THE AUSTRALIAN
Entomologist
published by
THE ENTOMOLOGICAL SOCIET OF QUEENSLAND
Volume 38, Part 1, 25 March 2011
Price: $8.00 per part
ISSN 1320 6133
THE AUSTRALIAN ENTOMOLOGIST
АВМ: 15 875 103 670. :
The Australian Entomologist is a non-profit journal published in four parts annually
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Queensland Museum (geoff.monteith@bigpond.com)
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Cover: A male of Canungrantmictis morindana Brailovsky 2002 (Heteroptera:
Coreidae). This large (25-30mm) coreid bug is spectacular in appearance but
extremely cryptic in the field. Adults hang ventral side upwards among foliage of its
food plant, the twining vine Morinda jasminoides (RUBIACEAE). It was known
from a single old specimen labelled "northern NSW" in the British Museum until the
1980s when discovery of its food plant allowed it to be reliably collected and
described. It is now known to occur from Taree to Brisbane with an isolated
population at Carnarvon Gorge.
Illustration by Geoff Thompson, Queensland Museum.
Australian Entomologist, 2011, 38 (1): 1-20 1
THE BENEFITS OF USING BOTH АР)
STONEFLIES (PLECOPTERA) IN ENVIR
AN EXAMPLE FROM NEW SOUTH WA]
OF THE AUSTRALIAN STO
WMENTAL SURVEYS:
3 ү
!NSW Department of Environment, Climate Change and Water, РО Box 29, Lidcombe,
NSW 1825
?Snowy Hydro Limited, PO Box 332, Cooma, NSW 2630
Abstract
Freshwater environmental assessment and monitoring studies rely heavily on aquatic insect
larvae because larvae are seen to directly reflect the aquatic environment they inhabit. However,
the species-level taxonomy of most aquatic insects is based on the diagnostic characters found in
adults. The uncertainty of correlating adult-based species with aquatic larvae means that aquatic
life stages are rarely identified further than generic level, especially where there are sympatric
congeners. However, Plecoptera adults are poor dispersers and generally remain near their site of
emergence. Therefore, stonefly adults have the potential to provide more finely resolved species-
level information about their nearby aquatic larval sites. Both adult and larval Plecoptera were
collected during sampling for macroinvertebrates in a monitoring project in the Snowy
Mountains, New South Wales. The results show that the collection and identification of adult
stoneflies has potential for finer scale delimitation of riverine ecosystems than information
provided by larvae alone. Also, adult stoneflies collected can provide information in related
studies of taxonomy, ecology, biogeography, biodiversity, conservation and climate change. An
updated check-list and an outline of taxonomic studies of Australian stoneflies, including all
primary literature, are also presented.
Introduction
There has been a long standing division in the study of aquatic insects by
ecologists and taxonomists. Whereas environmental and ecological studies
are based almost exclusively on larvae, partially because larvae are seen to
reflect directly the aquatic environment they inhabit, species-level taxonomy
of aquatic insects is based on adults, because they present diagnostic
characters such as wing venation and male genitalia. Aquatic entomologists
would ideally like to identify larvae collected in surveys to species-level and
thereby maximize biological and environmental information. However, in
reality they frequently must settle for coarse level generic identifications,
especially if the aquatic site hosts a number of sympatric, congeneric species.
Of all the entirely aquatic insect orders (Ephemeroptera, Odonata, Plecoptera,
Megaloptera and Trichoptera), plecopteran adults are the poorest flyers,
generally staying close to the emergence/breeding habitat and sometimes
even occurring in aquatic kick-net samples. Numerous stonefly species are
brachypterous, some even apterous, and adults of one North American
species never emerge from the water. Adult stoneflies may be collected in the
morning and during dull days as they emerge onto rocks protruding from the
streams, or at other times by beating foliage along the stream margin
(Theischinger 1991). Also, in winged species running, not flying is
2 Australian Entomologist, 2011, 38 (1)
commonly used to find food and mates. Since adults are often found in
association with congeneric and probably conspecific larvae, it is likely that
adults collected from vegetation, rocks and logs adjacent to a stream at a
particular site have emerged from that water body.
In spite of the availability of several comprehensive keys to the larvae of the
Victorian and New South Wales species (Hynes 1978, Yule 1997), species
level identifications of larvae are difficult because of a lack of diagnostic
characters, often made more uncertain because of the involvement of
different larval instars. Moreover, the larvae of many species are still
unknown.
The following study was based on surveys during 2009 for the program
‘Snowy Hydro Cloud Seeding Trial: Further Studies on Aquatic
Macroinvertebrates’, in the Snowy Mountains of New South Wales,
Australia. The monitoring team (the authors of this paper) were transported to
sites mainly by helicopter. A decision was made to collect stonefly adults in
addition to the normal aquatic samples, and thereby test if adults could
provide significant additional information for such projects in the region.
I. The Snowy Hydro Cloud Seeding Trial Monitoring Program
— Results for Plecoptera
Methods
Site locations
The sites were selected in 2007 for the Snowy Hydro Cloud Seeding Trial
Monitoring Program, and aquatic larvae at the sites have been sampled
annually since then. All sites are located within Kosciuszko National Park
and include sites of varying altitude that are within the cloud seeding areas,
plus control sites to the south of the seeding areas. The location of these sites
is shown in Fig. 1.
Sampling
In November 2009, kick-net samples were taken using the standard protocol
AusRivas (edge/riffle) (Turak and Wadell 2001). Subsequent to this up to 10
minutes per site were spent walking in the water and sweeping emergent and
trailing vegetation around the site for adult stoneflies, using an aerial insect
net (diameter 18"). All the collected stonefly material was emptied into small
plastic jars containing 70% ethanol, sorted in the lab and then identified using
the guide of Theischinger and Cardale (1987). A list was prepared including,
for each site, geographical coordinates (latitude and longitude in decimal
degrees) and altitude, numbers of males and females of adults collected by
adult sampling as well as adults and larvae collected by AusRivas kick-net
sampling in both edge and riffle habitats. The results were compared with the
2008 sampling results obtained from identification (by consultants) of larvae
only.
Australian Entomologist, 2011, 38 (1)
Macroinvertebrate Sampling Sites
EUCUMBENE RIVER
€ Sampling sites
— — Roads
Rivers
| 0 5,000 10,000 20,000 30,00
0
C SE ree Meters
Fig.1. Location of the study sites. Source: Snowy Hydro Limited.
A Australian Entomologist, 2011, 38 (1)
Analyses
The stonefly data were analysed to investigate whether they could provide
information on species distribution, dispersal and habitat constraints. The
location of specific stoneflies was compared with elevation and with known
distribution and habitat requirements by the senior author, based on personal
field experience and existing literature.
Stonefly assemblage data were also compared using the statistical software
PrimerE. Multidimensional scaling (MDS) using the Bray-Curtis measure of
similarity and presence/absence transformation was applied to the data.
Species that were responsible for dissimilarities between sites were
investigated using a simple ExcelO spreadsheet and the sites were ordered by
height above sea level to look for differences in species associations and
elevation (shown in Table 2).
Results
Sampling
Stoneflies (adults and larvae) that were collected at 16 sites for ‘Snowy
Hydro Cloud Seeding Trial; Further Studies on Aquatic Macroinvertebrates'
in November 2009 are shown in Table 1 below.
The following list provides the mostly confidently identified taxa that were
collected in the 2009 sample:
Austroperlidae
Acruroperla atra (Samal)
Austroheptura illiesi Hynes
Austroheptura sp.
Leptoperla / Riekoperla sp.
Newmanoperla thoreyi (Banks)
Riekoperla alpina McLellan
Riekoperla hynesorum Theischinger
Riekoperla karki McLellan
Eustheniidae Riekoperla reticulata (Kimmins)
Cosmioperla kuna (Theischinger)
Gripopterygidae
Gripopterygidae sp.
Dinotoperla brevipennis Kimmins
Dinotoperla christine McLellan
Dinotoperla eucumbene McLellan
Dinotoperla fontana Kimmins
Dinotoperla hirsuta McLellan
Dinotoperla thwaitesi Kimmins
Dinotoperla subserricauda Theischinger
Dinotoperla uniformis Kimmins
Dinotoperla sp.
Illiesoperla sp.
Leptoperla sp.
Riekoperla rugosa (Kimmins)
Riekoperla tuberculata McLellan
Riekoperla sp.
Trinotoperla montana (Riek)
Trinotoperla nivata Kimmins
Trinotoperla sp.
Notonemouridae
Notonemouridae sp.
Austrocerca tasmanica (Tillyard)
?Austrocerca ?tasmanica
Austrocercella hynesi Шіеѕ
?Austrocercella ?hynesi
Austrocercella ?tillyardi
Australian Entomologist, 2011, 38 (1) 5
Table 1. Plecoptera collected at sites for the Snowy Hydro Cloud Seeding Trial in
November 2009. Legend: @ = male adults, 9 = female adults; L = larvae; Au =
Austroperlidae, Eu = Eustheniidae, Gri = Gripopterygidae, No = Notonemouridae.
Edge/Riffle N
Coll: G. Theischinger
?Austrocerca ?tasmanica
°
o Austrocercella hynesi
Location data Family Species é 9 L
| Eu [Cosmipelakma ^ | [|]
SNO2 24/11/2009 | Gri | Dimotperlahirsua | if
Snowy River aboveBlue — | бі | Riekoperlahynesorums | | n |
D Gri
36420668 14833132E [ст — p Riekoperlasp. | M T
Biest Gi | Trinotperlasp. E
| 2Austrocerca tasmanica | | 1 |
| Austrocercellahynesi 12 | 3 |
ELIT. c ud
o
“к. ësst — — | J]
SNOS 241102009 Gri — | Riekoperla mesom — — | 16
Snowy River above
Spencers Creek Ori | Riekoperlasp. ||
36400608 14834314E | Gi | Trimorperap — ||
Eley. 1650 m asl, [— Ne _[?Ашшосетейа?уы | |
SE [No | Ansrocercela zallari — | — — |
Scis cca ata e a md)
е
| 18 |
| 10 |
шыг
EZ
EZ
==
SNO4 2311200 | Eu | Cosmiopelakna | TI
Kosciusko Creek above | Gi | героретар — | — | 1 |
Snowy River
3644278 5 148.30067 E
Elev. 1830 m asl. | Gi [Ж Ший» _ = | —]| 3 [| 24
Edge/Riffle Austrocercella hynesi Liew]
Coll:G.Theischinger — | No [s TE
== — tma =
SNO7 2311209 | Eu [| Cosmioperlakma | [ [ү
Snowy River above Ee aoa db dE |
Club Lake Creek [ Gi | Riekoperlahynesorum LI 5 | 4 | |
36.43138 S 14831946E | — Gri — | Riekoperlareticulata | 3 | 3 | |]
Elev. 1730 m asl. pu 1л» ee umasa: [ura cl
Edge/Riffle | No | Austrocercellahynesi | 1 | 4 | |
Coll: С. Theischinger | No | Austrocercella?iilyard | | 1 | |
ELILIEERENUEEIENM 7 01 UT T U T тын ООШ ANI
| Au | Austrohepturasp |] | [з |
| Eu [|Cosiopelakwa | | | 4 |
GEEL 24/11/2009 | Gri | Dimoperlafomama — | 7 | 10 | |
Gechi River above Gri — | Dimotoperlatwaitesi — | u | s | |
DG: | Gi | Dinotoperlasp, — | — | Tv"
“ZEE Hl Gri — | Trinotoperlamontana | 2 | | |
E AERE | Gri | Trinotoperlanivaa | — | 1 | |
Se o бй | Trinotoperlasp,§ [| | ^ | 1 |
кылыр ell
Eltre анаа а |ә |
6 Australian Entomologist, 2011, 38 (1)
Location data Family Species á
Austroheptura illiesi
Е]
Austroheptura sp. E=
m
E
A
=
DCK) 25/11/2009 Eu Cosmioperla kuna
Dicky Cooper Creek near Gri Dinotoperla hirsuta 3
Schlinks Hut ri Dinotoperla sp. sz
36.27524S 148.38231Е тї Riekoperla hynesorum
Elev. 1770 masl. ri Riekoperla karki
Edge/Rifle ri Riekoperla reticulata 20
Coll: G. Theischinger
z
e: IB.
Austrocerca tasmanica
N Austrocercella ?tillyardi
©
N
е
N
+
° Kä ЕЈ =
Austroheptura sp.
Cosmioperla kuna
Dinotoperla eucumbene
DCK2 25/11/2009 Dinotoperla fontana 3 3
Dicky Cooper Creek [он | Dinotoperla тае | 20 | 8 |
SE шн ПАТО ОП EE
SEA KOC | Gi | LeptoperlaRiekoperlasp | | | 1 |
UNO ox | Gri [Rikoeraapin — | 7 | 7 | — |
C reds EE acram mss [m
=. [zi
zi
[ Ea Tëscht | J T s]
[Gi | Dinotoperta brevipennis | 3 | 8 | |
ТОМ. 2999 [Gd | Dinovepertafomana TE — | 16 | m | |
E Dinotoperla hirsuta Г Sy | 2a | . 1
Semis Lagun EE Dinotopertatmeaiesi — | 9 | 9 |
ERU TENET он | Dinotopertasp. ра
Edge/Ri [ Gi meas. LL
ge/Riffle
Coll: G. Theischinger | O | Trinovoperlanivara | |||
[Gi e | — | IT]
Ee 35
Cosmioperla kuna
inotoperla brevipennis
inotoperla Fontana
Gri Dinotoperla hirsute
inotoperla thwaitesi
Q
a.
