111.

Proceedings of the Linnean Society of New South Wales 53: 90-144, 413-55 (1928); 54: 553-83

(1929).

TOFT, C.A. 1989a. Population structure and mating system of a desert bee fly (Lordotus

pulchrissimus; Diptera: Bombyliidae). 1. Male demography and interactions. Oikos 54: 345-358.

TOFT, C.A. 1989b. Population structure and mating system of a desert bee fly (Lordotus

pulchrissimus; Diptera: Bombyliidae). 2. Female demography, copulations and characteristics of

swarm sites. Oikos 54: 359-369.

YEATES, D.K. and DODSON, G. 1990. The mating system of a beefly (Diptera, Bombyliidae).

I. Non-resource-based hilltop territoriality and a resource-based alternative. Journal of Insect

Behaviour 3: 603-617.

YEATES, D.K. and GREATHEAD, D. 1997. The evolutionary pattern of host use in the

Bombyliidae (Diptera): a diverse family of parasitoid flies. Biological Journal of the Linnean

Society 60: 149-185, 5 figs.

Australian Entomologist, 2013, 40 (2): 93-98 93

THEMARA MACULIPENNIS (WESTWOOD) AND THEMARA

HIRTIPES RONDANI (DIPTERA: TEPHRITIDAE:

ACANTHONEVRINI): A CASE OF CONFUSED SYNONYMIES

DAVID L. HANCOCK

8/3 McPherson Close, Edge Hill, Cairns, Qld 4870

Abstract

Two species of the Oriental genus Themara Walker, viz. T. maculipennis (Westwood) and T.

hirtipes Rondani, are discussed, with T, enderleini Hering removed from synonymy with T.

hirtipes and placed as a new synonym of T. maculipennis, while Themara montina Enderlein is

removed from synonymy with T. maculipennis and placed as a new synonym of T. hirtipes.

Lectotypes for T. maculipennis and T. enderleini are designated. A note on biology is included.

Introduction

During a recent examination of Themara Walker specimens in the Natural

History Museum, London, it became clear that the synonymies in T.

maculipennis (Westwood) and T. hirtipes Rondani had become confused.

This is due to an undue reliance being placed on the presence or absence of a

dark medial vitta on the scutum for assigning specimens to one species or the

other (e.g. Perkins 1938, Hardy 1974, 1986). It is now known that this

character is intraspecifically variable (Hancock 2011), with the extent of

curvature in vein R+ in males being far more reliable as a specific character.

Themara yunnana Zia, previously placed as a synonym of T. hirtipes (e.g.

Hardy 1974, 1986, Hancock and Drew 1994), was shown to be a distinct

species by Hancock (2011). Two other cases of misplaced synonyms are

discussed here. Since the type series of one of these synonyms, T. enderleini

Hering, appears to be a mix of three separate species, a lectotype is

designated so that it may be properly defined. A lectotype is also designated

for T. maculipennis (Westwood) so that it, too, becomes taxonomically fixed.

Since biological studies are being undertaken on these species (e.g. Burkhardt

and de la Motte 1994, 1996), it is essential that they are properly identified.

The following abbreviations have been used for specimen depositories:

BMNH - The Natural History Museum, London; DEI — Senckenberg

Deutsches Entomologisches Institut; MCSNG - Museo Civico di Storia

Naturale *Giacomo Doria", Genoa; OUMNH - Oxford University Museum

of Natural History, Oxford; PAN — Polish Academy of Science Museum of

the Institute of Zoology, Warsaw; QDPI — Queensland Department of

Agriculture, Forestry and Fisheries, Brisbane; ZMHU — Museum für

Naturkunde, Berlin.

Systematics

Themara maculipennis (Westwood)

(Figs 1-3)

Achias maculipennis Westwood, 1847: 38, pl. 18 fig. 4. Type locality Java, Indonesia.

Lectotype 3 in BMNH, here designated; examined.

94 Australian Entomologist, 2013, 40 (2)

Achias horsfieldii Westwood, 1850: 235, pl. 23 fig. 9. Type locality Java, Indonesia.

Lectotype 3 in BMNH, designated by Hardy (1969); examined.

Acanthoneura maculipennis: Enderlein, 1911: 415 [partim]. Soekeranda and

Liangagas, Sumatra, Indonesia.

Themara enderleini Hering, 1938: 409. Type locality Liangagas, Sumatra, Indonesia.

Lectotype 3 in BMNH; here designated; examined. Syn. n.

