Records of the Australian Museum

Review of the Australasian genera of signal flies (Diptera: Platystomatidae).

..David K. McAlpine 113

Revision of Paralamyctes (Chilopoda: Lithobiomorpha: Henicopidae), with six new species

from eastern Australia. Gregory D. Edgecombe 201

A new seahorse species (Syngnathidae: Hippocampus ) from the Great Barrier Reef.

.Michelle L. Horne 243

Two new species of fairy shrimp (Crustacea: Anostraca: Thamnocephalidae: Branchinella )

from the Paroo, inland Australia. Brian V. Timms 247

Scaptodrosophila aclinata: a new Hibiscus flower-breeding species related to S. hibisci

(Diptera: Drosophilidae). Shane F. McEvey & J.S.F. Barker 255

VOLUME 53 NUMBER 2 12 September 2001

RECORDS OF THE AUSTRALIAN MUSEUM

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Records of the Australian Museum (2001) Vol. 53: 113-199. ISSN 0067-1975

Review of the Australasian Genera of Signal Flies
(Diptera: Platystomatidae)

David K. McAlpine

Australian Museum, 6 College Street, Sydney NSW 2010, Australia

Abstract. The distribution patterns of platystomatid genera in the 12 recognized provinces of the
Australasian Region are recorded. Notes are provided on biology and behaviour, including parasitism
by fungi and strepsipterans, and mimicry of other insects and spiders. Means of separation from other
acalyptrate families are provided. A key to Australasian genera is given. The subfamily Angitulinae is
placed in synonymy of Platystomatinae. The subfamily classification is briefly discussed. The following
new genera are described: Aetha, Bama, Eumeka, Hysma, Par, Phlyax, Signa, Tarfa, Terzia, Tomeus.
Gonga and Polimen are new subgenera of Naupoda and Bama respectively. The genus Lasioxiria Hendel
is a new synonym of Atopognathus Bigot. Chaetostichia Enderlein is a new synonym of Scholastes
Loew. Eopiara Frey, described as a subgenus of Piara Loew, is raised to generic status. The genera
Angituloides Hendel and Giraffomyia Sharp are reduced to subgenera of Angitula Walker. The following
new species are described: Aetha cowanae, Bama (Polimen) shinonagai, Eumeka hendeli, Hysma lacteum,
Paryphodes hospes, Signa mouldsi, Tarfa bowleyae, Terzia saigusai, Tomeus wyliei, Zealandortalis
gregi. Lule speiseri de Meijere, 1914 is a new synonym of Phasiamya metallica Walker, 1849. New
generic combinations are made as follows: Angitula austeni (Hendel, 1913) ( Angituloides ); Angitula
irregularis (Malloch, 1940) (Giraffomyia); Angitula regularis (Malloch, 1940) (Giraffomyia); Angitula
solomonensis (Malloch, 1940) (Giraffomyia); Angitula willeyi (Sharp, 1899) (Giraffomyia); Atopognathus
hirsutus (Hendel, 1914a) (Lasioxiria); Bama bipunctatum (Hendel, 1914a) (Euxestomoea); Bama
papuanum (Hennig, 1940b) (Xiria); Bama strigatum (Hennig, 1940b) (Xiria); Chaetorivellia tarsalis
(Walker, 1861c) (Ortalis); Cleitamoides trigonalis (de Meijere, 1913) (Cleitamia); Eopiara chrysoptera
(Frey, 1964) (Piara); Eopiara elegans (Frey, 1964) (Lamprogaster); Lamprophthalma egregia (de
Meijere, 1924) (Plagiostenopterina); Lamprophthalma medionotata (de Meijere, 1924)
(Plagiostenopterina); IMicroepicausta sangiensis (de Meijere, 1916) (Elassogaster); Neohemigaster
fascifrons (de Meijere, 1916) (Pterogenia); Neohemigaster guttata (Walker, 1856) (Lamprogaster, later
in Pterogenia); Parevitta (Malloch, 1939a) (Elassogaster); Phlyax simmondsi (Bezzi, 1928) (Naupoda);
Scholastes aduncivena (Enderlein, 1924) (Chaetostichia); Xiriella lunaris (de Meijere, 1916) (Lule).

McAlpine, David K., 2001. Review of the Australasian genera of signal flies (Diptera: Platystomatidae). Records
of the Australian Museum 53(2): 113-199.

114 Records of the Australian Museum (2001) Vol. 53

The Platystomatidae, recently termed signal flies, are
probably among the four largest families of acalyptrate
Schizophora in the Australasian Region, but in the Americas
and in the Palaearctic Region the representation is
comparatively small. There are about 119 known world
genera and nearly 1200 described species. In the
Australasian Region there are 54 recognized genera and c.
493 probably valid described species. At least 220 additional
undescribed Australasian species have been sighted in
collections, and I estimate that the total Australasian
platystomatid fauna is unlikely to include less than 900 species.

It is the main aim of this paper to provide means of
identification and general basic information on the genera
of Platystomatidae living in the Australasian Region,
including the Oceanian Region, as delimited by Evenhuis
(1989) and in the section on Geographic Distribution below.

Methods and terminology

In general I follow a traditional system with minimal use of
terms implying doubtful, unproved, or, for present purposes,
irrelevant homologies. Details are given by McAlpine
(1973a), with most terms also explained by Harrison (1959),
Crosskey (1973), and Colless & McAlpine (1991). Paired
bristles and other paired structures are described in the
singular, except where the context makes this inappropriate.
The antenna is treated as a six-segmented appendage and
the segments are numbered consecutively from the base
(Fig. 96). The greater part of the arista thus consists of
segment 6 and the very short segment 4 may not be visible
in dried specimens. The system of nomenclature of wing
veins (Fig. 1) is the simplest possible for one not concerned
with trans-ordinal homologies. Cell-4 index is defined as
the ratio of the length of the antepenultimate section of vein
4 to the full length of the discal cell along vein 4.

In using the keys and descriptions particular care should

Figure 1 . Euprosopia tenuicornis Macquart, base of wing of
female. Abbreviations: al, alula; ax, axillary lobe; c, costa; hb,
humeral break of costa; hm, humeral crossvein; pa, postalar callus
bearing postalar bristle; sc, subcosta; sq, squama; sr, suprasquamal
ridge; sv, stem vein (base of R); te, tegula (sexually dimorphic);
vl-v6, veins one to six.

be taken in interpreting the terms bristle, setula, hair,
pubescence, and pruinescence (see McAlpine, 1973a).

An effort has been made to make the key to genera
workable for all dried adults in good condition, provided
that a good stereo-microscope with magnifications up to
x75 is used. However, because of individual variation, great
diversity in some genera, and incomplete knowledge of the
fauna, it should not be too readily assumed that specimens
which do not key out necessarily belong in unrecorded
genera.

The following abbreviations refer to institutions housing
specimens:

AM Australian Museum, Sydney
AMST Zoological Museum, Amsterdam
ANIC Australian National Insect Collection, CSIRO,
Canberra

BM The Natural History Museum, Fondon
BPB Bernice R Bishop Museum, Honolulu
CNC Canadian National Collection, Agriculture
Canada, Ottawa

DEI Deutsches Entomologisches Institut, Eberswalde
FRIF Forest Research Institute, Fae
HEFS Zoological Museum Helsinki
KONE Department of Agriculture and Fivestock,
Konedobu, Port Moresby

MNB Museum of Natural Science at Humboldt
University, Berlin

MNM Hungarian Natural History Museum, Budapest
NAT Natal Museum, Pietermaritzburg
RMS Naturhistoriska Riksmuseet, Stockholm
NMWC National Museum of Wales, Cardiff
NSMT National Science Museum, Tokyo
OX University Museum, Oxford
PM Museum national d’Histoire naturelle, Paris
UQ University of Queensland Insect Collection,
Brisbane

USNM National Museum of Natural History, Washington
WM Natural History Museum, Vienna
ZMC Zoological Museum, Copenhagen

The following collectors’ names are abbreviated to the
initials: J.H. Barrett, D.J. Bickel, T.G. Campbell, G. Daniels,
B.J. Day, A.F. Dyce, E.D. Edwards, J.F. Gressitt, G.A.
Holloway, A. Hughes, J.W. Ismay, N.F. Krauss, D.K.
McAlpine, B.J. Moulds, M.S. Moulds, H. Roberts, J.
Sedlacek, M. Sedlacek, P. Shanahan, S. Shinonaga, B.J.
Sinclair, H.A. Standfast, F.H. Taylor, A. Walford-Huggins,
A.R. Wallace, T.A. Weir, F.R. Wylie.

Geographic distribution

The Australasian Region, the area covered in this review, is
the same as the Australasian and Oceanian Regions together,
of Evenhuis (1989), who proposed no dividing line between
the two. The Region is thus defined in the west by Weber’s
Fine (“original” version of Merrill, 1945), in the north by
the northernmost islands of Micronesia and in the east by
the easternmost islands of Polynesia. To the south, I am not
concerned with islands beyond Tasmania and the two main
islands of New Zealand, as such islands probably harbour

David K. McAlpine: Australasian Platystomatidae 115

no platystomatids. South Island, New Zealand, provides the
southernmost recorded habitat of playtstomatids, in view of
their apparent absence from southern parts of South America.

On the basis of present knowledge, it appears that
Australasia harbours the most diverse platystomatid fauna
of any biogeographic region. New Guinea is likely to have
a particularly large number of undiscovered species (see
discussion in McAlpine, 1994). About 80% of the species
sorted from New Caledonia remain undescribed, and it
would not be surprising if some other island groups prove
significant sources of new discoveries.

Of the Australasian genera 34 (64%) are endemic to the
Region. These are: Achias, Aetha, Angitula, Apactoneura,
Apiola,Asyntona,Atopognathus, Bama, Brea, Chaetorivellia,
Cleitamia, Cleitamoides, Duomyia, Eumeka, Euxestomoea,
Guamomyia, Hysma, Inium, Laglaisia, Lenophila,
Loriomyia, Loxoneuroides, Mesoctenia, Montrouziera, Par,
Pseudocleitamia, Pseudorichardia, Phlyax, Scotinosoma,
Signa, Tarfa, Terzia, Tomeus, Zealandortalis. Of the
remaining 20 genera all except Paryphodes extend to the
Oriental Region: Antineura, Conicipithea, Elassogaster,
Euprosopia, Lamprogaster, Lamprophthalma, Meringomeria,
Microepicausta, Naupoda, Neohemigaster, Plagiostenopterina,
Pogonortalis, Pseudepicausta, Pterogenia, Rhytidortalis,

Rivellia, Scholastes, Trigonosoma, Zygaenula.

Eight of the Australasian genera are shared with the
Afrotropical Region: Elassogaster, Lamprophthalma,
Naupoda, Paryphodes, Plagiostenopterina, Pseudepicausta,
Rivellia, Scholastes.

Five of the Australian genera are shared with the Palaearctic
Region: Elassogaster, Euprosopia, Lamprophthalma,
Rhytidortalis, Rivellia.

Only one Australasian genus occurs naturally in the
Americas (both Nearctic and Neotropical Regions), the
almost cosmopolitan Rivellia. However, an Australian
species of Pogonortalis is introduced in California. Duomyia
was erroneously recorded from Chile on the basis of a
mislabelled specimen (see McAlpine, 1973a).

For the purpose of recording generic distributions, I divide
Australasia into 12 provinces as indicated in Fig. 2. Delimitation
of these follows natural barriers and a degree of convenience,
rather than national boundaries. Thus the biogeographically
diffuse areas covered by the terms Indonesia, Papua New
Guinea, and Kiribati have no place in this system. The
boundaries between Micronesia and Polynesia have been
simplified, because ethnic zones are not relevant to
playtstomatid distributions, and because these zones are little
known and probably largely depauperate in platystomatids.

Figure 2. Provinces of Australasia which harbour platystomatids. 1, Micronesia. 2, Moluccas. 3, New Guinea. 4,
Bismarck Archipelago. 5, Solomon Archipelago. 6, Vanuatu. 7, New Caledonia. 8, Fiji. 9, Australia. 10, Norfolk
Island. 11, New Zealand. 12, Tropical Polynesia. 13, excluded (Oriental Region).

116 Records of the Australian Museum (2001) Vol. 53

The platystomatid faunas of many Australasian provinces
are still poorly collected and studied. Many of my statements
about distribution, diversity, and relationships may need
some revision as new data become available.

1. Micronesia. The four genera recorded for this province
are: Guamomyia, Pogonortalis, Scholastes (all from Guam,
Malloch, 1942), and Pseudorichardia (Tuvalu, in AM). No
endemic genus is known, but the fauna may be considerably
richer than indicated by the immediately available data.

2. MOLUCCAS. For present purposes this province covers
only the islands between Weber’s and Lydekker’s Lines,
the latter given as “Weber’s Line (modified)” by Merrill
(1945), and Aru is definitely excluded. It is thus not
coextensive with the Indonesian province of Maluku. The
17 recorded genera are: Angitula, Antineura, Asyntona,
Atopognathus, Brea, Chaetorivellia, Elassogaster, Euprosopia,
Lamprogaster, Lamprophthalma, Plagiostenopterina,
Pseudepicausta, Pterogenia, Rivellia, Scholastes,
Trigonosoma, Zygaenula. None of these is endemic to
the province.

This province provides the western limit for the ranges
of Angitula, Asyntona, Atopognathus, Brea, and Chaetorivellia,
all genera restricted to less remote tropical parts of the
Australasian Region. The placement of this province at the
western limit of the Region therefore has a degree of biogeo¬
graphic justification. The Oriental genera Lamprophthalma,
Trigonosoma, and Zygaenula penetrate no further eastwards
than this province.

3. New Guinea. I have previously discussed the platy¬
stomatid fauna of this province (MeAlpine, 1982). The
province includes the large island of New Guinea and the
following minor islands or groups: Aru, Waigeo, Salawati,
islands of Geelvink Bay, D’Entrecasteaux Group, Trobriand
Group, Woodlark, Louisiade Group. The inclusion of Aru
with New Guinea rather than Moluccas is justified by the
fact that its recorded platystomatids all belong in typical
New Guinea genera, and many of its species are shared
with mainland New Guinea.

The 31 recorded genera are: Achias, Angitula,
Antineura, Asyntona, Atopognathus, Bama, Brea,
Chaetorivellia, Cleitamia, Cleitamoides, Elassogaster,
Eumeka, Euprosopia, Euxestomoea, Guamomyia, Hysma,
Laglaisia, Lamprogaster, Loriomyia, Mesoctenia,
Microepicausta, Naupoda, Neohemigaster, Paryphodes,
Plagiostenopterina, Pseudepicausta, Pseudocleitamia,
Pterogenia, Rivellia, Scholastes, Tomeus. The following seven
genera are endemic to this province: Bama, Cleitamia,
Cleitamoides, Laglaisia, Loriomyia, Pseudocleitamia, Tomeus.
The more widely distributed genera Atopognathus,
Euprosopia, and Lamprogaster have very substantial
specific representation in New Guinea, and c. 95% of
species of the large genus Achias live in New Guinea.

I stated (McAlpine, 1973a) that New Guinea has the
greatest number of platystomatid species of any part of the
world of comparable area. Subsequent work (including

McAlpine, 1994, and much unpublished work) has
strengthened this view.

The record of Lenophila from New Guinea (McAlpine,
1982) was based on a misidentification.

4. Bismarck Archipelago. This province includes the
islands of the New Britain, New Ireland, and Manus
(Admiralty) districts of Papua New Guinea. The 18
recorded genera are: Achias, Angitula, Atopognathus,
Brea, Elassogaster, Eumeka, Euprosopia, Euxestomoea,
Guamomyia, Lamprogaster, Mesoctenia, Microepicausta,
Par, Plagiostenopterina, Pseudepicausta, Pterogenia,
Rivellia, Scholastes. Only Par is endemic to the province.

The Bismarcks provide the eastern limit for the genera
Achias, Brea, Euxestomoea, and Mesoctenia, and are in this
respect an eastern extension of the New Guinea province.

5. Solomon Archipelago. This province includes the
Bougainville district of Papua New Guinea and the islands
forming the present nation of Solomon Islands, including
the Santa Cruz Group. The 14 recorded genera are:
Angitula, Asyntona, Atopognathus, Elassogaster,
Euprosopia, Lamprogaster, Microepicausta, Naupoda,
Plagiostenopterina, Pseudepicausta, Pterogenia, Rivellia,
Scholastes, Terzia. The genus Terzia is endemic to the province.

The platystomatid fauna has been reviewed by Curran
(1936) and Malloch (1940).

6. Vanuatu. The platystomatids of this group of mostly
small islands are perhaps too little known for profitable
discussion. The two recorded genera are Euprosopia and
Pseudorichardia. Though both have wide distributions, this
is the only province known to harbour both genera.

7. New Caledonia. This province includes the main
island of New Caledonia, with its small satellites, and the
more removed Loyalty Islands. The seven recorded genera
are: Eumeka, Lamprogaster, Montrouziera, Rivellia,
Scholastes, Signa, Tarfa. Montrouziera, Signa, and Tarfa
are endemic, but, whereas Signa is so far known only from
the main island, Montrouziera and Tarfa are perhaps restricted
to the Loyalty Islands. The proportion of endemic genera (43%)
is greater than that of any other Australasian province.

Though the number of genera is only half that recorded
for the Solomon Archipelago, the preliminary species count
(material studied by me) is about as great, despite the much
smaller land area of New Caledonia. Lamprogaster (at least
14 species) and Signa (at least 10 species) are the genera
with largest representation, these species being all endemic
to the province, and perhaps to the main island. Most of
them remain undescribed. New Caledonia is already
recognized as a significant botanical hot spot (Jaffre et al.,
1998). Preliminary evidence indicates that it is also a hot
spot for platystomatid diversity.

8. Fiji. The Fijian platystomatids were reviewed by Bezzi
(1928), but some emendments to generic placement are now
made. The five recorded genera are Meringomeria, Phlyax,
Pseudorichardia, Rivellia, and Scholastes. Only Phlyax is

David K. McAlpine: Australasian Platystomatidae 117

endemic. The genera Duomyia, Lamprogaster, and Naupoda
have been recorded from Fiji in error, the first two from
mislabelled Australian material (see McAlpine, 1973a), and
Naupoda from generic misplacement of the type species of
Phlyax (q.v.).

The presence of Meringomeria and Phlyax is of unusual
interest. Meringomeria, which appears to have at least two
endemic Fijian species, is otherwise represented only in the
Oriental Region. Phlyax, though endemic to Fiji, is the only
Australasian genus of the subfamily Trapherinae. There is
no record of either Meringomeria or the Trapherinae from
any other province of the Australasian Region.

9. Australia. For purposes of this review, this province
includes the six Australian states, the Northern Territory,
Torres Strait Islands, and Lord Howe Island. The 26 genera
are: Achias, Aetha, Asyntona, Atopognathus, Brea, Duomyia,
Elassogaster, Eumeka, Euprosopia, Euxestomoea,
Guamomyia, Hysma, Inium, Lamprogaster, Lenophila,
Loxoneuroides, Mesoctenia, Microepicausta, Naupoda,
Plagiostenopterina, Pogonortalis, Pterogenia, Rhytidortalis,
Rivellia, Scotinosoma, Zealandortalis.

The following six genera are endemic to the province:
Aetha, Duomyia, Inium, Lenophila, Loxoneuroides,
probably Scotinosoma. Duomyia is much the largest of the
endemic genera with at least 106 species.

The platystomatid fauna of Tasmania is an extension of that
of mainland Australia, with no known endemic taxa. The seven
recorded genera are: Duomyia, Euprosopia, Lamprogaster,
Lenophila, Microepicausta, Rhytidortalis, Rivellia.

Lord Howe Island harbours the genera Duomyia,
Guamomyia, Naupoda, Pogonortalis, and Rivellia. The species
of Duomyia and Pogonortalis are very closely related to
mainland Australian species. Naupoda nudiseta (Bezzi) is
endemic to the island. Guamomyia, represented by one species
endemic to the island, is known elsewhere only from northern
Micronesia, New Guinea, and Bismarck Archipelago.

10. Norfolk Island. The only recorded genera are
Pogonortalis and Rivellia, each with one species. This is a
quite small island with, understandably, a minuscule
platystomatid fauna. The Rivellia sp. is very similar to some
Australian species of its genus, but the endemic Pogonortalis
hians Schneider & McAlpine is not very close to congeneric
species of the Australian mainland and Lord Howe Island.

11. New Zealand. This province can be interpreted
broadly to include the Kermadec, Chatham, and more
southerly island groups, but only the two main islands
(North and South) are known to harbour platystomatids.
The only recorded genus is Zealandortalis (see Harrison,
1959). This was thought to be endemic, but an Australian
species is now described in the genus (see below).

12. Tropical Polynesia. I include in this province all
the main island groups of the central-eastern Pacific, from
Tokelau, Samoa, and Tonga in the west to French Polynesia
and the Pitcairn Group in the east, and the Hawaiian Chain in
the north. The six genera recorded from the province are:
Apactoneura, Apiola, Plagiostenopterina, Pseudorichardia,

Rivellia, Scholastes. Apactoneura and Apiola are endemic.

The two endemic genera are recorded only from the
Samoan Islands, which harbour all six genera (Malloch,
1930b). Only Pseudorichardia and Scholastes seem to have
a wide distribution through the province (Malloch, 1932).

Biology

The flies of the family Platystomatidae, are biologically
diverse, particularly in the larval stages. Summaries of their
biology are given by McAlpine (1973a, 1998, 1999). The
most complete treatment of larval biology is that of Ferrar
(1988). Some additional biological and habitat data are
recorded below under generic headings. In particular,
parasitism of adults by Strepsiptera is mentioned under
Paryphodes, parasitism by laboulbenialean fungi under
Rivellia, and possible larval association with roots of Acacia
under Rivellia. Possible Batesian mimicry of blowflies
(Calliphorinae) is mentioned below under Lamprogaster,
of braconid or ichneumonid wasps under Tarfa, of ants
(Formicidae) under Inium, of pompilid wasps under
Antineura and Eumeka, of vespid wasps under Achias and
Lamprogaster, of chrysomelid beetles under Phlyax, and
of jumping spiders (Salticidae) under Atopognathus and
Lenophila. In general, these mimicry hypotheses need
testing by field observations, as effective mimicry often
depends on behaviour as well as morphology. No flightless
forms with reduced wings are known in the Platystomatidae,
but it should be noted that the family is apparently
unrepresented on far southern islands, which harbour such
forms in other dipterous families.

Sexual dimorphism, other than that in the organs of
copulation and oviposition, is very diverse in platystomatids,
a summary being given by McAlpine (1998), with additional
data by McAlpine (2000). Some functional aspects of sexual
dimorphism are considered by McAlpine (1973b, 1975,
1979, 2000). Further mention of sexual dimorphism is
made below under the numerous genera in which it has
been observed. A range of head modifications in male
platystomatids is shown in Figs. 3-9.

Family identification

Platystomatid adults may generally be identified by use of
the keys to fa mili es of Diptera given by Colless & McAlpine
(1991) or Oosterbroek (1998). Unfortunately platystomatid
larvae are so little known that it is unlikely that larval
characters diagnostic for the family will be recognized in
the foreseeable future.

For those desiring a more rigorous method of sorting to
family level, without going through an unnecessary priming,
I offer the four not necessarily simple steps for adult flies
suspected as belonging in the Platystomatidae.

Step 1: Segregation of the higher Diptera. Platystomatid
flies have the general features of the division (or series)
Schizophora or “higher Diptera.” Flies of this division
usually have three stout antennal segments (the first or basal
one shortest), and the large third segment has a fine bristle¬
like structure, the arista, arising from near the base of its

118 Records of the Australian Museum (2001) Vol. 53

Figures 3-9. Modified head capsules of male platystomatids. 3, Asyntona “sp. A” (Bougainville, AM). 4,
Atopognathus goniceps (Hendel). 5, Laglaisia “sp. 1” (Sewan, West New Guinea, AM). 6, Angitula (Giraffomyia)
“sp. A” (New Britain, AM). 7, Achias furcatus Hendel. 8, Achias wallacei McAlpine. 9, Mesoctenia “sp. 1” (Bainyik,
Papua New Guinea, AM).

David K. McAlpine: Australasian Platystomatidae 119

Figures 10-11. Hysma lacteum n.sp. 10, wing. 11, head of female.

dorsal surface (Fig. 96); also they generally have a curved
slit, the ptilinal fissure (Fig. 68, pt), on the head, immediately
above the sockets of the antennae. In the Schizophora the
number of wing veins and their branches is normally not
above that shown in Fig. 10.

Step 2: Identifying the setulose vein 1. The Platystomatidae
are one of the few families of Schizophora which have a
series of many setulae (macrotrichia) on the dorsal surface
of vein 1 of the wing, extending from approximately the
level of the humeral crossvein to the distal end of this vein
where it merges with the costa (Fig. 1). These setulae are
distinctly larger than the microtrichia or pubescence—very
fine hair-like processes that clothe much of the surface of
the wing membrane and the veins, but do not tend to form
a linear series on veins. Failure to appreciate the difference
between setulae and microtrichia sometimes leads
elementary students to classify Strongylophthalmyia spp.
(family Tanypezidae) as platystomatids. The only other
Australasian schizophoran families with such an
extensive series of setulae on vein 1 are the Tephritidae
(true fruit flies) and Pyrgotidae (nocturnal scarab-killing
flies), except on islands to the south of New Zealand, where
platystomatids are absent

The small flies of the family Lonchopteridae also have a
series of dorsal setulae on vein 1, but lonchopterids, which
do not belong among the Schizophora, have no ptilinal
fissure and the arista arises from the apex of antennal
segment 3. The genus Herina Robineau-Desvoidy (family
Otitidae) also has a series of dorsal setulae on vein 1 but
these are restricted to the distal part of the vein; Herina has
a well developed, isolated proepisternal (propleural) bristle,
which is always absent in Platystomatidae.

Step 3: Sorting platystomatids from tephritids.

Platystomatids consistently have no break, or incision in
the costa at the point where the subcosta meets it; the costa
is quite continuously developed and sclerotised through this
point, though there is usually a visible break indicating a
flexible point in the costa a little beyond the level of the
humeral crossvein (humeral break, see Fig. 1). Tephritids,
on the other hand, do have a break or incision in the costa
at the distal end of the subcosta (usually in addition to the
humeral break) which can be detected by using an
appropriately high magnification, carefully positioning the
wing, and using transmitted light. The great majority of
tephritids have one or more pairs of incurved lower fronto-
orbital bristles in front of the usual upper fronto-orbital
bristles nearer the inner vertical bristles. These paired
bristles are well differentiated by their size, thickness, or
regular placement, from any irregular covering of hairs or
setulae on the postfrons. Platystomatids have no incurved
lower fronto-orbital bristles, but one to three upper fronto-
orbital bristles of diverse inclination may be present just in
front of the inner vertical bristle on each side of the
postfrons, often on a differentiated fronto-orbital plate. The
vast majority of tephritid species have the anal crossvein
(free transverse section of vein CuA or CuA 2 ) strongly
indented so that the anal cell is acutely produced
posterodistally. In the platystomatids the anal crossvein is
straight, simply curved (e.g., Fig. 10), or strongly reflexed
(Fig. 79), but it is only strongly indented and tephritid-like
in the rarely collected genus Loriomyia Kertesz (Fig. 85).
In a few platystomatids (e.g., Lenophila achilles McAlpine
& Kim) the anal crossvein has slight sigmoid curvature and
the anal cell is posterodistally acute, but the crossvein is
scarcely indented.

120 Records of the Australian Museum (2001) Vol. 53

Step 4: Sorting platystomatids from pyrgotids. The

Pyrgotidae are a large and morphologically diverse family
having much in common with the Platystomatidae,
including the extent of dorsal setulae on vein 1. Most
Australasian pyrgotids have the first two abdominal sternites
fused into a single plate, though a transverse line of
demarcation may still be visible; but all known platy¬
stomatids have these sternites separated by a distinct
membranous zone. A significant proportion of Australasian
pyrgotids have no distinct ocelli on the head; all known
Australasian platystomatids have well-developed ocelli.
Thus, most pyrgotids can be distinguished from platy¬
stomatids by having either sternites 1 and 2 fused or no
visible ocelli. Many pyrgotids also have the anal crossvein
indented, approximately as in the Tephritidae. The residue
of pyrgotid species that have separate sternites 1 and 2 as
well as distinct ocelli is very small and these species, so far
as I am aware, are distinguishable from platystomatids by
having a break in the costa at the end of the subcosta (even
though the majority of other pyrgotids have no such break).
Female platystomatids generally have tergite 6 of the
abdomen very short and capable of being withdrawn
underneath the margin of tergite 5, or absent. Thus at most
only four tergal (dorsal) abdominal plates are visible in front
of the ovipositor sheath (the first two tergites being fused).
By contrast, female pyrgotids have tergite 6 well developed
and resembling tergite 5, though much shorter. Very few
Australasian platystomatid species have female tergite 6
permanently exposed (. Apiola , some Angitula spp., one
Duomyia sp.). The first two of these have characteristic
wings (Figs. 123, 88), and the latter localised species is
unlikely to be encountered before otherwise closely similar
species are seen.

While there are numerous other morphological features
of the Platystomatidae that can help to distinguish the family,
these may be largely dispensable to those simply wishing
to sort material. Reference may be made to the more detailed
work of Me Alpine (1973a) if required.

Subfamily classification

A classification of the Platystomatidae into five subfamilies
was proposed by McAlpine (1973a). Some aspects of this
classification seem unsatisfactory in the light of morpho¬
logical study of additional taxa.

Freidberg (1994, only printed abstract published) seems
to have raised once again the possibility of relationship
between “Phytalmiini (Tephritidae), [and] Angitulinae
(Platystomatidae)”. Hennig (1940a) disposed of this
supposed alliance as convergence, and our more recent
studies (McAlpine & Schneider, 1978) confirm the tephritid
affinities of the Phytalmiini, without revealing any evidence
of polyphyly for the Tephritidae.

In characterising Platystomatinae and Angitulinae as
separate subfamilies (McAlpine, 1973a), I inclined to regard
the relatively well-developed female tergite 6 as a
plesiomorphy in the ground plan of Angitulinae, and the
greater reduction of this tergite in Platystomatinae as a
unifying apomorphy (probable autapomorphy) for the
latter (though it is shared with the Scholastinae and

Plastotephritinae). Hence, with reference to the well
defined autapomorphies of Angitulinae, both could be
treated as probably monophyletic groups. Further study
convinces me that, even if the better developed tergite 6
is a groundplan condition for the Angitula alliance, it is
unlikely to represent a plesiomorphy relative to the
groundplan for Platystomatinae (sensu McAlpine, 1973a).

In relation to this problem, consideration of variation in
female tergite 6 in the genus Duomyia is instructive. I have
examined females of c. 90 of the species of this genus. With
one exception, all species seen have a much reduced tergite
6 which becomes concealed under tergite 5 in dried
specimens. The exceptional species (. Duomyia “sp.18” in
AM, numerous females examined) has a well-developed
tergite 6, which is non-retractile, quite broad laterally, with
setulose surface, and enlarged posterior marginal bristles.
Within the Platystomatidae, such a condition resembles the
presumed plesiomorphic state manifested in the outgroups
Pyrgotidae and Tephritidae. Three possible phylogenetic
interpretations are theoretically possible. (1) Duomyia
“sp.18” represents a primitive sister group to the greater
part of the Platystomatinae, which is synapomorphic in this
character. (2) Duomyia “sp.18” is related to other taxa with
closely similar general morphology, but these other taxa
(many of which are placed in the subfamily Platystomatinae
and a number in the genus Duomyia ) have attained the
apomorphic state by multiple convergence. (3) The
condition in Duomyia “sp.18” is due to an evolutionary
reversal in its own peculiar lineage, and is effectively a
species-level autapomorphy within the broad taxonomic
field of Platystomatinae, which are characterised by the
almost uniform homologous condition of a reduced,
retractile female tergite 6. I reject interpretation (1) as
extremely improbable in view of the apparently close
relationship of Duomyia “sp.18” to D. glebosa McAlpine
and a group of closely related undescribed species.
Interpretation (2) must be rejected as passing far beyond
the requirements of theoretical parsimony into the realm of
fantasy. Interpretation (3) must therefore be accepted by
elimination of alternatives.

Accepting that the reduction tendency in female tergite
6 of Platystomatidae has been markedly reversed within
Duomyia , I now regard it as possible/probable that the
relatively well-developed tergite 6 in the trapherine genera
Xiria Walker and Phasiamya Walker and the “angituline”
subgenera Angituloides and Giraffomyia is also a
secondarily derived condition. In those numerous
platystomatid groups which show heterogeneity in
development of female tergites 4-5, evolution of these
structures also may not necessarily be unidirectional.

Clearly, the Angitulinae must be incorporated into the
Platystomatinae for reasons given below in the discussion
of Terzia n.gen. This means that all taxa in which the
aedeagus is known to possess a pair of hollow terminal
filaments with terminal gonopores are now placed in the
Platystomatinae, and this is considered to be a groundplan
condition for the subfamily.

Delimitation of the remaining three subfamilies is
problematical. Australasian genera sometimes placed in the
Plastotephritinae are Chaetorivellia, Atopognathus, and

David K. McAlpine: Australasian Platystomatidae 121

Guamomyia, but A. Whittington has indicated (pers.
comm.), from his studies of this mainly Afrotropical
subfamily, that these assignments may be incorrect.
Chaetorivellia is here assigned to the Scholastinae, though
it is in incomplete agreement with the characterisation
previously given (D. McAlpine, 1973a). Atopognathus and
Guamomyia are here categorized as of doubtful subfamily
position, together with Apiola which is poorly known
morphologically and certainly not close to the “angituline”

genera, now placed in Platystomatinae. Thus no Australasian
genera now remain in the Plastotephritinae.

The Scholastinae and Trapherinae are discussed
separately below. As aedeagal structure seems to be of major
significance in the subfamily classification of platy-
stomatids, it is possible that correlative study of reproductive
biology may help interpretation of phylogeny. I also point
out that gross morphology of the female reproductive system
is known for very few genera.

Key to Australasian genera of Platystomatidae

1 Stem vein (base of R, Fig. 1, sv) dorsally before level of humeral

crossvein with numerous setulae. 2

-Stem vein without setulae before level of humeral crossvein

(setulose beyond). 5

2 Fronto-orbital bristles distinct; scutellum devoid of hairs; squama

very narrow; New Guinea. Bama part

-Fronto-orbital bristles absent or vestigial; scutellum with few or

many dorsal hairs; squama large and rounded (Fig. 1, sq). 3

3 Face with strongly raised flat-topped carina; mesopleural bristle
usually absent; 6 : aedeagus with pair of terminal filaments (e.g.,
Figs. 24-26); ?: abdominal tergite 5 long, or short but well

sclerotised and exposed. 4

Face without median carina; mesopleural bristle present; 6 :
aedeagus without paired terminal filaments (e.g., Fig. 106); ?:

abdominal tergite 5 vestigial; widely distributed. Pterogenia

4 Mesoscutum, mesopleuron, and abdominal tergites 2 to 5 largely
glossy and almost without pruinescence; second basal cell largely
bare; wing hyaline, with three small transverse brown stripes and
very small apical spot on vein 3; New Guinea (one aberrant

species). Lamprogaster part

-Mesoscutum, mesopleuron, and/or tergites 2 to 5 almost entirely

densely pruinescent; second basal cell usually largely micro-
trichose; wing with more extensive dark markings than indicated

above; widely distributed. Euprosopia

5 Eye densely haired; scutellum extensively haired dorsally; axillary
lobe with fringe of long hairs, each >4 times as long as pubescence
on vestigial squama; ?: aculeus broad and blade-like; widely

distributed. Atopognathus

-Eye bare or with sparse minute hairs; other characters not entirely

as above. 6

6 Metathorax with complete, deep, glossy postcoxal bridge (Fig.

90, pcb); humeral callus hairless or almost so; mesoscutum with
hairing mainly reduced to few longitudinal series; outer vertical,
ocellar, humeral, supra-alar, posterior intra-alar, dorsocentral, and
mesopleural bristles absent; legs long; mid-femur neither thickened

nor ventrally spinose; squama vestigial, without lobe. 7

-Metathoracic postcoxal bridge usually absent, if complete, dull-

pruinescent; other characters not entirely as above. 8

7 Scutellum without setiferous horns; pronotum very short and

inconspicuous; thorax fulvous; Solomon Archipelago. Terzia

Records of the Australian Museum (2001) Vol. 53

-Scutellum with pair of setiferous horns (Fig. 90, ss); pronotum

enlarged, variously modified; thorax shining (often metallic) black;

Moluccas, New Guinea, Bismarck and Solomon Archipelagos. Angitula

8 Mid femur thicker than other femora, with strong ventral spines;

other femora without ventral spines. 9

-Mid femur usually not thicker than other femora, if ventrally

spinose then fore femur and/or hind femur also spinose. 10

9 Pronotum transversely narrow-linear, recessed into front of thorax
and scarcely visible from above; mesoscutum much broader than
long; antennal sockets rather broadly separated by at least the

width of antennal segment 1; widely distributed. Mesoctenia

-Pronotum broadly visible from above, not recessed; mesoscutum

not broader than long; antennal sockets usually separated by less

than half width of segment 1; widely distributed. Brea

10 Fronto-orbital bristles absent or not distinctly differentiated. 11

-Fronto-orbital bristles distinct, though often small. 33

11 Arista densely whitish-pubescent for almost whole length; antenna
excluding arista longer than face on median line; face with minute

erect hairs (Fig. 62). 12

-Arista almost bare, or short-haired on basal part only, or loosely

haired to apex, or bipectinate; other characters not as above. 13

12 First basal cell broad at level of basal crossvein, narrowed near
middle by curvature of vein 4 (Fig. 61); penultimate section of

vein 4 very short, straight; Moluccas, New Guinea, Philippines. Antineura

-First basal cell very narrow for most of length, widened near

anterior crossvein; penultimate section of vein 4 longer, curved,

dipping into discal cell; Sulawesi, not Australasian. Philocompus

13 Mesopleural bristle absent. 14

-Mesopleural bristle present. 20

14 Anal crossvein strongly bent at anterior third of length; section of
costa on subcostal cell more than four times as long as that on
marginal cell; S : costa basally inflated, eliminating first costal

cell (Fig. 82); New Guinea. Cleitamoides

-Anal crossvein curved or almost straight; section of costa on

subcostal cell not much longer than that on marginal cell; d': costa

not inflated basally. 15

15 Arista long-haired, at least on mid section and usually for most of

length, hairs tending to alignment in a dorsal and a ventral series
(bipectinate condition); often some or all femora with ventral

spinescent bristles but mid femur always without posterior bristles. 16

-Arista at most short-haired on basal part (hairs not much longer

than basal diameter), usually bare on distal half, or, if relatively
long haired, then mid femur with posterior bristles on distal half
(few Duomyia spp.); femora usually without ventral spinescent
bristles, except sometimes for a posteroventral series on fore femur

17

David K. McAlpine: Australasian Platystomatidae

16 Antennal segment 2 rotund, cap-like; mid femur much thicker
than other femora; squama vestigial, not forming lobe; <$: eye
never stalked or laterally protruding; hom-like cheek process often
present; ?: abdominal tergite 6 well developed, exposed, but

shorter than tergite 5; Samoan Islands. Apiola

—— Antennal segment 2 subconical to subcylindrical, more or less
attenuated basally; squama forming a distinct broad to narrow
lobe; mid femur not thicker than other femora; S : eye often stalked
or laterally protruding; cheek process absent; $: tergite 6 vestigial,
generally concealed; New Guinea, Bismarck Archipelago,

northern Australia. Achias

17 Basal crossvein very oblique, its distal end aligned with
penultimate section of vein 5; postfrons usually with pronounced
central tubercle; antennal segment 3 usually potentially extending
far beyond lower margin of face on median line; second section
of vein 4 at least three times as long as first; Moluccas, Oriental

Region etc. Lamprophthalma

Basal crossvein transverse to somewhat oblique, meeting

penultimate section of vein 5 at an angle; other characters variable. 18

18 Squama vestigial, not forming a lobe; mesoscutum with transverse
depression crossing medial line (evident in profile), much longer
than wide; face somewhat saddle-like but without carina; scutellum
with two pairs of bristles, bare medially but setulose laterally;
suprasquamal ridge without erect hairs or setulae; mid femur
neither bristled nor more conspicuously haired on posterior

surface; widely distributed. Elassogaster part

-Squama forming a definite, often large lobe; mesoscutum without

complete transverse depression; other characters not in above

combination. 19

19 Suprasquamal ridge without hairs (sometimes with short
pubescence); supra-alar and prescutellar acrostichal bristles absent;

mainly New Guinea species. Lamprogaster part

-Suprasquamal ridge with erect hairs (Fig. 1, sr); supra-alar and

prescutellar acrostichal bristles often present; Australia. Duomyia part

20 Prelabrum very deep, ventrally much extended towards proboscis;
scutellar bristles only one pair or none; occiput very convex or

prolonged; habitus elongate, ant-like; Queensland. Inium part

-Prelabrum not thus prolonged ventrally; other characters not

entirely as above. 21

21 Antennal segment 3 with numerous black setulae on inner surface;
suprasquamal ridge with erect hairs; parafacial with fine hairs
(distinct from pruinescence) near centre; thorax markedly

elongate; Moluccas, Oriental Region. Conicipithea

—— Antennal segment 3 without setulae, with only the usual pile-like
(under low magnification) vestiture; suprasquamal ridge without

erect hairs (often pruinescent); other characters variable. 22

22 Face with strongly raised, sharply margined, flat-topped carina;
squama very broad and rounded; posterior bridge of hind coxa

without fine hairs; widely distributed. Lamprogaster part

-Without above combination of characters; face rarely with such

carina in which case squama is narrow or vestigial. 23

124 Records of the Australian Museum (2001) Vol. 53

23 Mesoscutum wider than long; $: abdominal tergites 4 and 5 much

reduced (less so in Asyntona) . 24

—— Mesoscutum not wider than long; $ : abdominal tergites 4 and 5

large and exposed. 27

24 Anal cell acute at posterodistal angle; vein 4 ending at extreme
apex of wing, its combined sections on discal cell much shorter

than section on second basal cell (Fig. 121); widely distributed. Asyntona part

-Posterodistal angle of anal cell not acute; vein 4 ending a little

behind apex, its sections on discal cell not shorter than that on

second basal cell. 25

25 Face with deep transverse central depression, highly convex and
glossy below; antennal segment 3 at least four times as long as
wide; abdominal tergites 2 and 3 with prominent median carina;

Moluccas, Oriental Region. Trigonosoma

-Face little depressed centrally, not noticeably convex below;

antennal segment 3 no more than three times as long as wide;

abdominal tergites 2 and 3 without median carina. 26

26 Antennal sockets separated by c. width of each or less; face almost
entirely densely pruinescent; lower end of facial ridge with group
of hairs not in a linear series; mesoscutum and dorsal surface of
scutellum minutely roughened or pruinescent between hairs; New

Guinea, Oriental Region. Neohemigaster

-Antennal sockets separated by c. twice width of each or more;

face largely smooth and glossy; lower end of facial ridge with
few hairs in a linear series; mesoscutum and dorsal surface of

scutellum shining, almost smooth between hairs; Moluccas. Zygaenula

27 First posterior cell not narrowed apically; distal section of vein 4
not curving forwards apically; parafacial with fine hairs near

middle. 28

-First posterior cell narrowed apically; distal section of vein 4

curved anteriorly towards apex; parafacial haired at most only on

upper extremity (often pruinescent). 29

28 Mesoscutum without pattern of numerous black spots; wing
generally with brown markings; mesopleuron generally grey-
pruinescent on entire posterior margin; widely distributed, not in

Australia. Pseudepicausta part

-Mesoscutum with many black spots on a grey-pruinescent field

(pattern not visible in greasy specimens); wing generally without
dark markings; mesopleuron grey-pruinescent only on upper part;

Australia, Oriental Region. Rhytidortalis part

29 Face with fine hairs on central part; postscutellum setulose at sides;
a dark longitudinal stripe covering first basal cell (Fig. 56), or

wing with extensive infuscation; widely distributed. Plagiostenopterina part

-Face not haired; postscutellum without setulae; wing with neither

dark stripe on first basal cell nor general infuscation. 30

30 Fore femur with strong dorsal bristles; arista plumose or
subplumose on basal part; scutellum not haired dorsally; Fiji,

Oriental Region. Meringomeria

-Fore femur without distinct dorsal bristles; arista bare or with short

pubescence; scutellum variable. 31

David K. McAlpine: Australasian Platystomatidae

31 Scutellum without hairs on central dorsal part, usually with few
lateral hairs; vein 4 usually ending slightly in front of extreme
apex of wing (Fig. 49); subcosta without ventral setulae on basal
section; S (where known): aedeagus with pair of terminal

filaments; widely distributed. Elassogaster part

-Scutellum extensively haired dorsally; vein 4 ending at least

slightly behind extreme apex; other characters variable. 32

3 2 Subcosta with a series of small black ventral setulae before humeral

crossvein; second basal cell less than half as long as discal cell;
distal section of vein 5 approximately aligned with penultimate
section; posterior bridge of hind coxa with fine hairs; 6 : aedeagus

with pair of subequal terminal filaments; Bismarcks. Par part

-Subcosta without ventral setulae (microtrichose only); second

basal cell more than half as long as discal cell; distal section of
vein 5 somewhat bent posteriorly from junction with discal
crossvein; posterior bridge of hind coxa hairless; 6 : aedeagus

with only one terminal filament (Fig. 48, f); widely distributed. Microepicausta

33 Sternopleural bristle present; $: abdominal tergite 5 reduced. 34

-Sternopleural bristle not distinct from surrounding setulae; $:

abdominal tergite 5 variable. 36

34 Scutellum strongly convex, entirely glossy black, without paler
markings, setulose only on dorsal surface above level of lateral

bristles; wing with large recurved black band (Fig. 94); Australia. Lenophila part

-Scutellum slightly convex, dorsally pruinescent, with yellowish

U-shaped stripe or more extensively yellowish tawny, setulose
laterally as well as dorsally; wing markings not including a

recurved black band. 35

35 Ventral margin of face not reflexed; scutellum without a
ventrolateral series of bristles; first section of vein 4 shorter than
second section (Fig. Ill); basal section of vein 5 bare; widely

distributed. Scholastes

-Ventral margin of face narrowly reflexed, thus facing ventrally;

scutellum with a ventrolateral series of bristles; first section of
vein 4 longer than second section (Fig. 108); basal section of vein

5 dorsally setulose; New Guinea, Afrotropical Region. Paryphodes

36 Part of vein 4 on discal cell shorter than that on second basal cell;

squama very broad; form subglobose. 37

—— Part of vein 4 on discal cell longer than that on second basal cell;

squama variable; form usually more elongate. 38

37 Anal cell acutely or subacutely pointed at posterodistal angle; vein
4 ending in wing apex (Fig. 121); posterior margin of scutellum
thin, with numerous bristles intergrading with setulae;
mesopleuron, in dorsal view, prominently gibbous anteriorly;

widely distributed. Asyntona part

-Anal cell obtuse posterodistally; vein 4 usually ending behind

wing apex (Fig. 117); posterior margin of scutellum thick, rounded;
scutellar bristles in 2 or 3 well differentiated pairs; mesopleuron
without dorsally visible anterior gibbosity; widely distributed ....

Naupoda

Records of the Australian Museum (2001) Vol. 53

38 Section of costa on subcostal cell more than five times as long as

that on marginal cell. 39

-Section of costa on subcostal cell not more than three times as

long as that on marginal cell. 40

3 9 Anal crossvein strongly bent near middle, its posterior part running
basad into vein 6 so that anal cell is without an angle at junction;

New Guinea. Cleitamia

-Anal crossvein recurved near middle, then flexed distad posteriorly

to form a short acute lobe to anal cell where it joins vein 6 (as in

most Tephritidae, Fig. 85); New Guinea. Loriomyia

40 First basal cell at least as wide as combined width of second basal
and anal cells; posterodistal angle of discal cell acute (Fig. 59);

New Guinea. Pseudo cleitamia

-First basal cell narrower than combined width of second basal

and anal cells; venation otherwise variable. 41

41 Hind femur much thicker than other femora, with two series of
stout ventral spines; scutellum with three pairs of marginal bristles

and no hairs. 42

-Hind femur not significantly thicker than other femora, without

seriate spines; scutellum with variable vestiture. 43

42 First posterior cell divided by a complete supernumerary crossvein;
second section of vein 4 (before anterior crossvein) much longer
than first (Fig. 54); fore femur with a series of stout ventral spines;

Samoan Islands. Apactoneura

-First posterior cell without supernumerary crossvein; second

section of vein 4 shorter than first (Fig. 20); fore femur without
ventral spines; southern Micronesia, eastern Melanesia, tropical

Polynesia. Pseudorichardia

43 All femora strongly thickened, fusiform, without ventral bristles
or spines; upper pleurotergite (above callus) with many long fine

hairs; lateral occipital bristle present; Sulawesi, not Australasian. Scelostenopterina

-Femora of normal shape, or only hind femur of male modified;

upper pleurotergite not haired, except in some Laglaisia spp. with

slender femora and no lateral occipital bristle. 44

44 Arista bipectinate (plumose) for most of its length. 45

-Arista with non-seriate short hairs or almost bare. 48

45 Anal crossvein strongly subangularly bent near middle (Fig. 79);
postfrons broader than long; scutellum usually with fine ventral

setulae; eyes of male usually stalked; New Guinea. Laglaisia

—— Anal crossvein simply curved or almost straight; postfrons usually
longer than broad; scutellum without ventral setulae, but often

densely pubescent; eyes not stalked. 46

46 Wing membrane with many pale dots on darker field; membrane
in discal cell with sharp longitudinal crease; one or two scapular
bristles present; S : aedeagus with pair of terminal filaments;

Sulawesi, not Australasian. Euthyplatystoma

-Wing membrane without pale dots, without sharp crease enclosed

in discal cell; scapular bristles absent; S : aedeagus without

terminal filaments. 47

David K. McAlpine: Australasian Platystomatidae

47 Squama narrow-linear, not forming a lobe; scutellum very broad,
without setulae, with metallic green reflections; vein 2 less than
half as long as vein 3; anterior crossvein meeting vein 4 near distal

end of discal cell (Fig. 122); Fiji. Phlyax

-Squama forming a distinct rounded lobe; scutellum of moderate

width, dorsally setulose, not metallic green; vein 2 more than half
as long as vein 3; anterior crossvein meeting vein 4 near mid¬
length of discal cell (Fig. 97); Moluccas, New Guinea, Solomon

Archipelago. Chaetorivellia

48 Median part of face with fine hairs; lateral occipital bristle present;
wing with dark longitudinal streak entirely covering first basal
cell (Fig. 56); mesoscutum markedly longer than wide; New

Guinea, Queensland, Moluccas. Plagiostenopterina part

—— Median part of face bare; other characters not entirely as above.49

49 Postscutellum with group of setulae on each side (Fig. 52, ps);
distal section of vein 4 from its origin to apex convergent with
vein 3; costal margin with black spot on end of vein 2 but none at

apex (Fig. 50); scutellum haired dorsally on entire width; Australia. Aetha

Postscutellum without setulae; other characters not entirely as

above. 50

50 Lower part of facial ridge with dense short black setulae; lower
anterior margin of cheek with a series of longer peristomial setulae,
the foremost approximating in position to a vibrissa (Fig. 126);
hind femur with long anteroventral bristles; axillary lobe with
fringe of long hairs, each >4 times as long as pubescence on the
vestigial squama; Lord Howe I, New Guinea, Bismarck

Archipelago, Micronesia. Guamomyia

Facial ridge with fine usually uniseriate hairs only; lower margin
of cheek with setulae mainly on posterior part (these often very

long in Pogonortalis)', other characters variable. 51

51 Anterior crossvein very oblique, no shorter than and almost aligned
with penultimate section of vein 4 (Fig. 28); face with flat-topped
sharply margined carina; arista rather densely short-haired for most

of length; mesoscutum as long as wide; Loyalty Is. Montrouziera

—— Anterior crossvein not as above or other characters not entirely as

above. 52

52 Prelabrum deep and much prolonged ventrally towards proboscis,
not arcuate in ventral view but sclerotised across full width; small,

shining black ant-like flies; Queensland. Inium part

-Prelabrum not thus prolonged towards proboscis, arcuate to

horseshoe-shaped in ventral view; habitus various. 53

53 Scutellum dull brown with yellowish U-shaped stripe, setulose
on entire dorsal surface and on ventrolateral surface below
marginal bristles; lower end of facial ridge with numerous non-
seriate setulae covering a broad field (Fig. 98); axillary lobe and
squama both moderately large; first section of vein 4 longer than

second section; New Guinea. Tomeus

Scutellum without yellowish U-shaped stripe; other characters not

entirely as above. 54

Records of the Australian Museum (2001) Vol. 53

54 Squama notably larger in area than axillary lobe. 55

-Squama vestigial or not much larger than axillary lobe. 57

55 First costal cell largely bare; face without central carina; 9: tergite

4 c. l A as long as tergite 3; tergite 5 still shorter; New Guinea. Bama part

-First costal cell entirely microtrichose; face usually with broad

flat-topped carina between antennal grooves; 9: tergites 4 and 5

not much reduced. 56

56 Suprasquamal ridge with erect hairs or setulae (Fig. X, sr);
mesopleural bristle usually absent, if present, small; 8 : abdomen

usually subcylindrical; Australia. Duomyia part

-Suprasquamal ridge with minute pubescence only; mesopleural

bristle well developed; 8 : abdomen usually broadly ovate; widely

distributed. Lamprogaster part

57 Scutellum haired on central part of dorsal surface (hairs reduced

to minute stumps in male of one Rivellia sp.). 58

-Scutellum without hairs on central part of dorsal surface,

sometimes with a few hairs at sides. 63

58 Marginal cell much attenuated on distal half; hind femur longer
than mesoscutum; second section of vein 4 (before anterior
crossvein) less than half as long as third section; distal (fourth)
section of vein 4 strongly converging with vein 3 apically (Fig.

18); Loyalty Is. Tarfa

-Marginal cell not much attenuated distally; other characters not

entirely as above. 59

59 Dark band on discal crossvein, when present, quite separate from
any more basally located dark band or zone; two pairs of scutellar
bristles present; 8 : shape of hind tibia not modified; 9: abdominal

tergites 4 and 5 large. 60

-Dark band from apex of vein 5 passing obliquely forward to costa

on subcostal cell where it connects with more basally located dark
band or zone (Figs. 13, 94); three pairs of scutellar bristles present;

8 : hind tibia usually strongly modified in shape; 9: tergites 4

and 5 variable. 62

60 Veins 3 and 4 apically convergent; subcosta with small ventral
setulae before level of humeral crossvein; posterior bridge of hind

coxa with small hairs; Bismarck Archipelago. Par part

-Veins 3 and 4 not apically convergent; subcosta without ventral

setulae; posterior bridge of hind coxa hairless. 61

61 Second section of vein 4 rather strongly concavely curved on distal
part (Fig. 14); postgenal fold absent; height of cheek generally
less than 0.2 of height of eye; antenna (without arista) about as
long as face on median line, at least in male; widely distributed,

but absent from New Zealand. Rivellia

-Second section of vein 4 with only slight curvature; deeply incised

vertical postgenal fold present (Fig. 17, pf); height of cheek greater
than 0.2 of height of eye; antenna much shorter than face; Australia,

New Zealand

Zealandortalis

David K. McAlpine: Australasian Platystomatidae

62 Ground-colour of thorax almost entirely black; lateral occipital
bristle absent; second costal cell partly hyaline; S : aedeagus

without terminal filaments (Fig. 95); ?: abdominal tergite 5
reduced; Australia.

. Lenophila part

-Ground-colour of thorax largely orange-tawny; distinct but fine

black lateral occipital bristle present; second costal cell entirely dark
brown; 3 : aedeagus with pair of terminal filaments (Fig. 12, f); $:
tergite 5 large; Australia.

. Loxoneuroides

63 Height of cheek immediately below eye less than one tenth height
of eye; hind femur with anteroventral keel or slightly developed
tooth at or near distal third; anterior crossvein meeting vein 4
before mid-length of discal cell (Fig. 53); widely distributed.

. Pogonortalis

—— Height of cheek below eye more than one tenth height of eye;
hind femur without anteroventral keel or tooth; anterior crossvein
generally meeting vein 4 at or beyond mid-length of discal cell.

. 64

64 Antennal segment 3 rounded oval, not more than c. twice as long
as deep on outer exposure (Fig. 11); mesoscutum scarcely longer
than wide; S : prelabrum creamy, and either hypertrophied,
anteriorly flattened and shield-like, or vestigial and largely
desclerotised, in the latter case palpus much broadened; New
Guinea, Australia.

. Hysma

-Antennal segment 3 c. three times as long as deep or longer;

mesoscutum variable in proportions; S : prelabrum with none of
above modifications.

. 65

65 Parafacial with fine socket-based hairs on central part, often in

addition to pruinescence.