S
°
TUM2 23/11/2009
Tumut River below
SE 8107Е бп E uniformis
Dinotoperla sp.
Elev. 1360 m asl. бп SE
Edge/Riffle ri iekoperla tuberculata
Trinotoperla montana
Coll: G.Theischinger
Trinotoperla nivata
Trinotoperla sp.
Australian Entomologist, 2011, 38 (1) 7
Location data Family Species 3 9 L
| Eu | Cosmioperlakuna | J J è|
| Gri | Dinotoperla brevipennis | | 2 | |
AUS oes IL a | бт j 1
Tumut RiveratRound ` [ Gi | Dinoroperta gene — — | 4 | ü | __
Mountain mn ri | Dinorperla tman — —| 8 | 8 | —]
ЕЗДЕ [ oi [Dimog а
Edge/Riffle L Gi | Trinotoperiamontana | | 3 | |
ZE en 98 [ Trinoroperlanivaa — — | 1 | 1 | _|
| Gd [Trmooperlasp. —— LL | s |
p Г ЛГ 10356]
| Aw | Austrohepturasp | | | 2 |
| Еш [|Comipelakma —— LL [о |
DBI 23/11/2009 | Gi | Dinotoperla Fontana | | 1 | |
Doubtful Creek at | Gri | Dinotoperlahirswe | 2 | 22 | |
McCallister | Gi jDimtoperlasp. — — LL [| 4 |
36.16371 S 14843515 E Gi [Жошы ` JL 13 | |
Elev. 1676 m asl. | Gi |Riekoperlahymsorum — — | 4 | 4 | |
Edge/Riffle Gri___| Riekoperlakarki | 2 | 2 | — |
Coll: G. Theischinger Gi — [Rikpelasp — — | — | — [| 53 |
Мо | Austrocercatasmanica —— | 1 | LI |
Ee РСА | TN | WNT [ШЕЛ ОЛЫ
DB2 23/11/2009 [ Eu [| Cosmioperlakuna | J |
Doubtful Creek at Cesjacks Gri | Dinotoperla Man | 4 | 6 |
36.14505 S 148.44429 Gri | Dinotoperla thwaitesi | | 1 |
Elev. 1650 m asl. Gri
Edge/Riffle Riekoperla sp.
Coll: G. Theischinger w е
|
=
—
A
Di
£
Q
S
š.
S
S
ç
š
Er
ES
š
à
i
=
s
Е
š
El
S
Q
>
š
š
š
8
3
5
DB3 23/11/2009
Doubtful Creek at Grey
Mares Trail
36.11081S 148.43168 E
Elev. 1540 m asl.
Edge/Riffle
Coll: G. Theischinger
o
=
S
Е
©
S
S
©
БЫ
=
š
=
ç
m omas O | GE EE
Tin Mine Creek at Hut Gri Dinotoperla brevipennis PWH J 2 J |
š Kee o 559Е агі Dinotoperla sp. EE
Se i
SE оп [ Rekpera s. Fes EUER Ense]
Coll: G. Theischinger
8 Australian Entomologist, 2011, 38 (1)
Location data Family Species á 9 L
| Eu [| Cosmioperlakuna | |
MURR 24112009 | Gri | Dinotoperla subserricauda | 1 |
Upper Murray at Cowambat
36.79249 S 148.16801E | бп | Dimooperlasp. | |
Elev. 1160 m asl. ой | Ийехоретаѕр УШЕШ
Edge/Riffle | Gi | Newmanoperlathoreyi | 1 |
Coll: G. Theischinger | Gri | Riekoperlakarki | 1 |
| Gri ]Rikoelargoa | 1 | 1 | |
[ Gi | Dinotoperla brevipennis | 3 | J |
[ Gi | Dinotoperlachristinae | | [| |
[ Gi | Dinoroperlafontana | — | 1 [| |
| Gri a — — — | — [| — [| 1 |
L Gr — [Rikpelargoa | 2 | — [| — |
EECH
ee Eee Ee)
ING 24/11/2009
Ingeegoodbee River above
access road
36.73327 S 148.26489 E
Elev. 1140 m asl.
Edge/Riffle
Coll: G. Theischinger
Analyses
The MDS plots of site Stonefly assemblage similarities indicated that site
differences were better defined using species level adult stonefly data (shown
in Fig. 3) rather than the genus/family level larval data (Fig. 2). The plot of
stonefly larvae could not differentiate between sites TUM2 and TUM3,
SNO2 and SNO3, and DB2, TIN and ING.
Transform: Presence/absence
Resemblance: S17 Bray Curtis similari
2D Stress: 0.06
DB3
A
5025 ж dicky cooper
TUMI A doubtful
T TUM2 y tin mine
2 + TUM3 MURR с ingeegoodbee
DB2 *
SNO7 ING & Оо
° SNO4 TIN GEE1
°
À
DB1
DCK1
*
Fig. 2. MDS plot of Stonefly larvae assemblages for the study sites.
Australian Entomologist, 2011, 38 (1)
Transform: Presence/absence -
Resemblance: $17 Bray Curtis similarity
MURR
x
DCK2
A
ct) 082
O A
TUM2 + gn
TUMI
ING
ü
TIN
V
Fig. 3. MDS plot of Stonefly adult assemblages for the study sites.
20 Stress: 0.1 || river
[© snowy
О geehi
|+ tumut
||x murray
|| Ж. dicky cooper
1А. doubtful
у їп mine
* [D Ingeegoodbee
Transform: Presence/absence
Resemblance: S17 Bray Curtis similari
GEE1 pon
vui
+
TUM3 A
2
MURR DB1
x А
TIN
V
ING
D
2D Stress: 0.09
SNO3
o
SNO2
° SNO7
°
їп ingeegoodbee
DCK1
*
ME SNO4
o
Fig. 4. MDS plot of aquatic larvae, identified to species level where possible, for the
study sites riffle habitats.
The stonefly adults plot, however, identified all sites to be different and
differences between rivers to be generally greater than differences between
sites within each river. The exception to this was Dicky Cooper Creek, where
10 Australian Entomologist, 2011, 38 (1)
the two sites were dissimilar. Note that there was only one site in the Murray
River, Geehi River, Ingeegoodbee River and Tin Mine Creek.
MDS plots of the aquatic larval assemblages collected from the riffle and
edge habitats showed a similar general site pattern to the stonefly adult plot.
The plot of the riffle larval fauna is presented in Fig. 4. The plots of the full
larval assemblages were able to differentiate all sites.
Discussion
Previously, when the same Snowy Mountain sites were sampled for the
monitoring program in 2008, the study only included the collection of larvae.
Without adult stoneflies, only three stonefly taxa were able to be identified
(by consultants) beyond family level: Acruroperla atra, Cosmioperla (? as
Sternoperla) sp. and Austroheptura (? as Tasmanoperla) sp. By comparison,
the 2009 study identified 23 different Stonefly species at the Snowy Hydro
monitoring sites using adults, thus allowing analysis of differences between
sites and, more importantly to the study, identification of change over time.
It appears that the geographical and habitat information from the 2009 Snowy
Cloud Seeding trials, paired with confident identities for many stonefly
species (see Tables 1 and 2), more than make up for the extra effort of
collecting and identifying stonefly adults. The use of adult insect
identifications where larvae identifications are not feasible could well prove
to be an invaluable tool for the assessment of river health, biodiversity,
conservation and climate change projects.
Outcomes of the study of adult stoneflies from the Snowy Hydro monitoring
sites are:
Five Dinotoperla species were found coexisting simultaneously at the same
site (TUM2) and finding diagnostic characters of these species, not only
apparent in the structure of the male genitalia but also in colour and pattern of
the forewings of both sexes, provides information on their ecology and may
help future taxonomic and behavioural studies. Species, and particularly
species assemblages, have greater indicator value for river health than family
or generic-level data.
The presence in numbers of Riekoperla alpina in DCK2 (Dicky Cooper
Creek at 1504 m), its absence in DCK1 (same creek at 1770 m) where its
likely sister species Riekoperla hynesorum was collected in numbers, and the
presence of Riekoperla hynesorum again in all sampled Snowy River sites
(SNO2, SNO3, SNO4, SNOWO7), all different in size, flow and substrate
from Dicky Cooper Ck at both elevations, provides more than just very
interesting ecological information. Table 2 shows clear species changes with
elevation. It probably also enables taxonomists to make the first promising
step to help distinguish the larvae of the Riekoperla alpina group
morphologically without the need for expensive DNA analysis. The
altitudinal detail may become significant for studies of climate change.
Australian Entomologist, 2011, 38 (1) 11
Table 2. Distribution of Stonefly species from the Snowy Hydro Cloud Seeding Trial.
The sites are arranged with increasing elevation from left to right. The shaded boxes
with crosses indicate the presence of the species at the site.
Stonefly adults
Acruroperla atra x
Austroheptura illiesi x
Cosmioperla kuna
Dinotoperla brevipennis x x
Dinotoperla hirsute
Dinotoperla еиситЬепе x
Dinotoperla christinae x
Dinotoperla fontana x x HO S Xe» X
Dinotoperla subserricauda x
Dinotoperla thwaitesi XD o D ея x
Dinotoperla uniformis
Riekoperla alpine
Riekoperla karki x o AN
Riekoperla hynesorum x o ow zi
Riekoperla reticulate x x x
Riekoperla rugosa ХХ х
Riekoperla tuberculata x x
Trinotoperla montana x x x x x
Trinotoperla nivata x CHE
Austrocercella hynesi
Austrocercella ?tillyardi x XS
Austrocerca tasmanica XIX x
Stonefly larvae
x
Austroheptura sp. x х
Dinotoperla sp. XB EX Mee XX XX XX Уа x
x
x
Leptoperla sp. x
Riekoperla sp. x x Rio x» OX äist Kë 23
Gripopterygidae sp. x
Trinotoperla sp.
Illiesoperla sp. x
Notonemouridae sp x
Cosmioperla kuna
elevation m asl.
1140
1160 | *
1230]х x * x
1290
1310 |х >
1360 | х ж
1400 |x х
1504 |х х
1540 | х
1650 [х >
1650 |х х
1670 |х х
1676 | * х
1730 | *
1770 |х х
1830 |х х
12 Australian Entomologist, 2011, 38 (1)
Changes in species assemblages and dominance situations may be indicative
of any sort of pollution and may be sensitive enough to show changes relating
to climatic conditions. For the family Gripopterygidae, our lists show
dominance of Riekoperla species of the alpina group in high alpine sites,
together with dominance of Dinotoperla species and the presence of some
species of the Riekoperla tuberculata and rugosa groups in lower altitude
sites. The existence of the Riekoperla alpina group in high altitude sites
probably results from warm periods making available formerly glaciated
areas. On the other hand, the presence of Dinotoperla species and species of
the Riekoperla tuberculata and rugosa groups in the cooler uplands of
northern New South Wales and Queensland is probably an outcome of cooler
periods in the past, facilitating dispersal northward to suitable habitats.
Similar cases of speciation were discussed by Peters and Theischinger (2007)
for dragonflies and by Watson and Theischinger (1984) for dragonflies,
stoneflies and other aquatic groups.
The fact that the appearance of adults of certain stoneflies in some regions is
highly seasonal, as found in the genus Austrocercella in the Snowy
Mountains (Theischinger 1982), is worth investigating in more detail. More
than one survey a year may significantly increase the number of species per
habitat (biodiversity). Changes in altitudinal and seasonal occurrence over
time of these seasonal species may prove indicative for monitoring pollutants
and ecological and climatic change.
The distribution of the species across the sampled sites generally supports the
existing information on the poor dispersal abilities of stoneflies. The sites that
have the most similar faunal assemblages are geographically close on a river
continuum (e.g. the four Snowy River sites: Tables 1 and 2). The Geehi River
site GEEI is close to, and connected by water to, the Dicky Cooper site
DCK2. The two sites on Dicky Cooper Creek, DCK1 and DCK2, are close
geographically and connected by water but have dissimilar assemblages; a
possible cause could be a barrier to migration created by the waterfall
between the two sites. Further sampling of these sites into the future is
expected to add information on stonefly dispersal in the Snowy Mountains
region.
By sampling adult stoneflies at sites, a clear picture of the diversity of species
at individual sites can be formed. As such, a plea is made for the inclusion of
adult stonefly collecting together with standard AusRivas sampling wherever
stoneflies make up a significant part of the aquatic fauna, in particular
Australian alpine and montane sites.
II. Summary of the Australian Plecoptera Fauna
This could also lead to a renaissance of taxonomic and ecological stonefly
studies that, after a boom in the early 1980s, have been rather stagnant for the
past two decades. In order to stimulate taxonomic research, an updated
check-list of the Australian Plecoptera, with state distribution data, is
Australian Entomologist, 2011, 38 (1) 13
presented below (Table 3). Currently, the Australian Plecoptera fauna
comprises 198 valid species (and subspecies) in 26 genera and 4 families.
