THE AUSTRALIAN ENTOMOLOGIST

ABN#: 15 875 103 670

The Australian Entomologist is a non-profit journal published in four parts annually

by the Entomological Society of Queensland and is devoted to entomology of the

Australian Region, including New Zealand, Papua New Guinea and islands of the

south-western Pacific. Articles are accepted from amateur and professional

entomologists. The journal is produced independently and subscription to the

journal is not included with membership of the society.

The Publications Committee

Editor: Dr D.L. Hancock Editorial Co-ordinator

Assistant Editors: Dr C.J. Burwell Dr F. Turco

Queensland Museum Business Manager

Dr F. Turco Dr G.B. Monteith

Queensland Museum (geoff.monteith@bigpond.com)

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Subscriptions are payable in advance to the Business Manager, The Australian

Entomologist, P.O. Box 537, Indooroopilly, Qld, Australia, 4068.

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ENTOMOLOGICAL SOCIETY OF QUEENSLAND

Membership is open to anyone interested in Entomology. Meetings are normally

held at the Ecosciences Precinct, Dutton Park, at 1.00pm on the second Tuesday of

March-June and August-December each year. Meetings are announced in the

Society's News Bulletin which also contains reports of meetings, entomological

notes, notices of other Society events and information on Members' activities.

Enquiries relating to the Society should be sent to the Honorary Secretary,

Entomological Society of Queensland, P.O. Box 537, Indooroopilly, Qld, 4068.

Cover: A male of Megacmonotus magnus (McLachlan 1871), one of the largest of

the Australian members of the lacewing family Ascalaphidae. Ascalaphids are

sometimes known as “owl flies” and many are partly daytime active. This species

has a wing length of up to 45 mm and is very widespread in Australia, being

recorded from all mainland states except Victoria and South Australia. The strange

process jutting up from the base of the abdomen is found in many male ascalaphids

and is of unknown function.

The illustration is reproduced by permission from CSIRO’s Insects of Australia and

is by the late Mary Quick, one of the many talented artists who worked in the 1960s

on the hundreds of new insect illustrations for the first edition of this classic work.

Australian Entomologist, 2012, 39 (4)

COURTENAY NEVILLE SMITHERS

29 August 1925 — 12 May 2011

210 Australian Entomologist, 2012, 39 (4)

COURTENAY SMITHERS MEMORIAL ISSUE

This special issue of the Australian Entomologist is dedicated to the life of

Courtenay Neville Smithers, who passed away in Sydney at the age of 85 on

May 12, 2011. He was a big, generous man whose life encompassed three

continents and whose interests in entomology were just as broad. Our

frontispiece portrait shows him as Deputy Director of the Australian Museum

in 1968, a position he relinquished to return to hands-on entomology.

To explore his life we have four biographical pieces. Courtenay’s wife (and

collecting companion) Aletta, always known as Smila, touchingly tells the

little-known story of his early life in Africa and England, and how his drive

to study natural history evolved. David Britton, who now sits in Courtenay’s

old office at the Australian Museum, describes Courtenay’s career at that

institution, how he revamped the displays and brought entomology to the

public in many ways. David also compiled the list of Smithers’ patronyms

and dug into AM archives for images of the camera-shy subject. Tim New,

from La Trobe University, himself a psocid worker, describes Courtenay’s

prodigious output of research on this group of tiny insects which he had

consciously chosen as a life-long study in Rhodesia in the early 1950s. John

Peters, a British lepidopterist who came to Australia a year after Courtenay,

describes his energetic attempts to record and explain the migratory patterns

of Australian insects. John, a close friend of Courtenay’s, also unearthed an

unpublished manuscript of Courtenay’s dealing with the uplift of butterflies

into atmospheric thermals and the implications of this on movement of

insects from Australia east into the Pacific. This is included.

The other seven papers are by friends and colleagues of Courtenay who

dedicate their studies to his memory. Five of them name new species after

Courtenay.

We thank all the authors for their contributions to this special volume. At 128

pages, it is the largest volume of the Australian Entomologist produced in its

38 years. This could not have been done without extra support. Thirty-seven

subscribers gave almost $1000 to help. Thank you all. Those who wished to

be named are: R. Bashford, I. Endersby, S. Ginn, R. Grandison, C. Hood, T.

Houston, J. King, R. Kohout, R. Lachlan, C. Lambkin, J. Majer, R. May, G. Maynard,

G. Monteith, B. Moore, M. Moulds, K. Murray, G. Owen, J. Peters, L. Willan, J.

Yates.

We are especially grateful to Courtenay’s home institution, The Australian

Museum in Sydney, for its most generous support of this special issue

honouring one of its most distinguished members of staff.

Australian

museum

nature culrure discover

Australian Entomologist, 2012, 39 (4): 211-220 211

COURTENAY SMITHERS — BEFORE AUSTRALIA

ALETTA (SMILA) SMITHERS

2 Wolsten Avenue, Turramurra. NSW 2074

Abstract

The entomologist, Courtenay Neville Smithers (1925-2011), was born in South Africa and

schooled both there and in England, developing an early interest in natural history. After serving

in World War II, he returned to South Africa and trained at universities in Pretoria and

Grahamstown, culminating in an MSc thesis on citrus entomology. While working in various

applied posts in Zimbabwe (then Rhodesia) and South Africa on tsetse fly, acacia entomology

and general agricultural pests, he developed an interest in taxonomy of Psocoptera and moved to

the Australian Museum in Sydney, as insect curator, in 1960.

Courtenay Neville Smithers was born on 29 August, 1925 in Pretoria, South

Africa. His family were of English descent and lived in Natal. His father,

Robert Francis Smithers, was a public servant, later magistrate, who was also

heavily involved with sports administration. At age eight his father died and

in 1938 his mother, Matilda Rebecca (Tilly), with her four daughters and

Courtenay made their home in England, where the eldest married daughter

lived. Courtenay’s four grown up brothers remained in South Africa. At this

time he was too old to sit for the exam to get him into a school where he

could qualify for university so he went to the local school. This particular

school had a teacher who was interested in soccer and the “new” boy was

soon in their first team and, as an under 13 team, they never lost a match. The

boys were excused class, no matter what the subject or time of the day, to

practise soccer. He was selected for special training by soccer scouts who

were searching for likely lads for professional soccer clubs when they

finished school. Courtenay’s mother did not approve of soccer as a career so,

with the help of the Methodist minister and a friendly headmaster, he was

sent to a small private school. (Courtenay’s passion for soccer remained with

him for the rest of his life). There was little money so his mother could not

afford the fees required for science lessons. He was a good student and

managed to pass the Oxford School Certificate. He was a prefect and,

continuing with his music lessons, played the piano for school assembly.

During the nights he had to be on fire watch and in the mornings go to class

as usual. In 1943, aged 18, he was called up (Fig. 1).

Courtenay was sent to Northern Ireland to train with the 1“ Battalion of the

Royal Ulster Rifles, 6" Airborne Division. Rigorous training across the

mountains and swamps of County Tyrone prepared Courtenay for his time

ahead. Transported in gliders on D-day (6 June 1944), the troops landed

behind the German coastal defences in Normandy. The following morning he

was captured by the Germans and, after many months in railway cattle

carriages and unbelievable hardship, eventually ended up in a POW camp in

Poland to work in a coal mine. When the Russian army was approaching the

212 Australian Entomologist, 2012, 39 (4)

whole camp started on a march to the south. Most of them died as it was in

the middle of a European winter with virtually no food. The march eventually

reached Czechoslovakia, where US soldiers released them in late 1945.

Courtenay’s survival was mainly because of the fitness he acquired while

training to be an airborne soldier and his stubbornness to make it to life after

the war. His brothers had fought with the South African forces.

As a schoolboy he became interested in natural history. His mother did not

approve of this because, as with soccer, she felt playing with insects was no

way to earn a decent living. He nevertheless roamed the English countryside

collecting insects and breeding them in the garage. He became an avid

birdwatcher and spent his spare time in the local library learning all he could

about animals in general. One of his sisters gave him a copy of A.D. Imms’

classic General Textbook of Entomology and with it he taught himself to use

the keys to identify the insects and learn general insect biology. By the time

he came out of the army in 1947 his mind was made up. He and his family

had returned to South Africa after the War so he worked at any sort of job,

mainly selling motorcar parts, to eke out enough money to go to university.

With a small loan from the ex-servicemen’s league and living with his

mother, he went to Pretoria University to study BSc Agriculture, a 4-year

course. After two years of trying to study in a language foreign to him

(Pretoria was then an Afrikaans-speaking university), he managed to pass but

realised that he knew enough entomology to teach his teacher, so he

transferred to Rhodes University in Grahamstown, Eastern Cape. His two

years at Pretoria did not qualify him for the formal science degree but they

allowed him Ist year. That year, 1949, he met Aletta du Preez, whose home

was in Rhodesia but was in her third year at Teachers’ Training College in

Grahamstown. She’d been known since babyhood as “Smila” because of her

happy disposition and the name stuck for life.

After he graduated he decided to try for an MSc funded by African

Explosives (who also made all the chemicals for agriculture). This degree had

very little course work; it was mainly practical research with a thesis at the

end and covered the years 1952-53. There is a very large citrus plantation

near Port Elizabeth where he was given accommodation and assistance to

undertake a study of all insect pests of citrus. During the season every tree

was covered by tarpaulin and cyanide gas was pumped into this “tent”.

Courtenay had placed covers on the ground beneath each tree to catch

everything that had been living in the canopy. When the covers were

removed next morning the catch was stored in jars to be sorted during the

day. Many hundreds of these samples were taken during the season. At the

time he (mentally) thanked Imms for preparing him for sorting all these

insects and this also stood him in good stead when he had a whole insect

collection in his charge at the Australian Museum in later life. Smila was

teaching in Rhodesia at the time and she travelled to South Africa during her

school holidays to type the thesis, having learned to touch type by

Australian Entomologist, 2012, 39 (4) 213

correspondence especially to do this. Courtenay had neither time nor money

to do it himself. Courtenay sorted the insects and wrote it all up, getting

specialists from all over the world to identify to species the insects in their

particular fields. ;

Fig. 1. Courtenay Smithers aged 18 and in British Army uniform, with his mother

Matilda in England in 1944.

214 Australian Entomologist, 2012, 39 (4)

Now the aim was to find employment. Having a 1" class pass in both degrees

in entomology and botany, Courtenay was offered some jobs, the first being

at the Horticulture Department just outside Pretoria. He turned this down, not

wanting to live too close to his authoritarian mother! The next one was a

better paying job but with an insecticide company. He turned this down for

obvious reasons. He then successfully applied to the Southern Rhodesian

Government for a post in the Tsetse Fly Eradication programme.

The reason for eliminating tsetse flies (Glossina spp.) was to enable large

tracts of lowveld (bush) country to become habitable for humans. Tsetse flies

spread sleeping sickness in humans and, as the veterinary disease nagana, it

kills most domestic animals. The local African population depends on cattle

and goats for their living so they could not live in this huge area of country

without them. The tsetse fly is small, much like an ordinary housefly, but it

has a bite similar to that of a tabanid. The idea at that time was to eradicate

all wild mammals that carry the disease but are immune to the sickness.

Large numbers of African men were employed to shoot all targeted

mammals. The entomologist in charge paid each man each month by

counting the tails of the dead animals. They were also given standard maize

meal and salt for the month and enough bullets to go on with. Their meat

supply came from the animals they shot. Courtenay was also employed, apart

from supervising the monthly gathering of all the hunters, to study the tsetse

fly populations, where they were most numerous and so on. These counts

were made by driving slowly (the pace of a walking mammal) with the

workers collecting all the flies that settled on the green coloured truck in

small butterfly nets (Fig. 2). Every tenth of a mile the truck stopped and the

collected flies were counted and sexed. There were usually no roads and they

drove through the bush following blaze marks on trees, often with one of the

African staff walking ahead to find the next marked tree. Their first car (Fig.

3) was understandably kept for town use!

For Courtenay this was a wonderful job. He saw all the animals in the bush at

close quarters. The elephants wreaked damage to plant life as they meandered

through the bush, warthogs dug holes, lions lived freely among the other

animals, antelopes of all types were plentiful and it was a wonderful life for a

man used to life in England or in a city in Africa. He was too busy doing his

daily chores to think about normal insects and in life afterwards he always

regretted not being able to collect insects in that uninhabited area of Africa.

Later he was moved to a more civilised part where aerial spraying was tried.

Here the country was more open and so long tracts of bush and valleys

considered suitable were measured out by compass bearings and marked by

long poles with balloons at the ends for small crop-dusting planes spraying

insecticide to kill the tsetse flies (and all other insects too). From earlier

studies it was found that tsetse flies spent daytime hours in these valleys and

could thus be more easily destroyed there instead of trying to spray the whole

Australian Entomologist, 2012, 39 (4) 215

Fig. 2. Courtenay Smithers at the wheel of the green truck used in testse fly transect

surveys at Sebungwe, near Bulawayo, Rhodesia in early 1950s with the help of

African assistants.

Fig. 3. Courtenay and Smila with their first car, outside the school where Smila taught

near Salisbury, Rhodesia, in the early 1950s.

216 Australian Entomologist, 2012, 39 (4)

countryside. There are people living in these places now, so presumably some

of these practices were successful.

Courtenay really loved the life (Fig. 4) and especially being in close contact

with the animals and the birds. He realised that there was no future in trying

to kill all mammals; it was wrong policy and he did not want to be involved

with that type of work. He was also not really a field worker by nature and

needed a more academic type of work. He applied for and got a post in Natal

at the Wattle Research Institute, affiliated to the University of

Pietermaritzburg. At that time Australian wattles (Acacia decurrens Willd.)

were planted in huge plantations all over the hills. The trees grew in rows for

miles and they were pruned by cutting off the side branches at the bottom, so

that the trees grew very tall and straight. After 10 years they were cut down

and the bark removed and the tannin extracted to be used in the leather

tanning industry. (These days artificial tannin is used). Courtenay was left in

charge of the Entomology Department while the chief entomologist went to

Europe on 6 months leave. This was an opportunity for Courtenay to do

something just for himself. He chose the main pest of the wattles, the

bagworm moth Kotochalia junodi (Heylaerts) and its parasitic wasp, the

ichneumonid Sericopimpla sericata (Kreichbaumer). The bagworm lives on

other acacia species in Africa but the wattle plantations were an ideal single

species planting and he found it an ideal life history study. These pest

bagworms are sometimes so numerous that they can defoliate whole trees,

and from the air patches can be seen where large numbers of trees have been

stripped.

The newly hatched larvae leave the parent bag and waft in the breeze on a

silken thread to a likely host tree where it spins a little cocoon. The wingless

female lives in this all her life, never emerging; she eats the wattle leaves and

uses the small stalks to build her bag, enlarging it as she grows. The male

lives in a similar bag but, when adult, it flies in search of a female. They

copulate through the end of her bag. When the eggs hatch, the larvae leave

the bag and waft away to find a suitable place to live. In the lab a great many

bagworms were cut open; when carefully done the insect inside continues life

apparently unconcerned. During the season the life history was completed

and a paper on the study was prepared and published in the journal of the

Entomological Society of Southern Africa. On his return the chief

entomologist was not too pleased that his junior did all this without his

knowledge. Anyway that was the end of Courtenay’s decision making.

Thereafter he had to toe the line and do as he was told. The work continued.

There was more aerial spraying of plantations and cooperation between the

wattle plantation owners. This was the second time that Courtenay was

sprayed with insecticide while supervising aerial spraying not wearing

protective clothing. Pietermaritzburg is a very pleasant town near the

Drakensberg Mountains and would have been a lovely place to make a home.

Australian Entomologist, 2012, 39 (4)

Fig 4. Courtenay with butterfly net and rifle in the field at Sebungwe.

218 Australian Entomologist, 2012, 39 (4)

Courtenay and Smila were married in Pietermaritzberg in 1954 (Fig. 5) and at

this time she taught in a beautiful school: St. Charles College. He became

more and more restless at work and frequently disagreed with his senior till

one day things became very heated and Courtenay went home and discussed

the situation with Smila. The next day he gave notice. They lived for six

months on her teacher’s salary while they looked around for a new post.

It was soon after they were married that they discussed his choice of an insect

group to study. He was drawn to parasitic wasps. His old professor, John

Omer-Cooper at Rhodes University, was a water beetle specialist and he was

keen on a student to follow in his footsteps. Smila really liked these insects

but Courtenay wanted a group that had not been “messed about” by

specialists. He decided on Psocoptera, a group that he thought was small in

size and in the number of genera. They spent every weekend in the country

searching for psocids, but soon began to find suitable methods of identifying

the sort of trees where they were likely to be found and methods of

collecting. While he was unemployed he was given laboratory space, use of

their library and a key to rooms at the Natal Museum. This was a marvellous

opportunity to begin his card indexes, to write to all the living people still

working in the group and to find papers of everything ever published. This

continued all his life.

Eventually he decided that he would like to live in Rhodesia, where Smila

had grown up. He went back to Salisbury (now Harare) in 1956 to work in

the Entomology Department and Smila started teaching at David Livingstone

School, a short walk away from the Research farm where all the agricultural

offices were and experimental work on crops was carried out. Rhodesia was a

wonderful country to live in while David Livingstone was a beautiful modern

school and life was full of promise. Work for Courtenay and the other

entomologists consisted of studying all pest species connected with

agriculture and also other duties like handing out permits for introducing new

plants, inspecting plant nurseries and so on. Work was pleasant but there is

always a problem area. Farmers would consult the department about pests:

army worm, tobacco beetle, maize stalk borer, water weeds on dams, even

ageing maggots on corpses and so many other problems. But the staff could

never run experiments on pests, usually only suggesting insecticide remedies

and hoping for the best. All this time psocids were the evening occupation for

both of them. Every weekend was spent collecting and sorting. Life was

good. All holidays were spent in game reserves, on rivers looking at water

birds, always with collecting materials and butterfly nets.

Courtenay was making a name for himself as a specialist in psocids. He was

also involved in bird watching groups, going on field trips with other

international zoologists, mammalogists, ornithologists and botanists, many of

whom stayed with them in Salisbury. It was a place where naturalists liked to

visit to see the wild life, to see the insect collections and life was good. The

Australian Entomologist, 2012, 39 (4) 219

—

hae, APS oe se

hoan o ai

Fig. 5. Courtenay and Smila at their wedding in Pietermaritzberg, South Africa in

1954.

220 Australian Entomologist, 2012, 39 (4)

climate in Salisbury was excellent, people were friendly, servants affordable

and entertaining was no problem, but there is always a niggling and the

problem was that Courtenay did not like taking orders and doing other

seniors’ bidding. He became restless once more and tried to find an

occupation somewhere where he could make his own decisions.

The Australian Museum was the answer! So once more they packed up,

Smila very reluctant to leave home to go into the unknown. At least in

Sydney people spoke the same language and Courtenay had his own

department. This was 1960 and was to be their final move. Museum work and

life in Australia suited them well.

Australian Entomologist, 2012, 39 (4): 221-230 221

COURTENAY NEVILLE SMITHERS — HIS CAREER IN

AUSTRALIA

DAVID R. BRITTON

Australian Museum, 6 College Street, Sydney, NSW 2010 (Email: dave.britton@austmus.gov.au)

Abstract

Courtenay Smithers was an entomologist on the staff at the Australian Museum from 1960 until

his retirement in 1985. He remained as a senior fellow with the Museum until his death in 2011.

During this time he was actively involved with the Australian entomological community in many

areas, as well as with the general public and in other areas of the natural sciences.

Museum life in the 1960s and 70s

Anthony Musgrave was the Curator of Insects at the Australian Museum

from 1920 until his unexpected death from a heart attack on June 4, 1959.

Musgrave had planned on retiring in 1960-61, so the Museum had advertised

to replace him in anticipation of this retirement. After a diverse early

entomological career in South Africa and Rhodesia (now Zimbabwe),

Courtenay Smithers was looking around for employment outside Africa (A.

Smithers 2012) and received responses from the Australian Museum and

CSIRO. The Museum offered: him a job while Musgrave was still alive, but

Courtenay never got to meet him. He took up the post of Curator of Insects at

the Australian Museum on February 12, 1960 (Anon. 1960).

Courtenay’s first impressions of Australia and Sydney were not at all

favourable. They landed in hot weather and were immediately driven from

the airport at Mascot, which he recalled as being ‘odoriferous’, to a hotel

booked by the Museum. The hotel was ‘... less than desirable could I say. It

was unliveable actually’. There were two hotels of the same name and the

Museum clerk had accidently booked them into the wrong one. The Smithers

endured one night there and then moved to a boarding house in Coogee,

where they stayed for several months until they could find a better

arrangement. After this things improved greatly.

At this time the Entomology Department staff comprised Courtenay, David

McAlpine, who was the Assistant Curator, and a technical assistant, Romola

Wilkinson. Subsequent technical assistants included Klara Kota (1962),

Dianne Raffles (1964), Janet O’Hare (1965), Janet Walsh (1965), Robyn

Jeffrey (née Pettett) (1967), Clare Trickett (1970), Lynn Hoskins (1971),

Robyn Brewer (née Spalwit) (1974), Christine Horseman (1974) and Barbara

Duckworth (1975). Because of the work load in the Department, Courtenay

was able to get a fourth position for a technical officer, a position which was

filled by Geoff Holloway in 1966 and later by Barry Day in 1978, when

Geoff became Collection Manager of Entomology. This position was largely

needed to assist with the numerous enquiries that arrived in the Department.

Other staff working as research assistants, often on specific projects, included

John Peters (1968), Margaret Schneider (1974), S.P. Kim (1974), Justine

222 Australian Entomologist, 2012, 39 (4)

O’Regan (1974) and Greg Daniels (1976). Mike Gray was appointed as

assistant curator for Arachnology in 1968. The staff members in 1981 are

shown in Figure 1.

Fig. 1. The Australian Museum Entomology Department staff in 1981. From left:

David McAlpine, Barry Day, Deborah Kent, Geoff Holloway, Robyn Brewer and

Courtenay Smithers.

Overall, there were far fewer staff at the Museum compared with the present

institution and the workplace had a strong family atmosphere. The Director

of the Museum at this time was John Evans, who was also an entomologist.

Evans had a strong vision for development of research and exhibitions at the

Museum and, under his leadership, the institution went on to become an

internationally recognised force in the natural history sciences. This climate

of cooperation and improvement was ideal for Courtenay and resulted in

significant growth of the insect collections (including by public donations:

e.g. Fig. 2) and improvement in the way they were housed (Britton 2011).

Courtenay also initiated the design of new modular cabinets for the collection

(Britton 2011). He noted that one of the key design criteria was that the

empty cabinet shell could be lifted by two average women, as Public Service

regulations held that women on staff weren’t allowed to lift above a certain

weight. This is why Australian Museum cabinets have a somewhat unusual

and elongate profile, with the drawers arranged in two tiers with each tier

holding seven drawers.

Australian Entomologist, 2012, 39 (4) 223

Fig. 2. Courtenay Smithers (right) at a function in the Australian Museum galleries in

1978 to receive the donated butterfly collection of David Rushworth (left).

One of the areas where development occurred was in field work. When

Courtenay arrived at the museum there was only one Museum vehicle, which

was a small delivery van. Shortly afterwards, the Museum managed to obtain

a 4WD vehicle, which was probably a first for a museum in Australia. A

collecting trip was planned for Cape York Peninsula and, fortunately for

Courtenay, one of the occupants pulled out leaving space for him to go. He

had only been in Australia for a few weeks and had no collecting equipment,

so he had to rush to equip himself. The trip was hard work and they were

away for almost three months. They followed the telegraph maintenance

track, getting as far as the then flooded Archer River (Cogger 1961).

Subsequent trips over the next few decades took him to Lord Howe Island,

Norfolk Island, New Zealand, Papua New Guinea and many other islands in

the Melanesian arcs. He saw no boundary between work and the rest of life,

so wherever he went he and his wife Smila collected. Indeed Smila was

probably the more productive when it came to collecting.

From 1967 to 1970, Courtenay took on the role of Deputy Director at the

Museum (Strahan 1979), plus running the Entomology Department and being

Secretary for the International Congress of Entomology Organising

Committee (Britton 2011). The work load and responsibility must have been

tremendous and Courtenay recalled ‘I don’t know that I lost all my hair then

but I certainly lost a lot of it. A lot of it went grey.’ The Deputy Director’s

role was largely administrative and Courtenay felt that his skills as an

224 Australian Entomologist, 2012, 39 (4)

entomologist, including over 15 years of research on Psocoptera, were wasted

in the Deputy Directorship. He characterised the administrative role as ‘Here

I am writing to the Public Works Department asking them to fix the toilets,

when I have now arrived at the point where I can make a contribution (to the

systematics of Psocoptera).’ In 1970 he stepped down and continued on as

Principal Curator until his retirement in 1985 (Britton 2011).

Courtenay’s contributions to insect taxonomy are discussed by New (2012),

but as well as publishing on Psocoptera, Neuroptera and Mecoptera he

managed to cover a diversity of other topics relating to natural history. He

published articles on all of the major insect Orders, covering topics such as

conservation, behaviour, life history, ecology, economic entomology and

taxonomy, as well as introductory manuals for insect study and collecting. A

full list of refereed publications can be found in Britton (2011), with extra

minor writings listed at the end of this volume. He was an early supporter of

the Australian Entomological Magazine (now Australian Entomologist) when

it was established by Max Moulds in 1974 and published many articles in this

journal.

Exhibitions and Public interaction

One of Courtenay’s early plans for the Museum was to prepare new insect

exhibits to take advantage of the innovative skills of a young, talented and

very active team working in Exhibitions and Preparation Departments at that

time. It was not until the 1980s that this goal was achieved. The project team,

consisting of a core of Courtenay, Bob Ross-Wilson, Roland Hughes and

Janette McLeod, commenced meeting in late 1982. The development and

construction took over two years, with the gallery opening in early 1985 with

a total of 45 display cases featuring insects, spiders, worms, molluscs and

other invertebrates (Fig. 3). Courtenay was charged with coordinating text

content as well as helping source specimens, either through field work with

Science and Exhibitions staff or by purchase from various specimen dealers.

The project team meeting notes at the time detail just how many different

people were drawn into the development of this gallery, including

entomologists, arachnologists, collectors and natural historians such as

Densey Clyne, Martyn Robinson, Glenn Hunt, David McAlpine, Mike Gray,

Geoff Holloway and George Hangay.

Part of this gallery is still on display and it is still well utilised, as it

represents one of the few places in Australia where you can see a

comprehensive display detailing all of the insect orders present in Australia,

along with appropriate scientific interpretation and, for many groups, real

specimens. The gallery was also considered very advanced at the time,

because it not only sought to outline the diversity of Australian insects and

invertebrates but also had sections dealing with ecology, environment,

behaviour, conservation, toxicology and other themes relating to these

organisms.

Australian Entomologist, 2012, 39 (4) 225

Fig. 3. A case on cicadas from the new Australian Museum insect gallery opened in

1985. It showed how models, real specimens, photographs and dioramas could be

combined with good scientific information to give a striking overall effect (Australian

Museum photo).

Courtenay’s research interests also expanded, including projects investigating

butterfly migration, the results of which are discussed by Peters (2012). This

project was notable in that it had a strong public involvement, with

‘collaborators’ from all over eastern Australia involved in both marking and

recapturing butterflies. They remember it fondly and it helped network a

large number of amateur enthusiasts, some of whom went on to become

entomologists later in life. Shane McEvey recalls that because he was on

Courtenay’s file as a collaborator, it meant that even as a teenager from

Melbourne he could visit Sydney, drop into the Museum and get to see the

collection and talk to real scientists.

Courtenay’s interest in migratory behaviour illustrated another facet of his

personality that contributed to his success as a scientist and natural historian.

He was a compulsive note taker and always jotted down details on insect,

bird and other natural events around him. He knew that it was not possible to

remember key details, such as the direction of a flight of butterflies, so all of

this got recorded. For example, he recorded daily the birds at the feeder at his

home in Turramurra.

Role in formation of the Australian Entomological Society

When Courtenay arrived in Australia from Rhodesia, he was surprised that

there was no national society equivalent to the Entomological Society of

Southern Africa and that there was no national forum for Australian

entomology (Marks and Mackerras 1972, Britton 2011). His Director, John

Evans, who was then the Australian representative on the Permanent

Committee of the International Congresses of Entomology, supported the

226 Australian Entomologist, 2012, 39 (4)

view that a national body was needed and discussed this with Bruce Champ

from the Entomological Society of Queensland (Marks and Mackerras 1972).

Courtenay and Champ arranged a meeting of entomologists during the

August meeting of the Australian and New Zealand Association for the

Advancement of Science (ANZAAS) in Sydney. It was attended by 30

entomologists and chaired by Courtenay. As recorded in Marks and

Mackerras (1972), discussion was particularly vigorous and Courtenay

somewhat fondly recalled one irate NSW supporter of ‘State rights’ referring

to him as ‘that bastard Smithers’.

Marks and Mackerras (1972) documented the subsequent formation of the

Australian Entomological Society (AES). Courtenay regularly attended the

AES conferences (Fig. 4), was very proud of his role in the Society’s birth

and was granted an Honorary Life Membership in 1983 (Britton 2011). He

acted as inaugural Editor of the AES News Bulletin (now Myrmecia) from

1965-68 and was AES President from 1977 to 1980.

Fig. 4. At Mt Field National Park, Tasmania, on an outing from the annual conference

of the Australian Entomological Society in 1985. From left: Margaret Schneider

(UQ), Courtenay Smithers (AM), Tom Weir (ANIC), Greg Daniels (UQ) and Geoff

Holloway (AM). All except Tom Weir had worked with Courtenay at the Australian

Museum (Photo: Geoff Monteith).

Role in the 14" International Congress of Entomology, Canberra

Courtenay attended the International Congresses of Entomology held in 1964

in London and in 1968 in Moscow (New 2012) and dryly noted that he

‘learnt how to run a congress’ at the former and ‘how not to run a congress’

Australian Entomologist, 2012, 39 (4) 227

at the latter. Courtenay and Doug Waterhouse (CSIRO Entomology) drafted a

formal submission on behalf of the Executive of the AES to the Council of

the Australian Academy of Science, requesting that the Academy issue a

formal invitation in conjunction with the AES to the Permanent Committee

of International Congresses of Entomology (Wharton and Marks 1968). The

Academy agreed to be joint sponsor for the congress and Waterhouse was

nominated to be the Australian representative on the Permanent Committee.

An Advisory Committee was appointed by the Academy, with Courtenay as

secretary to the Committee (Marks 1968). The recommendation that the 14"

Congress be held in Australia was formally accepted by the Permanent

Committee at the final plenary session of the Moscow Congress (Anon.

1968) and Courtenay became the Congress Secretary.

Much of the coordination of Congress, inviting speakers, drawing up a

program, promotion of the Congress and many other tedious and detailed

tasks, was done with the help of Smila. Courtenay set up a makeshift office in

a tiny garden shed in the backyard of their home in Turramurra for handling

Congress communications. He sent out a first promotional Circular on the

Congress to almost 10,000 recipients in early 1970, along with providing

advertisements to the major journals such as Nature and Science. Regular

updates on the progress of the Congress Committee were published in issues

of the AES News Bulletin. By mid-1971, almost 1,800 delegates had given

expressions of interest (Smithers 1971), all of whom received correspondence

back from the Smithers’ garden shed office. Courtenay shifted his office to

Canberra in early 1972 to be closer to the rest of the Congress working group

from CSIRO, Australian National University and other institutions in

preparation for the final stages in organising the Congress, which ran in late

August, 1972. A total of 1,323 attendees were present at the Congress, with

720 papers presented at over 21 Symposia (invited papers) and 15 Sections

(submitted papers) (Smithers 1972).