. 66

-Parafacial with hairs restricted to upper part near confluence with

postfrons, or absent.

. 68

66 Second section of vein 4 gently arched (Fig. 40); abdominal tergite

4 with broad lateral zone of dense non-directional white
pruinescence; except in New Caledonian species, vein 1 ventrally
near end of subcosta with a series of short black setulae, and alula
largely bare; Queensland, New Guinea, Bismarck Archipelago,

New Caledonia.

. Eumeka

-Second section of vein 4 not arched, straight or with concave

curvature; abdominal tergite 4 either without pale pruinescence,
or with pale pruinescence not covering lateral marginal zone and
usually unidirectional (i.e. pale zones not apparent from some
angles); vein 1 without ventral setulae; alula entirely microtrichose.

. 67

67 Mesoscutum extensively pale grey-pruinescent, with black dot at
base of each hair; hind basitarsus distinctly less than half as long
as hind tibia; wing, in Australasian species, without dark markings;

$: ovipositor sheath not much longer than tergite 5; Australia,
Oriental Region.

. Rhytidortalis part

-Mesoscutum not extensively pale grey-pruinescent, sometimes

with pale grey median stripe; hind basitarsus at least half as long
as tibia; wing with apical brown mark; 9: ovipositor sheath usually
at least twice as long as tergite 5; New Guinea, etc.

. Pseudepicausta part

130 Records of the Australian Museum (2001) Vol. 53

68 Mid femur with one isolated preapical posterior bristle; a
dorsocentral bristle situated almost halfway between scutellar
suture and transverse suture, and usually, behind this, a bristle
intermediate in position between dorsocentral and prescutellar

acrostichal; anterior and discal crossveins enclosed in the one dark

band (Fig. 72); New Guinea, Bismarck Archipelago, Queensland. Euxestomoea

If mid femur with posterior bristles, then these merging with series
of long hairs; dorsocentral bristle much closer to scutellar suture;

wing markings various. 69

69 Second section of vein 4 with sigmoid curvature on distal part;
anterior crossvein meeting vein 4 well beyond middle of discal
cell; capitellum of halter dark brown; mesoscutum without
noticeable grey pruinescence; $: abdominal tergites 4 and 5

vestigial, usually concealed in dried specimens; New Guinea. Bama part

Second section of vein 4 without sigmoid curvature; 9: abdominal
tergites 4 and 5 well developed, exposed; other characters not

entirely as above.70

70 Mesoscutum without noticeable grey pruinescence between hairs,
the interstices sometimes roughened; brown costal mark restricted
to subcostal cell or absent; a shining median quadrate sclerite
present between face and prelabrum (few aberrant species, e.g.,

Fig. 63); Australia. Duomyia part

-Mesoscutum with much grey pruinescence, often forming

longitudinal bands; brown costal mark well developed at wing
apex; sclerite between face and prelabrum absent or not distinctly

developed. 71

71 Section of costa on subcostal cell subequal in length to that on
second costal cell; distal section of vein 4 not perceptibly curved
forward apically (Fig. 38); posterior bridge of hind coxa hairless;
scutellum with two or three pairs of major bristles, progressively

longer posteriorly, and no setulae; Australia. Scotinosoma

-Section of costa on subcostal cell much longer than that on second

costal cell; distal section of vein 4 curved forward apically,
converging with vein 3 (Fig. 42); posterior bridge of hind coxa
with fine hairs; scutellum with two pairs of major bristles and one

or few setulae on each side; New Caledonia. Signa

Subfamily Platystomatinae Genus Hysma n.gen.

This subfamily is much the largest of the family. It is here
further expanded to include the subfamily Angitulinae,
which becomes a synonym, for reasons discussed above
and under genus Terzia.

The groundplan of the Platystomatinae includes the
characteristic aedeagus, with sclerotised, capsule-like glans,
and a pair of hollow terminal filaments, each with an apical
gonopore, but it is not certain if the presence of paired
terminal filaments is an autapomorphy for the subfamily,
as it is also feasible that the condition may have been
secondarily lost in taxa outside this subfamily. However,
the only known platystomatine taxa without this condition
have a structure clearly derived therefrom. For instance,
some species of Brea and Lamprogaster have three instead
of two terminal filaments; at least one species each of
Bromophila Loew and Duomyia have the two filaments
fused for all or most of their length, and in Micro epic austa
there is a single terminal filament.

Type species: Hysma lacteum n.sp.

Description (<J, $). Small or moderately small robust flies,
with legs of moderate proportions, as in stouter Rivellia spp.

Head much higher than long; face concave, without distinct
median carina, with slightly sunken antennal grooves; facial
ridge with single series of setulae on lower part only;
parafacial narrowed on central part,without setulae; occipital
region broadly convex below, slightly convex above;
postgenal fold absent; cheek rather shallow or of moderate
depth; the following bristles present: inner and outer
vertical, small or vestigial postvertical, variably developed
non-proclinate ocellar, two fronto-orbitals, postgenal; lateral
occipital sometimes present. Antenna much shorter than
face; segment 3 broadly oval, rounded distally; segment 6
with short pubescence of moderate density on entire length.
Prelabrum sexually dimorphic, in female large or

David K. McAlpine: Australasian Platystomatidae 131

moderately large, in male very large with anterior surface
flattened ( Hysma lacteum ), or reduced and apparently little
sclerotised; palpus moderate to large, in males of species
other than H. lacteum subtriangularly broadened.

Thorax. Mesoscutum c. as broad as long or slightly broader,
extensively setulose,with surface between hairs varying
from largely glabrous and glossy ( H. lacteum ) to extensively
pruinescent; scutellum rounded in outline, dorsally convex,
almost glabrous or pruinescent, without setulae; the
following bristles present: poorly differentiated scapulars,
humeral, 1+1 notopleurals, supra-alar, postalar, posterior
intra-alar, one dorsocentral, prescutellar acrostichal, three pairs
of scutellars, of which intermediate one slightly removed from
margin, mesopleural, sometimes pteropleural. Posterior bridge
of hind coxa without hairs; femora without ventral spines;
fore femur with dorsal and anteroventral bristles; mid femur
with or without a series of short posterior bristles; hind femur
usually with distinct distal dorsal bristles; mid tibia with
one large apical ventral spur. Wing generally similar to that
of Rivellia; veins 3 and 4 distally slightly divergent; cell-4
index = c. 0.6-0.8; second section of vein 4 less curved
than in Rivellia’, distal section of vein 6 well sclerotised
almost to margin, or sclerotisation discontinued at c. 0.7 of
distance to margin; wing membrane, including costal cells
and alula, microtrichose, except sometimes on much of second
basal and anal cells; squama forming a very small lobe; axillary
lobe slightly larger in area.

Abdomen. Tergite 5 unreduced in either sex, without strong
bristles. Female abdomen: spiracles 4 and 5, where investigated,
located in pleural membrane near posterior part of lateral
margin of respective tergites; aculeus very slender, obtuse.

Distribution. Eastern Australia: high rainfall areas of north
and south. Papua New Guinea: only known from Morobe
Province.

Notes. There are three or, perhaps, four species in the
available material of Hysma, but only the type species is
described at present.

Probably all species have remarkable sexual dimorphism
of the face, prelabrum, and palpus, which are much modified
in males. In H. lacteum the male has the prelabrum enlarged,
with flattened, conspicuously pale creamy anterior surface,
and the face similarly coloured (these parts being brown in
the female), the palpus brown-black but no larger than the
tawny-orange palpus of the female. In the other two species
for which the males are known, the face of the male is paler
and foreshortened below, relative to that of female, so that
the pale, translucent, anterior extension of the subcranial
membrane is broadly exposed above the small, desclerotised,
pale yellow prelabrum (these parts unmodified in female).
Also, these males have the palpus conspicuously broadened
and subtriangular, in contrast to the unmodified palpus of
the females.

A series of Hysma sp. from Bulburin (Monto district,
Queensland, AM) shows extraordinary difference in wing
pattern between the two males and the 12 females. Without
further study material I cannot be sure if this indicates

unusual sexual dimorphism, or the presence of two species,
each represented by one sex in the sample. In a pair of a
very similar species from near Bulolo, Papua New Guinea
(AM), the wing pattern of both sexes resembles that of the
Bulburin females.

In general, Hysma spp. are found only in rainforest, but
the specimen of H. lacteum from Bodalla State Forest was
probably taken in wet sclerophyll forest. Several have been
collected in faecal traps.

The generic name is Greek for rain, and is neuter.

Hysma lacteum n.sp.

Figs. 10, 11

Material examined. Holotype, 9, New South Wales:
Upper Allyn (Lister Park), near Eccleston, 9.iii. 1970,
G.A.H., D.K.M. (AM). Paratypes, New South Wales: 1
6, Dorrigo National Park, Glades area, Jan. 1988, D.J.B.
(AM); 2 9 9, Upper Allyn, Dec. 1969, Mar. 1970, G.A.H.,
D.K.M. (AM); 1 9, Mount Keira, near Wollongong, Jan.
1970, G.A.H. (AM); 1 9, Bodalla State Forest, near
Narooma, Jan. 1982, B.J.D. (AM).

Other material. Queensland: Longland’s Gap, near Atherton
(AM); Sluice Creek, near Millaa Millaa (AM).

Description (c£, 9). Coloration. Head of female largely
tawny; upper orbits and sides of vertex dark brown; occiput
largely blackish, with silvery pruinescence in part; cheek
suffused with brown; face tawny. Head of male with
postfrons largely dark brown; face pale cream. Antenna in
male brown, with base of segment 3 tawny, in female
segments 1-3 all tawny. Prelabrum of male pale cream on
anterior surface, dark brown laterally, in female entirely
brown; palpus brown-black in male, tawny in female.
Thorax largely glossy black, with faintly blue-green tinged
reflections. Legs, including coxae and tarsi, dark brown to
black. Wing with brown to blackish markings, as in Fig.
10, which are slightly more extensive in male, particularly
the suffusion behind vein 5; wing apex between veins 3 and 4
subopaque white. Halter dark brown, tawny-brown basally.
Abdomen black to dark brown; tergites 1-5 largely glossy.

Head. Height of cheek c. 0.22 of height of eye; fronto-orbital
and ocellar bristles small. Prelabrum in female large, slightly
convex anteriorly, in male larger, nearly as deep as face,
well sclerotised, with anterior surface vertically flattened;
palpus of moderate size, not enlarged in either sex.

Thorax. Wing: vein 3 setulose dorsally for most of length, bare
ventrally; cell-4 index = 0.69-0.76; second basal cell bare on
c. basal three quarters; anal cell with longitudinal bare zone.

Abdomen. Male postabdomen: epandrium with few enlarged
setulae near base of outer surstylus; outer surstylus
moderately stout, its apex exceeding that of inner surstylus,
produced into a slender point posteroapically; aedeagus
broken in only available male.

132 Records of the Australian Museum (2001) Vol. 53

Dimensions. Total length: 8 2.7 mm, 9 $ 4.6-5.6 mm; length
of thorax, 8 1.3 mm, $ 9 1.9-2.6 mm; length of wing, 8 3.3
mm, 9 9 4.5-5.2 mm.

Distribution. Queensland: only known from higher parts
of Atherton Tableland. New South Wales: coastal and
subcoastal areas from Dorrigo district to Narooma district.

Notes. Hysma lacteum is distinguished from other,
undescribed species of Hysma by having the mesoscutum
and scutellum shining black, almost without pruinescence,
and the halter largely dark brown, instead of pale yellowish.

The specific epithet is a Latin adjective meaning milky,
in reference to the opaque white wing apex.

Genus Loxoneuroides Hendel

Figs. 12, 13

Loxoneuroides Hendel, 1914a: 15, 80-81. Type species (original
designation) L. varipennis Hendel.

Description. A detailed description has been given by
Hendel, which I supplement as follows.

Male abdomen. Sternite 5 very deeply bilobed; base of
preglans with elongate lobe, but preglans otherwise not
much differentiated from stipe; glans short and stout,

desclerotised on one side; bulb rather large and lobed;
terminal filaments equal, tapered, each not much longer than
glans, partly ensheathed by membranous bulb; cerci largely
separate, lying in almost parallel planes.

Female abdomen. Tergites 3 and 4 large; tergite 5 slightly
shorter; tergite 6 vestigial; spiracles 4 and 5 larger than
others, situated close to posterior part of lateral margins of
respective tergites; ovipositor sheath short; aculeus slender,
obtuse, not compressed.

Distribution. Eastern Australia, from Cooktown district,
Queensland, to Taree-Gloucester district, New South Wales.

Notes. Loxoneuroides includes at least three superficially
similar species, of which only L. varipennis Hendel is
described. Evenhuis (1989) used the spelling L. variipennis
of the two original spellings given by Hendel (1914a).
However, Hendel (1914b) consistently used the spelling L.
varipennis, thus acting as first reviser under Article 24.2.4
of ICZN (1999).

These flies inhabit rainforest, including dryer rainforests
in some areas, but are not recorded from littoral rain forests.
They frequently rest on foliage (generally on the lower
surface of leaves) or bark, with the wings fully extended
laterally. They can be so conspicuous in this position, that
aposematic display is suspected. Adults are attracted to
mammalian faeces for feeding.

Figures 12-13. Loxoneuroides “sp. 2” (Mulgrave River, AM). 12, distal part of aedeagus (scale = 0.2 mm). 13,
wing. Abbreviations: f, terminal filaments; g, glans; pg, preglans; st, stipe.

David K. McAlpine: Australasian Platystomatidae 133

Genus Rivellia Robineau-Desvoidy

Fig. 14

Rivellia Robineau-Desvoidy, 1830: 729. Type species (designated
Rondani, 1869) Musca syngenesiae Fabricius.

See Soos, 1984 for details.

Description. The description by Hendel (1914a) remains
largely valid. The generic and specific descriptions by Namba
(1956) give a useful indication of the range of morphological
variation in the species of America north of Mexico.

Distribution. Almost cosmopolitan. Australasian Region:
Moluccas; New Guinea; Bismarck Archipelago; Solomon
Archipelago; New Caledonia; Fiji; Samoan Islands; Austral
(Tubuai) Islands; Norfolk Island; Australia—all states and
Northern Territory (including central Australia). Rivellia has
much the widest distribution of any platystomatid genus,
both on a world basis and in the Australasian Region.

Notes. Namba (1956) gave an excellent review of the
Nearctic species, and Lyneborg (1969) characterised the
two European species. Taxonomic work on other faunas
has a quite inadequate morphological basis, so that
reidentifications and subsequent distribution records of
described species should in general be queried. Future
taxonomic studies should record details of male genitalia,
including measurements of the glans, and positions of spiracles
4 and 5 in the female abdomen. Descriptions of additional
species without this information are to be avoided.

Evenhuis (1989) listed 27 named species for the
Australasian Region. Accumulated collections contain many
undescribed species, but the number cannot be estimated
without a thorough study. An arbitrary minimum of 45
species for the Region is here utilised.

Larvae of Rivellia spp. Are recorded as feeding on the
nitrogen-fixing root nodules of leguminous plants in
Australia (Diatloff, 1965) and several other countries
(summary in Bibro & Foote, 1986). In Australia, the very
general distribution of Rivellia spp. is probably related to a
similar distribution of these plants (family Leguminosae or

Fabaceae s. 1.), perhaps particularly, but by no means
exclusively, the genus Acacia (wattles, subjective synonyms
Zygmoloba, Racosperma ). Though I have no direct evidence
of Rivellia larvae living on Acacia roots, adults are found
on Acacia plants in many parts of Australia. For some years
adults of a Rivellia sp. were plentiful in my back yard at
Willoughby, Sydney, and seemed particularly to inhabit a
large tree of Acacia saligna. When this tree died several
years ago, the Rivellia population disappeared. The broad
treatment of arthropods associated with Acacia by New
(1984) seems to indicate that very little has been recorded
of those infesting the root systems. Damage to the plants
from injury to the roots has probably not been generally
diagnosed, and there is a possibility that Rivellia spp. may
have significant ecological impact on Acacia populations.
Perhaps larvae of some other platystomatine genera frequently
found in Acacia-dominated habitats, e.g., Duomyia,
Microepicausta, Rhytidortalis, and Zealandortalis, are
associated with Acacia roots.

A female specimen of Rivellia sp. from Jervis Bay, A.C.T.
(AM), carries an ascoma of an ectoparasitic fungus of the
order Laboulbeniales on its abdomen. I have seen a
somewhat similar fungus on a female of Senopterinafoxleei
Shewell from British Columbia, Canada.

Genus Zealandortalis Malloch

Zealandortalis Malloch, 1930a: 243. Type species (original
designation) Z. interrupta Malloch.

Description. Flies of medium to rather stout build and
moderately small size, with legs of moderate length.

Head markedly or only slightly higher than long; facial
carina distinct and narrow, or undeveloped; facial ridge with
a distinct series of hairs only near lower end; parafacial
haired only at upper extremity, or with few hairs extending
to middle; occiput broadly convex below, slightly concave
to planate above, with distinctly incised vertical postgenal
fold; cheek relatively deep (its height 0.24-0.38 of height
of eye in measured specimens); the following bristles

Figure 14. Rivellia “sp. A” (Iluka, AM), wing.

134 Records of the Australian Museum (2001) Vol. 53

present: inner and outer vertical, strongly divergent
postverticals, ocellar, two distinct fronto-orbitals, lateral
occipital, postgenal. Antenna, excluding arista, much shorter
than face; segment 6 minutely pubescent on most of length.
Prelabrum of moderate size in female, markedly smaller in
male; palpus and proboscis moderately developed.

Thorax. Mesoscutum c. 0.9-1.1 times as long as wide;
scutellum rounded in outline, dorsally convex, setulose
dorsally, with little or no pale pruinescence; the following
thoracic bristles present: one or two scapulars, humeral, 1+1
notopleurals, supra-alar, postalar, posterior intra-alar,
dorsocentral (sometimes reduced), prescutellar acrostichal
(sometimes absent), two pairs of scutellars, mesopleural.
Posterior bridge of hind coxa without hairs; femora without
ventral spines; fore femur with dorsal and posteroventral
bristles; hind femur with few dorsal bristles and, in
Australian species only, one or more anteroventral bristles.
Wing membrane, including anal cell and alula, largely
microtrichose; section of costa on subcostal cell shorter than
that on second costal cell; subcosta and vein 5 without
setulae; anterior crossvein approximately transverse,
meeting vein 4 slightly beyond mid-length of discal cell;
second section of vein 4 with only slight curvature
(compared with Rivellia)’, distal section of vein 4 slightly
arched, not apically convergent with vein 3; anal crossvein
slightly to rather strongly curved; axillary lobe small, with
marginal hairs longer than on the narrower squama.

Abdomen ovoid; tergite 5 unreduced. Male postabdomen:
apical part of outer surstylus much surpassing apex of inner
surstylus; aedeagus either of simple platystomatine structure
with short bulb and longer terminal filaments (New Zealand
species), or with strap-like bulb many times as long as glans
and terminal sclerotised lobe larger than either terminal
filament (Australian species). Female abdomen: spiracle 5
located below lateral margin of tergite 5 (at least in Australian
species); aculeus moderately or quite slender, obtuse.

Distribution. New Zealand: North and South Islands.
Australia: temperate eastern areas and south-west.

Notes. Zealandortalis has been previously used to include
two New Zealand species, and has been regarded as endemic
to that country (Harrison, 1959). After careful consideration,
I have decided to include the Australian Z. gregi (described
below) in Zealandortalis , despite the number of differences
from the New Zealand species. There is a reasonable

probability, but no certainty, that the more broadly defined
genus is monophyletic, and it is readily identified from
key characters. The placement of Z. gregi in another
monotypic genus, when cladistic considerations do not
necessitate it, is undesirable. I do not consider that
divergence in male genitalia characters is a stronger
indication of phylogenetic remoteness than is divergence
in other kinds of character, especially as it is possible that
the full range of morphological variation in the New Zealand
species is not yet recorded.

The Australian species is treated as belonging in a
separate informal group from the New Zealand ones,
characterised in the key given below. My limited material
from New Zealand shows a combination of the characters
for the two described species as given by Harrison, so that
the number of New Zealand species and their distinguishing
characters may need clarification.

In temperate eastern Australia Z. gregi lives in a range
of forest types from warm coastal lowlands to winter-cold
highlands. Specimens of Zealandortalis sp. from South
Island, New Zealand (B.J.S.), are labelled “ex nikau palms.”
I collected a specimen doubtfully referable to Z. interrupta
from low vegetation in Nothofagus fusca forest on Banks
Peninsula, there being no palms in the vicinity.

The female from Banks Peninsula was observed in vitro
to walk with the wings folded in a horizontal plane largely,
but not completely, overlapping. No wing-waving was
observed over a period of about five minutes’ activity.
Though the above behaviour was observed comparatively
briefly, it contrasts with that of Z. gregi, of which several
specimens of both sexes from near Church Point, Sydney
district, were observed in a plastic container over several
days. When at rest, the wings were held in a horizontal plane,
slightly spread in a V so as to avoid any overlap. But often
there was an apparently alert attitude, with wings spread
and vibrated rapidly between the fully spread and more
flexed positions. When the fly was walking, the wings were
sometimes held in a V-flexed position, but, for most of the
time, there was no marked wing vibration. Two intermediate
actions, not infrequent, were (1) vibrating one wing only,
with the other almost still; (2) holding the wings in a very
broad V and vibrating them with very small amplitude. A
pair was observed apparently in copula several times. The
male was clinging to the abdomen of the female in a position
rather like that described for Pogonortalis doclea (see
Me Alpine, 1973b). The wings of the female were widely
spread, while those of the male were almost fully flexed.

Key to groups in Zealandortalis

1 Ocellar bristle large, potentially reaching more than halfway to
ptilinal suture; facial carina strongly raised, narrow, with steep
sides; antennal segment 3 acute dorsoapically; hind femur with
one or more long antero ventral bristles, stronger in male; vein 3
with setulae distributed over most of length; abdominal tergite 5
(both sexes) with long posterior marginal bristles; aedeagus with
bulb strap-like, many times as long as glans, terminating in

sclerotised lobe much larger than each terminal filament. Australian group

David K. McAlpine: Australasian Platystomatidae 135

Ocellar bristle smaller, not nearly reaching halfway to ptilinal
suture; facial carina undeveloped; antennal segment 3 broadly
rounded distally; hind femur without differentiated anteroventral
bristles; vein 3 with few setulae, mostly near its base; abdominal
tergite 5 without posterior marginal bristles; aedeagus (where

known) with bulb little developed, lacking sclerotised lobe. New Zealand group

Figures 15-17. Zealandortalis gregi n.sp. 15, wing. 16, distal part of aedeagus (scale = 0.2 mm). 17, head of male.
Abbreviations: bu, bulb; f, terminal filaments (paired); g, glans; lo, lateral occipital bristle; pf, postgenal fold; pg,
preglans; st, stipe; tl, sclerotised terminal lobe of aedeagus.

Zealandortalis gregi n.sp.

Figs. 15-17

Material examined. Holotype, S, New South Wales:
Iluka rainforest reserve, 26.xi. 1970, D.K.M. (AM).
Paratypes, New South Wales: 100 S S, 18 ??, Iluka,
Nov. 1970, Jan. 1971, A.H., D.K.M. (AM, ANIC, UQ, BM,
BPB, RMS, USNM).

Other material. Queensland: Blackdown Tableland, Expedition
Range (AM); Bribie Island (AM, UQ). New South Wales: Mount
Gibraltar National Park (AM); Tucker’s Rock, near Repton (AM);
Urunga (AM); Boonanghi State Forest, near Kempsey (AM);
Cobark Picnic Area, Wilson River, Bellangry district (AM); Taree
(AM); Timor Rock, Warrumbungle Range (AM); Lahey’s Creek
Road, 12 km NW of Gulgong (AM); Wongarbon Nature Reserve,

17 km SE of Dubbo (AM); “Tuglo”, near Mount Royal (AM);
Mungo Brush, near The Broadwater, Myall Lakes (AM); Mooney
Mooney Creek, near Gosford (AM); Newnes, Lithgow district
(AM); Mount Wilson (AM); Katoomba (AM); Wentworth Falls
(AM); Yellow Rock Lookout, near Springwood (AM); Newport
(AM); Church Point (AM); Vaucluse (AM); Royal National Park
(AM); Heathcote (AM); Otford (AM); Stanwell Tops (AM);
Bendalong, Conjola district (AM); Bodalla State Forest, near
Narooma (AM). Australian Capital Territory: Condor Creek,
Brindabella Range (AM). Western Australia: 15 mi. (c. 24 km)
SE of Manjimup (AM).

Description (c£, $). Habitus and appearance suggestive of
a stout, dark Rivellia species, with few wing markings.
Coloration. Head fulvous to reddish brown; orbital plates
shining blackish brown; entire occipital region blackish
brown, with grey pruinescence. Antenna fulvous, with

136 Records of the Australian Museum (2001) Vol. 53

brown arista. Prelabrum tawny; palpus fulvous. Thorax
black, largely shining, with dark grey pruinescence mainly
on notopleural region, upper part of mesopleuron, and
posterior parts of pleura, none on scutellum or a minute
amount near scutellar bridge. Legs fulvous; hind tibia and
sometimes fore femur suffused with brown; tarsi variably
browned apically (flies in southern populations often with
more extensively browned legs). Wing hyaline; first costal
cell, base of second costal and stigmatal section of subcostal
cell browned; also a small brown spot in marginal cell near
fork of veins 2 and 3, and a large apical dark brown spot on
vein 3, usually not or only indistinctly reaching to veins 2
and 4; axillary lobe and squama creamy, with fulvous
margins. Halter creamy, with tawny scabellum. Abdomen
black; preabdominal tergites without obvious pruinescence.

Head slightly higher than long; height of cheek 0.25-0.38
of height of eye; face with pair of moderately deep antennal
grooves, separated by a high, narrow, steep-sided carina;
parafacial narrowed near middle, haired at upper end, the
hairs extending as a spaced single series to about middle;
occiput with well marked postgenal fold almost from lower
extremity of postgena to middle of posterior margin of eye;
anterior fronto-orbital much larger than posterior one, both
reclinate; ocellar bristle much longer than anterior fronto-
orbital. Antennal segment 3 relatively narrow, acute
dorsoapically.

Thorax. Hairing on scutellum and posterior part of
mesoscutum markedly longer in male than in female. Hind
femur of male with a series of c. four to seven well-
developed black anteroventral bristles, somet im es reduced
to one bristle in diminutives, in female with c. one to three
finer, usually yellow, but sometimes quite long anteroventral
bristles. Wing: vein 3 dorsally with well-developed spaced
setulae on most of length; cell-4 index = 0.57-0.64.

Abdomen. Tergites 2 to 5 with longer hairs in male than in
female; tergite 5 with several long posterior marginal
bristles, in male this tergite particularly broad and giving
preabdomen a broadly truncated outline; spiracle 5 located
below lateral margin of tergite 5 in both sexes. Male
postabdomen: distal section of outer surstylus considerably
exceeding apex of inner surstylus, tapering to a slender apex;
aedeagus with short, simple preglans; glans short, broadly
ovoid; bulb consisting of elongate tubular section, c. five
times as long as glans, and containing three pigmented
longitudinal strips, and broad, darkly sclerotised terminal lobe
almost as long as glans; terminal filaments very small, subequal,
arising from a distinct sclerite at base of terminal lobe.

Dimensions. Total length, 6 S 2.9-3.9 mm, $ ? 2.4-3.9
mm; length of thorax, S 6 1.1-1.7 mm, ? ? 1.1-1.7 mm;
length of wing, S 6 2.4-3.6 mm, 9 9 2.7-4.1 mm; length
of glans of aedeagus 0.20-0.24 mm.

Distribution. Queensland: apparently widely distributed
south of the tropic (few records). New South Wales: widely
distributed in coast and tableland districts, inland to Dubbo
district and Warrumbungle Range (common in many

districts). Australian Capital Territory: Canberra district.
Western Australia: far south.

The specific epithet refers to Greg Daniels who has
collected much useful material.

Genus Tarfa n.gen.

Type species: Tarfa bowleyae n.sp.

Description (9, 6 unknown). Moderately small flies;
habitus somewhat like that of Rivellia, but significantly more
elongate, not subcylindrical as in Eumeka, Microepicausta,
etc.; legs conspicuously longer than in the above-named genera.

Head markedly higher than long; face without median
carina, concave in profile except towards lower margin;
lower end of facial ridge with uniseriate fine hairs; parafacial
very narrow, without hairs; occiput convex below, slightly
concave above; the following bristles present: inner and
outer vertical, small divergent postvertical pair, vestigial
ocellar, one large fronto-orbital, moderately small postgenal;
lateral occipital bristle absent. Antenna, excluding arista,
slightly shorter than face on median line; segment 2 short
and rather rotund; segment 3 narrowly ovate; segment 6
slender, with numerous short hairs on entire length.
Prelabrum deep; palpus broad.

Thorax largely shining, with very little pruinescence;
mesoscutum c. 1.2 times as long as wide, with many short
non-seriate hairs; scutellum short, sub triangular, only
slightly convex dorsally, with numerous dorsal hairs; zone
between metathoracic spiracle and halter base with only
micro-pruinescence, no pile-like pubescence or setulae; the
following thoracic bristles present: one or two quite small
lateral scapulars, 1+1 notopleurals, supra-alar, postalar,
posterior intra-alar, two unequal pairs of scutellars,
mesopleural; paramedian scapular, humeral, dorsocentral,
and prescutellar acrostichal absent. Femora slender, without
distinct bristles; mid tibia with one long apical ventral spur;
tarsi long and slender, each with terminal segment depressed
but not dilated. Wing much narrowed basally; membrane
largely microtrichose, except for anal cell and limited bare
zones in second basal cell and base of discal cell; both costal
cells densely microtrichose; subcosta on distal part straight,
well sclerotised, running close to costa; subcostal cell very
narrow; vein 2 with sigmoid curvature, rendering marginal
cell exceedingly narrow on distal half; vein 3 at its origin
approximated to vein 4, arched distally, setulose on most
of length dorsally; stem of veins 2 and 3 and vein 5 bare;
anterior cross vein meeting vein 4 well before mid-length
of discal cell; penultimate section of vein 4 arched, distal
section with slight sigmoid curvature, strongly converging
with vein 3 distally; anal cell extending much further basad
than second basal cell, about half as long as discal cell;
distal section of vein 6 much shorter than preceding section;
alula quite narrow, entirely microtrichose; axillary lobe
narrow; squama very narrow.

Abdomen moderately narrowed basally; preabdomen
otherwise subovoid; tergite 5 well developed; tergite 6 quite

David K. McAlpine: Australasian Platystomatidae 137

small (not visible from above in dried material); aculeus
slender, apically depressed and obtuse.

Distribution. New Caledonia: Loyalty Islands only.

Notes. The unusual combination of wing characters makes
Tarfa easy to distinguish from other platystomatid genera:
marginal cell much attenuated distally; first basal cell
narrowed at origin of vein 3; anterior crossvein meeting
vein 4 within basal third of length of discal cell; etc. (see
Fig. 18). Determining its relationship to other genera is more
difficult. The habitus and appearance suggest a large,
elongate Rivellia- like fly, but Tarfa differs from Rivellia in
the absence of humeral, dorsocentral, and fore femoral
bristles, the single, large fronto-orbital bristle, the much
longer legs, and quite different contour and relations of vein
4. Thus, there is hardly any morphological evidence of close

relationship between these genera. A few features suggest
relationship to the loosely defined alliance known as the
cleitamiine genera (McAlpine, 1982). Of these, the
somewhat slender species of Cleitamoides resemble Tarfa
in the basally narrowed wing, narrow alula, narrow
subcostal cell, devious contours of veins 2 and 3, apically
convergent veins 3 and 4, partly concave face, absence of
humeral and dorsocentral bristles, and slender femora with
reduced bristling. Cleitamoides lacks the fronto-orbital
bristle, but the probably related genus Cleitamia usually
has one large fronto-orbital as in Tarfa. It is doubtful if these
resemblances to cleitamiine genera indicate genuine
relationship. Cleitamia, Cleitamoides , and Laglaisia seem
likely to form a monophyletic group or the major
components of such a group, and a deduced groundplan
for such group may not include many Tarfa- like character
states. Tarfa shows certain points of resemblance to Aetha,
as mentioned under that genus, but differs in the slender
thorax, elongate legs, more reduced bristling, more
attenuated subcostal, marginal, and first basal cells, different
relative positions of bases of second basal and anal cells,
and more reduced alula and squama.

Etymology. The generic name is a contraction of the Greek
tarphys, dense, in reference to the proximity of certain veins,
and is feminine.

Tarfa bowleyae n.sp.

Fig. 18

Material examined. Holotype, $ (unique), Loyalty
Islands: Fayaoue, Ouvea, xii.1968, N.L.K (BPB).

Description ($). Coloration. Head brown, darker towards
vertex; face and cheek fulvous; upper part of face,
parafacial, orbital margin of postfrons, and post-orbital

Figure 18. Tarfa bowleyae n.sp., holotype.

138 Records of the Australian Museum (2001) Vol. 53

margin whitish pruinescent. Antenna fulvous; arista brown.
Prelabrum and palpus fulvous. Thorax shining orange-
fulvous. Legs predominantly fulvous; fore coxa paler; mid
and hind femora brown; mid and hind tarsi tawny. Wing
clear, except for narrow brown costal band on entire length,
which fills both costal cells, and brown zone in distal part
of first basal cell, which extends over anterior crossvein.
Halter fulvous, with largely brown capitellum. Abdominal
tergites 2 and 5 shining black; tergite 1 fulvous; ovipositor
sheath brown-black.

Head. Eye 0.66 as long as high; height of cheek c. 0.08 of
height of eye. Longer hairs on antennal segment 6 slightly
longer than basal diameter of segment.

Thorax. Mesoscutum and pleura almost devoid of
pruinescence; scutellum slightly rugose and pruinescent on
dorsal surface; metapleuron slightly encroaching on
membranous postcoxal area. Wing: cell-4 index = 0.30.

Abdomen. Tergite 5 almost as long as tergite 4, with few
differentiated bristles on posterior margin.

Dimensions. Total length 5.2 mm; length of thorax 1.8 mm;
length of wing 5.0 mm.

Distribution. As given for genus.

Notes. Tarfa bowleyae superficially resembles a braconid
or small ichneumonid, and may be a hymenopterous mimic.

The specific epithet refers to Janis Bowley, whose
illustrations in this and other papers form a significant
contribution to Dipterology.

Genus Brea Walker

Fig. 19

Brea Walker, 1859 (February): 117. Type species (designated
Hendel, 1914a) B. contraria Walker.

Maria Bigot, 1859 (July): 311. Type species (monotypy) M.
caeruleiventris Bigot (= Brea contraria Walker).

Description. A detailed description has been given by
Hendel (1914a), to which the following notes are added.

Male postabdomen (examined in three species). Aedeagus
with preglans undifferentiated from stipe; glans elongate-
ovoid to elongate-subcylindrical; bulb very long and slender,
terminating in either two or three terminal filaments.

Female postabdomen. Ovipositor sheath moderately short
and broad; aculeus slender, obtuse.

Distribution. Moluccas: Kei Island. New Guinea: mainland;
Aru; Normanby Island. Australia: Queensland, N of 14°S.

Notes. Hendel (1914b) gave a key to species, which was
repeated in English by Malloch (1939a), but the status and
limits of the species are not yet well understood. I included
the Queensland population in Brea contraria Walker (as
indicated by Evenhuis, 1989), but further, incomplete
studies render this identification doubtful. Nine nominal
species, as listed by Evenhuis, may perhaps claim validity,
and there may be one or two undescribed species.

In my experience, Brea species have been found on
foliage in or at the margin of rainforest in Papua New Guinea
and Queensland.

Brea nouhuysi de Meijere has a remarkably broadened
head in the male, but such sexual dimorphism is absent or
barely noticeable in other species examined.

Genus Pseudorichardia Hendel

Fig. 20

Pseudorichardia Hendel, 1911: 21. Type species (original
designation) Richardia flavitarsus Macquart.

Description. See Hendel (1914a: 123-124). In the available
males from Tuvalu (AM), the long, paired terminal filaments
are fused for the greater part of their length.

Distribution. Outer Melanesia: Vanuatu (New Hebrides);
Fiji. Tropical Polynesia: Samoan Islands; Cook Islands;
Society Islands; Marquesas; Pitcairn Island. Micronesia:
Tuvalu.

Figure 19. Brea sp. (Claudie River, AM), wing.

David K. McAlpine: Australasian Platystomatidae 139

Notes. Steyskal (1952b) recognized four species and
provided a key. It is evident from the small amount of
material available to me that this is an over-simplification.
Characters of the aedeagus, and, in the females, abdominal
coloration and relative size of tergites, all omitted by
Steyskal, should be utilised. The number of known species
is likely to be increased by further collecting and study.

Almost no information on biology and behaviour is
available, but Bezzi (1928) recorded material “bred from
oranges” from Raratonga, Cook Islands.

Genus Lamprogaster Macquart

Figs. 21-26

Lamprogaster Macquart, 1843: 211. Type species (monotypy) L.
flavipennis Macquart.

Cruphiocera Macquart, 1843: 212. Type species (monotypy) C.
violacea Macquart.

Chromatomyia Walker, 1849: 801. Type species (designated
McAlpine, 1973a) C. formosa Walker (= Lamprogaster laeta
(Macquart)). Homonym (? senior) of Chromatomyia Hardy,
1849 (family Agromyzidae).

Ceratopelta Bigot, 1878b: 34. Type species (monotypy) C. tricolor
Bigot (= Lamprogaster patula Walker).

Cryphiocera.- Hendel, 1914a: 104. Variant spelling for
Cruphiocera, as syn. of Lamprogaster.

Liolamprogaster Enderlein, 1924: 128. Type species (original
designation) Liolamprogaster angusta Enderlein (? =
Lamprogaster semicyanea (Walker), see McAlpine, 1995a).

Description. Generic descriptions have been given by
Hendel (1914a) and McAlpine (1973a), to which I add the
following.

Male postabdomen. Aedeagus diverse; sometimes a long,
partly sclerotised process arising from between preglans
and glans; glans rounded-ovoid to elongate-cylindrical; bulb
seldom complex; terminal filaments usually two, but three
present in one group of species, usually of similar size,
occasionally very unequal.

Female abdomen. Tergites 4 and 5 not reduced in size;
spiracles 4 and 5 (where known) located in pleural
membrane below lateral margins of tergites; ovipositor
sheath usually broad, of moderate length; aculeus slender,
obtuse.

Distribution. Australasian Region: Moluccas; New Guinea;
Bismarck Archipelago; Solomon Archipelago; New
Caledonia; Australia—all states. Some species, e.g.,
Lamprogaster excelsa McAlpine and L. maculipennis
Macquart, live in winter-cold highlands of south-eastern
Australia. Oriental Region: Sulawesi; Philippines.
Lamprogaster elegans Frey, 1964, apparently the only
species recorded from continental Asia (Burma), is here
transferred to subfamily Trapherinae as Eopiara elegans
n.comb., from examination of the holotype (HELS).

Notes. McAlpine (1973a) reviewed the Australian species;
Malloch (1939a) keyed those of New Guinea and adjacent
islands that were known to him. In the Australasian Region
there are c. 38 valid described species, and a further 54
apparently undescribed species have been seen in collections.
Centres of diversity are New Guinea (especially), eastern
Australia, and New Caledonia.

Though Lamprogaster is generally a well defined taxon,
it may not be monophyletic (see McAlpine, 1994). A few
species have caused difficulty because of marginal
agreement in morphology. A remarkable undescribed
species or species complex (provisional coding “sp. BM”
from several localities in Papua New Guinea mainland and
on Normanby Island, AM and BPB) is a mimic of vespid
wasps. It has a petiolate abdomen, and can crease the wings
longitudinally as do those insects. I have decided to include
it in Lamprogaster because (1) it possesses the elongate
process on the aedeagus between preglans and glans,
characteristic of a number of Lamprogaster species, (2) there
is another little known New Guinea species morphologically
intermediate between this and more typical Lamprogaster
species, and (3) there is no reason for assuming that,
phylogenetically, it is not just a highly derived Lamprogaster

Figure 20. Pseudorichardia interrupta Bezzi, wing. After A.J.E. Terzi, in Bezzi (1928).

140 Records of the Australian Museum (2001) Vol. 53

Figures 21-23. 21 , Lamprogaster viola Malloch, wing. 22, Lamprogaster “sp. V” (Papua New Guinea, AM), wing.
23, Lamprogaster “sp. AN” (New Caledonia, AM), head.

species. These aberrant species are among the vey few
Lamprogaster species without a mesopleural bristle, and
should run to couplet 19 in my key to genera.

Though generally found in forested country, Lamprogaster
species live in a wide range of habitats. Adults of some
species rest on tree trunks, or on the lower surface of tree
foliage. They are often collected in traps baited with
mammalian dung.

I strongly suspect that Lamprogaster macrocephala
Hendel, L. nigrihirta McAlpine, L. patula Walker, L.

pumicata Wulp, and other species are Batesian mimics of
blow flies (Calliphoridae), with which the adults may be
associated at mammalian dung.

Previously (McAlpine, 1973a: 11), I recorded a specimen
of Lamprogaster corusca attacked by the fungus Empusa.
The name of the host is now corrected to Lamprogaster “sp. 1 ”
(AM), and the fungal genus is probably Entomophthora
(Zygomycotina: Entomophthorales).

Figures 24-26. Distal part of aedeagus of Lamprogaster spp. 24, L. relucens McAlpine (scale = 1 mm). 25, L.
rugifacies McAlpine (scale = 0.5 mm). 26, L. imperialis McAlpine (scale = 0.5 mm). Abbreviations: f, terminal
filaments; g, glans; pp, process of preglans.

David K. McAlpine: Australasian Platystomatidae 141

Genus Montrouziera Bigot

Figs. 27, 28

Montrouziera Bigot, 1860b: 224. Type species (monotypy) M. lifua
Bigot.

Description. The most detailed description is that of Hendel
(1914a), though based on meagre material. Further
morphological study of the only included species is needed.

Distribution. New Caledonia: perhaps Loyalty Islands only.
Evenhuis (1989) also mentions the main island of New
Caledonia, but I think this may be due to misinterpretation
of the term “New Caledonia”, which may have been used
in the broad sense to include the Loyalty Islands, which
come under the same French administrative area. Fairly
extensive modern collections from New Caledonia (main
island) do not include Montrouziera.

Notes. Only the type species of Montrouziera is known.
The wing pattern and venation (Fig. 28, based on Hendel’s
figure of type material) should enable ready identification
of M. lifua.

Montrouziera is probably closely related to Lamprogaster,
but differs from probably all species of that genus in having
a quite large fronto-orbital bristle. Though Lamprogaster
is represented in New Caledonia by numerous species
(mostly undescribed), apparently all these differ from
Montrouziera in the less hirsute lateral occiput, less
developed hairing of the arista, and different placement of
the anterior crossvein. The characters of Lamprogaster
species of New Guinea are, however, more variable.

Genus Achias Fabricius

Figs. 7, 8, 29-33

Achias Fabricius, 1805: 247. Type species (monotypy) A. oculatus
Fabricius.

Mystia Walker, 1861b: 249-250. Type species (monotypy) M.
attrahens Walker.

Achiosoma Hendel, 1914a: 100-101. Type species (original
designation) Achias dacoides Walker.

Description. See McAlpine (1994).

Distribution. New Guinea. Bismarck Archipelago.
Northern Australia. Mainland New Guinea is the centre of
diversity of the genus, but the limits of distribution in the
west are Waigeo, Aru, and Melville Islands, in the north the
equator, in the east New Britain and the Louisiade Group,
and in the south the Townsville district of Queensland (see
map, McAlpine, 1994: fig. 1).

Notes. The genus has been reviewed by McAlpine (1994).
There are 96 probably valid described species, and I have
seen about six additional species in collections.

The males of many species have the head capsule
prolonged laterally to form eye-stalks. Achias rothschildi
Austen probably has the widest head capsule of any insect
(up to 55 mm). The role of eye-stalks in agonistic behaviour
has been discussed by McAlpine (1979).

Several species are apparent Batesian mimics of vespid
wasps. This has been confirmed by my field observations
on Achias nigrifacies Malloch and A. minax McAlpine.

A female paratype of Achias xyrion McAlpine has a male
strepsipteran pupa protruding from the abdomen (AM).

Figures 27-28. Montrouziera lifua Bigot. 27, head. 28, wing. After Hendel (1914a).

142 Records of the Australian Museum (2001) Vol. 53

Figures 29-31. Achias kurandanus Hennig. 29, head of female. 30, head of male. 31, wing.
Illustrations by S.R Kim.

Genus Inium McAlpine

Figs. 34, 35

Inium McAlpine, 1995b: 252. Type species (original designation)
I. marine McAlpine.

Description. See McAlpine (1995b).

Distribution. North-eastern Queensland, from the Atherton
Tableland to the vicinity of Cape York.

Notes. Inium includes three rarely collected species, one
still undescribed (ANIC), of diverse appearance, though
probably all Batesian mimics of ants. The most noteworthy
common feature is the structure of the prelabrum, which
departs from the usual horse-shoe shape and is much
extended posteroventrally towards the proboscis and largely
medially sclerotised. It thus somewhat resembles the beak
of certain parrots. The two described species are
morphologically dissimilar in many respects (McAlpine,
1995b: table 1), but the third species is intermediate in
morphology, size, and geographic distribution.

Genus Rhytidortalis Hendel

Figs. 36, 37

Rhytidortalis Hendel, 1914a: 14, 66-68. Type species (original
designation) R. cribrata Hendel.

Description. See McAlpine (2000).

Distribution. Australasian Region: Australia—all states,
including Northern Territory and Tasmania. Oriental
Region: Taiwan; Vietnam. Palaearctic Region: Russian Far-
east. Most species have strictly coastal distributions.

Notes. I have recently reviewed this genus (McAlpine,
2000), but omitted R. nigripes Korneyev from the Russian
Far-east (Korneyev, 1991). The type species is the only
known Oriental species, but seven species live in Australia.
The Australian species originally described as Senopterina
rugifrons Thomson and Pseudepicausta solocifemur
Enderlein have been referred to Rhytidortalis, but are now
placed in Duomyia.

Adults are usually found on low vegetation on sand near
the sea shore.

David K. McAlpine: Australasian Platystomatidae 143

Figures 32-35. 32, Achias minax McAlpine, head of male. 33, Achias trivittatus McAlpine, wing.
34, lniurn simplum McAlpine, wing. 35, Inium mariae McAlpine, holotype.

O.S. KENT

144 Records of the Australian Museum (2001) Vol. 53

Figures 36-37. Rhytidortalis averni McAlpine. 36, head of male. 37, wing, showing approximate distribution of
microtrichia.

Genus Scotinosoma Loew
Fig. 38

Scotinosoma Loew, 1873: 45. No included species.

Scotinosoma Loew.-Hendel, 1914a: 16, 65-66. Type species
(subsequent monotypy) S. bistrigatum Hendel.

Description. Hendel gave a detailed description based on
the characters of the type species only. I add the following
descriptive notes from an examination of the available
species.

Head. Facial carina typically strongly elevated, rather
narrow, and rounded, sometimes reduced and more or less
flattened, always without hairs; facial ridge with few fine
hairs below; parafacial usually pruinescent, without hairs
except near upper extremity; lateral occipital and postgenal
bristles present, sometimes duplicated. Prelabrum generally
well developed, not joined to face by median shining
quadrate sclerite, but sometimes its sclerotisation extended
medially towards face, its lower anterior margin not
markedly receding.

Thorax. Me so scutum with extensive grey pruinescence,
often in part forming a pale median stripe, less frequently
generally distributed, not perforated by pattern of black
spots as in Rhytidortalis; posterior part of mesopleuron with
grey pruinescent zone, which extends forward on vicinity
of sternopleural suture to fore-coxal cavity; usually only
one true supra-alar bristle present (in contrast to most
Rhytidortalis species), but bristling of this region sometimes
modified in females; scutellum with two or three pairs of
bristles and no hairs. Fore coxa broad; femora without
spines; fore femur often with variably developed dorsal and
posteroventral bristles; hind basitarsus slender, sub-
cylindrical, as compared with Rhytidortalis. Subcosta and
ventral surface of vein 1 without setulae; second basal cell
less than half as long as discal cell; wing markings usually
simple, including a dark costal mark, and, in two species,
one or two obliquely transverse bands and a sub-basal mark.

Abdomen subcylindrical in males, usually more ovoid in
females. Male postabdomen (where known): preglans
forming separate sclerite from stipe; glans ovoid to
cylindrical; terminal filaments subequal in length, not very
long. Female abdomen: tergite 5 unreduced, with spiracles

Figure 38. Scotinosoma sp. (near S. erasum Malloch, Mount Mowbullan, AM), wing.

David K. McAlpine: Australasian Platystomatidae 145

close to its posterior margin; ovipositor sheath not much
longer than tergite 5 (in contrast to typical Pseudepicausta
species); aculeus slender, obtuse.

Distribution. Australia: eastern Queensland; north-eastern
New South Wales—Richmond River district northwards. I
am at present treating the one recorded Philippine species
as probably generically misplaced, and the genus as perhaps
endemic to Australia.

Notes. The first association of a specific name with
Scotinosoma is that of Dacus basalis Walker, a junior
synonym of Plagiostenopterina aenea (Wiedemann), by
Hendel (1912b: 14), but, as this combination was inferred
with doubt, it is not a valid type designation.

Four Australian species have been referred to Scotinosoma
and a key to these was given by Malloch (1939a). From
preliminary study of available material, I believe that there
are likely to be at least 12 Australian species, and that use
of secondary sexual characters not mentioned by Malloch
will be necessary to characterise some of these.

The type locality for the type species, S. bistrigatum
Hendel, 1914a, has been given as Cape York (Hendel,
1914b, Malloch, 1939a), but this is an error. The two original
specimens collected by Thorey in Queensland and described
by Hendel (1914b) in WM bear labels of similar format,
but that on the male says “Thorey 1868 Port Denison”, that
on the female “Thorey 1868 Cap York”. The supposition
that both are from Cape York arose from Hendel’s
carelessness in checking the two similar-looking labels. The
illustrations on which the original availability of S.
bistrigatum depends (Hendel, 1914a: pi. 5, figs. 103, 104),
were based on the male from Port Denison (now Bowen),
which is therefore the holotype. The female from Cape York,
which was also used in the subsequent description (Hendel,
1914b), is not a type specimen. There are slight obvious
differences between the male and female, and, as far as can
be determined without a detailed taxonomic study based
on adequate material, they may well belong to different
species. The collection sites are nearly 1200 km apart in a
virtual straight line.

Scotinosoma species live mainly in rainforests, including
dry rainforest and vine scrub. They are apparently absent
from most of the littoral rainforests of north-eastern New
South Wales and south-eastern Queensland. Adults have
been collected in dung-baited traps.

Genus Eumeka n.gen.

Type species: Eumeka hendeli n.sp.

Description (c3, $). Medium-sized to moderately large
elongate, dark flies, with legs of moderate length and wing
lightly marked.

Head markedly higher than long; face with narrow flat-
topped carina for most of its length (less elevated in New
Caledonian species), without hairs; facial ridge with several
or many fine hairs below; parafacial densely pruinescent,
with scattered hairs on entire length; occiput moderately

convex below, slightly concave above; the following bristles
present: inner and outer vertical, small postvertical, minute
ocellar, two medium-sized to small fronto-orbitals, small
lateral occipital often distinguishable; postgenal bristle
present or absent. Antenna, excluding arista, slightly shorter
to slightly longer than face on median line; segment 3 c.
four times as long as wide, apically rounded; segment 6
filiform, only slightly thickened basally, minutely pubescent
on basal part only. Prelabrum rather small, coarsely rugose,
its lower margin usually receding; palpus moderately small,
coarsely setulose.

Thorax. Mesoscutum 1.2-1.3 times as long as wide, with
pruinescent median stripe; scutellum moderately short and
convex, without hairs; pleura with pruinescent zones; the
following thoracic bristles present: a group of small lateral
scapulars, humeral, 1+1 notopleurals, supra-alar, postalar,
posterior intra-alar, small dorsocentral, prescutellar
acrostichal, three pairs of scutellars, mesopleural;
paramedian scapular absent. Posterior bridge of hind coxa
without hairs; femora without ventral spines; fore femur
with well-developed dorsal and posteroventral bristles; mid
femur without bristles; hind femur with anterodorsal bristles
near base (sometimes weak and pale) and dorsal bristles
distally; tibiae unarmed, except for rather short apical ventral
spur on mid one; tarsi elongate. Wing elongate, not
significantly narrowed basally; membrane with extensive
bare areas in first basal, second basal, and anal cells;
subcosta separate from vein 1 for whole length beyond
humeral crossvein; section of costa on subcostal cell slightly
shorter than that on second costal cell, much longer than
that on marginal cell; stem of veins 2 and 3 simple, nearly
straight, without setulae on either surface; veins 3 and 4
subparallel or slightly divergent apically; anterior crossvein
meeting vein 4 well beyond middle of discal cell; second
section of vein 4 gently arched, penultimate section almost
straight; second basal cell slightly over half as long as discal
cell; basal crossvein transverse; anal crossvein curved; distal
section of vein 5 slightly shorter than section on anal cell;
alula well developed, rounded; squama forming distinct,
narrowly rounded lobe, scarcely larger in area than the well-
developed axillary lobe.

Abdomen in male subcylindrical, in female more ovoid,
variably narrowed anteriorly, with tergite 5 unreduced;
ovipositor sheath shorter than tergite 3; aculeus very slender,
obtuse. Aedeagus, where known, with pair of moderately
short, equal terminal filaments.

Distribution. Australia: Queensland—north-east coast as
far south as Paluma. New Guinea: eastern and south-eastern
mainland. Bismarck Archipelago: Lavongai (New
Hanover). New Caledonia: main island.

Notes. Eumeka includes two very similar species or distinct
populations in north Queensland, one of which also occurs
in New Guinea. There is a probably distinct species on
Lavongai in the Bismarck Archipelago (ZMC) and another
more divergent species in New Caledonia (BPB), but I have
only seen single specimens of the two latter.

146 Records of the Australian Museum (2001) Vol. 53

Eumeka appears to be most closely related to Pseudepi¬
causta, and perhaps to Rhytidortalis and Conicipithea. All
these genera consist of more or less elongate, Senopterina-
like flies (Senopterinini sensu Hendel, 1914a, as Steno-
perinina), with veins 3 and 4 subparallel apically, the face
hairless, the parafacial haired near centre, and often a distinct
lateral occipital bristle. Eumeka species of Australia, New
Guinea, and the Bismarcks are unique among these genera
in having a series of setulae on the ventral surface of vein
1, which is usually most developed near the distal bend of
the subcosta. Because vein 1 tends to lie in a deep groove,
as seen from the ventral surface, the setulae are sometimes
difficult to see in dried material, but I have detected them
in all 12 available specimens of these species. Eumeka
further differs from Pseudepicausta in having the cheek
deeper, the second section of vein 4 longer and definitely
arched, and the thick, non-directional, almost paint-like
white pruinescence on the sides of tergite 4; also Eumeka
differs from the typical species of Pseudepicausta , from
New Guinea and nearby island groups, in having sternite 5
of the male compact and not deeply divided, and the
ovipositor sheath of the female not much longer than tergite
5. From Rhytidortalis, Eumeka differs in its consistently
larger size, vertically elongate eye, lack of the characteristic
mesoscutal pattern of that genus, more elongate hind
basitarsus, arched second section of vein 4, narrower anal
cell, and characteristic microtrichiation pattern of tergite 4.
Eumeka resembles Conicipithea in its more or less wasp¬
like appearance (both could be mimics of pompilids), the
head prominently produced at the bases of the antennae,
and the facial carina usually long and flat-topped; but it
differs in the absence or slight development of setulae on
antennal segment 3, the absence of setulae on the scutellum,
the absence of erect hairs on the suprasquamal ridge, and
the quite different venation; also the glans of the aedeagus
is much shorter than in the one available male of
Conicipithea.

The New Caledonian species (1 9 , Mount Koghi, BPB)
seems to form a distinctive group, distinguishable as
indicated in Table 1.

I have collected two males of Eumeka hendeli feeding at
sap exuding from recently cut stems of Calamus sp.
(Arecaceae) in lowland rainforest near Babinda, Queensland.

The generic name is from the Greek eumekes, of good
length, and is treated as feminine.

Eumeka hendeli n.sp.

Figs. 39-41

Material examined. Holotype, 6, Queensland: The
Boulders reserve, near Babinda, 27.L1991, B.J.D., D.K.M.
(AM). Paratypes, Queensland: 2 6 6, Whitfield Range,
near Cairns, Feb. 1978, A.W-H. (AM); 1 6 , The Boulders,
Jan. 1991, B.J.D., D.K.M. (ANIC); 1 6, Paluma, 2925 ft
(c. 890 m), Jan. 1970, G.A.H. (AM).