This list is followed by an outline of taxonomic research on the group
(including the primary taxonomic literature) from its beginning to the present.
Table 3. Check-list of the Australian stonefly species and their known distributions.
Legend: W = Western Australia, S = South Australia, T = Tasmania, V = Victoria,
N = New South Wales (including ACT), Q = Queensland.
Species Distribution
Acruroperla atra (Samal. 1921) +
Austroheptura campbelli Theischinger, 1993 +
+
+
Austroheptura illiesi Hynes, 1974
Austroheptura neboissi Illies, 1969
Austroheptura picta (Riek, 1973) Ee x
Austropentura hynesorum Theischinger, 1988 +
Austropentura victoria lllies, 1969 + +
Crypturoperla paradoxa Illies, 1969
Tasmanoperla larvalis (Illies, 1969)
Tasmanoperla thalia (Newman, 1839)
Cosmioperla australis (Tillyard, 1921)
Cosmioperla denise (Theischinger, 1983)
Cosmioperla kuna (Theischinger, 1983) +
Cosmioperla macrops (Theischinger, 1983)
Cosmioperla w. wongoonoo (Theischinger, 1983) +
Cosmioperla wongoonoo tropica (Theischinger, 1983)
Eusthenia costalis Banks, 1913 +
Eusthenia lacustris Tillyard, 1921 +
Eusthenia nothofagi Zwick, 1979 +
Eusthenia reticulata (Tillyard, 1921)
Eusthenia spectabilis Gray, 1832
Eusthenia v. venosa (Tillyard, 1921) +
Eusthenia venosa brachyptera (Tillyard, 1924) +
Thaumatoperla alpina Burns & Neboiss, 1957
Thaumatoperla flaveola Burns & Neboiss, 1957
Thaumatoperla robusta Tillyard, 1921
Thaumatoperla timmsi Zwick, 1979
Cardioperla diversa McLellan, 1971
Cardioperla edita Hynes, 1982
Cardioperla falsa Hynes, 1982
Cardioperla flindersi Hynes, 1982
Cardioperla incerta Hynes, 1982
Cardioperla lobata McLellan, 1971
Cardioperla media Hynes, 1982
Cardioperla nigrifrons (Kimmins, 1951)
Cardioperla spinosa Hynes, 1982
Dinotoperla arcuata Theischinger, 1982
Dinotoperla bassae Hynes, 1982 +
Dinotoperla brevipennis Kimmins, 1951 + +
Dinotoperla bunya Theischinger, 1982
Dinotoperla cardaleae Theischinger, 1982
Dinotoperla carnarvonensis Theischinger, 1982
Dinotoperla carpenteri Tillyard, 1921 +
Dinotoperla christinae McLellan, 1971 + +
dr
Dinotoperla cobra Theischinger, 1982
+++
+++
+
+++
++
+++++ +++ +++ ++
+
+
+++
+++ +++
14 Australian Entomologist, 2011, 38 (1)
Species EST
Dinotoperla dalrymple Theischinger. 1993 +
Dinotoperla dolichoprocta Theischinger, 1982
Dinotoperla duplex Theischinger, 1982
Dinotoperla eucumbene McLellan, 1971 +
Dinotoperla eungella Theischinger, 1982 +
Dinotoperla evansi Kimmins, 1951 +
Dinotoperla fasciata Tillyard, 1921 +
Dinotoperla fontana Kimmins, 1951 + +
Dinotoperla hirsuta McLellan, 1971 + +
Dinotoperla hybrida Theischinger, 1984 +
Dinotoperla inermis Theischinger, 1988 +
Dinotoperla kirrama Theischinger, 1982 +
Dinotoperla leonardi Theischinger, 1982 + +
Dinotoperla marmorata Hynes, 1976 +
Dinotoperla opposita (Walker, 1852) +
Dinotoperla parabrevipennis Theischinger, 1982 +
Dinotoperla pseudodolichoprocta Theischinger, 1982 +
Dinotoperla schneiderae Theischinger, 1982 +
Dinotoperla serricauda Kimmins, 1951 r
Dinotoperla spinosa Theischinger, 1982 +
Dinotoperla subserricauda Theischinger, 1988 +
Dinotoperla thwaitesi Kimmins 1951 + +
Dinotoperla uniformis Kimmins, 1951 ? P
Dinotoperla vulcanica Theischinger, 1982 Ki
Dinotoperla walkeri Dean & St Clair, 2006 +
Dundundra wanungra (Theischinger, 1982). s
Eunotoperla kershawi Tillyard, 1924 +
Illiesoperla australis (Tillyard, 1924) a» TE
Illiesoperla austrosimplex Theischinger, 1984 +
Illiesoperla barbara Theischinger, 1984 E
Illiesoperla brevicauda Theischinger, 1984 dr. x
Illiesoperla carnarvonensis Theischinger, 1984
Illiesoperla cerberus Theischinger.1984
Illiesoperla echidna Theischinger, 1984
Illiesoperla franzeni (Perkins, 1958)
Illiesoperla frazieri Theischinger, 1984
Illiesoperla mayi (Perkins, 1958) + + +
Illiesoperla tropica Theischinger, 1984
Kirrama abolos Theischinger, 1981
Kirrama naumanni Theischinger, 1993
Leptoperla alata Theischinger, 1984
Leptoperla albicincta Theischinger, 1981 +
Leptoperla angularis Theischinger, 1981
Leptoperla australica (Enderlein, 1909) d
Leptoperla beroe Newman, 1839 +
Leptoperla bifida Mclellan, 1971 +
Leptoperla bubalus Theischinger, 1980
Leptoperla cacuminis Hynes, 1974
Leptoperla collessi Theischinger, 1981 1
Leptoperla commoni Theischinger, 1981 +
Leptoperla curvata Theischinger, 1980 deo
Leptoperla dahmsi Theischinger, 1984 +
Leptoperla kalliste Hynes, 1974 +
Leptoperla kimminsi McLellan, 1971 +
+++
+
++++
+о+++++
+++
Australian Entomologist, 2011, 38 (1)
Species
Leptoperla longicauda Theischinger. 1988
Leptoperla magnicauda Theischinger, 1981
Leptoperla membranosa Theischinger, 1988
Leptoperla neboissi McLellan, 1971
Leptoperla primitiva McLellan, 1971
Leptoperla rieki Theischinger, 1981
Leptoperla rotunda Theischinger, 1984
Leptoperla rubiconis Theischinger, 1984
Leptoperla smithersi Theischinger, 1981
Leptoperla tasmanica Kimmins, 1951
Leptoperla thompsoni Theischinger, 1988
Leptoperla truncata Theischinger, 1980
Leptoperla uptoni Theischinger, 1981
Leptoperla varia Kimmins, 1951
Neboissoperla alpina McLellan, 1971
Neboissoperla monteithi Theischinger, 1982
Neboissoperla spinulata Theischinger, 2002
Nescioperla curtisae Theischinger, 1982
Newmanoperla exigua (Kimmins, 1951)
Newmanoperla hackeri McLellan, 1971
Newmanoperla prona Hynes, 1982
Newmanoperla thoreyi (Banks, 1920)
Riekoperla alpina McLellan, 1971
Riekoperla angusta Theischinger, 1985
Riekoperla barringtonensis Theischinger, 1985
Riekoperla citrea Theischinger, 1985
Riekoperla compressa Theischinger, 1985
Riekoperla cornuta Theischinger, 1985
Riekoperla darlingtoni (Illies, 1968)
Riekoperla elongata Theischinger, 1985
Riekoperla hynesorum Theischinger, 1985
Riekoperla intermedia Theischinger, 1985
Riekoperla isosceles Theischinger, 1985
Riekoperla karki McLellan, 1971
Riekoperla montana Theischinger, 1985
Riekoperla naso Theischinger, 1981
Riekoperla occidentalis Hynes & Bunn, 1984
Riekoperla perkinsi Theischinger, 1985
Riekoperla pulchra Hynes, 1982
Riekoperla reticulata (Kimmins, 1951)
Riekoperla rugosa (Kimmins, 1951)
Riekoperla serrata Theischinger, 1985
Riekoperla tillyardi McLellan, 1971
Riekoperla trapeza Theischinger, 1985
Riekoperla t. triloba McLellan, 1971
Riekoperla triloba regalis Hynes, 1982
Riekoperla tuberculata McLellan, 1985
Riekoperla williamsi McLellan, 1971
Riekoperla zwicki Theischinger, 1985
Trinotoperla comprimata Hynes, 1982
Trinotoperla groomi Perkins, 1958
Trinotoperla hardyi Perkins, 1958
Trinotoperla inopinata Hynes, 1982
Trinotoperla irrorata Tillyard, 1924
+++
+++++
+
+ ++++ + + ++
++
++
15
Trinotoperla maior Theischinger, 1982
16 Australian Entomologist, 2011, 38 (1)
Species —— STEE
Trinotoperla minima Theischinger. 1982
Trinotoperla minor Kimmins, 1951 + +
Trinotoperla montana (Riek, 1962) + +
Trinotoperla mouldsi Theischinger, 1982 +
Trinotoperla nivata Kimmins, 1951 + +
Trinotoperla sinuosa Theischinger, 1982 +
Trinotoperla tasmanica (McLellan, 1971) +
Trinotoperla woodwardi Perkins, 1958 +
Trinotoperla yeoi Perkins, 1958 + +
Trinotoperla zwicki McLellan, 1971
Austrocerca rieki Шіеѕ, 1975
Austrocerca tasmanica (Tillyard, 1924) +
Austrocercella alpina Theischinger, 1984
Austrocercella autumnalis Theischinger, 1984
Austrocercella christinae Illies, 1975 +
Austrocercella columbae Hynes, 1981 +
Austrocercella c. communis Theischinger, 1984 + +
Austrocercella communis obtusa Theischinger, 1984
Austrocercella distans Theischinger, 1984
Austrocercella elevata Theischinger, 1984
Austrocercella forcipula Theischinger, 1984
Austrocercella hynesi Illies, 1975
Austrocercella i. illiesi Theischinger, 1984 +
Austrocercella illiesi tarraensis Theischinger, 1984 +
Austrocercella mariannae Illies, 1975 +
+
+
+
+
+++
+++
+++
Austrocercella nivalis Theischinger, 1984
Austrocercella tillyardi (Kimmins, 1951)
Austrocercella verna Theischinger, 1984
Austrocercella weiri Theischinger, 1984
Austrocercoides bullata (Kimmins, 1951)
Austrocercoides kondu Theischinger, 1993
Austrocercoides neboissi Illies, 1975 +
Austrocercoides tunta Theischinger, 1993
Austrocercoides zwicki Illies, 1975
Kimminsoperla albomacula (Kimmins, 1951)
Kimminsoperla biloba Illies, 1975
Kimminsoperla hystrix ЇШез, 1975
Kimminsoperla kaputaris Theischinger. 1980
Kimminsoperla mcalpinei Theischinger, 1981
Kimminsiperla neboissi Theischinger, 1988 d
Kimminsoperla williamsi Шіеѕ, 1975 +
Notonemoura lynchi Illies, 1975 +
Notonemoura maculata (Weir, 1967) cme cur
Tasmanocerca bifasciata (Kimmins, 1951) +
Total number of species group taxa = 198 з 9 Sg 3D Ut G
++
+ +++++
++++
+++
Historical perspectives
The taxonomic history of Australian Plecoptera started with the description
of Eusthenia spectabilis from Tasmania (Gray 1832). Additions of new
species mostly came from contributions of one or two species each by
Newman (1839), Walker (1852), Enderlein (1909), Banks (1913, 1920),
Samal (1921), Burns and Neboiss (1957), Riek (1962, 1973), Weir (1967)
Australian Entomologist, 2011, 38 (1) 17
and Hynes and Bunn (1984). Early family/genus revisions allowed Tillyard
1921, 1924) to add twelve, Perkins (1957) to add six and Kimmins (1951) to
establish 18 more species. Revisions of all families, undertaken by eminent
overseas plecopterists between 1968 and 1982, provided 15 additional
species described by Illies (1968, 1969, 1975), 19 by McLellan (1971), two
by Zwick (1979) and 17 by Hynes (1974-1982). Theischinger (1980-2002)
added a further 94 species, mainly in generic revisions based on personal
collecting in the field (mainly New South Wales and Queensland) and from
museum holdings. The concluding 198th Australian species (Dinotoperla
walkeri) resulted from river health studies and was added by Dean and St
Clair (2006), suggesting a closer synergy between taxonomy and
environmental studies might be established. As recently as 1996, McLellan
established the genus Cosmioperla for the Australian species previously
included in Stenoperla McLachlan.
Theischinger (94)
Hynes (18)
number of species
8
8
McLellan (19)
Wes (15)
Kimmins (18)
Tillyard (12)
1830 1850 1870 1890 1910 E 1950 1970 1990 2010
year
Fig. 5. Historical record of establishment of species-group taxa in Australian
Plecoptera. In the above historical outline, taxa regarded as synonyms are not
included. Vertical lines mark the year of single publications, horizontal lines indicate
the time span between first and last publications of the author and numbers in brackets
indicate the number of new species added to the fauna.