Other interests

Before Courtenay and Smila left Rhodesia, they saw a Doberman bitch at an

obedience trial and fell in love with the breed. They decided that when they

got a dog it would be a Doberman. When they arrived in Australia, they

discovered that there were very few examples of the breed in the country and

it took a while to find a good one, but they eventually did. It was a

characteristic of Courtenay that he never just dabbled with a new interest; he

went in boots and all. He took a course on becoming a show judge for the

Doberman breed and, when overseas on entomological business, managed to

find time to visit Germany and talk to the German Doberman breeders. He

ended up doing the commentary at the Royal Easter Show in Sydney for the

Massed Obedience Display, which consisted of over 100 dogs in the arena at

one time doing obedience trials. This drew upon his military background, as

he had received training in radio broadcasting during WWII.

228 Australian Entomologist, 2012, 39 (4)

A similar result occurred when Courtenay started keeping bee hives in the

backyard at Turramurra. He tried them at Tuglo (see below), but they did not

do well and died out. When Smila became sensitized to bee stings he set up

his hives at his son’s property near Wollombi, where they thrived. The honey

was sold to a health food shop or used to “bribe” neighbours. As a result of

his developing interest in this area, he published ‘Backyard Beekeeping’

(Smithers 1987), a popular guide which was updated and reprinted in 1992

and 2011 under different publishers. He regularly lectured to bee clubs and

his phone number was listed as a catcher of feral bee swarms.

Courtenay loved classical music, which was always a loud accompaniment to

his desk work at home. He played piano for relaxation and, as Smila recalls,

‘whenever anything annoyed him he would take it out on the piano!’

‘Tuglo’, the family’s nature retreat

In 1972, Courtenay and Smila purchased ‘Tuglo’, a largely forested 200 ha

property on the southwestern slopes of Mt Royal, 36 km north of Singleton

and not far from Barrington Tops. The property became a regular retreat for

Courtenay and his family and he recalled spending 105 weekends there

between August 1976 and April 1979 (Smithers 1981). He took an interest in

almost all aspects of its natural history. He collected data on butterflies, birds

and mammals on the property from 1972 onwards, as well as collecting

various insects with Malaise traps and other methods. This resulted in some

large collections of insects, which are held in the Australian Museum

collections, as well as a series of papers on the ferns (Smithers 1978), birds

(Smithers 1980), Papilionoidea (Smithers 1981), Megaloptera, Mecoptera

and Neuroptera (Smithers 1993), Peripsocidae (Smithers 1994a) and

Hesperiidae (Smithers 1994b).

Courtenay regularly referred to the property as ‘Tuglo Wildlife Refuge’ in

print. Though it was always intended as a wildlife refuge and had National

Parks signs declaring it was a wildlife sanctuary, the property was never

formally gazetted as such. It was visited by many naturalists and natural

history societies over the years, adding to Courtenay’s own observations on

its plants, birds and other wildlife. When the children, Graeme and Hartley,

grew older they spent much time riding horses on the property and nearby

ranges. Courtenay and Smila also planted South African proteas on the

property. When their elder son, Graeme, graduated from his degree in

horticulture, he purchased the adjacent property to set up a commercial

flower farm growing proteas and other commercial species while living on

Tuglo. Graeme has since retired from supplying flowers to florists but is still

resident part-time on the property.

Retirement from the Museum

Courtenay retired at 60 in 1985 from his salaried position as Principal

Curator. As with many other entomologists, Courtenay’s retirement (Fig. 5)

Australian Entomologist, 2012, 39 (4) 229

from a paid position simply meant that he had much more time to get on with

his entomological research without the interruptions of administration and

other onerous tasks that come with paid work. He set up a lab on the back

verandah of the house in Turramurra and did most of his research work at

home. He always came in to the Museum for one day each week, where he

could access the research library and catch up with Museum life. He

published over 80 refereed articles and books after retirement (Britton 2011)

and continued his writing and research right up until his death on May ist

2011.

Fig. 5. Courtenay Smithers (centre) in retirement in 2006 with his wife Smila and

Barry Day, assistant in entomology at the Australian Museum for many years (Photo:

Max Moulds).

Acknowledgements

Details for this biography were largely drawn from autobiographical notes

that Courtenay left with Smila Smithers, from the Australian Museum

Archives Department, including transcripts of an oral history taken in July,

2004, and from personal conversations with Courtenay. In particular, I would

like to thank Vanessa Finney and Rose Docker from the AM Archives

Department and Carl Bento and James King from AM Photography for

scanning and preparing photographic material from the Archives. I would

also like to acknowledge the assistance of John Peters, Max Moulds, Shane

McEvey, Smila Smithers and Geoff Monteith in recalling details about

Courtenay and suggesting sources of information.

230 Australian Entomologist, 2012, 39 (4)

References

ANON. 1960. New Curator at Museum. Australian Museum Magazine March: 157.

ANON. 1968. XIV" International Congress of Entomology. Australian Entomological Society

News Bulletin 4(3): 50.

BRITTON, D.R. 2011. Dr Courtenay Neville Smithers 1925-2011. A tribute. General and

Applied Entomology 40: 25-34.

COGGER, H.G 1961. An expedition to Cape York Peninsula. Australian Museum Magazine 13:

362-367.

MARKS, E.N. 1968. Report from the executive. Australian Entomological Society News Bulletin

4(2): 24-27.

MARKS, E.N. and MACKERRAS, I.M. 1972. The evolution of a national entomological society

in Australia. Journal of the Australian Entomological Society 11(2): 81-90.

NEW, T.R. 2012. Courtenay Neville Smithers: his scientific contributions to the ‘small orders’

of insects. Australian Entomologist 39(4): 231-237.

PETERS, J.V. 2012. A tribute to Courtenay Smithers and his butterfly migration studies.

Australian Entomologist 39(4): 239-246.

SMITHERS, A. 2012. Courtenay Smithers — before Australia. Australian Entomologist 39(4):

211-220.

SMITHERS, C.N. 1971. 14th International Congress of Entomology. Australian Entomological

Society News Bulletin 7(3): 93-94.

SMITHERS, C.N. 1972. Report on Congress. Journal of the Australian Entomological Society

11(3): 177-184.

SMITHERS, C.N. 1978. A preliminary list of the ferns of Tuglo Wildlife Refuge. Hunter

Natural History 10(4): 189-191.

SMITHERS, C.N. 1980. A preliminary list of the birds of Tuglo Wildlife Refuge. Hunter

Natural History 11(2): 41-45.

SMITHERS, C.N. 1981. A preliminary note on the Papilionoidea (Lepidoptera) of Tuglo

Wildlife Refuge, New South Wales. Australian Entomological Magazine 7(6): 91-96.

SMITHERS, C.N. 1987. Backyard beekeeping in Australia and New Zealand. Angus and

Robertson. Sydney; 118 pp; republished 1992, Kangaroo Press, Kenthurst, New South Wales; 80

pp; republished 2011, Rosenburg Publishing, Kenthurst, New South Wales; 80 pp.

SMITHERS, C.N. 1993. A note on the Megaloptera, Mecoptera and Neuroptera of Tuglo

Wildlife Refuge, New South Wales. Australian Entomologist 20(2): 67-71.

SMITHERS, C.N. 1994a. A note on the Peripsocidae (Psocoptera) of Tuglo Wildlife Refuge,

Hunter Valley, New South Wales. Australian Entomologist 21(1): 7-10.

SMITHERS, C.N. 1994b. A note on the Hesperiidae (Lepidoptera) (skippers) of Tuglo Wildlife

Refuge, New South Wales. Australian Entomologist 21(3): 103-109.

STRAHAN, R. 1979. Rare and curious specimens. An illustrated history of the Australian

Museum 1827-1979. The Australian Museum, Sydney; 173 pp.

WHARTON, R.H. and MARKS, E.N. 1968. Annual Report of the Australian Entomological

Society. Australian Entomological Society News Bulletin 4(1): 7-12.

Australian Entomologist, 2012, 39 (4): 231-237 231

COURTENAY NEVILLE SMITHERS: HIS SCIENTIFIC

CONTRIBUTIONS TO THE ‘SMALL ORDERS’ OF INSECTS

T.R. NEW

Department of Zoology, La Trobe University, Vic 3086

Abstract

Courtenay Smithers’ contributions to the knowledge of Psocoptera and other orders, notably

Neuroptera and Mecoptera, are summarised. They extended over some 50 years and established

him as a leading authority on these orders. His contributions to systematics, biogeography and

biology of Psocoptera, particularly within the Australian region, were accompanied by

compilations of global catalogues and bibliographies of immense value to others. His many

publications assure him of a persistent place in the history of psocopteran studies.

Introduction

Courtenay Neville Smithers (1925-2011) was known and respected widely as

an excellent naturalist with a keen, life-long sense of curiosity and

wonderment that led to a variety of scientific interests, together with an

enduring enthusiasm about communicating his passions to others. These

personal traits rendered him a highly effective publicist for entomology and

led to his writing of several books to introduce insects to the wider public.

His 1971 ‘Australian Insects in Colour’, with Anthony Healy, was a pioneer

in all-colour popular publication, while his ‘Handbook of Insect Collecting’,

first published in 1981, remains a useful and readable resource. His

‘Backyard Beekeeping’ has just been republished 25 years after it first

appeared in 1987. However, his major formal scientific contributions focused

on studying the systematics and biogeography of a commonly ignored insect

order, the Psocoptera, augmented by original contributions on two other

‘small orders’, the Neuroptera and Mecoptera. His studies on these insects

assure him of a lasting place in their documentation.

Psocoptera

Smithers’ taxonomic interest in Psocoptera (the barklice and booklice)

flowed from the 1950s (A. Smithers 2012), with his first papers on these

insects (in 1956) the source of a stream that reached more than 150

contributions over more than half a century and established him firmly as a

leading world authority for much of that period.

When I started to work on British psocids (in 1965), I wrote to Courtenay at

the Australian Museum for his advice, and his kind and encouraging response

— together with a set of his reprints then available — marked the start of a long

friendship founded in our mutual interests in ‘the other orders’. I met him

first in England in 1967 and next at the XIIIth International Congress of

Entomology in Moscow (August 1968), an occasion that marked perhaps the

first major international gathering of psocid workers, with several meeting for

the first time (Fig. 1); it was not until the 1980s that such a meeting of

psocopterists, always few in number, was held again. Following my arrival in

Australia in 1970, we continually exchanged ideas on psocids and lacewings.

232 Australian Entomologist, 2012, 39 (4)

Fig. 1. Gathering of psocopterists outside Moscow University, August 1968. Left to

right: A.M. Nadler (USA), T.R. New (UK), A. Badonnel (France), I.W.B. Thornton

(Australia), C.N. Smithers (Australia) (Photo: V.N. Vishniakova).

While most of his published work deals with systematics, distributions and

relationships of psocids, Courtenay was always aware that he was dealing

with dynamic living organisms and, somewhat unusually for that era, his

taxonomic studies flowed in large part from material collected during his own

field studies, rather than from specimens mostly collected by others and

submitted for identification. Biological oddities intrigued him greatly. One

such was his discovery of the remarkable phragmotic nymphs of Psilopsocus

mimulus Smithers (Figs 2-3), which resemble small bark beetles and live in

twigs, blocking the entrance to their tunnels with their heavily sclerotised

abdomens (Smithers 1995). His knowledge spanned the world fauna.

Among his first major self-appointed tasks was to bring together the then

very widely scattered literature on Psocoptera into a single, easily available

and convenient format: his ‘Bibliography of the Psocoptera’ (Smithers 1965,

with drafts circulated to most specialists for comment and augmentation) was

of incalculable value to tyros such as myself. His comment in the

introduction that it was ‘an attempt to provide a working tool which will

relieve students ... of the drudgery of compiling their own bibliographies’

was indeed pertinent. The Bibliography was followed by a World Catalogue

of species (Smithers 1967). The two works together provided, for the first

time, a firm basis for later studies; subsequent progress in understanding

Australian Entomologist, 2012, 39 (4) 233

Psocoptera over the last half century is largely due to Smithers’ Herculean

efforts in producing these accounts. It is worth remembering that these

compilations were made long before availability of computers, email and

even routine use of photocopiers — so that obtaining and checking the

numerous obscure references for verification could entail weeks of patience

awaiting international postage, or laboriously copying out details by hand

during sporadic visits to major institutional libraries. They were followed

(Smithers 1972) by publication of his Ph.D. thesis, a global synthesis of

psocid classification and phylogeny, with illustrated diagnoses of all genera

and families then known and discussions of putative relationships that set a

basis for much later consideration. It remains a potent reference source of

earlier information and diagnoses.

Figs 2-3. The wood-boring psocid, Psilopsocus mimulus Smithers: (2) adult male

from Lindfield, NSW (AM Paratype K68224); (3) final instar nymph with abdomen

modified to plug burrow, from same series (Photos: D. Britton, Australian Musem)

These landmark publications were revised later, with new keys to families

and genera (Smithers 1990), a revised Bibliography (with Charles Lienhard:

Smithers and Lienhard 1992) and Smithers’ magisterial joint volume

(Lienhard and Smithers 2002), as a world catalogue and bibliography,

updating these to comprise perhaps the single most significant publication

ever issued on the order. The classification used there was essentially that of

Smithers (1990) and pre-dated current opinion that the order is not

monophyletic but intertwined intricately (as ‘Psocodea’) with the true lice,

Phthiraptera (Yoshizawa and Johnson 2006).

Psocoptera had become much better known between 1967 and 2002, in no

small part reflecting the impact of the publications noted above in

collectively facilitating the perspective needed for others to study these

insects. The known richness of Psocoptera rose from 1605 species (in 197

genera) in 1964 to 4408 species (371 genera) by 2000, together with an

increase from 31 to 41 generally recognised families. Smithers was an active

contributor to this increase. Alone or with his collaborators (notably the late

Ian Thornton and in associations that led to global dominance of Psocoptera

234 Australian Entomologist, 2012, 39 (4)

Studies by Australians for some two decades), he described somewhat more

than 350 new species of psocids, across about 25 families. Many of these

came to his attention though extensive field work and collecting in Australia

and — with Thornton — during a major Australian Research Grants-funded

Survey during the 1970s of the various islands, including New Guinea and the

Melanesian Arcs, to the north and east of Australia.

While his first major regional synopsis dealt with the psocids of Madagascar

(Smithers 1964), his first paper from the Australasian region (following his

move from South Africa to the Australian Museum in 1960) dealt with

Philotarsidae from Macquarie Island (Smithers 1962). However, the account

that founded his reputation in the Australasian fauna was a major paper on

the New Zealand fauna (Smithers 1969), with 15 new species bringing the

country’s total to 43 species. His early African experiences assuredly whetted

his appetite for biogeographical puzzles centred in the ‘southern continents’

and the regional fauna remained his primary focus for many years. From the

early 1960s he published notable species additions and family synopses of the

Australian psocids and, by the time he prepared the entry on Psocoptera for

the Zoological Catalogue of Australia (Smithers 1996), he had described well

over a third of the Australian species then known. These included records of

several families new to the country, as well as a number of novel genera that

hinted at the considerable complexity and high levels of endemism of the

fauna. His successive chapters in the original (1970) and revised (1991)

editions of ‘The Insects of Australia’ revealed an increase from 120 to 299

species over that period. Regional accounts, such as for South Australia

(Smithers 1984) and (jointly) for Norfolk Island (Smithers and Thornton

1974) and Lord Howe Island (Smithers and Thornton 1975), were augmented

by substantial family accounts for the Melanesian Arc psocids that

collectively set a new perspective for their diversity and evolution within the

region.

Families such as Philotarsidae, Calopsocidae, Myopsocidae and Psocidae

proved to be far more diverse, intricate and biogeographically informative

than suspected before these studies began. Elucidating their features did

much to enhance understanding of the Australian fauna and how this had

developed. Many of the places visited in Melanesia had never been explored

properly for psocids, although the few taxa described sporadically from New

Guinea and elsewhere suggested the likely richness of those areas. The

fieldwork in remote areas was sometimes hazardous, frequently adventurous

and innovative. Courtenay recalled some of these adventures in our tribute to

Ian Thornton (New and Smithers 2004). However, his dedication and care

resulted in the Australian Museum now housing one of the all-time greatest

and most representative collections of this order.

Smither’s seminal work on psocids has been commemorated by his

colleagues by seven species named ‘smithersi’. One of these he himself

Australian Entomologist, 2012, 39 (4) 235

synonymised, a step that accorded him some amusement. Members of the

genus Sigmatoneura Enderlein (Psocidae) are characterised by extensive

sexual dimorphism, so that males and females can be associated clearly only

by co-incidence. Before this sexual difference was appreciated, I had

described a male and female from Nigeria (collected at the. same general

locality but a year apart) as separate taxa (New 1973). Courtenay later

decided that the more parsimonious approach was to consider them

conspecific; unfortunately, the second described was ‘smithersi’. In the same

paper, though, he also sunk Scaphopsocus Smithers as a new synonym of

Sigmatoneura! Two other psocid species are named ‘smithersorum’,

acknowledging Smila’s continued encouragement and support throughout

their long partnership. Courtenay himself named a number of psocids

‘alettae’ in appreciation of Smila’s notable contributions. Two patronymic

genera also exist: Smithersia Thornton (Myopsocidae) and Smithersiella

Badonnel (Caeciliusidae).

Neuroptera

The psocopteran patronyms are augmented by five patronymic lacewings,

marking another of his favourite insect orders. Courtenay’s interest in

Neuroptera was also a long one and, although he did not publish as

extensively on them, his knowledge was broad and exceptional. He described

a number of new Australian species, across four families, and wrote a number

of biological notes. Sadly, his long-projected revision of Australia’s

spongeflies, Sisyridae — a project we talked about at our first meeting, when I

was able to show him one of the three British species in the field — was not

completed, although he described several new species in isolation, including

Sisyra pedderensis Smithers, an endemic to Tasmania’s Lake Pedder

(Smithers eż al. 2008) and the last insect he described (Figs 4-6).

Other Orders

His interests in ‘the other orders’ also included Zoraptera (the chapter in

‘Insects of Australia’) and Mecoptera, for which he described a remarkable

endemic genus, 7ytthobittacus, from the Blue Mountains (Smithers 1973)

and produced a synopsis of Australian taxa (Smithers 1987). A full

bibliography of his publications is given by Britton (2012).

In an era in which specialisation is the norm, Courtenay Smithers’

entomological interests remained broad, but never forgoing depth and

authority. His commitment to advancing knowledge of Australian insects was

coupled with an equally strong commitment to their conservation. His legacy

is enduring and few could claim to have changed perspective of any insect

order to the extent that his studies on psocids, in particular, have done. His

responses to queries were always generous and informative and the

encouragement he gave to me, and to other less experienced colleagues, will

be remembered with gratitude and affection.

236 Australian Entomologist, 2012, 39 (4)

Figs 4-6. Adult, host and habitat of the spongefly, Sisyra pedderensis Smithers from

Lake Pedder, Tasmania: (4) adult female (Photo: G.N. Forteath); (5) encrusting

freshwater sponge, Radiospongilla pedderensis Osborn, Forteath and Stanisic, larval

food of Sisyra pedderensis (Photo: A.W. Osborn); (6) Lake Pedder, Tasmania, type

locality of both species (Photo: A.W. Osborn).

Acknowledgements

I am very grateful to Dr David Britton and Professors Nigel Forteath and

Andrew Osborn for use of their photographs.

References

BRITTON, D.R. 2012. Dr Courtenay Neville Smithers 1925-2011. A tribute. General and

Applied Entomology 40: 25-34.

LIENHARD, C. and SMITHERS, C.N. 2002. Psocoptera. World catalogue and bibliography.

Instrumenta Biodiversitatis V. Muséum d’histoire naturelle, Genéve; 745 pp.

NEW, T.R. 1973. A collection of Psocoptera from Nigeria. Occasional Papers of the

Entomological Society of Nigeria, No 10: 22 pp.

Australian Entomologist, 2012, 39 (4) 237

NEW, T.R. and SMITHERS, C.N. 2004. Ian W.B. Thornton - notes on his life and work, with

list of publications. Pp 9-30, in: Garcia Aldrete, A.N., Lienhard, C. and Mockford, E.L. (eds),

Thorntoniana, a commemorative volume for Ian W.B. Thornton (1926-2002). Publicaciones

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SMITHERS, A. 2012. Courtenay Smithers — before Australia. Australian Entomologist 39(4):

211-220.

SMITHERS, C.N. 1962. Insects of Macquarie Island: Psocoptera: Philotarsidae. Pacific Insects

4: 929-932.

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africaines 70: 209-294.

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137-209.

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145.

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259-344.

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Memoirs 14: 1-349.

SMITHERS, C.N. 1973. New species and records of Australian Bittacidae (Mecoptera). Journal

of the Australian Entomological Society 12: 296-300.

SMITHERS, C.N. 1984. The Psocoptera (Insecta) of South Australia. Records of the South

Australian Museum 18: 453-491.

SMITHERS, C.N. 1995. Psilopsocus mimulus Smithers (Psocoptera: Psilopsocidae), the first

known wood-boring Psocoptera. Journal of the Australian Entomological Society 34: 117-120.

SMITHERS, C.N. 1987. Synopsis of Australian scorpion flies (Mecoptera). General and Applied

Entomology 19: 31-44.

SMITHERS, C.N. 1990. Keys to the families and genera of Psocoptera (Arthropoda: Insecta).

Technical Reports of the Australian Museum 2: 1-82.

SMITHERS, C.N. 1996. Psocoptera. Pp 1-79, 333-335, 363-372, in: Wells, A. (ed.), Zoological

catalogue of Australia. Vol. 26. Psocoptera, Phthiraptera, Thysanoptera. CSIRO Publishing,

Melbourne.

SMITHERS, C.N. and LIENHARD, C. 1992. A revised bibliography of the Psocoptera

(Arthropoda: Insecta). Technical Reports of the Australian Museum 6: 1-86.

SMITHERS, C.N. and THORNTON, I.W.B. 1974. The Psocoptera (Insecta) of Norfolk Island.

Records of the Australian Museum 29: 209-234.

SMITHERS, C.N. and THORNTON, I.W.B. 1975. The Psocoptera (Insecta) of Lord Howe

Island. Records of the Australian Museum 29: 453-471.

SMITHERS, C.N., FORTEATH, G.N. and OSBORN, A. 2008. A new species of Sisyra

Burmeister (Insecta: Neuroptera: Sisyridae) from Lake Pedder, Tasmania. Australian Journal of

Entomology 47: 77-79. f

YOSHIZAWA, K. and JOHNSON, K.P. 2006. Morphology of male genitalia in lice and their

relatives and phylogenetic implications. Systematic Entomology 31: 350-361.

238 Australian Entomologist, 2012, 39 (4)

LIST OF PATRONYMIC TAXA DEDICATED TO

COURTENAY SMITHERS

Compiled by David R. Britton

Two genera and 31 species of insects and arachnids are dedicated to

Courtenay Smithers as follows (all Insecta unless otherwise stated):

Genera

Smithersia Thornton & Browning, 1992 (Psocoptera: Myopsocidae)

Smithersiella Badonnel, 1977 (Psocoptera: Caeciliusidae)

Species

Allocaecilius smithersi Lee & New, 1992 (Psocoptera: Pseudocaeciliidae)

Anthicus (Aulacoderus) smithersi von Hille, 1984 (Coleoptera: Anthicidae)

Bagauda smithersi Wygodzinsky, 1966 (Hemiptera: Reduviidae)

Bittacus smithersi Londt, 1972 (Mecoptera: Bittacidae)

Capillopsocus smithersorum Mockford, 1978 (Psocoptera: Amphipsocidae)

Clematoscenea smithersi Endang & New, 2010 (Psocoptera: Psocidae)

Clinopsocus smithersi New, 1972 (Psocoptera: Elipsocidae)

Coniocompsa smithersi Meinander, 1972 (Neuroptera: Coniopterygidae)

Delias mysis smithersi Daniels, 2012 (Lepidoptera: Pieridae)

Drosophila (Sophophora) smithersi Bock, 1976 (Diptera: Drosophilidae)

Epicaecilius smithersorum Mockford, 1999 (Psocoptera: Caeciliusidae)

Heteroconis smithersi Meinander, 1969 (Neuroptera: Coniopterygidae)

Javapsocus smithersi Endang, Thornton & New, 2002 (Psocoptera: Psocidae)

Lachesilla smithersi Garcia Aldrete, 1990) (Psocoptera: Lachesillidae)

Leptoperla smithersi Theischinger, 1981 (Plecoptera: Gripopterygidae)

Limnophila (Elaeophila) smithersi Alexander, 1958 (Diptera: Limoniidae)

Mesopsocus smithersi Badonnel, 1982 (Psocoptera: Mesopsocidae)

Molophilus (Austromolophilus) smithersi Theischinger, 2000 (Diptera: Limoniidae)

Nanolpium smithersi Beier, 1964 (PSEUDOSCORPIONES: Olpiidae)

Phyllodes imperialis smithersi Sands, 2012 (Lepidoptera: Erebidae)

Plexitartessus smithersi F. Evans, 1981 (Hemiptera: Cicadellidae)

Polyrhachis smithersi Kohout, 2012 (Hymenoptera: Formicidae)

Rhachiberotha smithersi Tjeder, 1959 (Neuroptera: Berothidae)

Scaphopsocus smithersi New, 1973 (Psocoptera: Psocidae)

Sympherobius smithersi Tjeder, 1961 (Neuroptera: Berothidae)

Tarlina smithersi Gray, 1987 (ARACHNIDA: Araneae: Gradungulidae)

Teratomyza smithersi McAlpine, 2012 (Diptera: Teratomyzidae)

Thaumastopsaltria smithersi Moulds, 2012 (Hemiptera: Cicadidae)

Theristria smithersi Lambkin, 1986 (Neuroptera: Mantispidae)

Tridactylus smithersi Günther, 1978 (Orthoptera: Tridactylidae)

Ziridava smithersi Holloway, 1977 (Lepidoptera: Geometridae)

Australian Entomologist, 2012, 39 (4): 239-246 239

A TRIBUTE TO COURTENAY SMITHERS AND HIS BUTTERFLY

MIGRATION STUDIES

JOHN V. PETERS

245 Quarry Road, Ryde, NSW 2112

Abstract

The contribution of Courtenay Smithers to Australian butterfly migration studies is discussed

and a few noteworthy migrations not previously published are recorded.

Introduction

I first met Courtenay Smithers in 1961. I had recently arrived in Australia

from the United Kingdom and Courtenay had arrived from Rhodesia less than

twelve months previously. We shared an interest in British birds and both of

us had prior experience with butterfly migration. Courtenay had witnessed

the mass migrations of the African Migrant, Belenois aurota (Fabricius,

1793) and I had reported on records of migrant butterflies in Ireland.

In the 19th century, British lepidopterists found it difficult to accept that

butterflies migrated. They could not ‘account for an almost complete absence

one year followed by a superabundance the next. They somehow believed that

butterflies successfully ‘hibernated’ to re-emerge the following spring and

that, being very mobile, their presence in large numbers was the result of

movement from elsewhere in the kingdom’ (Salmon 2000).

Williams (1930) documented early studies on butterfly migration. Common

and Waterhouse (1972, 1981) devoted almost four pages to it, with special

reference to the work of the entomologists at the Australian Museum, in

particular the marking of the wings of the Monarch followed by release and,

hopefully, the eventual recapture of some of the specimens. Details were

given of the need for recording butterfly migrations: i.e. species name (and

specimens), locality, compass direction, time of day, date and the number of

individuals passing over a measured distance in a given time.

A migration is noted when ‘large numbers of butterflies may be seen flying in

one direction for hours on end or for several days’ (Common and Waterhouse

1972). These butterflies fly with a purpose and are rarely distracted from it;

they fly over buildings or trees rather than fly around them. On the other

hand, there are ‘some species of butterflies that may, in favourable seasons,

extend their distribution southward by hundreds of kilometres’ (Common and

Waterhouse 1972). Sydney is an ideal place to record these range extensions,

as many of Courtenay’s record show. Temporary range extension or full-on

migration must be important to the biology of each species involved and is a

part of butterfly biology that certainly needs further investigation.

Shortly after arriving in Australia, Courtenay started to bring together all the

records of migrating butterflies that he could find, along with new and current

records from members of the Entomology Section of the Royal Zoological

240 Australian Entomologist, 2012, 39 (4)

Society and from the many correspondents who were part of his Monarch

(Danaus plexippus (Linnaeus, 1758)) labelling project (Fig. 1). Much of this

information was published from 1961 to 1995 in the 28 papers he wrote

dealing with butterfly migration. He first published on insect migration in

Australia as early as 1961, followed by a paper on butterfly migration in

1963. His series of papers entitled ‘Migration Records in Australia, parts 1-5’

(Smithers 1970b, 1978, 1983a, 1983b, 1985) brought together most of the

previous records of Australian butterfly migration, plus those involving the

Odonata, Homoptera, Coleoptera, Diptera and Hymenoptera.

a j ai TF mA nN

Fig. 1. Volunteers Justine O’Regan and Alana Thomas helping Courtenay Smithers

tag and record Monarch butterflies in the 1960s.

Danaus plexippus, the Monarch butterfly

On 8 August 1962, Courtenay and I commenced the first marking of

Monarch butterflies (Danaus plexippus) in Australia, by using tiny

handwritten labels glued to the underside of a hind wing using Mendene

thinned with ethyl acetate (Fig. 2), with some successfully recaptured (Fig.

3). In 1963, the programme commenced in earnest, using small self-adhesive

labels produced by Avery Scales; however, this still required the numbers to

be handwritten. The cell of a forewing had the scales removed by gently

rubbing between finger and thumb and the label then folded and attached to

the wing (Fig. 4) — this meant that there was no need to use glue (a rather

messy procedure) and the label was more easily seen. This method had been

used earlier by Canadian researchers. Courtenay, through the Australian

Australian Entomologist, 2012, 39 (4) 241

Museum, gathered together a great many collaborators from all over

Australia, but mainly from NSW, and many thousands of Monarchs and other

species were labelled, with the results of this study published by Smithers

(1972, 1977).

Fig. 2. Monarch number 995 marked with a trial label glued to hindwing underside.

A landmark in Australian Monarch studies was when the first overwintering

aggregation sites were recorded by Smithers (1965) south-west of Sydney

(Fig. 5). I have a note recording that, on 23 April 1967, at the Razorback

(near Camden, NSW) overwintering site, a small group of us labelled 1,400

Monarchs in one day from the very large clusters that had assembled that

winter. I’m not sure how many butterflies were labelled during the entire

length of the programme but I still have a few unused labels and the numbers

on them are in excess of 69,000.

242 Australian Entomologist, 2012, 39 (4)

The Canadian Entomologist Frederick Urquhart pioneered the labelling of

Danaus plexippus and the results of his studies into the movements of the

Monarch in North America were published in his book (Urquhart 1960).

Courtenay was able to discuss his own studies on the Monarch with Urquhart

during a visit to Toronto, Canada in 1986. His Australian studies were

included in a review of Monarch migration in Australia by James (1993).