Material examined. INDONESIA: Lectotype 3 of Achias maculipennis [labelled HT],

Java (BMNH); 1 ¢ paralectotype [labelled ST], Java (BMNH); 1 ¢ paralectotype,

‘W? [ex Westwood Collection], no data (OUMNH); Lectotype $ and 1 3, 1 9

paralectotypes of Achias horsfieldii, Java (BMNH); 4 33, 2 99, West Java

(BMNH); Lectotype 3 of Themara enderleini [labelled PT], Liangagas, Sumatra,

Dohrn (also labelled *Themara enderleini, PT, det. M. Hering 1937' / *Purch. from

E.M. Hering, B.M. 1965-270’) (BMNH); 1 3, West Sumatra (BMNH); 1 d, Padang,

Sumatra (BMNH); 1 3, Lebang Tandal, West Sumatra (BMNH). SINGAPORE: 1 3,

Singapore, 1867 (OUMNH); 1 3, Singapore (BMNH). WEST MALAYSIA: 1 3,

1 9, Ulu Gombak, Serdang, 25.iii.1988, ILM. White & Ooi, C.S. (BMNH).

Notes. Themara enderleini was placed in synonymy with T. hirtipes by

Hardy (1974) but Enderlein’s (1911) series, removed from 7. maculipennis

and renamed by Hering (1938), is clearly mixed, containing representatives

of T. maculipennis, T. hirtipes (a female with the hyaline indentation in cell

m narrow — presumably the specimen referred to T. hirtipes below) and T.

ampla Walker (a female with no hyaline spot in cell r4,5 near the DM-Cu

crossvein). Enderlein (1911) noted the range of variation and had included T.

ampla within the synonymy of 7. maculipennis. Hering (1938) referred to a

‘Type 3’, a ‘Type 9" and several other specimens; these are thus all of

syntype status. The location of most of this material is uncertain (Norrbom ef

al. 1999). Hering (1938) stated ‘Stettiner Museum’ [now PAN], whereas

Hardy (1974) recorded the ‘Type 3” as in DEI. One pair of STs, however, is

in BMNH (from Hering’s personal collection) so it is evident that not all, if

any, of the specimens examined by Hering were returned to PAN [the

intervention of World War II might have prevented this]. In order to stabilise

the name and prevent further confusion, a lectotype is designated. The male

in BMNH (Fig. 1), although labelled PT, is selected since (a) it bears

Hering’s determination label and was definitely seen by him; (b) its location

is known; and (c) it is readily comparable with the lectotypes of T.

maculipennis and T. horsfieldii, both also in BMNH.

In order to maintain stability of the name T. maculipennis, a lectotype is also

designated for that species. Three syntype males are present in BMNH and

OUMNH, the specimen selected being the male in BMNH labelled ‘HT’.

Perkins (1938) also illustrated the male wing. The wing of a Javan female

attributed to this species is provided here (Fig. 2). The above specimens from

West Malaysia (Fig. 3) are smaller than typical specimens and the male has a

correspondingly narrower head but they otherwise appear to belong to T.

maculipennis and are included here.

Australian Entomologist, 2013, 40 (2) 95

Figs 1-3. Themara maculipennis. (1) Lectotype male of the synonym T. enderleini

from Sumatra; (2) wing of female from West Java; (3) male and female from West

Malaysia. Photos by K. Goodger O Natural History Museum, London.

96 Australian Entomologist, 2013, 40 (2)

This species may be separated from T. hirtipes by the more distinctly

undulate and curved vein R243, especially in males (where it meets the costa

before the mid point between veins R; and R4,;), the broader hyaline

indentation in cell m and the generally shorter and thicker ‘eye stalks’ in

males. From 7. ampla it differs in the presence of the male ‘eye stalks’ and

better developed hyaline spot in cell r4+5 near the DM-Cu crossvein. The dark

medial vitta on the scutum is often indistinct. In the otherwise similar Indo-

Chinese species T. yunnana, the male head is broad but not produced into

distinct ‘eye stalks’ (Hancock 2011, David and Ramani 2011).

T. maculipennis is known from Java, Sumatra, Singapore and West Malaysia.

Themara hirtipes Rondani

Achias maculipennis: Walker, 1856: 134. Sarawak, Malaysia. Misidentification.

Themara hirtipes Rondani, 1875: 435. Type locality Sarawak, Malaysia. Type 3 in

MCSNG; not examined.

Acanthoneura maculipennis: Enderlein, 1911: 415 [partim]. Soekeranda, Sumatra,

Indonesia. Misidentification.

Acanthoneura montina Enderlein, 1911: 416. Type locality Mt Gede, west Java,

Indonesia. Holotype in PAN; not examined. Syn. n.