Larval taxonomy was seldom extensively covered in revisions or descriptive
papers. There are, however, two comprehensive treatments by Hynes (1978)
and Yule (1997) of the larvae from Victoria, and New South Wales and
northern Victoria, respectively, and a more specialised publication (Tsyrlin
2001) which included only a key to the Australian stonefly genera and to the
species of Leptoperla from Victoria. An ‘Illustrated Guide to the Adults of
the Australian Stoneflies (Plecopteray, including brief descriptions and
18 Australian Entomologist, 2011, 38 (1)
illustrations of the adults of all Australian species, was presented by
Theischinger and Cardale (1987). Based largely on this publication, the then
known Australian stonefly fauna was catalogued by Michaelis and Yule
(1988).
Acknowledgements
Access to the remote sites was made possible by Snowy Hydro. Specifically,
John Denholm arranged for helicopter access to sites and accomodation for
the field team. The helicopter pilot, Colin De Pagter, assisted with sampling
at the sites. Theresa Dye of Cardno Pty Ltd provided information on site
locations and previous samplings. Klaus Koop of DECCW facilitated the
sampling program. The manuscript benefited greatly from critical comments
from Dr D.J. Bickel (Sydney), Dr К. Marchant (Melbourne), Dr A.L. Sheldon
(Crawfordville, Florida, USA) and last but not least from the editorial input
of Dr A.G. Orr. The contributions of these people are gratefully
acknowledged.
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Australian Entomologist, 2011, 38 (1): 21-28 21
A NEW SPECIES OF FORESTER MOTH FROM VICTORIA
(LEPIDOPTERA: ZYGAENIDAE: PROCRIDINAE)
AXEL KALLIES! and BERNARD MOLLET?
!The Walter and Eliza Hall Institute, 1G Royal Parade, Parkville; Vic 3050
(Email: kallies@wehi.edu.au)
216, Parc Vatonne, 91190, Gif-sur-Yvette, France
(Email: molleth@wanadoo fr)
Abstract
A new species of forester moth, Pollanisus marriotti sp. n., is described from Victoria. It is
similar to Pollanisus angustifrons Tarmann, 2004 and several closely related species from
northern Queensland but distinguishable by its size, the breadth of its head and by the structure
of the antennae. Pollanisus marriotti sp. n. is currently known from a single locality in the
vicinity of Gembrook, east of Melbourne, where it occurs together with Pollanisus lithopastus
Turner, 1926, from which it can be readily distinguished by the coloration of the abdomen and
the size of the head and compound eyes.
Introduction
The Zygaenidae of Australia were revised recently by Tarmann (2004). The
most speciose genus is the Australian endemic Pollanisus Walker, with
currently 20 species described. Pollanisus species are distributed mainly
along the eastern and south-eastern coast of Australia and Tasmania, with
relatively few species occurring in South and Western Australia.
Despite Tarmann’s (2004) revision, a relatively large number of species
remain unnamed due to the lack of sufficient material. This is especially true
in the case of Pollanisus from the northern part of the genus’ range. The
species from south-eastern Australia, in particular from Victoria and southern
New South Wales, however, have been collected extensively and can be
considered well-known; the discovery of a previously unknown species of
Pollanisus in the vicinity of Melbourne therefore came as a surprise. This
new species belongs to a group of species known previously only from the
tropical and sub-tropical north-east of Australia, and is clearly distinct from
all other species that occur in Victoria or New South Wales.
Methods
The terminology of the setal combination of the first abdominal segment of
the first instar larvae follows Efetov et al. (2000) and describes the position,
number and colour of setae. Abbreviations used in the description as are
follows: D (dorsal), SD (subdorsal), L (lateral), / (light), d (dark).
Material
The acronyms of the following depositories are given in parentheses: ANIC -
Australian National Insect Collection, CSIRO Entomology, Canberra,
Australia; CAKM -— Collection of Axel Kallies, Melbourne, Australia;
CBMG - Collection of Bernard Mollet, Gif-sur-Yvette, France.
22 Australian Entomologist, 2011, 38 (1)
Figs 1-10. Pollanisus spp. and host plant. (1-2) P. marriotti sp. п.: (1) male
(holotype), dorsal; (2) female (paratype), dorsal. (3-4) P. lithopastus: (3) male, dorsal;
(4) female, dorsal. (5) P. angustifrons (paratype) male, dorsal. (6-7) P. marriotti sp.
n., male: (6) head, ventral; (7) abdomen, dorsal. (8-9) P. lithopastus, male: (8) head,
ventral; (9) abdomen, dorsal. (10) Hibbertia empetrifolia (DC.) Hoogland, the
assumed host plant of P. marriotti sp. n. at the type locality, Gilwell Park, Gembrook,
Victoria.
Australian Entomologist, 2011, 38 (1) 23
Pollanisus marriotti sp. n.
(Figs 1, 2, 6, 7, 11-15)
Types. Holotype 3 (Fig. 1): ‘Australia, Victoria, E of Melbourne, Gembrook, Gilwell
Park, $37°26’ E145?39', 3.11.2008, lux, leg. A. Kallies & P. Marriott’ (to be deposited
in ANIC). Paratypes: 1 3, same data as holotype but 10.1.2008 (P. Marriott)
(CAKM); 2 é é, 1 9 (GP698), same data as holotype but 3.11.2008 (A. Kallies & Р.
Marriott) (CBMG); 2 3, 1 9, same data as holotype but 6.11.2009 (A. Kallies & A.
Young) (CAKM); 2 55, same data as holotype but 27.1.2009 (A. Kallies, S. & B.
Mollet) (CBMG); 4 55 (GP697), 1 9 (Fig. 2), same data as holotype but 8.11.2009
(A. Kallies, S. & B. Mollet) (CBMG); 1 $, same data as holotype but 7.11.2009 (M.
Марр) (CAKM); 3 AO. same data as holotype but 2.11.2010 (A. Kallies, P. Marriott &
M. Hewish) (CAKM); 1 d. same data as holotype but 21.11.2010 (A. Kallies)
(CAKM).
Etymology
marriotti — a noun in the genetive case: this new species is dedicated to Peter
Marriott, Bentleigh, Victoria, main author of the *Moths of Victoria book
series, who collected the first specimen.
Description
Male (Fig. 1). Forewing length: 8.5-9.5 mm. Head dorsally dark brown with a
bluish green sheen, with metallic green frontal scales and a narrow stripe of
shiny bluish green scales running along the margin of the black compound
eye; frons ca 1.2 x broader than breath of compound eye in frontal view,
significantly protruding beyond compound eyes in both lateral and dorsal
view; labial palps light brown without metallic scales; proboscis yellow to
light brown; distance of ocellus from compound eye 0.8 x diameter of
ocellus; chaetosemata dark brown, the anterior extension overreaching
ocellus and completely covering the space between compound eye and
ocellus. Antenna dark brown with a weak bluish green sheen on dorsal side
of the shaft; segments 1 to 29-31 bipectinate, segments 30-32 to 39-42
biserrate; at segment 15 pectination 6-6.5 x longer than breath of shaft in
dorsal view and 4-4.5 x longer at segment 25; sensory hairs on pecten very
short. Collar with bright metallic golden green scales.
Thorax dark brown with a weak bluish green sheen dorsally and with shiny
metallic bluish green scales laterally and ventrally; patagia dark brown with
golden green scales proximally. Legs dark brown, femur of hind leg metallic
green blue laterally. Abdomen dark brown on the first segment and mostly
metallic golden green dorsally on other segments, dark brown with a very
weak bluish green sheen ventrally.
Wings broad, forewing elongate triangular, hindwing almost rectangular,
rounded apically, posterior margin straight; forewing upperside dark brown
with a weak bluish green sheen and a patch of green metallic scales near the
cell (on fresh specimens); underside dark grey-brown without metallic scales;
hindwing upperside dark brown with a slightly translucent space between
24 Australian Entomologist, 2011, 38 (1)
veins CuP and Cu2; underside dark brownish grey with metallic bluish green
scales in a band between cell and costa and at anal angle. Fringe blackish
brown with a weak sheen.
Female (Fig. 2). Forewing length: 7-7.5 mm. Similar to male but with
narrower and more rounded wings. Antenna biserrate. Abdominal hair tuft
bright yellow.
Male genitalia (Figs 11-12). Valva pointed distally, slightly convex dorsally,
folded translucent central part triangular, ventral sclerotization broad, ventral
margin of valva straight with a proximal part lobed, saccus strongly
sclerotized. Aedeagus slightly tapered and upcurved, ca 4.5 x longer than
broad; cornutus large and slender, straight, pointed distally, its length ca 80%
of aedeagus.
Female genitalia (Figs 13-14). Sternite VIII not sclerotized, ductus bursae
short, translucent, wall of corpus bursae near point of insertion of ductus
bursae with a sclerotization bearing 2 small teeth. Ductus seminalis arising
near lumen of corpus bursae.
Description of the first instar larva
The L1 (first instar) is cream coloured and about 1 mm in length. It has a
combination of three anal combs, an arrangement known for the genus
Pollanisus and other genera of the tribe Artonini (Mollet and Tarmann 2010).
There are no brown lateral spots on the subdorsal part of the third thoracic
segment and on the second and fifth abdominal segments visible. The setal
formula of the first abdominal segment is: D: 1d; SD: 1d, 1/; L: 21. This
appears characteristic of the Artonini (Mollet and Tarmann 2010).
Diagnosis
Pollanisus marriotti sp. n. is similar and appears to be most closely related to
P. angustifrons Tarmann, 2004, P. eungellae Tarmann, 2004, P. eumetopus
Turner, 1926, P. acharon (Fabricius, 1775) and a number of unnamed
species, all of which occur in the northern parts of Queensland and the
Northern Territory. These all belong to a group of species characterized by
their dark brown forewings with sparse metallic scales, their bright metallic
green collar and a conspicuous green metallic mark that gradually widens
from a point at the centre of the 2nd tergite and covers most of the caudal part
of the abdomen. Within this group, P. marriotti sp. n. is remarkable in that it
occurs outside of the tropics in temperate sclerophyll forest in Victoria. P.
marriotti sp. n. can be distinguished from all other species of the group by its
relatively wider wings and larger size. In particular, P. angustifrons (Fig. 5)
is smaller, has a wider head, more extended pectination of the antennae (Fig.
17) and the forewings are relatively narrower. Pollanisus eungellae, P.
eumetopus and P. acharon are smaller and their forewings are relatively
narrower; P. eungellae and P. eumetopus have a broader frons.
Australian Entomologist, 2011, 38 (1) 25
11
\
13
| 4 0.1 mm
15
17
Figs 11-17. Pollanisus spp. (11-14) P. marriotti sp. n., genitalia: (11) ventral view of
male genitalia, aedeagus removed, left valve omitted; (12) aedeagus; (13) overview of
female genitalia, praebursa with spermatophore; (14) sclerotization at insertion of
ductus seminalis. (15-17) Pollanisus spp., male antenna with shaft, pectination on
only one side shown: (15) P. marriotti sp. n.; (16) P. lithopastus; (17) P. angustifrons.
26 Australian Entomologist, 2011, 38 (1)
Pollanisus marriotti sp. n. also shows similarities to P. subdolosa (Walker,
[1865]) and some related species. With these it shares the general
morphology such as wing and body shape and the distribution of metallic
scales on the abdomen. However, P. subdolosa and related species can be
readily distinguished by the colour of its abdomen and collar, which are
metallic copper-red, but never metallic green. Females of P. subdolosa also
differ by the lack of metallic scales on the abdomen, which are always
present in P. marriotti sp. n., by their relatively larger yellow anal tuft and by
their somewhat narrower wings. P. subdolosa has not been found at the type
locality of P. marriotti sp. n.; however, it occurs in other forests in the
vicinity of Melbourne and other parts of southern Victoria and is also
attracted to light. Furthermore, P. subdolosa has two generations per year
with adults being on the wing in November and December and again in
March, whereas P. marriotti sp. n. occurs only in a single generation in
summer.
Superficially, P. marriotti sp. n. is also similar to P. lithopastus Turner, 1926
(Figs 3-4) and both species occur syntopically at the type locality. P.
marriotti sp. n. differs from P. lithopastus as follows: head narrower, eyes
smaller and proboscis yellow (Fig. 6) (head wider, eyes larger and proboscis
dark brown in P. lithopastus, Fig. 8); dorsal side of the 1st abdominal
segment without metallic scales, with a conspicuous green metallic mark that
gradually widens from a point at the centre of the 2nd tergite and covers most
of the caudal part of the abdomen (Fig. 7) (dorsal side of abdomen
completely metallic blue in P. /lithopastus, Fig. 9); fore and hindwings
narrower (broader in P. /lithopastus); hindwings lighter (darker іп P.
lithopastus). Furthermore, the apical biserrate part of the antennae is
relatively longer in P. marriotti sp. n. (Figs 15-17).
The new species cannot be confused with any of the other Pollanisus species
(c.f. Tarmann 2004).