Fig. 5. Courtenay Smithers collecting Monarch butterflies at an aggregation in 1968.

Australian Entomologist, 2012, 39 (4) 243

Belenois java teutonia, the Caper White

A species conspicuously absent from Courtenay’s reviews was the Caper

White, Belenois java teutonia (Fabricius, 1775). He had so much material on

the migrations of this butterfly, much of it from his many correspondents as

well as his own records, that he often expressed frustration that the job of

making any sense of it seemed insurmountable, particularly as he was so

involved in his museum work as well as his research interests in, particularly,

the Psocoptera. As a result, Courtenay did not publish any records of the

migrations of the Caper White. However, he and I discussed the migrations

of this species at length and we concluded that the main breeding area of this

species is that vast area of southern Queensland and New South Wales west

of the Great Dividing Range, where larval food plants such as the Warrior

Bush Apophyllum anomalum F. Muell and the Native Orange Capparis

mitchellii Lindl (both Capparaceae) grow.

The native food plants occur naturally only north of Griffith, NSW, which

poses one of the key questions as to why the migrations seem to be so

maladaptive, given that the butterflies often fly in such huge numbers away

from potential breeding localities such as, for example, into all of Victoria.

The butterflies emerge in large numbers in late October-November, with

freshly emerged females soon mated and ready to lay eggs on any suitable

food plant they come across. They normally fly in a northeasterly direction

but their exact route varies annually, depending on the prevailing winds at the

time. A strong westerly can cause them to fly through Canberra and as far

eastwards as Sydney and Brisbane and beyond. During years when the

westerlies fail to coincide with their migrations, very few Caper Whites may

be seen in Sydney.

On 28 November 1969, Courtenay recorded ‘a tremendous “invasion” of

Caper Whites on the western side of Lord Howe Island (Smithers 1970a).

Reports from Australia indicated that a large-scale migration was taking

place in eastern Australia at that time, with the butterflies moving in a

northwesterly direction. These were the first Caper White butterflies recorded

from Lord Howe Island, suggesting that with appropriate conditions this

species is easily blown off course. Specimens were seen on Lord Howe

Island for only a few days and the drop in numbers by 30 November suggests

that they continued their migratory movement, leaving the island and

presumably becoming ‘lost at sea’. Similarly, on 24/25 November 1987, I

noted large numbers (‘a cloud’) of Caper Whites coming straight in to the

shore off the sea at Surfers Paradise, Queensland, flying just above wave

height. They continued up the beach and across the main road. Very few were

seen on the 26th and only one on the 27th. Common and Waterhouse (1981)

wrote of the Caper White butterfly: ‘Immense migratory flights of this

species frequently occur in southern Queensland and New South Wales, and

sometimes extending into Victoria. ... At Canberra it is not uncommon to see

this species flying steadily north to north-east during November’.

244 Australian Entomologist, 2012, 39 (4)

Discussion and new observations

The general public tend not to notice migrating butterflies unless the

movement is spectacular, such as the migrations of Caper Whites. Large

numbers of white butterflies moving in one direction for days on end are

difficult to ignore. My first encounter with this phenomenon was in 1963 at

the Sydney suburb of Ryde. The migration started as a trickle on 23

November with one butterfly seen at 10.00 h, nine from 11.30-12.30 h and,

from 12.45 to 13.45 h, 196 butterflies were recorded over a 15 metre front

flying in a NNE direction. This is the equivalent of over 13,000 per hour over

a 1 km front. This migration quickly petered out, with one Caper White

sighted on 24 November and another on the 29th. A much more spectacular

migration of Caper Whites occurred during November 1966. I observed this

at both Ryde and Chatswood (another Sydney suburb). All the butterflies

were flying in a northerly direction, first noticed on 1 November and

extending until 20 December. At its peak on 28 November, I recorded 180+

butterflies over a 25 metre front for just 5 minutes, an estimate of 86,400+

butterflies per km front per hour. It is not unusual for a migration to continue

from dawn until dusk suggesting that, in this example, as many as 7 million+

Caper Whites passed through Sydney in one day. Migrations such as this

have been observed from Sydney west to the Blue Mountains, a distance of

approximately 100 km!

An example of the general public failing to notice migrating butterflies

occurred during November 1986, while I and a number of teachers were

supervising children at Ryde swimming pool. For some time I watched

Painted Lady butterflies, Vanessa kershawi (McCoy, 1868), flying south

across the pool. After a while I asked the other teachers if they had noticed

the migrating butterflies. No, they hadn’t and they were astonished when they

realised what was happening around them and how unobservant they were.

Since the last of Courtenay’s papers on butterfly migration was published

(Smithers 1995), I have observed quite a number of migrations, particularly

at my daughter’s property at Tuchekoi, near Cooroy on the Sunshine Coast,

Queensland. There I have recorded 10 species migrating, one not previously

recorded and, on occasions, three species migrating in different directions at

the same time. On 22 October 2002, from 11.08 to 11.23 h, over a 33 metre

front, I observed 23 Belenois java teutonia flying in a NW direction, three

Catopsilia pomona (Fabricius, 1775) flying southeasterly and nine Elodina

angulipennis (P.H. Lucas, 1852) flying in a SE direction at a height of 1-3

metres. (Two voucher specimens of Æ. angulipennis were collected). Two

days previously, at the same locality, there were 54 B. j. teutonia flying to the

NW, seven C. pomona flying to the SE and three Junonia villida (Fabricius,

1787) flying to the west.

Also at Tuchekoi, on 25 and 27 December 1993 I observed three species of

butterfly in a spectacular southerly migration. Blue Tigers, Tirumala hamata

Australian Entomologist, 2012, 39 (4) 245

(W.S. Macleay, 1826), Lemon Migrants, C. pomona and Pale green

Triangles, Graphium eurypylus (Linnaeus, 1758) were observed flying over a

50 metre front during four 15 minute sessions over 3 days (Table 1). The

migration continued all day on 26 December until a quite. violent storm

arrived at 1500 h, with the migration continuing on the 27th. The Blue Tigers

flew in a leisurely but determined flight, while the Lemon Migrants and the

Pale green Triangles flew at a much faster pace. It is interesting to note that

‘resident’ Blue Tigers were not involved in the migration.

Table.1. Southerly migration of three butterfly species at Tuchekoi, near Cooroy, SE

Queensland, on 25 and 27 December 1993. Numbers were recorded flying over a 50

metre front.

Butterfly species 25 Dec. 27 Dec. Height

1030- nas 1530-—1100- Bove,

1045h 1230h 1545h — 1li5h FOUN

Blue Tiger 22 29 11 25 0.5-2 m

Tirumala hamata

Lemon Migrant 9 9 - 21 2-3 m

Catopsilia pomona

Pale green Triangle 1] 3 9 11 1-3 m

Graphium eurypylus

Temperature 29:0 B2 30°C 29°C

Many of the butterfly species that migrate also have a return migration but

usually in much smaller numbers. The major flights of B. java teutonia in

October-November are often spectacular, with reports reaching major Sydney

newspapers; however, the return flights the following September are

numerically very small and may only be observed by alert lepidopterists.

Although Ian Common worked extensively on bogong moth migration,

Courtenay was the only person to have recorded butterfly migration in

Australia seriously during the past 50 years. He explored what is perhaps the

most interesting and least understood aspect of butterfly biology. We owe

him a debt of gratitude for his work on Australia’s migratory butterflies and

other insects and, in particular, his studies of the Monarch, Danaus plexippus.

Apart from a posthumous paper in this memorial issue (Smithers 2012), it is

almost 20 years since his last publication on butterfly migration; thus there is

an opportunity for someone to continue to record and publish details of this

fascinating phenomenon. Courtenay’s paper records of Caper White

migrations are maintained in the Australian Museum archives and are

available to researchers.

246 Australian Entomologist, 2012, 39 (4)

References

COMMON, I.F.B. and WATERHOUSE, D.F. 1972. Butterflies of Australia. Angus ang

Robertson, Melbourne; [xii] + 498 pp, 41 pls.

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised edition,

Angus and Robertson, Melbourne; xiv + 682 pp.

JAMES, D.G. 1993. Migration biology of the monarch butterfly in Australia. Pp 189-200, in:

Malcolm, S.B. and Zalucki, M.P. (eds.), Biology and conservation of the Monarch butterfly.

Natural History Museum of Los Angeles County, Los Angeles; xii + 419 pp.

SALMON, M.A. 2000. The Aurelian legacy. University of California Press. Berkeley and Los

Angeles, California; 432 pp.

SMITHERS, C.N. 1961. Insect migration. Australian Museum Magazine 13: 350-353.

SMITHERS, C.N. 1963. Butterfly migration. Ausralian Natural History 14: 215-217.

SMITHERS, C. N. 1965. A note on overwintering in Danaus plexippus (Linnaeus) (Lepidoptera;

Nymphalidae) in Australia. Australian Zoologist 13(2): 135-136.

SMITHERS, C.N. 1970a. Observations on Lord Howe Island butterflies. Australian Zoologist

15: 377-379.

SMITHERS, C.N. 1970b. Migration records in Australia. 1. Odonata, Homoptera, Coleoptera,

Diptera and Hymenoptera. Australian Zoologist 15: 380-382.

SMITHERS, C.N. 1972. Observations on a breeding population of Danaus plexippus (L.)

(Lepidoptera: Nymphalidae) at Camden, New South Wales. Autsralian Zoologist 17: 12-148.

SMITHERS, C.N. 1977. Seasonal distribution and breeding status of Danaus plexippus (L.)

(Lepidoptera: Nymphalidae) in Australia. Journal of the Australian Entomological Society 16:

175-184.

SMITHERS, C.N. 1978. Migration records in Australia. 2. Hesperiidae and Papilionidae

(Lepidoptera). Australian Entomological Magazine 5: 11-14.

SMITHERS, C.N. 1983a. Migration records in Australia, 3. Danainae and Acraeinae

(Lepidoptera: Nymphalidae). Australian Entomological Magazine 10: 21-27.

SMITHERS, C.N. 1983b. Migration records in Australia. 4, Pieridae (Lepidoptera) other than

Anaphaeis java teutonia (F.) Australian Entomological Magazine 10: 47-54.

SMITHERS, C.N. 1985. Migration records in Australia. 5. Lycaenidae and Nymphalinae

(Lepidoptera). Australian Entomological Magazine 11: 91-97.

SMITHERS, C.N. 1995. The first record of Tirumala hamata hamata (W.S. Macleay)

(Lepidoptera: Nymphalidae), the blue tiger butterfly on Norfolk Island. Norfolk Nature Notes 11:

395-396.

SMITHERS, C.N. 2012. Thermals and butterfly (Lepidoptera) migrations from Australia to New

Zealand. Australian Entomologist 39(4): 247-252.

URQUHART, F. 1960, The Monarch butterfly. University of Toronto Press, Toronto; xxiv + 361

pp.

WILLIAMS, C.B. 1930. The migration of butterflies. Oliver and Boyd, Edinburgh & London; xi

+ 473 pp.

Australian Entomologist, 2012, 39 (4): 247-252 247

THERMALS AND BUTTERFLY (LEPIDOPTERA) MIGRATIONS

FROM AUSTRALIA TO NEW ZEALAND

C.N. SMITHERS

Deceased; formerly at Australian Museum, 6 College Street, Sydney, NSW 2010

Abstract

Two direct observations of butterflies at high altitudes in thermals are noted, supporting the

hypothesis that long distance migration of butterflies between Australia and New Zealand takes

place at high altitudes.

Introduction

The arrival of Australian insects in New Zealand from across the Tasman Sea

is a well known phenomenon. Perusal of papers such as those of Ramsay

(1954, 1971), Ramsay and Ordish (1966), Gibbs (1969), Fox (1978) and

Early et al. (1995), plus references provided by these authors, give details and

discussion of many such arrivals, especially of butterflies. Holloway (1977,

1996) discussed in great detail similar arrivals on Norfolk Island (especially

of moths), based on a remarkable long-term light-trapping program carried

out by Marge and Fred Jowett. Observations have been recorded of the

arrival of the butterflies Vanessa kershawi (McCoy) on Norfolk Island

(Smithers 1969), Belenois java teutonia (Fabricius) on Lord Howe Island

(Smithers 1970) and Tirumala hamata (W.S. Macleay) on Norfolk Island

(Smithers 1995). Tomlinson (1973) discussed meteorological aspects of

dispersal of insects across the Tasman Sea and Holloway (1977) discussed

possible mechanisms of transport of insects to Norfolk Island. Further afield,

recent work on Danaus plexippus (Linnaeus) in North America (Calvert

2001) and Vanessa cardui (Linnaeus) migrating from North Africa to Europe

(Stefanescu et al. 2007), has convincingly demonstrated the use of thermals

and high altitude synoptic scale winds to assist migration.

Opinions on long-distance butterfly migration over water

There has been some uncertainty, if not reluctance, as to acceptance of the

possibility of butterflies undertaking long-distance flights over water from

Australia to New Zealand in low temperature, fast-moving, high altitude air

streams. There appears to have been greater acceptance that the journey could

be made at low levels with wind assistance. Wind assistance is regarded as

essential by most authors, because ‘refuelling’ by nectar intake is not possible

over the sea and because of the physiological inability of the insects to fly for

the time needed to make the crossing without such help. Johnson (1969)

discussed the physiological needs of insects undertaking long-distance flights

and suggested that rapid high altitude flight was involved for some of them.

Ramsay (1954) pointed out that, during ten days before the recorded arrival

of specimens of Junonia villida (Fabricius) in New Zealand, ‘there were at

least three occasions on which these insects could have been carried across

the Tasman Sea’, implying that they were blown from Australia. The time

taken for such a journey was given as ‘about 3 days’. Ramsay and Ordish

248 Australian Entomologist, 2012, 39 (4)

(1966) discussed weather conditions in relation to an extensive influx of

Hypolimnas bolina nerina (Fabricius) in 1956, in terms which suggest that

the major factor enabling butterflies to make the journey was the wind. They

also estimated that ‘The time of transit for a non-active butterfly being

carried [my italics in both cases] from the Australian coast near Sydney to

New Zealand would have varied between one and four days ...’ and noted

that ‘Many of the butterflies arrived in remarkably good condition and

therefore must have had a relatively easy passage.’ We find also a suggestion

that ‘At 10,000 ft, where the wind direction was much more constant, the

speed varied between 20 and 50 knots, but mostly remained at near 30 knots.

Strong WNW winds occurred near ground level in the Sydney-Newcastle

area from 15-18 April and may have carried the butterflies up and out into the

trans-Tasman airflow.’ The interesting suggestion that winds at higher

altitudes might be involved is not followed up, probably because of lack of

direct evidence or disbelief that they could survive for long enough at the low

temperatures and high altitudes. ‘After 20 April the winds became southerly

... Thus, there was less chance of the butterflies being carried [my italics] to

New Zealand in this manner after 20 April, the period when most butterflies

were seen.’ Gibbs (1969) recorded an immigration of large numbers of

Vanessa kershawi in New Zealand in October/November 1968, at the same

time as this species was arriving on Norfolk Island (Smithers 1969). Gibbs

suggested, on the basis of smoke arriving from bushfires near Sydney, that

‘winds at the time of the main 1968 invasions were favourable for the

movement of butterflies’.

Smithers (1969), when recording the arrival of what was obviously part of the

same migrant population of V. kershawi on Norfolk Island, also with smoke

from Australian fires, pointed out that the ‘air conditions in many parts of

eastern Australia were such that strong updraughts from extensive bushfires

were common. Over wide areas, therefore, conditions were favourable for the

rapid lifting of specimens in flight to heights of 15,000 feet and more. At

these heights, winds were easily capable of transporting specimens to Norfolk

Island and the passive movement of torpid butterflies could have been quite

rapid; certainly a short enough period of time would be involved to permit

butterflies to descend unharmed.’ It is a frequently observed fact that partly

burned leaves and other debris are lifted high during a bushfire and carried a

long distance before returning to earth. Clearly, a butterfly raised to high

altitudes, where temperatures low enough to induce torpor are encountered,

could be carried at the high speeds of high altitude winds for considerable

distances before descending to warmer air. The fact that a butterfly was not

actively flying would not, alone, necessarily cause its descent.

Tomlinson (1973) discussed five means by which a butterfly might be raised

to high altitude. These included convective cloud, thermals, initial lift

provided by passage of a cold front, a small ‘dust devil’ equivalent to a small

‘tornado’ and the processes of turbulent transfer. When considering the April

Australian Entomologist, 2012, 39 (4) 249

1971 Hypolimnas bolina immigration into New Zealand, he estimated that

temperatures at 1000 metres were from 7-10°C and at 3000 metres they were

from -3-0°C. Even if conditions are not cold enough to induce torpor, the

wind speeds at high altitudes could carry a flying or gliding butterfly from

Australia to New Zealand in a matter of hours providing the specimen

remained airborne, something which would not require much action on the

part of the insect. It must be remembered that no damage is incurred by an

insect which is simply supported by a moving body of air, irrespective of the

speed at which the air mass is moving. Nine hours of flying time in a day is

not exceptional for a migrant over land in eastern Australia.

Observations of arrivals in New Zealand, Lord Howe Island or Norfolk

Island have, of course, all been made at ground level. The assertion is

sometimes made that, as arrivals have only been recorded near ground level,

high level arrival does not take place. This is an assumption which is difficult

to accept when little opportunity has been available to make observations at

high levels. A few authors (e.g. Johnson 1969, Smithers 1969, Tomlinson

1973, Holloway 1977) accepted that wind assisted high level travel could be

involved in over sea migrations. Such assistance could allow long distances

to be covered very quickly, considering that the winds at high altitudes are

often very strong, persistent and maintain their direction for long periods. On

the other hand, caution in accepting this has been justifiable in view of the

lack of direct, positive, observational evidence.

Two instances of direct evidence that butterflies can rise to high altitudes and

thereby have the opportunity to enter high level air streams are available.

Both have involved ‘thermals’ as the lifting force. The first, from an

unexpected source not likely to be noticed by entomologists (Welch and

Welch 1965), originates in an observation made in August 1931 in Germany

by W. Hirth, a German pioneer glider pilot. The first to make serious use of

thermals at a time when the art of ‘soaring’ was in its infancy and gliders

were slow-moving, Hirth (1938) wrote: ‘I was unable to find any new up-

currents in my immediate vicinity. However, I suddenly caught sight of some

butterflies which had obviously reached that height under thermal influences

and I hastened to make use of the same locality.” The observation was made

at a height of 2000 ft.

The second observation was made by G. Sutherland (pers. comm.) while

paragliding. Paragliders rely on thermals to obtain lift. Thermals frequently

carry up debris such as leaves, grass, seeds and other parts of plants. These

items are used by the flyers to detect thermals which, unless associated with

cloud formations or other visible signs, cannot be detected. Butterflies were

encountered (probably B. java teutonia) above 6000 ft; they were circling or

flying ‘aimlessly’ in a thermal at Manilla, New South Wales, at the end of

November 1997 and it is probable that most of them would have reached

altitudes of 10,000 ft if they remained in the thermal. It is significant that low

250 Australian Entomologist, 2012, 39 (4)

level migrations of B. java teutonia were recorded at several New South

Wales localities before and at the same time as the observations in the

thermals. These were at Ryde on 1.x.1997 (J.V. Peters), Turramurra on

7.xi.1997 (C.N. Smithers), Wallis Lake on 11-12.xi.1997 (A.B. Rose), Forster

on 15.xi.1997 (A.B. Rose) and between Sydney and Goulburn on 27-

28.x1.1997 (A.S. and C.N. Smithers). The last-mentioned was a continuous,

sometimes dense migration observed over a distance of 100 km. Although

there are no direct records of migration at ground level at Manilla itself at the

time of the observation in the thermal, it is obvious from the localities and

dates of the recorded migrations that there would almost certainly have been

movements in the Manilla district, or nearby, at the time.

Rising with the aid of a thermal is one way which would provide access to a

very strong and persistent high altitude wind system. Whether butterflies

actively fly upwards once within a thermal or are passively lifted within the

thermal is of little importance in the present context. In the case of winds

travelling from Australia over the Pacific, butterflies could be moved

consistently at speeds well in excess of 100 km/hour above 10,000 ft (c.f.

Holloway 1977, p. 158, fig. 111 and associated discussion) and greater

speeds at higher altitudes. This would enable a butterfly to complete the

journey from Australia to Norfolk Island in a matter of hours, depending on

the wind speed. It is relevant to note that all species of butterflies which have

been recorded as arrivals in New Zealand and Norfolk Island are habitual

migrants in Australia. They all cease flight at night and take nectar during

their migrations over land. This is not possible when flying over sea but

would not be necessary if their journey were fast enough and economical

enough in energy consumption.

There are, of course, many records of migrating butterflies being seen at sea

at low altitudes. Tirumala hamata was seen on 7 April 1995, two days before

appearing on both Norfolk Island and New Zealand on 9 April 1995

(Smithers 1995, Early et al. 1995). This observation does not preclude the

possibility of successful high altitude travel at the same time by parts of the

same migrating population. It would depend on the conditions in the area of

origin in Australia, which could be very extensive and variable. There is,

also, no reason to assume that only one method is ‘used’ by the butterflies. It

does seem, however, with the physiological constraints discussed by Johnson

(1969), that fast-moving, long-distance, high level migrating individuals have

an advantage and a better chance of successful long distance travel than low-

level travellers moving ‘under their own steam’ across areas where

‘refuelling’ is not possible. There are many records of individual butterflies

and other insects, some of which are small and possessed of very limited

energy reserves, having been seen hundreds of miles from land. The

turbulence caused by wind changing directions (as in the case of moths

arriving on Norfolk Island (Holloway 1977), mentioned below) could well

bring down high altitude butterflies before landfall, with the butterflies

Australian Entomologist, 2012, 39 (4) 251

making only the final part of the journey at lower altitudes. This would be

more likely in the case of New Zealand, where the land mass is much greater

than that of Norfolk Island. Whether one or two ‘methods’ are available for

successful long-distance movement will only become known through further

observation. In any event, it must be accepted that there will probably be

great losses of individuals on any long distance migration low over the sea

through physiological failure. Movement within a high level air mass would

not require exceptional energy expenditure; as long as the insect remains

airborne it could be transported at high speed. A torpid butterfly could be

carried for long distances, just as inanimate objects raised by bushfires are

carried. Higher level movement is likely to be more economical than low

level flight in terms of energy expenditure.

Holloway’s (1977, p. 152) data indicated that for Norfolk Island ‘Apart from

October 1971, when conditions may have been exceptional, rarely does a

peak of vagrant arrivals coincide with a period of strong and predominant

westerlies.’ Such peaks occurred when wind was ‘changing from westerly to

south-easterly or vice versa’ and the moths themselves were taken more

frequently in the traps on the eastern side of the island than the west. This

suggests an approach from the east into sheltered areas of the island against

the predominant westerly wind. The presence of the island near such wind

conditions would ‘create conditions favouring downfall on Norfolk as distinct

from vagrant transport over Norfolk’. As the moths are Australian species

they must, however, have originated in Australia. This apparent anomaly can

be explained if the moths are using a high level route provided by the almost

constant westerly winds which blow at high altitudes, irrespective of the low

level wind directions at the time, and are brought down by the turbulence

associated with wind direction change at low levels. At low level the winds

were easterly and hence the moths were being assisted by easterly winds for

the last part of the journey and approached the island from the east. Descent

initiated by a zone of turbulence, whether cause by wind change or by

presence of a land mass, need not be injurious to the insect.

It may require a different set of circumstances to occur for nocturnal insects,

e.g. moths, to be lifted initially by thermals, which are essentially diurnal

phenomena. Nocturnal migrants could well gain the required initial lift by

some other mechanism, such as the ‘cold front’ method described by

Tomlinson (1973). In the past, the hypothesis that butterflies (and other

insects) can reach the heights necessary to enable them to be transported

rapidly from Australia to New Zealand, Lord Howe and Norfolk Islands

(Smithers 1969, Holloway 1977) has rested on somewhat circumstantial

evidence. The two observations reported here lend strong, direct

observational support to that hypothesis.

Acknowledgements

I would like to thank G. Sutherland for his observations at Manilla,

252 Australian Entomologist, 2012, 39 (4)

John Peters and Tony Rose for their continued help in providing insect

migration records and Gerald Hyam for drawing attention to and providing a

copy of the publication by Welch and Welch.

References

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flight behavior and direction in relation to celestial and physiographic cues. Journal of the

Lepidopterists’ Society 55(4): 162-168.

EARLY, J.W., PARRISH, G.R. and RYAN, P.A. 1995. An invasion of Australian blue tiger

butterflies (Lepidoptera: Nymphalidae) in New Zealand. Records of the Auckland Institute and

Museum 32: 45-53.

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hypothesis on colonisation. New Zealand Entomologist 6(4): 368-380.

GIBBS, G.W. 1969. A large migration of the Australian painted lady butterfly, Vanessa

kershawi (McCoy), to New Zealand. New Zealand Entomologist 4(2): 14-21.

HIRTH, W. 1938. The art of soaring flight. Stuttgart. Translated by Naomi Heron-Maxwell.

(Quoted in Welch and Welch 1965, p. 86).

HOLLOWAY, J.D. 1977. The Lepidoptera of Norfolk Island, their biogeography and ecology.

Junk, The Hague; vi + 291 pp.

HOLLOWAY, J.D. 1996. The Lepidoptera of Norfolk Island, actual and potential, their origins

and dynamics. Pp 123-151, in: Keast, A. and Miller, S.E. (eds.), The origin and evolution of

Pacific Island biotas, New Guinea to Eastern Polynesia: patterns and processes. Academic

Publishing, Amsterdam.

JOHNSON, C.G. 1969. Migration and dispersal of insects by flight. Methuen, London; xvi +

763 pp.

RAMSAY, G.W. 1954. Records of the Australian butterfly Precis villida (Fabr.) from the

Wellington District. New Zealand Entomologist 1(4): 22-23.

RAMSAY, G.W. 1971. The blue moon butterfly Hypolimnas bolina nerina in New Zealand

during autumn, 1971. New Zealand Entomologist 5(1): 73-75.

RAMSAY, G.W. and ORDISH, R.G. 1966. The Australian blue moon butterfly Hypolimnas

bolina nerina (F.) in New Zealand. New Zealand Journal of Science 9(3): 719-729.

SMITHERS, C.N. 1969. A note on migrations of Vanessa kershawi (McCoy) (Lepidoptera:

Nymphalidae) in Australia, 1963-1968. Australian Zoologist 15(2): 188-194.

SMITHERS, C.N. 1970. Observations on Lord Howe Island butterflies. Australian Zoologist

15(3): 377-379.

SMITHERS, C.N. 1995. The first record of Tirumala hamata (W.S. Macleay) (Lepidoptera:

Nymphalidae), the blue tiger butterfly on Norfolk Island. Norfolk Nature Notes 11(3): 295-296.

STEFANESCU, C., ALARCON, M. and AVILA, A. 2007. Migration of the painted lady

butterfly, Vanessa cardui, to north-eastern Spain is aided by African wind currents. Journal of

Animal Ecology 76(5): 888-898.

TOMLINSON, A.I. 1973. Meteorological aspects of trans-Tasman insect dispersal. New Zealand

Entomologist 5(3/4): 253-268.

WELCH, A. and WELCH, L. 1965. The story of gliding. John Murray, London.

Postscript. The introduction was updated by Albert Orr — Ed.

Australian Entomologist, 2012, 39 (4): 253-260 253

REVISITING THE PSOCOPTERA (INSECTA) OF BARROW

ISLAND, WESTERN AUSTRALIA

N.R. GUNAWARDENE, C.K. TAYLOR and J.D. MAJER

Curtin Institute for Biodiversity and Climate, Department of Environment and Agriculture,

Curtin University, GPO Box U1987, Perth, WA 6845

(Email: n.gunawardene@curtin.edu.au)

Abstract

Courtenay Smithers recorded five species of Psocoptera on Barrow Island in 1982. Since then,

repeated surveys have been conducted as part of environmental impact assessments associated

with the Gorgon Project development on Barrow Island. This baseline information on

invertebrates is to be utilised as a component of surveillance programs in support of the Gorgon

Project’s quarantine detection system. These additional surveys on the island have yielded a

further 20 species of psocopterans, bringing the total fauna collected to 25 species. This includes

the first Australian record of the synanthropic species Dorypteryx domestica (Smithers).

Introduction

Courtenay Smithers originally recorded five species of Psocoptera collected

over 18 field days on Barrow Island during 1982 (Smithers 1984). Smithers

described a new genus and two new species from this collection event, one of

which was Barrowia insularis Smithers (Fig. 1). This psocid species has not

been recorded so far from any other locality in Australia. The initial

collection on Barrow Island was the start of the discovery of a surprising

diversity of Psocoptera on this arid island, which lies off the northwestern

coast of Australia and has become the site of Chevron Australia Pty Ltd’s

liquefied natural gas development, a part of the Gorgon Project (see

acknowledgements). As a result of the environmental impact assessments

associated with the development, extensive field surveys were conducted

with the aim of providing baseline information on invertebrates. This baseline

information is utilised as a component of a surveillance program in support of

the Gorgon Project’s Quarantine Management System. We present herewith a

list of species collected to date on Barrow Island and provide an analysis of

species distribution within the island. As data are lacking for the

corresponding mainland psocid fauna, it is difficult to place this island fauna

into a larger continental context. However, the existence of this body of work

will hopefully enable other researchers to gain an understanding of the

possible diversity of psocopterans in arid Australia.

This study forms part of a multi-million dollar biosecurity project which

provides essential data for protecting Australia’s biosecurity. More than 30

taxonomists were involved in identifying invertebrates from the baseline

invertebrate survey of Barrow Island (Callan ef al. 2011), including

Courtenay Smithers. It is interesting to reflect that six of the taxonomists are

retired scientists and four undertake taxonomic work only in their spare time.

The diminishing number of salaried taxonomists in this country is disturbing,

particularly considering the advanced age of many of those who assisted us,

as evidenced by the sad passing away of our friend, Courtenay Smithers.

254 Australian Entomologist, 2012, 39 (4)

Fig. 1. Lateral view of a male specimen of Barrowia insularis Smithers from Barrow

Island. This psocid species was collected by Courtenay Smithers and described in

Smithers (1984).

Materials and methods

Barrow Island is located approximately 60 km from the North-West coast of

Australia and is 234 km? in extent. The island receives, on average, 300 mm

of rainfall annually. However, this is highly variable from year to year, with

some years receiving less than 100 mm (e.g. 2009) and others receiving more

than 700 mm (e.g. 1973) (BOM 2012). The majority of this rain falls in the

hot, humid summer months of February and March, with a smaller amount

falling in the cooler months of July and August. The most dominant

vegetation complex on the island is Triodia hummock grasslands (Fig. 2).

These are interspersed with Acacia stands and small clumps of Ficus trees.

Mangroves (Avicennia and Rhizophora) occur in small patches along the

coast (RPS Bowman Bishaw Gorham and Mattiske Consulting 2005).

Intensive surveys of invertebrates were carried out on Barrow Island between

2006 and 2007 and the results of these surveys have been reported in Callan

et al. (2011), with species lists provided as an electronic appendix. This

report recorded 19 species of Psocoptera in ten families. A variety of methods

were used to capture invertebrates from the litter, ground surface, vegetation

and other structures in a range of habitats distributed across the island (Figs

3-5). An extensive description of these methods and sites can also be found in

Callan et al. (2011). All specimens from these surveys are lodged in the

Australian Museum, with voucher reference specimens held in Curtin

University Entomology Museum.