Description (c£, ? of type race unknown). Coloration.
Head with ground colour largely brown; fronto-orbital
margin (except posteriorly), parafacial, much of antennal
groove, posterior orbits (narrowly), and postgenal region
(diffusely) silver-grey pruinescent; facial carina with
translucent pruinescence, not concealing ground colour,
which is distinctly yellowish on upper part. Antenna tawny;
segment 3 variably suffused with brown distally; arista
brown basally, black elsewhere. Prelabrum brown or tawny-
brown; palpus tawny-brown, somewhat paler distally.
Thorax with dark brown or sometimes partly tawny-brown
ground colour; mesoscutum with rather narrow pale grey-
pruinescent median stripe, complete, except that it merges
posteriorly with pale transverse stripe along scutellar suture,
also with grey lateral presutural pruinescent zone, which is
denser and paler between notopleural bristles; scutellum
with thin greyish pruinescence dorsally; propleuron largely
greyish white-pruinescent; mesopleuron shining anteriorly,
with broad greyish-white pruinescent band on posterior
margin; sternopleuron with central greyish-white pruin¬
escent zone reaching sternopleural suture and expanded
ventrally; a pale-pruinescent mark covering pleurotergite
and part of meropleuron. Legs brown to tawny-brown; fore
coxa extensively whitish-pruinescent, except on much of
posterior surface. Wing: subcostal cell brown; brown costal
band extending from end of vein 1 to just behind vein 4,
narrowest near end of vein 2, broader and diffuse in first
posterior cell; marginal cell with small brown mark at origin
of vein 2; anterior and discal crossveins narrowly margined
with brown; a small brown mark on vein 5 between basal
and anal crossveins; axillary lobe and squama white. Halter
brown. Abdomen dark brown to black; tergite 2 with
transverse, medially narrowed, densely greyish white
pruinescent band across whole width on or near posterior

Table 1. Comparison of groups in Eumeka.

Hendeli group

New Caledonian group

• Postgenal bristle absent or poorly

differentiated

• Prelabrum small, receding

• Vein 1 setulose ventrally

• Both costal cells and alula with extensive

bare areas

• Anterior and discal crossveins with incon¬

spicuous separate brown marks, or none

• Abdominal tergite s 3 and 5 without

whitish-pruinescent markings

• Postgenal bristle well developed

• Prelabrum of moderate size, rather prominent

• Vein 1 bare below

• Both costal cells and alula almost

entirely microtrichose

• Anterior and discal crossveins enclosed

in the one oblique brown mark

• Abdominal tergites 3 and 5 with whitish-

pruinescent markings

David K. McAlpine: Australasian Platystomatidae 147

Figures 39-41. Eumeka hendeli n.sp. 39, head of male. 40, wing. 41, distal part of aedeagus (scale = 0.2 mm).

margin; tergite 4 with large triangular, densely greyish
white-pruinescent zone on each lateral margin; tergites 3
and 5 without pale pruinescence.

Head. Facial carina strongly raised, especially on upper
part, in centre markedly narrower than antennal groove,
slightly depressed and broadened below, not reaching
lower margin of face; height of cheek 0.33-0.40 of height
of eye. Antenna potentially extending for 0.7-0.8 of
length of face on median line.

Thorax. Cell-4 index = 0.70-0.75; alula microtrichose only
on narrow posterior marginal zone, or entirely bare (except
for marginal series of setulae).

Abdomen. Tergite 4 slightly longer than tergite 3; tergite 5
slightly shorter than tergite 3. Postabdomen: outer surstylus
obtuse, its apex only slightly exceeding that of inner
surstylus; stipe of aedeagus slender; preglans sharply
differentiated from stipe, sclerotically continuous with
glans; a small patch of pubescence on base of preglans,
extending on to stipe; glans elongate-ovoid; terminal
filaments rather short, of equal length, arising from a
common sclerotised base, the whole structure c. as long as
glans; cerci large, much exceeding surstyli, fused to
proctiger for their whole length.

Dimensions. Total length 8.9-10.9 mm; length of thorax
3.2-4.0 mm; length of wing 7.5-9.1 mm; length of glans of
aedeagus 0.24 mm.

Distribution. Queensland: Caims district to Paluma district
(typical populations). Populations in the Iron Range district,
Queensland (AM) and the Central and Oro Provinces of Papua

New Guinea (AM, BPB) appear to be morphologically distinct,
probably at infraspecific rank.

Notes. Eumeka hendeli is here interpreted broadly to include
populations in northern Cape York Peninsula and eastern
mainland New Guinea, as well as the type population. On
this interpretation, the species differs from the only other
recognized species of the hendeli group (from Lavongai)
in the complete but usually dorsally narrowed whitish-
pruinescent posterior marginal band on abdominal tergite
2, and in having the facial carina not broadened near mid¬
length nor very sharply margined. The type population of
E. hendeli differs from more northern populations in the
markedly shorter antenna (at least in males), slightly shorter
facial carina, and from at least some such populations in the
less distinctly petiolate abdomen. From the little available
material, I am not sure if all these populations are conspecific.

The specific epithet refers to Friedrich Hendel, master
worker on platystomatid flies.

Genus Signa n.gen.

Type species: Signa mouldsi n.sp.

Description ($, $). Flies of medium build or slightly
elongate, of small to rather large size, with moderately short,
rather slender legs.

Head markedly higher than long; face not very broad,
minutely roughened or pruinescent, without hairs, with
rather low, rounded, saddle-like carina; fine uniseriate hairs
along lower parafacial suture scarcely extending on to facial
ridge; parafacial with hairs near upper extremity only;
occiput variably convex below, somewhat concave above;

148 Records of the Australian Museum (2001) Vol. 53

the following bristles present: inner and outer vertical, small
ocellar, generally two reclinate fronto-orbitals, of which
anterior one is always well developed and at least slightly
larger than posterior one, lateral occipital (sometimes
reduced), large postgenal; postvertical bristle absent;
antennal sockets narrowly separated. Antenna, excluding
arista, slightly shorter than face on median line, slightly
shorter in female than in male; segment 3 generally subacute
dorsoapically; segment 6 long, filiform, but sometimes
preapically dilated in males, with variable amount of basal
pubescence. Prelabrum of moderate size, its lower margin
slightly anteriorly prominent; palpus moderately large,
elongate; proboscis stout.

Thorax. Mesoscutum c. 1.0-1.2 times as long as wide,
largely with blackish ground-colour and extensive covering
of grey pruinescence, a pair of variably developed black,
non-pruinescent stripes on or near dorsocentral lines,
sometimes broad and running almost full length of
mesoscutum, sometimes narrow, disjunct at suture, or
present only in front of suture; scutellum thick, rounded,
dorsally pruinescent, with few hairs restricted to lateral
margins, or sometimes quite hairless; pleura, except
sternopleuron, extensively greyish-pruinescent; sterno-
pleuron pruinescent along upper margin and along ventral
median margin, with extensive smooth zone between, which
extends broadly approximately to mid-coxal cavity; centre
of sternopleuron with a haired zone, which is almost smooth
between hairs, and isolated, except on dorsal margin, from
other haired or pruinescent zones; the following thoracic
bristles usually present: one short lateral scapular, humeral,
1+1 notopleurals (some of these reduced or modified in
females of some species), supra-alar, postalar, posterior
intra-alar, dorsocentral, two pairs of scutellars, mesopleural;
paramedian scapular and prescutellar acrostichal absent.
Posterior bridge of hind coxa with fine hairs; fore femur
with well-developed dorsal bristles and a series of long
posteroventral bristles on almost its entire length; mid femur
with several posterior bristles distally, merging into long
hairs, or posterior bristles scarcely differentiated from hairs;
hind femur distally with few dorsal bristles. Wing:
membrane microtrichose except for anal cell and much of
second basal cell; section of costa on subcostal cell
substantially longer than that on second costal cell and that
on marginal cell; subcosta and vein 1 without ventral setulae;
veins 3 and 4 converging apically; vein 4 terminating
distinctly behind wing apex, its basal section often with
weak point or flexure near distal end, its distal section
slightly or strongly curving forward only on apical half or
less; anterior crossvein meeting vein 4 near or slightly
beyond mid-length of discal cell; vein 5 without setulae, its
short distal section slightly bent towards posterior margin;
second basal cell on vein 4 c. one third as long as discal cell
or slightly less; anal crossvein curved on anterior part only;
axillary lobe of moderate to small size, with moderate marginal
fringe; squama forming a rounded lobe, slightly variable in
size, but not noticeably larger in area than axillary lobe.

Abdomen ovoid, not much narrowed basally. Male: tergite
5 usually rather large; outer surstylus usually extensively
spinulose on distal part (not examined in some species);
aedeagus with subcylindrical glans, variably developed
bulb, and pair of about equal-sized terminal filaments.
Female: tergite 5 well developed but always shorter than
tergite 4; spiracles 1 to 3 situated in pleural membrane;
spiracle 4 situated in intersegmental membrane near lateral
margins of tergites 4 and 5; spiracle 5 situated behind tergite
5, near its posterior margin; ovipositor sheath broad and
rather short; aculeus slender, obtuse.

Distribution. New Caledonia: only known from main island.

Notes. Signa is represented by at least 10 species in available
collections (mainly BPB, PM, and AM).

Signa belongs among the group of more or less elongate
platystomatine genera with hairless face, apically
convergent veins 3 and 4, distinct but rather small squamal
lobe, non-setulose suprasquamal ridge, and no femora
noticeably enlarged or ventrally spinose, this group
including Elassogaster, Microepicausta, Meringomeria, and
Par. Signa differs from Elassogaster in the presence of
fronto-orbital bristles, the termination of vein 4 behind the
wing apex, and the absence of a shallow groove connecting
the transverse sutures of the mesoscutum. It differs from
Microepicausta in the distinct fronto-orbital bristles,
presence of small hairs on the posterior bridge of hind coxa
and dorsal bristles on the fore femur, absence of hairs on
the central part of the scutellum, and in having paired
terminal filaments on the aedeagus. It differs from
Meringomeria in the presence of fronto-orbital bristles, non-
pruinescent central zone of the sternopleuron, slender
aculeus, and, particularly from the Melanesian species of
that genus, in the non-plumose arista and absence of a
distinct series of anterior bristles on the hind femur. Perhaps
the most closely related genus is Par , which, though also
Melanesian, is apparently geographically isolated from
Signa. Signa differs from Par in having larger fronto-orbital
bristles, centrally grey-pruinescent mesoscutum, scutellum
without dorsal setulae, sternopleuron centrally smooth, fore
femur with dorsal bristles, subcosta without ventral setulae,
anterior crossvein infuscated, abdominal pleura of female
without dense, pile-like covering (? microtrichia or
macrotrichia), outer surstylus extensively spinulose on distal
part (not checked for some Signa spp.).

Signa species have been collected in both lowland and
mid-mountain rainforest, while the long series of S. mouldsi
was obtained in dry vine forest or sclerophyllous forest
(M.S.M., pers. comm.). According to Jaffre et al. (1998),
this is the most fragmented and vulnerable natural vegetation
type in New Caledonia, and S. mouldsi is not represented
among the many available specimens of Signa from other
localities. Probably the numerous species occupy a wide
range of natural habitats in New Caledonia.

The generic name is derived from the Latin signum, a sign
or signal, and is feminine on account of the altered suffix.

David K. McAlpine: Australasian Platystomatidae 149

Signa mouldsi n.sp.

Figs. 42-45

Material examined. Holotype, 6 , New Caledonia: Pindai,
Nepoui Peninsula, ll.vi.1996, M.S.M. (PM). Paratypes,
New Caledonia: 28 S S, 38 9 9, same data as holotype
(AM, BM, BPB, PM, RMS, USNM). Additional material
in alcohol (AM).

Description (c?, 9). Dark fly, small for the genus, with
blackish apical wing spot. Coloration. Head tawny; antennal
groove brown, with whitish pruinescence; upper part of face
and parafacial yellow, with pale pruinescence; occipital
region extensively blackish, with grey pruinescence th inn er
on upper part, tawny-yellow zone extending over centre of
vertex on to central upper occiput, so that inner vertical
bristle stands on tawny zone, while outer vertical stands on
black zone. Antenna tawny; segment 3 largely greyish

brown; segment 6 blackish brown. Prelabrum fulvous
yellow, brownish only at posterolateral extremities; palpus
fulvous-yellow, greyish brown on a relatively small area
near base. Thorax black, with large part of cuticular surface
covered with thin grey pruinescence; mesoscutum with pair
of narrow anterior dorsocentral to intradorsocentral black
stripes, and postsuturally a pair of narrow extradorsocentral
black stripes; scutellum thinly grey-pruinescent dorsally,
glabrous on free margins. Legs black to brown-black,
including all tarsi. Wing: stigmatal section of subcostal cell
blackish; apical blackish spot on vein 3 relatively compact,
extending to vein 4, but not to vein 2; brown spot present
on anterior crossvein; discal crossvein unmarked; marks
on fork of veins 2 and 3 and on basal crossvein absent or
barely perceptible; axillary lobe and squama white. Halter
creamy. Abdominal tergites shining black.

Head. Height of cheek 0.11-0.14 of height of eye.

Figures 42-45. Signa mouldsi n.sp. 42, wing. 43, diagram of female abdominal tergites 3-6, showing positions of
spiracles. 44, aedeagus (scale = 0.2 mm). 45, epandrium and associated structures (scale = 0.2 mm).

150 Records of the Australian Museum (2001) Vol. 53

Thorax. Humeral and anterior notopleural bristles quite large
in both sexes, the latter inserted well above level of humeral
and posterior notopleural; usually one or few long setulae,
better developed in female, just above anterior notopleural
and in front of transverse suture; in female only, several
long setulae on anterior slope of posterior notopleural callus;
hairs on humeral callus fairly long and pale in male, short
and mostly pale in female; hairs on mesopleuron pale and
of moderate length in both sexes. Wing: distal section of
vein 4 only slightly curved apically; cell-4 index = 0.49-0.58.

Abdomen. Male: tergites with moderately developed hairing,
not much denser towards lateral margins; tergite 5 c. 1.3
times as long as tergite 4; outer surstylus only moderately
elongate, with many unevenly distributed mostly black
spinules, many of them directed basad, its distal section
much surpassing apex of inner surstylus, very obtuse,
obliquely compressed; inner surstylus with few, relatively
minute spinules, in addition to the usual two stout apical
prensisetae; preglans rather long, not very sharply
demarcated from stipe, but more uniformly sclerotised,
slightly expanding distally; glans moderately elongate,
without lobe; bulb quite short, enclosed in short sclerotised
tunic; each of two terminal filaments c. twice as long as
glans, not tapering, very slightly expanded apically. Female:
preabdomen broadly humped; tergite 5 not over half as long
as tergite 4, with hairs mostly pale, a little more crowded
than those on tergite 4, not especially developed on sides,
those near anterior margin much shorter and more crowded,
many of them reduced to minute stumps in relatively
conspicuous sockets; spiracles of segment 5 closer to each
other than to lateral margins of tergite; pleural membrane
with rather short yellow to brown setulae on upper part on
segments 2 to 5, also on segment 4 a few long setulae or
bristles, largely pale yellowish, often some black.

Dimensions. Total length, 6 S 4.1-4.2 mm, $ 9 3.5-4.4
mm; length of thorax, S S 1.6-1.8 mm, 9 9 1.6-1.9 mm;
length of wing, S A 3.7-4.3 mm, 9 9 3.8-4.2 mm; length
of glans of aedeagus 0.41-0.47 mm.

Distribution. Only known from the type locality on the
central west coast of New Caledonia.

Notes. A brief comparative survey of the available species
of Signa indicates that S. mouldsi is distinguishable by the
following character combination: small size; yellow palpus;
filiform arista in both sexes; long humeral bristle in both
sexes; high position of insertion of anterior notopleural
bristle, which is unreduced in both sexes; quite short hairs
on humeral callus in female (longer in male); quite narrow
non-aligned presutural and postsutural pairs of black, non-
pruinescent stripes on mesoscutum; compact apical wing
spot, not touching end of vein 2; absence of shading on
discal and anal crossveins; comparatively slight apical
curvature of vein 4; also, in the female, the vestiture of
tergite 5; and, in the male, the form and degree of spination
of the outer surstylus, and numerous details of the aedeagus.

The specific epithet refers to Maxwell S. Moulds, who
collected the type series.

Genus Par n.gen.

Type species: Elassogaster evitta Malloch.

Description ( 6, $). Somewhat elongate flies, but more robust
than most species of Elassogaster and Microepicausta, with
ovoid abdomen and moderately short legs.

Head markedly higher than long; face not very broad,
minutely roughened and pruinescent, but without hairs, with
rather low, rounded, dorsally narrowed, saddle-like carina;
facial ridge with a series of few fine hairs at lower end;
parafacial with hairs as upper extremity only; occiput
moderately convex below, concave and largely bare above;
the following bristles present: inner and outer vertical,
minute ocellar, one very small black fronto-orbital, large
lateral occipital, large postgenal; postvertical bristle absent;
antennal sockets very narrowly separated. Antenna,
excluding arista, as long as face (male) or distinctly shorter
(female); segment 3 moderately elongate, apically rounded;
compound segment 5+6 much longer than rest of antenna,
slender, with little basal pubescence and, in male only,
spatulate apex. Prelabrum of moderate size, its lower margin
not receding, with minute rugosity and pruinescence on
anterior surface; palpus and proboscis moderately developed.

Thorax. Mesoscutum c. 1.2-1.3 times as long as wide,
pruinescent laterally, but without noticeable pruinescence
on broad median zone; scutellum thick, rounded, dorsally
convex, in dorsal view longer than a semicircle, with rather
dense hairs on entire dorsal surface, and pubescence-
pruinescence towards scutellar suture; pleura extensively
pruinescent; sternopleuron pruinescent and largely short-
haired on at least posterior half; the following thoracic
bristles present: scapulars (in a small group on each side),
humeral (particularly long in male), 1+1 notopleurals, supra-
alar, postalar, posterior intra-alar, dorsocentral, two pairs
of scutellars, mesopleural; paramedian scapular and pre-
scutellar acrostichal absent. Posterior bridge of hind coxa
with fine hairs; femora without ventral spines; fore femur
without dorsal bristles, with long posterior bristles, mainly
on distal half; mid femur with posterior bristles intergrading
with hairs; hind femur with few dorsal bristles. Wing:
membrane almost entirely microtrichose, except for variable
zone in anal cell; first basal cell with microtrichia in basal
part much denser than those in distal part; section of costa
on subcostal cell at least 1.5 times as long as that on second
costal cell; subcosta with several small ventral setulae on
basal part before humeral crossvein; vein 1 with dorsal
setulae more numerous and crowded than in Elassogaster
and Microepicausta’, vein 4 terminating distinctly behind
wing apex, its distal section curving forward only in c. apical
third; anterior crossvein meeting vein 4 slightly before mid¬
length of discal cell; vein 5 without setulae, its short distal
section longitudinally placed, approximately aligned with
its penultimate section; second basal cell c. one third as
long as discal cell; anal crossvein curved on anterior part
only; axillary lobe small, with marginal fringe of hairs less
developed than that on squama; squama not larger than
axillary lobe.

David K. McAlpine: Australasian Platystomatidae 151

Abdomen ovoid, its anterior part not narrowed; tergite 5
unreduced. Male postabdomen: surstyli of moderate length;
preglans distinct, short and simple; glans subcylindrical,
nearly three times as long as its greatest diameter; bulb
undeveloped; terminal filaments two, of similar size, each
nearly twice as long as glans; cercus large, c. three times as
long as wide. Female postabdomen: spiracles not seen,
evidently not enclosed in tergites; ovipositor sheath not
much longer than wide; aculeus (apex only visible) obtuse,
apparently not compressed.

Distribution. Bismarck Archipelago: New Britain; Makada
Island, Duke of York Group.

Notes. The available material of Par (1 $ , Tari Creek, near
Kimbe, central northern New Britain, AM, 1 9, Mount
Sine wit, eastern New Britain, BPB, and 1 9, Makada Island,
assumed holotype of Elassogaster evitta Malloch, AM)
possibly represents a single species. The only feature that
suggests a difference at population level is the very largely
bare anal cell of the male, as compared with the more
extensively microtrichose anal cell of the two females.

I previously (McAlpine, 1973a) placed Par evitta in
Micro epic austa, but the latter is characterised by having
only a single terminal filament on the aedeagus, a condition
which I consider to be a generic autapomorphy, and P. evitta
further differs in detail from Microepicausta as indicated
in the key. Meringomeria is also rather similar, but has
strong dorsal bristles on the fore femur, no dorsal hairs on
the scutellum, and, typically, a compressed, blade-like
aculeus. There is possibly close relationship to Signa, q.v.
for comparison. The wing venation, scutellar vestiture, and,
to a degree, the aedeagus resemble those of the Oriental
and Lemurian genus Icteracantha Hendel, but Par lacks
the ventral femoral spines of that genus, has a more nearly
bare arista, the dorsal setulae near base of vein 3 much more
crowded, and the aculeus not broadly compressed. Par evitta
was originally described in Elassogaster, but it differs from
typical taxa of that genus (including the two available
Afrotropical species) in the absence of a transverse
depression across the centre of the mesoscutum, in the
extensive hairing on the mid-dorsal region of the scutellum,
in the termination of vein 4 distinctly behind the apical
extremity of the wing, and in having the apical section of
vein 5 almost aligned with the previous section. Par also
differs from these probably related genera in the series of

small ventral setulae near the base of the subcosta. Among
other taxa of Platystomatinae, I have noted this condition
only in a few species of Euprosopia, including E. impingens
(Walker). Par and Euprosopia are not very closely related,
and the subcostal setulae must have been acquired
independently in the two genera. Par has the arista 2-
segmented through fusion of segments 5 and 6, whereas
related genera, including Elassogaster, Icteracantha,
Meringomeria, Microepicausta and Signa, generally show
a suture between these segments.

Par is a Latin adjective meaning equal, in reference to
the two similar terminal filaments of the aedeagus. As it is
treated as a Latin noun of variable gender, it is masculine
under Article 30.1.4.2 of the ICZN. The specific epithet
evitta, though grammatically unusual, cannot be treated as
an adjective with variable suffix.

Par evitta (Malloch) n.comb.

Fig. 46

Elassogaster evitta Malloch, 1939a: 116, 153, pi. 4, fig. 16.
Microepicausta evitta (Malloch).-McAlpine, 1973a: 30, 188.

Type material. Apparent holotype, 9 , Makada I. off New
Britain, F.H.T. (AM, formerly in SPHTM), left wing mounted on
slide.

Malloch described this species from “Type, male, and
allotype, Makada Is.” etc., and it is apparent from his
description that he had both sexes. The only specimen of
the type series that I have traced is a female, with a typical
Malloch “Type” label, which should indicate a holotype,
though, from the above, the type should be a male. However,
the left wing of the female specimen is mounted on a slide
and was used for Malloch’s fig. 16, captioned as
“Elassogaster evitta, n.sp. type”. Lee etal. (1956: 316), in
cataloguing types and other material of J.R. Malloch, listed
only one specimen of E. evitta, which they categorized as
holotype, in SPHTM, undoubtedly meaning this female
specimen.

In view of (1) the contradictory statements in the original
publication, (2) the fact that Malloch labelled the female as
“Type”, (3) the assumption by Lee et al. that the female
was the holotype, and (4) the unknown fate of the original
male, it is considered best to regard the above female
specimen as the holotype.

Figure 46. Par evitta (Malloch), wing of holotype.

152 Records of the Australian Museum (2001) Vol. 53

Genus Micro epic austa Hendel
Figs. 47,48

Microepicausta Hendel, 1914a: 52-54. Type species (original
designation) M. gracilis Hendel.

Description. The description by Hendel gives much detail,
though based on a limited sample of one species. The
following statements supplement Hendel’s description.

Head. Face without hairs; carina varying from very low to
high and narrow, but not flat-topped; prelabrum large and
prominent to small and receding.

Thorax. Mesoscutum largely dark, with or without pale
grey-pruinescent median stripe, or largely grey-pruinescent;
anterior notopleural bristle present or absent; scutellum
haired dorsally across most of its width. Fore femur without
strong dorsal bristles, with variably developed, sometimes
undifferentiated, posteroventral bristles. Subcosta ventrally
without setulae; distal section of vein 4 usually gradually
curved forwards to meet costa at or slightly behind apex.

Abdomen. Male: aedeagus (where known) with short,
compact glans and single, long terminal filament. Female:
tergite 5 unreduced; spiracle 5 in membrane near posterior
margin of tergite 5; aculeus slender, its apex compressed
and very acute to relatively thick and obtuse.

Distribution. Australasian Region: New Guinea; Bismarck
Archipelago—New Britain, New Ireland; Solomon
Archipelago; Australia—eastern and southern coasts,
including northern Tasmania. Oriental Region: ?far western
Indonesia. In temperate Australia all records are from the
sea coast.

Notes. I have seen at least nine species of Microepicausta.
Described species known to me (not necessarily all valid)
are: M. albopilosa (de Meijere, 1915), M. gracilis Hendel,

1914a, M. marginalis (Malloch, 1940), M. lineata (de
Meijere, 1915), M. terraereginae (Malloch, 1928).
Elassogaster sangiensis de Meijere, 1916 from western
Indonesia (AMST) perhaps belongs in Microepicausta , but
the aedeagus is undescribed. Microepicausta evitta (Malloch,
1939a) is no longer included in Microepicausta (see above
under Par). There is no published key to species.

The single terminal filament of the aedeagus, now
confirmed for almost all species, is a remarkable condition
not known to me in other Platystomatinae. In one or two
species of Duomyia and in some specimens of the African
genus Bromophila Loew there appears to be a single
terminal filament, but this structure has two separate lumina
and is the product of fusion of the usual two filaments; this
is not the case in Microepicausta. Reduction to a single
terminal filament is probably a groundplan autapomorphy
for the genus Microepicausta.

Adults of Microepicausta are usually found in sandy
areas. In eastern Australia they are often found in coastal
dune scrub or on the grass Spinifex sericeus on or near
beaches. In northern Papua New Guinea I have found them
on low herbage at sandy stream margins far from the coast.
A specimen from New Britain, not determined to species,
is labelled “coastal area, rainforest”.

Genus Elassogaster Bigot
Fig. 49

Elassogaster Bigot, 1860a: 546. Type species (monotypy) E.
metallicus Bigot.

Myiodella Rondani, 1873: 290. Type species (monotypy) M.
brachialis Rondani.

Epicausta Loew, 1873: 46. Type species (designated Steyskal,
1980) Epicausta nigra Wulp (= Myiodella brachialis Rondani).

Description. With access to so few of the accepted species, I
am not prepared to give a formal description. Hendel (1914a)
simply listed the differences from Plagiostenopterina s.l., and
quoted Bigot’s uninformative original description.

Figures 47-48. 47, Microepicausta “sp. 2” (near Musgrave, AM), wing. 48, Microepicausta “sp. 1” (Nullica Beach,
AM), distal part of aedeagus (scale = 0.2 mm). Abbreviations: f, terminal filament (single); g, glans.

David K. McAlpine: Australasian Platystomatidae 153

Figure 49. Elassogaster linearis (Walker), wing.

Distribution. Australasian Region: Moluccas—Buru
(AMST); New Guinea; Bismarck Archipelago—New
Britain; Solomon Archipelago; northern and eastern
Australia. Afrotropical Region: Arabian Peninsula;
Madagascar; Mascarene Islands; Africa—widely distrib¬
uted. Palaearctic Region: Japan. Oriental Region: widely
distributed.

Notes. The type species, E. metallica Bigot, is unknown to
me (as it was to Hendel), but Hendel and all subsequent
authors have interpreted Elassogaster, possibly correctly,
as the generic concept that includes the well known E.
linearis (Walker) (syn. Dacus sepsoides Walker) and some
closely related African and Oriental species.

Elassogaster is much in need of a thorough taxonomic
revision. Five species have been transferred from
Elassogaster to Microepicausta (McAlpine, 1973a;
Evenhuis, 1989), in accordance with my view on the limits
of these genera, but E. evitta Malloch is now placed in Par
(q.v.). Hardy (1959) transferred Dacus sordidus Walker to
Elassogaster, and synonymised Dacus varialis Walker
therewith from examination of type material of both nominal
species (in BM). I am unable to evaluate this action at
present. Evenhuis (1989) listed six Australasian species of
Elassogaster, and I am not aware of any undescribed species
in this Region.

Australasian species of Elassogaster differ from those
of Plagiostenopterina in the absence of fine hairs on the
face, the longer anal cell which is not or only slightly shorter
than the distal section of vein 6, and the absence of a dark
longitudinal stripe covering the first basal cell. These
Elassogaster species differ from Microepicausta in having
the scutellar hairs restricted to the sides or absent, and two
terminal filaments instead of one on the aedeagus.

Larvae of Elassogaster linearis have been found
consuming the eggs of the locust, Locusta migratoria
(Linne) (Orthoptera: Acrididae), in the Philippines and
Papua New Guinea (AM). However, specimens that may
be conspecific have been reared from rotting potato tubers
in Western Highlands Province, Papua New Guinea, and
others are labelled as a secondary pest of taro roots (?
rhizomes, Colocasia esculenta ) in Milne Bay Province
(AM), and larvae have also been found “in rotting asparagus
crowns” near Katherine, Northern Territory, Australia.

Genus Aetha n.gen.

Type species: Aetha cowanae n.sp.

Description ($, 8 unknown). Flies of moderate build, with
anteriorly narrowed but not petiolate abdomen and
moderately short legs.

Head of moderate proportions, but markedly higher than
long; face concave in profile, without median carina; facial
ridge with few fine hairs only; parafacial pruinescent,
without hairs; occiput entirely convex, with many fine, pale
hairs on most of surface; the following bristles present: inner
and outer vertical, weakly differentiated postvertical, two
short fronto-orbitals, minute ocellar, postgenal; lateral
occipital bristle absent without trace. Antenna (excluding
arista) a little shorter than face on median line; segment 3
not attenuated, acute dorsoapically; segment 6 filiform,
minutely pubescent. Prelabrum and mouthparts normal.

Thorax. Mesoscutum c. as long as wide; scutellum extensively
haired dorsally; subscutellum with a well-developed series of
setulae on each side; the following thoracic bristles present:
scapular, humeral, 1+1 notopleurals, supra-alar, postalar,
posterior intra-alar, dorsocentral, two pairs of scutellars,
mesopleural; prescutellar acrostichal absent. Posterior bridge
of hind coxa bare; femora not spinose; fore femur with weakly
differentiated dorsal and long, fine posteroventral bristles; hind
femur with few dorsal bristles distally. Wing: subcosta without
ventral setulae, not strongly flexed near distal end; section of
costa on subcostal cell shorter than that on either side of it;
anterior crossvein located before mid-length of discal cell;
terminal section of vein 4 on its entire length converging with
vein 3; first basal cell entirely microtrichose; second basal cell
more than half as long as discal cell; axillary lobe with marginal
fringe of hairs not noticeably longer than those on squama;
squama forming a rather small lobe, slightly less in area than
axillary lobe.

Abdomen somewhat ovoid; tergite 1+2 somewhat narrowed
anteriorly; tergite 5 unreduced; tergite 6 small and more or
less concealed, well sclerotised; aculeus very slender, not
much compressed, with rounded apex.

Distribution. Australia: tropical and subtropical Queensland
and Northern Territory.

154 Records of the Australian Museum (2001) Vol. 53

Notes. Aetha is apparently monotypic, and only females
are known.

Aetha runs near Pogonortalis and Microepicausta in the
generic key of Me Alpine (1973a) and near Pogonortalis in
that of Malloch (1939a). It differs from Pogonortalis in the
absence of the lateral occipital bristle, the haired scutellum,
the virtual absence of noticeable areas of thoracic
pruinescence, the shorter stigmatal section of the costa (on
subcostal cell), and in the termination of vein 4 before the
wing apex. It differs from Microepicausta in the extensively
pubescent arista, broader thorax, shorter stigmatal section
of costa, more basally located anterior crossvein, and the
termination of vein 4 before the wing apex.

Aetha cowanae was initially sorted as a species of
Elassogaster, but it differs from that genus in its more
uniformly convex occiput, presence of distinct fronto-orbital
bristles, absence of the lateral occipital bristle, extensively
pubescent arista, broader thorax, more extensively haired
scutellum, relatively short fore coxa, and different contour
of the terminal section of vein 4. Aetha also has some
resemblance to Tarfa n.gen. of the Loyalty Islands,
particularly in contour and relations of vein 4. See under
that genus for comparison.

Aetha differs from all the above genera in possessing
a group of setulae on each side of the postscutellum. The
only other Australasian platystomatid taxon with setulae

in this position is the subgenus Plagiostenopterina of
Plagiostenopterina, but in the latter they are usually fewer
and smaller. Aetha differs from Plagiostenopterina in the
absence of the lateral occipital bristle, absence of hairs on
the face, broader thorax, reduced thoracic pruinescence,
shorter fore coxa, much longer second basal cell, largely
unpigmented first basal cell, and more extensive pubescence
on the arista. Aetha and subgenus Plagiostenopterina are
not so close morphologically, and presumably phylo-
genetically, as to allow any probability that the setulosity of
the postscutellum is a homologous condition in the two taxa.

The generic name is latinised from the Greek aithos,
burnt, in reference to the dark coloration, and is feminine.

Aetha cowanae n.sp.

Figs. 50-52

Material examined. Holotype, $, Northern Territory:
Mudginberri, near Jabiru, iii—iv. 1971, Manitoba trap,
A.L.D., H.A.S. (ANIC). Paratypes, Northern Territory: 2
$ 9, Black Point, Cobourg Peninsula, i, ii. 1977, E.D.E.,
T.A.W. (AM, ANIC); 1 9, Wollogorang (Homestead), c.
17°13'S 137°57'E, ix.1930, T.G.C. (ANIC).

Figures 50-52. Aetha cowanae n.sp. 50, wing. 5 1, head. 52, thorax (setulae of mesoscutum omitted); ps, postscutellar
setulae.

Other material. Queensland: 1 9 , Peach Creek, near Coen,
xi.1979, M.S.M., B.J.M. (AM); 1 9 , Charleville-Bollon Road,
70 km SE of Charleville, 26°52'S 146°35'E, x.1991, G.D. (AM).

Description ($). Appearance and habitus resembling those
of the familiar Pogonortalis doclea (Walker). Coloration
predominantly black and somewhat shining; major bristles
of head and thorax black; hairs of most parts whitish.
Postfrons, face, parafacial, anterior part of cheek, and
antenna brownish-tawny; eye almost encircled by narrow
whitish-pruinescent stripe. Prelabrum greyish brown; palpus
brownish, with tawny apex, with mainly black hairs above
and more numerous white hairs below. Pleurotergite largely
grey-pruinescent; thorax otherwise almost without pale
pruinescent zones. Legs black to blackish brown. Wing
hyaline; both costal cells, base of marginal cell, large blotch
at end of vein 2, and lighter zone near middle of first basal
cell brown; axillary lobe and squama white. Halter dark
brown. Abdomen without pale pruinescent zones on tergites.

Head. Eye 0.67-0.71 as long as high; height of cheek 0.17-
0.18 as high as eye; occipital region extensively haired, but
postocular setulae undifferentiated.

Thorax. Mesoscutum, humeral callus, notopleural region,
mesopleuron, pteropleuron, sternopleuron, and dorsal
surface of scutellum extensively haired. Wing: cell-4 index
= 0.34-0.37.

Abdomen. Tergite 5 only slightly shorter than tergite 4.

Dimensions. Total length 5.2-6.3 mm; length of thorax 2.0-
2.4 mm; length of wing 4.1-4.7 mm.

Distribution. Northern Territory: widely distributed in less
arid districts; Queensland: Cape York Peninsula and
Charleville district in southern inland (isolated records).

Notes. Aetha cowanae resembles Pogonortalis doclea in
superficial appearance, at least as dried material. In the field
it should be easily distinguished by the large preapical dark
wing spot and lack of an apical spot.

Locality data indicate at least seasonally dry habitats.
The Peach Creek locality includes riverine forest (M.S.
Moulds, pers. comm.).

The specific epithet refers to Sally Cowan who has
assisted in many of my dipterous projects.

Genus Pogonortalis Hendel

Fig. 53

Pogonortalis Hendel, in de Meijere, 1911: 370 (footnote). Type
species (monotypy) Pogonortalis uncinata de Meijere.
Pogonortalis.-Hendel, 1914a: 16, 84-85. Type species given twice
in error as Pogonortalis barbifera Hendel (not originally
included) = Trypeta doclea Walker.

Pogonortalis- Hendel, 1914b: 6. Type species given in error as
Pogonortalis barbata Hendel, sic (not originally included) =
Trypeta doclea Walker.

David K. McAlpine: Australasian Platystomatidae 155

Figure 53. Pogonortalis doclea (Walker), male. After Hendel
(1914a), retouched.

Description. Hendel (1914a) gave a detailed description,
but the particular characters of Pogonortalis similis Hendel
should be discounted, as that species is now placed in
Plagiostenopterina (see McAlpine, 1973a). The following
notes supplement Hendel’s description.

Male abdomen. Surstyli short; preglans sharply defined,
separated from the more strongly sclerotised stipe by a
constriction; glans simple, short, stoutly ovoid; bulb little
developed; paired terminal filaments long, subequal.

Female abdomen. Tergites 4 and 5 large; tergite 6 very short
and generally concealed; spiracles 3, 4, and 5 in pleural
membrane close to lateral margins of tergites; ovipositor
sheath rather short; aculeus slender, obtuse, not compressed.

Distribution. Australasian Region: Micronesia—Guam;
Australia—southern and eastern parts, Lord Howe Island;
Norfolk Island. Oriental Region: Java. Nearctic Region:
California (introduced).

Notes. I regard Hendel, not de Meijere, as the author of
Pogonortalis, because the latter quoted the descriptive
statements for the genus from a letter from Hendel, who
originated the name.

Previously (McAlpine, 1973a: 31) I expressed doubt as
to the relationships of the type species, Pogonortalis
uncinata de Meijere. Since then I have examined type
material (AMST) and confirmed its close relationship to
Pogonortalis doclea and the consequent stability of the
generic name.

A key to the species was given by Paramonov (1958),
but he included Pogonortalis similis Hendel, now placed

156 Records of the Australian Museum (2001) Vol. 53

in Plagiostenopterina, and the species Pogonortalis hians
Schneider & Me Alpine, 1979 should be added. Six species
are described, and I am not aware of any undescribed
species. Pogonortalis commoni Paramonov is still only
known to me from the holotype.

All the species have a low anteroventral keel at about
the distal third of the hind femur, but this may be little
developed in diminutive specimens. The male genitalia of
Pogonortalis doclea are illustrated by Steyskal (1961).

Pogonortalis doclea is very common in gardens in Sydney
and other Australian towns. Adults of P. doclea and P. howei
Paramonov are attracted to fresh mammalian dung for feeding.

Some observations on sexual behaviour of Pogonortalis
doclea have been made (MeAlpine, 1973b), and fighting
between males of this species has been recorded (Me Alpine,
1975).

Genus Apactoneura Malloch
Fig. 54

Apactoneura Malloch, 1930b: 223. Type species (original
designation) A. flavicornis Malloch.

Description. Medium-sized to rather large, moderately
elongate flies, reminiscent of Plagiostenopterina in habitus,
largely black, with heavy dark wing markings.

Head slightly higher than long; face with slight development
of median carina on upper part only, with low, rounded,
saddle-like median elevation, without hairs; facial ridge
almost without hairs, except where it meets peristomial
series; parafacial without hairs, but with well-developed
pubescence-pruinescence; the following bristles present:
inner and outer vertical, small postvertical and ocellar, two
moderately small reclinate fronto-orbitals, postgenal; lateral
occipital bristle absent. Antenna, excluding arista, much
shorter than face; segment 3 little more than twice as long
as wide; segment 6 with sparse, minute pubescence on basal
part only, in male apically spatulate. Prelabrum moderately
developed; palpus rather long and narrow.

Thorax. Mesoscutum c. 1.2 times as long as wide, without
pruinescent median stripe; scutellum slightly longer than a
semicircle, slightly flattened dorsally and finely rugose,

without hairs, pubescent on free margins; mesopleuron
pruinescent on upper part only; sternopleuron with pale
pruinescent stripe along upper margin, extending to fore-
coxal cavity; pteropleuron and posterior pleural sclerites
pruinescent; the following thoracic bristles present; humeral,
1+1 notopleurals, supra-alar, postalar, posterior intra-alar,
small dorsocentral (near scutellar suture), prescutellar
acrostichal, three long scutellars, mesopleural; scapular
bristles absent. Posterior bridge of hind coxa haired; fore
femur with an extensive series of thick posteroventral spines
and at most rudimentary anteroventral ones; mid femur with
few anteroventral and posteroventral spines distally; hind
femur longer and thicker than other femora, with a well-
developed series each of anteroventral and posteroventral
spines not extending to base. Wing: setulae on veins (other
than costa) restricted to dorsal surfaces of vein 1 (beyond
humeral crossvein) and vein 3; section of costa on subcostal
cell c. as long as that on second costal cell and c. twice as
long as that on marginal cell (Fig. 54); supernumerary
crossvein dividing first posterior cell meeting vein 4 just
beyond discal crossvein; anterior crossvein meeting vein 4
slightly beyond apical third of discal cell; basal cells and
alula entirely microtrichose; squama forming a rounded
lobe, only slightly greater in area than axillary lobe.

Abdomen. Male: tergite 5 as long as tergite 4; preglans little
differentiated; glans stoutly subcylindrical; bulb little
developed; paired terminal filaments subequal, each less
than half as long as glans. Female: spiracle 5 located in
pleural membrane, well below lateral margin of tergite 5;
tergite 6 small but well sclerotised; ovipositor sheath
moderately long, with long posterior marginal bristles;
aculeus slender, obtuse.

Distribution. Tropical Polynesia: Samoan Islands.

Notes. Apactoneura has one recognized species, but
possible differences between the populations on Savaii and
Upolu are yet to be evaluated.

Apactoneura is readily recognizable from its wing
venation and ventral spination on all femora, combined with
the enlargement of the hind femur. No other platystomatid
genus in this Region has a regular crossvein dividing the
first posterior cell. This condition is consistent in the five
specimens known to me, from both island populations.

Figure 54. Apactoneura flavicornis Malloch, wing. After A.J.E. Terzi, in Malloch (1930b).

David K. McAlpine: Australasian Platystomatidae 157

Figure 55. Meringomeria neurostigma (Bezzi), wing. After A.J.E. Terzi, in Bezzi (1928).

However, the relationship to other platystomatine genera is
obscure. Apactoneura resembles the Oriental genus
Icteracantha Hendel in the presence of stout ventral spines
on all femora, presence of well-developed hairs on the
posterior bridge of the hind coxa, and in the general habitus,
but differs in the lack of forward curvature in the distal
section of vein 4, the presence of three instead of two pairs
of bristles and the absence of dorsal hairs on the
scutellum. The genus also resembles Lamprogaster , but
no Lamprogaster species known to me has well-developed
femoral spines or so well-developed posterior hairing on
the hind coxa; also the squama is consistently larger in
Lamprogaster and the facial carina is distinctly margined
on each side. Apactoneura differs from Plagiostenopterina
in the lack of hairing on the face, and absence of apical
forward curvature of vein 4.

Genus Meringomeria Enderlein

Fig. 55

Meringomeria Enderlein, 1924: 108. Type species (original
designation) Dacus trivittatus Walker.

Charax Walker, 1861a: 325 (preocc. Charax Gronovius, 1763).

Type species (monotypy) C. planidorsum Walker.
Stenopterella Malloch, 1931: 13, 16 (as subgenus of Plagio¬
stenopterina). Type species (original designation) Dacus
trivittatus Walker.

Description. Flies of elongate habitus, with rather slender
legs of moderate length.

Head markedly higher than long; face pruinescent, without
hairs, typically with rounded, saddle-like carina, which is
depressed near middle (Oriental species), or with carina
broadly almost flat-topped on upper part (Fijian species);
facial ridge with fine uniseriate hairs near lower extremity
only; parafacial without hairs; occiput moderately convex
below, concave to almost flat above, haired mainly on
dorsolateral and lower parts; the following bristles present:
inner and outer vertical, minute ocellar, lateral occipital,
postgenal; fronto-orbitals minute or absent; postvertical
absent. Antenna about as long as face on median line, or
slightly longer in some males (Oriental species), or distinctly
shorter than face (Fijian species, females only known);
segment 3 elongate, usually distally tapered but not pointed;
segment 6 filiform, usually with basal pubescence (Oriental
species), or plumose on basal part (Fijian species).

Prelabrum moderately large and prominent, pruinescent and
often also finely rugose (Oriental species), or almost smooth
and shining (at least one Fijian species); palpus moderately
developed, broadly rounded at apex.

Thorax. Mesoscutum longer than wide, extensively
pruinescent, with or without pair of broad, dark, shining
longitudinal stripes (Oriental species), or with relatively
little pruinescence and largely shining (Fijian species);
mesopleuron extensively rather thinly pruinescent (Oriental
species), or with dense pale pruinescence restricted to
posterior part (Fijian species); scutellum somewhat convex,
more or less pruinescent, with few lateral hairs only; the
following thoracic bristles present: one well-developed
lateral scapular (Oriental species), or several small such
bristles (at least one Fijian species), humeral, 1 + 1
notopleurals, supra-alar, postalar, posterior intra-alar,
dorsocentral, two pairs of scutellars, mesopleural;
paramedian scapular present or absent. Posterior bridge of
hind coxa with fine hairs; fore femur with dorsal bristles,
with long posteroventral bristles (Oriental species), or latter
scarcely differentiated (one Fijian species); mid femur
without bristles; hind femur with dorsal bristles beyond mid¬
length; mid tibia with apical ventral spur. Wing with dark
markings quite restricted to costal region (Oriental species),
or (in Fijian species) either with more extensive markings
(Fig. 55) or completely without dark markings; membrane
largely microtrichose, except for variable bare zones in basal
and anal cells; section of costa on subcostal cell considerably
longer than that on second costal cell (Oriental species), or
scarcely longer (Fijian species); anterior crossvein meeting
vein 4 near or slightly beyond mid-length of discal cell;
distal section of vein 4 strongly curved forward near distal
extremity to meet costa at or slightly behind apex of wing;
vein 3 also curved to a smaller extent near apex to converge
with vein 4; vein 5 without setulae, its distal section short
(Oriental species), or very short (less than one quarter length
of discal crossvein) in Fijian species; second basal cell c.
one quarter to one third as long as discal cell; anal crossvein
curved, sometimes more strongly so on anterior part;
axillary lobe moderately developed; squama forming a short
lobe, scarcely greater in area than axillary lobe.

Abdomen ovoid to subcylindrical, not significantly narrowed
anteriorly. Male abdomen (unknown for Fijian species):
surstyli elongate, straight, apex of outer surstylus not
extending much beyond that of inner one; aedeagus with
preglans differentiated but simple; glans rather stoutly

158 Records of the Australian Museum (2001) Vol. 53

subcylindrical; bulb little developed; paired terminal
filaments present; cerci moderately large. Female abdomen
(Oriental species): spiracles 4 and 5 situated within
respective tergites and at least the latter visible dorsally;
ovipositor sheath well developed, scarcely tapered
posteriorly; aculeus very broad and compressed, obtuse.
Female abdomen (available Fijian specimen): spiracles not
situated in tergites, those of segment 5 somewhat
approximated close behind posterior margin of tergite 5;
ovipositor sheath very like that of Oriental species, but
aculeus not visible.

Distribution. Australasian Region: Fiji. Oriental Region:
southern China (Guangdong = Kwangtung Province;
Taiwan) and Sikkim to Palawan and Western Indonesia.
There appear to be no records from localities between
Wallace’s Line and Fiji.

Notes. Although Meringomeria (or Stenopterella) has often
been included as a subgenus of Plagiostenopterina, Malloch
(1931) found difficulty in defining such a diffuse genus, and
more recently Meringomeria has been given generic status
(McAlpine, 1973a; Evenhuis, 1989). Steyskal (1966) provided
a useful key to species, but it seems to me that one or two of
the categories keyed may be complexes of species rather than
single species. This is suggested by the wide distributions
accepted for some species, and a study of limited material,
representing c. six Oriental species now available to me.

There would seem to be at least two species, one
undescribed, in Fiji, as the only Fijian specimen at present
available differs from Meringomeria neurostigma (Bezzi,
1928) in the absence of wing markings (and perhaps in several
other characters, as suggested by Bezzi’s description). These
Fijian species are more similar to each other than to the
Oriental species, as indicated by my comparisons in the
above redescription, but to what degree they need taxonomic
segregation can only be decided when better study material
is available.

Genus Plagiostenopterina Hendel

Figs. 56, 57

Plagiostenopterina Hendel, 1912a: 3. Type species (original
designation) Dacus aeneus Wiedemann.

Stenopterosoma Malloch, 1939a: 114. As subgenus of Plagiosteno¬
pterina. Type species (monotypy) P. orbitalis Malloch (= Dacus
lativentris Walker).

Description. See McAlpine (1973a).

Distribution. Very widely distributed in the tropics of the
Old World. Australasian Region: extending from the Moluccas
to the Solomon Archipelago; Tropical Polynesia—Samoan
Islands; tropical Australia—coastal and subcoastal districts
of Queensland and Northern Territory.

Notes. In accordance with McAlpine (1973a) the taxa
Carolimyia Malloch, Meringomeria Enderlein, and
Peronotrochus Enderlein, which are sometimes given as
subgenera of Plagiostenopterina , are now excluded from
the genus. See Steyskal (1977) for synonymy of these taxa.
Steyskal (1980) has corrected the original citation of the
genus and the means of type designation. McAlpine (1973a)
reviewed the Australian species of the genus, but I am now
very doubtful if populations in New Guinea, Solomon
Archipelago, and Australia are correctly referred to P.
enderleini Hendel. Of the two subgenera represented in
Australasia, Plagiostenopterina s.str. is also widely
distributed in the Afrotropical and Oriental Regions, while
Stenopterosoma Malloch is known only from New Guinea
and northern Queensland.

Adults of subgenus Plagiostenopterina are often
numerous in disturbed vegetation or the interface of forest
and cultivation. Those of subgenus Stenopterosoma are
found mainly in rainforest. Adults of both subgenera are
attracted to fresh mammalian dung.

Figures 56-57. Plagiostenopterina (Stenopterosoma) claudiana McAlpine. 56, wing. 57, aedeagus (scale = 0.5 mm).

David K. McAlpine: Australasian Platystomatidae 159

Key to Australasian subgenera of Plagiostenopterina

1 Scutellum extensively setulose; postscutellum with fine setulae

on each side; fronto-orbital bristles absent. Plagiostenopterina s.str.

-Scutellum and postscutellum without setulae; two pairs of fronto-

orbital bristles present. Stenopterosoma

Figure 58. Pseudepicausta chalybea (Doleschall), wing.

Genus Pseudepicausta Hendel
Fig. 58

Pseudepicausta Hendel, 1912b: 14. Type species (designated
Hendel, 1914a) Herina chalybea Doleschall.

Description. See Hendel (1914a). That description, together
with characters cited above in my key, should enable
identification of the Australasian and Oriental species.

Distribution. Australasian Region: Moluccas; New Guinea;
New Britain; Solomon Archipelago. Also Oriental and
Afrotropical Regions.

Notes. Pseudepicausta seems to be a well defined genus
within the Australasian and Oriental Regions. Some
Afrotropical material (e.g., from Madagascar) is slightly
less typical, e.g., in having better developed fronto-orbital
bristles and shorter ovipositor sheath. However, I do not
have sufficient Afrotropical species available for invest¬
igation of their relationships.

Malloch (1939a) gave a key to species of New Guinea
and nearby islands, but included P. lagarosia Hendel, which
belongs in Pseudocleitamia (q.v.). Evenhuis (1989) listed
nine Australasian species, but the list includes P. rufitarsis
(Macquart, 1847, as Tephritis ). The latter is not recognizable
from the brief description, and the type locality (Nouvelle
Holland = mainland Australia) is beyond the known range
of Pseudepicausta. The holotype (Bigot collection, OX,
examined by author September 1998) is in exceedingly poor
condition with almost no significant features visible, but is
very doubtfully referred to the genus Rivellia. Without a
thorough revision of the genus, it is difficult to estimate
how many of the other listed species are valid. I have seen
at least one probably undescribed species from southern
Papua New Guinea (AM).

Genus Pseudocleitamia Malloch
Fig. 59

Pseudocleitamia Malloch, 1939a: 104. Type species (original
designation) P. setigera Malloch.

Description. A brief description was given by Malloch, and
the description of Pseudepicausta lagarosia Hendel (1914b:
118-119), now included in Pseudocleitamia , indicates some
characters of the male. I do not have on hand enough
material for a redescription. The genus is not represented
in BPB and Australian collections.

Distribution. New Guinea: Aru Islands; Papua New
Guinea—mainland (Madang and Oro Provinces).

Notes. The nominal species Dacus pompiloides Walker,
1859, Pseudepicausta lagarosia Hendel, 1914a, and
Pseudocleitamia setigera Malloch, 1939a, are referable to
this genus. I have examined the types of D. pompiloides
and Pseudocleitamia setigera (BM). Hendel (1914a: pi. 6,
fig. 121) and Malloch (1939a: pi. 4, fig. 5) each illustrated
the wing of their holotypes, showing the quite distinctive
pattern and venation of Pseudocleitamia. Among the more
distinctive characters mentioned by Malloch for the genus
are “mesopleura, sternopleura, pteropleura with numerous
short stout bristles”, and “the very narrow upper portion of the
frons”. The latter condition is not atypical of Pseudepicausta
species, but the multiple bristling of the pleura is not recorded
in that genus, nor is it present in the type of Pseudocleitamia
pompiloides (and probably that of P. lagarosia ), in which the
thoracic pleura have slender hairs and only the usual bristles.
The type of P. lagarosia has a series of broadened, lanceolate
posteroventral bristles on the fore femur, but the fore femoral
bristles of Pseudocleitamia setigera and P. lagarosia are not
thus modified. The types of P. pompiloides and P. lagarosia

160 Records of the Australian Museum (2001) Vol. 53

Figure 59. Pseudocleitamia setigera Malloch, wing of holotype.

are males, whereas that of Pseudocleitamia setigera is a
female. Hardy (1959) examined the type of D. pompiloides
and placed it in Pseudepicausta, but he did not make
comparison with Pseudocleitamia lagarosia nor mention
the genus Pseudocleitamia. Evenhuis (1989) made the
combination Pseudocleitamia pompiloides (on my
suggestion) and gave P. lagarosia as a junior synonym,
leaving P. setigera as a separate species.

From my studies of morphology of many platystomatine
taxa, I think that the presence of numerous stout pleural
bristles as in the type of Pseudocleitamia setigera is likely
to be female-restricted, whereas the presence of compressed,
lanceolate femoral bristles, as in the type of P. lagarosia , is
likely to be male-restricted. On the other hand, there are
differences which suggest that these species and P.
pompiloides are distinct from one another. Pseudocleitamia
setigera differs from the other two species in having a
marked sinuosity in vein 2 just before mid-length (not very
clearly shown in Malloch’s fig. 5, but see my Fig. 59 based
on same specimen). Pseudocleitamia setigera differs from
P. pompiloides in having the hyaline prediscal band quite
bare from the costa to halfway across the discal cell; also
the sub-basal hyaline zone in the submarginal cell, smaller
adjacent zone in marginal cell, and a substantial adjacent
zone in discal cell are bare, whereas these zones are entirely
microtrichose in the type of P pompiloides. The type of P
pompiloides lacks the modified lanceolate femoral bristles
present in that of P. lagarosia (confirmation by J. Chainey
in litt.), though both specimens are males.

With each of the apparent species of Pseudocleitamia
known to me from only one specimen, it is not possible
adequately to describe the range of morphological variation
in the genus nor fully to characterise the species.

Genus Lamprophthalma Portschinsky

Lamprophthalma Portschinsky, 1892: 16-17, pi. 1, figs. 7, 7a.

Type species (monotypy) L. metallica Portschinsky.

Description. The annotated description by Hendel (1914a:
45, largely quoted from Portschinsky) is probably adequate
for identification of this marginally or doubtfully
Australasian genus. I can add that the frontal tubercle is
quite variable in development among Oriental species, but

this variation may not indicate that these species are not
closely interrelated. The face has fine hairs (macrotrichia)
in some species, but not in others. The putative hind coxal
spine, figured and described by Portschinsky, is probably
really on the trochanter and male-restricted. It is absent in
some species. The statement “nervi omnes inermes” does
not apply to veins 1 and 3.

Distribution. Australasian Region: Moluccas—?Seram, see
below. Afrotropical Region. Palaearctic Region: Iran; China;
Japan. Oriental Region: widely distributed.

Notes. The only Australasian record for Lamprophthalma
known to me is that of the type specimen of Dacus
sepedonoides Walker, which I have not examined. Hardy
(1959: 181) corrected the type locality from “Waigiou”
Island to Ceram (Seram), and made the combination
Lamprophthalma sepedonoides in accordance with the
placement of the type in the BM collection. As this supposed
type is possibly not from the given type locality, and as Hardy
did not appear to vouch for its generic placement, the
Australasian occurrence of this genus needs confirmation.