Phenology and bionomics
The only known locality of this species is a semi-dry to wet eucalypt forest at
about 300 m altitude with a rich understorey of Leptospermum, Banksia and
Hibbertia. The locality is a mosaic of slopes and wet gullies and harbours a
rich lepidopterous fauna, including many species that apparently reach their
most southern and western distribution limit in this area.
At the type locality, P. marriotti sp. n. shares its habitat with Hestiochora
furcata Tarmann, 2004 and P. lithopastus, the latter being very common in
this locality. The adults of P. marriotti sp. n. fly from early January to late
February, whereas P. lithopastus has a longer flight period from late
November to early March. Although both species are also likely to be active
during the day, most specimens were observed or collected at the light
between 22.00h and 24.00h. Up to 50 specimens of P. lithopastus were
Australian Entomologist, 2011, 38 (1) 27
attracted to the light on warm and dark nights and hundreds were observed
over the course of the flight period, all but two being males. P. marriotti sp.
n. is comparatively rare, with typically only a few specimens attracted to the
light during one night. Despite extensive searching during the day, few
specimens of P. marriotti sp. n. were found, two flying at about 5 pm and one
male on the flowers of Leptospermum at about 10.00h. Despite extensive
fieldwork around Melbourne by one of us (AK), P. lithopastus was observed
only once during daytime, when a female was found resting on a grass stem
in the afternoon.
The hostplant of both P. lithopastus and P. marriotti sp. n. in Gilwell Park is
likely to be the Trailing Guinea-flower, Hibbertia empetrifolia (DC.)
Hoogland (Fig. 10). Larvae that hatched from eggs obtained from females of
both species started feeding on the leaves of this species. Larvae of P.
marriotti sp. n. also accepted Hibbertia scandens (Willd) Dryand as a
surrogate, but died subsequently, whereas the larvae of P. lithopastus refused
this plant. P. lithopastus seems to utilize different Hibbertia species as
hostplants as it can be common in places where Hibbertia empetrifolia is
apparently absent.
Distribution
Despite recent intensive collecting around Melbourne and occasional activity
in parts of Gippsland and East Gippsland, P. marriotti sp. n. was only found
at the type locality. However, it can be assumed that other colonies of this
species exist in sheltered coastal forests east of Melbourne. P. lithopastus, on
the other hand, is relatively widespread east of Melbourne and also occurs in
the Otway Ranges and near Nelson in the west of Victoria.
Conservation status
Pollanisus marriotti sp. n. is currently known only from a single locality,
Gilwell Park, in the vicinity of Gembrook. This is remarkable as light
trapping was frequently conducted in similar forests east of Melbourne.
Whereas P. lithopastus was found in most of these locations, P. marriotti sp.
n. was not. A similar distribution pattern was observed for various other moth
species (Kallies and Marriott, unpublished observations). Although the
reasons for the restricted distribution of these species are unknown, we
speculate that lack of fire is a major factor. In an area well known for its high
bushfire risk, Gilwell Park has escaped widespread fire damage for over 80
years and was not subject to ‘controlled’ back burning. This lack of fire may
well be the key factor for the high diversity observed in this area.
Although parts of Gilwell Park are frequently used by Scouts and other
groups for camping and other outdoor activities and the vegetation is
controlled by regular slashing, other parts of the park are relatively
undisturbed. Importantly, these activities do not seem to have obvious
negative impacts on plant and insect diversity. This shows that use of forest
28 Australian Entomologist, 2011, 38 (1)
for recreational activities when carefully managed is fully consistent with
protection of a diverse fauna and flora. It furthermore underlines the
importance of woodland and forest protected from bushfires and back
burning as refugia that ensure long-term survival of a diverse array not only
of birds and mammals but also of butterflies, moths and other insects. Gilwell
Park is a prime example of such a place. P. marriotti sp. n., due to its
restricted distribution and its potential susceptibility to fire, should be
considered a threatened species.
Acknowledgements
We would like to express our gratitude to Peter Marriott who collected and
provided the first specimens of this interesting new species, to Murray Vagg
who helped us with our fieldwork, and to the Gilwell Park authorities for
allowing us to conduct research on their premises and for allowing us to use
the facilities of the Park. We furthermore would like to thank the Victorian
Department of Sustainability and Environment for providing appropriate
research permits.
References
EFETOV, K.A., KEIL, T., MOLLET, B. and TARMANN, G.M. 2000. New data on the
chaetotaxy of the first instar larvae of Forester moths (Lepidoptera: Zygaenidae, Procridinae).
Nachrichten des entomologischen Vereins Apollo (N. F.) 21: 83-90, figs 1-34.
MOLLET, B. and TARMANN, G.M. 2010. Notes on the ecology, phenology, and distribution of
Pollanisus eumetopus Turner (Lepidoptera: Zygaenidae, Procridinae, Artonini). Australian
Entomologist 37(2): 63-67.
TARMANN, G.M. 2004. Zygaenid moths of Australia: A revision of Australian Zygaenidae
(Procridinae: Artonini). CSIRO Publishing, Collingwood; 248 pp, 64 col. pls, 448 text-figs, distr.
maps.
TURNER, A. 1926. Revision of Australian Lepidoptera: Drepanidae, Limacodidae, Zygaenidae.
Proceedings of the Linnean Society of New South Wales 51: 437-445.
Australian Entomologist, 2011, 38 (1): 29-36 29
THE NOMENCLATURE OF OGYRIS HALMATURIA (TEPPER, 1890)
(LEPIDOPTERA: LYCAENIDAE)
M.F. BRABY!2*, Е. DOUGLAS? and R.C. WILLAN!
!Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin, NT 0801
2Research School of Biology, The Australian National University, Canberra, ACT 0200
?PO Box 37, Rainbow, Vic 3424
*E-mail: michael.braby@nt.gov.au
Abstract
The purpose of this paper is to clarify and resolve the nomenclature of Ogyris halmaturia
(Tepper, 1890), a nationally threatened butterfly which has had a long and complex
nomenclatural history. This complexity has arisen because: (1) the species group name
halmaturia was based on a mixed series comprising two different species; (2) historically at least
six authors have attempted to resolve the nomenclature of halmaturia, but most failed to render a
valid and unambiguous lectotype designation; (3) one of these authors (N.B. Tindale) made a
particularly confusing lectotype designation in 1923; and (4) introduction of the name Ogyris
waterhouseri (Bethune-Baker, 1905). The proposal to treat O. waterhouseri as a junior synonym
of O. halmaturia is accepted. We argue that Tindale made the first valid and unambiguous
lectotypification in 1923. Consequently, we propose, with the intention of bringing closure to
this matter, that O. halmaturia is the senior synonym of O. waterhouseri and that Tepper's
syntype *female' is the lectotype male of O. halmaturia. Attention is drawn to ambiguity in
Article 74.5 (lectotype designation made before 2000) in the most recent edition of the
International Code of Zoological Nomenclature.
Introduction
The Australian endemic butterfly Ogyris halmaturia (Tepper, 1890) (Eastern
Bronze Azure) is an endangered species of heathland and mallee-heathland
habitats in coastal and semi-arid areas of South Australia and (formerly)
western and south-western Victoria (Braby and Douglas 2008). Its taxonomic
status is now agreed upon as a valid species, but a consensus on its
nomenclature needs to be resolved urgently to effectively underpin
conservation efforts.
The species has had a long and complex nomenclatural history (see Braby
and Douglas 2008 for review). This complex nomenclature has arisen in at
least four different ways. The first stems from the fact that Tepper (1890) had
a mixed series comprising four syntypes representing two species (Table 1).
The type specimens were all collected from near Kingscote (given as
‘Queenscliffe’), Kangaroo Island, South Australia, on 20-21 November 1886,
viz: three males of O. otanes (C. & R. Felder, 1865) and a male of O..
halmaturia, which Tepper incorrectly assumed represented the female of his
new species. Because Tepper (1890) did not designate a type specimen or
make reference to a type of any sort, a taxonomist must therefore determine
which specimen of Tepper's type material (i.e. his syntypic series) represents
the name-bearing ‘type’ in order to fix the name О. halmaturia to the species
in question.
30 Australian Entomologist, 2011, 38 (1)
According to Article 74 of the ICZN (1999), the fixation of a name from
syntypes is dependent on the designation of a lectotype; that specimen then
becomes the unique bearer of the name of the nominal species group taxon
and the standard for its application. And here lies the second issue, which is
at the core of this complex problem.
In historical times, at least six authors (Lower 1893, Tepper 1893,
Waterhouse 1903а, 1903b, Bethune-Baker 1905, 1916, Waterhouse and Lyell
1914, Tindale 1923) have attempted to resolve the taxonomy of O.
halmaturia but few attempted to do so definitively by making a lectotype
designation. Lower (1893) placed Tepper's male O. halmaturia under O.
otanes and Tepper's ‘female’ О. halmaturia in synonymy with O. idmo
(Hewitson, 1862) but did not refer to ‘types’ of any form. Tepper (1893)
himself maintained O. halmaturia as a species distinct from O. otanes and O.
idmo, and restricted its distribution to Kangaroo Island (and possibly on the
mainland at Port Lincoln on the tip of Eyre Peninsula, SA), but he did not
refer to type material and still failed to realise that he had a mixed series.
Waterhouse (1903b p. 249) concurred with Lower (1893) and listed O.
halmaturia as a synonym of О. otanes and remarked that ‘I almost certainly
agree with Lower who says that Tepper's O. halmaturia comprises O. otanes
3 and О. idmo 4’. He subsequently treated О. halmaturia as a synonym of
both О. otanes and О. idmo in his catalogue of Australian butterflies
(Waterhouse 19032), but again did not examine or make reference to a single
‘type’ specimen that would bear the name. Waterhouse and Lyell (1914) later
subsumed O. halmaturia under O. otanes in an attempt to resolve the
taxonomy, but did not examine type material to clear up the nomenclature.
Bethune-Baker (1905, 1916) and Tindale (1923) did, however, both refer to
‘types’, which we expand upon below. But here lies the third part of the
problem: Tindale's (1923) type designation was confusing in that he partly
synonymised Tepper's concept of O. halmaturia under O. otanes. Lastly, 15
years after the species was formally described, another name, О.
waterhouseri (Bethune-Baker, 1905), was introduced for it.
In attempting to resolve this complex nomenclatural problem, Braby and
Douglas (2008) traced and critically examined the type series (= syntypes) of
O. halmaturia (currently housed in the South Australian Museum (SAM) and
The Natural History Museum, London (BMNH)), reviewed the historical
literature and concluded that the name O. halmaturia is a junior synonym of
О. otanes, and consequently that Tepper's syntype of the second species is a
paralectotype of O. halmaturia. In contrast, Field (1999), and more recently
Grund (2010), proposed that O. waterhouseri is a junior synonym of O.
halmaturia. Grund's argument was based on three lines of evidence: (1) that
of Tepper's (1890) description of O. halmaturia and his original intention;
(2) crediting Tindale (1923) as the first taxonomist to validly select a
lectotype of O. halmaturia; and (3) espousing the premise of nomenclatural
stability and the presumption that the name waterhouseri was interfering with
Australian Entomologist, 2011, 38 (1) 31
Table 1. Tepper's (1890) syntypic series of Ogyris halmaturia and their type status.
MM M M M gege
Specimen, Complete label data, Repository, Type status and Current valid species name
— 1
9 O. halmaturia
“Queenscliffe, 1 mile N.W. very shy, 9, 20.11.86. Tepper” [in Tepper’s original handwriting],
“Ogyris halmaturia Tepper, Type female = 5, Kangaroo Island | n348, vide, TRSSA. 1923”,
“SAMA Database No. 31-001699"
SAM. Lectotype. О. halmaturia д
3 О. halmaturia
“Queenscliffe, @, 1 m. N.W. very shy, 21.11.86. Tepper” [in Tepper's original handwriting],
*Ogyris halmaturia Tepper, Type male, — not type, Kangaroo Island | vide TRSSA 1923, p.
389", “Ogyris otanes 2 not halmaturia", “SAM Database No. 31-001700”
SAM. Paralectotype. О. otanes д
3 О. halmaturia
“Queenscliffe, 1 m. N.W. very shy, 20.11.86. Tepper" [in Tepper's original handwriting],
*Ogyris halmaturia Tepper, Cotype male, Kangaroo Island | vide TRSSA 1923, p. 389",
“Ogyris otanes ĝ not halmaturia", “SAM Database No. 31-001701”
SAM. Paralectotype. О. otanes д
3 О. halmaturia
“Queenscliffe, 1 mile N.W., in scrub. $, 20.11.86. Tepper” [in Tepper’s original handwriting],
“Ogyris halmaturia, Queenscliffe, Kang. Island, Nov. 1886., legit J.G.O. Tepper”, “Bethune-
Baker Coll. B.M. 1927-471."
BMNH. Paralectotype. O. otanes 3
common usage. We discuss each of these components of evidence in turn and
show that, while the hypothesis to treat O. halmaturia as the senior synonym
is supported, two of Grund’s (2010) arguments are misguided on
nomenclatural grounds.