Australian Entomologist, 2012, 39 (4) 255

Figs 2-5. Environment and collecting techniques: (2) the east coast of Barrow Island

is typified by low, spinifex covered dunes and sandy beaches with tidal flats — in the

distance are five holding tanks for the oil that is pumped from below the island; (3)

one of the collecting methods for invertebrates is night hand collection — many night-

active flying insects are collected with this method; (4) another collection method

commonly used for Psocoptera — beating or sweeping the vegetation into a collection

tray or net often yields a number of invertebrate orders; (5) a leaf blower in suction

mode is used to vacuum small invertebrates inhabiting vegetation — as spinifex is

dense and spiky and unpleasant for hand collection, the blower vac is an effective

method for collecting psocopterans within these tussocks.

Psocoptera were collected from 12 undisturbed native vegetation sites

(classified as GP sites) in two seasons in 2006. The first collection was after

the hot wet season in March and the second was carried out after the cooler

dry season in September. Psocoptera were also collected from 13 disturbed

habitat sites (classified as NIS sites) in 2006, which were visited again in

2007. The latter sites were chosen to see whether any non-indigenous species

were present. A slightly modified collection protocol was carried out for the

disturbed habitats to take into account the built environment present at these

sites.

Australian Entomologist, 2012, 39 (4)

Table 1. The 25 Psocoptera species or morphospecies that have been collected from

Barrow Island to date, in both native vegetation (N), disturbed habitats (D) and the

built environment (B). Asterisked species were recorded by Smithers (1984) and

species with a hash are cosmopolitan species found opportunistically since 2007. All

species were identified by C. Smithers except Dorypteryx domestica and Liposcelis

bostrychophila, which were identified by C. Taylor.

Family

Amphientomidae

Caeciliusidae

Ectopsocidae

Lepidopscocidae

Liposcelididae

Peripsocidae

Philotarsidae

Pseudocaeciliidae

Psocidae

Psyllipsocidae

Trogiidae

indet.

Genus and Species

Amphientomid 1 sp. H

Caecilius sp. T *

Ectopsocus nr. erosus

(Enderlein)

Ectopsocus sp. B

Ectopsocus sp. N

Ectopsocus sp. P

Ectopsocus sp. R

Unknown Genus sp. Q

Pteroxanium sp. A

Liposcelid 1 sp. D

Liposcelid 2 sp. E

Liposcelid 3 sp. J

Liposcelid 4 sp. K

Liposcelid 5 sp. L

Liposcelis bostrychophila

Badonnel #

Liposcelis entomophila

(Enderlein) *

Liposcelis sp. O (complex)

Peripsocus fici Smithers *

Philotarsid 1 sp. G

Cladioneura foliata

Smithers *

Barrowia insularis

Smithers *

Dorypterx domestica

Smithers #

?Lepinotus sp. F

?Lepinotus sp. I

Gen. n. sp. M

Habitat

N, D

Vegetation type

Spinifex on limestone ridge and

floodplain

Rehabilitated site

Widespread on island

Spinifex on limestone ridge and

floodplain, coastal dunes

Spinifex on limestone ridge

Barge landing and rehabilitated

sites

Camp and Warehouse

Widespread on island

Coastal dunes

Coastal dunes,spinifex on

limestone ridgetop

Spinifex on limestone ridgetop

Spinifex on limestone floodplain

Camp

Widespread on island

Camp

Spinifex on limestone ridge

Coastal dune, spinifex on

limestone ridgetop

Widespread on island

Camp

Spinifex on limestone floodplain

Coastal dunes

Australian Entomologist, 2012, 39 (4) 257

Data were analysed as presence/absence. The collection protocol resulted in

large volumes of invertebrates being collected, which meant that counting

individuals was not time effective. For the GP sites, seasonal differences

(wet/dry) in species richness were tested using an independent t-test. This

was also done for species richness from two of the the collection periods

(2006/2007) of the NIS sites. The statistical program PASW Statistics 18.0.2

was used to carry out this analysis. The GP and NIS site-by-species matrices

were treated separately, as the collection protocols were not comparable. The

species matrices from each season or survey were combined and a similarity

matrix calculated for each site-by-species matrix using a Bray-Curtis

similarity index (Bray and Curtis 1957). Each similarity matrix was then

utilised to create a non-metric multidimensional scaling (NMDS) of the data

to provide a visual presentation of how similar each site’s species assemblage

was in comparison with another site. These analyses were carried out using

PRIMER-E v6.1.11.

Results

The 25 psocopteran species collected on Barrow Island from both native

vegetation and disturbed habitat sites are listed in Table 1. The two most

speciose families on the island are the Liposcelididae and Ectopsocidae.

Included in the table are three synanthropic species that have been collected

during surveys of the built environment on the island. These are Liposcelis

bostrychophila Badonnel, Liposcelis entomophila (Enderlein) and Dorypteryx

domestica (Smithers). Liposcelis bostrychophila and D. domestica have been

collected in the built environment (Chevron pers. comm.) since the GP and

NIS studies concluded. However, L. entomophila has not been collected since

its first collection on the island in 1982 (Smithers 1984). Images of 21 of the

psocopteran species of Barrow Island are publicly available and can be

accessed at the website [www.padil.gov.au/barrow-island]. |

There appeared to be no significant difference between seasons for species

richness at each GP site (P > 0.1). Species assemblage analysis did not yield

any discernible trends in terms of similarity between sites, even when the

assemblages from the two seasons were combined. Hence, the results from

the NMDS are not displayed. For the NIS sites, no significant difference in

species richness could be determined between the two surveys in 2006 and

2007 (P > 0.1). Here too, no discernible trends in species assemblages could

be observed within the NIS sites.

Discussion

There are 252 described species of Psocoptera listed for Australia by

Smithers (1996a), which is undoubtedly an underestimate due to the under-

sampling of the northwestern part of the country (Schmidt and New 2008).

For instance, Smithers (1996a) listed only four species of Liposcelididae for

this area, fewer than present on Barrow Island alone (three further species of

this family were recorded in the Western Australian wheatbelt by Smithers

258 Australian Entomologist, 2012, 39 (4)

1996b). Considering the remoteness of the northwestern part of Australia, the

Barrow Island invertebrate fauna has been relatively well studied. Barrow

Island has only slightly lower species diversity than other Australian offshore

islands. Psocopteran collections on Lord Howe Island yielded 27 species

(Smithers 2007) and 40 species were identified accumulatively on the four

larger Bass Strait Islands (Cole et al. 1989).

The collection of an undescribed species of Amphientomidae is of interest as

this is a rare family in Australia. Only three species have been described

previously from Australia, all from single specimens (Smithers 1989, New

1994). In contrast, the Barrow Island amphientomid is known from several

specimens, with both macropterous and brachypterous individuals present. A

description is currently in preparation.

Psocoptera are generally associated with microflora (e.g. fungi, lichen, algae)

and the highly seasonal arid environment on Barrow Island is not conducive

to supporting a diverse psocopteran fauna. Despite collecting only five

species, Smithers (1984) seemed to accept that such a low number of species

was to be expected from such an arid island. The collection of an additional

18 species between 2006 and 2007 shows that a large number of species can

be overlooked when only one method of collection is utilised. However, the

island’s highly seasonal environment and the generally accepted view that

Psocoptera are easily dispersed by wind (New 1987) can also complicate

species richness assessments.

Fig. 6. Dorypteryx domestica female collected in accommodation block on Barrow

Island: (a) lateral view; (b) close-up of forewing showing venation.

To date, only three widespread synanthropic species have been identified

from the island. One of these, Dorypteryx domestica (Fig. 6), has not

previously been recorded from Australia but is widely distributed, with

records from Europe, North America and Africa (Mockford 1993). Their

small size means that they easily escape detection in regions where they are

Australian Entomologist, 2012, 39 (4) 259

not previously known and their presence in Australia is not unexpected (T.R.

New pers. comm.).

With few exceptions, Dorypteryx species are known almost entirely from

synanthropic environments. Another species, Dorypteryx longipennis

Smithers, was first described from Australia as a quarantine intercept

(Smithers 1991), while D. longipennis and D. pallida Aaron are known to

have cosmopolitan distributions similar to that of D. domestica (Lienhard and

Schneider 1993, Mockford 1993).

Liposcelis bostrychophila and L. entomophila are also both widely distributed

cosmopolitan species (Mockford 1993). Despite regular pest inspections of

the built environment on Barrow Island and quarantine surveillance of the

surrounding vegetation, these species are collected very rarely. Liposcelis

entomophila is only known from its original collection in 1982. It is believed

that these known cosmopolitan species are not permanent inhabitants of the

built environment on Barrow Island and only occasionally re-occur within the

man-made structures, possibly being blown over or being carried from the

mainland. Their small size makes Psocoptera excellent wind dispersers, even

wingless species, and Liposcelis specimens have been collected in wind traps

on ships up to more than 1100 km from shore (Thornton and Harrell 1965).

For a comparison with regards to synanthropic species, Lord Howe Island has

four well known cosmopolitan species out of its 27 recorded species

(Smithers 2007) and Norfolk Island has seven cosmopolitan species out of 21

(Smithers et al. 1999).

The fact that there did not appear to be any seasonal or annual collection

differences in the species richness within the native vegetation and disturbed

area sites perhaps points to a more stable fauna for the island. Also, the high

ratio of un-named species to named species demonstrates the potential

uniqueness of the fauna. Smithers was not able to put names on the majority

of species from Barrow Island, which would indicate a high level of potential

endemicity either for the island itself or the north-west of Australia in

general.

Acknowledgements

The authors thank Chevron Australia for on-island support with logistics and

the Gorgon Joint Venture Partners for financial support to undertake the

project. The Gorgon Project is operated by an Australian subsidiary of

Chevron and is a joint venture of the Australian subsidiaries of Chevron

(approximately 47%), ExxonMobil (25%) and Shell (25%), Osaka Gas

(1.25%), Tokyo Gas (1%) and Chubu Electric Power (0.417%). Shae Callan,

Karl Edwards, Karen Edward, Rebecca Graham, Tim Lardner, Dr Anita

Lyons, Morgan Lythe, Natalie Randall, Brad Scanlon and Dusty Severtson all

assisted the project, either in the field and/or in the laboratory. Professor

Timothy New confirmed the identification of Dorypteryx domestica.

260 Australian Entomologist, 2012, 39 (4)

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Australian Entomologist, 2012, 39 (4): 261-272 261

THAUMASTOPSALTRIA SMITHERSI, A NEW CICADA FROM

NORTHERN QUEENSLAND WITH AN ANALYSIS OF ITS

PHYLOGENETIC RELATIONSHIPS (HEMIPTERA: CICADIDAE:

CICADETTINAE: CHLOROCYSTINI)

M.S. MOULDS

Entomology Department, Australian Museum, 6 College St, Sydney, NSW 2010

Abstract

Thaumastopsaltria smithersi sp. n. is only the second species of Thaumastopsaltria Kirkaldy to

be recorded from Australia. Its distinguishing features are documented together with notes on its

distribution and habitat. A revised phylogeny for the genus is provided and the phylogenetic

relationships of T. smithersi discussed.

Introduction

The genus Thaumastopsaltria Kirkaldy includes seven species found in

Waigeo and MisoGl Islands, mainland New Guinea, D'Entrecasteaux Islands,

New Britain, Umboi Island, northern Australia and, doubtfully, from Buka

Island and Bougainville (de Boer 1992). The genus has recently been

redefined and its distinguishing features summarised (Moulds 2012).

Thaumastopsaltria belongs to the tribe Chlorocystini Distant, 1905 and a

detailed analysis of relationships within the Chlorocystini was provided by de

Boer (1995a).

The single previously described Australian species of Thaumastopsaltria, T.

globosa (Distant), is known from along the eastern margin of Cape York

Peninsula from Banks Island in Torres Strait to Daintree, and from Groote

Eylandt, Northern Territory (Moulds 1990, as T. glauca, a synonym of T.

globosa). A second species is described here, with its distribution confined to

the rainforests south from Cooktown and with only a marginal overlap with

that of T. globosa. Phylogenetic relationships of this new species are

discussed below and a revised phylogeny for the genus is provided.

Terminology for morphological features follows Moulds (2005, 2012) and

for higher classification that of Moulds (2005). The following abbreviations

have been used for collections housing specimens: AE, collection of A.

Ewart, Golden Beach; AM, Australian Museum, Sydney; ANIC, Australian

National Insect Collection, Canberra; LP, -collection of Lindsay Popple,

Brisbane; MSM, collection of M.S. Moulds, Kuranda; ML, collection of Mark

Lane, Atherton; OM, Queensland Museum, Brisbane.

Thaumastopsaltria smithersi sp. n.

(Figs 1-9)

Type material. Holotype 6, QUEENSLAND: 6 Victor P1., Kuranda, 16°48'50"S

145°38'36"E, 18.xii.2008, Hill, Marshall, Moulds (AM). Paratypes (all northeastern

Queensland): 1 3, 2 29, Kuranda, 10.i.1983, 7.111982, 3.iii.1982, W.N.B. Quick;

1 9, 16.08S 145.26E, 2.5 km WSW of Noah Head, Cape Tribulation NP, 'Marrdja

Botanical Walk’, 2.xi.1993, D.C.F. Rentz & W.L. Lowe, stop 11 (ANIC); 1 ĝ, 1 9, nr

262 Australian Entomologist, 2012, 39 (4)

The Crater, 20 km S of Atherton, 21,29.xi.1987, D.A. Lane (AE); 1 4, 1 9, The

Creator, near Herberton, 3.i.1967, D.K. McAlpine & G. Holloway (AM); 1 ĝ, 1 9, nr

The Crater, 20 km S of Atherton, 29.xi.1987, D.A. Lane (LP); 5 dd, 1 9, Longlands

Gap, 20 km south Atherton, 19.xii.2001, D.A. Lane (ML); 1 9, Mt Hartley, nr

Rossville, S of Cooktown, 1.i.1984, M.S. & B.J. Moulds; 1 2°, Gap Ck, Mt Finlayson

Rg, S of Cooktown, 10.ii.1982, M.S. & B.J. Moulds; 2 34, Windsor Tableland, NW

of Mossman, 16.i.1988, M.S. & B.J. Moulds; 1 9, Mt Lewis via Julatten, 21.xii.2009,

Gilligan & Epstein; 5 £, 7 9°, Kuranda (Top of the Range), 19 Butler Dr., 335 m

elev., 16°48'S 145°38'E, 15-30.x.2004, 1-15.xi.2004, 1-15.xi.2006, 1-15.xii.2005, 1-

15.x11.2007, 16-31.x.2008, 1-15.xi.2008, 16-30.xi.2008, 1-15.xii.2008, 1-15.ii.2009,

1-15.i.2010, D.C.F. Rentz; 1 9, Kuranda, 10.xi.1987, 7.xii.1987, J. Hasenpusch; 1 3

(genitalia prep. THE3), Kuranda, xii.1985, R. Straatman; 1 ĝ, Kuranda, 5.ii.1987, R.

Straatman; 1 3, Whitfield Rg, nr Cairns, 2.xii.1986, M.S. & B.J. Moulds; 1 4, Hugh

Nelson Range, 20 km S of Atherton, 18.xi.1990, D.A. Lane; 2 3 (one genitalia prep.

THE4), 2 99, nr The Crater, 20 km S of Atherton, 21,29.xi.1987, D.A. Lane; 1 4,

The Crater, Atherton Tableland, 24.xii.2009, Gilligan & Epstein; 1 4, 'Chowchilla’,

Westcott Rd, Topaz, approx. 15 km SSE of Malanda, 18.xi.1990, W.T. Cooper; 1 9,

‘Chowchilla’, Westcott Rd, Topaz, approx. 15 km SSE of Malanda, 25.xi.2000, W.T.

Cooper; 1 9, Bartle Frere, 27.xii.2000, A. Polak; 1 2, Ravenshoe, i.1986, S. Lamond;

4 ő (one genitalia prep. THE2), 3 99, Polly Ck, Garradunga N of Innisfail,

20.x1.1987, 23.xi.1996, 20,23.x.2006, 9.1.2008, 12.xi.2008, J. Hasenpusch (MSM);

1 3, 1 Q, Kuranda (Top of the Range), 19 Butler Dr., 335 m elev., 16°48'S 145°38'E,

1-15.xii.2007, 1-15.xii.2008, D.C.F. Rentz; 1 ĝ, Kirrama Range, Douglas Ck Rd, 800

m, 7-12.xii.1986, Monteith, Thompson & Hamlet (QM).

Distinguishing features. Distinguished from all other Australian cicadas in

having 12 apical cells in the forewing (11 or 13 if aberrant, but usually so

only in one wing) and no subapical cells. Distinguished from 7. globosa by

the pair of black thoracic spots located near the ends of the anterior arms of

the cruciform elevation highlighting the scutal depressions (absent on T.

globosa), and the widely spaced ocelli that have the distance between them

considerably greater than their diameter (those of 7. globosa have the ocelli

close together, closer than their diameter). The male aedeagus is unique

within Thaumastopsaltria in having the apical gonopore sloping backwards

along the dorsal surface and terminating in a small dorsal crest (Fig. 5); other

species have a gonopore either sloping backwards along the ventral surface or

not sloping at all so that the aedeagus in lateral view has the apex square.

Description. Male (Figs 1, 3-8). Head light green, tending pastel in tone,

paler below than above, about as wide as lateral margins of pronotal collar;

eyes tan tending a little reddish; ocelli small and widely separated with the

distance between them much greater than their diameter; postclypeus and

anteclypeus a little paler than dorsal surface of head; postclypeus in lateral

view barely obtusely bent; in ventral view (Fig. 3) rostrum very pale brown

with darker tip, reaching near to bases of hind coxae. Thorax similar in colour

to head. Pronotum sometimes with hints of very pale orange centrally;

prothoracic collar mostly very pale orange across dorsal surface but

otherwise light green; paratnota rounded. Mesonotum usually with a very

Australian Entomologist, 2012, 39 (4) 263

Figs 1-2. Thaumastopsaltria smithersi sp. n.: (1) holotype male, dorsal view; (2)

paratype female, 19 Butler Drive, Kuranda, dorsal view.

pale orange fascia either side of midline aligned with distal ends of anterior

arms of cruciform elevation; scutal depressions highlighted as black spots.

Metathorax similar in colour to pro- and mesothorax. Wings (Figs 1-2)

hyaline. Forewing with 12 apical cells (often 11 or 13 if aberrant but if so

usually only in one wing); veins very fine, costal margin very pale yellowish

orange, veins otherwise pale green or black. Hind wing with seven apical

cells; distinct infuscation at distal ends of veins 1A and 2A spreading onto

wing margin; veins mostly very pale green but some pale brown or very pale

orange. Legs very pale green becoming very pale yellowish brown distally

from about mid length of tibia. Timbals (Fig. 4) with seven long ribs

spanning width of timbal plate and seven short ribs, the ribs occupying most

264 Australian Entomologist, 2012, 39 (4)

of timbal area; timbal plate narrow and mostly concealed. Opercula (Fig. 3)

very pale green, covering a little less than half tympanal cavity, transversely

broad and broadly rounded towards midline, clearly separated exposing most

of sternite I. Abdomen dorsally similar in colour to head and thorax, paler

ventrally, sometimes tending very pale bluish green and often with hints of

very pale brown around distal midline and sternite VIII.

Genitalia (Figs 5-8). Pygofer in ventral view (Fig. 8) with cavity tapering

towards base, in some individuals forming a distinct 'V' shape; pygofer upper

and basal lobes in lateral view (Fig. 7) weakly developed and broadly

rounded. Claspers well developed, claw-like, their apices slightly diverging

and slightly down-turned. Uncus degenerate, barely discernible. Aedeagus

with theca (Fig. 5) almost cylindrical, gradually tapering towards apex,

straight or gently curved in a shallow arc; gonopore apical but sloping

backwards along dorsal surface of theca, the opening at its proximal end

narrowing and forming a small crest in lateral view; basal plate almost

rounded in dorsal view (Fig. 6), stepped in lateral view (Fig. 5).

Female (Fig. 2). Similar to male. Abdominal segment 9 uniformly pale green,

along dorsal midline just a little longer than abdominal tergites 7 and 8

combined; ovipositor sheath terminating about level with caudal beak; caudal

beak sharply pointed.

Measurements. Range and mean (in mm) for 10 males and 10 females,

including smallest and largest available specimens. Body length: male 21.4-

27.2 (25.1); female 21.0-25.5 (23.3). Head width (including eyes): male 5.7-

6.7 (6.2); female 5.9-6.9 (6.5). Pronotum width (across lateral angles): male

6.7-8.8 (7.7); female 6.7-8.7 (7.9). Forewing length: male 28.3-37.3 (32.3);

female 27.5-35.1 (31.4). Forewing width: male 10.2-13.8 (11.9); female 9.7-

13.1 (11.5). Forewing ratio width/length: male 2.7 + 0.1; female 2.7 + 0.1.

Etymology. Named in memory of Dr Courtenay Smithers, a friend and

mentor, past Curator of Insects, Principal Curator and Deputy Director of the

Australian Museum. Courtenay had broad interests in entomology covering

most Orders and his extensive publications contributed significantly to our

knowledge of Australian insects.

Song. Singing occurs at dusk and is a high pitched, whistle-like, continuous

call.

Distribution and habitat (Fig. 10). Northeastern Queensland, south from

Mount Hartley (south of Cooktown near Rossville) to the Kirrama Range

(north-west of Cardwell). Most records are from mountain areas between

about 300 m and 800 m but there are records from much lower altitudes,

including Gap Ck south of Cooktown at 40 m, Marrdja Botanical Walk, Cape

Tribulation at 21 m and Polly Ck, Garradunga at 25 m. The species has been

found only in rainforest. Adults have been taken from mid October to mid

February but are most common during November and December.

Australian Entomologist, 2012, 39 (4) 265

~— anterior

3 4

Figs 3-6. Thaumastopsaltria smithersi sp. n.: (3) ventral view of body; (4) left timbal

showing ribs; (5) aedeagus in lateral view; (6) basal plate of aedeagus in dorsal view,

apex at bottom. op operculum; pc postclypeus; ro rostrum; ty tympanum.

266 Australian Entomologist, 2012, 39 (4

7 8

Figs 7-8. Thaumastopsaltria smithersi sp. n., male genitalia: (7) lateral view; (8)

ventral view with claspers spread apart. bp/ basal pygofer lobe; c/ clasper; gon

gonopore of theca; pyg pygofer; th theca of aedeagus; up/ upper pygofer lobe.

Phylogenetic relationships

De Boer (1995a) presented a cladistic analysis based on morphological

characters and incorporating all known species of the tribe Chlorocystini. He

found 600 equally parsimonious trees. The 50% majority rule tree for these

600 trees was found to contain nearly all previously recognized genera and

species groups as monophyletic groups and was chosen by de Boer as the

preferred tree showing relationships. De Boer's cladistic analysis has been

adopted here to determine the phylogenetic position of 7. smithersi sp. n.

within the genus Thaumastopsaltria. A cladogram was reconstructed from de

Boer's data set retaining only those characters suggesting phylogenetic

relationships within Thaumastopsaltria and the outgroup. De Boer found that

Species of the genus Mirabilopsaltria de Boer were sister to those of

Thaumastopsaltria. Mirabilopsaltria globulata de Boer and M. humilis

(Bléte) were chosen as outgroup taxa. Character and state numbers used by

de Boer (1995a) were retained to allow direct comparison with his analysis.

Data were analysed using the heuristic search parsimony algorithms

implemented with PAUP* version 4.0b2 (Swofford 1998). Tree searches

utilized the tree bisection reconnection algorithm (TBR) conducting 1,000

random addition searches (RAS) starting from random trees; other settings

were left at their default values. All characters were unweighted and all

multistate characters were treated as unordered. Unknown character states

Australian Entomologist, 2012, 39 (4) 267

were scored as ‘?’. The matrix of species and assigned states is given in Table

1, followed by the characters and character states used, adopted from de Boer

(1995a) using his character numbers and states but with minor modification

as stated in the notes below relevant characters.

Table 1. Character matrix for the eight known species of Thaumastopsaltria and two

species of Mirabilopsaltria used in the parsimony analysis (from de Boer 1995a).

Missing data and character states unknown are scored as '?' Character numbers are

those used by de Boer in his larger analysis of the Chlorocystini [reading downwards

in heading]. However, character states have been renumbered sequentially from zero

for the purpose of the current analysis in order to elucidate plesiomorphic states.

00000 00000 00000 00011 11111 111

Species 01122 33345 66667 89900 22344 555

52724 23514 15695 20503 09125 023

Mirabilopsaltria humilis (Blöte) 00000 00000 00000 00000 00000 000

Mirabilopsaltria globulata de Boer 00001 11110 10000 00011 00000 010

Thaumastopsaltria adipata (Stal) 10110 0120? 01012 00111 10100 0??

Thaumastopsaltria pneumatica de Boer 11102 01201 01011 01111 10101 020

Thaumastopsaltria spelunca de Boer 10112 11110 10011 11110 02211 120

Thaumastopsaltria globosa (Distant) 10102 11102 11112 11100 02211 121

Thaumastopsaltria sicula de Boer 10100 00010 10011 01100 02011 120

Thaumastopsaltria lanceola de Boer 10102 11000 01112 10100 01002 021

Thaumastopsaltria sarissa de Boer 11102 11000 01112 11101 01001 020

Thaumastopsaltria smithersi sp. n. 11103 00201 11712 10101 02020 201

Characters and character states

005.

012.

017.

022.

024.

032.

Postclypeus in lateral view: (1) not swollen; (4) protruding in obtuse angle.

Distance between lateral ocelli: (1) about equal to distance between lateral

ocellus and eye; (2) generally slightly larger than distance between lateral

ocellus and eye.

Paranotum (anterolateral side of pronotal collar): (1) rounded; (4) forming

continuous ridge.

Proximal spine of foreleg: (1) not shorter than distance to middle spine; (2)

distinctly shorter than distance to middle spine.

Colour of forewing: (1) hyaline, completely without colour; (2) opaque greenish

or reddish; (3) slightly reddish but still hyaline; (5) slightly greenish but still

hyaline.

NOTE: De Boer listed four states for this character but his state 4 was not

relevant to this analysis and is omitted. However, another state is here added

(state 5) to accommodate the unique forewing colour of T. smithersi.

Subapical cells of forewing: (1) absent; (2) present but not forming a continuous

band.

268

033.

035.

041.

054.

069.

075.

082.

090.

095.

100.

103.

120.

129.

131.

142.

Australian Entomologist, 2012, 39 (4)

Base of first apical area of forewing: (1) distally of base of third apical area; (3)

distinctly proximally of base of third apical area.

Number of apical cells in hind wing: (1) six; (3) six or seven; (4) seven or more,

NOTE: It is not clear why de Boer did not designate discrete character states.

Examination of material available to me suggests that some species have either

six or seven hind wing apical cells (often on the same specimen), while other

species consistently have either six or seven or more; this is the presumed

interpretation adopted here when scoring T. smithersi.

Timbal cavity: (1) broad; (2) narrow.

Distal part of female operculum: (1) rectangular oblong; (2) trapezoid; (3)

narrow sickle-shaped.

. Second tergite of male: (1) forming a distinct ridge along timbal cavity; (2) not

forming a distinct ridge along timbal cavity.

NOTE: The scoring of this character had been accidentally reversed in de Boer

(199Sa).

- Male auditory capsule: (1) swollen and protruding; (2) not swollen and not

protruding.

- Male pygofer and 9th sternite: (1) resting on 8th sternite; (2) elevated respective

to 8th sternite.

Male pygofer: (1) not curved to ventral position; (2) curved to ventral position

on abdomen.

Apex of male dorsal beak: (2) rounded; (3) truncate or weakly concave; (5)

broadly rounded.

NOTE: The scoring of T. smithersi for these subjective character states was

done by comparing the dorsal beak of smithersi with other available material

and adopting the scoring of the most similar species.

Protuberance on lateral lobe of male pygofer: (1) not triangularly swollen; (2)

triangularly swollen.

Male pygofer: (1) not swollen at ventral margin; (2) swollen at ventral margin.

Ventral half of male pygofer opening: (1) triangular and pointed; (2) broad and

rounded.

Clasper: (1) strongly bent and hook-shaped; (2) nearly straight and posteriorly

directed.

Clasper: (1) parallel; (2) diverging towards apices.

Dorsal edge of clasper: (1) not curved around aedeagus; (5) globularly rounded.

Clasper: (1) without ventromedial protuberance; (2) with broad and laminiform

ventromedial protuberance; (3) with narrow and spiny ventromedial

protuberance.

Clasper: (1) without lateral process; (2) with lobate lateral process; (3) with

lateral thorn.

Apical part of aedeagus: (1) not abruptly narrowing; (2) abruptly narrowing and

upcurved; (4) abruptly narrowing on dorsal surface.

Australian Entomologist, 2012, 39 (4) 269

NOTE: De Boer listed three states for this character but state 3 was not relevant

to this analysis and is omitted. However, another state is here added (state 4) to

accommodate the unique aedeagus of T. smithersi.

145. Aedeagus: (1) without dorsal crest; (2) with single dorsal crest; (3) with paired

dorsal crests.

150. Aedeagal pore: (3) apically pointed but not widening proximally; (5) rounded;

(6) apically wide and narrowing proximally.

NOTE: An additional state (state 6) is here added to accommodate the unique

aedeagus of T. smithersi.

152. Ovipositor sheaths: (1) reaching to apex of dorsal beak; (2) reaching just

beyond apex of dorsal beak; (3) reaching far beyond apex of dorsal beak.

153. Apex of female dorsal beak: (1) sharply pointed; (2) broadly rounded.

M. humilis

32 35 41 61 100 103

fmf}

H H

2>11>32>1 1>2 2>1 1>2

M. globulata

35 54 103

1>2 1>3 1>3

i i

2>11>5 1>33>2

T. pneumatica

Ši

62 2>12>31>21>3

1>4 4>12>31>22>1 5>22>1

T. sarissa

35 54 131 153

T. globosa

1>31>21>22>1

ini

1>23>21>25>3] 41 65 66 75

2>12>12312>3

3>11>23>11>2

Fig. 9. The single most parsimonious tree derived using the procedures described in

the text above (length 66, CI 57, RI 61) from an analysis of all Thaumastopsaltria

species, employing Mirabilopsaltria humilis and M. globulata as outgroups, with all

characters unordered and equally weighted. Numbers at nodes are bootstrap values

>50% from 1,000 replications. Character transformations (generated using Clados) are

represented by black bars = non-homoplasious forward changes; grey bars =

homoplasious forward changes; and white bars = reversals (whether homoplasious or

not). The analysis was run using the data set in Table 1 and state numbers modified

(after analysis) to correspond with those used by de Boer (1995a).

270 Australian Entomologist, 2012, 39 (4)

Results from running de Boer's (1995a) reduced data set produced three

equally parsimonious trees. However, by omitting de Boer's homoplasious

character (character 103) from the analysis only one shortest tree was

obtained. This tree was identical in topology to de Boer's 50% majority rule

tree for the genus Thaumastopsaltria. T. smithersi sp. n. was then scored and

the data added to the data set and the analysis re-run. Three additional

character states were added to accommodate the variant forewing colour of T.

smithersi (state 3 in character 24) and unique morphology of the aedeagus

(state 2 in character 142 and state 2 in character 150) (Table 1). Three equally

parsimonious trees were obtained but the relationships of T. smithersi were

largely inconclusive. However, by again eliminating de Boer's character 103,

just a single tree (Fig. 9) resulted, which was fully resolved and

topographically similar to de Boer's 50% majority rule tree.