I transfer the two following Oriental species from
Plagiostenopterina to Lamprophthalma from study of type
material (AMST): Lamprophthalma egregia (de Meijere,
1924: 41-42) n.comb.; Lamprophthalma medionotata (de
Meijere, 1924: 40-41) n.comb.

Genus Conicipithea Hendel
Fig. 60

Conicipithea Hendel, 1912b: 14. Type species (monotypy) Dacus
addens Walker.

Description. Hendel (1914a) gave a fairly detailed
description, which I supplement from a specimen of C.
addens from Kendari, south-east Sulawesi (AM).

Head. Face without hairs; parafacial with several short
setulae on central part.

Wing. Costa much narrowed but not broken at end of
subcosta.

David K. McAlpine: Australasian Platystomatidae 161

Figure 60. Conicipithea addens (Walker), wing of male (Sulawesi, AM).

Male postabdomen. Preglans differentiated, but not separated
from stipe by a constriction; glans narrowly elongate,
subcylindrical; paired terminal filaments very short.

Distribution. Australasian Region: Moluccas—Ambon.
Oriental Region: Sulawesi (Celebes).

Notes. Conicipithea is a little known but easily recognized
genus, the very long antenna with black setulae on segment
3 and the relatively long hairs on the suprasquamal ridge
provide a unique character combination among Australasian
platystomatids. The haired suprasquamal ridge and flat
facial carina suggest a relationship to Duomyia, but
Conicipithea differs from that diverse purely Australian
genus in the setulose antennal segment 3, the much more
extensively haired segment 6, the narrowed costa at the end
of the subcosta, and the relatively large mesopleural bristle.

It has been assumed that all available material of
Conicipithea represents C. addens , but no direct comparison
seems to have been made between Sulawesi and Ambon
specimens. The wing venation of my single specimen from

Genus Antineura Osten Sacken

Figs. 61, 62

Antineura Osten Sacken, 1881b: 134. Type species (designated
Hendel, 1914a) A. stolata Osten Sacken.

Antineura.-Hendel, 1914a: 12,40-42.

Adantineura Hendel, 1914a: 41 (as subgenus of Antineura , no
included species). Type species (designated Evenhuis, 1989)
Antineura biroi de Meijere.

Adantineura- Hendel, 1914b: 32-34 (four included species).

Description. Hendel (1914a) gave a detailed description,
but this is based mainly on species of subgenus Adantineura,
because the two species of Antineura s.str. from the
Philippines were known to him only from Osten Sacken’s
descriptions, as evidenced from the fact that he only quoted
Osten Sacken’s (1882) descriptions, without citing material
examined (Hendel, 1914b: 32-33). The following
supplementary data are derived from species of both
subgenera, but I only have females of subgenus Antineura.

Figures 61-62. 61, Antineura (Adantineura) sp. (Wewak, AM), wing. 62, Antineura (Adantineura) biroi de Meijere,
head.

162 Records of the Australian Museum (2001) Vol. 53

Head. Postfrons broadly convex on anterior part, depressed
behind convexity; face with broadly rounded, transversely
grooved, finely haired carina for whole length.

Thorax. Humeral bristle absent; mesopleural bristle well
developed (subgenus Antineura), reduced or absent
(subgenus Adantineura) . Wing: veins 3 and 4 distally
convergent; stem of veins 2 and 3 (base of radial sector)
with distinct flexure before (subgenus Antineura) or beyond
(subgenus Adantineura) mid-length, section beyond flexure
thicker and armed with a series of long setulae.

Male postabdomen. Aedeagus relatively simple, with slender
stipe, no differentiated preglans, simple rather compact glans,
and pair of rather short, equal terminal filaments.

Female postabdomen. Tergite 5 large; spiracle 5 located
laterally below margin of tergite; tergite 6 small and
separate, or relatively well developed and sometimes
laterally fused with sternite 6 to form an annular
tergosternite; segment 8 slender; aculeus small, flat, obtuse,
with rounded preapical expansion on each side and pair of
ventral bristles.

Distribution. Australasian Region: Moluccas; New Guinea.
Oriental Region: Philippines.

Of the two subgenera, Antineura is recorded only from
the Philippines. The subgenus Adantineura lives in the
Moluccas and New Guinea. Williston (1908: fig. 107)
published a photograph of a fly determined as “ Antineura
sp. Brazil.” This is apparently a richardiid of the genus
Setellia Robineau-Desvoidy.

Notes. There are six nominal species of subgenus
Adantineura listed as Australasian (Evenhuis, 1989), but
the status of those recorded from the western part of the
range needs reassessment. I have sorted five distinct species
from Papua New Guinea (mainland), apparently including
A. biroi de Meijere and A. kerteszi de Meijere. Specimens
determined by Malloch as A. kerteszi (see Malloch, 1939a:
104; pi. 4, fig. 4) are probably not that species.

Hendel (1914b) inadvertently referred A. sericata Osten
Sacken to Adantineura, though it was correctly excluded
from this subgenus in his key to species. Enderlein (1924:
110) treated Adantineura as a separate genus, but has not
been followed by others.

The appearance of Antineura species suggests mimicry
of pompilid wasps. The species of subgenus Antineura are
particularly large, conspicuous flies.

Genus Duomyia Walker

Figs. 63-67

Duomyia Walker, 1849: 800. Type species (designated Hendel,
1914a) D. obscura Walker.

Campigaster Macquart, 1855: 142. Type species (monotypy) C.
testaceus Macquart.

Euchalcota Loew, 1873:40. Type species (monotypy) Senopterina
decora Macquart.

Helocnemia Enderlein, 1924: 128. Type species (original

designation) Chromatomyia apicalis Walker.

Duomyza Malloch, 1929: 507 (as subgenus of Duomyia ). Type
species (original designation) Duomyia tomentosa Hendel.

Description. See Me Alpine (1973a).

Distribution. Australia: all states, including Torres Strait
Islands and Lord Howe Island—apparently endemic.

The species are very generally distributed. Numerous
species, mostly locally endemic, live in south-western
Australia, two in the Snowy Mountains of the south-east,
one in central Australia, one in the Torres Strait Islands,
and one on Lord Howe Island.

Notes. Duomyia was revised by Me Alpine (1973a), but I
have now seen at least 106 species, including two species
formerly placed in Rhytidortalis (McAlpine, 2000) and more
than 30 undescribed species. Although D. rugifrons
(Thomson) and D. solocifemur (Enderlein) are not
morphologically typical of Duomyia, the intermediate
characters of several species prevent generic separation on
the basis of present knowledge. These few aberrant species
could be mostly excluded by restricting Duomyia to species
with erect hairs on the suprasquamal ridge, but (1) I am
satisfied that the relationships of the species thereby excluded
are not with Rhytidortalis, and (2) I am not prepared to
introduce a new genus for such an arbitrary group of species.

Contrary to the use of Evenhuis (1989), the name
Duomyia punctifrons Malloch, 1929, is invalid under
Articles 10.6 and 59.1 of the ICZN (1999). The replacement
name D. montium McAlpine, 1973a, should be used.

The genus occupies a greater range of habitats than do
most other Australian platystomatid genera. Many species
inhabit seasonally wet to moderately dry Eucalyptus forest
or woodland. Some live only in sandy coastal vegetation,
and populations may be at risk from habitat destruction. Littoral
rainforests harbour more species than upland rainforests.

Adults of some species visit myrtaceous flowers for
feeding, e.g., D. decora (Macquart) at Leptospermum, D.
rugifrons (Thomson) at Kunzea, and D. lutea McAlpine at
Astartea. Adults of several species have been taken at fresh
mammalian dung, and many have been collected at light.

Genus Euprosopia Macquart

Figs. 1, 68-71

Euprosopia Macquart, 1847: 89-90. Type species (monotypy) E.
tenuicornis Macquart.

Pachycephala Doleschall, 1859:115-116 (43^44 in reprint), (preocc.
Pachycephala Vigors, 1825). Type species (designated Enderlein,
1924 for replacement name Oncoscelia ) Pachycephala mohnikei
Doleschall.

Oncoscelia Enderlein, 1924: 136. New name for Pachycephala
Doleschall.

Additional synonymy given by McAlpine (1973a: 132).

Description. See McAlpine (1973a).

Distribution. Australasian Region: Moluccas; New Guinea;
Bismarck Archipelago; Solomon Archipelago; Vanuatu;

David K. McAlpine: Australasian Platystomatidae

Figures 63-67. 63, Duomyia rugifrons (Thomson), male. 64, Duomyia “sp. 17” (Shoal Bay, AM), head of male. 65,
Duomyia sp. (Binnaway State Forest, AM), wing. 66, Duomyia chaetostigma McAlpine, aedeagus (scale = 0.5
mm). 67, Duomyia lonchaeina McAlpine, distal part of aedeagus (scale = 0.5 mm). Abbreviations: bu, bulb; c,
membranous caeca; f, terminal filaments (paired); g, glans; pg, preglans.

164 Records of the Australian Museum (2001) Vol. 53

Figures 68-71. 68, Euprosopia armipes McAlpine, head of male; pt, ptilinal fissure. 69, Euprosopia rete McAlpine,
wing. 70, Euprosopia kurandae McAlpine, wing. 71, Euprosopia acula McAlpine, distal part of aedeagus (scale =
0.5 mm).

Australia—all states, but absent from arid and subarid areas,
and in Western Australia restricted to far north. Oriental
Region: widely distributed. Palaearctic Region: eastern
provinces only.

Notes. The Euprosopia species of New Guinea were revised
by Malloch (1939a), those of the Solomon Archipelago by
Malloch (1940), and those of Australia by McAlpine (1973a).

This is one of the largest platystomatid genera. Evenhuis
(1989) lists 69 species of Euprosopia for Australasia. I have
sorted 38 undescribed species of New Guinea-Melanesia
and a further 15 undescribed Australian species. Steyskal
(1977) recorded 26 species for the Oriental Region, to which
should be added Euprosopia robusta de Meijere, 1916, from
Sumatra. Soos (1984) listed four species for the Palaearctic
Region, two of which are also in Steyskal’s Oriental list.
Thus 151 species (including undescribed ones) have been
recognized, of which 122 are Australasian, but numerous
others undoubtedly exist.

The only species common to the published Oriental and
Australasian lists is E. sexpunctata (Osten Sacken, 1882,
as Notopsila sp.) from the Philippines, with which Hendel
(1914b) synonymised Platystoma atomaria Walker, 1861c,
described from Batchian (Bacan, Moluccas). The latter is a
primary junior homonym of Platystoma atomarium Walker,
1860, from “Makessar” (Sulawesi), a synonym of
Euthyplatystoma rigidum (Walker, 1856). Hendel, from his
listed material, appears only to have seen material of E.

sexpunctata from the Philippines, and his synonymy of the
Moluccan P. atomaria was inferred from Walker’s
inadequate description, which omits many morphological
data relevant to species diagnosis in Euprosopia. I therefore
consider that there is at present no case (1) for assuming
the synonymy of P. atomaria under E. sexpunctata , and (2)
for recording any one species of Euprosopia for both the
Oriental and Australasian Regions.

There is confusion in the listing of Euprosopia miliaria
Hendel in the Australasian catalogue (Evenhuis, 1989:490).
The lectotype of this species (McAlpine, 1973a: 141) is the
holotype of Platystoma pectoralis Walker, 1862 (a
preoccupied name), and E. miliaria is only reliably reported
from the Moluccas (Gilolo or Halmahera), despite the
additional distribution given by Evenhuis. Other material,
recorded by Hendel (1914b) and Malloch (1939a) as E.
miliaria, is referable to other species (see McAlpine, 1973a).
Evenhuis also gave E. diminutiva de Meijere (1913) as a
new synonym of E. miliaria. However, de Meijere
established no such name; he simply misapplied the name
E. diminutiva (Walker) to material from West New Guinea,
which Hendel (1914a, 1914b) erroneously thought to be
conspecific with E. miliaria. The material mentioned by de
Meijere (1 6 ,1 9, Heuvelbivak, Lorentz River, West New
Guinea, AMST) is probably referable to E. rete McAlpine.

Sexual behaviour and its relation to sexual dimorphism
were described for a few Euprosopia species by McAlpine
(1973b).

David K. McAlpine: Australasian Platystomatidae 165

Genus Euxestomoea de Meijere

Fig. 72

Euxestomoea de Meijere, 1913: 377. Type species (designated
Hendel, 1914a) Ortalis prompta Walker.

Description. The description by Hendel (1914a) remains
largely valid, though the genus is now restricted by
transference of E. bipunctata Hendel to Bama. The principal
differences between Euxestomoea and Bama are listed under
the latter genus. Also, Euxestomoea consistently has the
base of vein 1 bare dorsally, and the second section of vein
4 without sigmoid curvature.

Distribution. New Guinea: widely distributed, including
Karkar Island. Bismarck Archipelago: New Ireland.
Australia: Queensland—Iron Range district (E. prompta
(Walker), AM).

Notes. The last published key to species is that of Malloch
(1939a). In addition to the described species E. prompta
(Walker) and E. discifera de Meijere, I have seen an
undescribed species from New Ireland (AM). Material from
mainland Papua New Guinea shows variation which is not
yet assessed taxonomically. Euxestomoea discifera appears
to have no prescutellar acrostichal bristle and two pairs of
dorsocentrals, but it is possible that the posterior
dorsocentral is the homologue of the apparent acrostichal
of the other species, which has migrated laterad.

Genus Bama n.gen.

Type species: Xiria papuana Hennig.

Description (6, $). Flies of rather stout to slightly
elongate build, with legs of moderate length. General
appearance and morphology reminiscent of species of
Cleitamia, Laglaisia (females), or Euxestomoea. Cuticle
of thorax and abdominal tergites sparsely or thinly
pruinescent, largely shining.

Head considerably higher than long, a little wider than high
but not sexually dimorphic in this respect; face bare, without
carina, in profile concave near or above middle, often
slightly convex below; facial ridge with minute hairs in a
single series; occiput slightly concave above, moderately
convex below; the following cephalic bristles present: inner
and outer vertical, two moderately large fronto-orbitals,
postgenal; postvertical and ocellar bristles small or poorly
differentiated. Antenna, excluding arista, much shorter than
face; segment 2 largely bare on medial surface; segment 3
usually apically rounded; segment 6 short-haired for most
of length, the hairs either irregularly placed or with tendency
to be aligned in a dorsal and a ventral series. Prelabrum
well developed but not very deep, its lower margin not
receding; palpus moderately to conspicuously long;
proboscis stout, with broad prementum and labella.

Thorax. Scutellum slightly longer than a semicircle, without
ventral marginal hairs; pleurotergite and postscutellum
without hairs or setulae; the following thoracic bristles well
developed: humeral, 1+1 notopleurals, supra-alar, postalar,
posterior intra-alar, dorsocentral, prescutellar acrostichal,
three pairs of scutellars, mesopleural; scapular bristle absent
or vestigial. Mid coxa with variable armature, which is
sometimes specifically distinctive; fore femur with
posterodorsal and posteroventral bristles, the latter
sometimes enlarged and spinescent; other femora without
ventral bristles and spines; mid femur without posterior
bristles; hind femur with some weak dorsal bristles; mid
tibia with one long apical spur, secondary ones less than
half as long; other tibiae without special armature. Section
of costa on subcostal cell shorter than to somewhat less
than twice as long as that on marginal cell; base of vein 1
(stem-vein, base of R), before humeral crossvein, often with
group of dorsal setulae (smaller than in Euprosopia, absent
in two species); vein 3 with few fine ventral setulae and
more numerous dorsal setulae; anterior cross vein often
longer than penultimate section of vein 4 (some exceptional
species); second section of vein 4 usually with sigmoid
curvature, this curvature sometimes very slight; anal
crossvein curved to variable degree on anterior part only;
squama forming a very short to moderate-sized lobe, its

Figure 72. Euxestomoea prompta (Walker), wing.

166 Records of the Australian Museum (2001) Vol. 53

marginal fringe usually longer than that on distal half of axillary
lobe, but fringe on sinus between lobes conspicuously longer.

Abdomen in male typical of Platystomatinae, rather narrowly
ovoid, with tergites 3, 4, and 5 all large, in female with
tergite 3 medium-sized to very large and tergites 4, 5, and 6
much reduced, often concealed in dried specimens. Male
postabdomen (where known): surstyli rather elongate,
typical of the Platystomatinae; aedeagus with short, more
or less ovoid glans, membranous lobe on bulb, and two
short, unequal terminal filaments. Female postabdomen:
aculeus slender, obtuse.

Distribution. New Guinea: apparently mainland only. All
records and available material are from the eastern half of
the island, i.e. mainland Papua New Guinea. I expect that a
similar number of species lives in West New Guinea, where
the montane Diptera are little known.

Notes. This genus includes three previously described species,
viz. Bama bipunctatum (Hendel, 1914a) (as Euxestomoea sp.)
n.comb., Bama papuanum (Hennig, 1940b) (as Xiria sp.)
n.comb., and Bama strigatum (Hennig, 1940b) (as Xiria
sp.) n.comb. I have seen type material of all three species.
An additional species is described below, and I have also
seen seven undescribed species belonging in Bama.

Euxestomoea differs from Bama in having longer,
strongly divergent postvertical bristles, lateral occipital
bristle present, the medial surface of antennal segment 2
extensively setulose, segment 6 bare on distal half, the
mesopleuron and sternopleuron conspicuously pruinescent
in part, the dorsocentral and intermediate scutellar bristles
differently placed, the mid femur with one stout, isolated
preapical posterior bristle, the mid tibia with two slightly
unequal apical ventral spurs, the anal cell almost entirely
microtrichose, tergites 4 and 5 of the female abdomen well

developed and exposed, the aedeagus with relatively short
preglans and two terminal filaments of similar length. Bama
may be less closely related to Euxestomoea than to Cleitamia
and Laglaisia, but the two latter have only one fronto-orbital
bristle, antennal segment 6 long-haired, no prescutellar
acrostichal bristle, more extensive setulae or hairs on the
scutellum and pleurotergite, base of vein 1 always without
dorsal setulae, the anal crossvein generally more abruptly bent
near mid-length, and, in the female, tergites 4 and 5 unreduced.

Hennig (1940b) referred two Bama species to Xiria
Walker. The latter genus has the diagnostic characters of the
subfamily Trapherinae as stated by McAlpine (1973a), whereas
Bama is undoubtedly referable to the Platystomatinae. Xiria
is not otherwise recorded from east of Wallace’s Line, and
is readily distinguished from Bama in having the facial
convexity much more prominent and complex, antennal
segment 2 setulose on its medial surface, antennal segment
6 very long-bipectinate, the scutellum extensively haired
dorsally, vein 5 setulose dorsally, anal cell entirely
microtrichose, in the male the aedeagus without glans and
terminal filaments, and, in the female, abdominal tergites 4
and 5 unreduced and the aculeus broad, blade-like.

The species B. shinonagai is so different from the
majority of Bama species, that I initially intended to place
it in a separate genus. Though it perhaps constitutes a sister
group to the other known species, some of its unusual
features occur singly among other Bama species, and it has
the distinctive aedeagal features of Bama. I therefore place
it in a separate subgenus of Bama.

A series of an undescribed Bama species (Moro, Madang
Province, BM) was obtained by M.E. Bacchus (pers, comm.)
at a point where the nearly dry stream bed is closely roofed
over by trees. A newly eclosed specimen of B. shinonagai
(Myola, Oro Province, J.W.I., AM), is labelled “bamboo”.

The generic name is Greek for dye, in reference to the
usually conspicuous pigmentation of the wing. It is neuter.

Key to subgenera of Bama

1 Scutellum without setulae, dorsally subshining, with covering of
fine pubescence; antennal segment 3 rounded distally; squama
forming very short lobe, its area scarcely larger than that of axillary

lobe. Bama

-Scutellum setulose laterally, dorsally glossy, without pubescence;

antennal segment 3 distinctly angular dorsoapically; squama larger,

almost semicircular. Polimen

Subgenus Bama s.str.

Figs. 73-75

Diagnostic description. Scutellum with dorsal fine,
translucent pubescence, without setulae; intermediate
scutellar bristle not shorter than anterior scutellar; base of
vein 1 with fine dorsal setulae (except in one undescribed
species); squama forming a a very short lobe, its area not or
only slightly larger than that of axillary lobe. Female:
abdominal tergite 4 greatly reduced, usually more or less
concealed in dried material (less reduced, dorsally exposed,
c. one sixth as long as tergite 3 in one undescribed species).

This subgenus includes all known species of Bama ,
described and undescribed, except B. shinonagai, described
below.

Subgenus Polimen n.subgen.

Type species: Bama shinonagai n.sp.

Diagnostic description. Scutellum setulose dorsolaterally,
bare medially, glossy, without dorsal pubescence or
pruinescence; intermediate scutellar bristle much smaller
than anterior scutellar; base of vein 1 without dorsal setulae;
squama forming a well-developed lobe, almost as full as a

David K. McAlpine: Australasian Platystomatidae 167

Figures 73-75. Bama (Bama) “sp. A” (near Bulolo, AM). 73, wing. 74, head. 75, distal part of aedeagus (scale =
0.2 mm). Abbreviations: bu, bulb; dl, distal lobe; f, terminal filaments; g, glans; Is, terminal sclerotised lobe of
stipe; pg, preglans.

semicircle. Female: abdominal tergite 4 c. one fifth as long
as tergite 3, dorsally exposed; tergite 5 shorter.

Notes. The subgeneric name is a Latin noun meaning polish,
in reference to the glossy thorax, particularly the scutellum.
It is neuter.

Bama (Polimen) shinonagai n.sp.

Figs. 76-78

Material examined. Holotype, 8, Papua New Guinea:
Gumi near Bulolo, Morobe Province, 2010 m, 6.viii.l979,
H.R. (AM). Paratypes, Morobe Province: 1 9 , same data
as holotype (AM); 1 8, Mount Kaindi, near Wau, 1500-
2200 m, 19.viii.1965, J.S., M.S. (BPB); 4 $ $, Nami Creek,
near Wau, 10.vi.1962, 30.xii.1981, J.S., S.S. (AM, BPB,
NSMT). Eastern Highlands: 1 $, 22 km SE of Okapa, 2100
m, 28.viii.1964, J.S., M.S. (BPB). Western and Southern
Highlands: 1 9, Mount Wilhelm, 14.x. 1957, J.H.B. (AM);
1 $, Mount Giluwe, N side Malgi, 2500 m, 25-30.V.1961,
J.L.G. (BPB). Oro (Northern) Province: 1 8 ,2 9 9, Myola,
2080 m, 13.ix.1985, 28-3l.iii. 1986, J.W.I. (AM, BM).

Description (8 , 9). A stouter and somewhat smaller fly
than most Bama species. Coloration largely blackish,
with blue-tinted reflections. Postfrons largely dull brown-
black; upper half of face, parafacial, and an extensive
postorbital zone with greyish white pruinescence.
Antenna tawny, with variable brown suffusion; segment
6 largely brown. Prelabrum dark brown; palpus orange
tawny. Legs dark brown to blackish. Wing largely
hyaline; stigmatal section of subcostal cell dark brown;
a brown stripe along costa from just before end of vein 2
to a little behind end of vein 3, narrow near vein 2,
broadened where it crosses vein 3; base of first basal
cell with brown mark; a dark brown transverse stripe
covering anal and basal crossveins and extending forward
to vein 1 near level of end of subcosta; a brown stripe
covering anterior crossvein and, in female, usually
extending forward to about confluence of vein 1 with
costa, in male generally discontinuous in front of vein 3;
a pale or indistinct brown stripe covering discal
crossvein; squama pale creamy-translucent. Halter pale
brown basally, largely blackish brown distally. Abdomen
black; tergites 1-5 largely glossy, with bluish reflections.

168 Records of the Australian Museum (2001) Vol. 53

Figures 76-78. Bama (Polimen) shinonagai n.sp. 76, wing. 77, head. 78, distal part of aedeagus (scale = 0.2 mm).
Abbreviations: bu, bulb; f, terminal filaments; g, glans; gp, process of glans; pg, preglans; st, stipe.

Head 1.4-1.6 times as long as as wide; postfrons almost
parallel-sided, c. 1.6 times as long as wide; height of cheek
0.16-0.18 of height of eye. Antenna: segment 3 mucronate
or at least somewhat angular dorsoapically; segment 6,
except at apex, with numerous hairs, many of them as long
as its basal diameter, but not regularly seriate. Palpus of
moderate length, rather broad.

Thorax. Mesoscutum c. as long as wide; scutellum more
rounded and convex than in other species. Fore femur with
numerous non-seriate posterior to posterodorsal bristles and
long setulae and a series of large but not spinescent
posteroventral bristles; mid femur with hairs on postero-
ventral surface much longer than elsewhere. Wing: section
of costa on subcostal cell about as long as that on marginal
cell; first costal cell almost bare; second costal cell
microtrichose to variable extent, particularly so anteriorly
and distally; first posterior cell largely microtrichose, often
with narrow bare zones; submarginal cell with bare zone
just beyond base; first basal cell with a bare zone on each
side of sub-basal band; first posterior and discal cells entirely
microtrichose; second basal and anal cells largely bare,
except on pigmented areas; alula microtrichose, except on
an anterior zone of variable size; cell-4 index = 0.65-0.68;
second section of vein 4 with slight concave curvature; anal
crossvein with abrupt but not angular bend approximately

at anterior third; distal section of vein 6 with sigmoid
curvature.

Abdomen. Male: tergite 4 slightly shorter than tergite 3;
tergite 5 at least as long as tergite 3; sternite 5 transverse,
not medially cleft. Female: tergites as given for subgenus.
Male postabdomen: surstyli elongate; outer surstylus with
broadly rounded, setulose terminal section beyond apex of
inner surstylus; the two prensisetae of inner surstylus with
numerous parallel ridges; distal part of stipe of aedeagus
with two series of overlapping foliose lobes, each with free
margin minutely serrulate; preglans distinct, sclerotised,
with a smooth-margined membranous wing on its whole
length; glans stoutly ovoid-cylindrical, distally with a
membranous lobe; the two terminal filaments very unequal,
longer one slightly shorter than glans; cerci much shorter
than surstyli, joined for their whole length.

Dimensions. Total length, S S 4.8-5.4 mm, 9 $ 5.0-6.0
mm; length of thorax, 6 S 2.5-2.6 mm, 9 9 2.5-3.0 mm;
length of wing, S S 6.2-6.7 mm, 9 9 6.4-7.2 mm; length
of glans of aedeagus 0.23 mm.

Distribution. New Guinea: highlands of mainland Papua
New Guinea up to 2500 m (specimen labelled “Mount
Wilhelm” perhaps from significantly greater altitude).

David K. McAlpine: Australasian Platystomatidae 169

Notes. Bama (Polimen) shinonagai, the only known species
of its subgenus, is distinguished from other species by the
subgeneric characters given above. It is the only species of
Bama that runs to couplet 55 in the key to genera.

The specific epithet refers to Dr S. Shinonaga, who
collected material of this species.

Genus Laglaisia Bigot

Figs. 5, 79

Laglaisia Bigot, 1878a: 25. Type species (monotypy) L. caloptera

Bigot.

Description. See Hendel (1914a). An additional feature of
all species available to me is the presence of fine setulae
ventrally on the free margin of the scutellum, as in
Cleitamia. In the few species for which the aedeagus is
known, the preglans is simple, the tunic is elongate, and
the paired terminal filaments are moderately short.

Distribution. New Guinea: mainland only. These flies are
widely distributed and occur through a considerable range
of altitudes, but most seem to be narrowly localised and are
infrequent in collections.

Notes. The generic name was introduced as new several
times by Bigot with variable spelling. Evenhuis (1989) has
determined the earliest description and spelling as given
above, though I have previously used the spelling Laglaizia
(McAlpine, 1973a; 1982).

The key to species by Malloch (1939a) is an elaborated
translation of that of Hendel (1914b). Laglaisia includes six
described species, and I have seen c. nine additional species.

Males of Laglaisia generally have the head-capsule
laterally produced into eye-stalks, which are usually less
attenuated than in Fig. 5. An undescribed species (1 S , BPB)
is exceptional in lacking eye-stalks.

The habits and biology are almost unknown, but Parsons
(1984) records L. biroi Hendel as resting on the uppersides of
leaves of low-growing plants in well shaded primary forest,
and provides excellent colour photographs of the living flies.

Genus Cleitamoides Malloch

Figs. 80-82

Cleitamoides Malloch, 1939a: 106-107. Type species (original
designation) Cleitamia kerteszi Hendel.

Description. This genus was briefly characterised by
Malloch. Osten Sacken (1881a) and Hendel (1914a) did
not separate the group from Cleitamia , though Hendel noted
several points of disagreement of C. kerteszi and C. lituratus
(Walker) with other Cleitamia species. Because there are
numerous diagnostic differences between Cleitamoides and
Cleitamia not mentioned by Malloch, I indicate these in
Table 2.

Male postabdomen of type species (C. kerteszi ): outer
surstylus of moderate length, its free apical section
moderately slender, obtuse, slightly exceeding apex of inner
surstylus; aedeagus with rather long, glabrous stipe; preglans
complex, well sclerotised; glans moderately long, irregularly
subcylindrical, at apex with large, elongate, partly
sclerotised lobe sheathing a slender process; tunic well
sclerotised, with short, pubescent terminal lobe; terminal
filaments subequal, stout, distally tapered, each slightly
shorter than glans; cercus broad, apically emarginate,
extensively setulose on outer surface, with anteroapical
graduated comb of rather short to long setulae.

Distribution. New Guinea: mainland; Aru; Normanby Island.

Notes. Three species of Cleitamoides are listed by Malloch
(1939a) and by Evenhuis (1989). To these is now added
Cleitamoides trigonalis (de Meijere, 1913) n.comb. (from
Cleitamia ). This generic position is clear from the
description and comparison given by de Meijere. There
appear to be two additional probably undescribed species
in Papua New Guinea (Oro Province; Normanby Island;
both in AM), but a careful study of individual and geographic
variation and sexual dimorphism will be necessary to determine
more precisely the number of valid species.

Figure 79. Laglaisia “sp. 5” (Gumi, AM), wing.

170 Records of the Australian Museum (2001) Vol. 53

Table 2. Characters separating Cleitamia and Cleitamoides.

Cleitamia

Cleitamoides

• Fronto-orbital bristle present

•

Fronto-orbital bristle absent

• Humeral bristle present

•

Humeral bristle absent

• Mesopleural bristle present

•

Mesopleural bristle absent

• Scutellum with two or three pairs of
large bristles

•

Scutellum with one pair of

large and one of small bristles

• Scutellum with minute setulae below
free margin

•

Scutellum without setulae below
margin

• Membranous zone between hind coxae
and abdominal sternite 1 broad

•

Membranous zone between hind coxae
and sternite 1 narrow

• Vein 2 distally converging with vein 1, but
most closely approximated to vein 1
shortly before terminating in costa

•

Vein 2 most closely approximated to vein 1
where it meets costa (most males), or not
closely approximated to vein 1 distally (females)

• Anal cross vein strongly bent at or only
slightly in front of mid-length

•

Anal cross vein strongly bent at about anterior
third of length

• Male: basal part of costa not swollen

•

Male: basal part of costa inflated, obliterating
first costal cell

• Male: posterior margin of tergite 5 with
two pairs or more extensive fringe
of curved bristles, some at least half
as long as tergite

•

Male: posterior margin of tergite 5 with slightly
differentiated fringe of small bristles less
than a quarter as long as tergite

Figures 80-82. Cleitamoides kerteszi (Hendel). 80, distal part of aedeagus (scale = 0.2 mm). 81, head. 82, wing of
male. Abbreviations: dl, distal lobe; f, terminal filament; g, glans; gp, process of glans; pg, preglans; st, stipe; tu,
tunic. Fig. 82 after Hendel (1914a), retouched.

Genus Cleitamia Macquart

Figs. 83, 84

David K. McAlpine: Australasian Platystomatidae 171

Cleitamia Macquart, 1835: 440. Type species (monotypy) Ortalis
astrolabei Macquart (note that Boisduval’s publication of the
name O. astrolabei is slightly later than that of Macquart, see
Evenhuis, 1989).

Poticara Walker, 1861b: 248-249. Type species (monotypy) P.
triarcuata Walker (= Ortalis astrolabei Macquart).

Description. The description by Hendel (1914a) is largely
adequate, providing that the particular characters attributed
to Cleitamia liturata (Walker) and C. kerteszi Hendel are
excluded, as these are now placed in Cleitamoides. See also
Table 2 for some additional characters.

Male postabdomen. Aedeagus (where examined) with
glabrous stipe; preglans sharply differentiated from stipe,
without lobes; glans ovoid, with terminal lobe long or
vestigial; terminal filaments short, equal or subequal, with
basal tunic.

Distribution. New Guinea: widely distributed, but
apparently endemic.

Notes. Cleitamia includes more than 20 species, several of
which are undescribed. The last published key to species is
that of Malloch (1939a). Cleitamia trigonalis de Meijere
does not belong in this genus (see under Cleitamoides).
Cleitamia astrolabei (Fig. 83) is a widely distributed and
conspicuous insect of New Guinea forests.

Figure 83,84. 83 (above), Cleitamia astrolabei (Boisduval), male;
after Hendel (1914a), retouched. 84 (below), Cleitamia astrolabei,
distal part of aedeagus (scale = 0.2 mm).

Genus Loriomyia Kertesz

Fig. 85

Loriomyia Kertesz, 1899: 567. Type species (monotypy) L.

guttipennis Kertesz.

Loriomyia- Hendel, 1914a: 75-76.

Description (<5, 9 unknown). Hendel’s description is
excellent, and, as no material is available to me, I can only
add that the stem vein (base of vein 1) is bare dorsally (L.
Papp, in litt.).

Distribution. New Guinea: only known from Moroka
(Meroka), E of Sogeri, Central Province, Papua New
Guinea.

Notes. The type and only included species is, to my
knowledge, not represented in recent collections, and is only
known from the type material (MNM). This is surprising,
as the insect is of conspicuous appearance, and the Sogeri
district is one of the entomologically better known parts of
Papua New Guinea.

Figure 85. Loriomyia guttipennis Kertesz, wing. After Hendel (1914a).

172 Records of the Australian Museum (2001) Vol. 53

Despite the resemblance to certain tropical tephritids in
coloration and the contour of the anal crossvein, I think
that Hendel (1914a) was probably correct in placing
Loriomyia near Cleitamia in the Platystomatinae. Loriomyia
differs from Cleitamia in having two pairs instead of one
pair of fronto-orbital bristles, much reduced outer vertical
bristles, a pair of prescutellar acrostichal bristles, no setulae
on the scutellum, and the anal cell acutely produced. In
these features of thoracic vestiture, Loriomyia resembles
Bama, but differs from the latter in having only two pairs
of scutellar bristles, no posteroventral bristles on fore femur,
veins 1 and 2 approximated shortly before their distal ends,
and the anal cell acutely produced. In some undescribed
Bama species the anal crossvein has a somewhat sigmoid
course, which suggests an early stage in the development
of the condition present in Loriomyia. However, these Bama
species have the characteristic dorsal setulae on the stem
vein, which are absent in Loriomyia, and the venation is
not otherwise very like that of Loriomyia.

Loriomyia guttipennis should be readily distinguishable
from other Australasian platystomatids by the wing
markings (Fig. 85).

Genus Terzia n.gen.

Type species: Terzia saigusai n.sp.

Description (6, 9 unknown). Medium-sized, quite
elongate, rather long-legged flies.

Head slightly higher than long, with generally reduced
hairing; face concave in profile, without median carina;
lower end of facal ridge with few minute hairs; parafacial
narrow, without hairs; occipital region broadly, strongly
convex; the following bristles present: well-developed inner
vertical, one very small fronto-orbital, small postgenal; the
following bristles absent: outer vertical, postvertical, ocellar,
lateral occipital. Antenna, excluding arista, about as long
as face on median line; segment 2 short and subconical;
segment 3 narrowly ovate; segment 6 slender, with short
pubescence near base only. Prelabrum moderately large,
anteriorly convex; palpus moderately large.

Thorax deep and rather narrow, largely shining, with very
little pruinescence and reduced hairing; mesoscutum with
almost all hairs restricted to five longitudinal series; humeral
callus with one or two small hairs only; scutellum
moderately small, slightly convex dorsally, with few fine
hairs; postscutellum very deep; metathoracic postcoxal
bridge deep and well sclerotised; zone between metathoracic
spiracle and halter with pile-like pubescence and several
setulae other than those on margin of spiracle; the following
bristles present: pair of outstanding, approximated
paramedian scapulars, 1+1 notopleurals, postalar, large
apical and small sublateral scutellar, one ventral sterno-
pleural in front of mid coxa; other thoracic bristles absent.
Legs all quite long, fore one least so; femora slender, but
less so than in Angitula, with reduced hairing, except
towards apices. Wing elongate, slightly narrowed basally;
membrane bare on much of first costal cell, smaller basal

zone of submarginal cell, anal cell, alula; section of costa
on subcostal cell much shorter than that on first costal cell
or marginal cell; humeral break of costa very indistinct;
second basal cell more than half as long as discal cell,
towards its distal end wider than either first basal or anal
cell; vein 3 with scattered dorsal setulae, more crowded
basally; anterior crossvein meeting vein 4 slightly before
mid-length of discal cell; distal section of vein 4 arched,
converging with vein 3 from level of discal crossvein, but
slightly diverging from vein 3 apically; vein 5 without setulae;
axillary lobe undeveloped; squama vestigial, without lobe.

Abdomen slender, only slightly narrowed basally. See under
Terzia saigusai for further details.

Distribution. Solomon Archipelago: New Georgia.

Notes. Terzia is known to me from a unique male example
of the type species. This is unfortunate, as the taxon is of
unusual phylogenetic interest (Saigusa, unpublished note
in BPB cabinet).

My study of the general morphology of Terzia indicates
that it is a moderately derived taxon of the subfamily
Platystomatinae. If one were to eliminate the more obvious
apomorphies (discussed below), the resulting character
combination would be not very different from that of the
genus Rivellia, with, perhaps, Tarfa occupying an
intermediate stage. Though I have not examined the
aedeagus, the outer surstylus of Terzia, with its elongate,
subtriangular apical section, is remarkably like that found
in numerous platystomatine species (e.g., in the genera
Achias, Bama, Duomyia, Laglaisia, Plagiostenopterina,
etc.), but I have seen no similar condition in other
subfamilies.

Terzia is remarkable for the number of character states
that it shares with the “angituline” genus Angitula (now
including Angituloides and Girajfomyia), including absence
of the outer vertical bristle, great reduction of the hairing
of the humeral callus, mesoscutum, and mesopleuron, the
deep, glossy metathoracic postcoxal bridge, the virtual
absence of the axillary lobe, and the presence of both pile¬
like pubescence and setulae on the narrow sclerite between
the metathoracic spiracle and base of the halter. These
conditions I judge to be all or nearly all synapomorphies
indicating close relationship of Terzia to Angitula. However
it lacks certain of the peculiar features of Angitula, viz. the
very prolonged occipital region of the head (though it has a
more convex occiput than most platystomatine genera), the
loss of the normal anterior convexity of the mesoscutum,
the enlargement of the pronotum, the presence of a pair of
elongate setiferous horns on the scutellum, the ventral
prolongation of the membranous cleft in an almost straight
line, and the greater attenuation of the femora. This
constitutes a set of distinctive apomorphic character states
defining the clade formerly known as “Angitulinae” (e.g.,
Malloch, 1939b, Hennig, 1940a, McAlpine, 1973a).

From the above combinations of apomorphic character
states, I deduce that Terzia is the sister group of the genus
Angitula s.l. (including Angituloides and Girajfomyia). But
the general morphology of Terzia does not admit of its

David K. McAlpine: Australasian Platystomatidae 173

removal from from the Platystomatinae. Therefore the
“angituline” genera, including Terzia , are considered to form
a subordinate group within the subfamily Platystomatinae.

The generic name recalls A.J. Engel Terzi, formerly of
the British Museum (Natural History), whose illustrations
are a significant contribution to dipterology.

Terzia saigusai n.sp.

Figs. 86, 87

Material examined. Holotype, S (unique), Solomon
Archipelago: Munda, New Georgia Island, 1-30 m,
20.vii.1953, J.L.G. (BPB).

Description ($). Coloration. Head and thorax orange-
fulvous. Vertex with large dark brown semicircular median
blotch, immediately behind ocelli. Thorax without dark
markings. Legs fulvous; tarsi brownish tawny, slightly
darker distally. Wing hyaline, with uniform yellowish tinge;
distal section of subcostal cell yellowish brown; large brown
apical spot on costa extending from vein 2 to vein 4. Halter
fulvous with brown apical spot. Abdomen shining fulvous;
tergite 5 dark brown on about posterior two-thirds.

Head. Eye c. 0.71 as long as high; height of cheek c. 0.18
of that of eye.

Thorax. Fore femur with two stoutly spinescent postero-
ventral bristles on distal third; mid tibia with elongate apical
ventral spur c. 1.3 times as long as greater diameter of tibia.
Wing: anal cell on vein 4 c. 0.59 times as long as discal cell
on vein 4; cell-4 index = 0.47.

Abdomen. Syntergite 1+2 c. 0.47 times as long as tergite 3;
tergite 5 c. 1.2 times as long as tergite 3, slightly longer
than tergite 4. Outer surstylus considerably exceeding inner
surstylus in length, with elongate, subtriangular apical
section.

Dimensions. Total length 6.7 mm; length of thorax 2.3 mm;
length of wing 6.1 mm.

Distribution. As given for genus.

Notes. The specific epithet refers to Toyohei Saigusa, of
Fukuoka, Japan, whose percipient comment on relationships
of this species is noted above.

Genus Angitula Walker

Angitula Walker, 1859: 123. Type species (monotypy) A.
longicollis Walker.

Anguitula- OstenSacken, 1881a: 481^-83. Error or emendation—
5 usages.

Giraffomyia Sharp, 1899: 391-392. Type species (monotypy) G.
willeyi Sharp. N.syn., but retained as subgenus.

Angituloides Hendel, 1913: 345. Type species (original
designation) A. austeni Hendel. N.syn., but retained as
subgenus.

Angitulina Enderlein, 1936a: 227, 228. Type species (original
designation) Elaphomyiapolita Saunders (probably = Angitula
longicollis Walker).

Hammatopelma Enderlein, 1936a: 227,228. Type species (original
designation) H. nigra (sic) Enderlein.

Meachina Enderlein, 1936b: 241. Type species (original
designation) M. violacea Enderlein.

Description. Medium-sized to rather large elongate flies,

somewhat resembling micropezids, with glossy black thorax

and abdominal tergites, and reduced hairing and bristling.

Figure 87. Terzia saigusai, wing.

174 Records of the Australian Museum (2001) Vol. 53

Head generally longer than high, with pair of either
epistomal processes or cheek processes in males of some
species; eye rounded, but sometimes ventrally emarginate
in males; postfrons slightly excavated between summits of
eyes, extending only slightly in front of eyes in profile;
face in part concave in profile, without distinct median
carina, either slightly convex below without prominent
epistomal margin, or with epistomal margin prominently
protruded and, in some males, greatly expanded, or, in male
of A. nigra , with a long, spatulate lobe on each side; inner
vertical bristle large to quite small; outer vertical small or
minute but usually discernible; postgenal bristle minute or
absent; fronto-orbital and postvertical bristles absent;
supracervical setulae strongly developed, on a transverse
cervical tubercle. Antenna: segments 1 and 2 short; segment
3 c. two to three times as long as wide or longer, not
potentially attaining centre of epistomal margin; segment 6
bipectinate, with rather long rays, or short-haired, or partly
bare. Prelabrum moderately shallow, well separated from
epistomal margin by membranous zone; palpus variably
developed; proboscis stout, with broad labella.

Thorax pyriform, slender anteriorly, remarkably deep
posteriorly; prothorax, especially pronotum, together with
lateral cervical sclerites and associated membranous
surfaces forming a variably long neck, which holds head
well away from mesoscutum; pronotum otherwise variable
in structure and sometimes sexually dimorphic; prostemum
anteriorly truncate or variably narrowed, without precoxal
bridge; mesoscutum broad posteriorly, somewhat narrowed
anteriorly; scutellum with pair of elongate setiferous horns
(as in Diopsidae); subscutellum little developed; post-
scutellum deep, convex; anterior section of sternopleural
suture aligned (almost vertically) with membranous cleft
(as in Micropezidae-Eurybatini), posterior section
sometimes detached from anterior section; metathorax
produced behind the halteres dorsally and ventrally into a
short posterior cylinder, which supports the abdomen;
metathoracic postcoxal bridge very deep and strongly
sclerotised; metapleura and metasternum together forming
a prominence to which hind coxae are attached (as in
Megamerinidae, Syringogastridae, etc.); thoracic cuticle
almost devoid of pruinescence, but with densely pubescent
zone between metathoracic spiracle and halter; hairing of
thorax greatly reduced, but dorsocentral series of small hairs
present; scutellum usually with few fine hairs; thoracic
bristles all very small or undifferentiated from hairs, except
for moderate-sized bristle at apex of each scutellar horn.
Legs long and slender (fore coxa particularly so) without
bristles, except for long apicoventral spur on mid tibia; mid
femur with a series of variably developed suberect posterior
hairs, except in A. willeyi. Wing elongate, basally narrowed,
with few markings, mainly along costa; subcosta distally
straight, free from costa, strongly sclerotised; anterior

crossvein meeting vein 4 near mid-length of discal cell;
veins 3 and 4 apically subparallel, somewhat arched; second
basal and anal cells each much more than half as long as
discal cell; anal crossvein curved throughout, variable in
length; alula, axillary lobe, and squama virtually absent.

Abdomen elongate, sometimes clavate and anteriorly
petiolate; tergites largely or partly with reduced hairing.
Male: tergites 4 and 5 generally about as long as tergite 3;
outer surstylus simple, somewhat exceeding inner surstylus;
stipe distally with linear band of short or long pubescence
within longitudinal groove; glans short and stout, with one
or two pairs of membranous distal lobes; paired terminal
filaments subequal in length, varying from very short and
stumpy to much longer than glans (in A. nigra), arising from
sclerotised tunic. Female: tergites 4 and 5 either almost as
long as tergite 3, or much shortened; spiracles 4 and 5
situated close below lateral margins of their tergites; tergite
6 short but sclerotised; ovipositor sheath moderately long;
aculeus slender, obtuse.

Distribution. Moluccas. New Guinea. Bismarck Archipel¬
ago. Solomon Archipelago. Further details are given under
the subgenera.

Notes. Angitula, as here delimited, was divided into six
genera by Enderlein (1936a, 1936b), who included these
taxa in his subfamily Angitulinae of the Phytalmiidae.
Hennig (1940a), in dissolving the family Phytalmiidae,
referred the Angitulinae as a whole to the Platystomatidae.
He listed the species according to Enderlein’s generic
placements, but stated in his discussion of Angitulinae:
“There is therefore to be recognized only one genus,
Angitula Walk., with at most several subgenera” (my
translation). Malloch (1939b), Steyskal (1950, 1952a),
Me Alpine (1973a), and Evenhuis (1989) accepted three
angituline genera, Angitula, Angituloides, and Girajfomyia.
Korneyev (1994), in excluding “the Girajfomyia group”
from the Angitula alliance (as Angitulinae), was surely
misled into interpreting certain character states (given by
Me Alpine, 1973a) as relevant to higher classification. I here
treat these as three subgenera of the genus Angitula, on
account of: (1) the unambiguous monophyly of Angitula
s.l.; (2) the necessity of downgrading the subfamily
Angitulinae sensu Enderlein to infra-tribal status, taking
into consideration the apparent phylogenetic proximity of
the outgroup Terzia\ (3) the range of variation in other
polytypic platystomatid genera, including variation in aedeagal
and female abdominal structure (examples in Duomyia,
Chaetorivellia, Euprosopia, Lenophila, Rhytidortalis and
Senopterinaf, (4) the constant, easily recognized diagnostic
characters of Angitula s.l.; and (5) the distinctive general
appearance for the whole genus.

Key to subgenera of Angitula

1 Inner vertical bristle very small, or at least not much larger than
outer vertical; vein 2 meeting costa well beyond level of discal
crossvein; anterior extremity of abdominal tergite 1+2 with pair
of conical or thorn-like processes, each sclerotised all round.

Angitula

David K. McAlpine: Australasian Platystomatidae 175

Inner vertical bristle large, in contrast to vestigial or undiffer¬
entiated outer vertical; vein 2 meeting costa near or before level
of discal crossvein; anterior extremity of tergite 1+2 with pair of

low, rounded gibbosities, each open to thoraco-abdominal

membrane on anterior side. 2

2 Pronotum deeply bilobed, without ante-pronotum; mesoscutum
with anterior median finger-like process, bearing pair of apical

bristles and projecting between lobes of pronotum (Fig. 92). Angituloides

-Pronotum not very deeply bilobed, with, at least in male, distinct

ante-pronotum; mesoscutum without anterior median process. Giraffomyia

Subgenus Angitula s.str., n.stat.

Figs. 88-90

Angitula Walker, 1859: 123. Type species (monotypy) A.
longicollis Walker.

Angitulina, Hammatopelma, and Meachina are synonyms, as
detailed for the genus.

Description (principal diagnostic features only). Inner
vertical bristle very small, or at least not much longer than
outer vertical; antennal segment 3 usually c. three times as
long as wide or longer; segment 6 bipectinate, with longer
rays c. half as long as width of segment 3 or longer; palpus
rather long, reaching approximately to anterior margin of
prelabrum; pronotum moderately prolonged in front of

mesoscutum, its anterior margin generally rounded, entire,
and slightly curved upwards, without well defined ante-
pronotum; mesoscutum without anterior median process;
hind trochanter without ventral tubercle; vein 2 meeting
costa well beyond level of discal crossvein; submarginal
cell much wider than marginal cell at and beyond level of
anterior cross vein; second basal cell at broadest part about
as wide as anal cell; syntergite 1+2 at anterior extremity
with pair of slender, strongly raised tubercles; abdominal
spiracles 4 and 5 each with opening concealed by a tuft of
pubescence; male: glans of aedeagus (where known) without
terminal, sclerotised scoop-like lobe; paired terminal
filaments each more than half as long as glans; female:
tergite 6 usually quite short and, except in species with
tergite 5 reduced, concealed by tergite 5.

Figures 88-90. Angitula (Angitula) longicollis Walker. 88, wing. 89, distal part of aedeagus. 90, thorax. Abbreviations:
as, anterior section of sternopleural suture; cy, metathoracic cylinder; g, glans; gp, process of glans; lc, lateral
cervical sclerite; me, membranous cleft; peb, metathoracic postcoxal bridge; pg, preglans; pn, pronotum; pr,
postnotopleural ridge; ss, scutellar spine (paired); st, stipe.

176 Records of the Australian Museum (2001) Vol. 53

Distribution. Moluccas: Halmahera, Bacan (Batchian),
Seram. New Guinea: widely distributed on mainland,
Misool, Aru, Normanby Island.

Notes. The subgenus Angitula includes those species listed
by Evenhuis (1989) under the genus Angitula. Steyskal
(1952a) gave a key to the species that he could identify
from literature, but the species level taxonomy remains very
unclear. Material of A. (Angitula) longicollis Walker here
illustrated is identified from Steyskal’s key, not from
examination of type material. I am not yet convinced that
A. polita (Saunders) is a synonym of A. longicollis. If the
Bacan record of A. longicollis is due solely to Walker
(1861c), it is based on a misidentification. Material from
Bacan in AM and NMWC, probably identical with Walker’s,
belongs to a quite distinct species.

Secondary sexual modifications of the male head capsule
vary greatly in this subgenus. Some species, e.g., A.
longicollis, have little or no modification. Males of A.
krombeini Steyskal have the epistomal margin of the face
expanded into a broad plate. Those of A. nigra (Enderlein)
have the epistomal margin produced into a pair of large,
spatulate processes (erroneously referred to as cheek
appendages). The only recorded male of A. polita (Saunders)
has a pair of very small cheek processes.

Subgenus Angituloides Hendel n.stat.

Figs. 92, 93

Angituloides Hendel, 1913: 345 (as genus). Type species (original
designation) A. austeni Hendel.

Description. Inner vertical bristle large; outer vertical
vestigial; antennal segment 3 c. twice as long as wide;
segment 6 irregularly short-haired dorsally, mainly on basal
third, almost bare ventrally; palpus shorter than in Angitula
s.str., less reduced than in Giraffomyia\ pronotum very
deeply divided into two subtriangular lobes, without ante-
pronotum; mesoscutum with median finger-like process
bearing pair of apical bristles, and projecting anteriorly
between lobes of pronotum; hind trochanter with tooth-like
ventral process in male, present as vestige in female; vein 2
meeting costa slightly before level of discal crossvein;
submarginal cell not wider than marginal cell near level of
anterior crossvein; second basal cell at broadest part
narrower than anal cell; syntergite 1+2 at anterior extremity
with pair of low, rounded gibbosities; abdominal spiracles
4 and 5 each with almost glabrous sclerotised ring; male:
stipe of aedeagus with very long, brush-like pubescence;
glans with sclerotised scoop-like distal lobe; paired terminal

Figures 91-93. 91 , Angitula (Giraffomyia) willeyi (Sharp), distal part of aedeagus (scale = 0.2 mm). 92, Angitula
(Angituloides) austeni (Hendel), anterior part of thorax, dorsolateral view; 93, distal part of aedeagus (scale = 0.2
mm). Abbreviations: amp, anterior median process of mesoscutum; bu, bulb; dl, distal lobe of glans; f, terminal
filament (paired); g, glans; gp, processes of glans; pg, preglans; pn, lobes of pronotum; st, stipe.

David K. McAlpine: Australasian Platystomatidae 111

filaments quite small, partly concealed by lobe of glans;
female: tergites 4 and 5 large; tergite 6 quite short but
apparently permanently exposed.

Distribution. Solomon Archipelago: Bougainville; New
Georgia; Santa Isabel; Malaita; Guadalcanal; San Cristoval
(Makira). There is a likelihood that populations also live
on other islands of this Archipelago.

Notes. Only one species of Angituloides, Angitula
(Angituloides) austeni (Hendel) n.comb., has been
recognized, despite the wide distribution of the subgenus.
Hendel’s holotype was without adequate localisation
(“Salomons-Inseln”). At present I have before me only
material from Bougainville (3, $) and Guadalcanal (3
$ $). These specimens agree closely, the only characters
which appear likely to differentiate these insular populations
being the black orbital stripe and fine occipital spotting of
the Bougainville specimens, which are absent in those from
Guadalcanal. A further study of geographic variation is
desirable, but, on present data, such variation seems likely
to be infraspecific.

The few recorded males show no modifications of the
head capsule, but the hind-trochanteral process is distinctive.

Subgenus Giraffomyia Sharp n.stat.

Figs. 6, 91

Giraffomyia Sharp, 1899: 391-392 (as genus). Type species

(monotypy) G. willeyi Sharp.

Description. Inner vertical bristle large; outer vertical bristle
vestigial; antennal segment 3 c. twice as long as wide or
somewhat less; segment 6 irregularly short-haired on almost
entire length; palpus very small; thorax generally more
elongate than in other subgenera; pronotum always
prominently developed, usually longer in large males than
in females, not deeply divided as in Angituloides, generally
with freely projecting anterior part differentiated as ante-
pronotum (Speight, 1969), less distinct in some females;
mesoscutum without trace of median anterior process and
associated bristles; hind trochanter without ventral process
(in available material); vein 2 meeting costa near or slightly
before level of discal crossvein; submarginal cell not or
only slightly wider than marginal cell near level of anterior
crossvein; second basal cell at broadest part narrower than
anal cell; syntergite 1+2 at anterior extremity with at most
pair of low, rounded gibbosities; abdominal spiracles 4 and
5 each with almost glabrous sclerotised ring (in available
material); stipe of aedeagus with band of minute pubescence
only; glans with sclerotised scoop-like distal lobe; paired
terminal filaments very short and stout; female: tergites 4
and 5 large; tergite 6 quite short but apparently permanently
exposed.

Distribution. Bismarck Archipelago: Manus; Los Negros;
New Ireland; New Britain. Solomon Archipelago: Vella
Lavella; Malaita (or Mala); Florida; Guadalcanal; Ugi (Uki);
Ulawa.

Notes. New combinations for nominal species resulting
from reduction of Giraffomyia to subgeneric status are:
Angitula (Giraffomyia) nigripes (Steyskal), Angitula
(Giraffomyia) regularis (Malloch), Angitula (Giraffomyia)
irregularis (Malloch), Angitula (Giraffomyia) solomonensis
(Malloch) (the above all from Solomon Archipelago), and
Angitula (Giraffomyia) willeyi (Sharp) (from New Britain,
Bismarck Archipelago).