Tepper’s description of O. halmaturia
Grund (2010) argued that Tepper (1890) intentionally gave first priority in his
description to the ‘female’ of О. halmaturia (= 3 О. halmaturia) because
Tepper made reference to its similarity with the underside of O. oroetes
(Hewitson, 1862). ‘This was the normal way of describing new species
during this historical time period, viz. describing the important reference
specimen first, be it male or female’ (Grund 2010 p. 115). However, Tepper
(1890) actually made no reference to the underside of O. oroetes, he just
stated ‘It comes nearest to O. oroetes, Hew., but differs from various details
from Hewitson’s figure’. More importantly though, Tepper (1890) did not
describe the ‘female’ first, but in fact described the two supposed sexes
together: after first giving approximate size measurements of the ‘female’
and male, he then described in some detail the upperside of both sexes (our
emphasis) simultaneously’; he then proceeded to describe the underside of
the ‘female’ and then the underside of the male. Whether Tepper (1890)
intended to give priority to the ‘female’ of O. halmaturia or not is irrelevant
in terms of nomenclature under the ICZN (1999) because Tepper (1890,
1893) did not refer to the specimens before him at the time of description (i.e.
32 Australian Entomologist, 2011, 38 (1)
his type material). As noted above, in such cases where there are two or more
syntypes, especially where two or more species are involved, a lectotype
must be selected from the type series in order to fix the name of the nominal
species group taxon (Article 74.1) (ICZN 1999).
Designation of lectotype of O. halmaturia
Article 74.5 of the ICZN (1999) stipulates that *In a lectotype designation
made before 2000, either the term ‘lectotype’, or an exact translation or
equivalent expression (e.g. ‘the type’), must have been used or the author
must have unambiguously selected a particular syntype to act as the unique
type of the taxon.' That is, the Code appears to recognise three different
situations or criteria under which a lectotype may be designated. In this case,
three authors (Bethune-Baker 1905, 1916, Tindale 1923, Field 1999)
potentially qualify as having undertaken acts of lectotypification under this
ruling.
In his revision of the genus Ogyris, Bethune-Baker (1905 pp 276-277)
remarked under the taxon O. otanes that ‘Mr. Waterhouse has kindly sent me
for examination two specimens from Kangaroo Island with a query as to
whether they are Felder's insect, but after a very careful comparison I believe
them to be distinct, and they are the form named by Tepper halmaturia. І
have now before me the type of this species as well as Felder's type (I must
here express my best wishes to Mr. Tepper for the loan of it); and I consider
that they are distinct forms; more material may prove them to be sub-species,
but they differ sufficiently to warrant them being named.' In other words,
Bethune-Baker (1905) is saying that O. otanes (from the South Australian
mainland) and O. halmaturia (from Kangaroo Island) are closely related
species, but further research may prove them to be conspecific. Indeed, he
later remarked *... in the closing sentence of p. 277 of my monograph I
broadly hint at the possibility of halmaturia being a form of otanes, Felder,
and I am quite willing to concede it as a race of that species? (Bethune-Baker
1916 p. 390). This later statement was made in response to comments by
Waterhouse and Lyell (1914), who did not recognise O. halmaturia,
subsuming it (ie. the Kangaroo Island population) under the Species O.
otanes. Bethune-Baker (1905) was unaware of the fact that Tepper had a
mixed series until much later (see Bethune-Baker 1916); he examined only
one of Tepper's syntypes (= @ О. otanes) and referred to that specimen as a
‘type’, “Т regard Tepper's species as distinct from otanes, Feld., both of
which types are now before me’ (Bethune-Baker 1905 p. 275) and ‘I only had
the 3 type of this insect before me’ (Bethune-Baker 1916 p. 390).
Edwards et al. (2001) interpreted Bethune-Baker’s (1905) reference to a type
as an intentional and valid lectotype designation; however, Braby and
Douglas (2008) and Grund (2010) did not consider this to be the case because
Bethune-Baker (1905 p. 277) used the term ‘type’ rather vaguely to describe
all the syntypes of O. halmaturia, ‘The types from Kangaroo Island are in the
Australian Entomologist, 2011, 38 (1) 33
S. Australian Museum. Mr. Waterhouse also has specimens from the same
locality.” and as such he did not intentionally or explicitly make a formal type
designation, at least not one that could be deemed *unambiguous' in the sense
of Article 74.5. Although there is some uncertainty in the current edition of
the ICZN (1999) in relation to Article 74.5, particularly with interpretation of
the second criterion ‘an exact translation or equivalent expression (e.g. ‘the
type’), must have been used’ in a lectotype designation made before 2000,
many taxonomists would interpret the use of the word “уре” by Bethune-
Baker for one of Tepper's syntypes to be construed as a valid lectotype
designation by inference, provided the specimen could be identified and
unambiguously located. The specimen is currently located in the BMNH and
was identified and illustrated by Braby and Douglas (2008 Figs 7-9 p. 319),
who considered it to be a paralectotype of O. halmaturia. On the other hand,
an alternative interpretation of Article 74.5 is that a lectotype designation
must satisfy all three criteria: that of being intentional, unambiguous and
based on a single or unique type specimen (C. Thompson pers. comm. 2010).
Although it is clear that Bethune-Baker (1905) had only one of Tepper's
syntypes available to him, and on two occasions in that publication he
referred to that specimen as the “уре” (see also remarks by Tindale 1923), he
was not intentionally selecting that specimen among the type series to be the
unique type specimen and, moreover, he used the word “type” in different
senses, both in the singular and plural. Bethune-Baker (1916) again referred
to that specimen as ‘the male type’, but this was prefixed by the phrase ‘I
only had...’ (our emphasis), indicating that he was aware of other ‘types’ (i.e.
Tepper's syntypic series); hence, here again there is ambiguity as to whether
or not he was intending the loaned syntype to be the primary type of
halmaturia.
We now reconsider the work of Tindale (1923) because this was the second
line of evidence used by Grund (2010) to synonymise the name О.
waterhouseri. Tindale's publication is interesting because of the confusing
way it was written. Tindale (1923 p. 347) considered O. halmaturia and O.
otanes to be conspecific and synonymised O. halmaturia under O. otanes in
part. Tindale also illustrated one of Tepper's male syntypes (in SAM) from
Kangaroo Island in Plate 24, Figure 16, and in the figure caption (p. 354)
referred to that specimen as “Ogyris halmaturia, Tepper, Type male,
Kangaroo Island = ofanes, Felder.’ Braby and Douglas (2008) considered
Tindale's (1923) action on p. 347 to be an intentional designation of a
lectotype on the basis that: (1) he referred to one of Tepper's syntypes as the
‘type’ in the figure caption (p. 354); and (2) he illustrated that specimen
(Plate 24), but these authors overlooked the fact that Tindale explicitly wrote
‘(part)’ at the end of the synonymy line. Tindale (1923) did the same for О.
halmaturia on the next page (p. 348) on which he redescribed and illustrated
Tepper's ‘female’ syntype in Plate 24, Figure 20 and referred to it as the
‘type’ of O. halmaturia in the figure caption (p. 354). Tindale (1923 p. 348)
34 Australian Entomologist, 2011, 38 (1)
stated ‘The type male is a typical specimen of O. otanes, Feld.; the ‘female’
is the male of a species very close to O. waterhouseri, Bethune-Baker and, as
in the original description, the *female' is mentioned and described first, the
name halmaturia will stand.’ Grund (2010) considered this action by Tindale
to be evidence in support of an intentional lectotype designation; that is,
Tindale deliberately selected the ‘female’ to be the primary type or name-
bearer of O. halmaturia. In considering this particular aspect we concur with
Grund (2010) of Tindale's intent; it is likely that Tindale (1923 p. 347) was
merely referring to, and illustrating, the syntype male of О. halmaturia (= 3
O. otanes) to show that it belonged to a different species, rather than
attempting to synonymise the whole of Tepper's concept of O. halmaturia
with O. otanes.
Tindale's (1923) action on p. 348 in which he explicitly refers to Tepper's
*female' type specimen therefore, in our opinion, fixes the name O.
halmaturia to the taxon. Because the lectotype of О. halmaturia is the same
species as the lectotype of O. waterhouseri, which was described 15 years
later, the species group name waterhouseri is therefore a junior synonym of
O. halmaturia. Accepting Tindale (1923 p. 348), rather than Tindale (1923 p.
347) or Bethune-Baker (1905, 1916), as the first taxonomist to validly and
unambiguously designate a lectotype for the nominal species group name
halmaturia, renders the subsequent action of Field (1999), who also
designated Tepper's “female? syntype as a lectotype of О. halmaturia, as an
incorrect subsequent lectotype designation. That is, once a lectotype has been
validly designated, all subsequent lectotypifications have no validity (Article
74.1.1) and, moreover, this designation permanently deprives all other
specimens that were formerly syntypes of that status in that they
automatically become paralectotypes (Article 74.1.3). Tepper's three syntype
males (= 3 О. otanes) thus all qualify as paralectotypes of О. halmaturia
(Table 1).
Nomenclatural stability
Grund (2010) argued that because the name O. halmaturia has been in usage
for the past 86 years (ie. since Tindale 1923) at one level or another it
qualifies for nomenclatural "protection" in some way. It is true that the name
has been in common usage for a long period, but it is also true that the name
O. waterhouseri has had continuous usage for an almost equally long period,
from 1905 to 1972 (67 years) (see synonymic list and review of literature in
Braby and Douglas 2008). In terms of actual usage, waterhouseri has
appeared unambiguously with full species status four times or as a subspecies
of O. idmo eight times; halmaturia has appeared unambiguously with full
species status only twice or as a subspecies of O. idmo 20 times; and once
they have appeared ambiguously with both names combined, as O.
halmaturia waterhouseri (Tindale 1923). Moreover, the name waterhouseri
has appeared unambiguously as valid in a major checklist of type material in
Australian Entomologist, 2011, 38 (1) 35
the Australian Museum, Sydney (Peters 1971) and in several books
(Common 1964, D'Abrera 1971, McCubbin 1971), as well as in the
perceptive paper by Quick (1972).
The ICZN (1999) does allow for the automatic reversal of precedence of a
long-unused senior synonym under its Reversal of Precedence provision
(Article 23.9), but then only under two strict conditions. If either of these
conditions cannot be met completely, then an author must refer the case to
the Commission of the ICZN for a formal decision requesting existing usage
of the junior synonym to be maintained instead of acting unilaterally. A
recent entomological example under this provision is that by Jendek (2007),
wherein the name Buprestris angustulata Illiger, 1803 was given precedence
by the ICZN (2009) over B. pavida Fabricius, 1793. There is no way this
provision by itself could be invoked to conserve halmaturia over
waterhouseri because halmaturia is not the junior synonym.
In closing, our hope in resolving and clarifying the nomenclature of Ogyris
halmaturia — that it is a senior synonym of O. waterhouseri based on
interpretation of Tindale (1923 p. 348) as making the first valid lectotype
designation — is that it will meet acceptance amongst the wider entomological
community, and that it will be the name adopted by students of Lepidoptera,
government agencies and non-government organizations in attempts to
improve the conservation status of the species, which ranks as one of
Australia’s most threatened butterflies (Braby and Douglas 2008). We deem
this more preferable than attempting to decipher confusion inherent in Article
75.4 and prepare an Application to the International Commission of
Zoological Nomenclature to reject O. halmaturia in favour of O.
waterhouseri.
Acknowledgements
We thank Pete Cranston and Penny Gullan (UC Davis, USA), Ted Edwards
(ANIC, Canberra), Tim New (La Trobe University, Melbourne), Graham
Brown (MAGNT, Darwin), Alan Beu (GNS Science, New Zealand) and
past/present ICZN Commissioners Chris Thompson (Smithsonian Institution,
USA), Bruce Halliday (ANIC, Canberra) and Gary Rosenberg (ANSP, USA)
for their wise council on this note.
References
BETHUNE-BAKER, G.T. 1905. A monograph of the genus Ogyris. Transactions of the
Entomological Society of London 1905: 269-292, pl XV.
BETHUNE-BAKER, G.T. 1916. Notes on the synonymy of the genus Ogyris. Annals and
Magazine of Natural History (8) 17(101): 386-390.
BRABY, M.F. and DOUGLAS, F. 2008. The nomenclature, taxonomy and conservation status
of Ogyris waterhouseri (Bethune-Baker, 1905) stat. nov. (Lepidoptera: Lycaenidae), a threatened
butterfly from southern Australia. Australian Journal of Entomology 47: 315-329.
36 Australian Entomologist, 2011, 38 (1)
EDWARDS, E.D., NEWLAND, J. and REGAN, L. 2001. Lepidoptera: Hesperioidea,
Papilionoidea. Zoological Catalogue of Australia. Vol. 31.6. CSIRO Publishing, Melbourne; x +
615 pp.
FIELD, R.P. 1999. A new species of Ogyris Angas (Lepidoptera: Lycaenidae) from southern
arid Australia. Memoirs of Museum Victoria 57: 251-259.
GRUND, R. 2010. The taxonomy of Ogyris halmaturia (Tepper, 1890) stat. nov. (Lepidoptera:
Lycaenidae). Australian Journal of Entomology 49: 114-120.
ICZN. 1999. International code of zoological nomenclature. International Trust for Zoological
Nomenclature, London; xxix + 126 pp.