Although the single tree obtained here agrees with that of de Boer's 50%

majority rule tree, it should be kept in mind that both trees are based on the

same data set. Only the node distinguishing Thaumastopsaltria and one

internal node are well supported statistically. Molecular data is currently

unavailable for Thaumastopsaltria species but such data may provide

confidence for internal nodes.

Relationships of Thaumastopsaltria smithersi sp. n.

The fully resolved tree obtained (Fig. 9) shows that T. smithersi sp. n. forms a

monophyletic group together with T. adipata (Stal) (known from just a single

specimen from Misodl Island) + T. pneumatica de Boer (known only from

Mt Dayman in eastern Papua New Guinea). This group of three is sister to all

other Thaumastopsaltria species, including the only other Australian species,

T. globosa (Distant).

Thaumastopsaltria globosa has a wide distribution throughout southern parts

of Papua New Guinea, eastern Queensland south to Daintree, and Groote

Eylandt on the Northern Territory side of the Gulf of Carpentaria (Fig. 10), a

distribution accounted for when sea levels were much lower during several

Pliocene-Pleistocene ice ages (between 5-3 and 0.01 Mya), when Queensland

was connected to New Guinea via a land bridge and the Gulf of Carpentaria

was largely dry (Pigram and Davies 1987, de Boer 1995b). T. globosa is not a

wet rainforest species and would not have been inhibited in its spread by this

specific habitat requirement. In contrast, T. smithersi is restricted to wet

tropical rainforest between Cooktown and Cardwell (Fig. 10), a habitat that

would have shrunk during those past ice ages when conditions were much

dryer (Hopkins eg al. 1993),

It is difficult to explain how T. smithersi could have become so isolated from

its New Guinean sister species, none of which now inhabit the southern

regions of that large island anywhere near Australia. This isolation must have

originated a long time ago, possibly as long as 25 Mya when other, now

Australian Entomologist, 2012, 39 (4) 271

endemic, Australian Chlorocystini are thought to have entered Australia

following the Sepik Arc collision (Daly et al. 1991, de Boer 1995b).

Banks Island

a globosa

e smithersi

& Groote poko

Eylandt

Fig. 10. Distribution of Thaumastopsaltria smithersi sp. n. and T. globosa.

On the monophyly of Thaumastopsaltria Kirkaldy, 1900

De Boer (1992) distinguished Thaumastopsaltria on the basis of the shape of

the postclypeus and long female ovipositor. The postclypeus in lateral view is

obtusely bent at about mid point and thereafter straight to anteclypeus, and

the lateral margins show several rows of short parallel ridges. Further, de

Boer pointed out that the male pygofer upper lobes in most species are

distinctly inflated towards the ventral margin, a feature believed to be unique

within Thaumastopsaltria.

The monophyly of Thaumastopsaltria (incorporating T. smithersi sp. n.) is

strongly supported in the cladistic -analysis by seven non-homoplasious

synapomorphies and a bootstrap of 86%. However, three of those

synapomorphies have one or more reversals within Thaumastopsaltria

(characters 24, 65 and 75). The combination of the remaining four

synapomorphies (characters 5, 17, 69 and 95 unambiguously define the genus

as follows: (a) shape of the postclypeus as mentioned above; (b) paranotum

(the lateral side of pronotal collar) rounded; (c) the male pygofer not curved

to ventral position; (d) the ventral half of pygofer opening triangular and

272 Australian Entomologist, 2012, 39 (4)

pointed rather than broad and rounded. The combination of these four

synapomorphies clearly characterises the genus although some of these may

have been homoplasious in de Boer's (1995a) large analysis (shared with

nodes in other parts of his tree). As de Boer (1992) pointed out, the shape of

the postclypeus appears to be the most reliable distinguishing feature. Note

that the long female ovipositor, considered by de Boer as a defining attribute

for the genus, is not included among the seven synapomorphies identified by

the present cladistic analysis (character 152). This is because the ovipositor

of T. smithersi is short, terminating at about the distal end of abdominal

segment 9 and, as T. smithersi is near the base of the tree, the analysis treats

the character as homoplasious.

Acknowledgements

For helpful comments on the manuscript I am especially grateful to Dr Tony

Ewart and Dr Lindsay Popple. A number of people kindly provided

specimens for study as detailed in the list of types; to all I extend my sincere

thanks. The photographs for Figs 1 and 2 were kindly taken by David Rentz.

The line drawings for Figs 3-8 were done by Ivan Nozaic. Financial

assistance for the figures was provided by the National Science Foundation,

grant number DEB 09-55849.

References l

BOER, A.J. de 1992. The taxonomy and biogeography of the genus Thaumastopsaltria Kirkaldy,

1900 (Homoptera, Tibicinidae). Beaufortia 43: 17-44.

BOER, A.J. de 1995a. The phylogeny and taxonomic status of the Chlorocystini (Homoptera,

Tibicinidae). Bijdragen tot de Dierkunde 65: 201-231.

BOER, A.J. de 1995b. Islands and cicadas adrift in the west-Pacific. Biogeographic patterns

related to plate tectonics. Tijdschrift voor Entomologie 138: 169-244.

DALY, M.C., COOPER, M.A., WILSON, I., SMITH, D.G. and HOOPER, B.G.D. 1991.

Cenozoic plate tectonics and basin evolution in Indonesia. Marine and Petroleum Geology 8: 1-

213

HOPKINS, M.S., ASH, J., GRAHAM, A.W., HEAD J. and HEWETT R.K. 1993. Charcoal

evidence of the spatial extent of the Eucalyptus woodland expansion and rainforest contractions

in North Queensland during the late Pleistocene. Journal of Biogeography 20: 357-372.

MOULDS, M.S. 1990. Australian cicadas. New South Wales University Press, Kensington; 217

pp, 24 pls.

MOULDS, M.S. 2005. An appraisal of the higher classification of cicadas (Hemiptera:

Cicadoidea) with special reference to the Australian fauna. Records of the Australian Museum

57: 375-446. http://dx.doi.org/10. 3853/).0067-1975.57.2005.1447

MOULDS, M.S. 2012. A review of the genera of Australian cicadas (Hemiptera: Cicadoidea).

Zootaxa 3287: 1-262. http://mapress.com/zootaxa/2012/f/zt03287p262.pdf

PIGRAM, C.J. and DAVIES, P.J. 1987. Terranes and the accretion history of the New Guinea

orogen. BMR Journal of Australian Geology and Geophysics 10: 193-212.

SWOFFORD, D.L. 1998. PAUP* 4.0. Phylogenetic analysis using parsimony. Beta version

4.0d100. Smithsonian Institution.

Australian Entomologist, 2012, 39 (4): 273-276 273

A NEW SUBSPECIES OF DELIAS MYSIS (FABRICIUS)

(LEPIDOPTERA: PIERIDAE) FROM THE GULF OF

CARPENTARIA, QUEENSLAND, AUSTRALIA

G. DANIELS

School of Biological Sciences, University of Queensland, St Lucia, Qld 4072

Abstract

Delias mysis smithersi subsp. n. is described from Karumba and Kowanyama, southeastern Gulf

of Carpentaria, Queensland.

Introduction

Delias mysis (Fabricius) is a common species along the eastern coast of

Queensland (subsp. D. m. mysis) and less common in the Northern Territory

(subsp. D. m. aestiva Butler). A new subspecies, collected enroute to the

Royal Geographical Society of Queensland Flinders 2002 Expedition to

Sweers Island, is from the lower part of the Gulf of Carpentaria, Queensland

and is geographically midway between the eastern and western subspecies.

Delias mysis smithersi subsp. n.

(Figs 1-6)

Types. Holotype 3, AUSTRALIA (QUEENSLAND): Karumba, Qld, 17°29'S

140°50'E, 9.x.2003, G. and A. Daniels, urban garden. Paratypes: 1 4, Karumba, Qld,

17°29'S 140°50'E, 12.xii.2002, G. Daniels, urban; 1 ĝ, Mitchell River, Q.,

26.x.[19]71.1.S.R. Munro, I.S.R. Munro Collection (all in Australian Museum).

Description. Male. Wingspan 56-60 mm. Upperside. Forewing white, with

narrow black costa, apex black enclosing five large white subapical spots.

Hind wing white, with narrow black terminal band. The red and black

markings on the underside of the hind wing are faintly visible. Underside.

Forewing similar to upperside but with the first subapical spot faintly yellow.

Hind wing white, with base and dorsum broadly yellow and with a narrow

black terminal band bordered by a much narrower scarlet subterminal band

which ends at cell Rs; the inner margin of the subterminal band is bordered

by an extremely narrow black band which is absent in cell Rs and may be

absent in cell M).

Female unknown.

Etymology. I am pleased to dedicate this paper to the memory to my late

friend, Courtenay Smithers, who gave me much help and encouragement

when starting out in entomology many years ago.

Taxonomy

The reduced inner black subterminal band is similar to that of Delias doylei

Sanford & Bennett from Papua New Guinea but that species has much of the

hind wing underside suffused yellow and the subapical spots of the forewing

underside are yellow.

274 Australian Entomologist, 2012, 39 (4)

Figs 1-8. Delias mysis males. (1-6) D. mysis smithersi subsp. n. (1-2) Holotype from

Karumba: (1) underside; (2) upperside. (3-4) Paratype from Karumba: (3) underside;

(4) upperside. (5-6) Paratype from Mitchell River: (5) underside; (6) upperside. (7) D.

mysis aestiva from Northern Territory. (8) D. mysis mysis from Queensland.

Australian Entomologist, 2012, 39 (4) 275

Davenport and Mastrigt (2008) reviewed the Delias mysis group of species

and raised Delias mysis lara (Boisduval) from New Guinea to species level,

based upon the sympatric distribution of D. mysis lara and D. mysis nemea

Fruhstorfer in the Merauke District of Papua. Davenport and Mastrigt

distinguished the two species by the subapical spots on upperside of forewing,

being well developed and streak-like, forming a white band with black veins in

D. mysis, while in D. lara they are white, quite poorly developed and even

sometimes absent. Delias mysis smithersi agrees with the characters given by

Davenport and Mastrigt for Delias mysis.

The upperside subapical forewing spots of D. m. smithersi are larger and the

black apical area is less extensive than in D. mysis mysis (Fig. 8). The narrow

terminal black band of the hind wing upperside is narrower than in D. mysis

mysis. The hind wing underside has the scarlet subterminal band ending in

cell Rs; this feature is shared with D. mysis aestiva (Fig. 7).

Delias mysis smithersi differs from D. mysis aestiva by the much narrower

black terminal band of the hind wing underside and, on the hind wing

underside, the black band on the inner margin of the scarlet subterminal band

is much wider in D. m. aestiva (Fig. 7).

Although some specimens of D. mysis mysis have reduced black markings on

the upperside, none had a corresponding reduction to the black and scarlet

markings on the underside of the hind wing (n = 82) and none had the inner

subterminal black band anywhere near as reduced.

The three specimens show little variation. In two specimens the scarlet band

in the hind wing cell Rs is present as a spot; one specimen has the apical

black area on the upper- and undersides much smaller with a corresponding

reduction in the size of the subapical white spots. All specimens are worn,

which could indicate a winter flight period.

Distribution

The nearest populations of Delias mysis to Karumba and Mitchell River can

be found some 500 km to the east, along the eastern coast of Queensland

(Braby 2000) and a similar distance to the north-east at Weipa (Hancock and

Monteith 2004), where adults are associated with rainforest. Karumba is on

the northern edge of the vast savannah-eucalypt woodlands where there are

no large tracts of rainforest. Adults of the Northern Territory subspecies, D.

mysis aestiva, can be found near mangroves and melaleuca swamps (Ted

Fenner pers. comm.) and this is perhaps the habitat where the Karumba

population is more abundant, as the type locality is situated approximately

500 m from the mangrove areas of the mouth of the Norman River.

Collecting there is another matter!

The two Karumba specimens were collected in the grounds of Matilda's End

Motel. The first specimen collected was feeding at eucalyptus blossom.

Within 200-300 m of the motel grounds are numerous eucalypts festooned

276 Australian Entomologist, 2012, 39 (4)

with mistletoe. Despite observing the mistletoe for several days no Delias

mysis were observed, although Delias argenthona (Fabricius) was found to

be exceedingly abundant.

Talbot (1937) examined a male specimen from Groote Eylandt (coll. Bodley,

ex coll. Joicey) in the northwestern area of the Gulf of Carpentaria. Talbot

placed this specimen in subsp. D. m. mysis, stating: ‘This specimen should

belong to aestiva; but only differs from mysis in having a narrow black outer

border on the hind wing beneath.’ An examination of the collection of the

Natural History Museum, London by R. Eastwood in 2004 failed to locate

this specimen and its whereabouts and subspecific placement remain

unknown. Apart from this problematic Groote Eylandt record, D. m. aestiva

is only known to come as far eastwards in the Northern Territory as the

Cobourg Peninsula (fide Braby 2000).

Acknowledgements

The first specimen from Karumba was collected during the Royal

Geographical Society of Queensland Flinders 2002 Expedition to Sweers

Island and I thank the Society for its support. Rod Eastwood kindly examined

material in the Natural History Museum, London. Max Moulds (Australian

Museum), kindly loaned material in his care. Fred Gerrits provided a copy of

Davenport and Mastrigt’s publication.

References

BRABY, M.F. 2000. Butterflies of Australia. Their identification, biology and distribution. 2

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

DAVENPORT, C. and MASTRIGT, H. von 2008. Revision of Delias mysis (Fabricius, 1775)

and closely related species (Lepidoptera: Pieridae). Suara Serangga Papua 3(2): 15-31.

HANCOCK, D.L. and MONTEITH, G.B. 2004. Some records of butterflies (Lepidoptera) from

western Cape York Peninsula, Queensland. Australian Entomologist 31: 21-34.

TALBOT, G. 1937. A monograph of the pierine genus Delias. Part VI. Pp i-v, 261-656, pls 8-36,

44-53, 60-67, 71. British Museum (Natural History), London.

Australian Entomologist, 2012, 39 (4): 277-280 277

NOTES ON THE BIOLOGY OF NACADUBA NIUEENSIS LACHLAN

(LEPIDOPTERA: LYCAENIDAE) FROM NIUE, SOUTHWESTERN

PACIFIC OCEAN

R.B. LACHLAN

Entomology Department, Australian Museum, 6 College St, Sydney, NSW 2010

Abstract

Additional specimens of Nacaduba niueensis Lachlan, 2012 are recorded from the Pacific island

of Niue and notes on behaviour of both sexes are recorded for the first time. Fresh females are

illustrated and the original description augmented.

Introduction

On a second trip to the Pacific island of Niue between 30 March and 13 April

2012, a further 37 specimens of Nacaduba niueensis Lachlan, 2012 were

collected. This included 33 males but only 4 females.

Prior to this second survey, N. niueensis was known only from two

specimens, a male and female described by Lachlan (2012). These two

specimens had been collected some 30 years apart.

On this recent trip, from one to six specimens were encountered on each day

of the survey. The survey was conducted towards the end of the rainy season

and all the days were at least partly sunny.

This paper is dedicated to the memory of a good friend, Courtenay N.

Smithers, who took an active interest in the butterflies of Pacific islands.

Discussion

Despite careful examination of numerous sites around much of the island,

specimens of N. niueensis were only ever seen or collected in the far north-

west, even though suitable forest habitat is found around much of the island.

There were no obvious reasons why this species would be confined to this

one area of the island.

During the survey, most males and half the females were collected while

alighting on the common forest tree Alphitonia zizyphoides (family

Rhamnaceae), which was in flower. The males were often observed flying

quite rapidly, for short periods, as they arrived at the upper reaches of this

tree, then alighting on either the tips of leaves among the clusters of small,

white flowers or the flowers themselves, usually facing outwards. They often

engaged and drove off other lycaenids, bees and wasps that were attempting

to feed on the flowers. They would then fly off over the top of the canopy.

Rarely were any other specimens sighted or collected away from this species

of tree and none was observed on any other species of tree in the forest or

cultivated areas around the island.

Ten males and two females were collected away from this flowering tree in

the far north of the island, while flying in, or near, a short section of a partly

278 Australian Entomologist, 2012, 39 (4)

enclosed forest trail. Six of these males and the two females were collected

inside the shaded forest adjacent to the trail. They were all flying slowly

between 1 and 2 metres above the ground.

Figs 1-2. Nacaduba niueensis, female: (1) upperside; (2) underside.

The author was lucky to come across two of these flowering trees about 80

metres apart, which allowed relatively easy access to the upper reaches of

each tree with a long-handled net. Alphitonia zizyphoides trees often reach

heights in excess of ten metres, so it is not possible to reach the tops of most

of these trees to observe or collect N. niueensis, even if they are present.

Australian Entomologist, 2012, 39 (4) 279

It is now clear that N. niueensis primarily inhabits the canopy of the forest on

Niue, which largely explains why only two specimens were collected in the

previous 30 years and it also appears, at present, to be rather localised in its

distribution, for unknown reasons. The vast majority of specimens collected

showed varying degrees of wing damage, indicating that they had been flying

for some time. There were very few fresh specimens on the wing. This may

be indicative of the end of the season for this species. In addition to the

specimens collected, close to that number were sighted but not collected,

mostly around the two flowering trees mentioned. Therefore, N. niueensis

appears to be relatively common, at least locally, but very difficult to collect.

Lachlan (2012) described the paratype female from the only specimen known

at that time but it was slightly damaged and 32 years old. Additional notes

from fresh females are provided below.

Description. Female (Figs 1-5). Upperside, unicolorous very dark brown as in

Figs 1, 3-5; on three of the four specimens the basal region of the forewing

on either side of the cubitus vein is lightly irrorated with bright, bluish-purple

scales to varying degrees; the hind wing is also irrorated with bluish-purple

scales but to a lesser degree and only from the basal and subbasal regions.

This scaling is more evident when viewed obliquely. In both sexes, on all

specimens collected, there is a clearly visible patch of white scaling on the

basal third of the inner margin of both hind wings, as in Figs 1, 3-5. This

white scaling was not seen as clearly on the holotype or at all on the paratype

female. The forewing termen is more rounded than in the male.

Fig. 3. Nacaduba niueensis, female upperside showing distinct bluish scaling.

280 Australian Entomologist, 2012, 39 (4)

Figs 4-5. Nacaduba niueensis, female uppersides: (4) showing a trace of bluish

scaling; (5) with no bluish scaling.

Acknowledgements

I am very grateful to Ted Edwards (ANIC, Canberra) and Dr Max Moulds

(Research Fellow, Australian Museum, Sydney) for their very helpful

comments on the manuscript. I also wish to acknowledge the assistance of Dr

David Britton (Collection Manager, Entomology Department, Australian

Museum, Sydney) for his production of the digital images of the females.

R.G. Coveny (curator of the Rhamnaceae, The Royal Botanic Gardens,

Sydney) is also sincerely thanked for identification of the flowering tree.

Reference

LACHLAN, R.B. 2012. A new species of Nacaduba Moore (Lepidoptera: Lycaenidae) from

Niue, southwestern Pacific Ocean. Australian Entomologist 38(2): 49-54.

Australian Entomologist, 2012, 39 (4): 281-292 281

REVIEW OF AUSTRALIAN PHYLLODES IMPERIALIS DRUCE

(LEPIDOPTERA: EREBIDAE) WITH DESCRIPTION OF A NEW

SUBSPECIES FROM SUBTROPICAL AUSTRALIA

D.P.A. SANDS

CSIRO Ecosystem Sciences, PO Box 2583, Brisbane, Old 4001

Abstract

The subspecies of Phyllodes imperialis Druce are reviewed: P. i. meyricki Olliff from tropical

northeastern Australia and New Guinea is validated and compared with the nominotypical P. i.

imperialis from the Solomon Islands and P. i. dealbata Holloway from New Caledonia. P.

imperialis smithersi subsp. n., from subtropical southeastern Queensland and northeastern New

South Wales, is described and figured.

Introduction

Several genera of large moths previously referred to the subfamily

Catocalinae (Noctuidae), including Phyllodes Boisduval (1832), were

elevated to the family Erebidae (Calpinae: tribe Phyllodini) by Zahiri et al.

(2011), Holloway (2011) and Zahiri et al. (2012). Many species in this group

belonging to endemic Australian genera (Common 1990) have remained

unplaced (Edwards 1996) since the review of Noctuidae by Kitching (1984).

Taxa then included in the family Noctuidae have since been rearranged by

Zahiri et al. (2011), with Phyllodini now including Phyllodes, Xylophylla

Hampson, Oporophylla Hampson, Lobophyllodes Hampson, Minoides

Guenee and Minophyllodes Joannis. Of these genera, Phyllodes is the only

genus occurring in Australia.

Poole (1989) listed 11 species (with synonyms) of Phyllodes and regarded P.

imperialis Druce, 1888, P. meyricki Olliff, 1889 and P. papuana Hampson,

1913 as separate species. Earlier authors, e.g. Olliff (1889) listed 12 species

of Phyllodes, Seitz (1923) listed 13 and Hampson (1913) provided a key to

11 species. Phyllodes i. imperialis was described from the Solomon Islands

(Druce 1888) and this subspecies is also known from Bougainville, Papua

New Guinea (ANIC unpublished). Several other populations of P. imperialis

include the distinctive subspecies P. i. dealbata Holloway from New

Caledonia (Holloway 1979), while other island populations from Vanuatu

(Viette 1950), New Britain and New Ireland (Sands unpublished) may also be

distinct. Phyllodes i. meyricki Olliff, from northern Queensland and New

Guinea, is recognised here as a subspecies differing morphologically from P.

i. imperialis and P. i. dealbata. The population previously referred to as a

southern subspecies of P. imperialis by Sands (1999) is listed both federally

(EPBC 2002, Clark and Spier-Ashcroft 2003) and in New South Wales

(TSCA 1995) as an endangered subspecies. The habitat for this subspecies,

subtropical lowland rainforest, is listed federally as a critically endangered

ecosystem (EPBC 2002; endorsed November 2011).

This southern population of P. imperialis from eastern Australia is described

here as new; it, P. i. meyricki and P. i. imperialis are illustrated in Figs 1-12.

282 Australian Entomologist, 2012, 39 (4)

Figs 1-3. Phyllodes imperialis, male uppersides: (1) P. i. meyricki; (2) P. i. smithersi

subsp. n.; (3) P. i. imperialis.

Australian Entomologist, 2012, 39 (4) 283

Figs 4-6. Phyllodes imperialis, male undersides: (4) P. i. meyricki; (5) P. i. smithersi

subsp. n.; (6) P. i. imperialis.

284 Australian Entomologist, 2012, 39 (4)

Figs 7-9. Phyllodes imperialis, female uppersides: (7) P. i. meyricki; (8) P. i. smithersi |

subsp. n.; (9) P. i. imperialis.

Australian Entomologist, 2012, 39 (4) 285

Figs 10-12. Phyllodes imperialis, female undersides: (10) P. i. meyricki; (11) P. i.

smithersi subsp. n.; (12) P. i. imperialis.

286 Australian Entomologist, 2012, 39 (4)

Abbreviations used are: ANIC - Australian National Insect Collection,

CSIRO, Canberra; AM - Australian Museum, Sydney; BMNH - Natura|

History Museum, London; QM - Queensland Museum, Brisbane; Qld -

Queensland; NSW - New South Wales; fwl - forewing length; hwl - hind

wing length.

Phyllodes imperialis meyricki Olliff, 1889, stat. rev.

(Figs 1, 4, 7, 10)

Phyllodes meyricki Olliff, 1889: 114; Hampson, 1913: 392; Seitz 1923 (Vol. 11): 470.

Phyllodes papuana Hampson, 1913: 392; syn. by Seitz 1923 (Vol. 11): 470.

Phyllodes imperialis Druce; Common 1990: 454; Edwards 1996: 307-308; Zborowski

and Edwards 2007: 15.

Types. Lectotype ° (here designated), AUSTRALIA (QUEENSLAND): labelled ‘Mt

Bellenden Ker, Cairns, Q.’, ‘Phyllodes meyricki Oll.’, ‘AM registration number

K351914’; 1 Paralectotype 2, labelled ‘Daintree Riv N. Queensland, Pres. C. French

89.1’, ‘Phyllodes meyricki Oll. Type, AM registration number K183621’ (both in

AM: photographs examined). Olliff (1889) described this taxon from two specimens

in the Australian Museum, which he indicated as from ‘Mt Bellenden-Ker, near

Cairns, and Daintree River, Queensland’, but did not designate a Type. A lectotype is

designated here in order to stabilise the nomenclature, the specimen selected being the

one first mentioned by Olliff (and in better condition).

Other material examined. PAPUA NEW GUINEA: 2 36, 4 99, Kiunga, Lae; 1

labelled ‘Kiunga 65/516, AUST. NAT. INS. COLL’ (NEW GUINEA) (ANIC).

AUSTRALIA (QUEENSLAND): northern Queensland: 14 33, 6 99 (ANIC), 2 3d,

3 99 (Sands); male genitalia slides: 1 labelled ‘NOCT 3 18608 Phyllodes imperialis

17°18'S, 145°35'E, Lake Eacham Q, 24 May 1989 IFB Common’ (ANIC).

Diagnosis. Phyllodes i. meyricki (Figs 1, 4, 7, 10) can be distinguished from

P. i. imperialis (Figs 3, 6, 9, 12) by its narrower forewings with its margins

not so strongly bowed as in the nominotypical subspecies. P. i. meyricki is

overall larger and the forewings longer (44 fwl: 74-78 mm, n = 5; 29: fwl:

75-85 mm, n = 5) than in P. i. smithersi ($8 fwl: 58-69 mm, n = 5; 2° fw:

59-69 mm, n = 6) and the pink hind wing band of P. i. meyricki, although

variable, is not as wide as in P. i. imperialis and extends further towards the

apex than in P. i. smithersi. Phyllodes i. meyricki can be distinguished from

P. i. dealbata from New Caledonia by the presence of white sub-triangular

spots at the vein ends of the hind wing termen, absent in P. i. dealbata.

Distribution. The island of New Guinea. (Indonesian West Papua and

mainland Papua New Guinea) and northern Queensland, Australia (Hunter

1939). In Papua New Guinea recorded from Aroa River (Poole 1989) and

Kiunga (ANIC). In Queensland subspecies P. i. meyricki is recorded from

Mount Bellenden-Ker, Daintree River (Olliff 1889), Atherton, Mutarnee

(ANIC), Paluma, Ingham, Tully, Innisfail (L. Ring), Dunk Island, Kuranda,

Cooktown, Claudie River and Bamaga (unpublished data and ANIC records).

Australian Entomologist, 2012, 39 (4) 287

One specimen from near Proserpine, labelled ‘Airlie Beach, N.Q, Sept. 1976,

A.W. Smith, CGL Gooding Coll.’ (in ANIC) is probably this subspecies but

more specimens are required to confirm its status; none of the food plants of

P. imperialis are known to occur in this region of Queensland.

Larval food plant. The life history and the immature stages of P. i. meyricki

were described by Hunter (1939). The moth breeds in the heavily-shaded

understorey of rainforest, where females oviposit on low, young growth of

the vines Pycnarrhena novoguineensis Miq. (= P. australiana) (Hunter 1939,

Fay 1996) and P. ozantha Diels (Menispermaceae). Eggs are parasitised by

an unidentified Ooencyrtus sp. (Hymeoptera: Encyrtidae).

Phyllodes imperialis smithersi subsp. n.

(Figs 2, 5, 8, 11, 13-15, 16-21)

Phyllodes imperialis ‘southern subspecies’: TSCA 1995; Sands 1999: 386; EPBC

2002 (ANIC Ref. No. 3333), Clark and Spier-Ashcroft 2003: 99; Sands 2012: 38-

39.

Types. Holotype 3, AUSTRALIA: labelled ‘QUEENSLAND. Mary Cairncross Pk nr

Maleny, 26°44'S, 152°52'E, e.p. 27.vi.1988, DPA Sands & R. Broe, larva on Carronia

multisepalea col. 16.iii.1988’ (in ANIC). Paratypes: (QUEENSLAND): 4 3, as

above but e.p. 13.v.1988, col. 9.iii.1988; e.p. 24.i11.1988, col. 16.1i1.1988; e.p.

6.vi.1988, col. 16.iii.1988, ANIC genitalia slide No. 20391 (all ANIC); 5 29, as

above but e.p. 4.vi.1988, col. 9.iii.1988, e.p. 7.vi.1988, col. 9. iii. 1988, e.p. 6.vi.1988,

col. 13.iii.1988, e.p. 14.v.1988, col. 9. iii. 1988 and e.p. 8.vi.1988, col. 5. iv. 1988 (all

ANIC); 1 3, Mary Cairncross Pk nr Maleny, e.p. 22.i.2004, DPA Sands, larva on

Carronia multisepalea, col. 22.xii.2003 (QM); 1 9, Maleny, 26°44'S, 152°52' E, nr

Mary Cairncross Park, to light, 3.xi.2001, A.M. Stabler & A.G. Orr (QM); 1 ĝ,

labelled ‘Phyllodes imperialis Lamington PN, 16.iii.07 Antoine Levegne leg’ (QM);

1 9, Upper Currumbin, 4.vi.32, L. Franzen (ANIC). (NEW SOUTH WALES): 1 9,

Dorrigo National Park, Dorrigo, 18.xi.1990, A.A. Calder (ANIC); 1 3, Dorrigo,

27.iii.73, D.S.P 1. ARCH. (ANIC); 1 9, ANIC Uni. of New England Coll. donated

1983 (ANIC); 1 ĝ, Bellingen Island Reserve, 30°26'55”S, 102°53'47E, egg coll.

13.xi.2006, D. Britton & P Richards, ex pupa 25.i.2007, Australian Museum K243319

(AM); | ĝ, same except ex pupa 26.i.2007, Australian Museum K243460 (AM).

Description. Male (Figs 2, 5). Head, palpus, antenna and thorax light brown;

eyes grey-brown, large, rounded, setae obscure; antenna less than half

forewing length, slender with short fine setae; palpi upturned, segment 2 very

broad, flattened, apically squared, segment 3 short, pencil-like, arising from

proximal edge of segment 2; proboscis long, setae very fine, short, without

serrations. Forewing length (holotype) 66 mm, more than twice width (x 2.1-

2.2), ovate (leaf-like appearance), apex produced, costa convex, sub-apically

convex; termen obliquely curved, inner margin convex basally; above grey-

brown, white area at two thirds length from base at costa, obscure widely-

spaced transverse darker bands reaching costa, from sub-apex to base, a dark

brown median line from below apex to base of M3; termen narrowly dark

brown, a broad paler terminal band, inwardly broadly toothed, extending

288 Australian Entomologist, 2012, 39 (4)

from median line to tornus; subcentral, post-cell reniform near discocellular

vein, shape resembling a leaf miner scar, white edged brown and white, broad

basally, angled and narrow towards costa. Hind wing longer than wide, apex

obtuse, termen convex; above black, costa and base brown, outer margins

with 8 triangular subterminal white spots at vein ends, decreasing in size

from apex to tornus; a central pink band less than half hind wing width,

indented at vein 1A+2A, from M; to inner margin above tornus. Beneath

grey-brown, forewing with central subtriangular grey-black area, from sub-

base to tornus and sub-costa, submarginally-crenate, area with 3 submedian

greyish-white areas between tornus and costa. Hind wing veins M; to M3

dark towards termen, with 2 black median transverse and submedian bands

M; to costa; broad dark grey-brown area to base and inner margin at tornus;

overlain by a broad oval postmedian pink band extending about two thirds

length of wing, from the tornus to M; and reaching Sc, indented at 1A + 2A

and Cu Aj.