I have on hand very little material from the Solomons
and the above is a largely non-critical list. However, I do
not think that Evenhuis (1989) is likely to be correct in
raising Giraffomyia regularis var. perfecta Malloch to
specific status. The variation in size of the male cheek
processes described by Malloch in A. (G.) regularis is to be
expected among individuals of the one species population,
and is also seen in the large series of A. (G.) willeyi in AM.
Also Evenhuis’s synonymy of A. (G.) irregularis under A.
(G.) solomonensis, perhaps on the basis of an ambiguous
comment by Steyskal (1950: 96), seems unwarranted
without further evidence, the two names applying to separate
insular populations.

The numerous specimens of subgenus Giraffomyia
available from the Bismarck Archipelago appear to represent
at least four species, of which only A. (G.) willeyi is yet
described.

Elongate cheek processes of various structure are
present in larger males of perhaps all species of this
subgenus (e.g., Fig. 6).

Subfamily Scholastinae

The subfamily has been characterised by McAlpine
(1973a). While there remains an element of doubt as to
whether the group is monophyletic, its distinction from
the Platystomatinae is unambiguous.

Because the genera Asyntona and Chaetorivellia include
both species with female tergite 5 much reduced and species
with tergite 5 well developed, it is possible that the reduced
condition is not in the groundplan of the subfamily. In
Lenophila, female tergite 6 is small but by no means
vestigial, so it appears that either absence of tergite 6 is not
in the groundplan of the subfamily, or that Lenophila is
misplaced here.

The Australasian genera included at present are: Lenophila,
Chaetorivellia, Tomeus, Pterogenia, Neohemigaster,
Paryphodes, Scholastes, Trigonosoma, Zygaenula,
Mesoctenia, Naupoda, Asyntona.

Genus Lenophila Guerin-Meneville

Figs. 94-96

Lenophila Guerin-Meneville, 1843: 200; McAlpine & Kim, 1977:

314-314. Type species (monotypy) Ceratitis dentipes Guerin-

Meneville = Ortalis dentipes Macquart.

Celetor Loew, 1873: 4; Hendel, 1914a: 113-115. Type species

Ortalis dentipes Macquart = Tephritis strigipennis Macquart

(designated Hendel, 1914a).

Description. Hendel (1914a) gave a detailed description,
which I supplement as follows.

178 Records of the Australian Museum (2001) Vol. 53

Figures 94-95. Lenophila secta McAlpine & Kim. 94, wing. 95, aedeagus (scale = 0.25 mm). Abbreviations: g,
glans; pg, preglans; ps, distal process of stipe; st, stipe. Illustrations by S.R Kim.

Figure 96. Lenophila coerulea (Macquart), antenna. al-a6,
antennal segments one to six.

Legs. Hind tibiae with male-restricted modifications of form
and vestiture which differ in each species (McAlpine & Kim,
1977: figs. 9-14).

Male postabdomen. Aedeagus: stipe with terminal appendage
tapering to a fine point; preglans distinct; glans large, with
complex sclerotisation, without terminal filaments.

Female postabdomen. Tergites 5 and 6, often also tergite 4,
variably reduced; ovipositor sheath (tergostemite 7) moderately
long; aculeus slender, not depressed and blade like.

Distribution. Australia: apparently endemic.

The species are widely distributed in less arid parts of
temperate Australia, including all states, but apparently are
not present in Northern Territory. Three species are of
limited occurrence in tropical Queensland. Lenophila
achilles McAlpine & Kim extends to southern Tasmania
(Bruny Island, AM).

Notes. The genus, which includes six known species, was
reviewed by McAlpine & Kim (1977).

The bold pattern of the wing strongly resembles that of
certain tephritid genera, particularly Ceratitella Malloch
(see Hardy, 1967), and also the otitid Dyscrasis Aldrich (see
Hernandez-Ortiz, 1988). This pattern, which is usually
found in combination with a convex, glossy, black
scutellum, also occurs in two undescribed species of
Atopognathus. The common features of these flies could
be due to mimicry of jumping spiders (Salticidae), an idea
first mentioned to me by G. Daniels (see also Mather &
Roitberg, 1987).

Adults of five of the species are almost invariably found
on leaves of Xanthorrhoea spp. (Xanthorrhoeaceae) and
for some it has been demonstrated that larvae live in the
damaged or rotting trunk of this plant. On the other hand L.
dentipes (Macquart) is not found on Xanthorrhoea and there
is evidence of association with Eucalyptus sp. (Myrtaceae)
and Macrozamia sp. (Zamiaceae). C.E. Chadwick (pers.
comm.) reared numerous adults of L. dentipes from larvae
living in the caudex of Macrozamia communis damaged by
larvae of the weevil Melanotranes internatus (Pascoe)
(Curculionidae).

David K. McAlpine: Australasian Platystomatidae 179

Genus Chaetorivellia de Meijere

Fig. 97

Chaetorivellia de Meijere, 1913: 376; Hendel, 1914a: 128-129.
Type species (monotypy) Ortalis punctifascia Walker.

Description. Hendel (1914a) keyed the genus and gave a
detailed description, which I supplement as follows:

Male postabdomen. Aedeagus: stipe without terminal
appendage; preglans distinct; glans not forming a simple
capsule, but with complex sclerotisation and no terminal
filaments.

Female postabdomen of two markedly different types (a
and b). (a): tergite 5 large, at least as long as tergite 4; aculeus
slender, not blade-like, (b): tergite 5 much reduced, more
or less concealed in dried specimens; aculeus broad, blade¬
like. (a) and (b): aculeus with pair of small, terminal scale¬
like processes.

Distribution. Moluccas: Ternate; Halmahera (Gilolo);
Bacan (Batchian); Ambon (Amboina). New Guinea: widely
distributed on mainland. Solomon Archipelago: Bougain¬
ville. There are no records for Australia and the Bismarck
Archipelago.

Notes. Hendel (1914a) gives Ortalis trifasciata Doleschall
(described from Amboina) as the type species of Chaetorivellia,
but C. punctifascia (Walker) (described from Gilolo) is the
only nominal species originally included by de Meijere.
Hendel considered these two nominal species as synonyms,
but I do not consider the synonymy to be demonstrated as
yet. The nominal species Ortalis tarsalis Walker, 1861c is
also referred to Chaetorivellia (n.comb.) from examination
of holotype (Batchian or Bacan, A.R.W., OX).

There are probably at least four distinct species of
Chaetorivellia in collections, but species-level taxonomy
in the genus needs further study. The great variation in
female abdominal morphology is remarkable, but there is
comparable variation in some larger genera of the family.

Genus Tomeus n.gen.

Type species: Tomeus wyliei n.sp.

Description (<?, $). Stout flies, conforming to the
characterisation of the Scholastinae (McAlpine, 1973a: 26)
except in features of the aculeus. Cuticle of head, thorax
and abdomen largely dull and densely pruinescent.

Head much higher than long in profile, slightly wider than
high; eye much higher than long; face slightly concave near
centre, slightly convex below; parafacial on lower half with
numerous non-seriate hairs; the following bristles present: outer
vertical, quite small inner vertical, two fronto-orbitals,
postgenal; postvertical bristle absent. Antenna, excluding arista,
about half as long as face; segment 6 with non-seriate hairs on
whole length. Prelabrum small; palpus of moderate size.

Thorax. Mesopleuron without strong gibbosity on upper
part; scutellum rounded in outline, longer than a semicircle,
somewhat depressed, extensively haired on dorsal surface
and much of lateral surface; subscutellum convex, but small;
the following thoracic bristles present: humeral, 1+1 or 0+1
notopleurals, small supra-alar, large postalar, posterior intra-
alar, sometimes prescutellar acrostichal, two (posterior) pairs
of scutellars, mesopleural. Legs moderately short and stout;
femora not notably thickened; fore and hind femora with a
few weak dorsal bristles, other femoral bristles undiffer¬
entiated; mid tibia with rather short, stout terminal spur.
Wing essentially similar to that of Scholastes; vein 1 without
setulae before level of humeral crossvein; vein 5 without
setulae dorsally; all veins without ventral setulae; squama
moderately small, almost semicircular, its area not greater
than well-developed axillary lobe.

Abdomen ovoid, broadest across segment 2. Male
postabdomen: tergite 5 c. as long as tergite 4. Female
postabdomen: tergite 4 large; tergite 5 much reduced; tergite
6 apparently reduced to pair of minute plates; aculeus broad,
strongly depressed, obtuse.

Distribution. New Guinea: Papua New Guinea—mainland.

Figure 97. Chaetorivellia sp. (Laloki, Papua New Guinea, AM), wing.

180 Records of the Australian Museum (2001) Vol. 53

Notes. Tomeus belongs in the subfamily Scholastinae and
is most similar to Neohemigaster, Pterogenia, and
Scholastes. It differs from all these in its smaller squama
and broad, flattened ovipositor complex. It further differs
from Neohemigaster in the bare vein 5, more strongly
curved anal crossvein, and large, exposed tergite 4 of the
female abdomen. It differs from Pterogenia (at least the
Australasian species) in the short, non seriate hairing of the
arista, bare base of vein 1 before humeral crossvein, bare
vein 5, and more curved anal crossvein. It also differs from
Scholastes in the arista as above, in the absence of
dorsocentral, sternopleural, and anterolateral scutellar
bristles, the undifferentiated posteroventral bristles of the
fore femur, the apically desclerotised subcosta, and the large,
exposed tergite 4 of the female abdomen. In general
appearance Tomeus is reminiscent of some species of the
mainly African genus Paryphodes, but is distinguishable
from the only known Australasian species by the absence
of the sternopleural and anterior scutellar bristles, the simple
epistomal margin of face, the much less prominent
subscutellum, the non-incrassate hind femur, the simpler
contour of the posterior crossvein, the absence of setulae
on vein 5, etc.

The generic name is from the Greek tomeus, a cutting
instrument, in reference to the aculeus, and is masculine.

Tomeus wyliei n.sp.

Figs. 98-103

Material examined. Holotype, 6 , Papua New Guinea: Stony
Logging Area, near Bulolo, Morobe Province, 765 m,
17.vii.1979, H.R. (AM). Paratypes, Papua New Guinea: 1
8 , same data as holotype (AM); 1 9, Upper Manki Logging
Area, near Bulolo, Morobe Province, 5000 ft (c. 1500 m),
iii.1973, F.R.W., PS. (AM); 1 9, Brown River, near Port
Moresby, Central Province, x.1960, J.L.G., (on) palm (BPB).

Description (<J, 9). Coloration. Head largely ochraceous;
ocellar spot black; postfrons and face with brown markings
as in Fig. 98; occiput with brown suffusion. Antenna
ochraceous, with brown suffusion on segments 2 and 3.
Prelabrum and palpus brown. Thorax predominantly dull
blackish brown; mesoscutum with dark to mid-grey
pruinescence forming three very diffuse longitudinal bands;
scutellum with broad U-shaped yellow band; humeral callus
with yellowish zone entending as a band along upper margin
of mesopleuron; lower part of mesopleuron with slight
tawny suffusion. Fore and mid coxae predominantly brown;
hind coxa predominantly ochraceous; femora ochraceous
with brown suffusion most strongly developed on distal part
of ventral surface; tibiae pale ochraceous with brownish
sub-basal zone and usually trace of preapical zone; tarsi
pale yellow. Wing with brown markings and suffusion as
in Fig. 100. Halter pale yellow. Abdominal tergites 1 to 4
dull tawny with variable median and lateral marginal brown
suffusion; tergite 5 of male brown laterally, tawny medially;
ovipositor sheath brown to tawny.

Head as given for genus; height of cheek 0.15-0.19 of height
of eye.

Thorax. Mesoscutum c. 0.86 times as long as wide, very
densely pruinescent-pubescent and extensively setulose.
Wing: entire membrane microtrichose; length of second
basal cell c. 0.72 of length of discal cell.

Abdomen. Male postabdomen: outer surstylus broad, simple,
ensheathing inner surstylus, with few fine setulae; inner
surstylus shorter, with two large prensisetae; stipe of
aedeagus broad, ribbon-like; preglans almost undifferen¬
tiated; glans with large rounded concave lobe (not visible
in Fig. 103), and solid, sclerotised apical horn-like process;
cavity of glans with lining partly densely papillose-rugose;
cercus with numerous long, non-seriate setulae.

Dimensions. Total length, 8 5.7 mm, 9 9 5.1-6.3 mm;
length of thorax, 8 8 2.5-2.6 mm, 9 9 2.4-2.9 mm; length
of wing, 8 5.5 mm, 9 9 5.4-6.0 mm; length of glans
(excluding entire apical process) 0.23 mm.

Distribution. As given for genus.

Notes. The specimens from Stony Logging Area were found
on leaves of Musa (native banana).

The specific epithet refers to F. Ross Wylie, who collected
material of this and other interesting Diptera in Papua New
Guinea.

Genus Pterogenia Bigot
Figs. 104-106

Pterogenia Bigot, 1859: 312. Type species (designated Hendel,
1914a: 20) P. singularis Bigot.

Agastrodes Bigot, 1859: 311. Type species (monotypy) A.
niveitarsis Bigot.

Elachigaster Rondani, 1875: 431-432. Type species (monotypy)
E. albitarsis Rondani (= Agastrodes niveitarsis Bigot).
Ditomogaster Rondani, 1875: 431,433. Type species (monotypy)
D. xanthomera Rondani (= Pterogenia dayak Bigot).

Description. The detailed description by Hendel (1914a:
141-144) remains valid, so long as the particular characters
now attributed to Neohemigaster spp. are omitted. I
supplement Hendel’s description as follows.

Head. Area between antennal sockets forming a narrow
ridge; face generally without rounded gibbosity at each
lateroventral extremity, at least in females; inner vertical
and fronto-orbital bristles generally absent. Antenna
generally reaching not much more than halfway from basal
socket to centre of epistomal margin of face. Prelabrum
generally attenuated anteriorly, sometimes joined to face
by small median sclerite; palpus usually well developed,
sometimes very broad, in one species minute.

Thorax. Prosternum usually larger and less truncated
anteriorly than in Neohemigaster, subscutellum large,
usually markedly less convex than in Neohemigaster. Wing:
vein 1 always with group of dorsal setulae on basal part,
before humeral crossvein (as in Euprosopia ).

Abdomen. Tergite 2 variable, but usually without median

David K. McAlpine: Australasian Platystomatidae

Figures 98-103. Tomeus wyliei n.sp.. 98-99, head. 100, wing. 101, female postabdomen, dorsal view. 102, epandrium
and associated structures, posterolateral view. 103, distal part of aedeagus (scale for Figs. 102, 103 = 0.2 mm).
Abbreviations: ac, aculeus; c, cerci; g, glans; gp, terminal process of glans; is, inner surstylus; o, ovipositor sheath
(tergosternite 7); os, outer surstylus; seg6, seg8, abdominal segments six and eight; st, stipe.

182 Records of the Australian Museum (2001) Vol. 53

Figures 104-106. 104, Pterogenia nubecula Hendel, wing; 105, Pterogenia “sp. A” (Mulgrave River, AM), head;
illustrations by S.R Kim. 106, Pterogeniapectoralis Hendel, distal part of aedeagus (scale = 0.1 mm). Abbreviations:
g, glans; gp, terminal process of glans; pg, preglans; st, stipe.

tubercle on posterior margin. Female: tergite 4 usually well
sclerotised, but shorter than tergite 3; tergite 5 vestigial;
ovipositor sheath generally short, well sclerotised,
subcircular in cross-section; aculeus attenuated.

Distribution. Australasian Region: Moluccas; New Guinea;
Bismarck Archipelago (New Britain, AM); Solomon
Archipelago (Bougainville, AM); Australia (Queensland,
New South Wales). Oriental Region (widely distributed).

Notes. Numerous Australasian species are listed by
Evenhuis (1989) and Oriental species by Steyskal (1977).
Some of these are now transferred to Neohemigaster (q.v.),
and the generic position of others remains to be determined.

The keys to species of Hendel (1914b) and Malloch
(1939a) are very incomplete. Frey (1930) gave a key to the
Philippine species.

Pterogenia is differentiated from other Australasian
genera of Scholastinae by the presence of setulae on the
stem vein (base of vein 1 before level of humeral crossvein).
It most resembles Neohemigaster , q.v. for comparative data.

Adults of several Australian species are found on rotting
logs or tree stumps infested with boring Coleoptera in rain
forest. These may be oviposition sites.

Genus Neohemigaster Malloch
Fig. 107

Hemigaster Rondani, 1875: 431. Type species (monotypy) H.

albovittatus Rondani. Preoccupied Brulle, 1846 (Hymenoptera).
Neohemigaster Malloch, 1939a: 100, 126-127. New name for
Hemigaster Rondani. Type species (automatic) H. albovittatus
Rondani.

Description. Resembling Pterogenia , as redescribed by
Hendel (1914a). The following are the most notable features:

Head somewhat anteroposteriorly compressed, largely
pruinescent and non-shining; facial ridge with few setulae
at lower end; area between antennal sockets flat or slightly
concave, margined on each side by a slight ridge which is
decurrent on to face for short distance; face without median
carina, but slightly convex medially on upper part, with a
rounded gibbosity at each lateroventral extremity; outer
vertical bristle well developed; inner vertical present or
absent; postvertical and fronto-orbital bristles absent.
Antenna extending c. halfway from basal socket to centre
of epistomal margin of face; arista usually subplumose for

David K. McAlpine: Australasian Platystomatidae 183

Figure 107. Neohemigaster “sp. 1” (Awala, Papua New Guinea, AM), wing.

most of length. Prelabrum small, but well sclerotised; palpus
well developed, usually rather broad.

Thorax largely dull, pruinescent; prosternum usually very
short and broad; subscutellum usually large and very
convex; the following bristles generally present: humeral,
1+1 notopleurals, supra-alar, postalar, posterior intra-alar,
usually three pairs of unequal scutellars situated on posterior
part of scutellum, mesopleural; dorsocentral small or absent;
prescutellar acrostichal bristle absent. Legs: fore femur with
short or rudimentary dorsal and posteroventral bristles.
Wing with very diverse brown markings; vein 1 with dorsal
setulae only beyond level of humeral crossvein, bare basally;
vein 5 setulose dorsally on basal section, sometimes also
setulose more distally; squama larger than axillary lobe.

Abdomen short and rather broad; posterior margin of tergite
2 with median raised tubercle. Male: tergites 4 and 5 well
developed. Female: tergite 3 very large; tergites 4 and 5
minute or desclerotised; ovipositor sheath depressed, keeled
along each side; aculeus very slender, not depressed.

Distribution. Australasian Region: New Guinea. Oriental
Region: widely distributed in Indonesia and Malaysia,
possibly also Philippines and Taiwan.

Notes. Species of this genus were included in Pterogenia
in all the writings of de Meijere and Hendel, and characters
for separating these genera were first given by Malloch
(1939a). It has therefore been difficult to place some of the
described species without examination of types.

In my experience, species of Neohemigaster can be
separated from those of Pterogenia by the absence of dorsal
setulae on the basal part of vein 1 (stem vein), the flattened
or slightly concave zone between the antennal sockets, the
small rounded gibbosity, present even in females, at each
lateroventral extremity of face, the great reduction of
abdominal tergite 4 in the female, and the depressed,
laterally keeled ovipositor sheath. The greater convexity of
the subscutellum in Neohemigaster may be a valid
difference, but needs checking for more species. The
presence of the inner vertical bristle in Neohemigaster and
the absence of the median tubercle on abdominal tergite 2
in Pterogenia are not consistent enough for generic

segregation. In the male of an undescribed species of
Neohemigaster (“sp.l” in AM) from Papua New Guinea the
postgenal region on each side bears a stout, forwardly directed
lamella, reminiscent of that of Pterogenia singularis Bigot.

The type species, Neohemigaster albovittata (Rondani),
was redescribed from the holotype female by Hendel
(1914b: 322-324) and the head and wing figured (1914a:
pi. 12, fig. 231; pi. 13, fig. 232). Originally from Sarawak,
East Malaysia, Malloch (1939a: 127) recorded a male of N.
albovittata from Sandakan, Sabah. My knowledge of this
species derives from the publications of Hendel and Malloch.

In addition, I have seen the following species referable
to Neohemigaster. Neohemigaster fascifrons (de Meijere,
1916) n.comb., Java (AMST); Neohemigaster guttata
(Walker, 1856) n.comb., as determined and redescribed by
de Meijere (1916: 85-86), Singapore and Sumatra (AMST);
Neohemigaster “sp.l”, Papua New Guinea (Morobe, Oro,
and Central Provinces, AM); Neohemigaster “sp.2”,
Sumatra (AM). Malloch (1939a: 127) mentioned two
unnamed species of Neohemigaster from Sibuyan Island,
Philippines. From the original descriptions, it seems
probable that Pterogenia eurysterna Hendel (1914b: 319—
321) from Taiwan, and P. glabrina Hendel (1914b: 318—
319) from Sarawak are referable to Neohemigaster. Thus it
appears that as many as nine species of Neohemigaster exist
in collections. The described Oriental species are listed by
Steyskal (1977) under Neohemigaster and Pterogenia.

I have collected females of Neohemigaster “sp.l” in
a rubber plantation in Papua New Guinea on trunks of
Hevea brasiliensis, at wounds probably made by boring
Coleoptera. H. Roberts found both sexes of the same
species in a natural rainforest habitat, the specimens
being labelled “ex Xanthophyllum” (AM).

Genus Paryphodes Speiser

Paryphodes Speiser, 1911: 251. Type species (original designation)
P. omega Speiser.

Systellodiscus Enderlein, 1912: 372. Type species (original
designation) S. perforatus Enderlein.

Simomesia Enderlein, 1912: 373. Type species (original
designation) Simomesia tigrina Enderlein (= Scholastes
nepticula Loew).

Description. See Hendel, 1914a.

184 Records of the Australian Museum (2001) Vol. 53

Distribution. Australasian Region: New Guinea—north¬
east, new record for Region. Afrotropical Region: mainland
Africa—mainly tropical; Madagascar (specimen of a typical
Paryphodes in NMWC; a further species doubtfully referred
here in AM).

Notes. Hendel (1914b) gave a key to species, and Steyskal
(1980) listed the species with additions and revised
synonymy. The latter recognized 17 species, all from Africa.

Judging from the numerous Afrotropical specimens that
I have examined, the only New Guinea species is reasonably
typical of the genus. In addition to characters given in the
key to genera, it differs from the species of Scholastes, which
have a similarly developed sternopleural bristle, in having
the postfrons c. twice as long as wide (much less than twice
as long in Scholastes ), and without the anterior transverse
ridge present in Scholastes", two fronto-orbital bristles
(instead of one); antennal segment 6 with numerous, quite
short, non-seriate hairs on most of length (instead of being
relatively sparsely long-bipectinate); dorsocentral bristles
vestigial (instead of in one or more well-developed pairs);
fore femur with short posteroventral bristles, scarcely longer
than dorsal bristles (instead of much longer than any dorsal
bristles); hind femur much thicker than mid femur (instead
of scarcely thicker); vein 2 not undulated (instead of strongly
undulated near mid-length); basal section of vein 5 dorsally
setulose (instead of bare); anal crossvein nearly straight
(instead of distinctly curved); tergite 5 of male as long as
tergites 3 and 4 together (instead of no longer than tergite 3).

The record for Papua New Guinea is unexpected in the
absence of any records from the Oriental Region. As the
species are relatively small and inconspicuous for
platystomatids, it is possible that they have been overlooked
in some countries. As far as I can judge, the species from
New Guinea is distinct from any of the described African
ones. The possibility that it may be introduced by man is
acknowledged.

Paryphodes hospes n.sp.

Fig. 108-110

Material examined. Holotype, 6 (unique), Papua New
Guinea: Bainyik, near Maprik, East Sepik Province,
21.xii.1963, D.K.M. (AM).

Description (6, 9 unknown). Coloration. Head dull
fulvous; postfrons with pair of parallel brown stripes clear
of orbits on posterior two-fifths, dilated and connected
posteriorly but not quite reaching vertex; a pair of less
distinct brownish marks at anterior third of postfrons; face
with brown mark near centre, not extending its full width;
occiput largely dark brown. Antenna fulvous, with slight
brownish suffusions; segment 6 brown except at fulvous
base. Palpus and prelabrum dull fulvous. Thorax dark
brown, largely pruinescent and little shining; humeral callus
with two yellow marks, upper one smaller and a little
extended posteriorly above anterior notopleural bristle,
lower one extending over upper part of mesopleuron;
mesoscutum with very indistinct pair of grey dorsocentral

stripes; scutellum with dull yellow arcuate marginal zone
not extending on to anterior half. Femora brownish tawny,
fore and mid ones a little darker except towards apices; fore
tibia yellow, with tawny suffusions; mid tibia tawny, with
brownish band beyond middle and yellow apex; hind tibia
largely yellowish, with tawny base; tarsi pale yellow,
without apical darkening. Wing hyaline or subhyaline except
as follows: a pale uneven brownish suffusion in both costal
cells; stigmatal section of subcostal cell yellowish brown,
slightly paler distally; marginal cell with basal streak, a small
brown blotch at base of vein 2, two larger brown blotches
near middle, and a small brownish blotch near distal end;
submarginal cell with two brown blotches adjoining large
brown blotches in marginal cell, a less intense brownish
mark adjoining distal blotch in marginal cell and an
indistinct brown mark between last and apex; first basal
cell tinged with brown on basal half, distally with two brown
blotches, one of which is terminal; first posterior cell
unevenly suffused with brown on basal third, with pair of
brownish blotches about halfway between dark basal zone
and apex and a faintly indicated blotch between this pair
and apex; second basal and anal cells with faint suffusion
only, but the latter with brown streak along vein 5; discal
cell with brownish blotch at about basal quarter and
extensive irregular distal brown suffusion which is clear of
vein 5 except near discal crossvein; second posterior cell
suffused with brown mainly near discal crossvein and with
sub-basal blotch near vein 4, with very indistinct suffusion
distally; third posterior cell with faint general suffusion and
an indistinct blotch beyond middle near vein 5; axillary lobe
and squama dull creamy. Halter yellowish, with brown
capitellum. Abdominal tergites largely thinly pruinescent
and somewhat shining, tawny to brownish tawny; pleural
membrane fulvous.

Head. Postfrons evenly convex in profile, its length
(antennal sockets to vertex) about twice as great as width
near mid-length; ptilinal suture narrowly arched; parafacial
bare; facial ridge not expanded below, where it has few
setulae in two irregular series; face with no distinct carina,
somewhat explanate on central lower part, slightly
projecting below, but with epistomal margin sharply
reflexed to face ventrally; antennal grooves broad and
shallow; upper postocular area very narrowly visible in
profile, margined by a very regular series of closely placed
postocular setulae; the following cephalic bristles well
developed: inner and outer vertical, two pairs of fronto-
orbitals, postgenal; postvertical small; ocellar and lateral
occipital absent. Antenna about half as long as face on
median line; segment 6 long and slender, with very short
hairs on whole length. Prelabrum deeply sclerotised at sides,
much attenuated anteriorly; palpus moderately broad.

Thorax. Mesoscutum c. 0.85 times as long as wide;
scutellum somewhat flattened dorsally, with numerous
dorsal and some lateral setulae; mesopleuron moderately
convex on upper part; subscutellum very convex and
prominent (tachinid-like); the following thoracic bristles
well developed: humeral, 1+1 notopleurals, supra-alar,
postalar, posterior intra-alar, mesopleural, sternopleural,
three pairs of normal scutellars, and a series of c. nine

David K. McAlpine: Australasian Platystomatidae 185

Figures 108-110. Paryphodes hospes n.sp. 108, wing. 109, head. 110, distal part of aedeagus (scale = 0.05 mm).
Abbreviations: g, glans; gp, medial process of glans; st, stipe.

ventrolateral shorter bristles or enlarged setulae on each
side of scutellum; dorsocentral vestigial; prescutellar
acrostichal absent. Fore and mid legs rather short; hind leg
longer; fore femur with numerous non-seriate dorsal bristles,
and a series of moderately long posteroventral bristles; mid
femur more slender, with a series of postero ventral bristles;
hind femur longer and much stouter than other femora, not
curved, without strong bristles; mid tibia with one
moderately large apical ventral spur; mid basitarsus
elongate, much more than half as long as whole tarsus.
Wing: entire membrane microtrichose; both costal cells
unusually broad, with costa more arched on these cells than
usual (compare figs. 50, 52, 53, 55-57 of Hendel, 1914b);
vein 2 almost straight, except at origin; vein 3 with faint
undulation near middle of distal section; length of second
basal cell c. 0.87 of length of discal cell, both measured on
vein 4; vein 4 index = 0.47; second section of vein 4 with
very slight curvature; distal section of vein 4 with slight
sigmoid curvature, becoming parallel with vein 3 apically,
2.0 times as long as penultimate section; discal crossvein
moderately oblique, with sigmoid curvature, c. 1.7 times
as long as distal section of vein 5; anal crossvein with almost
imperceptible curvature; basal section of vein 5 with nine
or ten dorsal setulae; axillary lobe and squama both
moderately large and rounded.

Abdomen. Tergite 2 setulose mainly near posterior margin,
with small median group of larger setulae on margin and
more extensive lateral group of larger setulae; tergites 3
and 4 short; tergite 5 larger, almost uniformly setulose,
somewhat narrowed posteriorly, as long as tergites 3 and 4
together. Aedeagus (Fig. 110): stipe moderately long,
slightly expanded distally, but otherwise without differ¬
entiated preglans; glans moderately compact but constricted
and partly desclerotised near middle, distal section with
complex, asymmetrical sclerotisation, but no elongate
processes, basal section with cup-like cavity containing
complex fibrous structure (detail not shown in figure).

Dimensions. Total length 4.4 mm; length of thorax 2.3 mm;
length of wing 3.9 mm; length of glans of aedeagus 0.24 mm.

Distribution. Papua New Guinea—lowlands of East Sepik
Province.

Habitat. Tall rainforest remnant. I am informed that this
forest at the Agriculture Station, Bainyik, was later clear-
felled.

Notes. In the key to Paryphodes species of Hendel (1914b:
253-254), P. hospes runs imperfectly to P. tigrinus
Enderlein, but differs in having the thorax predominantly
dark brown instead of clay yellow and the second section

186 Records of the Australian Museum (2001) Vol. 53

of vein 4 shorter and much less curved. (P. tigrinus is a
junior synonym of P. nepticula (Loew), according to
Steyskal, 1980). Of the other species with yellowish marking
on the scutellum, P. duus Steyskal has the second section
of vein 4 longer and more curved and the wing much more
extensively shaded with brown; P. conspurcatus Hendel has
the distal section of vein 4 relatively short, the costa less
arched over the costal cells, and the wing markings quite
different (Hendel, 1914b: fig. 56); P. compticeps Enderlein
has extensive yellow striping on the me so scutum, black-
brown abdomen (male), and a broad dark brown band
covering “pterostigma” and anterior crossvein, all in
disagreement with P hospes. From the above it seems very
probable that P. hospes is distinct from any of the described
African species of Paryphodes, though most of these were
also described from very little material. Of the Afrotropical
species of Paryphodes that I have examined (in NMWC,
PM, and other collections), none has the setulae on vein 5
or the well differentiated series of ventrolateral bristles on
the scutellum seen in P. hospes, but there are numerous
described species not yet examined for these features.

The holotype has a female strepsipteran or stylops
projecting from the pleural membrane of the abdomen.
Parasitism does not appear to have affected the morphology
of the host fly, as is also the case in other stylopised
platystomatids examined. Perhaps these parasites are
referable to the family Dipterophagidae (see Drew &
Allwood, 1985).

The specific epithet is Latin for stranger or visitor, on
account of the unexpected occurrence of this Afrotropical
genus in New Guinea.

Genus Scholastes Loew
Figs. Ill, 112

Scholastes Loew, 1873: 38. Type species (generally accepted as
original designation, but see note below) Platystoma cincta
Guerin-Meneville.

Chaetostichia Enderlein, 1924: 133-134. New synonym. Type
species (original designation) C. aduncivena Enderlein.
Chaestotichia- Enderlein, 1924: 133. Variant spelling of above,
here interpreted as an incorrect original spelling.

Description. See Hendel (1914a). I add the following.

Wing. Stem vein without dorsal setulae before level of
humeral crossvein.

Male postabdomen. Epandrium with thick lobe on each side,
which overlaps base of outer surstylus; aedeagus with long
stipe; preglans undifferentiated; glans rather small, with
complex apical structure and no terminal filaments.

Female abdomen. Tergite 3 very large; tergite 4 vestigial;
tergites 5 and 6 vestigial or absent; aculeus slender, not
compressed.

Distribution. Australasian Region: Micronesia; Moluccas;
New Guinea; Bismarck Archipelago; New Caledonia; Fiji;
Tropical Polynesia; probably not established in Australia.
Oriental Region. Afrotropical Region: Seychelles only.

Notes. I follow universal usage in accepting Platystoma
cincta Guerin-Meneville as the type species, though this is
at least questionable under the ICZN. Although Loew (1873)

Figures 111-112. Ill, Scholastes taylori Malloch, wing. 112, Scholastes cinctus (Guerin-Meneville), distal part of
aedeagus (scale = 0.5 mm). Abbreviations: g, glans; gp, terminal process of glans; pg, preglans; st, stipe.

David K. McAlpine: Australasian Platystomatidae 187

stated that this was his type species, he simultaneously
indicated that Scholastes was introduced as a replacement
name because Pachycephala Doleschall was preoccupied.
Such dual implications are to be resolved by Article 66.8 of
ICZN (1999), especially as P. cincta was not among the
species originally included in Pachycephala Doleschall. If
Pachycephala mohnikei Doleschall is the type species of
Scholastes (through designation by Enderlein, 1924 as type
species of Pachycephala Doleschall, for which both
Scholastes and Oncoscelia Enderlein are replacement
names, see above under Euprosopia), then Scholastes
becomes a junior subjective synonym of Euprosopia
Macquart. The disused name Chaetostichia Enderlein would
then become the valid name for the genus which includes
P. cincta and allied species. As Scholastes auctorum is the
type of a currently used subfamily name, this substitution
is undesirable.

Scholastes, Paryphodes, and Lenophila are the only
Australasian platystomatid genera with a sternopleural
bristle (sometimes undeveloped in Lenophila). Scholastes
is distinguished from the other two by having an irregular
transverse ridge on the postfrons a little above the ptilinal
suture, and by having vein 2 strongly undulated opposite
the termination of vein 1. For other distinguishing characters
see under Paryphodes and the key to genera.

Evenhuis (1989) lists 11 Australasian species of
Scholastes. A further four possibly valid, mainly Oriental
species have been listed (Steyskal, 1977 and 1980). I am
not aware of any undescribed species in collections. I have
examined type material of Chaetostichia aduncivena
Enderlein in MNB, labelled “?Java”. It is typical of
Scholastes (n.comb.), but I have not determined its specific
status. Keys to species have been given by Hendel (1914b),
Curran (1936), and Malloch (1939a).

The larvae of Scholastes lonchifer Hendel (often
incorrectly given as lonchifera or lonchiferus ) and S.
bimaculatus Hendel are recorded as infesting coconuts
(Cocos nucifera; Auberton & Cheesman, 1929; McAlpine,
1973a).

Genus Trigonosoma Gray
Fig. 113

Trigonosoma Gray, in Griffith & Pidgeon, 1832: 774. Type species
(monotypy) T. perilampiformis Gray (correctly perilampiforme).
Tropidogastrella Hendel, 1914a: 18, 134-136. Type species
(original designation) T. tropida Hendel.

Figure 113. Trigonosoma cristiventre (Gerstaecker), wing. After
Hendel (1914a).

Description. See Hendel (1914a).

Distribution. Australasian Region: Moluccas—Ambon.
Oriental Region: widely distributed. Steyskal (1977) gives
also Ceram (or Seram), but he is mistaken in including
Ceram in the unrestricted type locality of T. cristiventre
(Gerstaecker). Gerstaecker (1860) gives “Amboina” (=
Ambon) only.

Notes. Trigonosoma cristiventre (Gerstaecker) from the
Moluccas is the only recorded Australasian species. The records
of this species from the Philippines by Frey (1930) need careful
checking. Steyskal (1971) gave a key to species.

Genus Zygaenula Doleschall
Fig. 114

Zygaenula Doleschall, 1859: 117. Type species (monotypy) Z.
paradoxa Doleschall.

Gorgopis Gerstaecker, 1860: 180-184. Type species (original
designation) G. bucephala Gerstaecker.

Gorgopsis Schiner, 1868: 229. Unjustified emendation for
Gorgopis Gerstaecker.

Figure 114. Zygaenula paradoxa Doleschall, wing. After Hendel
(1914a).

Description. The redescription by Hendel (1914a: 138-139)
is generally apt, even though he included species now
referred to Mesoctenia. I have no material on hand for a
revised description.

Distribution. Australasian Region: Moluccas (Ambon).
Oriental Region: Philippines. The ranges of Zygaenula and
Mesoctenia appear to be separated by Lydekker’s Line.

Notes. Zygaenula closely resembles Mesoctenia , but the
mid femur of the former is neither much thickened nor
ventrally spinose. Only the type species, Z. paradoxa
Doleschall (= Gorgopis bucephala Gerstaecker) is
recognized.

188 Records of the Australian Museum (2001) Vol. 53

Genus Mesoctenia Enderlein
Figs. 9, 115, 116

Mesoctenia Enderlein, 1924: 130-131. Type species (original
designation) M. ralumensis Enderlein (= Zygaenula coalescens
Hendel).

Description. See Me Alpine (1973a).

Distribution. New Guinea: widely distributed on mainland;
Aru. Bismarck Archipelago: New Britain; Manus. Australia:
Queensland.

Notes. I have seen at least six species of Mesoctenia , and
provided a provisional key to five of these (McAlpine, 1973a).

Hendel (1914a, 1914b) included Mesoctenia spp. in the
genus Zygaenula , but Enderlein established the separate
genus Mesoctenia for species with thickened, ventrally
spinose mid femur. Malloch (1939a) accepted Enderlein’s
genus. Without material of the type species of Zygaenula
for study, I am uncertain if this generic separation of
Mesoctenia on a single character is justified. The wing
illustration of Zygaenula paradoxa by Hendel (1914a, here
copied as Fig. 114) suggests some venational differences
from available Mesoctenia spp., but I am unable to confirm
these at present.

G.A. Holloway and the author collected more than 20
specimens of Mesoctenia australis McAlpine on a young
palm (Arecaceae) at Thornton Range, Queensland, in 1967.
As a majority of these were soft, newly eclosed specimens,
it is likely that the immature stages were also associated
with the palm.

Genus Naupoda Osten Sacken
Figs. 117-120

Naupoda Osten Sacken, 1881b: 135. Type species (monotypy) N.
platessa Osten Sacken.

Description. See Hendel (1914a) for a detailed description,
to which I add the following.

Male abdomen. Tergites 4 and 5 well sclerotised, but shorter
than tergite 3; epandrium without distinct lateral lobe; outer
surstylus with pair of moderately small terminal prensisetae;
aedeagus with moderately long stipe; preglans forming
separate sclerite; glans well developed, with complex,
variable sclerotisation; terminal filaments absent; cercus
with moderately long, rather few to numerous, irregularly
placed setulae.

Female abdomen. Tergite 3 large; tergites 4 and 5 completely
desclerotised (probably so in all true Naupoda spp.);
ovipositor sheath depressed, keeled on each lateral margin;
aculeus slender.

Distribution. Australasian Region: New Guinea; Solomon
Archipelago—Choiseul, Malaita; Australia—east coast as
far south as Richmond River district, Lord Howe Island.
Oriental Region: widely distributed. Afrotropical Region:
widely distributed in African tropics; Madagascar.

Notes. The Fijian species Naupoda simmondsi Bezzi was
misplaced in this genus and is here transferred to the new
genus Phlyax, q.v. The Oriental species N. strigifera de
Meijere, N. ypsilon van der Wulp, and N. ypsilonoides de
Meijere are apparently misplaced in Naupoda, from
examination of type material (AMST), and their generic
placement needs further study.

Figures 115-116. Mesoctenia australis McAlpine. 115, wing. 116, male external genitalia, setulae omitted from
epandrium and cerci (scale f= 0.5 mm). Abbreviations: aa, aedeagal apodeme; c, cercus; ep, epandrium; g, glans; is,
inner surstylus; os, outer surstylus; pg, preglans; st, stipe.

David K. McAlpine: Australasian Platystomatidae 189

Figures 117-120. Naupoda nudiseta (Bezzi). 117, wing. 118, distal part of aedeagus (scale = 0.2 mm). 119, scales
from lining of cavity of glans, acuminate ones nearest rim (scale = 0.025 mm). 120, Naupoda platessa Osten
Sacken (?) (Luzon, BPB), distal part of aedeagus (scale = 0.1 mm). Abbreviations: bu, bulb; cv, cavity of glans; g,
glans; gp, terminal process of glans; lg, sclerotised lobe of glans; pg, preglans; pu, pustulose sclerite; st, stipe.

The most typical species of Naupoda available to me
are the Oriental N. platessa Osten Sacken, N. imitans de
Meijere, and an undetermined species from West Malaysia.
I place these in the subgenus Naupoda s.str. The known
Australasian species all differ from Naupoda s.str. in several
significant characters as shown in Table 3, and are here
placed in the new subgenus Gonga. The subgeneric
placement of the Afrotropical species (see Steyskal, 1980)
and the few remaining Oriental species (see Steyskal, 1977)
remains to be worked out. I have examined the aedeagus in
N. (Naupoda) platessa Osten Sacken (or near, Luzon), N.
(Gonga) nudiseta (Bezzi), and N. (Gonga) regina Hendel
(Queensland population), and have found great structural
differences between all three. Naupoda (G.) nudiseta differs
from the other two in that the basal part of the glans contains
a capacious, distally open cup-like cavity lined with several
hundred densely packed scales (Figs. 118, 119), which is
absent in the other two (compare Fig. 120).

Adults of Naupoda regina Hendel have been observed

several times on foliage of rainforest trees in Australia. Bezzi
(1928) quoted label data for N. nudiseta (Bezzi) from Lord
Howe Island as said to do damage to peaches and bananas.
The report may be due to confusion with Dacus (or
Bactrocera) spp. (Tephritidae) which abound on the island
(author’s observations).

Subgenus Gonga n.subgen.

Type species: Pterogenia nudiseta Bezzi.

Description. See Table 3 for characters distinguishing
Gonga from Naupoda s.str. In addition, species of Gonga
have the anterodorsal gibbosity of the mesopleuron less
pronounced than in Naupoda (Naupoda) platessa, the anal
crossvein more markedly oblique so that the posterodistal
angle of the anal cell is much more obtuse than the
anterodistal angle, the basal section of vein 5 not setulose
(condition variable in Naupoda s.str.), the conjoined cerci

190 Records of the Australian Museum (2001) Vol. 53

Table 3. Subgeneric characters in genus Naupoda.

Subgenus Naupoda Subgenus Gonga

• Fronto-orbital bristle absent

• Epistomal margin of face narrowly

reflexed to form a marginal ridge, which
diverges from margin on each side

• Peristomial series of setulae

undifferentiated

• Subcosta with very slight apical curvature

• Discal crossvein bent abruptly just before

anterior end

• Basal section of vein 4 with weak, flexible

point near distal third, sclerotised

beyond this point

• Fronto-orbital bristle present

• Epistomal margin of face simple

• A differentiated series of fine anterior

peristomial setulae present

• Subcosta with strong apical curvature

• Discal crossvein curved, but without

abrupt bend

• Basal section of vein 4 weakened

along much of distal quarter

of the male mounted on distal end of the non-sclerotised
proctiger (instead of occupying the whole of the sides of
the proctiger as in N. platessa). The squama is much larger
than the axillary lobe, as in Naupoda s.str., but in contrast
to some species referred, perhaps incorrectly, to the genus
Naupoda.

Notes. This subgenus includes the following species:
Naupoda (Gonga) nudiseta (Bezzi) (syn. N. insularis
Paramonov) from Lord Howe Island; N. (Gonga) regina
Hendel from New Guinea and eastern Australia, and N.
(Gonga) ventralis Curran from the Solomon Archipelago.
These are all the Australasian species of the genus Naupoda
known to me.

The subgeneric name is a contraction of the Greek
gongylos, a sphere, and is feminine because of the
termination.

Genus Asyntona Osten Sacken

Figs. 3, 121

Asyntona Osten Sacken, 1881b: 135. Type species (monotypy) A.
doleschalli Osten Sacken.

Description. See Hendel (1914a).

Distribution. Moluccas: Ambon; Kai Islands. New Guinea:
mainland; Aru; Trobriand Islands. Solomon Archipelago:
Bougainville; Tulagi (Florida Group); Guadalcanal.
Queensland: Coen district northwards.

Malloch (1939a) recorded Asyntona tetyroides (Walker)
from Philippine Islands, probably because Osten Sacken
(1882) redescribed the supposed synonym, A. doleschalli,
in a work on Philippine Diptera, which Malloch cited.
However, Osten Sacken clearly gave the locality as
“Amboina” (i.e. Ambon, Moluccas) for this species. I know
of no other records of Asyntona from the Oriental Region,
and I think the genus is endemic to the Australasian Region.

Notes. Hendel (1914b) and most subsequent authors have
generally accepted only two species of Asyntona, viz. A.
flavipes Hendel, with yellow head, and A. tetyroides
(Walker) with predominantly blackish head, the names A.
doleschalli Osten Sacken and A. paradoxa de Meijere being
treated as synonyms of A. tetyroides. This interpretation is
almost certainly not wholly correct. I have examined at least
three dark-headed species (all in AM), but females and,
perhaps, some diminutive males may be hard to place. The
figures of Hendel (1914a: figs. 247-249, locality not given)
may represent a fourth such species. Males (but perhaps
not diminutives) of these species differ in armature of
antennal segment 2 and of the hind trochanters. Further

Figure 121. Asyntona “sp. A” (Wewak, Papua New Guinea, AM), wing.

David K. McAlpine: Australasian Platystomatidae 191

study of type material and comparative morphology is
needed to enable specific identifications.

Asyntona resembles Naupoda but has a larger alula, more
complex folding of the wing (Malloch, 1939a), and tergites
4 and 5 of the female abdomen well sclerotised. The fringe
of numerous marginal scutellar bristles is distinctive.

A specimen of Asyntona sp. from Trobriand Islands
(J.W.I., AM) is labelled “behind beach, forest”. Specimens
of two distinct Asyntona spp. from Papua New Guinea (AM)
have labels indicating that they were associated with
Pandanus plants (Pandanaceae). Malloch (1940) recorded
a specimen of A. flaviceps Hendel from Guadalcanal
associated with Pandanus.

Subfamily Trapherinae

Description. Fronto-orbital bristles well developed; lower
end of facial ridge often broadened and with field of
irregularly placed setulae; face usually with transverse
convexity below, or with transverse ridge or gibbosity near
middle; cheek often with linear stripe of outstanding
pubescence along lower margin of eye. Arista usually long-
bipectinate (with only short hairs in Eopiara). Palpus usually
broad and rather short. Wing: subcosta usually nearly
straight and gradually approaching costa distally or with
very short, often desclerotised transverse distal section;
second basal cell often much broader than first basal cell,
which often has a pronounced bend near junction of basal
crossvein with vein 4; vein 7 usually represented beyond
alular incision by a long, sharp crease, often accompanied
by pigment; squama forming a small lobe, or vestigial and
without lobe. Male postabdomen (studied in only a few
genera): aedeagus without glans and terminal filaments,
sometimes very short for the family (e.g., Xiriella sp.), or
long, strap-like, and rolled up when not in use (e.g., Xiria
spp.); cerci usually short, joined to membrane for most of
their length, usually numerously and quite irregularly
setulose (except in Phlyax). Female abdomen: tergites 4
and 5 unreduced; tergite 6 usually broad and, though always
short, often permanently exposed, sometimes vestigial;
ovipositor sheath usually rather short; aculeus often broad
and blade-like.

Notes. Very little has been published on the taxonomy and
morphology of this subfamily, apart from simple character¬
isations of included taxa. Some brief attempts to define and
justify the group have been made by Hendel (1914a),
Steyskal (1965, 1990), and McAlpine (1973a). The above
characterisation is intended to aid decisions as to included
taxa, though it has an admittedly weak basis. Some genera
are still known from very few specimens, e.g., Aglaioptera
and Eopiara , which are only represented in HELS
collection, so far as I am aware.

Previously described genera of Trapherinae include: Lule
Speiser, Microlule Steyskal, Piara Loew, Seguyopiara
Steyskal, Traphera Loew, in the Afrotropical Region;
Aglaioptera Frey, Eopiara Frey new stat. (originally as
subgenus of Piara), Phasiamya Walker ( Phasiamyia,
subsequent misspelling by Hendel, 1914a; syn. Pachymyza
Frey), Poecilotraphera Hendel, Xiria Walker, Xiriella Frey new

stat. (originally as subgenus of Xiria) in the Oriental Region.

Oriental records of the genera Lule and Piara (see
Steyskal, 1977) are due to misplacement of species of
Phasiamya, Xiriella, and Eopiara. In this connection,
Xiriella lunaris (de Meijere, 1916) and Eopiara chrysoptera
(Frey, 1964) are new combinations. Lule speiseri de Meijere,
1914 is a new synonym of Phasiamya metallica Walker,
1849, from my examination of types of these Oriental
species. Also, Eopiara elegans (Frey, 1964) is a new
combination (from Lamprogaster).

The new genus Phlyax, evidently endemic to Fiji, is the
first known trapherine in the Australasian Region. The species
recorded from New Guinea as Xiria papuana Hennig and Xiria
strigata Hennig belong in the platystomatine genus Bama, q.v.

Genus Phlyax n.gen.

Fig. 122

Type species: Naupoda simmondsi Bezzi.

Description (S, $). Small very rotund dark metallic flies,
with short moderately slender legs; habitus reminiscent of
Naupoda.

Figure 122. Phlyax simmondsi (Bezzi), wing. Modified from
A.J.E. Terzi, in Bezzi (1928).

Head somewhat anteroposteriorly compressed; median zone
of postfrons bare on entire width, with a series of incurved
setulae on each side of bare zone; face largely smooth,
transversely concave near middle, with lower margin
prominently projecting, and median carina reduced to a short
tubercle between antennae; facial ridge slightly expanded
below, where fine setulae form a narrow field instead of a
single series; peristomial setulae relatively long, forming a
single series and becoming shorter where they approach
bottom of facial ridge; parafacial narrow, finely setulose at
upper extremity; occiput convex on each side below, very
broadly concave above; the following bristles present: inner
and outer vertical, very small postvertical, small ocellar,
two well-developed reclinate fronto-orbitals, weakly
differentiated postgenal; lateral occipital bristle absent.
Antenna (excluding arista) much shorter than face; segment
3 elongate-oval; segment 6 long-bipectinate on most of
length. Palpus broad, of moderate length.

Thorax very broad, dorsally convex; mesoscutum 0.71-0.75
times as long as wide; scutellum very broad, without setulae,
densely pubescent on free margins; posterior margin thin,

192 Records of the Australian Museum (2001) Vol. 53

but not sharp; the following bristles present: humeral, 1+1
notopleurals, supra-alar, postalar, posterior intra-alar,
prescutellar acrostichal, mesopleural, three pairs of
scutellars of which those of apical pair are longer and
separated from each other by nearly half width of scutellum;
scapular, dorsocentral, and sternopleural bristles absent.
Fore femur with a series of posteroventral bristles; femora
otherwise without differentiated bristles or spines; mid tibia
with long apical ventral spur and shorter anteroventral spur;
tibiae otherwise unarmed. Wing with oblique crease
enabling flexure of apical part as in Naupoda and Asyntona,
but crease not entering discal cell; section of costa on second
costal cell horizontally dilated, with microtrichia on dorsal
surface in transverse rows, appearing like fine transverse
striations; subcosta almost straight distally; vein 1
terminating close to subcosta; anterior crossvein short,
meeting vein 4 near distal end of discal cell; discal crossvein
strongly bent near anterior extremity; second basal cell much
broader than anal cell; vein 5 without setulae; anal crossvein
only slightly curved; distal section of vein 6 shorter than
preceding section, not reaching margin; squama much
reduced, without any lobe.

Abdomen broad, dorsally convex; tergites 4 and 5 unreduced
in both sexes. Male postabdomen: epandrium with rounded
posterolateral lobe on each side; outer surstylus slender,
with minor setulae only; inner surstylus short and stout with
two massive prensisetae; aedeagus shorter than epandrial
complex, broad, largely membranous, with little apical
sclerotisation and no glans or terminal filaments; cerci
elongate, joined by membrane for entire length, each with
only c. six setulae in a spaced longitudinal series. Female
postabdomen: tergite 6 broad but very short, usually partly
concealed in dried specimens; ovipositor sheath short and
rather broad; aculeus with somewhat dilated basal section
and parallel-sided, obtuse apical section.

Distribution. Fiji: Viti Levu Island (numerous localities,
BPB, BM).

Notes. Phlyax simmondsi (Bezzi, 1928) n.comb., the only
known species of Phlyax, was described in the genus Naupoda.

Phlyax differs from Naupoda in having: longer seriate
peristomial setulae (these short or undifferentiated in
Naupoda)’, arista long-bipectinate (instead of quite short-
haired); scutellum without setulae (instead of extensively
setulose dorsally); mid tibia with two unequal apical ventral
spurs (instead of only one spur); section of costa on second
costal cell broadly compressed (instead of slender,
cylindrical); vein 1 closely approximated to end of subcosta
(instead of diverging from subcosta distally); vein 3 bare
(instead of dorsally setulose); anterior crossvein meeting
vein 4 near distal end of discal cell (instead of, typically,
near mid-length of discal cell); basal section of vein 4
without weak, flexible point (basal section of vein 4 with
obvious flexible point or region beyond mid-length);
oblique crease of wing membrane not entering discal cell
(instead of intersecting discal cell); shorter, differently
shaped second basal cell (compare Figs. 122 and 117),
squamal lobe absent (instead of quite large), aedeagus short,

without glans (instead of relatively long with well-
developed glans), each cercus in male with setulae restricted
to a single, spaced series (instead of having many irregularly
placed setulae); abdomen of female with tergites 4 and 5
well developed (instead of quite desclerotised).

Naupoda ypsilon Wulp, 1899 from Java resembles
Phlyax in the proximity of the anterior and discal crossveins,
but the venation does not otherwise resemble that of Phlyax.
It and certain related species are misplaced in Naupoda, as
noted under that genus.

Similarity between Phlyax and Naupoda may be due to
convergence in features related to mimicry of Coleoptera,
perhaps particularly of alticine (galerucine) chrysomelids.
The partly folded distal section of the fly’s wing suggests
the exposed tip of an imperfectly folded beetle’s wing.

Naupoda is typical of the subfamily Scholastinae, but
certain features of Phlyax seem to indicate a relationship to
the Trapherinae rather than the Scholastinae, notably the
reduced squama, as in the trapherine genera Aglaioptera
Frey and Poecilotraphera Hendel, unreduced tergites of the
female abdomen, and the very short aedeagus lacking the
glans, which is rather like that of the one examined species
of Xiriella. The only platystomatids known to lack the glans
belong in the Trapherinae, but it is not yet known if this is
the case in all included genera.

The venation of Phlyax is somewhat similar to that of
the Afrotropical trapherine genus Lule Speiser, from which
Microlule Steyskal is doubtfully generically distinct, in my
opinion. Phlyax differs from Lule in the termination of vein
1 close to the subcosta, the absence of vein 7, even as a
slightly pigmented crease, and the strong divergence of
veins 3 and 4 distad of the discal crossvein. Phlyax has
some resemblance to the Oriental genera Phasiamya Walker
and Xiriella Frey, but differs from these in the dorsally
hairless scutellum, bare vein 5, absence of visible vestige
of vein 7 beyond the alula, and absence of the squamal lobe.

The name Phlyax (Latinised stem phlyac-) is Greek for
jester. The gender is masculine.

Genera unplaced to subfamily

Genus Apiola McAlpine

Fig. 123

Apiola McAlpine, 1973a: 31. Replacement name for Xenognathus
Malloch. Type species (automatic) Xenognathus bryani
Malloch.

Xenognathus Malloch, 1930b: 225-226 (preocc. Xenognathus
Gilbert, 1915). Type species (original designation) X. bryani
Malloch.

Description. See Malloch (1930b). I have added (McAlpine,
1973a) that tergite 6 of the female abdomen, though much
shorter than tergite 5, is well developed and not capable of
being concealed below the latter, thus resembling Angitula
(subgenus Giraffomyia ) and Xiria.

Distribution. Tropical Polynesia: Western Samoa.

David K. McAlpine: Australasian Platystomatidae 193

Figure 123. Apiola bryani (Malloch), wing. After A.J.E. Terzi, in Malloch (1930b).

Notes. Malloch (1930b) described two sympatric species,
but expressed doubt as to their specific status. Lack of
material has prevented further investigation of this question.

The male postabdominal structures of Apiola are
undescribed, and, as there is no obvious close relationship
to any other genus, its systematic position is hard to
determine. The antennal features, including the form of
segment 2, and the details of the ovipositor sheath and
aculeus are reminiscent of some Atopognathus species, but,
in the latter, tergite 6 is quite vestigial. With our present
imperfect knowledge of the morphology of many
platystomatid genera, I can only conjecture that its
relationships may lie with such genera as Atopognathus and
Guamomyia, which are difficult to place in the broader system.