Themara enderleini: Hering, 1938: 409 [partim]. Soekeranda, Sumatra, Indonesia.

Misidentification.

Themara palawanica Hering, 1938: 410, fig. 2. Type locality Binaluan, Palawan,

Philippines. Holotype in ZMHU; not examined. Synonymised by Hardy (1974).

Themara maculipennis: Hardy, 1986: 141 [partim]. Sabah, Malaysia.

Misidentification.

Themara maculipennis: Hancock & Drew, 1994: 564. Sarawak, Malaysia.

Misidentification.

Themara maculipennis: Chua, 2002: 46. Brunei. Misidentification.

Material examined. INDONESIA: 1 9 [labelled PT of Themara enderleini],

Soekeranda, Sumatra, Dohrn (BMNH). EAST MALAYSIA: 17 dd, Kuching,

Sarawak (OUMNH); 1 d, Tarat Agric. Station, near Kuching, Sarawak, 10.viii.1992,

S. Leong (QDPI); 2 33, Sg. Poyan, Limbang, Sarawak, vi.1993, S. Leong (QDPI).

Several additional specimens from Sarawak, of both sexes, in BMNH.

Notes. Acanthoneura montina was placed in synonymy with 7. maculipennis

by Hardy (1986) but Enderlein’s (1911) description indicates a very broad

head and states 'r;,; endet sehr wenig ausserhalb der Mitte zwischen r; und

T4+5” (vein Ras; ends a little outside [beyond] the mid point between veins R;

and R45), thus indicating a weakly undulate vein that does not curve sharply

towards the costa. The synonymy of T. palawanica is evident from Hering’s

(1938) description and illustration. At least one specimen from Enderlein’s

(1911) series (the ‘PT’ of T. enderleini from Soekeranda, Sumatra noted

above) appears to belong to T. hirtipes. A third species included in the

synonymy of T. hirtipes by Hardy (1974), T. yunnana Zia, was shown to be a

distinct species by Hancock (2011) and removed.

Australian Entomologist, 2013, 40 (2) 97

This species was illustrated by Perkins (1938), Hancock and Drew (1994, as

T. maculipennis) and Hancock (2011). It differs from both 7. maculipennis

and T. ampla in the less undulate vein R23 in both sexes, the narrower

hyaline indentation in cell m and the generally more elongate and slender

‘eye stalks’ in males. The dark medial vitta on the scutum may be present or

absent and the amount of black on the scutellum is variable in extent.

T. hirtipes has been recorded from southeast China (Hainan), Laos, Thailand,

southern Burma, Sumatra, Java, West Malaysia, Sarawak, Sabah, Brunei and

Palawan. The note following the Singapore record of T. maculipennis in

Hardy (1986) possibly refers to one of the other listed specimens.

Biology

Perkins (1938) recorded specimens of Themara hirtipes collected on the bark

of felled trees in old secondary forest in Sarawak, Malaysia and this species

is now known to breed in rotting logs, with females ovipositing in holes made

by bark beetles or other insects (Burkhardt and de la Motte 1994, 1996).

Perkins (1938) also recorded specimens of T. hirsuta (Perkins) on bark of

felled trees and it is likely that all species of Themara breed in rotting logs. It

is also likely that the closely related genera Acanthonevra Macquart (as

restricted by Hancock 2011, 2012b), Chaetomerella de Meijere and

Freyomyia Hardy also utilise this host. A presumed association with bamboo

(Hancock 2011) for these genera is likely to be no more than casual, whereas

other genera in the Acanthonevra complex, such as Ptilona van der Wulp and

Rioxoptilona Hendel, breed in dead bamboo culms or decaying bamboo

shoots respectively (Hancock 2011).

The breeding biology of Themara is very similar to that of Rioxa

sexmaculata (van der Wulp) [a species in the related Rioxa complex], which

also oviposits in holes made by bark beetles or other insects in rotting logs

(Kovac et al. 2010) and log-breeding might be a basal character in the

Acanthonevra complex, with utilisation of decaying bamboo a secondary

characteristic (and possibly independently derived in Ptilona and

Rioxoptilona, which use different parts of the bamboo plant). In the similarly

related Sophira complex, it appears to be living bamboo stems and older

shoots that are utilised (Hancock 2012a).

Acknowledgements

I thank Kim Goodger (BMNH) and James Hogan (OUMNH) for the

photographs and/or access to specimens in their care. Damir Kovac

(Forschungsinstitut Senckenberg, Frankfurt am Main) kindly brought the

studies on Themara biology to my attention and supplied a copy of his paper.