Male genitalia (Figs 13-15). Vinculum and tegumen with prominent median

junction, saccus subtriangular in dorsal aspect, tegumen apically

subtriangular, tapered at base of gnathos; gnathos heavily sclerotized, curved

and hook shaped, with apex between apices of valvae clothed in long setae;

valvae broadly subtriangular, dorsal edge straight, apex strongly bowed,

tapered ventrally to a triangular point, ventral margin weakly convex,

ampulla fold with slender, ventrally-directed and pencil-like apically pointed

process; juxta hood shaped with short, apical bifurcate sclerotized flanges;

aedeagus broadly tubular, base of subzonal sheath rounded, apex truncated,

cornuti with median short group of sclerotized cornuti and short apical group

near orifice (vesica retracted); prezonal sheath apically broad, with two, rod-

like sclerites,

Figs 13-15. Phyllodes imperialis smithersi, male genitalia: (13) lateral view; (14)

posterior view; (15) aedeagus. (14-15 slide mounted).

Australian Entomologist, 2012, 39 (4) 289

Female (Figs 8, 11). Colour similar but often darker than male, wings usually

Jonger and broader than male, forewing costa and inner margin more strongly

convex.

Etymology. Named to honour the late Courtenay Smithers.

Variation. The ground colour of both sexes varies from pale grey-brown to

dark reddish brown and is usually paler than in ssp. meyricki. The reniform

mark on the forewing (sensu Hampson 1913) is usually white edged dark

brown, sometimes brown, or rarely obscure in females.

Diagnosis. P. i. smithersi is smaller and the forewings ($4 fwl: 58-69 mm, n

= 5, 2° fwl: 59-69 mm, n = 6) are significantly shorter that in other

subspecies of P. imperialis, including P. i. meyricki ($8 fwl: 74-78 mm, n =

5; 99: fwl: 75-85 mm, n = 5). The pink hind wing band (measured apex -

inner margin) of P. i. smithersi (Figs 2, 8) is less than half the width of the

hind wing (0.34-0.47:1 hwl) and smaller than in P. i. meyricki (0.55-0.62:1

hwl). The termen of the forewings of P. i. smithersi (Figs 2, 5, 8, 11) are

narrower and the inner margins not as strongly convex as in P. i. meyricki

(Figs 1, 4, 7, 10). The male genitalia are smaller than in P. i. meyricki and P.

i. dealbata (Holloway) and the apical cornuti (Fig. 15) are less dense and

proportionally smaller than in P. i. meyricki. Although both subspecies are

variable, differences in the colour, shape and forewing length enable P. i.

smithersi from subtropical eastern Australia to be easily distinguished from

P. i. meyricki from northern Queensland and mainland Papua New Guinea.

Distribution. Queensland: P. i. smithersi occurs at Kin Kin Creek, southeast

from Gympie to the NSW-Qld Border Ranges; it has been observed at

Conondale (E. Weir pers. comm.), Conondale Range and Witta (R. Thomas

pers comm.), Maleny, Blackall Range (R. Broe, N. Clancy, A. Orr),

Bellthorpe, Mount Mee (J. Moss and R. Kendall 2007), Lamington (AM) and

Springbrook (R. Bell pers. comm.). New South Wales: P. i. smithersi larvae

or adults have been photographed or collected at Mount Warning and on the

Border Ranges, Richmond Range, Richmond River (unpublished),

Billinudgel (K. Vale), Dorrigo (ANIC), Rosewood River (Britton 2006),

Bellingen, Huonville (N. Hart) and Bellinger Island (V. Jones, T. Deane pers.

comm.).

Larval food plant: Carronia multisepalea F. Muell. (Menispermaceae)

(Sands 1999, Britton 2006). C. multisepalea is an endemic subtropical vine,

mostly restricted to ‘old growth’ subtropical rainforests on the coast and

ranges (< ca 1,000 m), from Kin Kin Creek, Qld to the Bellinger River,

NSW. The vine frequently occurs in association with another vine,

Pararistolochia praevenosa (Aristolochiaceae), the principal food plant for

the Richmond birdwing butterfly, Ornithoptera richmondia (Gray).

Life history. Since discovery of the life history of P. i. smithersi and its food

plant (Sands 1999), images of the early stages (Figs 16-21) on its food plant

290 Australian Entomologist, 2012, 39 (4)

C. multisepalea, including the remarkable larva, have been published on

several occasions (e.g. Moss 2002, Moss and Kendall 2007, Britton 2006,

Zborowski and Edwards 2007). Adults have been observed feeding on the

damaged fruit of Ficus spp. and a Syzygium sp.

Figs 16-21. Phyllodes imperialis smithersi, immature stages: (16) egg deposited

beneath leaf of Carronia multisepalea; (17) first instar larva beneath leaf of Carronia

multisepalea; (18) third instar larva; (19) fifth instar larva, resting position; (20) fifth

instar larva, alarmed position; (21) pupal case.

Comments

Phyllodes i. imperialis from the Solomon Islands and Bougainville are here

considered con-subspecific. Similarly, P. imperialis from Lae and Kiunga,

Australian Entomologist, 2012, 39 (4) 291

Papua New Guinea and from localities from north of Townsville to the tip of

Cape York Peninsula, northern Queensland, are indistinguishable, confirming

the synonymy of P. i. papuana and P. i. meyricki as proposed by Seitz

(1923). The subtropical subspecies P. imperialis smithersi is confined to

notophyll vine forest where the food plant, C. multisepalea, is an uncommon

endemic vine in eastern Australia, growing on rich volcanic slopes and

riparian or alluvial soils. The leaf-like forewings of most Phyllodes spp.

(Seitz 1923) are similar, with the reniform marking resembling leaf miner

scars (M. Horak pers. comm.).

Acknowledgements

My special thanks to Ted Edwards (CSIRO) for his extensive taxonomic

advice, guidance and encouragement during preparation of this manuscript.

For early biological studies on the moth, I thank Rose Broe, Martina Schotz

and Veronica Brancatini (formerly CSIRO) and for helpful discussions,

photographs and field records, I am grateful to Marianne Horak and the late

Ian Common, Ted Edwards (CSIRO), David Britton (Australian Museum,

Sydney), Paul Grimshaw, Mark Graham, Kath Vale, Noel Hart, Albert Orr,

Vivian Jones, Trevor Deane, Nick Clancy, Robyn Bell, Bob Brown, Les Ring

and Alan Wynn. You Ning kindly prepared the figures for publication and

Susan Sands sketched the male genitalia. Mark Bonner and earlier managers

(Caloundra Municipal Council) are thanked for permission to work at the

Mary Cairncross Scenic Reserve, Maleny. CSIRO studies on this and related

moths (1987-1997) that led to discovery of the new subspecies of P.

imperialis were supported by the Australian Centre for International

Agricultural Research.

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Australian Entomologist, 2012, 39 (4): 293-304 293

FERN FLIES OF AUSTRALIA: THE GENUS TERATOMYZA S.L.

(DIPTERA: TERATOMYZIDAE)

DAVID K. McALPINE

Australian Museum, 6 College Street, Sydney, NSW 2010

Abstract

The broad concept of the genus Teratomyza Malloch (sensu McAlpine and de Keyzer 1994) is

retained so that Vitila McAlpine & de Keyzer, syn. n. and Poecilovitila Papp, syn. n. are no

longer utilised at either generic or subgeneric level. Teratomyza is here provisionally divided

into seven informal species groups. Teratomyza smithersi sp. n. and T. pappi sp. n. (both from

Queensland) are described. The following new generic combinations are made for species

originally described in Poecilovitila Papp: Teratomyza barbata (Papp, 2011), comb. n.;

Teratomyza brevicornis (Papp, 2011), comb. n.; Teratomyza bulbiscapus (Papp, 2011), comb. n.;

Teratomyza defecta (Papp, 2011), comb. n.; Teratomyza elegans (Papp, 2011), comb. n.;

Teratomyza erugata (Papp, 2011), comb. n.; Teratomyza hindustanica (Papp, 2011), comb. n.;

Teratomyza japonica (Papp, 2011), comb. n.; Teratomyza taiwanica (Papp, 2011), comb. n.;

Teratomyza thaii (Papp, 2011), comb. n.; Teratomyza variegata (Papp, 2011), comb. n.

Introduction

The Teratomyzidae or fern flies have a mainly south-temperate distribution

(South America, Australia, New Zealand), but one of the seven here

recognised genera (Teratomyza Malloch s.l.) has a wider distribution,

extending from New Zealand to eastern Asia via Australia and New Guinea.

As the Asian species of the family share a set of apomorphic character states

with certain of the temperate Australasian taxa, I consider that all these

should be included in the apparently monophyletic, though polytypic, genus

Teratomyza, as delimited by McAlpine and de Keyzer (1994).

This paper makes known two undescribed species of Teratomyza of

Queensland. Some additional undescribed species of New Guinea and the

Oriental Region are mentioned below under Teratomyza groups 2-6.

The following abbreviations are used for institutions holding collections:

AM, Australian Museum, Sydney; ANIC, Australian National Insect

Collection, CSIRO, Canberra; BPB, B.P. Bishop Museum, Honolulu; CNC,

Canadian National Collection, Agriculture Canada, Ottawa; HELS,

Zoological Museum, University of Helsinki, Helsinki; QM, Queensland

Museum, Brisbane.

Notes on morphology

The nomenclature of the bristles (enlarged and individually differentiated

macrotrichia) on the upper back of the head in schizophoran flies has been

confused in the literature. I have attempted (McAlpine 2007) more accurately

to systematise the identification and terminology of these bristles, and this

system is here relevant because of taxonomic variation in the bristle pattern

in Teratomyza s.l. The postvertical bristles are reduced in the Teratomyzidae

and are often scarcely differentiated from the nearby setulae (small

macrotrichia of irregular placement). In Teratomyza group 1 (including T.

294 Australian Entomologist, 2012, 39 (4)

neozelandica Malloch and perhaps one or two closely similar species), the

series of postocular setulae terminates dorsomedially below the level of the

outer vertical bristle. Widely separated from that series and located behind

and slightly mesad of the inner vertical bristle (Fig. 2), is the bristle I identify

as the paravertical bristle, as it corresponds in position to that bristle in other

families (e.g. Heteromyzidae and Pseudopomyzidae). In other species groups

(e.g. group 7, Fig. 3), there is no such isolated paravertical bristle and the

series of postocular setulae often extends further mesad.

In some teratomyzid taxa (numerous examples figured by McAlpine and de

Keyzer 1994 and Papp 2011), some of the macrotrichia on the surstylus of

the male are short, very stout, and peg-like or tooth-like (see Figs 5, 6, 8). I

refer to these as blunt spinules, to distinguish them from the slender, fine-

tipped setulae, often also present on the surstylus.

Genus Teratomyza Malloch

Teratomyza Malloch, 1933: 113-114. Type species T. neozelandica Malloch (original

designation).

Vitila McAlpine & de Keyzer, 1994: 321 (as subgenus of Teratomyza). Type species

T. (Vitila) undulata McAlpine & de Keyzer (original designation). Syn. n.

Poecilovitila Papp, 2011: 11. Type species P. elegans Papp (original designation).

Syn. n.

Diagnostic description. Head. Postfrons not setulose anteriorly; cheek region

commonly with upper and lower series of setulae; palpus absent or very

minute. Thorax. Dorsocentral bristles two pairs; mesopleural (anepisternal)

and pteropleural (anepimeral) bristles absent; costa distally with variably

reduced setiferous tubercles; anal cell (cup) not strongly enclosed on

posterior side; alular lobe and alular incision absent; fringe of alular setulae

reduced or absent. Male postabdomen. Aedeagus prominent, somewhat

elongate, asymmetrical. Female abdomen. Tergite 7 and sternite 7 broadly

fused, annular.

Notes. The genus Teratomyza sensu McAlpine and de Keyzer (1994) has

been divided into three genera by Papp (2011), viz. Teratomyza (s.str.), Vitila

and Poecilovitila. My own studies indicate that these categories are less well

defined than indicated by Papp and that his restricted genus Teratomyza, still

including both Asian and New Zealand species and defined only by

plesiomorphic character states, is very probably paraphyletic. I find

significant morphological differences between the available Asian and New

Zealand taxa (the latter including the type species), but both categories lack

the obvious apomorphic wing features that characterise his so-called genera

Vitila and Poecilovitila. Several taxa not considered by Papp possess

character combinations which tend to link his major groupings, but some are

still very incompletely known. I consider that the apparently monophyletic

genus Teratomyza s.l. (e.g. Fig. 1) is best divided into seven informal species

Australian Entomologist, 2012, 39 (4) 3 295

groups on the basis of present knowledge, and that such formal categories as

subgenera should be avoided.

Fig. 1. Teratomyza smithersi sp. n., male from Upper Tully River.

Key to species groups of Teratomyza s.l.

1 Paravertical bristle usually present behind and slightly mesad of inner

vertical bristle and widely separated from largest postocular setula, latter

located behind but not mesad of outer vertical bristle (Fig. 2); male:

surstylus arising well within border of epandrium; cercus vestigial or

apparently absent; New Zealand ..................ccccececeneeeeseeeeees group |

296 Australian Entomologist, 2012, 39 (4)

- Paravertical bristle absent; largest postocular setula located at least

slightly laterad of inner vertical bristle and not widely separated from rest

of postocular series (Fig. 3); male (where known): surstylus articulated

with lateral margin of epandrium; cercus well developed, setulose; not in

NewZealand fr: ees re Sar ee ye LE rare ete eer nena ee en niet

2 Postfrons with longitudinal stripes; wing membrane with four or five

white spots between costa and vein 2; sternopleural bristles two, large;

CASTEINFAUStral 1a eee emte mene teen fetes ements E teers group 7

- Postfrons without longitudinal stripes; wing without series of white spots

between costa and vein 2; only one sternopleural bristle well developed;

notin A ustraliac. oc. fect oe A E E ete RNC E Cates

3 Vein 2 near its mid-length closely approximated to costa, but usually

diverging from costa again before distal termination .....................65 4

- Vein 2 not closely approximated to costa and then diverging before distal

Terminationys. 2 ies. ells see I ae eee rts teres erect cet ears:

4 Anterior crossvein (r-m) at least twice as long as discal crossvein (dm-

cu); basal section of vein 4 strongly arched, making second basal cell

much broader than first basal cell in this region; New Guinea ..... group 6

- Anterior crossvein not much longer than discal crossvein; basal section of

vein 4 not arched, thus second basal cell not broader than first basal cell;

Oriental and eastern Palaearctic Regions ..............0.ccceseeeeseeeseee ences 5

5 Wing membrane with some brownish shading and usually a hyaline spot

in submarginal cell beyond end of vein 2; vein 5 with complex curvature

(sometimes slight) immediately beyond discal crossvein (Papp 2011: figs

112-121); transverse ridge of vertex strongly elevated laterally so that, in

exact profile, outer vertical bristle arises much above upper margin of eye

group 5

- Wing membrane quite clear; vein 5 without such complex curvature;

transverse ridge of vertex relatively little elevated laterally, outer vertical

bristle arising little above upper margin of eye ....................084 group 4

6 Ocellar bristle small, not over half as long as fronto-orbital bristle; eye

markedly longer than high; vein 2 rather strongly curved; thoracic pleura

with dark brown longitudinal stripe; New Guinea ................... group 3

- Probably not agreeing in above characters (only limited material and

limited published information available); Oriental and eastern Palaearctic

REBiONS aa a E a aeo rece e terete R tarsi group 2

Teratomyza group 1

Included species: Teratomyza neozelandica Malloch; possibly one or more

closely related undescribed species.

Australian Entomologist, 2012, 39 (4) 297

Distribution: New Zealand (North and South Islands).

Notes. This group is not adequately studied, but my observations suggest that

it can be distinguished from other groups by the characters given in the key.

Of the 29 specimens examined, only one lacks the paraverticals shown in Fig.

2. The surstylus lacks blunt spinules.

Figs 2-3. Vertex of head, posterior view of: (2) Teratomyza nr neozelandica (group

1); (3) Teratomyza smithersi sp. n. (group 7). iv = inner vertical bristle, oc = ocellar

bristle, ov = outer vertical bristle, pa = paravertical bristle, po = postocular setulae, pv

= postvertical bristle.

Teratomyza group 2

Included species: Teratomyza chinica Yang Chikun; T. formosana Papp; T.

sp. undescr. (Nepal, CNC).

Distribution: Oriental Region—Vietnam, Taiwan, China, Nepal.

Notes. Representatives of this group have principally been made known by

Papp (2011), who described and figured the male postabdominal structure for

298 Australian Entomologist, 2012, 39 (4)

two species. The one species available to me (Nepal, CNC) has the

mesoscutum anteriorly simply convex, no dark pleural stripe on the thorax,

the postfrons very densely pruinescent, all in contrast to the condition in

group 3. This species has few large terminal blunt spinules on the slender

surstylus, in contrast to the two species treated by Papp.

Teratomyza group 3

Included species: Teratomyza sp. undescr. (Myola, Oro Province, AM).

Distribution: New Guinea (only yet known from Owen Stanley Range, Oro

Province, Papua New Guinea).

Notes. The specimens (two females) show the distinctive characters given in

the key to groups, but, until males become known, an adequate review of

relationships is not possible. The two female specimens, collected at the same

locality by J.W. Ismay, show slight differences in wing venation and

chaetotaxy, and there is a slight possibility that they are not conspecific. In

any case, the species cannot be accurately characterised without knowledge

of the male postabdomen.

Teratomyza group 4

Included species: Teratomyza sp. undescr. (Nepal, CNC).

Distribution: Oriental Region—Nepal.

Notes. The group is known to me from one male specimen. It resembles some

species of group 5 (‘Poecilovitila’) in having vein 2 on a substantial part of

its length thickened and closely approximated to the costa. Otherwise it is

without the apomorphic wing conditions of that group and superficially

resembles examples of groups 1 and 2.

Teratomyza group 5

Included species: Teratomyza barbata (Papp, 2011) comb. n.; Teratomyza

brevicornis (Papp, 2011) comb. n.; Teratomyza bulbiscapus (Papp, 2011)

comb. n.; Teratomyza defecta (Papp, 2011) comb. n.; Teratomyza elegans

(Papp, 2011) comb. n.; Teratomyza erugata (Papp, 2011) comb. n.;

Teratomyza hindustanica (Papp, 2011) comb. n.; Teratomyza japonica (Papp,

2011) comb. n.; Teratomyza taiwanica (Papp, 2011) comb. n.; Teratomyza

thaii (Papp, 2011) comb. n.; Teratomyza variegata (Papp, 2011) comb. n.;

Teratomyza sp. undescr. (NE Burma, HELS); Teratomyza sp. undescr.

(Fukien Province, S. China, BPB); Teratomyza sp. undescr. (Luzon,

Philippines, BPB); Teratomyza sp. undescr. (Sabah (North Borneo),

Malaysia, CNC). A few other specimens could represent additional Oriental

species.

Distribution: Oriental Region and adjacent parts of Palaearctic Region—

Japan, China, Taiwan, Nepal, India, Philippines, Vietnam, Burma, Thailand,

Malaysia.

Australian Entomologist, 2012, 39 (4) 299

Notes. Group 5 is apparently the largest group in the genus, with at least 15

species, but none is known from east of Wallace’s Line. It appears from

Papp’s study that all species probably have a well developed series of blunt

spinules on the distal margin of the surstylus, a feature that is shared with

group 7.

This group includes the type species of Poecilovitila Papp.

Teratomyza group 6

Included species: Teratomyza sp. undescr. (Papua New Guinea: mainland).

Distribution: New Guinea.

Notes. The one included species resembles those of groups 4, 5, and 7 in

having vein 2 undulate, thickened, and closely approximated to the costa well

before its distal end, but differs from these in its own specialised wing

features as indicated in the key. A much reduced series of blunt spinules is

present on the surstylus. None of these three other groups is known from

New Guinea, but the smaller flies of that island are still very incompletely

known.

Teratomyza group 7

Included species: Teratomyza undulata McAlpine & de Keyzer; Teratomyza

smithersi sp. n., Teratomyza pappi sp. n.

Distribution: eastern Australia, including Tasmania.

Notes. Group 7 is easily distinguished by the presence of two or three

undulations in vein 2, presence of many white spots on the wing membrane

including four or five between the costa and vein 2, and the presence of two

large sternopleural bristles.

This group includes the type species of subgenus Vitila McAlpine & de

Keyzer, Teratomyza undulata McAlpine & de Keyzer.

Key to species of Teratomyza group 7

1 Anterior and discal crossveins separated by not more than length of discal

crossvein, or these two crossveins contiguous; first posterior cell (behind

distal section of vein 3) with more than eight white spots; area behind

vein 6 with two white spots; male: surstylus as in Fig. 6 ...............068

AE. E E IA EAT IARA ET! undulata McAlpine & de Keyzer

- Anterior and discal crossveins separated by at least twice length of discal

crossvein; first posterior cell with fewer than eight white spots; only one

white spot behind vein 6; male: surstylus differently shaped

2 Vein 2 usually with three marked undulations (Fig. 4); usually five large

white spots present between costa and vein 2; submarginal cell (behind

distal section of vein 2) usually with seven to nine white spots; male:

surstylus narrowed beyond base, so that spinulose distal margin is not

300 Australian Entomologist, 2012, 39 (4)

more than half as long as basal margin (Fig. 5); cercus more than half as

longiasisurstylusmersseeesceetis beet E meets nets en since es smithersi sp. n.

- Vein 2 with two marked undulations and sometimes an indistinct one

distally (Fig. 7); usually four white spots present between costa and vein

2, or sometimes a smaller additional spot distally; submarginal cell

usually with six white spots; male: surstylus somewhat narrowed towards

mid-length, but expanded again distally, so that spinulose distal margin is

as long as basal margin (Fig. 8); cercus not more than half as long as

SUISTY US Peete eee ee ee ee nee TATAA EL crt rt: pappi sp. n.

Teratomyza undulata McAlpine & de Keyzer

(Fig. 6)

Teratomyza (Vitila) undulata McAlpine & de Keyzer, 1994: 321-324, figs 36-43.

Vitila undulata (McAlpine & de Keyzer), —Papp 2011: 10.

Description. See McAlpine and de Keyzer (1994).

Distribution. Mainly higher rainfall areas from Eungella (Dalrymple Heights)

district, Queensland, to eastern Victoria and western Tasmania. See

McAlpine and de Keyzer for details.

Teratomyza smithersi sp. n.

(Figs 1, 3-5)

Types. Holotype 6, QUEENSLAND: Crawford’s Lookout, Palmerston Highway,

17.61°S 145.79°E, 8.viii.2010, S.F. McEvey, J. Weiner (AM, K300569, S.McE

28392). Paratypes. QUEENSLAND: 2 3, 1 9, Crawford’s Lookout, Palmerston

Highway (AM); 1 4, Mossman Gorge, April 1967, D.H. Colless (ANIC); 1 2, 5-8 km

on Mount Lewis Road off Mossman-Mount Molloy Road, April 1967, D.H. Colless

(ANIC); 2 9,3 km NNE of Julatten, Sept. 1980, D.H. Colless (ANIC); 1 2, Kuranda

Range State Forest, Black Mountain Road, April 1967, D.H. Colless (ANIC); 1 9,

Kuranda, May 1958, D.K. McAlpine (AM); 1 ĝ, Barron Falls, near Kuranda, May

1958, D.K. McAlpine (AM); 4 3, 2 2, Mount Edith, 4-7 miles (c. 6-11 km) off

Danbulla Road, April 1967, D.H. Colless (ANIC); 6 3, 4 2, Mount Edith Forest road,

1-1.5 miles off Danbulla Road, May 1967, D.H. Colless (ANIC); 1 2, Wongabel

State Forest, near Atherton, May 1967, D.H. Colless (ANIC); 2 3,3 9, The Boulders,

near Babinda, May 1967, July 1971, D.H. Colless, Z.R. Liepa (ANIC); 15 4, 18 9,

The Crater [Ringrose National Park or Mount Hypipamee], Jan, May, Dec. 1961-

1972, D.H. Colless, G.A. Holloway, D.K. McAlpine (AM, ANIC, QM); 1 9,

Palmerston National Park, 23 km NE of Ravenshoe, Nov. 1981, D.H. Colless (ANIC);

1 ĝ, summit, Walter Hill Range, Cardstone-Ravenshoe Road, Jan. 1967, D.K.

McAlpine, G.A. Holloway (AM); 1 3, upper Tully River valley, 17.77°S 145.65°E,

Aug. 2010, S.F. McEvey (AM); 1 4, Kirrama rain forest [Kirrama Range vicinity

between Tully and Herbert Rivers], Aug. 1976, I.R. Bock, P.A. Parsons (ANIC).

Other material. QUEENSLAND: Paluma vicinity, including Birthday Creek (AM,

ANIC).

Australian Entomologist, 2012, 39 (4) 301

Figs 4-6. Teratomyza spp.: (4) T. smithersi sp. n., wing; (5) T. smithersi, left lateral

view of epandrium and associated structures, setulae on epandrium omitted; (6) T.

undulata, right surstylus, medial view. Scale for Fig. 5 = 0.1 mm. c = cercus, ep =

epandrium, ss = surstylus, v2-v6 = veins 2-6.

Description (male, female). A small slender fly, generally resembling the

well known T. undulata (see McAlpine and de Keyzer 1994 for detail).

Coloration. Head pale tawny-yellow to creamy; postfrons with three broad

dark brown to blackish longitudinal stripes on its full length, with whitish

302 Australian Entomologist, 2012, 39 (4)

intermediate zones and lateral margins; face, cheeks, and occiput without

darker markings. Antenna grey-brown. Thorax pale tawny, with paler mostly

creamy pleura, often slightly darker towards upper margin or with distinct

brown upper marginal stripe; scutellum usually yellow at extreme apex. Wing

membrane grey-brown, darker anteriorly, with numerous white spots, fewer

than in T. undulata; marginal cell with five large white spots between costa

and vein 2, and a smaller pale zone in extreme base; submarginal cell usually

with eight or nine white spots. Halter tawny-yellow to grey-brown. Legs

yellowish; femora sometimes slightly darker apically. Abdominal tergites

tawny-brown, in male usually without markings except for yellow lateral

zones on tergite 6, in female often with yellow lateral zones on most tergites.

Head and thorax structurally similar to those of 7. undulata. Wing: vein 2

with three distinct undulations on which it is thickened and approximated to

costa; vein 6 with single undulation or region of curvature; anterior and discal

crossveins separated by at least twice length of discal crossvein.

Male postabdomen. Surstylus rather broad basally, much narrowed beyond

base, so that spinulose distal margin is much shorter than basal margin

articulating with hypandrium; pregonite broadly ovate, larger than surstylus;

cercus rather large and stout, with long and short setulae.

Dimensions. Total length, ¢ 1.7-1.9 mm, Q 1.9-2.2 mm; length of thorax, 4

0.74-0.78 mm, 2 0.93-1.00 mm; length of wing, 3 2.2-2.4 mm, 2 2.8-2.9

mm.

Etymology. The name refers to my late friend Courtenay N. Smithers,

formerly of the Australian Museum.

Distribution. Tropical coast of Queensland between 16°S and 19°30’S. It is

thus apparently geographically isolated from populations of 7. undulata but

sympatric with 7. pappi.

Notes. Teratomyza smithersi resembles T. undulata but differs in characters

of wing venation, as indicated in the above key, and in the shape of the

surstylus and pregonite. For comparison with 7. pappi see under that species.

Teratomyza pappi sp. n.

(Figs 7-9)

Types. Holotype 6, QUEENSLAND: Ringrose National Park [The Crater or Mount

Hypipamee], 5.v.1967, D.H. Colless (ANIC). Paratypes. QUEENSLAND: 1 ĝ, same

data as holotype (AM); 1 3, 5-8 km up Mount Lewis Road, from Mossman-Mount

Molloy Road, April 1967, D.H. Colless (ANIC); 3 ĝ, Mount Edith, 4-7 km off

Danbulla Road, Atherton district, April 1967, D.H. Colless (ANIC).

Australian Entomologist, 2012, 39 (4)

303

Figs 7-9. Teratomyza pappi sp. n.: (7) wing; (8) right lateral view of epandrium and

associated structures, setulae on epandrium omitted; (9) right pregonite and

epiphallus. Scale for Figs 8 and 9 = 0.1 mm. el = epiphallus, pg = pregonite.

Description (male only known). A small fly closely resembling T. undulata

and T. smithersi, agreeing with description of the latter, except as indicated

below.

Coloration. Head and thorax as in T. smithersi; thoracic pleura with definite

brown upper marginal stripe. Wing pattern resembling that of T. smithersi;

usually only four white spots between costa and vein 2 (one specimen

showing a smaller distal additional white spot); submarginal cell with six

304 Australian Entomologist, 2012, 39 (4)

white spots; only one pale spot between vein 6 and wing margin. Halter with

dark grey capitellum (mature specimens).

Head and thorax structurally similar to those of T. undulata and T. smithersi.

Wing: vein 2 with two marked undulations, thickened along their summits,

and at most a trace of more distal undulation.

Male postabdomen. Surstylus narrowed towards mid-length, but expanded

distally so that distal margin is as long as basal articulating margin and

posterodistal angle is subacute; pregonite much shorter than surstylus,

slightly narrowed basally, broadly rounded distally; cercus much smaller than

in T, smithersi.

Dimensions. Total length, 1.8-1.9 mm; length of thorax, 0.75-0.80 mm;

length of wing, 2.2-2.3 mm.

Etymology. The name refers to László Papp, who has made the major

contribution to knowledge of Oriental Teratomyzidae.

Distribution. Northern Queensland—Atherton to Mossman districts. The

distribution is included within that of 7. smithersi.

Notes. Teratomyza pappi is very similar to T. smithersi but is distinguished

by the contour of vein 2, the distribution of pale wing spots and, in the male,

by details of the surstylus, pregonite and cercus.

Acknowledgements

Donald Colless, John Ismay, Shane McEvey, Richard Vockeroth and Jean

Weiner supplied significant material. Helen Smith reviewed the manuscript

and aided in its preparation. Scott Ginn provided the colour photograph.

References

MALLOCH, J.R. 1933. A remarkable anthomyzid from New Zealand (Diptera). Stylops 2: 113-

114.

McALPINE, D.K. 2007. The surge flies (Diptera: Canacidae: Zaleinae) of Australasia and notes

on tethinid-canacid morphology and relationships. Records of the Australian Museum 59: 27-64.

McALPINE, D.K. and DE KEYZER, R.G. 1994. Generic classification of the fern flies (Diptera:

Teratomyzidae) with a larval description. Systematic Entomology 19: 305-326.

PAPP, L. 2011. Oriental Teratomyzidae (Diptera: Schizophora). Zootaxa 2916: 1-34.