Apiola is easily identified by the slender, ichneumonid-
like habitus and obvious wing features (Fig. 123), in
combination with the long-bipectinate arista, presence of
ventral spinescent bristles on all femora, and absence of
postgenal, mesopleural, and dorsocentral bristles.

The altitudes given by Malloch (2000-4000 feet = c.
600-1200 m) indicate a mountain habitat.

Genus Atopognathus Bigot
Fig. 4, 124

Atopognathus Bigot, 1881a: 47-48; 1881b: 24. Type species
(monotypy) A. platypalpus Bigot.

Dasiortalis de Meijere, 1913: 378 (exHendelMS). Synonymised
Hennig, 1940a. Type species (monotypy) Ortalis contigua
Walker [= Atopognathus complens (Walker)].

Dasyortalis- Hendel, 1914a: 129-131. Incorrect subsequent
spelling of above.

Lasioxiria Hendel, 1914a: 12, 37-38. Type species (original
designation) L. hirsuta Hendel. N.syn.

Lasiopsila Curran, 1936: 54. Type species (original designation)
L. fasciata Curran [= A. complens (Walker)].

Description. Hendel (1914a) keyed the genus (as
Dasyortalis) and gave a detailed description. Although this
description did not take into consideration the characters
of the species he included in Lasioxiria, it indicates much
of the variation present in the genus, and is generally
adequate. I add the following data to Hendel’s description:

Antenna. Segment 6 well haired for most of length, either
bipectinate or with irregularly placed hairs.

Thorax. Dorsocentral bristles one to about four pairs.
Axillary lobe with fringe of long hairs, each >4 times as
long as pubescence on vestigial squama, and not restricted
to lower (squamal) region of lobe.

Abdomen. Male postabdomen: aedeagus with complex,
sclerotised glans, without terminal filaments. Female
postabdomen: tergite 5 large; tergite 6 much reduced,
usually concealed; aculeus broad and blade-like, often acute,
but sometimes obtuse or truncate.

Distribution. Moluccas: Ternate; Ambon. New Guinea:
mainland—widely distributed; Aru; Normanby Island.
Bismarck Archipelago: New Britain; Makada Island (Duke
of York Group); New Ireland. Solomon Archipelago:
Bougainville; Guadalcanal; Matema Island (Santa Cruz
Group). Australia: Queensland.

Figure 124. Atopognathus “sp. C” (Queensland, AM), wing.

194 Records of the Australian Museum (2001) Vol. 53

Notes. Hennig (1940a) examined type material of
Atopognathus platypalpus Bigot, and, as a result,
synonymised “ Dasyortalis ” under Atopognathus, noting that
Bigot’s species seemed to be distinct from any treated by
Hendel (1914b). Malloch (1939a) suggested the possibility
of synonymy between Lasioxiria and “ Dasyortalis ”, but in
his key used the presence of a pair of “rounded or oval
elevations” below the antennal foveae in Lasioxiria to
distinguish it from “Dasyortalis”. I find these elevations to
be differentiated to a variable extent among the species of
Atopognathus , but the variation does not provide a basis
for sharp group distinctions. The type species of Lasioxiria
is more elongate in habitus, wing, and cell proportions than
A. complens (Walker) and closely related species, but again
this variation does not clearly define major groupings.

One undescribed species (“sp. M” in AM, female only
known) lacks setulae on vein 5 and has a series of long
black bristles on the anterodorsal margin of antennal
segment 3, but this species is linked through a related
species, which is normal for Atopognathus in these features,
to the rest of the genus. Several species with relatively
narrow postfrons also have the fronto-orbital plates narrow.
Bigot (1881b) described the male of A. platypalpus as “with
the cheeks anteriorly, below the eyes, on each side
uniappendiculate, with the appendages compressed,
moderately erect, obovate ...” (my translation). The only
male of the type series (OX) with head still present shows
no such appendages, nor have I seen such structures in other
Atopognathus species. Males of several species have
broadened heads (Fig. 4).

A relatively large Atopognathus species (sp. J in AM),
which lives in highlands of PNG, is very similar to
Lenophila spp. in appearance, with its dark, shining body,
glossy black scutellum, and characteristic wing stripes. The
similarity is perhaps due to convergent mimicry of jumping
spiders. It is distinguishable from Lenophila by its distinctly
haired (ommatrichose) eyes, vestigial squama, sinuate vein
2, setulose vein 5, and, in the female, its unreduced tergite
5 and blade-like aculeus.

Atopognathus includes at least 20 species, many of them
undescribed. Atopognathus hirsutus is a new combination
for Lasioxiria hirsuta Hendel.

Hendel (1914a, 1914b) associated Ortalis tarsalis
Walker, 1861c, from Batjan (Bacan, Moluccas), and O.
leucomera Walker, 1864, from Misool (New Guinea), with
“Dasyortalis” , without seeing material. Ortalis tarsalis
belongs in Chaetorivellia (q.v.), but generic placement of

O. leucomera remains doubtful, as type material has not
been seen.

J. Ismay has reared A. complens (Walker) from larvae
living in a ginger stem damaged by a lepidopterous larva in
Papua New Guinea (label data in AM). Adults of an
undescribed Atopognathus sp. are commonly found on the
large leaves of Alocasia brisbanensis (family Araceae) in
or at the margins of rainforest on the Atherton Tableland,
Queensland (author’s observations).

Genus Guamomyia Malloch
Figs. 125-127

Guamomyia Malloch, 1942: 206. Type species (original
designation) G. fascipennis Malloch.

Description. The following data amplify the description
by Malloch.

Head. Postfrons coarsely but not densely setulose; inner
vertical bristle smaller than and situated in advance of outer
vertical; lower end of parafacial with patch of non-seriate
setulae; cheek with a series of longer peristomial setulae,
one of which corresponds in position to a vibrissa.

Thorax. Scutellar bristles two or three pairs. Wing: subcosta
with distal section bent away from vein 1 at an obtuse angle,
meeting costa at a very acute angle; anal crossvein strongly
and rather evenly curved to almost straight.

Male postabdomen. Aedeagus with compact sclerotised
glans, without terminal filaments.

Female postabdomen. Tergite 5 unreduced and exposed;
tergite 6 apparently absent, though sternite 6 present; aculeus
slender, not depressed, obtuse.

Distribution. Micronesia: Guam. Papua New Guinea:
mainland. Bismarck Archipelago: Mussau; Lavongai (or
New Hanover). Australia: Lord Howe Island. The genus is
apparently absent from the Australian mainland, and the
known distribution is discontinuous.

Notes. I have seen c. four species of Guamomyia but only
one is yet described. The genus was previously mentioned
and keyed as “genus A” (McAlpine, 1973a).

Figure 125. Guamomyia “sp. A” (Lord Howe Island, AM), wing.

David K. McAlpine: Australasian Platystomatidae 195

Figures 126-127. Guamomyia “sp. A”. 126, head. 127, distal part of aedeagus (scale = 0.1 mm).

The late Z.R. Liepa reported in litt. (29.xii.1972) on her
observations on the Lord Howe Island species: “They were
all taken on Pandanus roots, and most were on the juicier,
not yet earthed roots, and resembled the black spots on the
roots so much that, unless they moved, you couldn’t see
them.”

Acknowledgments. J. Bowley, S.P. Kim, D.S. Kent, J. Allen,
M. Weight, and R. Calvert are thanked for providing many of the
illustrations. The following provided important study material:
D.J. Bickel, G. Daniels, B.J. Day, G.A. Holloway, J.W. Ismay, R.
de Keyzer, M.S. Moulds, H. Roberts. The following provided
access to important collections: K. Arakaki and N.L. Evenhuis
(BPB), J. Chainey and B.R. Pitkin (BM), PS. Cranston and D.H.
Colless (ANIC), G. Daniels and M. A. Schneider (UQ), J.C. Deeming
(NMWC), R. Meier (ZMC), L. Matile (PM), A.C. Pont (OX). D.J.
Bickel, J. Chainey, B.J. Moulds, L. Papp, J. Tamawski, and A.E.
Whittington helped with information, discussion, or in other ways.

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Batchian, Kaisaa, and Makian, and at Tidon in Celebes, by
Mr. A.R. Wallace, with descriptions of new species. Journal
of the Proceedings of the Linnean Society of London, Zoology
5:270-303.

Walker, F., 1862. Catalogue of the dipterous insects collected at
Gilolo, Ternate, and Ceram, by Mr. R. Wallace, with
descriptions of new species. Journal of the Proceedings of the
Linnean Society of London, Zoology 6: 4-23.

Walker, F., 1864. Catalogue of the dipterous insects collected in
Waigiou, Mysol, and North Ceram by Mr. A.R. Wallace, with
descriptions of new species. Journal of the Proceedings of the
Linnean Society. Zoology 7: 202-238.

Williston, S.W., 1908. Manual of North American Diptera, 3rd
edn., pp. 405. New Haven: James T. Hathaway.

Wulp, F.M. van der, 1899. Aanteekeningen betreffende Oost-Indische
Diptera. Tijdschrift voor Entomologie 41: 205-223, pi. 10.

Manuscript received 12 April 2000, revised 12 September 2000 and
accepted 14 September 2000.

Associate Editor: D.J. Bickel.

David K. McAlpine: Australasian Platystomatidae 199

Index to genus group names

Pagination for the principal reference is bold, reference is also made to keys (italics), and to illustrations
(*); abbreviations: nA = apparently not Australasian, sg = subgenus, syn = invalid generic synonym.
Couplet numbers in the key to Australasian platystomatid genera (pp. 121-130) are given with page
numbers between square brackets [page: couplet number].

Achias, 116, 117, 118*, [123: 16],
141, 142*, 143*

Achiosoma syn, 141
Adantineura sg, 161, 162
Aetha, [127: 49], 137, 153, 154*
Agastrodes syn, 180
Aglaioptera nA, 191, 192
Angitula, 118*, 120, [122: 7], 172,
173, 174, 175*, 176*, 192
Angitulina syn, 173, 175
Angituloides n.stat., sg, 120, 172-
175, 176, 177
Anguitula syn, 173
Antineura, 117, [122: 12], 161*, 162
Apactoneura, [126: 42], 156*, 157
Apiola, 117, 120, 121, [123: 16],
192, 193*

Asyntona, 118*, [124: 24], [125:

37], 111, 190*, 191, 192
Atopognathus, 117,118*, 120, [121:

5], 178, 193*, 194
Bama, [121: 2], [128: 55], [130: 69],
165,166, 167*, 168*, 169,172,
191

Brea, [122: 9], 130, 138*
Bromophila nA, 130, 152
Campigaster syn, 162
Carolimyia, 158
Celetor syn, 177
Ceratopelta syn, 139
Chaestotichia syn, 186
Chaetorivellia, 116, 120, 121, [127:

47], 177, 179*, 194
Chaetostichia syn, 186, 187
Charax syn, 157
Chromatomyia syn, 139
Cleitamia, [126: 39], 137,166,169,
170, 171*, 172

Cleitamoides, [122: 14], 137, 169,
170*, 171

Conicipithea, [123: 21], 146, 160,
161*

Cruphiocera syn, 139
Cryphiocera syn, 139
Dasiortalis syn, 193
Dasyortalis syn, 193, 194
Ditomogaster syn, 180
Duomyia, 115, 117, 120, 122, [123:
19], [128: 56], [130: 70], 142,
152, 161, 162, 163*

Duomyza syn, 162
Elachigaster syn, 180
Elassogaster, [123: 18], [125: 31],
148, 150, 151, 152, 153*, 154
Eopiara nA, 191
Epicausta syn, 152
Euchalcota syn, 162

Eumeka, 117, [129: 66], 136, 145,
146, 147*

Euprosopia, 116, [121: 4], 151, 162,
164*, 165, 180, 187
Euthyplatystoma nA, [126: 46]
Euxestomoea, [130: 68], 165*, 166
Giraffomyia n.stat., sg, 120, 172-
176, 177, 192
Gong a n. subgen., 189, 190
Gorgopis syn, 187
Gorgopsis syn, 187
Guamomyia, 117, 121, [127: 50],
193, 194, 195*

Hammatopelma syn, 173, 175
Helocnemia syn, 162
Hemigaster syn, 182
Hysma, 119*, [129: 64], 130, 131,
132

Icteracantha nA, 151, 157
Inium, 117, [123: 20], [127: 52],
142, 143*

Laglaisia, 118, [126: 45], 137,165,
166, 169*

Lamprogaster, 116, 117, [121: 4],
[123: 19] [22], [128: 56], 130,
139, 140*, 141, 157, 191
Lamprophthalma, [123: 17], 160
Lasiopsila syn, 193
Lasioxiria syn, 193, 194
Lenophila, 116, 117, [125: 34],
[129: 62], 177, 178*, 187, 194
Liolamprogaster syn, 139
Loriomyia, 119, [126: 39], 171*,
172

Loxoneuroides, [129: 62], 132*
Lule nA, 191, 192
Maria syn, 138
Meachina, 173, 175
Meringomeria, 117, [124: 30], 148,
151,157*, 158

Mesoctenia, 116, 118*, [122: 9],
187, 188*

Microepicausta, [125: 32], 130,136,
148, 150, 151, 152*, 153, 154
Microlule nA, 191, 192
Montrouziera, 116, [127: 51], 141*
Myiodella syn, 152
Mystia syn, 141

Naupoda, 117, [125: 37], 188, 189*,
190-192

Neohemigaster, [124: 26], 177,180,
182, 183*

Oncoscelia syn, 162, 187
Pachycephala syn, 162, 187
Pachymyza nA syn, 191
Par, 116, [125: 32], [128: 60], 148,
150, 151*, 152, 153

Paryphodes, 115, 117, [125: 35],
180, 183, 184, 185*, 186, 187
Peronotrochus sg, 158
Phasiamya nA, 120, 191, 192
Phasiamyia syn, 191
Philocompus nA, [122: 12]

Phlyax, 116, 117, [127: 47], 188,
191*, 192
Piara nA, 191

Plagiostenopterina, [124: 29], [127:

48], 152-157, 158*, 159, 160
Poecilotraphera nA, 191, 192
Pogonortalis, 115, 117, [127: see
50], [129: 63], 154, 155*
Polimen n.sg., 166
Poticara syn, 171

Pseudepicausta, [124: 28], [129: 67],
145, 146, 159*, 160
Pseudo cleitamia, [126: 40], 159,
160*

Pseudorichardia, 116, 117, [126:
42], 138, 139*

Pterogenia, [121: 3], 177, 180, 182*,
183

Rhytidortalis, [124: 28], [129: 67],
142, 144*, 146, 162
Rivellia, 115, 117, [128: 61], 130,
131, 133*, 134-137, 159, 172
Scelostenopterina nA, [126: 43]
Scholastes, 116,117, [125: 35], 179,
180, 184, 186*, 187
Scotinosoma, 117, [130: 71], 144*,
145

Seguyopiara nA, 191
Senopterina, 146, 174
Signa, 116, [130: 71], 147, 148,
149*, 150, 151
Simomesia syn, 183
Stenopterella syn, 157, 158
Stenopterosoma sg, 158, 159
Systellodiscus syn, 183
Tarfa, 116,117, [128: 58], 136, 137*,
154, 172

Terzia, 116, 120, [121: 7], 130, 172,
173*, 174

Tomeus, [127: 53], 177, 179, 180,
181*

Trigonosoma, [124: 25], 187*
Tropidogastrella syn, 187
Xenognathus syn, 192
Xiria nA, 120, 166, 191, 192
Xiriella nA, 191, 192
Zealandortalis, 117, [128: 61], 133,
134, 135*

Zygaenula, 116, [124: 26], 187*, 188

© Copyright Australian Museum, 2001

Records of the Australian Museum (2001) Vol. 53: 201-241. ISSN 0067-1975

Revision of Paralamyctes
(Chilopoda: Lithobiomorpha: Henicopidae),
with Six New Species from Eastern Australia

Gregory D. Edgecombe

Australian Museum, 6 College Street, Sydney NSW 2010, Australia
greged @ austmus. gov. au

Abstract. Paralamyctes Pocock, 1901 has a classic Gondwanan distribution, with species previously
known from New Zealand, southern Africa, Madagascar and Chile. The genus as hitherto conceived is
paraphyletic, with Triporobius Silvestri, 1917 (India) and Haasiella Pocock, 1901 (New Zealand,
Tasmania) nesting within the group. Species from Argentina and New Caledonia originally assigned to
Paralamyctes belong to Analamyctes Chamberlin, 1955. Revision of Paralamyctes based upon
monophyletic subgroups recognizes four subgenera: P. (Paralamyctes), P (Haasiella), P. (Nothofagobius)
n.subgen. and P. (Thingathinga) n.subgen. The eastern Australian occurrence of Paralamyctes is
demonstrated by six new species. Paralamyctes (Thingathinga) grayi n.sp. is distributed from the Blue
Mountains to the Budawang Range, New South Wales; P. (Thingathinga) from the Barrington Tops may
be specifically distinct. Paralamyctes (T.) grayi and the allied P. (I) hornerae n.sp. from northern New
South Wales have affinities to the New Zealand P. (T.) validus. Paralamyctes (Nothofagobius) cassisi
n.sp. occurs in northern New South Wales, while the closely related P. (N.) mesibovi n.sp. inhabits
northern Tasmania. Australian P. (Nothofagobius) is sister to the Patagonian P (N.) chilensis (Gervais),
which is illustrated based upon new collections. Paralamyctes (Paralamyctes) monteithi n.sp. is
widespread in Queensland; P. (P.) neverneverensis n.sp. is endemic to the Dorrigo region, northern New
South Wales. In addition to these two Australian species, the nominate subgenus occurs in New Zealand,
southern Africa, Madagascar and India, the latter record based on the junior subjective synonym
Triporobius. All new species are included in a key to Australian Lithobiomorpha.

Edgecombe, Gregory D., 2001. Revision of Paralamyctes (Chilopoda: Lithobiomorpha: Henicopidae), with six
new species from eastern Australia. Records of the Australian Museum 53(2): 201-241.

Most native Australian species of lithobiomorph centipedes
belong to the family Henicopidae. Published systematic
work on eastern Australian henicopids was undertaken long
ago (Newport, 1845; Pocock, 1901; Chamberlin, 1920)
based on scant material. Like most of the Australian
chilopod fauna except for the Scolopendridae, the few
henicopid species formalised in the literature are in need of

modern revision, and most of the fauna is undescribed. This
study initiates a systematic survey of Australian Henicopidae
by documenting the genus Paralamyctes.

Pocock (1901) erected Paralamyctes for a new species,
P. spenceri, from Durban, South Africa. Additional species
were subsequently assigned from Argentina (P. andinus
Silvestri, 1903), South Africa (P. asperulus and P. weberi

202 Records of the Australian Museum (2001) Vol. 53

Silvestri, 1903; P laevigatus and P. tabulinus Attems, 1928),
New Zealand [P. validus Archey, 1917 and its junior
synonym P. dubius Archey, 1917 (Archey, 1921, 1937); P.
harrisi Archey, 1922], New Caledonia (P humilis Ribaut,
1923) and Madagascar (P. tridens and P. quadridens
Lawrence, 1960). The Chilean species Henicops chilensis
Gervais in Walckenaer & Gervais, 1847, was recognized
as Paralamyctes by Silvestri (1905), a reassignment
followed by subsequent workers. Attems (1928) and Archey
(1937) revised previously named taxa from South Africa
and New Zealand, respectively, and provided keys to species
of the genus. Further work on the southern African fauna
by Lawrence (1955a,b) recognized P. weberi as a junior
synonym of the widespread P. spenceri, which occurs in
the Cape region, Swaziland-Eastern Lowveld and
throughout KwaZulu-Natal, as well as in Madagascar

(Lawrence, 1960). Chamberlin (1955) removed five species
from Paralamyctes (P. andinus, P. asperulus, P. humilis, P.
laevigatus and P. tabulinus ), assigning them to a new genus
Analamyctes.

Though species from most fragments of Gondwana have
been recognized as Paralamyctes , none have as yet been
reported from Australia. Herein, six new species of this
genus are described from eastern Australia (see Fig. 1 for
distributions). These and other species accepted as members
of Paralamyctes are accommodated in four subgenera that
are diagnosed based on shared derived characters. The
systematic revision recognizes Haasiella Pocock, 1901, as
a clade within Paralamyctes , and Triporobius Silvestri,
1917, as a synonym of the nominate subgenus P.
(Paralamyctes). This revision identifies Paralamyctes in
India, but rejects its occurrence in northern Argentina and

• P. monteithi
o P. cassisi
a P. hornerae
□ P. neverneverensis
o P. ?grayi
☆ P. grayi
+ P. mesibovi

Figure 1. Map of eastern Australia, showing records of Paralamyctes species. Detail of northern New South Wales
indicates sympatric occurrences of P cassisi/P. hornerae and P. cassisi/P. neverneverensis.

Edgecombe: Australian Paralamyctes 203

New Caledonia. Paralamyctes is unknown from Western
Australia. Species of Dichelobius, Henicops and Lamyctes
described by Attems (1911) represent all known henicopid
diversity in the southwest (examination of the Western
Australian Museum collection by the author, March 2001).

Methods and terminology

Each species in this work is illustrated by electron
microscopy, with a particular focus on the largely neglected
mouthparts. This method has proven useful in examining
and figuring characters for henicopid phylogeny. Details
of the mandible in particular emerge as new taxonomic
characters for identifying clades within Paralamyctes. The
electron micrographs in the present paper accompany
illustrations of non-Australian species of Paralamyctes and
other genera of Henicopidae in a cladistic analysis based
on morphological and molecular sequence data (Edgecombe
et al., 2001). Specimens were air-dried, and photographed
on a Leo 435VP using a Robinson backscatter detector.

In all descriptions, length of specimens is measured from
the margin of the head shield to the end of the telson; because
this measure is affected by telescoping, length of the head
shield is cited as a measure of body size (Andersson, 1978).
Nomenclature for the segments of the second maxillary
telopod and the maxillipede follows Borucki (1996: figs.
41 and 57, respectively). The so-called Spingriffel (Rilling,
1968: fig. 17) is called a penis, following Eason (1964).
Terminology applied to the anogenital region is that used
by Eason (1964: figs. 279, 280). Antennal sensilla are
described as by Lewis (1981: figs. 81, 88). In descriptions
of the mandible, the “sickle-shaped bristles” (Attems, 1928)
that comprise the pectinate lamella are referred to as
aciculae, following Chamberlin (1912). The cluster of
bristles on the dorsal edge of the mandible is called a furry
pad, after Attems (1928) (= pulvillus of Crabill, 1960: 15).
Rows of scale- or peg-like structures on the mandibular
teeth ( denticules accessoires of Lawrence, 1960: 93) are
called accessory denticles.

The following abbreviations are used for repositories of
specimens examined:

AM

ANIC

BM/BMNH

CAS

FMHD

IAEP

MCZ

QM

QVMAG

SAM

TMAG

ZMUC

Australian Museum, Sydney
Australian National Insect Collection, Canberra
Department of Entomology, Natural History
Museum, London

California Academy of Sciences, San Francisco
Division of Insects, Field Museum of Natural
History, Chicago

Instituto di Entomologia e Zoologia Agraria,
Universita degli Studi di Napoli Federico II,
Portici

Museum of Comparative Zoology, Harvard

University, Cambridge

Queensland Museum, Brisbane

Queen Victoria Museum and Art Gallery,

Launceston

South African Museum, Cape Town
Tasmanian Museum and Art Gallery, Hobart
Zoological Museum, University of Copenhagen

Abbreviations for collectors are: DKY, D.K. Yeates; GBM,
G.B. Monteith; GC, G. Cassis; GDE, G.D. Edgecombe; GT,
G. Thompson; MRG, M.R. Gray; ZJ, Z. Johanson.

Systematics

Chilopoda Latreille, 1817
Order Lithobiomorpha Pocock, 1902
Family Henicopidae Pocock, 1901
Subfamily Henicopinae Attems, 1914
Tribe Henicopini Chamberlin, 1912

Paralamyctes Pocock, 1901

Type species. Paralamyctes spenceri Pocock, 1901; by
original designation.

Diagnosis. Henicopini with relatively large, bell-shaped
sternite on first maxilla bordered by unsclerotised inner edge
of coxa; suture between coxa and sternite confined to a
narrow contact at posterior edge of maxilla; coxa separate
in front of sternite, median suture lacking. Median furrow
on head shield well impressed, continuous to transverse
suture (= Paralamyctes + Haasiella of previous workers).

Discussion. Attems’ (1928) detailed diagnosis of Paralamyctes
accurately describes most features of new Australian species,
and only a few modifications need noting. The number of
aciculae (“sickle-shaped bristles”) on the mandible was cited
by Attems as 10, but ranges up to 15 in P. grayi n.sp. A
purportedly diagnostic presence of four lobes on the claw
(pretarsus) of the second maxilla is in fact atypical for the
genus, whilst Archey’s (1937) citation of a trifid claw as
diagnostic of this genus does not apply to P. spenceri , any
Australian species, or the New Zealand P. validus. Most
species of Paralamyctes , including representatives of each
of the four subgenera recognized herein (e.g., P monteithi
n.sp., Fig. 6B; P grayi , Fig. 14L; P cassisi n.sp., Fig. 201;
P mesibovi n.sp., Fig. 23L; P chilensis. Fig. 25G) have a
five-part claw, as is also shared by Haasiella trailli
(Archey, 1917). The details of branching of the claw are
identical to those in Lithobiidae (e.g Australobius scabrior
Chamberlin, 1920), with two slender digits interspersed
between three larger digits. As such, a five part claw is
considered to be plesiomorphic, probably a general
character for Lithobiomorpha. Attems listed 2+2 genital
spurs in the female gonopod as diagnostic of Paralamyctes.
Because this number is plesiomorphic for Henicopidae
(indeed, for Lithobiomorpha), the discovery of species with
3+3 spurs (P. cassisi n.sp. and P. mesibovi n.sp.) must be
identified as an apomorphy, and cannot serve to exclude these
species from Paralamyctes if monophyly is to be maintained.

Whether Paralamyctes sensu Attems (1928) and Archey
(1937) is monophyletic is not adequately established. Like
the second maxillary claw and 2+2 gonopod spurs noted
above, most purportedly diagnostic characters are
symplesiomorphies. Abisegmented tarsus on all trunk legs,
perhaps the most obvious character used to define
Paralamyctes , is shared with Analamyctes Chamberlin,
1955, Zygethobiini and Lithobiidae, and may therefore be

204 Records of the Australian Museum (2001) Vol. 53

plesiomorphic relative to the single tarsal segment found
in some legs of other Henicopini (e.g., Lamyctes Meinert,
1868; Lamyctinus Silvestri, 19Q9, HaasiellaPocock, 1901).
Attems (1928) cited a labrum “with numerous ramifying
bristles on the inside” in the diagnosis of Paralamyctes.
Despite a thorough survey by electron microscopy, no aspect
of labral bristling emerges as diagnostic of Paralamyctes
sensu Attems; Henicops Newport, 1844, for example, has a
distribution and branching structure of the labral bristles
completely within the range of Paralamyctes species. Other
symplesiomorphies cited by Attems (1928) are as follow: a
single ocellus (general for Henicopidae); presence of
posterior and lateral borders on the head shield (general for
Lithobiomorpha); stigmata on segments 1, 3, 5, 8, 10, 12
and 14 (general for Henicopini); gonopod segmentation
typical of all Henicopidae; a simple claw on the female
gonopod (general for Henicopidae, probably general for
Lithobiomorpha, and perhaps for Chilopoda); a three-
segmented telopod of the second maxilla (general for
Pleurostigmophora); “simple and branched bristles” on the
tarsus of the second maxilla (general for Lithobiomorpha);
an undivided sternite on the first genital segment (general
for Chilopoda, the division being informative only for
Henicops + Lamyctes)', and a row of coxal pores on legs
12-15 (general for Lithobiomorpha). These characters
cannot serve as synapomorphies within Henicopini.

Attems (1928) noted a first maxillary sternite being
“triangular, partially fused with the coxae” in the diagnosis
of Paralamyctes. The relative enlargement and bell
(“triangular”) shape of the sternite in all species of
Paralamyctes (Figs. 6F, 14E,G, 18F, 20G, 23G) is, however,
shared with Haasiella Pocock, 1901 (=Wailamyctes Archey,
1917; Johns, 1964), and is considered synapomorphic for
this broader group. Outgroup lithobiids (e.g., Lithobius )
possess a smaller, triangular sternite that lies at the end of a
median suture, similar to that of Henicops (Edgecombe et
al., 2001: fig. 8B), Lamyctes and Analamyctes. A large, bell
shaped sternite is observed in Haasiella insularis (=
Wailamyctes munroi : Archey, 1937: pi. 22, fig. 5) as well
as H. trailli (Edgecombe et al., 2001: fig. 8D). In addition
to the size and shape, the sternite of Paralamyctes and
Haasiella is distinctive for the less complete fusion of
its sutures than in other lithobiomorphs, the sternite lying
against an unsclerotised strip along the inner margin of
the coxa. A sutural contact with the coxa is confined to
the posterolateral extent of the sternite. The only
comparable development of the sternite in Henicopidae
is in Esastigmatobius japonicus Silvestri, 1909 (tribe
Zygethobiini) (Edgecombe et al., 2001: fig. 8E). Other
Zygethobiini, such as Zygethobius pontis Chamberlin, 1911,
have the typical, smaller sternite, such that the similarity
between Paralamyctes + Haasiella and Esastigmatobius may
be convergent.

Another character that serves as a synapomorphy for
Paralamyctes + Haasiella is a pronounced median furrow
on the head shield (Figs. 13A, 15H). Attems (1928)
accurately noted “median furrow deep” in his diagnosis of
Paralamyctes, but the same condition is present in
Haasiella. The status of this character as a synapomorphy
is revealed by outgroup comparison with Henicops +

Lamyctes, Zygethobius and Lithobiidae, in which the median
furrow is confined to the anteriormost part of the head shield
or forms a shallow depression rather than a sharp furrow.
The only instances of a median furrow resembling that of
Paralamyctes and Haasiella within the Henicopidae are
within Anopsobiinae (e.g., in Anopsobius), in which the
median furrow likewise extends to the transverse suture,
and in Esastigmatobius japonicus, in which the median
furrow extends about three-quarters the length to the
transverse suture (Edgecombe et al., 2001: fig. 1C).

All species of Paralamyctes described here possess a
cluster of small basiconic sensilla between the coxal process
and telopod of the first maxilla (Fig. 141). Attems (1928:
55) cited this “little cluster of spinules” as a diagnostic
character of Lamyctes, but this must be dismissed based on
its ubiquitous presence in Paralamyctes (e.g., see Archey,
1937: pi. 22, fig. 5 forP. (Haasiella) insularis). The sensilla
cluster does not provide evidence for a Lamyctes!
Paralamyctes clade because it is widely present throughout
Henicopidae and Lithobiidae (pers. obs. for Henicops,
Zygethobius, Anopsobius, Lithobius, Australobius) and thus
appears to be a general character for Lithobiomorpha.

A taxonomic issue of particular biogeographic significance
is the status of Triporobius Silvestri, 1917, a monotypic
genus based on T. newtoni Silvestri, 1917, from India. I
have examined the holotype of this species (IAEP
collection). Silvestri (1917) distinguished Triporobius from
Paralamyctes based on a single character, the restriction of
coxal pores to legs 13-15 in Triporobius. Given that this
state is certainly an autapomorphy for T. newtoni,
Triporobius falls within the cladistic structure of
Paralamyctes. It shares the large, bell-shaped sternite and
long median furrow in the head shield that define the
Paralamyctes + Haasiella clade, and additionally has certain
characters unique to Paralamyctes (Paralamyctes) as defined
below. As such, biogeographic hypotheses that regard
Paralamyctes as absent from India are based on a taxonomic
artifact. Attems’ (1928) statement that Triporobius possesses
eight ocelli on each side of the head is puzzling; the holotype
of T. newtoni bears a single ocellus per side, as described
and figured by Silvestri (1917). This anomaly may be
explained if Attems misread Silvestri’s paper, which has a
description of a species of Archilithobius with eight ocelli
printed just above the figure of Triporobius newtoni.

Some morphological details in Silvestri’s (1917)
illustrations of Triporobius newtoni require emendation.
Silvestri (1917: fig. 5.8) depicted only simple setae on the
tarsus of the telopod of the second maxilla, whereas the
holotype possesses eight or nine plumose setae, and thus
conforms to Paralamyctes and other henicopids. More
significantly, Silvestri (1917: fig. 5.3,5.4) drew the aciculae
with notches along their dorsal sides, but each of these eight
or nine aciculae is actually pinnulate along its distal half.
An arrangement of pinnules along the dorsal side of the
aciculae is elsewhere observed in Paralamyctes species
from South Africa (P. spenceri; Attems, 1928: fig. 448;
Edgecombe et al., 2001: fig. 5E), Madagascar (P.
quadridens; Lawrence, 1960: fig. 28B), New Zealand (P.
harrisi Archey, 1922; Edgecombe et al., 2001: fig. 5F),
Queensland (P. monteithi n.sp.: Fig. 51,J) and New South

Edgecombe: Australian Paralamyctes 205

Wales (P. neverneverensis n.sp.: Fig. 11A,J) (see discussion
of that subgenus for evidence that pinnules confined to the
dorsal side of the aciculae is a synapomorphy for this group).

Several species that were originally assigned to
Paralamyctes were reassigned to the new genus Analamyctes
by Chamberlin (1955). Chamberlin distinguished the two
genera based on a single character, the number of antennal
articles being fixed at 19-20 in Paralamyctes versus 23-43
in Analamyctes. Although antennal segmentation is
taxonomically useful at some levels in Henicopini,
Chamberlin’s classification is incongruent with a larger
body of character evidence that retains some supposed
Analamyctes within a monophyletic Paralamyctes while
grouping typical Analamyctes with Lamyctes, Lamyctinus
and Henicops. The reliability of Chamberlin’s two character
states for antennal segmentation (19-20 versus 23-43) is
weakened by new Australian species, some of which have
article counts that fall between these two states (e.g., 20-
21 in P. neverneverensis, 20-22 in P. hornerae ), and others
that range between both states (18-25 in P. monteithi). I
have examined two specimens that represent two named
species of Analamyctes, the type species, A. tucumanus
Chamberlin, 1955, and A. andinus (Silvestri, 1903). BM
1903.7.2.0 (Fig. 2), a male labelled as “Tucuman 450 m”,
represents A. tucumanus, the holotype being from Cerro
San Xavier, Tucuman, Argentina. The type locality for A.
andinus is Punta de Vacas, Mendoza, Argentina; a specimen
from 25 km north of Villaviciencio, Mendoza (ZMUC
collection: Fig. 3) is assigned to this species. Based on these
specimens, Analamyctes lacks both synapomorphies of
Paralamyctes + Haasiella\ it has no median furrow on the
head shield (Fig. 2), and the sternite of the first maxilla is
small, lying behind a median suture. Antennal segmentation
with short, ring-like articles occurring in pairs between
series of longer articles in both species (Fig. 2) resembles
Lamyctes (Edgecombe et al., 2001: fig. ID) and Henicops.
The subtriangular shape and dentition of the maxillipede
coxosternite suggest closest affinities to Lamyctes and
Lamyctinus ; a minute, conical seta set in a prominent socket
at the anterolateral edge of the dental margin in both A.
tucumanus and A. andinus (Edgecombe et al., 2001: fig.
3J) is certainly homologous with the “pseudoporodont” of
Lamyctinus (Negrea & Matic, 1996: fig. 3; Edgecombe et
al., 2001: fig. 3K) and some species of Lamyctes (“ectodont”
of Chamberlin, 1955; “porodont” of Zalesskaja, 1994). A
porodont is not differentiated in Paralamyctes. Mandibular
morphology, examined for A. andinus (Fig. 3), also most
closely resembles Lamyctes, Lamyctinus and Henicops.
Detailed similarities include: bipinnulate aciculae in which
the pinnules are relatively long (Fig. 3B); an abrupt
differentiation of the branching bristles along the length of
the fringe, with multifurcating bristles splaying from scale¬
like bases on the dorsal part of the fringe, and pectinate
bristles on the ventral part of the fringe (Fig. 3C); accessory
denticles near the fringe of branching bristles on the dorsal
part of the mandible being developed as flattened,
multifurcating scales (Fig. 3D) rather than simple conical
elements as in other Henicopinae. Also similar to Lamyctes
and Lamyctinus is the lack of posterolateral projections on
any tergites in A. andinus and A. tucumanus (Fig. 2).

Figure 2. Analamyctes tucumanus Chamberlin, 1955. 6 BM
1903.7.2.0, Tucuman, Argentina. Scale 1 mm.

At least one of the South African species reassigned by
Chamberlin (1955) from Paralamyctes to Analamyctes, P.
asperulus, is more appropriately retained in Paralamyctes
[see discussion of P. (Paralamyctes) below]. The status of
two others, P. laevigatus and P. tabulinus is uncertain; no
specimens have been illustrated and descriptions are
imprecise. However, maxillipede dentition of these species
(6+6 teeth in both fide Attems, 1928) is more consistent

206 Records of the Australian Museum (2001) Vol. 53

Figure 3. Analamyctes andinus (Silvestri, 1903). Scanning electron micrographs of mandible. ZMUC, sex
undetermined (genital region damaged), 25 km N of Villaviciencio, Mendoza, Argentina. A, gnathal lobe; B, aciculae;
C, fringe of branching bristles; D, teeth and furry pad. Scales 5 pm except A, 10 pm.

with membership in Paralamyctes than Analamyctes, and
P. laevigatas was described by Attems (1928) as having a
deep median furrow on the head shield (synapomorphy of
Paralamyctes ). Of the non-Argentine species referred to
Analamyctes by Chamberlin (1955), only the reassignment
of Paralamyctes humilis Ribaut, 1923, from New Caledonia
is accepted here. This placement accounts for its triangular,
few-toothed maxillipede coxosternite, large number of
antennal articles, and weak definition of an articulation
between tarsomeres. With the exclusion of A. andinus and
A. humilis from Paralamyctes, the occurrence of the latter
genus in South America is limited to P. chilensis in northern
Patagonia (Chile and Argentina), and Paralamyctes is not
known to occur in New Caledonia.

Paralamyctes (Paralamyctes) Pocock, 1901
= Triporobius Silvestri, 1917

Diagnosis. Paralamyctes with mandibular aciculae bearing
elongate pinnules along dorsal side of acicula only;
accessory denticle field intergrading with furry pad; antenna
with relatively few (typically 17-20) elongate, tubular
articles; articulations between tarsomeres strong; longi¬
tudinal median furrow on sternites well defined.

Assigned species. Paralamyctes asperulus Silvestri, 1903;
Paralamyctes weberi Silvestri, 1903; Triporobius newtoni
Silvestri, 1917; Paralamyctes harrisi Archey, 1922;
Paralamyctes quadridens Lawrence, 1960; Paralamyctes
tridens Lawrence, 1960; Paralamyctes (Paralamyctes)
monteithi n.sp.; Paralamyctes (Paralamyctes) never-
neverensis n.sp.

Distribution. South Africa (KwaZulu-Natal, Western Cape
Province, Eastern Lowveld, Northern Province), Swaziland,
Madagascar, southern India, Australia (Queensland,
northern New South Wales), New Zealand (North Island).

Discussion. Several apparently apomorphic characters are
shared between Paralamyctes monteithi from Queensland,
P. neverneverensis from the Dorrigo region of New South
Wales, P. newtoni from India, P. spenceri and P. weberi from
southern Africa and P. harrisi from New Zealand. These
species all have pinnules on the mandibular aciculae
branching from a single side (dorsally). The same
distribution of the aciculae was illustrated by Lawrence
(1960: fig. 28B) in P. quadridens from Madagascar. Other
species of Paralamyctes have either bipinnulate aciculae
(branching from both sides), for example P (Nothofagobius)
chilensis (Fig. 251,K) and P (Haasiella) trailli (Edgecombe

Edgecombe: Australian Paralamyctes 207

etal., 2001: fig. 5B), or simple (non-pinnulate) aciculae, as
in P. validus (Edgecombe et al., 2001: fig. 5G), P. grayi
(Fig. 13F) and P hornerae n.sp. (Fig. 18K). The bipinnulate
condition is regarded as plesiomorphic, based on the
presence of bipinnulate aciculae in Henicops (Edgecombe
et al., 2001: fig. 5C), Lamyctes (Edgecombe et al., 2001:
fig. 5A), Lamyctinus, Analamyctes (Fig. 3B), Zygethobiini
(. Esastigmatobius, Zygethobius', Edgecombe et al., 2001:
fig. 5H), Anopsobiinae ( Anopsobius ) and Fithobiidae
C Lithobius, Australobius, Bothropolys [Edgecombe et al.,
2001: fig. 5D]). Species with pinnules along only one side
of the aciculae, P (Paralamyctes), also share an inter¬
gradation between the accessory denticles on the mandible
and the furry pad, the distal projections of the furry pad
merely being progressively elongated (Fig. 5F,M). In other
Henicopini and in the Fithobiidae, the furry pad is well
differentiated from the accessory denticles. This involves
either an abrupt elongation of the elements of the furry
pad (Figs. 13E, 15M), or an intervening smooth region
that lacks bristle-like morphology (Fig. 20E,F).
Paralamyctes (Paralamyctes) as defined by morpho¬
logical (largely mandibular) characters is also identified
as a monophyletic group based upon molecular sequence
data (Edgecombe et al., 2001).

Species of P. (Paralamyctes) share strong joints between
the tarsomeres on all legs, with a small condyle commonly
developed at the articulation dorsally. The status of this
character as a synapomorphy [relative to the weak
articulations in P. (Thingathinga )] is uncertain. Most
lithobiids have strong articulations, so this state is possibly
plesiomorphic for Paralamyctes. Ambiguity is caused by
the inapplicability of the character in some relevant
outgroups (e.g., Lamyctes and Anopsobiinae, in which the
tarsi lack articulations). Another character shared by all
species of P. (Paralamyctes) by which they are distinguished
from other Paralamyctes is the elongation of the
longitudinal median furrow on the sternites. This is especially
pronounced on the posterior sternites, running along most of
the length of the stemite (Fig. 6F). The character is variably
developed in outgroups, and its polarity is uncertain.

Among South African species referred to Paralamyctes
by Attems (1928), I have examined P. spenceri (Natal
Museum collections), its alleged junior synonym P. weberi
(including its holotype), and the holotype of P asperulus
Silvestri, 1903 (IAEP collection). Fawrence’s (1955b)
synonymy of P. weberi is unconvincing. Fawrence cited
little material from the Cape region and claimed to “have
never seen a specimen which could be ascribed to weberi ”
(1955b: 18). I have observed material from Table Mountain
(MCZ 28614) that conforms to the holotype of P. weberi in
having four blunt, widely-separated teeth on each margin
of the maxillipede coxosternite and having a distal spinose
projection on the tibia of leg 14, and follow Attems (1928)
in considering it and P. spenceri as distinct species. Contrary
to Chamberlin’s (1955) reassignment of P. asperulus to
Analamyctes, this species can be confirmed as a member of
Paralamyctes based on its deep median furrow on the head
shield and the large, bell-shaped sternite of the first maxilla.
An elongate distal part of the maxillipede tarsungulum and
knob-like teeth on the dental margin of the coxosternite

indicate membership in P. (Paralamyctes), P. (Haasiella)
or ( Thingathinga ). The shape of the maxillipede coxo¬
sternite resembles that of P. spenceri, and the strong
articulations defining the tarsomeres on all legs are also
like that species (but strikingly different from Analamyctes,
in which tarsal articulation are faint). Paralamyctes
asperulus is apparently referable to P. (Paralamyctes).
Antennal articles are mostly elongate, resembling species
of that subgenus. The description by Fawrence (I960) is
sufficient to indicate that P. quadridens from Madagascar
is a member of P. (Paralamyctes). Two males of this species
(FMHD 94-60) from Reserve Anjanaharibe-Sud, Befingotra,
confirm the single, dorsal row of pinnules on the aciculae,
and the accessory denticles intergrade with the furry pad
as in other P. (Paralamyctes). The other endemic
Malagasy species, P. tridens Fawrence, 1960, is
provisionally assigned to P. (Paralamyctes) based on
Fawrence’s (1960) assertion that it bears very close
resemblance to P. (P.) spenceri, and description of a single
row of eight or nine thick “teeth” (pinnules) on the distal
part of the aciculae (Fawrence, 1960: 93).

Paralamyctes (Paralamyctes) monteithi n.sp.

Figs. 4-7, 8A,B, 9A

Etymology. For Geoff Monteith, Queensland Museum,
whose collections in the Wet Tropics include much of the
material of this species.

Diagnosis. Paralamyctes (Paralamyctes) usually with 20
long, tubular antennal articles; body relatively elongate, long
tergites extended (e.g., TT3 and 5 as long as wide); 3-6
(very rarely 7) small teeth on dental margin of maxillipede
coxosternite; cephalic pleurite constricted just behind
Tomosvary organ, which lies on ventral margin of head
shield; groove in accessory denticle field lacking on
mandibular teeth; 3-6 coxal pores (usually 4-6); posterior
margin of sternite 15 of male typically rounded.

Type material. Holotype QM S42683, 6 (Fig. 4A,C),
south end of Bluewater Range, 45 km NW of Townsville,
Queensland, 19°ir51"S 146°24’16"E, 750 m, GBM, GT
and S. Hamlet, 7 Dec 1986-16 Feb 1987, pitfall, rainforest.
Paratypes: QM S42682,1 9, QM S42684, \6, Bluewater
Range, same locality as holotype, 700-800 m, GBM, GT
and S. Hamlet, 6-8 Dec 1986.

Other material. NE Queensland (Wet Tropics): QM S42652,
26 6, Cardwell Gap, 18°31'53"S 146°H'E, R. Raven, P. and E.
Lawless andM. Shaw, 24 Sep-25 Nov 1992, pitfall; QM S42685,
2 6 6, 1 9, Cardwell Range, Upper Broadwater Creek Valley,
18°19'15"S 145°58'34"E, 750 m, GBM, GT and S. Hamlet, 18
Dec 1986-14 Jan 1987, pitfall, rainforest; QM S42680 1 6, S42681
1$, Mt Elliot, summit, 19°29'29"S 146°58'00"E, 1150 m, A.
Graham, Jan-26 Mar 1991, pitfall and intercepts, fern glade; QM
S42686, 1$, Emerald Creek, Lamb Range, 17°05'06"S
145°36'06"E, 950 m, GBM, DKY and GT, 11 Oct 1982, sieved
litter, rainforest; QM S42691, 1 9, Douglas Creek, Lamb Range,
17°06'51"S 145°37'26"E, 900 m, 12 Oct 1982, GBM, DKY and
GT, 12 Oct 1982, Pyrethrum, rainforest; QM S42687, 19, Mt
Hypipamee NP, tower near the Crater, 17°27'23"S 145°29T2"E,

208 Records of the Australian Museum (2001) Vol. 53

1230 m, GBM and J. Hasenpusch, 7 Mar-15 May 1995, intercept
trap; QM S42689, lc?, Mt Misery Rd, 15°52'39"S 145°12'58"E,
730 m, 6 Dec 1990-17 Jan 1991, Queensland Museum and
ANZSES, pitfall; QM S42690, 2 c? 6 and 1 9, North Bell Peak,
Malbon Thompson Range, 1000 m, GBM and GT, 20-22 Nov
1990, pitfall; QM S42692, 16, Mt Tyson, 2 km W of Tully,
17°55'35"S 145°54'34"E, 650 m, DKY, 7 May 1983, sieved litter,
rainforest; QM S42693,1 6, Malaan SF, 17°35'30"S 145°36'45"E,
R. Raven, P. and E. Lawless and M. Shaw, 25 Jul-26 Nov 1992,
pitfall; QM S42694, 1 $, Mt Fisher, Kjellberg Rd, 7 km SW of
Millaa Millaa, 17°32'34"S 145°33'31”E, 1000 m, GBM and DKY,

3 May 1983, sieved litter, rainforest; QM S42688, 1 6, Mt Bartle
Frere, 0.5 km N of South peak, 1500 m, 6-8 Nov 1981, Earthwatch/
Queensland Museum; QM S42695, 2 6 6, summit, Mt Bellenden
Ker, 17°15'40"S 145°51'25"E, 1560 m, GBM, H. Janetzki and D.
Cook, 8 Oct 1991; QM S42696, 16, S42704, 1 $, Bellenden Ker
Range, Cableway base station, 100 m, Earth watch/Queensland
Museum, 17 Oct-7 Nov 1981, pitfall; QM S42697, 1 9, S42699,
lc?, Bellenden Ker Range, Cable Tower 5, 17°16'32"S
145°53'00"E, 532 m, Earthwatch/Queensland Museum, 1-7 Nov
and 17-24 Oct 1981; QM S42700, 16, S42705, 1 9, summit TV
station, Mt Bellenden Ker, 1560 m, S. Montague, Apr-Oct 1982,
pitfall (S42700), GBM and DKY, 29 Apr-2 May 1983, moss on
ground (S42705), rainforest; QM S42701, 26 6, several
immatures, S42703, 2c? 6, 3 9 9 (Figs. 4B, 6A-C,G,J), summit,
Mt Bellenden Ker, 1560 m, Earthwatch/Queensland Museum, 17
Oct-5 Nov 1981, pitfall; QM S42706, lc?, S45147, larva,
Bellenden Ker Range, Cable Tower 3, 1054 m, Earthwatch/
Queensland Museum, 17-31 Oct 1981; AM KS 57900, 2c? c?, 19,
Black Mountain Rd, 28.1 km NW of Kuranda, 16°40'25"S
145°30'08"E, GDE, 28 Apr 1998, sifted litter, rainforest; AM KS
57901, 19 (DNA voucher specimen), KS 57902, 2 9 9, Mt
Hypipamee NP, The Crater, 17°25'29"S 145°29'00"E, GDE and
G. Milledge, 25 Apr-2 May 1998, sifted litter and pitfall, rainforest;
AM KS 57903, 1 c?, Danbulla SF, 1.9 km E of Mobo Creek Crater,
GDE, 27 Apr 1998, sifted litter, rainforest; ANIC, 1 9, Kuranda,
16°45'S 145°35'E, 430 m, R.W. Taylor, 29 Jul 1977, rainforest;
ANIC, lc?, Tully Falls SF, 17°46'S 145°33'E, A. Walford-Huggins,

4 Oct 1978, rainforest; ANIC, lc?, Koombooloomba, 17°50'S
146°36'E, 750 m, R.W. Taylor and J. Feehan, 4 Jul 1971, Berlese,
wet sclerophyll; ANIC, 69 9, Mission Beach, 17°50'S 146°06E,
10m, M. Cemak, 2 Sep-1 Oct 1996; ANIC, 2c? c?, 29 9, Mission
Beach, 17°52'S 146°04'E, 40 m, M. Cemak, 2 Sep-4 Nov 1996,
pitfall; ANIC, 19,5 kmWof Paluma, 19°01'S 146°10’E, ca. 950
m, R.W. Taylor and J. Feehan, 13 Jul 1971, Berlese, rainforest;
CAS, lc?, Tinaroo Lake near Kairi, 700 m, 8 Nov 1962, E.S.
Ross and D.Q. Cavagnaro; CAS, 19,1 mile NE of Ravenshoe,
975 m, 7 Nov 1962, E.S. Ross and D.Q. Cavagnaro; MCZ 34824,
1 6, Mt Spurgeon, 3500-4000 ft, Jul 1932, G.H. Curry.

Central eastern Queensland (Eungella/Rockhampton): AM
KS 57904, lc? (Figs. 7A, 8A,B), KS 57905, 1 9 (Fig. 7B), KS
57906, 1 9 (Figs. 7C, 9A), KS 57907, 1 9 (Figs. 5A-J, 6H,I,K-
M), KS 57908, 19 (DNA voucher specimen), KS 57909, lc?,
3 9 9, Eungella NP, Dalrymple Rd, 1.7 km NE of Snake Rd, 21 °04'S
148°34'30"E, GDE, S. Davis and G. Milledge, 18 and 21 Apr 1998,
rainforest litter; QM S45184, 1 c?, Eungella, Schoolhouse, 21°08'S
145°29E, R. Raven and J. Gallon, 13 Feb 1986, sieved litter, rainforest;
ANIC, larva, presumed fourth larval stadium (12 legs plus three limb-
buds), Cammoo Caves, 23°10'S 150°28'E, R.W. Taylor and A. Weir,
25 Oct 1976, dense low closed forest.

SE Queensland—Kenilworth/Imbil region: AM KS 57910,
19 (Figs. 5K-M, 6D-F), KS 57911,19 (DNA voucher specimen),
Kenilworth SF, Sunday Creek Rd, 9.8 km W of Charlie Moreland
Park, 26°40'H"S 152°36'35"E, GDE, S. Davis and G. Milledge,
6-7 May 1998, sifted litter, wet sclerophyll; QM S42679, lc?,

Amamoor Creek via Imbil, 26°21'45"S 152°37'56"E, 91 m, GBM
and S.R. Monteith, 1975-1976, pitfall, rainforest; QM S42678,
lc? (Fig. 7D), Cold Creek via Imbil, 26°27'20"S 152°37'26”E,
122 m, GBM and S.R. Monteith, 1974-1975, pitfall, rainforest;
QM S42677, 19, Deer Reserve SF via Kilcoy, 26°59'29 M S
152°28'33"E, 457 m, GBM and S.R. Monteith, 1974-1975, pitfall,
rainforest. Lamington region: QM S42669, 1 9, Lamington NP,
Mt Bithongabel, 28°15'55"S 153°10T3"E, 1160 m, GBM and S.R.
Monteith, 1975-1976, pitfall, rainforest; QM S42675, 19,
Repeater Station, Springbrook, 28°14'23"S 153°15'58"E, 1000 m,
GBM and S.R. Monteith, 1975-1975, pitfall, rainforest; QM
S42676, 26 6, Lamington NP, Nagarijoon, V.E. Davies and R.
Raven, 5 Apr 1976.

Description. Length up to 27 mm; length of head shield up
to 2.4 mm. Colour (based on specimens in absolute ethanol):
antennae dark orange; head shield orange with chestnut
mottled network; tergites pale orange with dark mottling
concentrated in longitudinal median band and near margins;
maxillipedes bright orange; sternites yellow/pale orange,
becoming darker orange with more abundant chestnut
mottling posteriorly; prefemur to tibia pale orange or yellow
with purple mottling; tarsi orange; genital sternite and
gonopods orange.

Head shield smooth. Frontal margin with strong median
notch; posterior margin transverse. Median furrow extends
back to transverse suture, length about 30% that of head
shield, deep throughout. Antenna extends back to tergite 4
or 5; 18-25 antennal articles, overwhelming majority of
specimens with 20; all articles substantially longer than
wide; basal two articles only slightly enlarged relative to
adjacent ones (Fig. 6A), with gradational rather than abrupt
changes in setation (increased abundance of short trichoid
sensilla from about fifth article, fewer longer setae).
Posterior side of polygonal area surrounding hair socket of
trichoid sensilla on antenna raised as a semicircular stop
( Kragen ) (Fig. 8B); two or three finger-shaped thin-walled
basiconic sensilla at anterior edge of antennal articles
adjacent to band of arthrodial membrane (Fig. 8A,B).
Ocellus whitish, moderately domed. Cephalic pleurite
usually inclined, narrowed to a slender band behind
Tomosvary organ (Figs. 5H, 6D), which lies on ventral
margin of head immediately posteroventral to ocellus (Fig.
5G), encircled by sutures that define anterior limits of
pleurite; Tomosvary organ moderately large, elliptical.

Tergites faintly to distinctly wrinkled, more so on
posterior segments. T1 generally trapeziform, considerably
smaller than T3, slightly narrower than head shield (Fig.
4A), about 75% width of widest tergite (T8), posterior
angles rounded and posterior margin transverse; lateral
borders subparallel in TT3 and 5, posterior angles rounded,
posterior borders transverse or T5 faintly concave; TT1, 3
and 5 bordered posteriorly; lateral borders of TT7-14
convex; border of T7 incomplete posteriorly, posterior
margin with shallow transverse median embayment,
posterior angle rounded; TT8-14 (variably T6) bordered
laterally; posterior margin of T8 gently concave, posterior
angle rounded, TT10 and 12 elongate, posterior borders
weakly to gently concave, posterior angles form obtuse,
blunt corners; TT9, 11 and 13 embayed posteriorly, with
largely transverse median extent, posterior angle of T9 blunt,
TT11 and 13 sharp but not toothed; T14 longer than wide,

Edgecombe: Australian Paralamyctes 209

Figure 4. Paralamyctes (Paralamyctes) monteithi n.sp. A,C, holotype S QM S42683, south end of Bluewater
Range, northeastern Queensland; C, terminal segments and gonopods; scales 2 mm, 0.5 mm. B, $ QM S42703,
terminal segments and gonopods, summit of Mt Bellenden Ker, northeastern Queensland; scale 0.5 mm.

posterior border usually gently concave in both sexes, with
blunt posterior angles (Fig. 6M); in males from Mt
Bellenden Ker, posterior margin of TT12 and 14 transverse
to weakly convex and posterior angles rounded, posterior
margin of T13 gently concave. Short, slender setae along

lateral borders of all tergites; anterior tergites with few
additional setae, concentrated anterolaterally; short setae
evenly scattered over surface of posterior tergites (Fig. 6M).