References

BURKHARDT, D. and de la MOTTE, I. 1994. Tephritid flies. In: Earl of Cranbrook and

Edwards, D.S. (eds), A tropical rainforest. The nature of biodiversity in Borneo at Belalong,

Brunei. The Royal Geographical Society and Sun Tree Publishing, Singapore; 389 pp.

98 Australian Entomologist, 2013, 40 (2)

BURKHARDT, D. and de la MOTTE, I. 1996. Stalk-eyed flies: caprices of evolution? Pp 169-

174, in: Edwards, D.S., Booth, W.E. and Choy, S.C. (eds), Tropical rainforest research —

current issues. Monographie Biologicae 74. Kluwer Academic Publishers; 566 pp.

CHUA, T.H. 2002. New records of Trypetinae from Brunei Darussalam (Diptera: Tephritidae).

Malayan Nature Journal 56(1): 43-48.

DAVID, K.J. and RAMANI, S. 2011. An illustrated key to the fruit flies (Diptera: Tephritidae)

from peninsular India and the Andaman and Nicobar Islands. Zootaxa 3021: 1-31.

ENDERLEIN, G. 1911. Trypetiden-Studien. Zoologische Jahrbucher. Abteilung für Systematik,

Oekologie und Geographie der Tierre 31: 407-460.

HANCOCK, D.L. 2011. An annotated key to the species of Acanthonevra Macquart and allied

genera (Diptera: Tephritidae: Acanthonevrini). Australian Entomologist 38(3): 109-128.

HANCOCK, D.L. 2012a. Bamboo-stem flies: an annotated key to the species of the Sophira

complex of genera (Diptera: Tephritidae: Acanthonevrini). Australian Entomologist 39(1): 5-32.

HANCOCK, D.L. 2012b. A note on the identity of ‘Acanthonevra’ inermis Hering (Diptera:

Tephritidae: Acanthonevrini). Australian Entomologist 39(3): 195-196.

HANCOCK, D.L. and DREW, R.A.I. 1994. New species and records of Asian Trypetidae

(Diptera: Tephritidae). Raffles Bulletin of Zoology 42(3): 555-591.

HARDY, D.E. 1969. Lectotype designations for fruit flies (Diptera: Tephritidae). Pacific Insects

11: 477-481.

HARDY, D.E. 1974. The fruit flies of the Philippines (Diptera: Tephritidae). Pacific Insects

Monograph 32: 1-266, 6 pls.

HARDY, D.E. 1986. Fruit flies of the subtribe Acanthonevrina of Indonesia, New Guinea, and

the Bismarck and Solomon Islands (Diptera: Tephritidae: Trypetinae: Acanthonevrini). Pacific

Insects Monograph 42: 1-191.

HERING, [E].M. 1938. Neue palaearktische und exotische Bohrfliegen. Deutsche

Entomologische Zeitschrift 1938: 397-417.

KOVAC, D., DOHM, P. and FREIDBERG, A. 2010. Field observations on the mating behaviour

of the Oriental Rioxa sexmaculata (van der Wulp) (Diptera: Tephritidae) and a review of the

reproductive behaviour patterns in Acanthonevrini. Biosystematica 4(1): 5-14.

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: ix + 524 pp.

PERKINS, F.A. 1938. Results of the Oxford University Expedition to Sarawak (Borneo), 1932.

Diptera, Trypetidae. Annals and Magazine of Natural History (11) 2: 401-409, pl. xv.

RONDANI, C. 1875. Muscaria exotica Musei Civici Januensis. Fragmentum III. Species in

Insula Bonae fortunae (Borneo), Provincia Sarawak, annis 1865-1868 lectae a March. J. Doria et

Doct. O. Beccari. Annali del Museo Civico di Storia Naturale, Genova 7: 421-464.

WALKER, F. 1856. Catalogue of the dipterous insects collected at Sarawak, Borneo, by Mr. A.

R. Wallace, with descriptions of new species. Journal of Proceedings of the Linnean Society of

London, Zoology (1857) 1: 105-136.

WESTWOOD, J.O. 1847. Diptera. Pp 37-38, pl. 18, in: The cabinet of Oriental Entomology. W.

Smith, London; 2 2 + 88 pp (1848).

WESTWOOD, J.O. 1850. Diptera nonnulla exotica descripta. Transactions of the Entomological

Society of London (1849) 5: 231-236, pl. xxiii.

Australian Entomologist, 2013, 40 (2): 99-100 99

FURTHER AUSTRALIAN LOCALITY DATA FOR CTENOLEPISMA

ROTHSCHILDI SILVESTRI (ZYGENTOMA: LEPISMATIDAE)

GRAEME SMITH! and JOHN IRISH?