Australian Entomologist, 2012, 39 (4): 305-320 305

SYSTEMATIC AND DISTRIBUTIONAL NOTES ON SOME

AUSTRALASIAN AND AFRICAN SPECIES OF PLATENSINA

ENDERLEIN AND DICHENIOTES MUNRO (DIPTERA:

TEPHRITIDAE: TEPHRITINAE), WITH DESCRIPTION OF A NEW

SPECIES OF DICHENIOTES FROM KENYA

DAVID L. HANCOCK

8/3 McPherson Close, Edge Hill, Cairns, Old 4870

Abstract

The tephritine genera Platensina Enderlein and Dicheniotes Munro are discussed, with several

new distribution records and keys to all known species provided. The Australian Platensina

trimaculata Hardy & Drew and SE Asian P. qguadrula Hardy are returned to Platensina; the East

Asian P. assimilis (Shiraki), comb. n. and P. shirouzui (Ito), comb. n. are transferred from

Bezzina Munro; P. voneda (Walker) is placed as a new synonym of P. acrostacta (Wiedemann)

and its type locality presumed to be Bengal, India; P. fukienica Hering is placed as a new

synonym of P. tetrica Hering; P. platyptera Hendel, stat. rev. (= P. malaita Curran, syn. n.; = P.

dubia Malloch, syn. n.) and P. dilatata Hering are removed from synonymy with P. amplipennis

(Walker), with P. dilatata placed as a new synonym of P. ampla de Meijere; a record of

‘Pliomelaena sp. B’ from Papua New Guinea and all records of P. amplipennis from the

Australian Region are referred to P. platyptera. The primarily Afrotropical Dicheniotes aeneus

(Munro), D. alexina (Munro), D. asmarensis (Munro), D. enzoria (Munro), D. parviguttatus

(Hering), D. sokotrensis (Hering) and D. ternarius (Loew) are transferred as new combinations

from Pediapelta Munro. Dicheniotes kakamegae sp. n. is described from western Kenya.

Introduction

Hancock (2001) placed the tephritine genus Bezzina Munro in the Platensina

group of genera, within the tribe Dithrycini, subtribe Platensinina (=

Oedaspidina), a distinctive assemblage of flies known to form stem galls on

various species of Asteraceae, Goodeniaceae and Onagraceae. However, a

molecular investigation by Han et al. (2010 and pers. comm.) has indicated

that the type species of Bezzina, the Afrotropical B. margaritifera (Bezzi),

appears to be much more closely related to Chipingomyia manica Hancock

(provisionally referred to the Campiglossa group in tribe Tephritini by

Hancock 2006) than to platensinines such as Oedaspis Loew and Platensina

Enderlein. Accordingly, the four non-African species assigned to Bezzina by

Hancock (2001) are currently misplaced. This error is corrected below and

notes on several other Platensina species are included, including the removal

of all current synonyms of P. amplipennis (Walker) to other species.

Hancock (2010) noted that all except the type species currently included in

the African genus Pediapelta Munro (with one record from SE Queensland),

in tribe Tephrellini, appeared to be better placed in Dicheniotes Munro, a

possibility initially suggested by Munro (1947) when describing Pediapelta.

Examination of most of the included species has supported this suggestion.

Abbreviations: AQIS - Australian Quarantine and Inspection Service, Cairns;

BMNH - Natural History Museum, London; OUMNH - Oxford University

Museum of Natural History, Oxford.

306 Australian Entomologist, 2012, 39 (4)

This paper is dedicated to the memory of Courtenay N. Smithers who, like

the present writer, experienced the pleasures of working in both Africa and

Australia. We both worked, at different times, on tsetse flies and plant pests

for the then Rhodesian Departments of Veterinary Services (Tsetse &

Trypanosomiasis Control Branch) and Research and Specialist Services

(Plant Protection Research Institute) before becoming Museum curators.

Systematic and distributional notes

Tribe Dithrycini (subtribe Platensinina)

Platensina Enderlein, 1911

Reevaluation of the morphology and relationships of the four non-African

species included in Bezzina by Hancock (2001) suggests that they properly

belong in Platensina, even though their wings are not as broad as is usual in

that genus. In all species the scutum is densely greyish pubescent and covered

with flattened, subrecumbent, yellow-white setulae, the apical scutellar setae

are about half the length of the basal pair and the hyaline indentations in wing

cell m are all short, as in typical species of Platensina. They differ from

Collessomyia Hardy & Drew in the frequent presence of a small, marginal

hyaline indentation in cell rə and in a non-elongate glans lacking a long,

flagellum-like and microsetose apical rod. Accordingly, these four species are

returned or newly transferred to Platensina, as P. assimilis (Shiraki), comb.

n., P. quadrula Hardy, stat. rev., P. shirouzui (Ito), comb. n. and P.

trimaculata Hardy & Drew, stat. rev. Platensina amita Hardy, from Luzon

(Philippines), also has a relatively narrow wing and is possibly related.

Platensina fukienica Hering, described from Fujian Province, China (Hering

1939b) is treated here as a new synonym of P. tetrica Hering, described from

Tamil Nadu, India (Hering 1939a), based on examination of a paratype

female and recently collected male of P. fukienica from Fujian (in BMNH) in

comparison with material of P. tetrica from West Malaysia (also in BMNH).

Trypeta voneda Walker, first placed in Platensina by Norrbom et al. (1999),

is treated here as a new synonym of P. acrostacta (Wiedemann), based on

examination of the lectotype female (Fig. 1) and a paralectotype female in

BMNH. Its stated type locality of ‘Bahia, Brazil’ was regarded as possibly

erroneous by Foote (1964) and that is certainly the case. The type labels bear

the data ‘Brazil, Bahia, ?Collector’. The true type locality is likely to be

‘Bengal, India’, which is also the type locality of Trypeta stella Walker,

another synonym of P. acrostacta described at the same time (Walker 1849),

The small hyaline spot near the apex of cell rı present in the lectotype of P.

voneda occasionally occurs, on one or both wings, in other specimens of P.

acrostacta from India and Sri Lanka.

Platensina dilatata Hering, P. dubia Malloch, P. malaita Curran and P.

platyptera Hendel are removed from synonymy with P. amplipennis

(Walker) (Fig. 2). These taxa are discussed below.

Australian Entomologist, 2012, 39 (4) 307

The only known host record for Platensina is of P. acrostacta from stem

galls on Ludwigia (= Jussiaea) (Onagraceae) in southern India (Hardy 1973;

specimens in BMNH: 2 99, Kodaguhalli [Kodihalli, Bangalore], 7.v.1963,

larvae causing galls on Jussiaea sp.).

Figs 1-6. Platensina spp., wings: (1) Lectotype female of P. voneda, a synonym of P.

acrostacta; (2) female of P. amplipennis from West Java; (3-4) P. ampla: (3) female

from Papua New Guinea; (4) male from Solomon Islands; (5-6) P. platyptera: (5)

Natural History Museum, London.

Platensina ampla de Meijere (Figs 3-4)

Platensina ampla de Meijere, 1914: 217. Type localities Batavia [Jakarta] and

Semarang, Java, Indonesia.

Platensina dilatata Hering, 1941b: 63, fig. 11; syn. n. Type locality Stephansort

[Bogadjim], Astrolabe Bay, Papua New Guinea.

Material examined. PAPUA NEW GUINEA: 1 Q, Laloki, Central Province,

23.iii.1986, J.W. Ismay (BMNH). SOLOMON ISLANDS: 1 ĝ, British Solomons,

1.1933, R.J.A.W. Lever (BMNH).

308 Australian Entomologist, 2012, 39 (4)

Platensina dilatata, described from Astrolabe Bay in Papua New Guinea

(Hering 1941b), is removed from synonymy with P. amplipennis and placed

as a new synonym of P. ampla. This species is distinguished by the presence

of two hyaline marginal indentations in cell r2;3 and isolation of the hyaline

discal spots in that cell. Newly recorded from Solomon Islands.

Platensina amplipennis (Walker) (Fig. 2)

Trypeta amplipennis Walker, 1860: 159. Type locality Makassar, Sulawesi.

Material examined. INDONESIA (SULAWESI): Lectotype 92, Macassar, Celebes,

W.W. Saunders, B.M. 1868-4 (BMNH). INDONESIA (JAVA): 1 9, Preanger,

Wynkoops Bay, West Java, iii.1935 (BMNH). MALAYSIA (WEST): 1 9, Wang

Tangga, Perlis, 18.iii.1936, ex FMS Museum (BMNH).

Most records of P. amplipennis from countries other than Indonesia and

Malaysia (including Australia) belong to P. platyptera; others (e.g. Hardy

1973) require confirmation. Length of the apical scutellar setae and wing

characters, particularly the shape and orientation of the hyaline indentations

in cells r+ r2, and m (c.f. Figs 2-6), separate it from similar species.

Platensina euryptera (Bezzi)

Tephritis euryptera Bezzi, 1913: 162. Type locality Tenasserim, Burma.

Platensina extincta Hering, 1952: 47, fig. 4. Type locality Wai Lekabe, Baing, east

Sumba I., Indonesia. Synonymy by Hardy 1988.

Material examined. VIETNAM: 1 ĝ, Indo-China, R.V. de Salvaza, 1918-1 (BMNH).

Newly recorded from Vietnam.

Platensina platyptera Hendel, stat. rev. (Figs 5-6)

Platensina platyptera Hendel, 1915: 461. Type locality Taihorin, Taiwan.

Platensina malaita Curran, 1936: 29, pl. 1; syn. n. Type locality Tai Lagoon, Malaita,

Solomon Islands.

Platensina dubia Malloch, 1939: 459; syn. n. Type locality Gordonvale, Qld,

Australia.

Platensina amplipennis: authors, nec Walker, 1860. Misidentifications.

Material examined. AUSTRALIA (QUEENSLAND): 1 2, Warnambool St, Trinity

Park, Cairns, 16°48'S 145°42'E, 28.iv.2010, J. Olive (AQIS). VANUATU: 1 9,

Nombur, Gaua, Santa Maria I., Banks Is, 15.x.1922, T.T. Barnard (BMNH); 1 ĝ,

native garden near Hog Harbour, Elephant I., Espiritu Santo, 0-50', 17.iv.1927, J.R.

Baker & Percy Sladen (OUMNH). SOLOMON ISLANDS: | ĝ, Solomon Is, xi.1932,

R.A. Lever (BMNH); 2 34, Lingatu, Russel I., 26.viii.1936, R.A. Lever (BMNH).

INDONESIA (FLORES): 1 4, Wae Rana, W. Flores, 26.i.1927 (BMNH).

MALAYSIA (SARAWAK): 1 9, R. Kapah trib. of R. Tinjah, 5.x.1932, undergrowth,

B.M. Hobby & A.W. Moore, Oxford Univ. Expd. (BMNH). BURMA: 1 9, Rangoon,

23.xii.[19]04-3.i.[19]05, Brunetti (BMNH).

Platensina platyptera, described from Taiwan (Hendel 1915), is also

removed from synonymy with P. amplipennis, from which it differs in wing

pattern characters and the shorter and weaker apical scutellar setae (about

Australian Entomologist, 2012, 39 (4) 309

a quarter length of basals, rather than half). P. platyptera closely resembles P.

zodiacalis (Bezzi) and, like that species, is widespread; however, P.

zodiacalis lacks apical scutellar setae.

Hardy (1954) also recorded this species from Espiritu Santo (as P. malaita)

and Hering (1941a) previously recorded the Flores specimen. It also occurs at

Tapini in Papua New Guinea (Hardy 1988, as Pliomelaena sp. B), Andaman

Islands, India (K.J. David pers. comm., photograph examined) and Ryukyu

Islands, Japan (Wang 1998, as P. amplipennis). Records from Thailand,

Laos, Vietnam and Micronesia (Hardy 1973) probably also belong here but

confirmation is required; his illustration is of P. amplipennis. Illustrations in

Hardy and Drew (1996) are also of P. amplipennis and the Trinity Park

female (Fig. 6) appears to be the first Australian specimen illustrated.

Platensina zodiacalis (Bezzi)

Tephritis zodiacalis Bezzi, 1913: 163. Type locality Calcutta, India.

Material examined. INDIA: 1 9, ex Brunetti (BMNH). NEPAL: E. shore of R. Arun

below Tumlingtar, Arun Valley, c1800', 23.xii.1961, swept from Ricinus communis L.

(BMNH). BURMA: 1 ĝ, Rangoon, 23.xii.[19]04-3.i.[19]05, Brunetti (BMNH).

CHINA: 1 3, Xishuangbanna, Yunnan, 650 m, 6.iv.1958, L.Y. Zhang & S.P. Hong

(BMNH). THAILAND: 1 ĝ, Sathorn Rd, [Bangkok], 26.xi.1933, W.R.S. Ladell

(BMNH). SINGAPORE: 1 3, Singapore, H.N. Ridley, 99-126 (BMNH).

Newly recorded from Burma and Singapore.

Tribe Tephrellini

Dicheniotes Munro, 1938

Examination of material in BMNH has confirmed the view that all except the

type species of Pediapelta, the South African Pediapelta spadicescens Munro

[3 females from Katberg examined], should be transferred to Dicheniotes; it

differs from all other species included in Pediapelta by Munro (1947) and

Hancock et al. (2003) in significant wing pattern characters (wing with base

largely infuscated including middle of cell c, not with base and middle of cell

c broadly hyaline; R-M crossvein aligned with middle of basal hyaline

indentation across cell rı, not between the two; the large hyaline spot in cell

r45 lies on line of outer hyaline indentation across cell r; and before, not

beyond, line of DM-Cu crossvein), head shape (lower occiput distinctly

swollen) and a larger, more robust body. In addition, cell dm is with or

without a subapical spot placed just beyond the line of R-M crossvein and the

postpronotal lobes are dark fulvous with a fuscous tint to entirely fuscous.

The affinities of P. spadicescens are uncertain but the dark band in cell c and

the position of the hyaline spot in cell r4;5 suggest it belongs in tribe

Tephritini; unfortunately only females have been recorded.

All other species are referable to Dicheniotes, considered here to comprise 19

species, including the following seven new combinations, all transferred from

Pediapelta [all species examined]: D. aeneus (Munro), D. alexina (Munro),

310 Australian Entomologist, 2012, 39 (4)

D. asmarensis (Munro), D. enzoria (Munro), D. parviguttatus (Hering), D.

sokotrensis (Hering) and D. ternarius (Loew). One new species is described.

The pale thoracic pubescence or ‘dust’ varies from fine and sparse to coarse

and relatively dense; the postocular setae also vary from black to reddish-

brown, yellowish or creamy-white (often mixed). In the examined material,

the ‘dust’ appears coarsest and the postocular setae palest in D. parviguttatus

and D. sokotrensis. Such variation, particularly in the colour of the postocular

setae, also occurs in other tephrelline genera such as Metasphenisca Hendel

and Pristaciura Hendel. Dicheniotes dispar (Bezzi) has been bred from

flowers of Becium obovatum (Lamiaceae) and others found associated with,

but not bred from, Ocimum suave (Lamiaceae) (Munro 1947).

Fig. 7. Dicheniotes kakamegae sp. n., wing of holotype female. Photo by K. Goodger

Dicheniotes kakamegae sp. n. (Fig. 7)

Type. Holotype 9, KENYA: Kakamega Forest, 5200 feet, 20.xii.1970, A.E. Stubbs,

B.M. 1972-211 (in BMNH).

Descrption. Female. Length of body (excluding oviscape) 3.0 mm, of wing

3.2 mm. Head oval, a little higher than long, largely black with face and

antennae fulvous; lower occiput not distinctly swollen; frons brown, paler

anteriorly; antennae shorter than face, with 3rd segment apically rounded,

arista pubescent; 2 pairs orbital and 2 pairs frontal setae, all black; postocular

and genal setae thin and black; ocellar setae as long as frontals; epistome

slightly protruding; palpi and labellum fulvous.

Thorax shining black; scutal pubescence very fine, dark and sparse; scutum

with a brownish tinge; pleura dark brown, pubescence coarser and pale

Australian Entomologist, 2012, 39 (4) 311

ventrally and on lower margin of katepisternum; dorsocentral setae slightly in

front of line of supra-alar setae, about half way between supra-alars and

suture; apical scutellar setae well developed, nearly as long as basals.

Legs fulvous except fore femora brown, mid and hind femora and basal 2/3

of hind tibiae dark brown to black; mid tibiae with an apical black spine.

Wing (Fig. 7) similar to that of D. enzoria but 2nd hyaline indentation in cell

rı beyond stigma not crossing all of cell r233 and cell m with an outer hyaline

spot present (c.f. D. aeneus); both indentations in cell cu; crossing cell and of

approximately even width (c.f D. alexina). Halteres cream-coloured;

squamae with a brownish tinge.

Abdomen shining black with fine, sparse pubescence that is longer, denser

and pale ventrally; tergite VI about 0.7 length of tergite V. Oviscape shining

black, length 0.6 mm, about as long as tergites IV-VI combined, narrowing

posteriorly; aculeus fulvous, apically acute, narrow and needle-like.

Etymology. Derived from the type locality.

Distribution. Known only from the Kakamega Forest in western Kenya.

Comments. This species resembles D. enzoria and, like that species, the

scutum has no discernible ‘dust’. It differs from D. enzoria in the slightly

broader and less elongate wing coupled with wing pattern differences, and

from both it and D. alexina in having 3 hyaline spots in cell m.

Key to Platensina species

Modified from Hardy (1973, 1974, 1988), Hardy and Drew (1996) and Wang

(1998) by combination and inclusion of subsequently assigned species. An

additional Asian species described by Wang (1998), plus the African species

included by Munro (1947) and Norrbom et al. (1999), were transferred to

Pseudafreutreta Hering (in tribe Pliomelaenini) by Hancock (2001) and

Hancock et al. (2003) respectively.

1 Wing cell c and basal two-thirds of stigma hyaline; hyaline marginal spots

(including 2 in cell m and 3 in cell cu,) present but pale discal spots

absent; head with 1-2 pairs of frontal setae; wing relatively narrow, not

distinctly angled posteriorly near apex of cell cu; [Philippines (Luzon)] ...

a e haber k S P. amita Hardy, 1974

— Wing cell c not entirely hyaline; stigma with at most a hyaline basal spot;

pale discal spots usually present; head with 3 pairs of frontal setae; wing

often relatively broad and angled posteriorly near apex of cell cu, ........ 2

2 Wing without hyaline discal or marginal spots or indentations except for a

pair of small costal spots at bases of stigma and cell rı adjacent to veins

Se and R; respectively; wing broad and almost circular beyond basal

third, the apex evenly rounded and entirely dark [Philippines (Luzon)] ...

BL ES OG nCcO AY cer ESTERS O I Soaccbucsaate P. bezzii Hardy, 1974

Australian Entomologist, 2012, 39 (4)

Wing usually with hyaline discal and submarginal spots or indentations;

wing often broad but distinctly longer than wide, the apex at least slightly

produced and with at least a hyaline spot at apex of cell r4+5 ...........24+ 3

Wing with hyaline apical spot extending across veins R4;5 and M; cell cu,

with 2 elongate hyaline marginal indentations; cell r; without a hyaline

preapical spot [Taiwan; male unknown] .......... P. apicalis Hendel, 1915

Not as above; if hyaline apical spot crosses veins R45 and M and cell cu;

with 2 elongate hyaline indentations, then apical spot extends at least

halfway into cells r,3 and m and cell r; with a hyaline preapical spot .... 4

Wing with hyaline apex a crescentic band from cell 143 to cell m,

crossing apices of both veins R45; and M; cell dm without a hyaline

subapicalispotions wrest etecce tute ae ere Miri nee rae ce tate et ee

Wing with hyaline apex an oval or quadrate spot confined to cell ras; cell

dm normally with a hyaline subapical spot ........cc.cccccccessessceeeseesseeseeseens 7

Male wing without hyaline spots or indentations apart from a small

indentation from costa in cell r; at apex of vein R; and the apical band;

female wing cell r; with 2 elongate hyaline indentations from costa and a

small preapical spot, cell dm with a hyaline basal spot, cell r4,5 without a

basal spot and cell cu; with 1 or 2 small round indentations from wing

margin [China (Yunnan)] ....................06++ P. nigripennis Wang, 1998

Male wing cell r; with 2 narrow or 1 broad hyaline indentation from costa

beyond stigma and a small preapical spot, cells r4+s and dm either with or

without distinct basal spots and cell cu, with a pair of short or elongate

indentations from posterior margin; female unknown ....................08 6

Male wing cell r; with 2 narrow hyaline indentations crossing vein R243

into cell r243, cells r4+s and dm each with a hyaline basal spot and cell cu,

with a pair of elongate transverse indentations from wing margin almost

reaching vein Cu; [NE Burma] ................ P. alboapicalis Hering, 1938

Male wing cell r; with 1 broad hyaline indentation crossing vein R243 into

cell r243, cells r4+s and dm without hyaline basal spots and cell cu, with a

pair of small round indentations from wing margin [Australia (NE

Queensland) | Poraa A TE merce eee P. parvipuncta Malloch, 1939

Wing with 2 elongate hyaline indentations from costa in cell rı, both

crossing vein R43 into cell r233, no marginal preapical hyaline spots in

cell ro43, large hyaline spots near base of cell r4;; and near base and apex

of cell dm, 1 marginal spot in cell m near apex of vein Cu), 2 indentations

in cell cu; and 1 or 2 spots along margin of anal lobe; stigma black,

Withoufanyalines basalispOteemetentamernt en LLT TLE D TTA 8

Wing markings not as above; marginal preapical hyaline spots in cell r243

usually present; cell m usually with 2 or 3 hyaline marginal spots; stigma

ofeniwithialhyalinei basalis poti een rr Err EEE T TETTA 9

Australian Entomologist, 2012, 39 (4) 313

Male with face largely black in male, yellow in female; hyaline

indentations in cell cu; of approximately equal length, almost crossing

cell but the basal spot sometimes medially divided; basal marginal

hyaline spot in anal lobe much smaller than second marginal spot or

absent [India, Sri Lanka, Bangladesh, Burma, China (Yunnan), Thailand

and Cambodia; Ensina guttata Wiedemann, 1824, Trypeta stella Walker,

1849 and Trypeta voneda Walker, 1849 are regarded as synonyms] ......

Wa a E TAEAE och S AAA tone, P. acrostacta (Wiedemann, 1824)

Male with face yellow, female unknown; basal hyaline indentation in cell

cu; much smaller than second indentation, not almost crossing cell; the

two marginal hyaline spots in anal lobe of approximately equal size [India

(Maharashtra) | a E a a a P. fulvifacies Hering, 1941

Wing cell r; with a large, single hyaline indentation in basal portion that

crosses vein R343 and almost all of cell r243; cells m and cu; each with a

single marginal indentation, that in cell cu, crossing cell [Philippines

UZON a e E E cee reer E mentee R P. aptata Hardy, 1974

Wing cell r; with no or 2 hyaline indentations in basal portion, the outer

sometimes reduced to a rounded spot or largely united with the basal one;

cells m and cu; not both with a single marginal indentation .............. 10

Wing cell r33 with 2 hyaline marginal indentations from costa; cell r;

with indentations in basal portion often medially constricted or reduced to

marginal spots and spots in cell r243 below them isolated [Indonesia (Java,

Ambon), Papua New Guinea and Solomon Islands; P. dilatata Hering,

1941 is regarded as a new synonym] ............ P. ampla de Meijere, 1914

Wing cell ro; with at most a single hyaline marginal indentation from

costa; cell r; with indentations in basal portion distinct and crossing cell,

reduced to isolated costal spots or absent ............. ccc cece eceee eect ee A 11

Two scutellar setae, the apical pair absent [India (Bihar, Karnataka, West

Bengal: type locality), Nepal, Sri Lanka, Bangladesh, Burma. China

(Yunnan, Guangdong, Hainan), Thailand, Laos, Cambodia, Philippines

(Luzon, Mindoro), West Malaysia, Singapore, Indonesia (Java, Maluku)

and Australia (NT, Qld)] ......... ce eeeeeee ee ee ee P. zodiacalis (Bezzi, 1913)

Four scutellar setae, the apical pair distin ta aA A EEST 12

Wing relatively broad, distinctly angled posteriorly near apex of cell cuj;

cell m with at most 2 hyaline marginal indentations ....................05 13

Wing relatively narrow, evenly rounded posteriorly and not distinctly

angled near apex of cell cu;; cell m with 2 or 3 hyaline marginal

indentations 18

Comme eer cere creer reer rere eee se ese ereaee assesses seseeesreesseseeeee

Wing cell 14,5 with a very small hyaline spot at apex [Indonesia (Java,

Sumba); type species of Platensina] .......... P. sumbana Enderlein, 1911

314 Australian Entomologist, 2012, 39 (4)

— Wing cell r45 with a broad or elongate hyaline spot at apex, crossing all

onmostioficell 4. Ae is See er eae, Pee Ay sie ee 14

14 Wing often without hyaline spots except along margin, the discal area at

most with hyaline spots in cells dm and base of 14:5; beyond R-M

crossvein; second hyaline indentation in basal portion of cell r; narrow

and often united with basal one, leaving an isolated dark costal spot; anal

lobe brown, the hyaline marginal spots absent or vestigial [S Burma,

Thailand, Vietnam, Indonesia (Sumba); P. extincta Hering, 1952 is

regarded as a SYNONYM] .............eseceeeeee eens P. euryptera (Bezzi, 1913)

Wing usually with distinct hyaline or subhyaline spots, at least in cells dm

and base of r4:s; hyaline indentations in basal portion of cell rı distinct,

reduced or absent; anal lobe with hyaline marginal ae eae, distinct

15 Wing with discal spots often subhyaline; cell r; with 0-2 small Wir:

indentations from costa in basal portion beyond stigma, often neither

partly fused nor crossing cell (especially in males); cell cu; with 3 small,

isolated, hyaline marginal spots and with or without additional small,

isolated discal spots [India (Tamil Nadu), China (Guangxi, Fujian),

Taiwan, Vietnam and West Malaysia; P. fukienica Hering, 1939 is

regarded as a new synonym] .................2.0e0eeee P. tetrica Hering, 1939

Wing with distinct discal hyaline spots; hyaline indentations in basal

portion of cell r; with at least the inner one broad and crossing cell in both

sexes; cell cu; with 2 or 3 hyaline indentations, the basal pair normally

elongate but often medially divided into 2 separate spots ................. 16

16 Wing cell m with at most 1 small hyaline marginal spot; cell cu; with 2

undivided indentations almost crossing cell [Thailand, Cambodia and

Vietnam E r ase ys seas siars sts wt es tasieeres pan cone P. intacta Hardy, 1973

— Wing cell m with 2 hyaline marginal spots; cell cu; with 2 or 3 hyaline

marginal spots, the basal pair short or divided into separate spots, the

Outemspotottentreducedionabsenthesmestenntraterstsvatcetessecrt TE 17

17 Basal hyaline indentations in wing cells r) and r233 more or less

convergent, those in r23 aligned with those in rı; hyaline apical spot

relatively narrow and filling entire apex of cell 14:5; basal hyaline

indentation in cell m narrow, elongate and perpendicular; anal lobe with

hyaline marginal spots vestigial or absent; apical scutellar setae distinct,

about half length of basals [?S Thailand, West Malaysia, ?Singapore and

Indonesia (Java, Sulawesi)] .................. P. amplipennis (Walker, 1860)

— Basal hyaline indentations in wing cells r; and 143 more or less parallel,

those in rə small and off centre with those in rı; hyaline apical spot

relatively broad and not filling entire apex of cell r4+s; basal hyaline

indentation in cell m often short and broad; anal lobe with hyaline

Australian Entomologist, 2012, 39 (4) 315

marginal spots round and distinct; apical scutellar setae weak, about a

quarter length of basals [India (Utranchal, Andaman Is), Burma, Japan

(Ryukyu Is), Taiwan, Northern Marianas, Guam, Micronesia, ?Thailand,

?Laos, ?Vietnam, Malaysia (Sarawak), Indonesia (Flores), Papua New

Guinea (Admiralty Is, Central Province), Solomon Is (Malaita, Russel),

Vanuatu (Espiritu Santo, Banks) and Australia (Queensland); P. malaita

Curran, 1936 and P. dubia Malloch, 1939 are regarded as synonyms] ...

E E A P R AEE TTT P. platyptera Hendel, 1915, stat. rev.

Wing cells m and cu; each with 3 hyaline indentations from margin, those

intcelllcurallismalllandisolated me mmm enn E E ENO 19

Wing cell m with 2 and cell cu; with 2 or 3 hyaline indentations from

margin, often with at least one of those in cell cu; elongate and crossing

most of cell 20

Wing cell dm with 3 rounded hyaline spots; cell m with 2 hyaline spots in

anterobasal half [Australia (NE Queensland)] ................cccceseee eee eees

ee ROS ead FOS STRETTON Oe NS, p P. trimaculata Hardy & Drew, 1996

Wing cell dm with 2 rounded hyaline spots; cell m with 1 hyaline spot in

anterobasal half [Japan (Ryukyu Is), Taiwan and China (Sichuan,

Guangxi) | ees tee meen P. assimilis (Shiraki, 1968), comb. n.

Wing cell cu; with 3 hyaline indentations from margin, with at least the

basal pair broad and crossing cell; anal cell with a transverse hyaline

indentation crossing vein Cu,+A, into cell cu,; face with silvery spots in

male [India, Thailand, Cambodia, Vietnam] ..... P. guadrula Hardy, 1973

Wing cell cu; with 2 or 3 narrow hyaline indentations from margin, the

basal pair constricted or divided into two separate spots; anal cell with

only small, round marginal hyaline spots; face without silvery spots

[Japan (Ryukyu Is) and China (Sichuan); a male from Indonesia (West

Papua), illustrated by Hardy (1988) as Pliomelaena sp. A, is possibly this

species ea N ANR RNE e eretetes P. shirouzui (Ito, 1984), comb. n.

Key to Dicheniotes species

Modified from Munro (1947) by inclusion of subsequently described or

assigned species. * = new country records based on material in BMNH.