Small transverse seta projects medially from pit in labral
sidepiece (Fig. 5B). Labral margin with rounded shoulder

210 Records of the Australian Museum (2001) Vol. 53

Figure 5. Paralamyctes (Paralamyctes) monteithi n.sp. Scanning electron micrographs. A-J, $ AM KS 57907, Eungella NP, central
eastern Queensland. A, ventral view of head, scale 200 pm; B, clypeus and labrum, scale 50 pm; C, branching bristles on labral margin,
scale 10 pm; D, gnathal lobe of mandible, scale 50 pm; E, fringe of branching bristles on mandible, scale 20 pm; F, mandibular teeth and
furry pad, scale 20 pm; G, cephalic pleurite, scale 100 pm; H, Tomosvary organ, scale 20 pm; I,J, mandibular aciculae, scales 20 pm. K-
M, $ AM KS 57910, Kenilworth SF, southeastern Queensland. K, ventral part of mandible, scale 20 pm; L, mandibular teeth, scale 20
pm; M, accessory denticles and furry pad on mandible, scale 10 pm.

beside midpiece, with pronounced break in curvature where
dense, long fringe of branching bristles overhangs margin;
bristles branch as irregular, elongate bifurcations or
multifurcations (Fig. 5C).

Maxillipede coxosternite trapezoidal to subsemicircular,
dental margin broad, each half gently convex (Figs. 6G,H,

7); median notch lacking; teeth small, blunt bulbs, ranging
from 3+3 to 6+6 (one specimen with 7+7), set off by furrow
that parallels dental margin. Coxosternite bearing relatively
few large, scattered setae, usually with distinctly denser
setation behind dental margin and anterolaterally (Fig. 6H).
Tarsungulum with long, slender pretarsal section (Fig. 6G).

Edgecombe: Australian Paralamyctes 211

Figure 6. Paralamyctes (Paralamyctes) monteithi n.sp. Scanning electron micrographs. A-C,G,J, $ QM S42703, summit of Mt Bellenden
Ker, northeastern Queensland. A, proximal part of antenna, scale 200 pm; B, pretarsus of second maxillary telopod, scale 10 pm; C,
coxal processes of first maxillae, scale 20 pm; G, maxillipede, scale 400 pm; J, coxa and telopod of second maxilla, scale 200 pm. D-
F, $ AM KS 57910, Kenilworth SF, southeastern Queensland. D, Tomosvary organ, scale 20 p; E, dental margin of maxillipede
coxosternite, scale 50 pm; F, first maxillae, scale 50 pm. H,I,K-M, $ AM KS 57907, Eungella NP, central eastern Queensland. H,
dental margin of maxillipede coxosternite, scale 100 pm; I, gonopods, scale 90 pm; K, distal part of telopod of second maxilla, scale 50
pm; F, ventral view of terminal segments, scale 300 pm; M, dorsal view of terminal segments, scale 300 pm.

Setae on forcipule concentrated on inner part of trochantero-
prefemur, femur, tibia and tarsal part of tarsungulum, these
setae relatively long, of fairly uniform diameter.

Mandible: Four paired teeth (Fig. 5D). Ten aciculae; up
to 10 finger-shaped pinnules aligned against each other on
distal third of dorsal side of each acicula (Fig. 51,J). Fringe

of branching bristles skirts aciculae, evenly shortening
dorsally to very short fringe beneath furry pad (Fig. 5F,M);
bristles narrow-based, evenly branching along entire length
(Fig. 5E), bifurcate or pauciramous distally. Accessory
denticle field without grooves between rows of denticles
or at margin of denticle field (Fig. 5F,L); largest accessory

212 Records of the Australian Museum (2001) Vol. 53

A

D

Figure 7. Paralamyctes (Paralamyctes) monteithi n.sp. Maxillipede coxosternite, showing variation in number of teeth on dental
margin. A-C, Eungella NP, Dalrymple Rd, central eastern Queensland. A, S AM KS 57904, 3+3 teeth; B, $ AM KS 57905, 4+4 teeth;
C, $ AM KS 57906, 5+5 teeth. D, S QM S42678, 6+6 teeth, Cold Creek via Imbil, southeastern Queensland. All scales 0.25 mm.

denticles scale-like, flattened, grading into rod-shaped scales
then small elongate scales near fringe of branching bristles
(Fig. 5M); slender, rod-like accessory denticles grade into
furry pad; furry pad with many elongate, simple bristles,
some bifid or pauciramous bristles.

First maxilla: Bell-shaped sternite of typical size for
genus (Fig. 6F), anterolateral margins set in arthrodial
membrane, posterolateral edge delimited from coxa by well-
defined suture. Coxal process triangular, with 8-12 simple
setae on anteromedial edge (Fig. 6C), this cluster usually
separated from a few setae aligned on inner edge of coxal
process. Cluster of minute, barb-like basiconic sensilla
between coxal process and telopod. Distal article of telopod
with two rows of long, plumose setae along inner margin;
branches developed along distal half of these setae; short,
simple setae on membranous strip alongside inner margin;
main, more sclerotised field of distal article of telopod
largely barren of setae, a few along outer margin.

Second maxilla: Sternite small, fused with coxae. Band
of short setae along anterior part of coxa not prominent
(Fig. 6J). Tarsus bearing numerous simple setae on outer
surface, dense cluster of plumose setae on membranous patch
on inner surface (Fig. 6K); branches on plumose setae mostly
confined to distal half. Claw small, composed of five digits,
median and outer pair long, thick, with shorter, needle-like
digit between median and each outer digit (Fig. 6B).

Strong, pointed distal projections with sclerotised tips
on tibiae of legs 1-14. Setae on legs of relatively uniform
diameter, gracile. Premur with few short setae on anterior
and posterior faces, numerous setae of varied lengths
dorsally, some longer setae on ventral edge of prefemur;
prefemur lacking ventral recess; longest setae on femur and
tibia equal to or slightly shorter than longest on prefemur;
a few slightly to indistinctly thickened setae encircling distal
margin of femur; seta at ventrodistal edge of tibia
distinguished by slightly darker (orange-brown) pigment¬
ation but not significantly thickened; tarsal setae uniformly
slender, of fairly even length. Articulation between
tarsomeres strong on all legs, with small condyle dorsally.
Distitarsus about 45% length of basitarsus on anal leg; anal
leg basitarsus 9.5-11 times longer than broad (Fig. 9A).
Pretarsal claws curved; pair of accessory claws symmetrical,
about 25% length of main claw (Fig. 9A).

Longitudinal median furrow shallow on anterior half of
sternites, extending to or behind midlength on sternites 14
and 15 (Fig. 6L), sometimes present along entirety of
sternite. Short setae scattered across anterior third of
sternites and along lateral margin, with strongest marginal
setae anterolaterally; sternal setation similarly developed
along length of trunk.

Coxal pores round or transversely ovate (Fig. 4B,C),
commonly 4555/4555 or 5666/5666 in females, maximum

Edgecombe: Australian Paralamyctes 213

Figure 8. Antennal sensilla. A,B, Paralamyctes (Paralamyctes) monteithi n.sp. AM KS 57904, 6, Eungella NP, Queensland. A, nearly
dorsal view of second- to fourth-last antennal articles, showing trichoid sensilla, pair of thin-walled basiconic sensilla, scale 20 pm; B,
seventh-last antennal article, anterior to left, showing trichoid sensilla with stop ( Kragen ) basally, three thin-walled basiconic sensilla,
scale 5 pm. C,D, Paralamyctes (Thingathinga) grayi n.sp. 6 AM KS 58466, Mt Barrengarry, NSW. C, fourth- and fifth-last antennal
articles, showing pair of thin-walled basiconic sensilla at left, cluster of three thick-walled basiconic sensilla at right, scale 5 pm; D,
detail of thin-walled basiconic sensilla, scale 5 pm.

6666/6666, minimum 3444/3444; males most commonly
4444/4444, maximum 5555/5555, not uncommonly 3333/
3333. Coxal pore field delimited from anteroventral face
of coxa by rounded edge in most specimens, or with variably
developed fold, when present, distal coxal pores may be
partly concealed in ventral view by fold (Fig. 6L). Anal
pores large in both sexes (Fig. 4C).

Male (Fig. 4C): Sternite of segment 15 rounded
posteriorly/posterolaterally. Sternite of first genital segment
small, undivided, with transverse posterior margin, bearing
numerous short, evenly scattered setae. Gonopod of three
articles and tapering, seta-like terminal process, the three
articles each bearing a few short setae; maximum length of
penis equal to that of gonopod exclusive of terminal process,
in some specimens only as long as first article of gonopod.

Female (Fig. 4B): Sternite of segment 15 transverse or
convex posteromedially. Tergites of first genital segment
and telson usually well sclerotised. Sternite of first genital
segment large, posterior margin concave between condyles
of gonopods, surface evenly scattered with short setae. First
article of gonopod bearing two conical spurs of equal size;
spurs gently curved such that tip points up; first and second
articles of gonopod with mix of short and moderately long

setae (Fig. 61); third article with a few small setae. Claw
undivided.

Discussion. Comparison with the only other Australian
species of P. (Paralamyctes), P. (P.) neverneverensis n.sp.,
is made in discussion of that species below.

Paralamyctes (Paralamyctes) monteithi appears to be
most closely related to P. (P.) harrisi Archey, 1922, from
North Island, New Zealand. These taxa are particularly
similar in the elongation of the antennomeres (all being
longer than wide versus some subequally-proportioned
antennomeres in other species of Paralamyctes, e.g., Figs.
13B, 15C), the identical structure of the mandibular
aciculae, a much elongated pretarsal component of the
maxillipede tarsungulum, particularly dense setation on the
inner part of the forcipule (e.g., numerous long setae on the
tibia and femur; Fig. 6G), and the shape of the maxillipede
coxosternite. In both species, the dental margin of the
coxosternite is wide and usually biconvex, and the outer
part of the margin is conspicuously devoid of teeth (Fig.
7). Paralamyctes (P.) harrisi also resembles P. monteithi in
having the Tomosvary organ shifted lateral to the main
surface of the cephalic pleurite, such that the organ comes

214 Records of the Australian Museum (2001) Vol. 53

Figure 9. Anal legs of Paralamyctes (Paralamyctes), with details of pretarsi. A, P. (P.) monteithi n.sp., $ AM KS 57906, Eungella
NP, Queensland. B, P. (P.) neverneverensis n.sp., 8 AM KS 57955, Dorrigo NP, NSW. C, P. (P.) harrisi Archey, 1922, 9, CAS,
White Pine Bush, Hawkes Bay, New Zealand. D, P. (P.) spenceri Pocock, 1901, 8 , Natal Museum, Town Bush, Pietermaritzburg,
KwaZulu-Natal, South Africa. Scales 1 mm except B, 0.5 mm.

to lie on the ventral margin of the head (Fig. 5A,G). In other
species of Paralamyctes and in other Henicopidae, the
Tomosvary organ is situated on the flat surface of the
pleurite (Figs. 11G, 14J, 20L, 211, 25M). These characters
provide evidence for a sister species relationship between
the Queensland and New Zealand taxa. Paralamyctes (P.)
monteithi is distinguished from P (P.) harrisi by usually
having more antennal articles (typically 17 in P. harrisi,
but occasionally as many as 20), having a less setose

anterolateral part of the maxillipede coxosternite, and by
having a constriction of the cephalic pleurite behind the
Tomosvary organ (see discussion below).

Paralamyctes monteithi has a broad geographic range,
but it has not proven possible to consistently differentiate
populations from different parts of the range based on
external morphology. All Queensland samples share a
typical development of 20 elongate antennomeres and have
the Tomosvary organ at the ventral margin of the head. The

Edgecombe: Australian Paralamyctes 215

cephalic pleurite that bears the Tomosvary organ is
relatively steeply inclined, whereas it more consistently lies
horizontally in other species of Paralamyctes. The suture
that delimits the cephalic pleurite displays a most peculiar
course around the Tomosvary organ, the pleurite narrowing
sharply behind the organ (Figs. 5G,H, 6D). An elongate
body, expressed by the relative lengthening of the head
shield and long tergites, is developed throughout the range
of the species. These characters indicate that all of the
Queensland material is closely related (a monophyletic group).

Within this widespread species, the most distinctive
variant is observed from the summits of Mt Bellenden Ker
and Mt Bartle Frere, in which the posterior margins/angles
of tergites 12-14 exhibit a measure of sexual dimorphism
that is otherwise not observed in the species (or in other
Paralamyctes , in which dimorphism in the tergites is usually
limited to the tergite of the intermediate segment being
relatively wider in females). Males on Bellenden Ker have
less concave posterior margins of TT12-14 and more
rounded posterior angles than do females, or males from
other localities. The Bellenden Ker sample has either 3+3
or 4+4 teeth on the dental margin of the maxillipede
coxosternite, whereas material from other parts of the range
(including other localities in the Wet Tropics region) usually
has more teeth. Specimens from the southern part of the
range (Lamington and Kenilworth/Imbil regions) have 5+4,
5+5 or 6+6 teeth (Fig. 7D), and the dental margin is more
trapezoidal than semicircular (compare Fig. 6E,H for
maximum difference). However, these same characters are
present in specimens from the Bluewater Range and
Cardwell Range/Cardwell Gap in the Wet Tropics, so the
southeastern Queensland samples cannot be segregated on
this basis. Indeed, the sample from Eungella confounds
distinguishing geographic groupings within P. monteithi
based on tooth number or shape of the coxosternite, having
3+3,4+4, and 5+5 teeth about equally represented in a single
population (Fig. 7A-C), and the dental margin varies from
wide/biconvex (Fig. 1C) to approximately semicircular
(Figs. 6H, 7A).

Paralamyctes (Paralamyctes) neverneverensis n.sp.

Figs. 9B, 10, 11

Etymology. For the Never Never, Dorrigo NP, where the
species occurs.

Diagnosis. Paralamyctes (Paralamyctes) with Tomosvary
organ on cephalic pleurite, without modification of sutures
around it as in P. (P.) monteithi; dental margin of maxillipede
coxosternite relatively narrow, with 5+5 or 6+6 large,
pointed teeth; pretarsal section of tarsungulum relatively
short; spinose distal projection on tibia of leg 15.

Type material. Holotype AM KS 61063, $ (Fig. 10A,B),
Nana Creek SF, 5 km ENE of Lowanna, NSW, 30 o lF51"S
152°56'53"E, MRG, G. Milledge and H. Smith, 10-23 Nov
1999, pitfall. Paratypes: QM S42666, \ S (Fig. 11),
S42667,1 $, Dorrigo NP, NSW, Never Never, 700 m, GBM,
1980-1981, pitfall, rainforest; AM KS 57955,1 S (Figs 9B,
10C; DNA voucher specimen), Dorrigo NP, Wonga Walk,

200 m E of Tristania Falls, 30°22’S 152°44’E, G. Giribet, M.K.
Nishiguchi and Y.-y. Zhen, 29 Mar 2000, rainforest litter.

Description. This species is sufficiently similar to P. (P.)
monteithi , described in full above, to limit description to
features that differ. Description of colour in a freshly
collected specimen is also provided because this information
is often lacking in preserved material.

Length of head shield up to 1.4 mm. Head with irregular
chestnut mottling, bright orange band along transverse
suture; antennae dark orange, lighter distally; maxillipede
bright orange; tergites light orange with chestnut median
longitudinal patch and posterior and lateral margins; stemites
with irregular purple and chestnut network; proximal part of
legs light yellow with blue patches, tarsi orange.

Antenna with 20-21 articles; basal two articles
considerably larger than succeeding ones, several articles
of subequal length and width, including on distal part of
antenna (Fig. 11H). Inner margin of labrum gently
shouldered. Tergite shapes as for P. (P.) monteithi except
for slightly shorter TT12 and 14 (Fig. 10A). Maxillipede
coxosternite trapezoidal (Fig. 11K); dental margin convex,
each half sloping posteromedially; narrow band along dental
margin sclerotised to same extent as teeth; median notch
shallow or lacking (Fig. 11M); pretarsal part of tarsungulum
slightly longer than tarsal part (Fig. 11L). About five setae
on anteromedial edge of coxal process of first maxilla (Fig.
111). Distitarsus about 55% length of basitarsus on anal leg;
anal leg basitarsus about 10 times longer than broad (Fig.
9B). Coxal pores round; 2344/2344, 2443/2344 in females;
2222/2222, 2333/2233 in males. Genital sternite and
gonopods of both sexes (Fig. 10B,C) indistinguishable from
P. (P.) monteithi.

Discussion. Paralamyctes neverneverensis co-occurs with
P. (Nothofagobius) cassisi in the Dorrigo region. They are
most readily distinguished by the former’s fewer antennal
articles, more dentate margin of the maxillipede, larger Tl,
and two (versus three) spurs on the female gonopod, without
an extension on the spur-bearing segment. Mandibular
characters also provide obvious distinction (e.g., strongly
pinnulate versus simple aciculae).

Paralamyctes (Paralamyctes) neverneverensis is
distinguished from the Queensland species P. (P.) monteithi
by having the Tomosvary organ on the surface of the
cephalic pleurite (Fig. 11G) rather than on the ventral margin
of the head. The peculiar sutural course around the
Tomosvary organ is unique to P (P.) monteithi, whereas P.
(P.) neverneverensis retains the typical pleurite morphology
that is primitive for Paralamyctes (indeed, general for
Henicopidae). Further distinction between the New South
Wales and Queensland species is provided by the larger
teeth on the maxillipede coxosternite in the former, the
narrower dental margin, the shorter tarsungulum on the
maxillipede (Fig. 11K-M), and the presence of some short
antennal articles (Fig. 11H). The differences cannot be
attributed to ontogeny (all specimens of P. neverneverensis
being relatively small) because the specific differences can
be discerned in equivalent sized and even smaller specimens
of P. monteithi. Even the larval stage with 10 legs and two
limb-buds (equivalent to the third larval stadium of

216 Records of the Australian Museum (2001) Vol. 53

Figure 10. Paralamyctes (Paralamyctes) neverneverensis n.sp. A,B, holotype $ AM KS 61063, Nana Creek SF,
NSW; B, terminal segments and gonopods; scales 1 mm, 0.25 mm. C, paratype <5 AM KS 57955, terminal segments
and gonopods, Dorrigo NP, NSW; scale 0.25 mm.

Lithobius variegatus : Eason 1964) already has a long
maxillipede tarsungulum and only 3+3 blunt teeth on the
dental margin in P. monteithi (QM S45147).

The maxillipede coxosternite in P. (P.) neverneverensis
is similar in shape to that of P. (P.) spenceri from southern
Africa, and these species also share larger, more pointed

teeth than are developed in other species of the subgenus.
Paralamyctes neverneverensis is distinguished from P.
spenceri by having distal spinose projections on the tibiae
of all trunk legs [see Fig. 9 for comparison of anal legs in
P. (Paralamyctes )] and by having a shorter maxillipede
tarsungulum.

Edgecombe: Australian Paralamyctes 217

Figure 11. Paralamyctes (Paralamyctes) neverneverensis n.sp. 6 QM S42666, Never Never, Dorrigo NP, NSW. Scanning electron
micrographs. A, fringe of branching bristles on mandible, scale 10 pm; B,C, mandibular teeth and furry pad, scales 10 pm; D, branching
bristles on labral margin, scale 10 pm; E, telopod of first maxilla, scale 20 pm; F, second maxillae, scale 100 pm; G, cephalic pleurite,
scale 50 pm; H, distal articles of antenna, scale 20 pm; I, coxal processes of first maxilla, scale 20 pm; J, mandibular aciculae, scale 10
pm; K, maxillipedes, scale 100 pm; L,M, dorsal and ventral views of maxillipede dental margin, scales 100 pm, 20 pm.

Paralamyctes (Thingathinga) n.subgen.

Etymology. Thingathinga, the Yaurorka name for centipede
(Johnston, 1943).

Diagnosis. Paralamyctes with simple (non-pinnulate)
mandibular aciculae; dental margin of maxillipede
coxosternum wide, subtransverse, bearing numerous (6-
12) small, knob-like teeth; coxal process of first maxilla
gently tapering, blunt, with cluster of setae concentrated
on tip; articulation between tarsomeres of anterior legs

developed ventrally but fused dorsally; longitudinal median
furrow confined to anterior part of sternites.

Type species. Paralamyctes (Thingathinga) grayi n.sp.

Assigned species. Paralamyctes validus Archey, 1917 (=
P. dubius Archey, 1917); Paralamyctes (Thingathinga)
hornerae n.sp.

Distribution. Australia (New South Wales), New Zealand
(North and South Islands).

218 Records of the Australian Museum (2001) Vol. 53

Discussion. Thingathinga n. subgen. is endemic to New
South Wales and New Zealand. The simple mandibular
aciculae that unite species of Thingathinga (Figs. 13F,
18K; Edgecombe et al., 2001: fig. 5G) are elsewhere
observed in Henicopidae only in Australian species of P.
(Nothofagobius), but a majority of character data suggests
independent origins of simple aciculae from bipinnulate
aciculae in these two groups. Congruent with simple
aciculae in Thingathinga are a wide, subtransverse dental
margin of the maxillipede that bears at least six small teeth
(Figs. 15B,I, 18E), and a blunt and weakly tapering coxal
process on the first maxilla (Figs. 14G,H, 15K, 18G). The
setae on the tip of the coxal process are segregated from
those along the medial margin of the process more so than
in the other subgenera of Paralamyctes, in which the setae
are developed along a sloping anteromedial edge of the
coxal process (compare Figs. 14H and 111, 20G). Species
of P. (Thingathinga ) have weaker tarsal articulations than
other Paralamyctes , the articulation in anterior legs having
a narrow fused region along the dorsal edge. The relatively
incomplete median furrow on the stemites in P. (Thingathinga)
provides a distinction from P. (Paralamyctes ) in particular.

The subdued tarsal articulations of P. (Thingathinga)
invite comparison with Lamyctopristus validus Attems,
1928, from South Africa, in which a faint trace of a joint is
sometimes present on legs 1-12. Assignment of Thinga¬
thinga to Lamyctopristus can be dismissed based on the
deep median notch on the maxillipede coxosternite of L.
validus and L. granulosus Lawrence, 1955b, and presence
of only three large teeth on the dental margin in both species
of Lamyctopristus. I have examined the sole specimen of
the type species (SAM ENW X7515). Attems (1928) did
not describe a small conical node lateral to these teeth in
the holotype of L. validus that corresponds to the
pseudoporodont of Lamyctes, Lamyctinus and Analamyctes.
The median furrow on the head of Lamyctopristus validus
terminates just behind the median notch, thus excluding
this species from the Paralamyctes/Haasiella clade. An
alliance between Lamyctopristus and Lamyctes, Lamyctinus
and Analamyctes is indicated by short antennomeres
occurring in pairs between groups of longer antennomeres
in the proximal part of the antenna, the subtriangular outline
of the maxillipede coxosternite (narrow, curved dental
margin), and presence of a pseudoporodont (characters 4(1),
11(0), and 14(0), respectively, of Edgecombe et al., 2001).

Paralamyctes (Thingathinga) grayi n.sp.

Figs. 8C,D, 12-14, 15A-G,H-M?, 16A?, 16B

Paralamyctes n.sp.-Edgecombe et al., 1999: 295.

Paralamyctes n.sp.-Edgecombe et al., 2000: fig. 3C,D.

Etymology. For Michael R. Gray, in recognition of his
career in terrestrial arthropods at the Australian Museum,
and for collecting this species throughout much of its range.

Diagnosis. Paralamyctes (Thingathinga ) with antenna of
22-29 articles; ocellus weakly convex; mandible with up
to 15 simple aciculae; coxal process of first maxilla bearing
up to 25 simple setae, usually about 15; 6-12 small teeth

on dental margin of maxillipede coxosternite (most
commonly 7+7 at type locality); ventral edge of prefemora
commonly recessed; five to eight coxal pores on each of
legs 12-15, set in deep grooves and largely concealed by
anteroventral face of coxa; coxal pores usually ovate or
figure eight-shaped; distal spinose projection on tibia of
leg 14 but not 15; short setae abundant on genital sternite
and female gonopods; penis small; terminal process on male
gonopod short.

Type material. Holotype AM KS 57912, 6 (Fig. 12A), from
former Scout camp, Mt Keira Fauna Reserve, NSW, 34°24'S
150°50'E, C. Horseman, 20 Nov 1978-17 Jan 1979, pitfall.
Paratypes: all from type locality, C. Horseman and MRG, pitfalls:
Australian Museum KS 35607,4c? c?, 6 2 $, KS 57913, 1 $ (Fig.
15A-E), 14 Nov 1974; KS 35833, lc?, KS 35835, 4c?c?, 22 2,
KS 35836, 2c?c?, 12,KS 57916, lc?,KS 57917, lc?,22 2,KS
57918, 3c? c?, 23 Nov-20 Dec 1978; KS 35834, 1 c?, 23 Jun-20
Dec 1978; KS 35837, 2c? c?, KS 35838,1 c?, KS 35839,7c? c?, KS
35840, lc?, 17 Jan-14 Feb 1979; KS 35841, 1 c?, 12,7 Sep-4
Oct 1979;KS35858, lc? (Fig. 15G),KS57914,1 c? (Figs. 13A,B,
14A-C,J,K, 15F), KS 57915, 12 (Figs. 13F, 14D-F,L), 31 Jan
1979; KS 35860, 1 c?, KS 35863, 1 c?, KS 35866, 1 c?, KS 35868,
22 2, 14 Mar-11 Apr 1979; KS 57919, 2c?c?, 12, KS 57920,
6c? c?, 12,20 Nov 1978-17 Jan 1979; KS 57921, 6c? c?, 12,23
Nov-20 Dec 1978; KS 41430, 12 (Fig. 16B), KS 57922 (Figs.
13C-E,G-M, 14G-I), KS 57923, lc? (Fig. 12C), KS 57924, 1 2
(Fig. 12B), KS 57925, 9c?c?, 62 2, KS 57926, lc? (Fig. 14M),
KS 57927,6 c? c?, 12,20 Nov 1978-17 Jan 1979; KS 57928,6c? c?,
62 2, 14 Feb 1979; KS 57929, lc?, 14 Mar-11 Apr 1979; KS
57930, 1 c?, 12, 11 Apr-17 Jun 1979; ANIC, 1 2, Mt Keira Scout
camp, 320 m, L.C. Calder, 4-5 Mar 1981, litter Berlese.

Other material. Illawara: AM KS 57931, 2 2 2, KS 57932,
2c? c?, 12 (DNA voucher specimens), KS 58466, 9c? c? (Fig.
8C,D), 62 2, Mt Barrengarry, 6.8 km N of Hampden Bridge,
34°42'S 150°30'E, GDE and ZJ, 2 Mar 1997, 13 Nov 1997, 30
Sep 2000, rainforest; CAS, 5c? c?, 5 2 2 , Robertson, 760 m, E.S.
Ross and D.Q. Cavagnaro, 8 Dec 1962. Budawang Range: AM
KS 57933, 2c?c?, Budawang NP, Clyde Mountain, 35°33'S
149°57'E, GDE and ZJ, 2 Jan 2000, hand collected in litter, open
eucalypt forest. Blue Mountains: AM KS 35600,1 2, KS 35826,
lc?, KS 35828,1 c?, KS 35861,2c? c?, KS 35865,2 c? c?, KS 57934,
5 c? c?, Cathedral of Ferns, Mt Wilson, 33°30'S 150°23'E, MRG
and C. Horseman, Jun 1978-Jun 1979, pitfall; KS 30391, lc?,
Kanangra-Boyd NP, Boyd Plateau, 33°44'S 150°25'E, MRG, 15
May 1971; AM KS 57935, 1 c? and 1 2, road to Ingar picnic area,
33°46'05"S 150°24'30"E, Australian Museum Business Services,
3 Oct 1996; AM KS 57936 (DNA voucher specimen), Ingar picnic
area, 33°46'S 150°28'E, GDE, G. Giribet and ZJ, 21 Apr 2000.
Barrington Tops: AM KS 57937, 1 2 (Fig. 16A), Chichester
SF, Mt Allyn Rd, adjacent to Mt Allyn Forest Park, 32°07'04"S
151°25'33"E, 970 m, GC and MRG, 4 Feb-9 Apr 1993, pitfall;
AM KS 57938, lc?, from Chichester SF, 1.8 km N of ford on
Karuah River on Karuah River Rd, 32°05'58"S 151°43'18"E, GC
and MRG, 4 Feb-9 Apr 1993, pitfall; AM KS 57939, lc?,
Barrington Tops SF, W of junction of Thunderbolts Track and
Devils Hole Track, 31°54'S 151°28'E, 1420 m, MRG and GC, 4
Feb-9 Apr 1993, pitfall; AM KS 57940, 1 2 (Fig. 15H-M; DNA
voucher specimen), Chichester SF, Mt Allyn Forest Park, GDE
and ZJ, 16 Mar 1999, rainforest.

Description. Length up to 32 mm; length of head shield up
to 3.3 mm. Antennae and maxillipedes bright orange,
antennae lighter distally; head shield and tergites mottled
orange-brown, with darker pigment concentrated near

Edgecombe: Australian Paralamyctes 219

Figure 12. Paralamyctes (Thingathinga) grayi n.sp. Mt Keira Fauna Reserve, NSW. A, holotype 6 AM KS 57912.
Antennae incomplete distally; scale 2 mm. B, paratype 9 AM KS 57924, terminal segments and gonopods; scale
0.5 mm. C, paratype 6 AM KS 57923, terminal segments and gonopods; scale 0.5 mm.

lateral and posterior borders; sternites mottled, predom¬
inantly yellowish or light orange in anterior segments,
darker orange-brown in posterior segments; legs relatively
uniformly orange, pigmentation of tarsus only slightly
deeper than tibia.

Head shield smooth. Frontal margin with strong, short
median notch (Fig. 12A); posterior margin gently concave.
Median furrow extends back to transverse suture, length

about 30% that of head shield, deep throughout. Antenna
extending back to T5 or T6 (Fig. 12A); 22-29 antennal
articles, most commonly 23 at Mt Wilson (range from 22-
28), 25 at Mt Keira (range 23-29); basal two articles large,
succeeding six or so of similar length and width, some wider
than long (Figs. 13B, 15C), more distal articles considerably
longer than wide; distal article variably longer than
preceding one, typically about 40% longer; antennal articles

220 Records of the Australian Museum (2001) Vol. 53

Figure 13. Paralamyctes (Thingathinga) grayi n.sp. Mount Keira Fauna Reserve, NSW. Scanning electron micrographs. A,B, 8 AM
KS 57914. A, head shield, scale 300 pm; B, proximal antennal articles, scale 200 pm. C-E,G-M, AM KS 57922. C, gnathal lobe of
mandible, scale 50 pm; D, fringe of branching bristles and aciculae of mandible, scale 30 pm; E, furry pad on mandible, scale 10 pm; G,
ventral view of head, scale 300 pm; H,I, Tomosvary organ, scales 10 pm, 5 pm; J,K, labral margin and detail of branching bristles on
margin, scales 30 pm, 10 pm; L,M, distal part of telopod of second maxilla, scales 60 pm, 30 pm. F, 9 AM KS 57915, mandibular
aciculae, scale 10 pm.

densely, finely setose, with distinctly longer setae (ridged
trichoid sensilla) encircling distal edge (Fig. 13B); basal
articles with sparser, more robust setae (Figs. 13G, 14A).
One to three (usually two) finger-shaped thin-walled
basiconic sensilla (Fig. 8D) and one to four (usually two or
three) shorter, conical thick-walled basiconic sensilla at

anterior edge of antennal articles (Fig. 8C); cluster of thick-
walled basiconic sensilla (blasse Borsten ) at tip of distal
article; stop ( Kragen ) at base of trichoid sensilla as described
for P. (P.) monteithi. Ocellus whitish, gently bulging.
Tomosvary organ moderately large, ovate, with outer edge
at margin of cephalic pleurite (Figs. 13H, 14J), membranous

Edgecombe: Australian Paralamyctes 221

Figure 14. Paralamyctes (Thingathinga) grayi n.sp. Mount Keira Fauna Reserve, NSW. Scanning electron micrographs. A-C,J,K, 6
AM KS 57914. A, ventral view of head, scale 200 pm; B,C, first maxillae and detail of distal article of telopod, scales 100 pm; J,
cephalic pleurite, scale 100 pm; K, pectinate setae on inner margin of distal article of first maxillary telopod, scale 30 pm. D-F,L, $ AM
KS 57915. D, maxillae, scale 200 pm; E, first maxillae and coxosternite of second maxillae, scale 100 pm; F, coxa of second maxilla,
scale 100 pm; L, tarsus and claw of second maxilla, scale 10 pm. G-I, AM KS 57922. G, first maxillae, scale 100 pm; H, coxal processes
of first maxillae, scale 50 pm; I, basiconic sensilla between coxal process and telopod of first maxilla, scale 5 pm. M, <5 AM KS 57926,
gonopod flagellum, scale 10 pm.

medially, numerous small pores (openings of gland ducts)
scattered around edges of inner surface (Fig. 131). Head
shield sparsely setose, two pairs of setae on frontal margin
outside median notch, a few setae around ocellus and along
lateral border.

Tergites nearly smooth. T1 trapeziform, smaller than T3,
very slightly narrower than head shield (Fig. 12A), about
90% width of widest tergite (T8), posterior angles rounded
and posterior margin faintly concave; lateral borders
subparallel anteriorly in TT3 and 5, posterior angles

222 Records of the Australian Museum (2001) Vol. 53

Figure 15. A-G, H-M?, Paralamyctes (Thingathinga) grayi n.sp. A-G, Mount Keira Fauna Reserve, NSW. A-E, $ AM KS 57913. A,
ventral view of head, scale 200 pm; B, dental margin of maxillipede coxosternite, scale 50 pm; C, proximal part of antenna, scale 200
pm; D, sternite of first genital segment and gonopods, scale 100 pm; E, gonopods, scale 40 pm. F, $ AM KS 57914, tibia, tarsus and
pretarsus of leg 1, scale 60 pm. G, $ AM KS 35858, tergites 10-14, scale 400 pm. H-M, 9 AM KS 57940, Chichester SF, Barrington
Tops, NSW. H, head shield, scale 200 pm; I, dental margin of maxillipede coxosternite, scale 100 pm; J, ventral view of head to show
maxillae, scale 200 pm; K, telopods and coxal processes of first maxillae, scale 50 pm; L, gnathal lobe of mandible, scale 50 pm; M,
accessory denticles and furry pad on mandible, scale 10 pm.

rounded, posterior borders weakly concave; TT1, 3 and 5
bordered posteriorly; lateral border of T7 gently convergent
posteriorly, border incomplete posteriorly, posterior margin
with pronounced embayment, transverse or slightly convex
across medial quarter of width; TT8-14 bordered laterally;
posterior margin of T8 gently concave, posterior angle
rounded, TT10 and 12 more concave posteriorly, posterior
angles form obtuse, sharp corners; TT9 and 11 embayed,

with transverse posteromedial third; posterior angle of T9
blunt, TT11 and 13 angular; lateral margin of T14 convex,
posterior margin concave (Fig. 15G); tergite of intermediate
segment elongate subhexagonal, slightly wider in females,
posterior margin concave in both sexes; no tergites toothed
at posterior angles. TT3, 5 and 7 with most abundant setae
on anterior/anterolateral part, sparsely setose elsewhere
except for numerous short setae along lateral borders

Edgecombe: Australian Paralamyctes 223

Figure 16. Anal legs of Paralamyctes (Thingathinga ) and P. (Nothofagobius), with details of pretarsi. A, P. (Thingathinga) Igrayi n.sp.,
$ AM KS 57937, Chichester SF, Barrington Tops, NSW. B, P. (Thingathinga) grayi n.sp., $ AM KS 41430, Mt Keira Fauna Reserve,
NSW. C, P. (Nothofagobius) mesibovi n.sp., QVMAG 23/23009, holotype 9, Wombat Hill, Waratah, Tasmania. D, P. (Nothofagobius)
cassisi n.sp., 6 QM S42670, Gibraltar Range NP, NSW. E, P. (Nothofagobius) chilensis (Gervais in Walckenaer & Gervais, 1847),
9 ZMUC , Colonfa Suiza, San Carlos de Bariloche, Argentina. Scales 1 mm except E, 0.5 mm.

(present on all tergites); setae more evenly scattered over
surface of posterior long tergites (Fig. 15G).

Transverse seta projects medially from labral sidepiece.
Labral margin with rounded shoulder beside midpiece, with
marked break in curvature where dense fringe of branching
bristles overhangs margin (Fig. 13G,J); bristles generally
branch as uneven, elongate bifurcations along nearly their
entire length (Fig. 13K).

Dental margin of maxillipede coxosternite broad, each
half approximately straight, gently sloping backwards
laterally, set off by transverse furrow (Fig. 15B); teeth small,
blunt, 6-10 teeth per margin, most commonly 7, in Mt Keira
sample; 7-12 teeth per margin, most commonly 9, in Mt
Wilson sample; median notch in coxosternite shallow or
lacking; maximum length of coxosternite 60% of its
maximum width (Fig. 15A); setae concentrated on anterior/

224 Records of the Australian Museum (2001) Vol. 53

anterolateral part of coxosternite, longer anterolaterally.
Duct of poison gland short, extending to anterior one-third
of tibia, tubular. Tarsungulum long, slender (Fig. 15A).
Setae on forcipule concentrated on inner part of trochantero-
prefemur, femur and tibia and scattered over tarsal part of
tarsungulum.

Mandible: Trunk with single fissure defining lamina
condylifera. Four paired teeth (Fig. 13C), small, unpaired
fifth tooth dorsally on one side. Outer fringe of bifid bristles
skirts aciculae, evenly shortening dorsally to a narrow band,
with branches along entire length of each bristle (Fig. 13D).
12-15 curved aciculae, all lacking marginal branchings (Fig.
13F). Accessory denticle field delimited by strong groove
on two or three dorsal teeth; largest denticles tubercle-like,
grading into rod-shaped scales then small elongate scales
near fringe of branching bristles. Furry pad strongly
differentiated from accessory denticles / gnathal lobe,
developed as an isolated cluster of bristles with a few short
branchings (Fig. 13E).

First maxilla: Bell-shaped sternite of typical size for
genus, anterolateral margins against narrow unsclerotised
strip on inner edge of coxa, sternite and coxa separated by
strong suture just in front of posterior margin of maxilla
(Fig. 14B,E,G). Coxal process with blunt, transverse tip
bearing dense cluster of 14-25 simple, ridged setae (Fig.
14H); cluster of about seven tiny, barb-like basiconic sensilla
between coxal process and telopod (Fig. 141). Distal article
of telopod with two rows of long, plumose setae along inner
margin (Fig. 14C); branches developed along distal half of
these setae (Fig. 14K); numerous relatively short, simple
setae on inner half of distal article, longer simple setae on
lateral part, separated by a non-setose patch (Fig. 14C).

Second maxilla: Sternite small, trapezoidal, fused with
coxae (Fig. 14D,E). Short setae along anterior part of coxa
relatively weakly developed (Fig. 14F). Tarsus bearing
straight, ridged setae on outer surface, many plumose setae
on inner surface (Fig. 13F); branches on plumose setae mostly
along less than half of length (Figs. 13M, 14F). Claw composed
of four or five digits, with median digit enlarged (Fig. 14F).

Strong, sharp distal spinose projection on tibiae of legs
1-14. Premur and femur with rather evenly scattered setae
on anterior and posterior faces as well as dorsally, setae
longest against ventral prefemoral recess; fringe of
numerous setae encircling distal margin of femur; tibial
setae mostly shorter, of varied thickness, slightly thicker
seta at ventrodistal end of tibia; tarsal setae fine, of varied
length, aligned as two rows on ventral margin without
precise pairing of larger setae. Articulation between
tarsomeres relatively strong on anterior legs (Fig. 15F),
indistinct along a narrow extent on dorsal edge, typically
preserved with substantial flexure. Distitarsus 50 to nearly
60% length of basitarsus on anal leg; anal leg basitarsus 9-
10 times longer than broad (Fig. 16B). Pretarsal claws
relatively long, curved; pair of accessory claws symmetrical,
25-30% length of main claw (Fig. 16B).

Fongitudinal median furrow confined to anterior part of
sternite, not extending past anterior third even in
posteriormost sternites. Sternites sparsely setose, most
consistent setae in a transverse band across anterior third
and along anterolateral and posterolateral margins. Posterior
margins of sternites usually convex.

Coxal pores: 5-8 pores (most commonly 6 or 7) on each
of legs 12-15; most coxal pores ovate, in larger specimens
some pores slightly constricted medially or figure eight¬
shaped; coxal pore field in deep groove, separated from
anteroventral face of coxa by sharp fold (Fig. 12B,C).

Male (Fig. 12C): Sternite of segment 15 transverse or
gently convex posteriorly, with rounded posterolateral
corners. Sternite of first genital segment bearing numerous
short setae scattered on posterior two-thirds, more abundant
along posterior edge, with well-defined transverse row of
setae just in front of posterior margin; sternite undivided
medially. Gonopod of three articles and tapering, seta-like
(exceptionally bifurcate) terminal process, the three articles
bearing 5-7, 4-5, and 4-8 short setae (proximally to
distally); setae on proximal two articles in a transverse row;
terminal process short, about as long as third article of
gonopod, with numerous slender barb-like projections
concentrated in a narrow band along one side (Fig. 14M). Penis
a small, conical projection, extending about as far back as
articulation between first and second articles of gonopod.

Female (Fig. 12B): Sternite of segment 15 transverse
posteromedially. Tergite of first genital segment and telson
strongly sclerotised, telson typically with a lightly
sclerotised band medially, scattered with short setae. Sternite
of first genital segment large, with posterior margin concave
between condyles at which gonopods articulate (Fig. 15D);
short, rounded medial bulge variably developed in this
concavity; nearly entire surface of sternite bearing abundant,
short setae. First article of gonopod bearing two bullet¬
shaped spurs, the inner spur slightly smaller; gonopods
bearing many short setae, including a few on third article.
Claw undivided (Fig. 15E), with numerous elongate ridges
externally and internally.

Discussion. The most distinctive characters of this species
are the abundance of small, blunt teeth on the maxillipede
coxosternite, the weakly convex ocellus (flattened in P.
homerae\ more domed in P. validus), small penis, short
terminal process on the male gonopod, and the deep grooves
in which the coxal pores are set.

The only consistent difference that has been detected
between specimens from the Illawara region (Mt Keira and
Mt Barrengarry) and those from the Blue Mountains (Mt
Wilson and Boyd Plateau) is the yellow-brown colour of
the preserved specimens from the Blue Mountains, versus
orange-brown for the Illawara specimens. Blue Mountains
specimens have a higher modal number of teeth on the
maxillipede coxosternite (nine teeth is most common versus
seven in the Illawara material), though the observed range
overlaps completely.

Specimens of Paralamyctes (Thingathinga ) from the
Barrington Tops share several distinctive (derived)
characters with typical P. grayi, notably the abundant small
teeth on the maxillipede coxosternite (6-10 in Barrington
Tops specimens; Fig. 151), the strong embayment of the
posterior margin of the seventh tergite, and the weakly
convex ocellus (Fig. 15H). However, the embedding of the
coxal pores in a groove that is observed in other parts of
the species’ range is inconsistently developed in specimens
from the Barrington Tops. The only Barrington Tops
specimen with the anal leg preserved displays a distal

Edgecombe: Australian Paralamyctes 225

spinose projection on the tibia (Fig. 16A), a feature
otherwise unrecorded in P. grayi (Fig. 16B), and both males
from the Barrington Tops have fewer coxal pores than any
specimens from the Blue Mountains, Illawara or Budawang
Range (3455/4554 and 3444/2444). It is likely that the
Barrington Tops material represents a distinct species
closely related to P. grayi, but the few available specimens
do not permit confident diagnosis, particularly given the
limited data on the anal leg. The description of P grayi
above excludes the Barrington Tops specimens.

Among previously described species, most closely allied
to Paralamyctes (Thingathinga) grayi is P. validus Arc hey,
1917, a widespread species in New Zealand (ranging from
the Waitakere Ranges and islands off Auckland on North
Island to Queenstown on South Island). With P. grayi, P.
validus shares coxal pores that are set in deep grooves,
largely concealed by the anteroventral face of the coxa, and
a proliferation of small teeth on the dental margin of the
maxillipede coxostemite. Compared to P. validus, P. grayi
has stronger articulations between tarsomeres (Fig. 15F),
blunter teeth on the maxillipede dental margin (Fig. 15B),
a narrower dorsal section of the fringe of branching bristles
on the mandible (Fig. 13E), and a distinctive embayment
of the margin of tergite 7 (Fig. 12A).

The morphology and distribution of basiconic sensilla
on the antenna described for P. (T.) grayi and P. (P.)
monteithi (Fig. 8) are identical with those in Lithobius
(Rilling, 1968: fig. 36C,D; Lewis, 1981: fig. 88). Such
similarity between Henicopidae and Lithobiidae suggests
that details of these sensilla may provide synapomorphies
for Lithobiomorpha (e.g., discrete clusters of thick- and thin-
walled basiconic sensilla at the anterior margin of the
antennal articles).

Paralamyctes (Thingathinga) hornerae n.sp.

Figs. 17, 18

Etymology. For B. Elizabeth Horner, who sponsored the
description of this species.

Diagnosis. Paralamyctes (Thingathinga) with 20-22
antennal articles; ocellus flattened; dental margin of
maxillipede coxostemite nearly transverse, lacking median
notch, with 6-8 small teeth on each side; few (4-6) setae
on tip of coxal process of first maxilla; faint trace of
articulation defining tarsomeres on legs 1-12.

Type material. Holotype AM KS 57941, S (Fig. 17A,C),
bottom end of Cliffs Trail, about 3.8 km NE of Oxley Rd,
Styx River SF, NSW, 30°33’S 152°2FE, 1180 m, MRG and
GC, Feb 4-Apr 9 1993, pitfall. Paratypes: AM KS 57942
(Fig. 17B), 57943 (Fig. 18), 2 9 9, from type locality, same
collection.

Description. Length up to 17 mm; length of head shield up
to 1.4 mm. Tergites (of specimens in 70% ethanol) yellowish
brown or mauve, darker posteriorly; antennae and
maxillipedes yellow; prefemur to tibia mauve, particularly
on posterior legs of trunk; tarsi yellow.

21/22 and 21/20 antennomeres in specimens with

complete antennae; basal two articles stout, with long setae,
rest of antenna with uniform dense, shorter setae, longer
setae encircling distal end of each article; many articles not
substantially longer than wide, even in distal part of antenna.
Median furrow deep to transverse suture. Ocellus indicated
by faint convexity and subdued pigmentation, margins obscure.
Tomosvary organ at anterolateral edge of cephalic pleurite,
moderately large (size, position and shape as for P. grayi).

Tergites smooth; shapes, proportions and borders (Fig.
17A) as described for P. grayi.

Labral margin transverse adjacent to midpiece, then
abruptly flexed backward where fringe of bristles overhangs
margin (Fig. 18C); bristles irregularly pectinate (Fig. 18J),
branching along entire fringe.

Dental margin of maxillipede nearly transverse, each half
faintly convex forwards, bearing 6-8 small, blunt teeth,
median notch absent (Fig. 18E); dental margin delimited
by pronounced transverse furrow; coxostemite broadly
trapezoidal, maximum length 60% of maximum width (Fig.
18D); anterolateral comer of coxostemite bearing a few long
setae, shorter setae on anteromedial part; median suture
shallowed anteriorly; tarsungulum relatively long and slender.

Mandible: Four paired teeth (Fig. 18A). Outer fringe of
pectinate bristles with short branches along entire length of
each bristle (Fig. 18B); eight simple, sickle-shaped aciculae
lacking marginal pinnules (Fig. 18K); furry pad composed
of dense seta-like processes that birfurcate or trifurcate near
their tips.

First maxilla: Sternite triangular, size and sutures as in
P. grayi (Fig. 18F). Coxal process with cluster of 4-6 simple
setae clustered at tip, one strong seta towards base (Fig.
18G). Distal article of telopod with evenly scattered simple
setae, pairs of plumose setae aligned on inner margin.

Second maxilla as described for P. grayi; pretarsus of
five digits (Fig. 18H).

Distal spinose projection on tibia of legs 1-14; anal leg
unknown. Tarsi 1—12 entirely unflexed, faint trace of
articulation at 55-60% of length. Setae on legs as in similar
sized specimens of P. grayi except for slightly finer tibial
setae (Fig. 18L); tarsal setae uniformly fine. Pretarsal claws
relatively long, curved; pair of accessory claws symmetrical,
about 40% length of main claw (Fig. 18M).

Longitudinal median furrow extends to midlength of
posterior sternites.

Coxal pores on legs 12-15 round, 2344/2444 in male,
3455/4555 and 3445/3555 in females; pore rows in shallow
grooves.

Male (Fig. 17C): Short setae evenly scattered on sternite
of first genital segment. Three setae encircling distal end of
first segment of gonopod; one seta on second article; third
article slightly elongated, bearing a few setae; terminal
process long, flagelliform. Penis small, conical, about as
long as first article of gonopod.

Female (Fig. 17B): Two spurs on basal article of
gonopod; spurs bullet-shaped, pointed, dorsal surface
concave; inner spur smaller than outer one (Fig. 181); third
article of gonopod lacking setae.

Discussion. Paralamyctes hornerae differs from all
congeneric species in having only faint tarsal articulations
on legs 1-12. These articulations do not demonstrate any

226 Records of the Australian Museum (2001) Vol. 53

Figure 17. Paralamyctes (Thingathinga) hornerae n.sp. Cliffs Trail, Styx River SF, NSW. A,C, holotype 6 AM
KS 57941; C, terminal segments and gonopods; scales 1 mm, 0.25 mm. B, paratype 9 AM KS 57942, terminal
segments and gonopods; scale 0.25 mm.

flexure and are evidently non-functional. A possible
relationship to Haasiella could be considered based on the
weak tarsal jointing, though in Haasiella the tarsus is
entirely unjointed. Also suggestive of Haasiella is the
transverse dental margin of the maxillipede (Fig. 18E) (more
curved in most other henicopines). However, the shape of
the coxostemite differs markedly from New Zealand species
of Haasiella. The overall proportions are considerably
broader, and it lacks the prominently narrowed extension
that bears the teeth in the New Zealand species. As well, a

median notch is lacking (present in New Zealand species).
In each of these respects P. hornerae resembles species of
Paralamyctes (Thingathinga). A transverse, unnotched
margin of the maxillipede is similarly developed in P.
(Thingathinga) from the Barrington Tops (Fig. 151).
Membership in Thingathinga is indicated by the aciculae
of the mandible being simple (Fig. 18K), rather than
bipinnulate. The simple aciculae are not widely distributed
in the Henicopinae, but are shared with P grayi and P.
validus. In Haasiella trailli the aciculae are bipinnulate

Edgecombe: Australian Paralamyctes 227

Figure 18. Paralamyctes (Thingathinga) hornerae n.sp. Paratype $ AM KS 57943, Cliffs Trail, Styx River SF, NSW. Scanning
electron micrographs. A,B, gnathal lobe of mandible, scales 30 pm, 20 pm; C, labrum and right mandible, scale 50 pm; D,E, maxillipede
and detail of dental margin, scales 100 pm, 60 pm; F,G, first maxilla and detail of coxal processes, scales 50 pm, 30 pm; H, tarsus and
claw of second maxilla, scale 20 pm; I, spurs on female gonopod, scale 30 pm; J, branching bristles on labral margin, scale 5 pm; K,
mandibular aciculae, scale 10 pm; L, posterior view of leg 12, scale 200 pm; M, pretarsus of leg 13, scale 20 pm.

(Edgecombe et al., 2001: fig 5B), and they are likewise
bipinnulate (with short barbs) in Tasmanian Haasiella (see
below). The weakly tapering coxal process of the first
maxilla in P. hornerae , with its concentration of setae on
the blunt tip (Fig. 18G), is typical of P. (Thingathinga).

The flattened eye of P. hornerae resembles the condition
in other New South Wales species of Paralamyctes

(Thingathinga), such as the Barrington Tops material
assigned to P. (T.) grayi. However, also closely comparable
in having a flattened eye is an undescribed species from
Tasmania ( Wailamyctes sp. of Mesibov, 1986) that is
resolved as sister to New Zealand species of Haasiella
(Edgecombe et al., 2001: figs. 15, 16). Maxillipede
morphology is similar between the Tasmanian species and

228 Records of the Australian Museum (2001) Vol. 53

P. hornerae, notably in the proportions of the coxosternite.
The Tasmanian species resembles New Zealand Haasiella
in having the tarsi of legs 1-12 fully fused, without the
feeble joint that is present in P. hornerae, as well as in having
the median furrow of the head shield extended behind the
transverse suture. Archey (1937) cited the posterior
extension of the median furrow as a diagnostic character of
Haasiella (= Wailamyctes).

Paralamyctes (Thingathinga ) species from New South
Wales (P. grayi and P. hornerae) are united by an apparent
synapomorphy in the shape of tergite 7. In both species, T7
is strongly embayed, a transverse posteromedial sector being
flanked by curved posterolateral sectors, with a flange-like
ventral expansion along the median embayment. Another
character that may prove to be of value for grouping the
New South Wales species of P. (Thingathinga) is the size
of the penis, though this is affected by the extent of its
retraction above the sternite. In both P. (T.) grayi and P. (T.)
hornerae, the penis is comparatively small (about the length
of the first article of the gonopod). Australian species
belonging to P. (Paralamyctes) and P. (Nothofagobius) have
relatively larger penes (Figs. 4C, 19C), as does P.
(Thingathinga) validus (Archey, 1937: pi. 21, fig. 4). The
larger penis appears to be the plesiomorphic state for
Paralamyctes (seeAttems, 1928: text-fig. 14 for Lamyctes).

Paralamyctes (Nothofagobius) n.subgen.

Etymology. Compounding Nothofagus and the common
lithobiomorph suffix— obius, for the trans-Antarctic
distribution of this clade and occurrence of species in
Nothofagus forest.

Type species. Paralamyctes (Nothofagobius) cassisi n.sp.

Diagnosis. Paralamyctes with small Tl, distinctly narrower
than head and T3; maxillipede coxosternite with narrow,
curved dental margin bearing 4 or 5 large teeth; tarsi slender,
relatively sparsely setose; sternite of first genital segment
of male with posterior margin convex between gonopods;
basal article of female gonopod extended as a short process.

Assigned species. Henicops chilensis Gervais in Walckenaer
& Gervais, 1847; Paralamyctes (Nothofagobius) mesibovi n.sp.

Distribution. Chile and Argentina (northern Patagonia),
Tasmania, northern New South Wales.

Discussion. A female gonopod bearing three spurs is
synapomorphic for Paralamyctes cassisi from northern New
South Wales and P. mesibovi from Tasmania (rather than
the plesiomorphic condition of two spurs seen in all
congeners, as well as most other henicopid genera). In both
species the basal article of the gonopod is extended as a
process. The only other species showing a similar (though
shorter) extension of the gonopod is P. chilensis (Gervais)
from Patagonian Chile/Argentina (Fig. 24B), and this
character is interpreted as a synapomorphy. A particularly
narrow anterior part of the trunk (i.e., body narrowed across
a small Tl) is also shared by P. cassisi, P. mesibovi and P
chilensis. The homology can be described by Tl being

distinctly narrower than the head and T3 in these three
species, versus Tl nearly equally wide as the head and T3
in other species of Paralamyctes and in relevant outgroups
(e.g., Lamyctes, Lamyctinus, Lamyctopristus and Henicops).
A relative elongation of the tarsi in P. (Nothofagobius)
(compare Figs. 16C-E with Figs. 9,16A,B) may be expressed
in terms of the proportions of the anal leg basitarsus. In P
(Nothofagobius), the basitarsus is 12-17 times longer than its
proximal width, versus a length 9-11 times width in P.
(Paralamyctes) and P. (Thingathinga).

Paralamyctes chilensis also resembles the Australian P.
mesibovi and P. cassisi in having the posterior margin of
the first genital sternite of the male convex between the
gonopods, versus approximately transverse in other
Paralamyctes. This posterior bulge in the sternite may
account for the concealment of the penis in specimens of P.
mesibovi (Fig. 22C) and P. chilensis (Fig. 24C), the penis
presumed to be retracted above the sternite.

Paralamyctes (Nothofagobius) cassisi n.sp.

Figs. 16D, 19, 20, 21A-H

Etymology. For Gerasimos Cassis, Australian Museum,
whose 1993 survey uncovered this species, and who
accompanied me in the hunt for Paralamyctes in Queens¬
land and Western Australia.