ICZN. 2009. Opinion 2221 Buprestris angustulata Illiger, 1803 (currently Agrilus angustulatus,
Insecta, Coleoptera): specific name given precedence over the specific Buprestris pavida
Fabricius, 1793. Bulletin of Zoological Nomenclature 66: 98-99.
JENDEK, E. 2007. Buprestris angustulata Illiger, 1803 (Insecta, Coleoptera): proposed
precedence of the specific name over that of Buprestris pavida Fabricius, 1793. Bulletin of
Zoological Nomenclature 64: 178-181.
LOWER, O.B. 1893. List of South Australian Rhopalocera. Transactions, Proceedings and
Reports of the Royal Society of South Australia 17: 1-12.
PETERS, J.V. 1971. A catalogue of the type specimens of the Hesperioidea and Papilionoidea
(Lepidoptera) in the Australian Museum. Australian Entomological Press, N.S.W.; 36 pp.
QUICK, W.N.B. 1972. Collected Ogyris idmo waterhouseri. Victorian Entomologist 2: 9-10.
TEPPER, J.G.O. 1890. Common native insects of South Australia: a popular guide to South
Australian entomology. Part II. Lepidoptera, or butterflies and moths. Wigg & Son, Adelaide; iv
+ 65 pp.
TEPPER, J.G.O. 1893. Notes and remarks on South Australian Rhopalocera. Transactions,
Proceedings and Reports of the Royal Society of South Australia 17: 281-286.
TINDALE, N.B. 1923. On Australian Rhopalocera. Transactions and Proceedings of the Royal
Society of South Australia 47: 342-354.
WATERHOUSE, G.A. 1903a. A catalogue of the Rhopalocera of Australia. Memoirs of the New
South Wales Naturalists' Club 1: 1-49, i-ii.
WATERHOUSE, G.A. 1903b. Notes on Australian Rhopalocera: Lycaenidae. Part III
Revisional. Proceedings of the Linnean Society of New South Wales 28: 132-275.
WATERHOUSE, G.A. and LYELL, G. 1914. The butterflies of Australia. A monograph of the
Australian Rhopalocera. Angus and Robertson, Sydney; vi + 239 pp.
Australian Entomologist, 2011, 38 (1): 37-48 37
MATERNAL CARE, FOOD PLANTS AND DISTRIBUTION OF
AUSTRALIAN ONCOMERINAE (HEMIPTERA: HETEROPTERA:
TESSARATOMIDAE)
G.B. MONTEITH
Queensland Museum, PO Box 3300, South Brisbane, Qld 4010
(Email: geoff.monteith(a)bigpond.com)
Abstract
New information is given on maternal care, host plants and distribution for 9 Australian
Tessaratomidae: Oncomerinae in the genera Cumare Blóte, Garceus Distant, Lyramorpha
Westwood, Musgraveia Leston & Scudder, Oncoscelis Westwood, Peltocopta Bergroth,
Plisthenes Stal and Stilida Stal. Corrections to some recorded distributions and host plants are
discussed and a summary table of data for all 15 Australian species is presented.
Introduction
The subfamily Oncomerinae of the Tessaratomidae comprises 15 genera of
large, often colourful, shield bugs. It has its greatest generic diversity (12
genera) in Australia (Sinclair 2000). Species of Oncomeris Laporte are the
largest shield bugs in the world, one of which, O. flavicornis (Guérin), occurs
in northern Queensland. Cassis and Gross (2002) listed 18 species from
Australia but three of these (Lyramorpha perelegans Vollenhoven, 1868, L.
diluta Stàl, 1863 and Oncomeris ostraciopterus (Montrouzier, 1855)) are
based on literature records from the 1800s which have never been confirmed,
so the fauna is probably only the 15 species listed in Table 1. They can be
readily identified using the generic key of Sinclair (2000) and the species
keys of Leston and Scudder (1957). АП species are restricted to the eastern
parts of Queensland and New South Wales. Old records from Tasmania,
Victoria and South Australia (Gross 1975, Cassis and Gross 2002) have not
been verified in modern times and are not considered here. All species except
Cumare pallida Blóte, 1945 and Agapophyta bipunctata Guérin, 1831 are
virtually restricted to rainforests.
The group is well known for the parental care of eggs and nymphs shown by
some species, first observed by the pioneer Australian naturalist F. P. Dodd
(Dodd 1904, 1916), with later records by Kumar (1969) and Sinclair (2000).
Previous observations were summarised by Monteith (2006), who gave new
information on several species with photographs of parental behaviour and
discussion of its significance. In the most advanced forms (Cumare Blóte,
Garceus Distant and Peltocopta Bergroth), the female carries the nymphs on
her abdomen for a significant period of time. This so-called *nymphal
phoresy’ is also seen in certain SE Asian Tessaratomini (Gogala et al. 1998).
Oncomerines feed on the sap of woody plants, especially from apical shoots
when in flush growth. All species have a rather narrow host range.
Musgraveia sulciventris (Stal, 1863), the Bronze Orange-Bug, which feeds
on native species of Citrus L. (Rutaceae), has become a pest of cultivated
38 Australian Entomologist, 2011, 38 (1)
Figs 1-6. Oncomerinae. (1-4) Lyramorpha parens: (1) female and large nymphs
feeding at stem internode of Castanospora alphandii; (2) female brooding 2nd instar
nymphs; (3) gregarious group of 2nd and 3rd instar nymphs on non-food plant,
Passiflora; (4) adults feeding on Jagera pseudorhus (5-6) Oncoscelis australasiae on
Medicosma cunninghamii: (5) female brooding egg clutch; (6) eggs in clutch of 14.
Photos: A. Gillanders (1, 3), D. C. F. Rentz (2), K. Aland (4), J. Wright (5-6).
Australian Entomologist, 2011, 38 (1) 39
citrus and is well studied (Cant er al. 1996a, 1996b). Recorded food plants for
other Australian species are given by Kumar (1969), Sinclair (2000), Cassis
and Gross (2002) and Monteith (2006). A North Dakota State University
website (Rider 2010) lists all recorded food plants but without assessing
veracity. Some literature records of food plants are clearly simply resting
records and caution is needed in recording food plants in the absence of
actual feeding.
This paper records new information and/or photographs for nine species of
Australian Oncomerinae. Several doubtful food plant records are excluded
and a summary table of details for all Australian species is presented (Table
1). Plant nomenclature is taken from Henderson (2002). Vouchers for many
of the insects mentioned are in the Queensland Museum (QM), Brisbane.
Cumare pallida В1бїе, 1945
Eggs have been recorded (Monteith 2006) but not measured. Those from a
preserved clutch from Auburn River NP (in QM) are barrel shaped, 1.2 mm
high and 1.1 mm in diameter. Micropyles not countable.
Garceus fidelis Distant, 1893
Eggs in a hatched clutch of 14 eggs from Garradunga (in QM) are 1.5 mm
high and 1.7 mm in diameter. Micropyles not countable.
Lyramorpha rosea Westwood, 1837
Lyramorpha Westwood has two species in Australia: L. rosea in NSW and
southern Queensland with 4 antennal segments and L. parens Breddin in
northern Queensland with 5 antennal segments. L. rosea has been recorded
from five genera of the Sapindaceae (Table 1), four of them native plus the
cultivated lychee (Litchi chinensis Sonn.). Kumar (1969) found brooding
females on Alphitonia Reissek ex Endl. (Rhamnaceae) and Flindersia К. Br.
(Rutaceae) in Brisbane but this record has not been repeated for these
common trees in 40 years and they are deleted as host plants pending
confirmation. Similarly, the early record from Synoum glandulosum (Sm.) A.
Juss. (Meliaceae) by Musgrave and Whitley (1931) requires confirmation as
this is not a "beach plant” as described and might be a misidentification of the
confirmed food plant Cupaniopsis anacardioides (A. Rich.) Radlk., which is
a beach plant at the site and has similar leaves and fruit.
Lyramorpha parens Breddin, 1900 (Figs 1-4)
No native food plants have been recorded for this species. During April 2010,
Mr Alan Gillanders recorded it from the following six species of rainforest
Sapindaceae at three localities on the northern Atherton Tableland: Arytera
divaricata F. Muell. (Atherton), A. pauciflora S.T. Reynolds (Yungaburra),
Castanospora alphandii (F. Muell.) F. Muell. (Yungaburra), Guioa acutifolia
Radlk. (Lake Eacham), С. /asioneura Radlk. (Yungaburra), Sarcotoechia
serrata S.T. Reynolds (Yungaburra). Mr Garry Sankowsky recorded it from the
A0 Australian Entomologist, 2011, 38 (1)
Table 1. List of confirmed Australian Oncomerinae showing recorded information on
distribution, host plants, egg clutch size and aspects of brood care. Queensland is
divided into 4 latitudinal zones: SQ, from NSW border to 22°S; NQ, from 16-22°S;
CYP, from 16°S to tip of Cape York; TS, Torres Strait Islands. Blank cells indicate no
ЕЕЕ
SPECIES AND SOURCE AUSTRALIAN AUSTRALIAN FOOD
LITERATURE DISTRIBUTION PLANT FAMILIES &
GENERA
Agapophyta bipunctata NQ, CYP, TS CAESALPINIACEAE:
Guérin, 1831 (1,3,6) Cassia
Cumare pallida Blóte, 1945 SQ, NQ, CYP EUPHORBIACEAE:
(3,4,6,8) Petalostigma
Erga longitudinalis NSW, SQ FABACEAE:
(Westwood, 1837) (1) Austrosteenisia
Garceus fidelis Distant, NQ ELAEOCARPACEAE:
1893 (3,4,6) Elaeocarpus
Lyramorpha rosea NSW, SQ SAPINDACEAE:
Westwood, 1837 (1,3,4,5) Alectryon, Atalaya,
Cupaniopsis, Guioa, Litchi
Lyramorpha parens NQ, CYP, TS SAPINDACEAE:
Breddin, 1900 (6,7,8) Alectryon, Arytera,
Castanospora, Cupaniopsis,
Elattostachys, Guioa, Jagera,
Litchi, Nephelium, Sarcotoechia
Musgraveia antennatus CYP, TS RUTACEAE: Citrus
(Distant, 1880) (8)
Musgraveia sulciventris NSW, SQ RUTACEAE: Citrus
(Stal, 1863) (1,3,6)
Oncomeris flavicornis NQ, CYP -
(Guérin, 1831)
Oncoscelis australasiae NSW, SQ RUTACEAE: Acronychia,
Westwood, 1837 (3,8) Medicosma, Melicope
Peltocopta crassiventris NSW, SQ EUPHORBIACEAE:
(Bergroth, 1895) (1,6,8) Mallotus
Plisthenes australis NQ, CYP MELIACEAE: Aglaia
Horváth, 1900 (3,6,8)
Stilida indecora Stàl, 1863 NSW, SQ, NQ SAPINDACEAE
(1,3,6) Alectryon, Arytera,
Cupaniopsis, Guoia
Stilida sinuata Stàl, 1870 СҮР, -
Tibiospina darlingtoni NQ -
Sinclair, 2000
т
Australian Entomologist, 2011, 38 (1) 41
information. Literature sources for each species are as follows: 1, Kumar 1969; 2,
Cant et al. 1996a, b; 3, Sinclair 2000; 4, Cassis and Gross 2002; 5, Waite and Hwang
2002; 6, Monteith 2006; 7, Astridge 2006; 8, Monteith, this paper.
EGG EGG BROODING BROODING NYMPHAL
CLUTCH BROODING IST 2ND PHORESY
INSTARS INSTARS
14 Yes No
14 Yes Yes Yes Yes
24-26 Yes No
14 Yes Yes - Yes
up to 42 Yes - - No
40-42 Yes Yes Yes No
14 No No No No
14 Үез ?Yes No
24-39 Yes Yes Yes Yes
42 Australian Entomologist, 2011, 38 (1)
following seven additional Sapindaceae planted on his property, 8.5 km
NNW of Atherton: Alectryon coriaceus (Benth.) Radlk., 4. semicinereus
(F. Muell) Radlk., Cupaniopsis anacardioides (A. Rich.) Radlk., С.
diploglottoides Adema, С. flagelliformis (F.M. Bailey) Radlk. var.
flagelliformis, Elattostachys megalantha S.T. Reynolds, E. microcarpa S.T.
Reynolds. The author recorded it feeding on Jagera pseudorhus (A. Rich.)
Radlk. (Sapindaceae) at Iron Range in December 2010 (Fig. 4). Lyramorpha
parens also feeds on the exotic sapindaceous fruit trees lychee (specimens in
the Mareeba DPI collection from Innisfail) and rambutan, Nephelium
lappaceum L. (Astridge 2006), but is not of pest status. Thus both Australian
species of Lyramorpha apparently feed exclusively on Sapindaceae. Fig. 1
shows an adult and several late stage nymphs feeding together at an internode
of Castanospora alphandii.
Gende and Kumar (2001) listed Dodonea viscosa Jacq., plus a variety of non-
sapindaceous plants, from label data of New Guinea specimens identified as
Lyramorpha parens. However, both the identity of the Lyramorpha and the
feeding status of the plant records need confirmation and will not be pursued
for this Australian treatment.