Stigma with a subhyaline basal spot from costa; apex of cell rə, with a

hyaline spot near tip of vein R243; outer of 3 hyaline indentations in cell m

reduced to an isolated, rounded, submarginal spot; postocular setae

largelyswhite weirs Wetec eterna es iss. cere coins Ree, ce Paes eee be eve 2

Stigma without a subhyaline spot from costa; apex of cell r3, with or

without a hyaline spot near tip of vein Rz; outer of 3 hyaline

indentations in cell m, when present, often elongate and crossing most of

Cellapostoculansetaepalerondark arara Eea E E rears 4

Australian Entomologist, 2012, 39 (4)

Wing cell m with middle marginal spot large and quadrate; cell cu; with 2

hyaline spots, the basal spot large and quadrate; cell r; without an

additional subapical spot crossing cell; submarginal spot in cell r243 large

and situated just below tip of vein R23; female oviscape as long as

abdomen Kenyaa A E nsec: D. polyspilus (Bezzi, 1924)

Wing cell m with middle marginal spot small and round; cell cu; with 3

hyaline spots, the basal spot divided into two; cell r; with an additional

subapical spot crossing cell; submarginal spot in cell r3, small and

situated nearer midline of cell; female oviscape much shorter than

abdomen Fees cscs ccces foes Sac soshed wot scee Meee RE Ee See Nene te A Tatas

Wing cell r, with inner hyaline indentation narrow and strap-like and the

middle indentation small and confined to costal margin [Ethiopia] .....:

D. parviguttatus (Hering, 1952), comb. n.

Wing cell r; with inner hyaline indentation broad and subquadrate and the

middle indentation large and quadrate [Kenya, Tanzania, Saudi Arabia

and&Y/emen | peetusecsh snr eee D. multipunctatus Merz & Dawah, 2005

Wing cell m with 2 large hyaline spots, the inner one marginal, the outer

submarginal; cell rə, with 2 small hyaline spots near tip of vein Roi3;

postocular setae largely white [Sokotra] Ea a AE E ee eees

D. sokotrensis (Hering, 1939), comb. n.

Wing cell m mostly brown or with 2 or 3 hyaline indentations, the outer

often reduced to a small, rounded spot, if 2 then both are marginal and

cell rə without a small hyaline spot near tip of vein R23; postocular

setaepalerorndarke marnan AE A e E tees meena etc tain cece

Wing cell r23 with a round, hyaline spot near the tip of vein Rz+, that does

not reach the wing margin; cell m with 3 hyaline indentations from wing

margin, the outer one often reduced to a small spot; cell dm with a pair of

isolated and well separated hyaline spots e a AAEE ATO

Not as above; wing cell r2, without a round, hyaline spot near the tip of

vein R +3; cell m usually with at most 2 hyaline indentations from wing

margin; cell dm sometimes without a pair of isolated hyaline spots ....... 9

Outer hyaline indentation from costa in cell r; confined to that cell, not

crossing vein R243 [Sudan, Saudi Arabia and Yemen] .....................4..

bids. acicc LOAA wate aac te dear’ D. angulicornis (Hendel, 1931)

Outer hyaline indentation from costa in cell r; crossing vein R243 at least

halfswayrintoscellirss3 era eet tate eae E sce uhh rem en ay 7

Outer hyaline indentation in cell m elongate and crossing most of cell

[Uganda] Pate Pere ee ee aren, Les T ss D. acclivis Munro, 1947

Outer hyaline indentation in cell m no more than a small, rounded,

Marginalforsubmareinal spotemeretereacter tests cae A EE eae se 8

Australian Entomologist, 2012, 39 (4)

—

_—

317

Labellum greatly enlarged; wing cell r243; with a hyaline basal spot near

base of stigma; femora blackened [Uganda and Kenya] ...................05.

Soren orth pone E A E tare stat retrace narod ty D. turgens Munro, 1947

Labellum not greatly enlarged; wing cell r,3; without a hyaline spot near

base of stigma; fore femur often fulvous, others black [Kenya and

(Lanzania | Bese eee ett teen tee D. aeneus (Munro, 1947), comb. n.

Wing cell m with 3 elongate hyaline indentations, the outer 2 both

crossing vein M into and across cell r4;5 [Eritrea] .................0.00005

bees es ORENS ATS TEEN D. asmarensis (Munro, 1955), comb. n.

Wing cell m with at most 2 elongate hyaline indentations (sometimes

mostly brown) and with or without a small outer spot ..................06 10

Femora fulvous; wing cell m with outer hyaline indentation broadly

crossing vein M into cell r4;5 to or almost to vein R4+5; cell cu; with 2

hyalineundentations per e I aA a A ee 11

Femora largely black; wing cell m with outer hyaline indentation often

confined to cell, if crossing vein M into cell r4+s then cell cu, with a single

broadihyalinetindentationteesrmcssctesn E E T eens AE teen EAE, 13

Wing cell dm without a pair of hyaline spots [Kenya] ..................665

r AAT Aa Ae Hea AAAS CA s R Ta LA AT D. sexfissatus (Becker, 1909)

Wing cell.dm with a pair of hyaline spots

Wing cell dm with a pair of isolated and well separated hyaline spots

[Tanzania, Zimbabwe and South Africa; records from Namibia are errors

(Hancock 2000); Brachyaciura discoguttata Hering, 1941 is regarded as a

SYDONYIN a E AATA ETAS D. distigma (Bezzi, 1924)

Wing cell dm with a pair of large hyaline spots united with the

indentations in cell cu; [Democratic Republic of Congo, Uganda, Kenya

andklanzania | pnn m Nn aer sett renee tee aan eee D. erosa (Bezzi, 1924)

Sexes distinctly dimorphic; wing largely brown with diffuse longitudinal

pale streaks and patches (some males) or cell r; with 1 broad or 2

transverse hyaline indentations from costa, cell cu; with a single, broad

hyaline indentation extending broadly into: cell dm in females, not in

males; cell m with 2 hyaline indentations, the outer one crossing vein M

into cell r4;5 and cell dm without a pair of isolated hyaline spots ......... 14

Sexes not distinctly dimorphic; wing cell r; with 2 narrow hyaline

indentations from costa; cell cu; with 2 narrow hyaline indentations not

extending into cell dm; cell m with 2 narrow hyaline indentations and at

most a small outer spot, none crossing vein M into cell r4+s; cell dm with a

pair of well separated hyaline spots an rriena Ear AEA a TEA T 15

318

Australian Entomologist, 2012, 39 (4)

Wing cell m almost entirely filled (including apex at tip of vein M) by 2

broad hyaline indentations separated by at most a dark transverse band;

the 2 hyaline indentations from costa in cells r; and 1.43 combined into a

single broad band in males, separated in females [South Africa; type

SPECIESJONDICHENIOIES |pecne cect nc nny D. dispar (Bezzi, 1924)

Wing cell m largely brown with at most a pale anterobasal streak and a

diffuse posterobasal spot and cell cu, with hyaline indentation divided

into 2 separate spots (males), or with the outer of the 2 hyaline

indentations in cell m narrow and not almost filling apex of cell (females)

[Eritrea and Ethiopia (2463, 329, Simien, ravine on W side of Mai Shaba

valley, 9000', 14.xii.1952, H. Scott*)] ....... D. tephronotus (Bezzi, 1908)

Wing cell m with inner indentation more or less parallel with DM-Cu

crossvein and with a small rounded outer spot in addition to basal and

medial indentations [Kenya, Tanzania, Yemen (24.4, Wadi Doreija, W of

Dhala, 4500", xi.1937*) and South Africa] ....... D. katonae (Bezzi, 1924)

Wing cell m with inner indentation oblique, converging with DM-Cu

crossvein anteriorly and meeting wing margin posteriorly beyond apex of

vein Cu, and with or without a small rounded outer spot .................. 16

Wing cell m with 3 hyaline indentations, the outer spot present [Kenya]

aeee D. Kakamegae sp. n.

Wing cell m with only 2 hyaline indentations, the outer spot absent .... 17

Wing cell r; with the outer hyaline indentation not crossing vein R2+3 into

cell r3 [Kenya, Zimbabwe and South Africa; one record from Australia

(SE Queensland), presumably introduced (Hancock and Drew 2003)] ...

D. ternarius (Loew, 1861), comb. n.

Wing cell r; with the outer hyaline indentation crossing vein R43 into cell

Wing with line of DM-Cu crossvein meeting costa on the outer hyaline

indentation in cell r; or close to its outer margin; squamae with a brown to

blackish tinge [Uganda] ............... D. enzoria (Munro, 1947), comb. n.

Wing with line of DM-Cu crossvein meeting costa well beyond the outer

hyaline indentation in cell rı, at least its width away; squamae yellow; a

row of 3 hyaline discal spots in cells r2, and r4+s present or absent

[Zimbabwe and South Africa (1¢, Eshowe, KwaZulu-Natal, vi.1926,

Ree Mliurnery) |persct emer eee ces D. alexina (Munro, 1947), comb. n.

Acknowledgements

I thank Sally Cowan (AQIS), Kim Goodger and Nigel Wyatt (BMNH) and

Zoé Simmons and James Hogan (OUMNH) for the loan of or access to

specimens in their care, Kim Goodger (BMNH) for Figs 1-5 & 7, K.J. David

(National Bureau of Agriculturally Important Insects, India) for photographs

Australian Entomologist, 2012, 39 (4) 319

and new records of P. platyptera and P. quadrula from India, Anthony Rice

(AQIS, Cairns) for assistance with producing Fig. 6 and Albert Orr for

helpful discussions on nomenclature.

References

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Memoirs of the Indian Museum 3: 53-175, pls 8-10.

CURRAN, C.H. 1936. The Templeton Crocker expedition in the western Polynesian and

Melanesian islands, 1933. No. 30. Proceedings of the California Academy of Sciences (4) 22: 1-

67.

de MEIJERE, J.C.H. 1914. Studien über siidostasiatische Diptern IX [concl.]. Tijdschrift voor

Entomologie 57: 169-275.

FOOTE, R.H. 1964. Notes on the Walker types of New World Tephritidae (Diptera). Journal of

the Kansas Entomological Society 4: 316-326.

HAN, H.-Y., RO, K.-E. and FREIDBERG, A. 2010. Phylogeny and classification of the

Tephrellini (Diptera: Tephritidae: Tephritinae). P. 105, in: 7th International Congress of

Dipterology, San José, Costa Rica. Abstracts Volume: 296 pp.

HANCOCK, D.L. 2000. Tephritidae (Diptera: Tephritoidea). Pp 239-248, in: Kirk-Spriggs, A.H.

and Marais, E. (eds), Daures — biodiversity of the Brandberg Massif, Namibia. Cimbebasia

Memoir 9. National Museum of Namibia, Windhoek; iv + 389 pp.

HANCOCK, D.L. 2001. Systematic notes on the genera of Australian and some non-Australian

Tephritinae (Diptera: Tephritidae). Australian Entomologist 28(4): 111-116.

HANCOCK, D.L. 2006. Taxonomic updates in some genera and species of Afrotropical

Tephritinae (Diptera: Tephritidae). African Entomology 14(2): 391-394.

HANCOCK, D.L. 2010. A review of the fruit fly tribe Tephrellini (Diptera: Tephritidae:

Tephritinae) in the Indo-Australian Region. Australian Entomologist 37(1): 1-6.

HANCOCK, D.L. and DREW, R.A.I. 2003. A new genus and new species, combinations and

records of Tephritinae (Diptera: Tephritidae) from Australia, New Zealand and the South Pacific.

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HANCOCK, D.L., KIRK-SPRIGGS, A.H. and MARAIS, E. 2003. New records of Namibian

Tephritidae (Diptera: Schizophora), with notes on the classification of subfamily Tephritinae.

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Pacific Insects Monograph 31: 1-353, pls 1-8.

HARDY, D.E. 1974. The fruit flies of the Philippines (Diptera: Tephritidae). Pacific Insects

Monograph 32: 1-266, pls 1-6.

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Islands (Diptera: Tephritidae). Bishop Museum Bulletin in Entomology 1: vii + 1-92.

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Tephritidae). Invertebrate Taxonomy 10(2): 213-405.

HENDEL, F. 1915. H. Sauter’s Formosa-Ausbeute. Tephritinae. Annales Musei Nationalis

Hungarici 13: 424-467, pls viii-ix.

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143-147.

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und Taxonomische Entomologie aus Berlin-Dahlem 8(1): 24-45.

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Naturforschenden Gesellschaft in Basel 63: 41-48.

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identification system and systematic information database. Myia 9: ix + 524 pp.

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Australian Entomologist, 2012, 39 (4): 321-329 321

A NEW SPECIES OF THE SUBGENUS CAMPOMYRMA WHEELER

OF THE GENUS POLYRHACHIS FR. SMITH FROM THE

AUSTRALIAN CAPITAL TERRITORY

(HYMENOPTERA: FORMICIDAE: FORMICINAE)

RUDOLF J. KOHOUT

Biodiversity Program, Queensland Museum, PO Box 3300, South Brisbane, Qld 4101

(Email: rudolf:kohout@qm.qld.gov.au)

Abstract

Polyrhachis smithersi, a new species of the hexacantha complex of the subgenus Polyrhachis

(Campomyrma) Wheeler, is described from the Australian Capital Territory. A key

distinguishing it from the three other described species of the complex is provided. All four

species are illustrated and their distribution data summarised.

Introduction

The hexacantha complex of Polyrhachis (Campomyrma) Wheeler comprises

four species: P. fuscipes Mayr, P. hexacantha (Erichson), P. semipolita

André and a new species described below. All four are rather similar and

share many characters, including a distinctly slender mesosoma, a propodeal

dorsum with two slender, posteriorly directed spines and a petiole with

elongated lateral spines and a pair of shorter intercalary spines or teeth.

Taylor (1989) also considered these species to be closely related and ‘similar

and distinguished primarily by sculptural differences. P. fuscipes is less

regularly and coarsely sculptured than P. hexacantha, while P. semipolita has

all body surfaces remarkably smooth and strongly reflective. The 3 species

seem to be similarly distributed (except that P. hexacantha ranges north to

the New England Tableland) and are likely closely sympatric in parts of

southeastern Australia and Tas.’ [Tasmania]. His opinion on their sympatric

association has been confirmed, with three species of the complex collected

by the author in close proximity at the type locality of the new species,

Smokers Gap in the Australian Capital Territory.

Methods

Photographs of the specimens were taken with a digital camera attached to a

stereomicroscope and processed using Auto-Montage (Syncroscopy, Division

of Synoptics Ltd, USA) and Adobe Photoshop CS2 (Adobe Systems Inc.,

USA). Images of P. smithersi sp. n. depict the holotype, those of P.

hexacantha, P. fuscipes and P. semipolita depict the types or type-compared

specimens from the ANIC collection. All specimens were photographed by

Dr Steve O. Shattuck (ANIC).

The standard measurements and indices mainly follow those of Kohout

(2008): TL = Total length (the necessarily composite measurement of the

outstretched length of the entire ant measured in profile); HL = Head length

(the maximum measurable length of the head in perfect full face view,

measured from the anterior-most point of the clypeal border or teeth to the

322 Australian Entomologist, 2012, 39 (4)

posterior-most point of the occipital margin); HW = Head width (width of the

head in perfect full face view, measured immediately in front of the eyes); CI

= Cephalic index (HW x 100/HL); SL = Scape length (length of the antennal

scape, excluding the condyle); SI = Scape index (SL x 100/HW); PW =

Pronotal width (greatest width of the pronotal dorsum, measured behind the

pronotal teeth); MTL = Metathoracic tibial length (maximum measurable

length of the tibia of the hind leg). All measurements were taken using a

Zeiss (Oberkochen) SR stereomicroscope at 20x and 32x magnifications with

an eyepiece graticule calibrated against a stage micrometer. All

measurements are expressed in millimetres (mm).

Abbreviations. General: acc. — accession/s; ACT — Australian Capital

Territory; for. — forest; N.P. — National Park; NSW — New South Wales; Rd —

Road; rf. — rainforest; sclero. — sclerophyl forest; TAS — Tasmania; VIC —

Victoria; w — worker/s.

Institutions (with names of cooperating curators): AMSA — Australian

Museum, Sydney, NSW, Australia (Drs D. Britton, D. Smith); ANIC —

Australian National Insect Collection, CSIRO Entomology, Canberra, ACT,

Australia (Dr S.O. Shattuck); BMNH — The Natural History Museum,

London, UK (B. Bolton, S. Reider); CASC — California Academy of

Sciences, San Francisco, CA, USA (Dr B.L. Fisher); MCZC — Museum of

Comparative Zoology, Harvard University, Cambridge, MA, USA (Dr S.P.

Cover); MNHU — Museum fiir Naturkunde, Humboldt-Universitat, Berlin,

Germany (Dr F. Koch); MVMA — Museum Victoria, Melbourne, VIC,

Australia (Dr K. Walker); NHMW -— Naturhistorisches Museum, Wien,

Austria (Drs H. Zettel, D. Zimmermann); OXUM — Hope Entomological

Collections, University Museum, Oxford, UK (Dr D.J. Mann); QMBA —

Queensland Museum, Brisbane, QLD, Australia (Dr C.J. Burwell); ZMSG —

Zoologische Staatssammlung, Miinchen, Germany (Dr E. Diller).

Systematics

Genus Polyrhachis Fr. Smith, 1857

Polyrhachis Fr. Smith, 1857: 58. Type species: Formica bihamata Drury, 1773, by

original designation.

Subgenus Campomyrma Wheeler, 1911

Campomyrma Wheeler, 1911: 860 (as subgenus of Myrma Billberg, 1820 =

Polyrhachis Fr. Smith, 1857). Type species: Polyrhachis clypeata Mayr, 1862

(junior synonym of Polyrhachis exercita Walker, 1859), by original designation.

Key to workers of P. hexacantha complex

1 Smaller species (HL < 1.81); lateral petiolar spines only weakly divergent

in dorsal view, forming a continuous line with sides of petiolar node (Fig.

DO GLANO NLI AIET erred sip novel tese P. smithersi sp. n.

- Larger species (HL > 1.95); lateral petiolar spines widely divergent,

strongly extending laterally before curving posteriorly (e.g. Fig. 9) ........ 2

Australian Entomologist, 2012, 39 (4) 323

2 Body very smooth, highly polished

~emeBodyitinelyisculptured eE rier ren a ees 3

3 Dorsum of mesosoma very finely and somewhat longitudinally striate;

sculpturation of head distinctly finer, semipolished; intercalary spines of

petiole only moderately long (Figs 6-7) „s.es P. fuscipes Mayr

- Dorsum of mesosoma regularly reticulate-punctate, opaque; intercalary

spines of petiole distinctly longer (Figs 9, 11-12) ...cccescseseeeeesseseeees

mamennou, P. hexacantha (Erichson)

Polyrhachis smithersi sp. n.

(Figs 1-2, 4-5)

Types. Holotype worker: AUSTRALIAN CAPITAL TERRITORY, Smokers Gap,

Corin Dam Rd., 35°31’S, 148°54’E, 1240 m, 18-19.ii.2001, dry open forest, at night,

R.J. Kohout acc. 01.3. Paratypes: 9 workers, queen, data as for holotype; 7 workers,

as for holotype except 28-29.x.1973, R.J. Kohout acc. 73.138; 7 workers, as for

holotype except 20.1.1982, R.J. Kohout acc. 82.40. Type distribution: Holotype, 5

paratype workers and paratype queen in ANIC; 8 paratype workers in QMBA; 2

paratype workers each in AMSA, BMNH, CASC, MCZC and MVMA.

Description. Worker. Dimensions (holotype cited first): TL c. 7.41, 6.60-

7.41; HL 1.81, 1.65-1.81; HW 1.53, 1.37-1.53; CI 84, 81-85; SL 1.87, 1.78-

1.87; SI 122, 122-134; PW 1.12, 1.06-1.15; MTL 2.18, 2.00-2.18 (6

measured).

Mandibles with 5 teeth. Anterior clypeal margin widely truncate medially,

truncate portion somewhat irregularly denticulate and flanked by distinct

angles. Clypeus in profile weakly concave, with median carina distinctly

raised towards anterior and basal margins. Frontal triangle distinctly

impressed; frontal carinae sinuate with narrowly and weakly raised margins;

central area relatively wide with distinct frontal furrow. Sides of head in front

of eyes rounding towards mandibular bases in weakly convex line; behind

eyes sides rounding onto evenly convex occipital margin. Eyes convex, in

full face view clearly breaking lateral cephalic outline. Ocelli lacking,

position indicated by shallow pits in cephalic sculpture. Mesosomal dorsum

elongated; pronotal dorsum with distinct humeral teeth; lateral margins

evenly rounded into distinctly impressed promesonotal suture. Mesonotum

with lateral margins distinctly converging towards rather flat, metanotal

groove. Propodeal margins subparallel, terminating posteriorly in obliquely

raised, subparallel, acute spines; propodeal dorsum flat, descending abruptly

in medially uninterrupted line into steeply concave declivity. Petiole in side

view with anterior and posterior faces virtually parallel from base; dorsum

armed with slender, strongly raised, divergent and acute lateral spines; bases

of spines continuous medially onto narrow dorsum and merging into two

widely separated, blunt intercalary teeth. Anterior face of first gastral

segment only weakly concave, widely rounding onto dorsum.

324 Australian Entomologist, 2012, 39 (4)

Figs 1-7. Polyrhachis (Campomyrma) spp: (1-2, 4-5) P. smithersi sp. n.; (3, 6-7) P.

fuscipes Mayr. (1, 3) head in full face view; (2) petiole in front view; (4, 6) dorsal

view; (5, 7) lateral view.

Mandibles finely striate at bases, rather irregularly rugose towards

masticatory borders with shallow pits. Head rather finely, dorsum of

mesosoma more coarsely reticulate-punctate, with former somewhat

semipolished; sides of mesosoma very finely reticulate-wrinkled. Gaster

finely sculptured, opaque; sculpturation somewhat finer and semipolished in

some specimens.

Mandibles with numerous suberect, golden hairs near masticatory borders.

Anterior clypeal margin with a few long setae medially; clypeus and frontal

carinae with paired, medium length, golden hairs; hairs absent from vertex,

dorsum of mesosoma, petiole and most of gastral dorsum. Several longer,

golden, semierect hairs on apex and venter of gaster; occasional erect hairs on

venter of coxae and fore femora. Closely appressed pubescence very

sparingly distributed on various dorsal surfaces, notably on head and gaster.

Australian Entomologist, 2012, 39 (4) 325

Black; mandibles, except teeth, distinctly orange-red; legs orange-red to

reddish-brown, coxae and tarsi a shade darker. Antennae medium reddish-

brown; funiculi reddish-brown at base, with segments progressively lighter

towards apices. Gaster reddish-brown with margins of segments lined a shade

darker.

Queen. Dimensions: TL c. 7.51; HL 1.65; HW 1.37; CI 83; SL 1.68; SI 123;

PW 1.28; MTL 1.96 (1 measured).

The single available queen is rather small, about the size of a small worker.

However, it displays the usual characters identifying full sexuality, including

three ocelli, complete thoracic structure and wings. Pronotal humeri bluntly

angular; mesoscutum virtually as long as wide in dorsal view with lateral

margins strongly converging anteriorly into rather narrowly rounded anterior

margin; median line distinct; parapsides flat; mesoscutum in profile with

relatively low anterior face rounding onto flat dorsum. Mesoscutellum flat,

not elevated above dorsal plane of mesosoma. Propodeal dorsum weakly

convex, descending into steep, weakly concave declivity in medially

uninterrupted line; propodeal spines shorter than in worker, weakly divergent.

Petiole virtually identical to that in worker, except for absence of intercalary

teeth. Sculpturation of body, pilosity, pubescence and colour virtually as in

worker.

Male and immature stages unknown.

Etymology. It is a pleasure to name this species for the late Courtenay

Smithers, a generous and productive entomologist who left a lasting legacy of

insect studies in Australia.

Remarks. Polyrhachis smithersi is so far known only from the type locality,

with all specimens collected foraging at night on Eucalyptus tree trunks in

open forest.

Polyrhachis fuscipes Mayr, 1862

(Figs 3, 6-7)

Polyrhachis fuscipes Mayr, 1862: 679. Holotype worker. Type locatity: TASMANIA

(as Van Diemensland), NHMW.

Polyrhachis fuscipes Mayr; Roger, 1863: 9. Junior synonym of P. hexacantha.

Polyrhachis fuscipes Mayr; Dalla Torre, 1893: 263. Junior synonym of P. hexacantha.

Polyrhachis hexacantha subsp. fuscipes Mayr; Emery, 1925: 179. Combination in P.

(Campomyrma) and revived from synonymy as subspecies of P. hexacantha.

Polyrhachis fuscipes Mayr; Taylor and Brown, 1985: 134. Revived status as species.

Polyrhachis semipolita subsp. hestia Forel, 1911: 295. Holotype worker. Type

locality: Australia (Bates), ZMSG (examined).

Polyrhachis fuscipes Mayr; Taylor, 1989: 23. Senior synonym of P. hestia.

326 Australian Entomologist, 2012, 39 (4)

Additional material examined. NEW SOUTH WALES (including Australian Capital

Territory): Thredbo, 30.ii.1982 (A.N. Andersen) (w); 12 km of Kanangra Walls N.P.,

2.111977, sclero. (B.B. Lowery) (w); Smokers Gap, 35°31’S, 148°54’E, 1240 m, 28-

29.x.1973, dry open forest, at night (R.J. Kohout acc. 73.139) (w); ditto, 20.1.1982

(RJK acc. 82.40) (w); ditto, 18-19.ii.2001 (RJK acc. 01.8) (w); Jenolan Caves (J.C.

Wilburt) (w). VICTORIA: Mt Oberon, Wilson’s Promotory, 24.11.1982 (A.N.

Andersen) (w); Forrest (H.W. Davey) (w); Victoria (no further data) (w).

TASMANIA: 15 km W of Swansea, xi.2003 (N. Meeson) (w); Hobart (A.M. Lea)

(w).

Worker. Dimensions: TL c. 7.81-8.87; HL 1.96-2.12; HW 1.62-1.81; CI 81-

85; SL 2.10-2.37; SI 126-136; PW 1.28-1.47; MTL 2.43-2.65 (9 measured).

Remarks. Polyrhachis fuscipes is known from the mountainous parts of

southeastern NSW and ACT, and south through the Victorian Alps to

Tasmania. It was collected in sympatry with P. smithersi and P. hexacantha

at Smokers Gap in ACT.

Polyrhachis hexacantha (Erichson, 1842)

(Figs 8-9, 10-11)

Formica hexacantha Erichson, 1842: 260. Syntype worker, queen (lectotype and

paralectotype designated by Taylor, 1989: 24). Type locality: TASMANIA,

?Woolnorth (?A. Schayer), MNHU (worker examined).

Polyrhachis hexacantha (Erichson); Fr. Smith, 1858: 74. Combination in Polyrhachis.

Polyrhachis hexacantha (Erichson); Emery, 1925: 179. Combination in P.

(Campomyrma).

Polyrhachis froggatti Forel, 1910: 89. Syntype workers. Type locality: NEW SOUTH

WALES, Bombala (W.W. Froggatt), ANIC, QMBA (examined).

Polyrhachis hexacantha (Erichson); Taylor, 1989: 24. Senior synonym of P. froggatti

Forel.

Additional material examined. NEW SOUTH WALES (including AUSTRALIAN

CAPITAL TERRITORY): NSW (no further data) (W.M. Wheeler) (w); Mt

Kosciusco, The Creel, 3000’, 14-15.xii.1931 (W.M. Wheeler, Harvard Aust. Exp.)

(w); Mt Kosciusco, viii.1957 (Darlingtons) (w); Kosciusko NP, Island Bend,

26.xi.1952 (A. Musgrave) (w); ditto, 24.1.1975 (P. Ward #597) (w); Jenolan Caves

(J.C. Wilburt) (w). ACT, Blundells Creek, 2600’, 18.xii.1931 (W.M. Wheeler,

Harvard Aust. Exp.) (w); Smokers Gap, 35°31’S, 148°54’E, c. 1240 m, 7 &

10.xi.1973 (RJK accs 73.196 & 204) (w). VICTORIA: Mt Buffalo, 4500’ (F.E.

Wilson) (w); ditto, 5000 ft, 8.ix.1958 (B.B. Lowery) (w). TASMANIA: Tasmania (no

further data) (A.M. Lea) (w); Derby (41°08’S, 147°47’E), 10.1.1949 (T. Greaves) (w);

Oatlands (42°18’S, 147°22’E), 10.x.1956 (J. McAreavey) (w); University Reserve,

xi.2003 (N. Meeson) (w); Trevallyn, 19.iii.1928 (V.V. Hickman) (w).

Worker. Dimensions (syntypes of hexcacantha cited first, froggatti second):

TL c. 7.81, 9.27, 7.81-9.27; HL 1.96, 2.25, 1.96-2.25; HW 1.53, 1.87, 1.53-

1.87; CI 78, 83, 78-83; SL 2.18, 2.46, 2.18-2.46; SI 142, 131, 131-142; PW

1.22, 1.56, 1.22-1.56; MTL 2.65, 2.96, 2.59-3.03 (2+8 measured).

Australian Entomologist, 2012, 39 (4) 327

Figs 8-14. Polyrhachis (Campomyrma) spp: (8-9, 11-12) P. hexacantha (Erichson);

(10, 13-14) P. semipolita André. (8, 10) head in full face view; (9) petiole in front

view; (11, 13) dorsal view; (12, 14) lateral view.

Remarks. Polyrhachis hexacantha is very similar to P. fuscipes but differs by

the sculpturation of the body consisting of very closely spaced reticulate

punctations that gives specimens a rather dull, opaque appearance. In

contrast, the sculpturation of the body in P. fuscipes is distinctly finer,

notably on the head, and on the mesosomal dorsum it forms more-or-less

longitudinal striae with a somewhat semipolished appearance. The intercalary

spines in P. hexacantha are rather long, while in P. fuscipes they are

distinctly shorter. Both species co-occur with P. smithersi at Smokers Gap in

ACT.

Polyrhachis semipolita André, 1896

(Figs 10. 13-14)

Polyrhachis semipolita André, 1896: 251. Syntype workers. Type locality:

AUSTRALIA, Victorian Alps, ANIC (examined).

328 Australian Entomologist, 2012, 39 (4)

Polyrhachis hexacantha subsp. semipolita André; Emery, 1898: 228. Subspecies of P.

hexacantha.

Polyrhachis hexacantha subsp. semipolita André; Emery, 1925: 179. Combination in

P. (Campomyrma).

Polyrhachis semipolita André; Clark, 1934: 72. Revived status as species.

Additional material examined. NEW SOUTH WALES (including AUSTRALIAN

CAPITAL TERRITORY): Buccleuch St. Forest (35°21’S, 139°53’E), S of Wee

Jasper, 3300 ft, dry sclero, carton nest, 15.x.1995 (B.B. Lowery) (w); Mt Gemini, c.

6000 ft, 22.xii.1975, under snow gums at edge of sphagnum moss swamp (B.B.

Lowery) (w). VICTORIA: Emerald, 37°56’S, 145°26’E, 9.xii.1934 (J. Clark) (w);

Victoria (no other data) (Narri-Warren) (w). TASMANIA: Mt Nelson, 42°55’S,

147°20°E, 10.xii.1956 (J. McAreavey) (w); ditto, 2.11991, dry sclero, ex carton

material mound (B.B. Lowery) (w); 4 mi N of Dover (43°20’S, 147°00’E), 15.1.1949

(T. Greaves) (w); 15 km W of Swansea xi.2003 (N. Meeson) (w); Cunningham,

x1.2003 (N. Meeson) (w).

Worker. Dimensions: TL c. 7.91-8.87; HL 2.00-2.18; HW 1.68-1.87; CI 84-

90; SL 2.25-2.43; SI 126-134; PW 1.43-1.62; MTL 2.59-2.90 (8 measured).

Remarks. With its highly polished body, P. semipolita is a very easily

recognised member of the hexacantha complex. Like the other members it

occurs in the mountainous parts of southeastern NSW and the ACT to the

Victorian Alps and as far south as Tasmania.

Acknowledgements

I am very grateful to the Harvard University Grant committee for three Ernst

Mayr Grants, allowing me to travel and study specimens in the Museum of

Comparative Zoology and other museums and institutions in the USA and

Europe. I am much indebted to Drs Steve Shattuck (ANIC), Stefan Cover

(MCZC), Barry Bolton (BMNH) and Daren Mann (OXUM) for unlimited

access to the collections in their care. My sincere thanks go to my colleagues,

Drs Chris Burwell, Ms Susan Wright and Ms Karin Koch (all QMBA), for

their support during the course of this study. My special thanks are due to Dr

Steve Shattuck (ANIC) for his patience and care in preparation of the digital

images used for illustrations. Finally, my thanks go to Dr Chris Burwell

(QMBA) for reading and commenting on a draft of the manuscript.

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