!246 Alfred St, Narraweena, NSW 2099 (Email: le_gbsmith@optusnet.com.au)

?PO Box 30061, Windhoek, Namibia (Email: jirish@mweb.com.na)

Abstract

Further locality records for Ctenolepisma rothschildi Silvestri are given, confirming its presence

in Australia since at least 1960.

Introduction

Smith (2012) reported the presence of this semi-cosmopolitan silverfish

species at Longreach in Queensland and used that material to redescribe the

species according to current criteria. Although not previously reported in the

literature, its presence at a relatively remote inland town suggested it had

been in Australia for some time.

Recently, one of us (GS) became aware of older material in the Australian

National Insect Collection (ANIC), which had been identified by the other

(JI) as Ctenolepisma rothschildi Silvestri. Contact was established and JI

provided details of this and other material of C. rothschildi examined from

the Museum and Art Galleries of the Northern Territory (NTM) and the

Entomology Department of the University of Queensland (EUQ), which

confirms the species’ presence in Australia since at least 1960. This short

note records the material determined by JI from Queensland, the Northern

Territory and Western Australia. EUQ material is now housed in the

Queensland Museum, Brisbane.

New records

Family Lepismatidae Latreille

Ctenolepisma rothschildi Silvestri, 1907

Material examined. QUEENSLAND: 2 99, 1 3, Ingham Res. Station, 13.iv.1960,

K.L.S. Harley (EUQ); 1 9, Townsville Common, base of A. laurensis mound,

25.vii.1966, F.J. Gay (with two specimens of Lepisma sp. (sensu lato)) (ANIC).

NORTHERN TERRITORY: 1 3, Darwin, August 1983, M. Malipatil (NTM); 1 9,

Darwin, 30.iv.1985, M.B. Malipatil (NTM); 1 9, Darwin, August 1988, M.B.

Malipatil (NTM); 1 9, Howard Springs, 2 km down Virginia Rd, 9.ix.1990, K.

Coombes (NTM); 1 unsexed in many pieces, Radon Ck, WL PT A/3, 8.viii.1975

(NTM). WESTERN AUSTRALIA: 2 99, 1 ĝ, 2 unsexed, Camballin, 5.iii.1968,

J.A.L. Watson (ANIC).

Conclusion

These records confirm that Ctenolepisma rothschildi has been present in

Australia since at least 1960 and has since been found at a wide range of

locations across the tropical northern region. Many of these locations are

hundreds of kilometres from any port and in lightly populated areas,

100 Australian Entomologist, 2013, 40 (2)

suggesting that the species has become established in northern Australia and

is now widespread. Due to limited efforts to collect silverfish in much of

Australia, no comment can be made as to its abundance.

Acknowledgments

We thank Drs M. Schneider, A. Wells and the late J.A.L. Watson for

arranging the loan of this material to JI from EUQ, NTM and ANIC

respectively.

Reference

SMITH, G. 2012. Ctenolepisma rothschildi Silvestri (Zygentoma: Lepismatidae) in Australia.

Australian Entomologist 39(1): 39-47.

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THE AUSTRALIAN

Entomologist

Volume 40, Part 2, 21 May 2013

CONTENTS

BARTLETT, J.S. AND SZITO, A.

Opetiopalpus scutellaris Panzer (Coleoptera: Cleridae: Korynetinae) established in the

Western Australian wheatbelt

FERGUSON, D.J. AND YEATES, D.K.

The courtship behavior of the bee fly Meomyia vetusta Walker (Diptera: Bombyliidae)

GERSTMEIER, R. AND SEITNER, M.

Kanaliella gen. n., a new genus of Cleridae from the Melanesian region (Coleoptera: Cleridae:

Clerinae)

HANCOCK, D.L.

Themara maculipennis (Westwood) and Themara hirtipes Rondani (Diptera: Tephritidae:

Acanthonevrini): a case of confused synonymies

HOUSTON, T.F. AND PIKE, D.T.

Aerial brood cells constructed by some Australian Resin Bees (Hymenoptera: Megachilidae)

and a case of gregarious nesting

MARINOV, M. AND RICHARDS, S.

Notes on the female colour forms of Synthemis miranda Selys, 1871 (Odonata: Synthemistidae)

in New Caledonia

SMITH, G. AND IRISH, J.

Further Australian locality data for Crenolepisma rothschildi Silvestri (Zygentoma: Lepismatidac)

ISSN 1320 6133