The only breeding behaviour noted previously has been of a female brooding
a clutch of 40 eggs (Monteith 2006). A photograph taken at Kuranda by Dr
David Rentz (Fig. 2) shows a female standing guard over about 20 uniformly
sized nymphs. First instars in Oncomerinae are invariably subglobose in
shape and become flattened when they moult to second instar (Kumar 1969,
Monteith 2006). In L. rosea the nymphal colour pattern changes from
chequered red and black in first and second instars to uniform red in the older
instars (Kumar 1969); this also seems to be the case in L. parens. Based on
the size of the nymphs in Fig. 2, their flattened form and their colour pattern,
it can be assumed that they are second instars. This is evidence that L. parens
broods its young beyond moulting to the second instar.
Observations by Alan Gillanders show that the older nymphs of L. parens
form massed groups, which feed together and frequently move in close-
packed groups to new feeding sites when old sites are exhausted. They are
often to be found on non-host plants while in transit (Fig. 3). Clearly, their
brilliant aposematic colours reinforce the group defence they enjoy from their
dorsal defence glands during these exposed journeys. This feeding strategy
contrasts with that of oncomerines such as Cumare and Peltocopta, which
have solitary, camouflaged and rather sedentary later-stage nymphs
(Monteith 2006).
Musgraveia antennatus (Distant, 1880)
The southern citrus pest, Musgraveia sulciventris, feeds solely on native and
cultivated species of Citrus (Rutaceae), including species of ‘native limes'
previously included in the genera Eremocitrus Swingle and Microcitrus
Australian Entomologist, 2011, 38 (1) 43
Swingle, now placed in Citrus (Henderson 2002). The rare Musgraveia
antennatus is found only in Torres Strait (Moa Island) and northern Cape
York Peninsula. Sinclair (2000) recorded it from cultivated West Indian Lime
(Citrus aurantiifolia (Christm.) Swingle) and there is a record from cultivated
Pomelo (Citrus maxima Merr.) at Lockhart River in the AQIS collection,
Cairns (Sally Cowan pers. comm.). It overlaps with the native Citrus
garrawayi F.M. Bailey in the southern part of its range (Iron Range) and this
plant is a candidate for a natural host, but in the northern part of its range
(Lockerbie, Torres Strait) there are no native Citrus (fide Australia's Virtual
Herbarium, http://www.ersa.edu.au/avh/ and Garry Sankowsky pers. comm.)
and it must have a non-Citrus native host there. Mr Sankowsky suggests this
may be the rutaceous Zanthoxylum rhetsa (Roxb.) DC, which is common in
the region and used there by various citrus-breeding papilionid butterflies.
Oncoscelis australasiae Westwood, 1837 (Figs 5-9)
Cassis and Gross (2002) restored the original generic name Oncoscelis
Westwood for this species, instead of Rhoecus Bergroth or Rhoecocoris
Bergroth which have usually been used in modern times (e.g. Leston and
Scudder 1957, Sinclair 2000).
There are no published observations on the biology of this large but
uncommon species apart from records on Melicope micrococca (F. Muell.)
T.G. Hartley (Rutaceae) by Sinclair (2000). It has also been taken on
Melicope elleryana (F. Muell.) T.G. Hartley on Stradbroke Island (pers.
obs.). A specimen (in QM) from near Harrington, NSW, was taken on
Acronychia oblongifolia (А. Cunn ex Hook.) Endl. ex Heynh (Rutaceae) by
Geoff Williams, who reports that it also occurs there on Acronychia
imperforata F. Muell. In February 2009, Chris Burwell (QM) discovered a
colony feeding and breeding on Medicosma cunninghamii (Hook.) Hook. f.
(Rutaceae) along upper Enoggera Creek near Brisbane.
Several adults from this last collection were brought to the Queensland
Museum and maintained on the food plant in plastic bags. On 7 February, a
female laid a batch of 14 eggs on the underside of a leaf and took up a
guarding position over them (Fig. 5). The eggs were arranged in the standard
3,4,4,3 pattern (Fig. 6), which has been described for several other species
(Monteith 2006). The eggs were large (3.2 mm diameter), spherical, with 46-
48 micropyles, the largest number known for any oncomerine. The female
abandoned the eggs during photography two days after deposition. They
hatched on 20 February to give convex, non-feeding, white nymphs (Fig. 7)
which clustered together in an immobile group until the night of 27 February,
when all moulted to highly flattened, semi-transparent second instars (Fig. 8)
which dispersed. This post-hatching nymphal behaviour indicates that the
female would almost certainly have continued to brood the first instars until
their moult, as is usually the case (Monteith 2006).
A4 Australian Entomologist, 2011, 38 (1)
Figs 7-14. Oncomerinae. (7-9) Oncoscelis australasiae on Medicosma cunninghamii:
(7) eggs hatching to 1st instar nymphs; (8) newly hatched 2nd instar and exuvium; (9)
male showing enlarged hind legs. (10-12) Peltocopta crassiventris: (10) female from
Toogoom; (11) 2nd instar on underside of Mallotus discolor leaf; (12) female being
predated by grey butcher bird. (13-14) Plisthenes australis: (13) male (L) and female
on Aglaia meridionalis; (14) two adults and a nymph on presumed Aglaia sapindina.
Photos: J. Wright (7-9), M. Robinson (10-12), M. Cermak (13), G. Monteith (14).
Australian Entomologist, 2011, 38 (1) 45
Peltocopta crassiventris (Bergroth 1895) (Fig 10-12, 15)
This extremely rare species has been known from a few coastal localities
over a latitudinal range of 160 km between Iluka in New South Wales and
Surfers Paradise in Queensland (Monteith 2006). Its only food plant is
Mallotus discolor F. Muell. ex Benth. (Euphorbiaceae) and it exhibits the
most advanced maternal care known in the family. In late March 2010, Mrs
M. Robinson recorded the species from her garden at Toogoom
(25.258°S152.696°E), which is on the coast 330 km further north. Females
(Fig. 10) carrying both first and second instar nymphs (Fig. 11) were present.
The author visited and confirmed that the host plants were M. discolor which
was very abundant in this area of partly cleared remnant rainforest. This is a
major range extension for this unique species.
In December 2010, Mrs Robinson noted many bugs on the same trees at
Toogoom, with specimens commonly falling from overhanging Mallotus
trees onto the driveway and verandahs of the house. This coincided with a
heavy wet season and parallels the population outbreak of this usually rare
species described by Monteith (2006) at Surfers Paradise in 1961 and for
several years thereafter. Other observations by Mrs Robinson included
several instances of the unique copulation posture adopted by this species,
including one involving a female that was brooding a batch of 33 eggs at the
time (Fig. 15). Although Pe/tocopta has the same acrid defense secretions
shared by other oncomerines, a successful predation of a female by a grey
butcher bird, Cracticus torquatus (Latham), was also noted (Fig. 12).
Fig. 15. Peltocopta crassiventris: male and female in copulation above a clutch of 33
eggs, which the female was brooding when copultaion was initiated. Note the bright
colours of the male (left) compared with the pallid female. Photo: M. Robinson.
46 Australian Entomologist, 2011, 38 (1)
Plisthenes australis Horváth, 1900 (Figs 13-14)
Sinclair (2000) listed the locality of Byfield (22.858°S, 150.695°E) as the
southern limit for this otherwise far northern tropical species. The specimen
on which this is based has proved to be Oncoscelis australasiae, so the
southern limit for Plisthenes australis is now confirmed as being 740 km
further north at the Mulgrave River (17.34S°).
The only foodplant listed for this species is Citrus, in the Rutaceae (Sinclair
2000). This record is based on a single specimen taken from a planted West
Indian Lime tree at Iron Range in 1978 by M. DeBaar (now in the
Queensland Forestry Collection). This same individual tree is usually host for
the related citrus-feeding Musgraveia antennatus (pers. obs.) Citrus is
commonly available within the range of Plisthenes Stal so, without other
feeding records, it seems best to treat this single specimen as just a resting
record. Two confirmed host records are now available for P. australis on
Aglaia Lour. in the Meliaceae: Michael Cermak photographed it on Aglaia
meridionalis Pannell at Cape Tribulation in 2004 (Fig. 13) and the present
writer took it feeding with nymphs on a plant identified by experienced field
botanist Garry Sankowsky, from the May 1973 photograph taken at
Lockerbie (Fig. 14), as almost certainly Aglaia sapindina (F. Muell.) Harms.
Stilida indecora Stàl, 1863
Monteith (2006) recorded and pictured a female brooding a clutch of 42 eggs
which hatched into first instars. These nymphs dispersed because the food
plant was dead, so it was not possible to be sure that normally they would
have been brooded until the critical moulting from first to second instar. This
has now been confirmed: in February 2007, a female (in QM) was taken on a
species of Arytera in vine scrub at ‘Toomba’ station (19.966°S, 145.582°E),
brooding a mass of nymphs on top of a hatched clutch of 36 eggs. There were
12 first instars and 20 seconds, demonstrating that female brooding does
persist to the second instar.
Discussion
With the additions and corrections presented here we now have a relatively
full body of reliable information on the biology and distribution of the 15
accepted species of Australian Tessaratomidae (Table 1). Host plants are
known for 12 species and at least partial breeding behaviour is recorded for
10 species. Nothing of this behaviour is known for the giant, lowland,
northern Queensland species Oncomeris flavicornis (Guérin, 1831), or for the
curious endemic monotypic genus and species Tibiospina darlingtoni
Sinclair, 2000, known from high altitudes in the Wet Tropics. The author
would welcome observations on these or other species.
Australian oncomerines feed on a relatively limited range of plant taxa,
comprising only seven families in four plant orders. All genera of
oncomerines are restricted to a single plant family. In the Malvales, Garceus
Australian Entomologist, 2011, 38 (1) 47
occurs on Elaeocarpaceae; in the Fabales, Agapophyta Guérin and Erga
Walker occur on the legume families Caesalpiniaceae and Fabaceae
respectively; in the Euphorbiales both Peltocopta and Cumare feed on
Euphorbiaceae; in the Sapindales, Lyramorpha and Stilida Stàl feed on
Sapindaceae; Oncoscelis and Musgraveia Leston & Scudder feed on
Rutaceae; and Plisthenes on Meliaceae.
The Oncomerinae show five progressive levels of maternal care: (1), those
that lay eggs and abandon them (Musgraveia sulciventris); (2), those in
which the female broods the eggs until hatching (Erga and Agapophyta); (3),
those in which the female continues to brood the sedentary first instars until
they moult to second instars (Stilida and Oncoscelis); (4), those in which
females continue to shepherd mobile second instars (Lyramorpha parens);
(5), those in which the brooding female is modified to carry the first and
second instars on her body after hatching (Peltocopta, Cumare and Garceus).
This study confirms that more species than previously suspected guard their
young until the second instar. This supports the idea proposed by Monteith
(2006) that maternal care is largely a device to protect the vulnerable, usually
non-feeding, sub-globose first instars (Fig. 7) until they moult to the
flattened, camouflaged second instars, which disperse and begin to feed (Figs
8, 11).
As evidence for maternal care in more species accumulates, the complete
lack of parental care in Musgraveia sulciventris becomes more unusual.
Maternal care slows the potential for rapid population increase because
females invest time and energy in one egg clutch. Two of the three natural
food plants of M. sulciventris are typical of dry vine forests (Citrus
australasica F. Muell.) or of inland plains (C. glauca (Lindl.) Burkill), where
rainfall and plant growth are seasonal and unreliable. For M. sulciventris, the
loss of maternal care and the ability to produce multiple eggs clutches (Cant
et al. 19962) might give populations the ability to multiply rapidly in
temporarily favourable conditions. This characteristic of the species may
have pre-adapted it for the pest status it gained when cultivated citrus became
available.
Acknowledgements
I am grateful to the following for allowing me to include their observations:
Chris Burwell (QM, Brisbane), Michael Cermak (Cairns), Murdoch DeBaar
(Brisbane), Alan Gillanders (Yungaburra), David Rentz (Kuranda), Meg
Robinson (Toogoom), Garry Sankowsky (Tolga) and Geoff Williams
(Lansdowne). Photos are by Michael Cermak, Meg Robinson, Alan
Gillanders, Jeff Wright (QM, Brisbane) and David Rentz. Harry Fay
(DEEDI, Mareeba) helped with information about lychee feeders.
48 Australian Entomologist, 2011, 38 (1)
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THE AUSTRALIAN
Entomologist
Volume 38, Part 1, 25 March 2011
CONTENTS
BRABY, M. F., DOUGLAS, F. and WILLAN, R. C.
The nomenclature of Ogyris halmaturia (Tepper, 1890)
(Lepidoptera: Lycaenidae)
KALLIES, А. апа MOLLET, B.
А new species of forester moth from Victoria (Lepidoptera:
Zygaenidae: Procridinae)
MONTEITH, G. B.
Maternal care, food plants and distribution of Australian Oncomerinae
(Hemiptera: Heteroptera: Tessaratomidae) 37
THEISCHINGER, G., MILLER, J., TANG, C., KROGH, M. and POPE, E.
The benefits of using both adult and larval stoneflies (Plecoptera)
in environmental surveys: an example from New South Wales with
a summary of the Australian stonefly fauna
ISSN 1320 6133