Diagnosis. Paralamyctes (Nothofagobius) with 24-29
antennal articles; cephalic pleurite short; dental margin of
maxillipede coxosternite bearing four (exceptionally five)
robust teeth; trunk tergites gently wrinkled; three spurs on
female gonopod, set on moderately long projection; terminal
process on male gonopod as long as rest of gonopod.

Type material. Holotype QM S42670, d (Fig. 19A), Gibraltar
Range NP, NSW, 10 km W of northern park headquarters,
29°32'45"S 152°14'22"E, 990 m, GBM, 1980-1981, pitfall,
rainforest. Paratypes: QM S42671 Id, QM S48111 IS (Fig.
19C), from type locality, collection details as for holotype.

Other material. Northern NSW pitfall collections, MRG and GC,
4 Feb-9 Apr 1993: AM KS 57944,1 $ (Fig. 19B),AMKS 57945,
4 S S, 2 9 $, Dorrigo NP, Wonga Walk, about 600 m N of Tristania
Falls, 30°22'S 152°44'E, 730 m; AM KS 57946, 1 $ (Figs. 20,
21A-H), AM KS 57947, 5 S S, 2 9 9, Dorrigo NP, Wonga Walk,
about 200 m SW of Hardwood Lookout, 30°22'S 152°44'E, 630
m; AM KS 57948, 1 $, Styx River SF, bottom end of Cliffs Trail,
about 1.3 km from Oxley Rd, 30°33'S 152°20F, 1080 m; AM KS
57949, 1$, Mount Hyland Nature Reserve, 1.9 km along
Chaelundi Rd from Big Bull Creek Rd, 30°08'S 152°26'E, 1160
m; AM KS 57950, 2SS, Marengo SF, Big Bull Creek, 2.7 km
NE along Foamy Creek Rd from Chaelundi Rd, 30°07'S 152°25'E,
920 m; AM KS 57951, Id, Marengo SF, Opossum Creek,
upstream of Foamy Creek Rd, 30°06'S 152°25'E, 830 m; AM KS
57952,1 S, Washpool NP, track off Cedar Trail, 29°28'S 152°20'E,
950 m; AM KS 57953, Id, Washpool NP, Cedar Creek, Cedar
Trail, 920 m; AM KS 57954, Id, Boorook SF, 1 km E of main
road, 28°49'S 152° 11'E, 900 m. ANIC, Id, Richmond Range SF,
NSW, 28°48'S 152°59'E, ca 600 m, T. Weir and A. Calder, 13-14
Feb 1983; QM S42666, 1 $, Dorrigo NP, NSW, Never Never, 700
m, GBM, 1980-1981, pitfall, rainforest; QM S42668, Id, New
England NP, NSW, 200 m from Tom’s Cabin towards summit,
30°29'48"S 152°23'48"E, 1300 m, GBM, 1980-1981, pitfall,

Edgecombe: Australian Paralamyctes 229

Figure 19. Paralamyctes (Nothofagobius) cassisi n.sp. A, holotype 6 QM S42670, Gibraltar Range NP, NSW;
scale 1 mm. B, 9 AM KS 57944, Dorrigo NP, Wonga Walk, terminal segments and gonopods; scale 0.25 mm. C,
paratype 6 QM S42811, Gibraltar Range NP, terminal segments and gonopods; scale 0.25 mm.

Nothofagus rainforest; QM S42672, S 42673 ,26 6, Poverty Point,
SE of Tenterfield, NSW, 29°06'S 152°19'E, 1160 m, GBM, 1978-
1979, pitfall, rainforest; QM S42674, 1 6, Poverty Point, NSW,
GBM, 22 Feb 1979.

Description. Length up to 20 mm; length of head shield up
to 1.8 mm. Colour known only from specimens preserved
in 70% ethanol: head red-brown, trunk usually lighter
yellow-brown, often with purplish tinge; maxillipede pale
orange, with brownish coxosternite; stemites and coxae light
brown, darker in posterior segments; prefemur, femur and tibia
may be purplish, tarsi and distal part of antennae yellow.

Head shield smooth. Frontal margin with strong median
notch; posterior margin of head shield weakly concave.
Median furrow deep to transverse suture. Antennae
extending back to TT5-7 (Fig. 19A); 24-29 antennal
articles, basal two moderately larger than succeeding
articles; most of articles 3-9 of about equal length and width,
distal articles substantially longer than wide; setation fine,
dense from third article, mix of short and moderately long
setae. Ocellus strongly bulging. Tomosvary organ
moderately large, ovate, at anterolateral corner of cephalic
pleurite (Fig. 20L); openings of gland ducts marked by pores

230 Records of the Australian Museum (2001) Vol. 53

Figure 20. Paralamyctes (Nothofagobius) cassisi n.sp. 9 AM KS 57946, Wonga Walk, Dorrigo NP, NSW. Scanning electron micrographs.
A, gnathal lobe of mandible, scale 30 pm; B,C, mandibular aciculae viewed externally and internally, scales 10 pm; D, mandibular
teeth, scale 10 pm; E,F, furry pad on mandible, scales 10 pm, 5 pm; G,H, first maxillae, scales 20 pm, 50 pm; I, tarsus and claw of
second maxilla, scale 10 pm; J, second maxilla, scale 60 pm; K, distal part of telopod of second maxilla, scale 20 pm; L, cephalic
pleurite with Tomosvary organ, scale 20 pm; M, fringe of branching bristles skirting mandibular aciculae, scale 5 pm.

scattered around outer groove of Tomosvary organ. Pleurite
relatively short, length slightly more than twice width. Main
setae on head shield consist of two pairs on frontal margin,
a pair at anterior end of ocellus, one seta beneath rear edge
of ocellus, a few on posterolateral border.

Posteromedial part of clypeus with usual transverse row
of four setae (Fig. 21 A). Seta projecting transversely from
labral side piece. Inner margin of labrum gently concave

backwards between midpiece and termination of bristle
fringe, without a marked inflection or shoulder; bristles
overhanging margin moderately long (Fig. 2IB), branching
into several rami a short distance outside labral margin, these
rami often bifid (Fig. 21C).

Trunk tergites wrinkled with short, linear ridges and
grooves having weak relief. T1 trapeziform, considerably
smaller than T3, distinctly narrower than head shield (Fig.

Edgecombe: Australian Paralamyctes 231

Figure 21. A-H, Paralamyctes (Nothofagobius) cassisi n.sp. $ AM KS 57946, Wonga Walk, Dorrigo NP, NSW. A-C, labrum, with
details of branching bristles on margin, scales 50 pm, 10 pm, 5 pm; D,E, maxillipede and detail of dental margin, scales 200 pm, 50 pm;
F-H, gonopod, showing detail of spurs, ventral view and ventrolateral view, scales 30 pm, 100 pm, 100 pm. I-M, Paralamyctes
(Nothofagobius) mesibovi n.sp. Paratype 9 QVMAG 23/23010, Wombat Hill, Waratah, Tasmania. I, cephalic pleurite, scale 50 pm; J,
Tomos vary organ, scale 20 pm; K-M, maxillipede and details of dental margin, scales 200 pm, 40 pm, 20 pm.

19A), about two-thirds width of widest tergite (T8),
posterior angles strongly rounded and posterior margin
transverse medially or convex; lateral borders parallel in
TT3 and 5, posterior angles rounded, posterior borders
nearly transverse; TT1, 3 and 5 bordered posteriorly; TT7-
14 bordered laterally; lateral borders of T7-T14 convex;
posterior margins of TT7-8 gently concave, T7 more so,
posterior angles rounded; TT10, 12 and 14 progressively

more concave posteriorly; posterior angle of T10 an obtuse
corner, T12 angular, T14 sharply angular but without a
tooth-like extension; posterior margins of TT9, 11 and 13
deeply concave, posterior angles sharp, not toothed. Setae
on long tergites in anterior part of trunk sparse, mostly
confined to an anterior band and on/against lateral border;
setae evenly scattered over surface of posterior tergites,
including a row along posterior margin on TT7-14.

232 Records of the Australian Museum (2001) Vol. 53

Dental margin of maxillipede coxosternite convex,
sloping posteromedially, with moderately deep median
notch (Fig. 21E); teeth robust, most specimens having 4+4
teeth, a few having 4+5 (with small inner tooth); width
across dental margin half maximum width of coxosternite;
setae scattered on anterior and medial part of coxosternite.
Tarsungulum relatively small, tarsal and pretarsal sections
equal in length (Fig. 2ID).

Mandible: Four paired teeth (Fig. 20A), three ventral
tooth pairs bearing a third, triangular cusp on outer edge of
larger tooth; three dorsal teeth with abundant, scale-like,
triangular accessory denticles (Fig. 20D); accessory denticle
field without grooves on teeth, continuous between teeth.
Fringe of branching bristles with branches along entire
length of each narrow-based bristle (Fig. 20M); fringe
evenly narrowing dorsally, evenly grading into multi-
furcating scales on dorsal part of mandible (Fig. 20E). 10-
11 simple, curved aciculae, lacking marginal branchings
(Fig. 20B,C); basal part of aciculae with suture-like
striations, distal half with a few strong linear ridges, well
developed on inner surface of aciculae. Band of polygonal
scales between accessory denticles and furry pad (Fig. 20E);
furry pad composed of short, terminally-branching bristles
that grade into multifurcating scales (Fig. 20F).

First maxilla: Bell-shaped sternite distinctly set off from
coxa by sutures at posterolateral margin. Coxal process
conical, bearing 8-9 simple setae on tip and scattered along
medial edge, without significant clustering at tip (Fig. 20G);
three additional setae more posteriorly; cluster of tiny, barb¬
like basiconic sensilla between coxal process and telopod.
Distal article of telopod with two rows of long, plumose
setae along inner margin; branches developed along distal
half of these setae; rest of distal article evenly scattered
with numerous simple setae, slightly shorter than those on
coxal process (Fig. 20H).

Second maxilla (Fig. 20I-K) as described above for P.
(Paralamyctes) monteithi (no differences in proportions,
setal patterns or structure, or pretarsus form observed).

Strong, sharp distal spinose projections on tibia of legs
1-15. Prefemur, femur and tibia fairly evenly scattered with
distinctly pigmented (orange-brown) setae of varied length,
at most moderately long; several setae encircling distal
margin of femur; tarsal setae more uniformly slender; weak
development of two rows of setae ventrally along tarsus,
without pairing on each side of leg. Articulation between
tarsomeres relatively weak on anterior legs, continuous
dorsally, anterior tarsi usually preserved unflexed.
Distitarsus about 50% length of basitarsus on anal leg; anal
leg basitarsus about 12 times longer than broad (Fig. 16D).
Pretarsal claws relatively long, gently curved; pair of accessory
claws symmetrical, 30% length of main claw (Fig. 16D).

Sternites with longitudinal median furrow not well
defined on anterior segments, usually lightly impressed on
anterior third of sternite from mid trunk, not significantly
developed behind midlength on posterior segments.
Posterior margins of most sternites convex, subtransverse
on posterior few segments in both sexes. Sternal setation
consists of a few fairly long setae on anterolateral and
posterolateral margins, transverse band at about one-quarter
length of sternite; sternal setae of equivalent diameter to
largest setae on legs.

Coxal pores on legs 12-15. Coxal pore field set off from
anteroventral face of coxa by curved edge, without fold or
groove. Pores usually round, some slightly ovoid (Fig.
19B,C); most females with 4444/4444 coxal pores, one with
4333/3333; most males with 3333/3333 pores, sometimes
one or a few coxae with two pores.

Male (Fig. 19C): Tergite of intermediate segment broadly
barrel-shaped, with transverse posterior margin, relatively
narrower than in female; other tergites without sexual
dimorphism. Sternite of first genital segment undivided,
setae evenly scattered, with a transverse row just in front of
posterior margin; margin convex backwards between
gonopods. Gonopod of three articles and tapering, needle¬
like terminal process, the first two articles each bearing three
or four relatively long setae, those on first article in a transverse
row, third article bearing two or three setae; terminal process
as long as rest of gonopod. Penis a prominent cone.

Female (Fig. 19B): Tergite of intermediate segment with
more concave posterior margin than in male. Tergites of
first genital segment and telson fairly well sclerotised,
though much less so than other tergites; first genital tergite
bearing several setae along its posterolateral margin; telson
tergite with numerous slender setae posterolaterally. Sternite
of first genital segment equal in width to S15; margin
concave between gonopod condyles, with rounded
posteromedial bulge; most of sternite bearing moderately
dense, short setae (Fig. 21G), several longer setae at
posterolateral margin; first article of gonopod bearing three
or four pointed spurs (five specimens with 3+3 spurs, one
with 3+4), all about equal in size, curved such that tip points
dorsally (Fig. 2IF,FI); many fairly long setae on first article,
about seven similarly long setae on second article (Fig.
21H); third article lacking setae.

Discussion. Close affinity to the Tasmanian P. mesibovi n.sp.
is most obviously indicated by female gonopod morphology.
Distinction between the two species is addressed under P.
mesibovi.

Paralamyctes (Nothofagobius) mesibovi n.sp.

Figs. 16C, 21I-M, 22, 23

Etymology. For Robert Mesibov, guru of Tasmanian
myriapodology, who discovered this species.

Diagnosis. Paralamyctes (Nothofagobius) with narrow
dental margin of maxillipede coxosternite bearing four teeth;
simple aciculae on mandible; distal spinose projection on
tibia of legs 14 and 15; first article of female gonopod with
three spurs (all shared with P. cassisi ). Nearly all antennal
articles substantially longer than wide; cephalic pleurite
elongate; posterior margin of tergite 7 relatively deeply
sinuate; female gonopod spurs on a long, slender projection.

Type material. Holotype Queen Victoria Museum and Art
Gallery (QVMAG) 23/23009, $ (Figs. 16C, 22A), Wombat Hill,
Waratah, Tasmania, CQ 702 064, 41°29'00"S 145°26'42"E, 680
m, R. Mesibov, 19 Sep 1990. Paratypes: QVMAG 23/41725,
1 $ (Fig. 22B), from type locality; collection details as for
holotype; QVMAG 23/23010, 1 $ (Figs. 21I-M, 23), Wombat
Hill, Waratah, CQ 704 066 (ca. 250 m from type locality),
41°28'53"S 145°26'51"E, R. Mesibov, 30 Sep 1990.

Edgecombe: Australian Paralamyctes 233

Figure 22. Paralamyctes (Nothofagobius) mesibovi n.sp. A, holotype $ QVMAG 23/23009, Wombat Hill, Waratah,
Tasmania; scale 1 mm. B, paratype $ QVMAG 23/41725, Wombat Hill, Waratah, Tasmania, terminal segments
andgonopods; scale0.25mm. C, S Q VM AG 23/11817, Western Creek, Tasmania, terminal segments and gonopods;

scale 0.25 mm.

Other material. QVMAG 23/11817 IS (Fig. 22C), Western
Creek, Tasmania, DP 581 848,41°41T4"S 146°29'47"E, 1150 m,
R. Mesibov, 8 Aug 1993; QVMAG 23/21593, 19, Simons Rd,
Tasmania, EQ 435 212, 41°21'33"S 147 o 31T2"E,800 m, P.
Greenslade, 8 Mar 1989; QVMAG 23/21594, 1 9, Mt Michael,
Tasmania, EQ 845 406, 41°10'53"S 148°00'26"E, 740 m, H.
Mitchell, 29 Nov 1989; QVMAG 23/41377, IS (DNA voucher
specimen), Coldstream River, Tasmania, CQ 768027, 600 m,
41°31'03"S 145°31'24"E, R. Mesibov, 9 Mar 2000.

Description. Because this species is identical in most
respects to P. cassisi, described in full above, only features
of potentially diagnostic value and standard meristic
characters are described here, as well as colour for a specimen
in near absolute ethanol (these data lacking for P. cassisi ).

Length of head shield up to 1.7 mm. Head and tergites
orange, with red-brown mottled network concentrated in
longitudinal median band and near borders; antennae deep

234 Records of the Australian Museum (2001) Vol. 53

Figure 23. Paralamyctes (Nothofagobius) mesibovi n.sp. Paratype $ QVMAG 23/23010, Wombat Hill, Waratah, Tasmania. Scanning
electron micrographs. A, gnathal lobe of mandible, scale 20 pm; B,M, mandibular aciculae and fringe of branching bristles, scales 20
pm, 5 pm; C, accessory denticles on mandibular teeth, scale 10 pm; D, furry pad on mandible, scale 10 pm; E,F, ventral view of head,
showing clypeus and labrum, scales 200 pm, 60 p; G, first maxillae, scale 60 pm; H,I, coxal processes and distal segments of telopod of
first maxillae, scales 20 pm; J, branching bristles on labral margin, scale 5 pm; K, second maxilla, scale 60 pm; L, tarsus and claw of
second maxilla, scale 5 pm.

orange; maxillipede pale orange; sternites yellow, without
significant purple/chestnut mottling; tarsi of similar pale
orange/yellow colour to proximal part of leg. 25-28 antennal
articles, all but one or two articles substantially longer than
wide. Eye strongly domed, white. Cephalic pleurite
elongate, of typical size and shape for genus (Figs. 211,
23E). Anal leg basitarsus 13-17 times longer than broad
(Fig. 16C). Four coxal pores on legs 13-15 in females, two

to five on leg 12; large male with three coxal pores on each
of legs 12-15. Penis apparently concealed above sternite
of first genital segment in both known males. Terminal
process on male gonopod equal in length to second and
third articles of gonopod (Fig. 22C). Female gonopods as
for P. cassisi except for longer spur-bearing process on first
article (Fig. 22B).

Edgecombe: Australian Paralamyctes 235

Discussion. The morphology of the female gonopods is
particularly distinctive in P. mesibovi, it and P. cassisi n.sp.
being the only Paralamyctes with three (rather than the
plesiomorphic two) spurs. In both species the spurs are based
on a longer projection than is developed in P. (N.) chilensis,
this peculiar character certainly being a synapomorphy.
These species are similar in several other characters that
are rarely developed or unique within Paralamyctes.
Probable synapomorphies include: four large teeth on the
maxillipede coxosternite (Fig. 21L,M); a relatively short
fringe of labral bristles, with the bristles branching into
several rami (Figs. 21C, 23J) rather than having many
branches arising as pectinations from a main shaft (Figs.
11D, 13K, 18J); a continuous field of accessory denticles
on the mandible, with the denticles being a distinctive
triangular shape (Figs. 20D, 23C); simple, non-pinnulate
aciculae on the mandible, with a few strong lineations on
the sides of the aciculae (Figs. 20B, 23M) and; a distal
spinose projection present on the tibia of leg 15.

Distinction between the Tasmanian and northern New
South Wales taxa can, in fact, be made based on only a few
characters. Paralamyctes mesibovi is distinguished by its
more elongate cephalic pleurite (Fig. 20L versus Fig. 211),
more embayed posterior margin of tergite 7 (Fig. 19A versus
Fig. 22A) and by the longer, more slender projection of the
first article of the female gonopod (Figs. 19B, 21G,H versus
Fig. 22B). The pleurite of P. mesibovi is identical to that of
most other species of Paralamyctes , and certainly represents
the plesiomorphic state relative to the peculiarly shortened
pleurite of P. (N.) cassisi. The Tomosvary organ is of the
same absolute size in the two species, but it occupies a larger
area of the pleurite in P. (N.) cassisi. Paralamyctes (N.)
mesibovi has fewer short antennal articles, only a few articles
being of subequal length and width versus several in P.(N.)
cassisi. The two species also appear to differ with respect
to the length of the terminal process on the male gonopod
(Fig. 19C versus Fig. 22C), but since only one gonopod is
known from a relatively small male of P. mesibovi (the
gonopods being unexposed on the second, larger male
QVMAG 23/41377), the significance of this difference
should not be overstated. Paralamyctes cassisi has a
relatively longer terminal process than any other Australian
Paralamyctes , whereas the terminal process of P. mesibovi
is considerably shorter (compare Figs. 19C, 22C).

As noted above, the affinities of P. mesibovi and P. cassisi
appear to be with P. chilensis from southern Chile and Argentina
(see treatment of that species below). Paralamyctes mesibovi
and P. cassisi are both distinguished from P. chilensis by
their longer antennae, with a larger number of articles
(usually 19 in P. chilensis), a larger Tomosvary organ, the
absence of a groove delimiting the accessory denticle field
on the mandible (Figs. 20D, 23C versus Fig. 25H), simple
(Figs. 20A-C, 23M) versus bipinnulate (Fig. 251,K)
aciculae, a typically higher coxal pore count, presence of a
distal spinose projection on the tibia of the anal leg, and a
longer spur-bearing process on the female gonopod.

Based on the distribution of the P. mesibovi/P. cassisi
clade in Tasmania and northern New South Wales, a
predicted occurrence of the group might be in Nothofagus
forests in southern Victoria. A comprehensive sampling of
Nothofagus forests in the Otway Range, Strezlecki Range,

and Victorian Central Highlands by the Museum of Victoria
has not yet uncovered any Paralamyctes (examination of
this collection by the author in February 2000).

Paralamyctes (Nothofagobius) chilensis
(Gervais in Walckenaer & Gervais, 1847)

Figs. 16E, 24, 25

Henicops chilensis Gervais in Walckenaer & Gervais, 1847: 239.
Henicops chilien.-G ay, 1849: 65, fig. 6a,b.

Henicops chilensis- Silvestri, 1899: 242.

Paralamyctes chilensis- Silvestri, 1905: 749.

Paralamyctes chilensis.-Attems, 1928: 67.

Paralamyctes chilensis. -Archey, 1937: 81, 82.

Paralamyctes chilensis. -Chamberlin, 1955: 54.

Type material. Syntypes have not been identified; Gervais
(1847) had access to two specimens, collected by M. Claude
Gay from an unspecified locality in Chile. Two specimens
(6 and ?: BMNH [E] 200408 Chilo.l910-.6.17.39) from
Patagonia in the Natural History Museum, London, were
labelled as probable syntypes by R.E. Crabill. This
determination is unlikely because antennal segmentation
of one specimen disagrees substantially with the original
description of the species, and the specimens are labelled
as part of the Silvestri collection, dated 1900. The types
have not been identified in the Museum national d’Histoire
naturelle, Paris (J.-J. Geoffroy, pers. comm. 2000).

Material examined. TMAG collection by R. Mesibov, 1983—
Chile, Xth region: 19, Proyecto Polincay saw mi ll complex, under
bark of fallen Drimys winteri along road to sawmill, ca. 20 km
NE of Puerto Montt, 13 May 1983; juvenile, Proyecto Polincay,
lightly cut-over Nothofagus dombeyi/Saxegothea conspicua forest
above sawmill, ca. 20 km NE of Puerto Montt, 10 May 1983;
2 9 9 (Fig. 24A,B), 1 S, Correntoso, under bark of fallen trees in
heavily cut-over forest S of park office (Parque Alerce Andino), 5
Jun 1983; 29 9 , Lago San Antonio, Isla Chiloe, under bark of
fallen tree near creek in forest owned by I. Martini of Quellon, 9
Jun 1983; 2<S 6 (Figs. 24C, 25), Lenca, under bark on track, cut¬
over rainforest ca. 3 and 5 km E of sawmill, 14 Jun 1983; 1 6 and
1 9, Maullfn, lightly cut-over nadi forest ca. 7 km along road to
Carelmapu, 15 Jun 1983. ZMUC collection—Chile: 19,
Osorno 53: Parque Nacional Puyehue, Anticura, 300 m, Mision
Cientffica Danesa, 7-8 Mar 1979; 29 9 , Osorno 37: Parque
Nacional Puyehue, Antillanca, 1100-1300 m, Nielsen and
Karsbolt, 14Nov 1981. Argentina: 1 9 (Fig. 16E),RioNegro7:
S.C. de Bariloche, Colonfa Suiza, 810 m, Mision Cientffica
Danesa, 22 Nov 1978; sex undet., Neuquen 18: Lago Tromen,
Rodeo Grande, 900 m, Mision Cientffica Danesa, 12 Mar 1979.

Diagnosis. Paralamyctes (Nothofagobius) with relatively
short antenna of 17-21 (usually 19) articles; Tomosvary
organ relatively small; 4 or 5 teeth on dental margin of
maxillipede coxosternite; short, sharp pinnules on both sides
of mandibular aciculae; distal spinose projection on tibia
of legs 1-14; two or three coxal pores on legs 12-15 in
both sexes, fourth pore small when present; two spurs on
female gonopod set on short process.

Discussion. Paralamyctes (Nothofagobius) chilensis is
poorly known from published accounts, and previous
attempts to decipher its morphology are in part contra-

236 Records of the Australian Museum (2001) Vol. 53

Figure 24. Paralamyctes (Nothofagobius) chilensis (Gervais in Walckenaer & Gervais, 1847). A,B, 9 TMAG,
Correntoso, Parque Alerce Andino, Xth Region, Chile; B, terminal segments and gonopods; scales 1 mm, 0.25 mm.
C, S TMAG, Lenca, Xth Region, Chile, terminal segments and gonopods; scale 0.25 mm.

dictory. The description by Gervais (1847) notes few
potentially diagnostic attributes, many characters being
general for henicopids. Gervais described 17 antennal
articles, whereas the illustration of a specimen in Gay (1849)
depicted 17 articles on one antenna and 18 on the other.
Following a description by Silvestri (1905), Attems (1928)
listed purportedly diagnostic characters that disagree with
the description and illustrations by Gervais and Gay (i.e.,
posterior margin of T7 with a deep angular notch, the angles
toothed; antenna 19 jointed).

The only locality data published for Paralamyctes chilensis
are Silvestri’s (1905) record of it at San Vicente (Talcahuano).
A single species of Paralamyctes (Nothofagobius) is the only
henicopid represented in collections made by R. Mesibov from
near Puerto Montt and Maullfn and on Isla Chiloe in the
Xth region of Chile (see list of sites in “Material examined”);
this material is conspecific with the BMNH specimens cited
under “Type material”, and the same species occurs in
adjacent parts of the Argentinian Lake District (Fig. 16E).
Most specimens have 19 antennal articles, thus matching

Edgecombe: Australian Paralamyctes 237

Figure 25. Paralamyctes (Nothofagobius) chilensis (Gervais in Walckenaer & Gervais, 1847). 8 TMAG, Lenca, Xth Region, Chile.
Scanning electron micrographs. A,B, ventral views of head, showing clypeus and labrum, scales 50 pm; C,D, branching bristles on
labral margin, scales 10 pm, 5 pm; E, dental margin of maxillipede coxosternite, scale 50 pm; F, second maxillae, scale 100 pm; G, claw
of second maxilla, scale 5 pm; H, gnathal lobe of mandible, scale 20 pm. I,K, mandibular aciculae, scales 10 pm, 5 pm; J, fringe of
branching bristles on mandible, scale 5 pm; L, tarsus and claw of second maxilla, scale 20 pm; M, cephalic pleurite, scale 20 pm.

Silvestri’s (1905) description, though the margins of T7 are
untoothed and usually not deeply notched (Fig. 24A). Only
one specimen (ZMUC: San Carlos de Bariloche, Argentina)
has a subangular notch in the posterior margin of T7. The
only specimen with more than with more than 19 antennal
articles is one of the two identified by Crabill as probable
syntypes; it has 21/19 articles. Silvestri (1905) cited 5+5

coxosternal teeth on the maxillipede in P. chilensis ;
specimens examined in this study have 4+4 or 5+5 teeth
occurring in about equal frequency (Fig. 25E). Because a
single species of Paralamyctes is present in the examined
Patagonian collections, I consider it likely that this species
is P chilensis, though the vagaries of prior taxonomic
treatments do not make this certain. Regardless of

238 Records of the Australian Museum (2001) Vol. 53

nomenclatural ambiguities, the species documented here
confirms the presence of Paralamyctes in Patagonia.

Paralamyctes (Haasiella) Pocock, 1901

= Wailamyctes Archey, 1917

Type species. Henicops insularis Haase, 1887 (=
Wailamyctes munroi Archey, 1923).

Diagnosis. Paralamyctes with median furrow on head shield
extending behind transverse suture; mandibular aciculae
bipinnulate; tarsi of legs 1-12 fused into single article.

Assigned species. Wailamyctes halli Archey, 1917;
Wailamyctes trailli Archey, 1917; undescribed species from
Tasmania ( Wailamyctes sp. of Mesibov, 1986).

Distribution. New Zealand (South Island, Stewart Island,
Auckland Islands, Campbell Island), Tasmania.

Discussion. Pocock (1901) erected Haasiella for the
aberrant Henicops insularis Haase, 1887, from the Auckland
Islands. Johns (1964) indicated that a subsequently-named
Auckland Islands species, Wailamyctes munroi Archey,
1923, is a subjective synonym of H. insularis, the holotype
of which merely has unregenerated anal legs. With this
synonymy, Wailamyctes Archey, 1917, which had served
(e.g., Archey, 1937) to group the New Zealand clade
composed of H. insularis (= W. munroi ), H. trailli (Archey,
1917), and H. halli (Archey, 1917), is recognized as
Haasiella. Haasiella is employed as a subordinate taxon
(subgenus) within Paralamyctes because its removal renders
Paralamyctes paraphyletic; molecular sequence data
indicate a particularly close relationship between P.
(Haasiella) and P. (Thingathinga) (Edgecombe etal., 2001).
Description of the only known Australian member of P.
(Haasiella), the Tasmanian species identified by Mesibov
(1986), is in preparation.

Key to Australian genera of Lithobiomorpha and species of Paralamyctes

To assist in the identification of species described in this work and place these species in the context of
Lithobiomorpha known from Australia, the following key is offered. Characters are specific to taxa
occurring in Australia (i.e., large genera like the introduced Lithobius may deviate from certain of these
characters in other regions).

1 Cluster of ocelli. Pleurite of maxillipede segment interrupted
ventromedially, not forming a band between maxillipede
coxosternite and stemite of first pedigerous segment. Tibia lacking
distal spinose projection. One or more articulated spurs at distal
ends of at least a few podomeres on all legs. Female gonopod
with dorsal and ventral denticle on each side of main claw. Male
gonopod small, single segmented. Anal pores absent except in

larva and early post-larval stages. 2 (Lithobiidae)

-Ocellus single or absent. Pleurite of maxillipede segment

continuous ventromedially, forming a band between maxillipede
coxosternite and sternite of first pedigerous segment. Distal
spinose projection on tibia of at least legs 1-11. No articulated
spurs at distal ends of podomeres. Female gonopod simple, lacking
denticles against main claw. Male gonopod flagelliform, composed
of three articles and seta-like terminal process. Anal pores present

in adult. 3 (Henicopidae)

2 23 or more antennal articles. Outer tooth on maxillipede
coxosternite of similar size to inner tooth/teeth, more anteriorly
placed than inner tooth/teeth. Porodont at outer edge of maxillipede
tooth row or between teeth on outer part of row. Tergites smooth

or gently wrinkled. Lithobius

-17-22 (overwhelmingly 20) antennal articles. Outer tooth on

maxillipede coxosternite smaller and more posteriorly placed than
inner teeth. Porodont among the maxillipede teeth on inner half
of tooth row, typically between second and third teeth (as
numbered from medial to lateral). Tergites with pronounced
longitudinal wrinkling

Australobius

Edgecombe: Australian Paralamyctes 239

3 14-17 antennal articles. Spiracle absent on first pedigerous

segment. Coxal pores lacking on legs 12-13. Coxa of leg 15 with

long, lobate process ending in a spine. Prefemur of leg 15 with a
single ventral spur.

. 4 (Anopsobiinae)

-18 or more antennal articles. Spiracle present on first pedigerous

segment. Coxal pores present on legs 12-15. Coxa of leg 15
lacking distal spinose process. Prefemur of leg 15 lacking a ventral
spur.

. 5 (Henicopinae: Henicopini)

4 Spiracle present on segment 8.

. Anopsobius

-Spiracle absent on segment 8.

. Dichelobius

5 Median furrow absent or confined to anterior part of head shield.

Small, triangular sternite on first maxilla. First genital sternite of
male divided longitudinally.

. 6

-Median furrow on head shield deep, continuous to transverse

suture. Large, bell-shaped sternite on first maxilla. First genital
sternite of male undivided.

. 8 (Paralamyctes)

6 All tergites with rounded posterior angles. Tarsi of legs 1-12

unjointed, tarsi of legs 13—15 bipartite. Distal spinose projection
absent on tibia of legs 13-14.

. 7

-Several tergites with projections on posterior angles. Tarsi of legs

1-12 bipartite or tripartite, tarsi of legs 13-15 at least tripartite.
Distal spinose projection present on tibia of legs 13-14 .

. Henicops

7 Ocellus absent. Most antennal articles evenly short.

. Lamyctinus

-Ocellus present. Antennal article length uneven, with short, paired

articles interspersed between groups of longer ones.

. Lamyctes

8 Tarsi of legs 1-12 unjointed. Median furrow on head shield extends

behind transverse suture. Aciculae on mandible bipinnulate.

. Paralamyctes (Haasiella)

-Tarsi of legs 1-12 bipartite. Median furrow on head shield

terminates at transverse suture. Aciculae on mandible simple or
with pinnules along dorsal side only (not bipinnulate).

. 9

9 Tergite 1 markedly narrower than head shield. Basal article of

female gonopod extended as a process bearing three spurs.. 10 (. Paralamyctes (Nothofagobius))

-Tergite 1 of similar width to head shield. Basal article of female

gonopod not extended, bearing two spurs. 11

10 Several antennal articles as wide as long. Posterior margin of tergite

7 gently concave. Spur-bearing process on female gonopod
moderately long.

. P. (Nothofagobius) cassisi

-Nearly all antennal articles substantially longer than wide.

Posterior margin of tergite 7 deeply sinuate. Spur-bearing process
on female gonopod long, slender.

.. P. (Nothofagobius) mesibovi

11 Maxillipede tarsungulum relatively short, with pretarsal part equal

in length to tarsal part. Teeth on maxillipede coxosternite large,

pointed. P. (Paralamyctes) neverneverensis

Maxillipede tarsungulum long, slender, with pretarsal part
substantially longer than tarsal part. Teeth on maxillipede
coxosternite small, blunt.

12

240 Records of the Australian Museum (2001) Vol. 53

12 All antennal articles longer than wide. Ocellus bulging. Tomosvary
organ on margin of head, with cephalic pleurite constricted behind
organ. Aciculae of mandible with pinnules along dorsal side.

Posterior margin of tergite 7 with shallow median embayment . P (Paralamyctes) monteithi

-Some antennal articles as wide as long. Ocellus gently convex or

flattened. Tomosvary organ on surface of cephalic pleurite.

Aciculae of mandible simple, lacking pinnules. Posterior of margin

of T7 with strong, medially-transverse embayment. 13

13 Ocellus gently convex. Dental margin of maxillipede gently
sloping backwards laterally. Strong joints between tarsomeres on

legs 1-12 . P. (Thingathinga) grayi

-Ocellus flattened. Dental margin of maxillipede nearly transverse.

Faint trace of joints between tarsomeres on legs 1-12 . P. (Thingathinga) hornerae

Acknowledgments. Collection studies were hosted by M.
Colloff (Australian National Insect Collection), T. Crosby and G.
Hall (New Zealand Arthropod Collection), P. Johns (University
of Canterbury), P. Lawless (Queensland Museum), J. Margerison
(Natural History Museum) and K. Walker (Museum of Victoria).
H. Enghoff, G. Giribet, C. Griswold, M. Hamer, R. Mesibov, P.
Parrillo and G. Viggiani kindly arranged loans. I thank G. Cassis,
S. Davis and G. Milledge for assistance collecting in Queensland,
and G. Giribet, M. Nishiguchi and Y.-y. Zhen for collecting with
me in northern New South Wales. R. Mesibov is thanked for heroic
collecting in Tasmania and Chile. I am grateful to B. Elizabeth
Horner for sponsoring taxonomy in the Australian Museum.
Collaboration with G. Giribet sharpened ideas about homologies.
S. Lindsay prepared material and operated the SEM. S. Bullock
and Y.-y. Zhen skillfully prepared drawings and plates,
respectively. For advice and comments on the manuscript, I am
grateful to R. Mesibov, W. A. Shear and M. Zapparoli. My wife, Zerina,
is thanked for being a good sport on innumerable collecting trips.

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Manuscript received 25 August 2000, revised 22 October 2000 and
accepted 15 November 2000.

Associate Editor: D.J. Bickel.

© Copyright Australian Museum, 2001

Records of the Australian Museum (2001) Vol. 53: 243-246. ISSN 0067-1975

A New Seahorse Species
(Syngnathidae: Hippocampus )
From the Great Barrier Reef

Michelle L. Horne

Department of Marine Biology & Aquaculture, James Cook University, Townsville Queensland 4811, Australia

Michelle .Horne @ j cu.edu. au

Abstract. A new seahorse, Hippocampus queenslandicus (family Syngnathidae) is described from
northern Queensland, Australia. Diagnostic characters include meristics: 15-18 dorsal-fin rays, 16-17
pectoral-fin rays, 10-11 trunk rings, 34-36 tail rings, and the presence of body and tail spines, as well as
a moderately low coronet with five distinct spines.

Horne, Michelle L., 2001. Anew seahorse species (Syngnathidae: Hippocampus ) from the Great Barrier Reef.
Records of the Australian Museum 53(2): 243-246.

Seahorses, pipefishes and seadragons collectively belong
to the family Syngnathidae. Syngnathids occur in coastal
waters of temperate and tropical regions of the world in
habitats ranging from sand, seagrass beds to sponge,
algae, rubble and coral reefs (Vincent, 1997; Kuiter,
2000). A recent revision of the seahorses, genus
Hippocampus , recognizes 32 species world-wide (Lourie
et al., 1999). The number of valid Australian seahorse
species has been estimated at seven (Gomon, 1997) and
13 (Lourie et al., 1999).

Materials and methods

A total of 226 specimens (height range: 56-143 mm, 111
males, 115 females) of anew Hippocampus species, together
with five specimens of H. zebra Whitley, 1964, and four of
H. dahli Ogilby, 1908, were collected between October 1997
and December 1998. Seahorses found dead in trawl nets
were immediately frozen. Upon return to shore, frozen

seahorses were placed in FAACC (formaldehyde-acetic
acid-calcium chloride fixative) for 48 hours then removed
to 100% ethanol.

Macroscopic description of seahorses included sex,
number of body segments and colour morphs. Standard
seahorse measurement protocol was followed (Lourie et al.,
1999). Meristic values were recorded to within 0.1 mm using
dial callipers and include; height (measured from top of
crown to tip of tail, HT), wet weight, head length (HL),
snout length and snout depth, eye diameter (horizontal
measurement of the left eye), tail length, dorsal-fin length
(length of fin base) and abdominal width. In males, the
pouch length (measurement from top of opening slit to
ventral point where pouch meets tail) was also recorded.
Using a stereo dissector the number of rays in the dorsal,
anal and pectoral fins of each individual were counted. All
specimens examined, including types, were collected by
the author. Types are deposited in the Museum of Tropical
Queensland in Townsville (QM), Museum Victoria (NMV)
and the Australian Museum, Sydney (AMS).

244 Records of the Australian Museum (2001) Vol. 53

Hippocampus queenslandicus n.sp.

Hippocampus species 4 (Kuiter, 2000)

Holotype, QM 124445, <3, HT 103.9 mm, collected by
trawl, 50 m depth off John Brewer Reef, 18°37'S 147°03'E,
October 1999. Paratypes: QM 124446 $, HT 102.8 mm,
collected with holotype; NMV A 21578 6, collected by
trawl, 50 m depth off Britomart Reef, 18°14'S 146°35'E,
February 1999; NMV A 21579 4 $$, collected by trawl, 50
m depth off Britomart Reef, February 1999; AMS 1.40832-
001 $, HT 100 mm, collected by trawl, 40 m depth off Bait
Reef, 19°47'S 149°06'E, December 1998; AMS 1.40833-
001 <5", HT 111.2 mm, collected by trawl, 50 m depth, off
John Brewer Reef, 18°37’S 147°03’E, November 1998.

Non-type material: 217 specimens, 115$?, Ill SS, (HT
range 56-143 mm), collected by trawl, 30-50 m depth, north
Queensland, Great Barrier Reef, October 1997-December
1998.

Diagnosis. Number of dorsal fin-rays 15-18, (17 in 67%
of specimens). Pectoral fin-rays 16-17, (17 in 77% of
specimens). Snout length 35.9-54.2%, snout depth HL 8.2-
11.2%; coronet moderately low HT 2.3 mm (2.2-2.4 mm)
with 5 spines; inferior tail ridge with moderate spines along
pouch section of males, filaments absent; nasal spine absent;
1-2 cheek spines, length 1.4 mm (1.2-1.8 mm); eye spine
present, 1.1 mm (0.9-1.5 mm) in length; first trunk ring
spine large, 2.8 mm (2.4-2.9 mm) in length; lateral head
spine 1.8 mm (1.17-2.1 mm) in length; trunk rings 10 or
11; tail rings 34-36.

Description. Values are for the holotype, with ranges given
between parentheses based on other material examined
(Table 1). Holotype height (HT) 103.9 mm (56-143.4 mm).
Wet weight of holotype 6.89 g (1.8-14 g). The head
prominent and relatively mobile, maintained at an angle of
approximately 90° to the axis of the trunk. Head length
(HL) 27.9% HT (30.3-35.8). Snout length 47% HL (35.9-

first trunk spine
pectoral fin
trunk rings
cheek spine

dorsal fin

tail rings

Figure 1 . Composite diagram of male Hippocampus queenslandicus.
Scale 1 cm.

54.2). Snout depth 9.3% HL (8.2-11.2). Eye diameter 11.1%
HL (5.2-16.4). Abdominal width 14.5% HT (12.8-23.0).
Tail length 51% HT (37.8-64.4). Holotype pouch length
23% HT (11.2-27.7). Colour patterns vary among
individuals (see colour photos in Kuiter, 2000: 38-39);
holotype orange; colour range among paratypes: red, purple,
white, brown, silvery grey and orange.

Table 1 . Morphometric values for Hippocampus queenslandicus holotype and opther specimens (paratypes and non-type material).

holotype

(mm)

holotype

proportion

n

range

(mm)

range

proportion

height (HT)

103.9

—

226

56-143.4

—

head length (HL)

29.0

27.9% HT

226

17-39.7

30.3-35.8% HT

snout length

13.6

47.0% HL

226

6.1-18.6

35.9-54.2% HL

snout depth

2.7

9.3% HL

226

2.3-3.1

8.2-11.2% HL

eye diameter

3.2

11.1% HL

226

1.2-4.8

5.2-16.4% HL

abdominal width

15.1

14.5% HT

226

7.2-28.9

12.8-23% HT

tail length

53.2

51.0% HT

226

28-81.3

37.8-64.4% HT

pouch length

23.8

23.0% HT

111

11.2-33

11.2-27.7% HT

wet weight (g)

6.9

—

226

1.8-14.0

—

Horne: new Great Barrier Reef seahorse 245

Table 2. Selected meristic values for three Hippocampus species; data from Lourie et al. (1999) marked by asterisk.

species

dorsal fin-rays
(mode)

pectoral fin-rays
(mode)

anal fin-rays

trunk rings
(mode)

tail rings
(mode)

H. queenslandicus

15-18 (17)

16-17 (17)

4

10-11 (10)

34-36

H. taeniopterus *

17-18

15-18 (16)

4

11

34-38

H. spinosissimus *

16-20

16-19 (17)

4

11

33-39

The holotype coronet is moderately low and has five
distinct, pointed spines, forming a pentamerous crown (Fig.
1). A single distinguishable spine occurs above each orbit,
termed “eye spine” (length in holotype 1.1 mm). A single
prominent lateral spine on the sides of the head preceding
the coronet, termed “head spine” is apparent (length in
holotype 1.8 mm). Holotype has one large cheek spine, some
paratypes with 2, length 1.4 mm. The first spine on the dorsal
first trunk ring is extremely prominent, (length 2.8 mm in
holotype, Fig. 1). Inf erior tail ridge with moderate spines
along pouch section of males. Nasal spine is absent in this
species. Filaments are absent in this species.

Holotype possesses 10 trunk rings (range 10-11) and 34
tail rings (range 34-36), with the trunk of moderate length.
The male pouch occupies most of the abdominal cavity and
is ventral to the last two trunk and first seven tail segments.
The dorsal fin is relatively small with 17 rays (range 15-
18) and a length of 8.0 mm (range 5-10.5 mm). The number
of pectoral fin-rays in holotype is 17 (range 16-17) and
there are 4 anal fin-rays.

Distribution and ecology. Hippocampus queenslandicus
was recorded from Feather Reef (17°00'S 146°10'E), East
of Innisfail in the northern section of the Great Barrier Reef,
to Bait Reef (19°47'S 149°06'E). The highest recorded
numbers over the 14-month sampling period were at John
Brewer Reef (18°37'S 147°4E), East of Ingham. The species
was collected by trawl, at depths between 30 and 50 m and
was usually caught in a sponge or seagrass habitat, often
attached to hard and soft coral species.

Comparisons. Species in the genus Hippocampus are often
distinguished by fin-ray counts together with number of
body, dorsal and tail rings (Whitley & Allen, 1958; Lovett,
1969; Yari, 1982; Lourie et al., 1999).

Along with this new species there are a number of other
seahorse species occurring in the north Queensland waters
of the Great Barrier Reef, namely Hippocampus zebra
Whitley, 1964, H. bargibanti Whitley, 1970, H. dahli
Ogilby, 1908, H. taeniopterus Bleeker, 1852, and an
undescribed species: the wing-spined seahorse (Lourie et
al., 1999; Kuiter, 2000). Hippocampus zebra is a small
species (HT range in this study 79.2-88.4 mm), with
distinctive black (or dark brown) and white stripes over its
external body and head, clearly differentiating it from H.
queenslandicus. Hippocampus bargibanti differs from H.
queenslandicus by its small size (adult height < 20 mm)

and its “fleshy” appearance, mostly without recognizable
body rings and its extremely short snout. The external
surface of H. dahli is relatively smooth, with reduced, low
spines that form ridges around each body segment, in
contrast to the spiny external surface of H. queenslandicus.
Hippocampus dahli is long, slender and thin, with a narrow
head, flattened body shape and with the coronet laterally
flattened. The wing-spined seahorse often displays filaments
on the larger spines of the head and back, clearly differing
from H. queenslandicus which completely lacks filaments.

Hippocampus queenslandicus superficially resembles
two other species of the genus, H. taeniopterus and H.
spinosissimus Weber, 1913. Hippocampus queenslandicus
differs in its spinous ornamentation and fin-ray counts
(Table 2). Hippocampus taeniopterus a species often
confused with Hippocampus kuda Bleeker, 1852 (see Kuiter,
2000), is a commonly occurring inshore species in the
northern section of the Great Barrier Reef. Hippocampus
taeniopterus differs from H. queenslandicus in having 17-
18 dorsal fin-rays, 15-18 (mode of 16) pectoral fin-rays,
11 trunk rings and 34-38 tail rings (Table 2). Hippocampus
taeniopterus has a deep cheek area, a thick snout and a deep
abdominal area giving it a more rounded and stout
appearance than H. queenslandicus. Hippocampus
taeniopterus lacks well-developed spines, appearing
relatively smooth, with a number of irregularly spaced,
rounded tubercles covering the external surface (Dawson,
1986; Lourie et al., 1999; Kuiter, 2000), in contrast to the
spinous exterior of H. queenslandicus. The coronet of H.
taeniopterus is more rounded than the five-spined coronet
of H. queenslandicus and the coronet points backwards on
the head of H. taeniopterus. Male H. taeniopterus are often
drab coloured with striations over the head and small black
spots over the trunk; females are often yellow with
several large dark spots on the trunk. Male and female
H. queenslandicus often are bright orange or red. Another
notable difference between these two species is the
differing sizes of the young: the average height of
juvenile H. queenslandicus is 5.63 mm, whereas juvenile
H. taeniopterus examined by Rick Brayley (pers. comm.,
1999) are almost three times larger.

Hippocampus spinosissimus has similar meristic values
to those of H. queenslandicus (Table 2). However, H.
spinosissimus has a large eye spine, a medium to high
coronet with four or five spines, a generally spinous exterior
and strongly developed, blunt spines bordering the pouch
in males (Lourie et al., 1999). By comparison, H.

246 Records of the Australian Museum (2001) Vol. 53

queenslandicus, has a relatively reduced spinous exterior,
lacks long pouch spines and has a shorter coronet with four
or five spines. Hippocampus spinosissimus has not been
recorded in the northern waters of the Great Barrier Reef
to date.

In summary, H. queenslandicus differs in meristic
values and physical appearance from other seahorse
species found in the northern section of the Great Barrier
Reef and from similar species found elsewhere (Lourie
et al., 1999; Kuiter, 2000). This species seems restricted
to the northeast coast of Queensland, Australia, hence
the name H. queenslandicus.

Acknowledgments. I am grateful to GBRMPA and James Cook
University for their financial support throughout this project. I
am most grateful to Col Meyers and the Townsville trawlermen
for their collection assistance. The illustration was generously
provided by Natale Snape. I thank A. Vincent and R. Kuiter for their
expertise and assistance. Finally 1 am most grateful to Andrew Beer.

References

Bleeker, P., 1852. Bijdrade tot de kennis der icthyologische fauna
van Singapore. Naturrkundig. Tijdschrift Nederlandsch Indie
3:51-86.

Dawson, C.E., 1986. Syngnathidae. In Smith's Sea Fishes , eds.
M.M. Smith & P.C. Heemstra, pp. 445-458. Johannesburg:
Macmillan.

Gomon, M.F., 1997. A remarkable new pygmy seahorse
(Syngnathidae: Hippocampus ) from south-eastern Australia,
with a redescription of H. bargibanti (Whitley) from New
Caledonia. Memoirs of the Museum of Victoria 56(1): 245-
253.

Kuiter, R.H., 2000. Seahorses. Pipefish and Their Relatives. A
Comprehensive Guide to Syngnathiformes. Chorleywood
(United Kingdom): TCM Publishing.

Lourie, S.A., A.C.J. Vincent & H.J. Hall, 1999. Seahorses: An
Identification Guide to the World’s Species and Their
Conservation. London: Project Seahorse.

Lovett, J.M., 1969. An introduction to the biology of the seahorse
Hippocampus abdominalis. Unpublished Honours thesis.
University of Tasmania, pp. 19.

Ogilby, J.D., 1908. New or little known fishes in the Queensland
Museum. Annals of the Queensland Museum 9: 1-41.

Vari, R., 1982. The seahorses (subfamily Hippocampinae). In
Fishes of the Western North Atlantic. Part 8. Order
Gasterosteiformes, Suborder Syngnathoidei. Syngnathidae
(Doryrhamphinae, Hippocampinae), eds. J.E. Bohlke, D.M.
Cohen, B.B. Collette, W.N. Eschmeyer, R.H. Gibbs, T.W.
Pietsch Jr, W.J. Richards, C.L. Smith & K.S. Thomson. Memoir
Sears Foundation for Marine Research 1(8): 173-189.
Vincent, A.C.J., 1997. Sustainability of seahorse fishing.
Proceedings of the 8th International Coral Reef Symposium 2:
2045-2050.

Weber, M., 1913. Die Fische der Siboga-Expedition. Leiden: E.J.
Brill.

Whitley, G.P, 1964. Fishes from the Coral Seas and the Swain
Reefs. Records of the Australian Museum 26(5): 145-195.
Whitley, G.P., 1970. [Bargibant’s seahorse from New Caledonia].
Abstract of Proceedings, Ordinary Gen. Meeting, 20th
November. Proceedings of the Linnean Society of NSW 94(3):
292-295.

Whitley, G., & J. Allen, 1958. The Seahorse and its Relatives.
Melbourne: Griffin Press.

Manuscript received 7 September 2000, revised 1 May 2001 and accepted
7 May 2001.

Associate Editor: J.M. Leis.

© Copyright Australian Museum, 2001

Records of the Australian Museum (2001) Vol. 53: 247-254. ISSN 0067-1975

Two New Species of Fairy Shrimp
(Crustacea: Anostraca: Thamnocephalidae: Branchinella )
from the Paroo, Inland Australia

Brian V. Timms

School of Geosciences, University of Newcastle, Callaghan NSW 2265, Australia
ggbvt@alinga.newcastle.edu.au

Abstract. Two new species of fairy shrimp, Branchinella budjiti and B. campbelli, are described
from intermittent wetlands in northwestern NSW. Both have very distinctive frontal appendages, second
antennae and other features that defy classification into established groups within the genus.

Timms, Brian V., 2001. Two new species of fairy shrimp (Crustacea: Anostraca: Thamnocephalidae: Branchinella )
from the Paroo, inland Australia. Records of the Australian Museum 53(2): 247-254.

The genus Branchinella Sayce, 1903, is one of the most
speciose genera of fairy shrimps in the world with 33 species
so far described (Belk & Brtek, 1995). Geddes’ (1981)
review of Australian Branchinella recognized 18 species,
but Belk & Brtek (1995) elevated three of his subspecies to
specific rank, enlarging the Australian fauna to 20 species,
all endemic. Forty percent of these are known from a very
limited area or just the type locality in the arid zone. It is
not surprising then that intensive collecting in one such area,
the middle Paroo catchment of northwestern NSW and
southwestern Qld, has yielded many new distribution
records and two new species (Hancock & Timms, in review;
Sanders, 1999; Timms, 1993, 1997). In addition, a new
genus in the family Branchipodidae and a new species of
Parastreptocephalus (Streptocephalidae) have been found
(Timms, unpublished data). It is the purpose of this paper
to describe the new species of Branchinella and to attempt
to place them into the groupings of Branchinella proposed
by Geddes (1981).

Taxonomy

Thamnocephalidae
Branchinella Sayce, 1903

Branchinella budjiti n.sp.

Fig. 1

Type material. Holotype: S, Australian Museum (AM
hereafter), AM P58543, length 16.1 mm; allotype: 1 $,
AM P58544, length 13.9 mm; paratypes ASS (length
15.9+0.6 mm) and 3 $$ (length 13.3+0.6 mm) AM P58545.

Type locality. An unnamed claypan c. 500 m NW of Muella
Station homestead, 130 km NW of Bourke, NSW, 29°31’S
144°56'E, altitude 120 m a.s.l., collected by the author, 6
December 1999.

248 Records of the Australian Museum (2001) Vol. 53

Other material. 20 66, 10?? from Sues Pan, Bloodwood
Station, 132 km NW Bourke, NSW, AM P58546; collected
by the author 17 November 1999. Unnamed Yapunyah
swamp, 22 km NE of Currawinya homestead on Eulo Rd,
Currawinya National Park, Queensland, Queensland
Museum (QMB hereafter) W25187; collected by the author
18 May 1996. Collections by author from 18 sites, some at
multiple dates during 1988-2000, on Bloodwood, Muella
and Tredega Stations, 130-150 km NW Bourke. Collections
by author from 28 sites, some at multiple dates during 1987-
2000, at Currawinya National Park, via Hungerford, Qld.
Collections by author from 5 sites during 1995-2000 on
Rockwell Station, 145 km SW of Cunnamulla, Qld.

Etymology. Named in honour of the Budjiti tribe, first
human inhabitants of the middle Paroo country.

Description of male

Length 16.1 mm. Cephalothorax slightly longer than the
abdomen minus the cercopods. Antenna I reaches a little
beyond the apex of the basal segment of antenna II (Fig.
1A). Antenna II (Fig. IB) basal segment at c. 70° to body
axis (instead of c. 20-40° as in typical Branchinella species).
This segment with two raised pads anterodistally bearing a
few short thin setae, and an outgrowth at its posterolateral
corner. This outgrowth lamellar, smooth and serrate on the
anterior margin and about one third to half the length of the
distal segment. Distal segment a little longer than basal
segment, sclerotized, flattened and bending medially c. 20°.
Medial margin with a medial swelling and bearing small
serrations. Apex tapering and truncated.

Frontal appendage (Fig. 1C) large and elaborate, carried
curled beneath the thorax and reaching up to half its length
and doubling the depth of the thorax. Basal section fused
into a trunk bearing three long papillae on the lateral
margins. This supports two large wing-like branches, each
a little longer than wide and bearing an expanded distal
margin, particularly at the posterolateral corners.
Numerous papillae on the lateral margins and lower
surface; those on a central thickened area underneath each
branch longer than the rest, except for c. 7 long papillae
on the outer distal corners of each branch. Density of
papillae varies, least on the ventral area of the fused trunk
and inner area of the branches, and most numerous towards
the outer distal corners.

Thoracopods (Fig. IF) with endopodite smoothly
rounded (but first thoracopod has a shallow notch
anterodistally) and bearing long setae. On thoracopod 5,
but not thoracopods 1 and 11, the bases of the anterodistal
region of endopodite with a rounded knob on one side.
Endites 1 and 2 very setose and each with an anterior setae,
the first about half the length of the second. Endites 3-4
each bear 2 anterior setae and endite 5 bears 1 anterior setae
on phyllopods 1, 5 and 11. Exite longer than endopodite
but narrower and more setose. Epipodite sausage-shaped
and smooth. Preepipodite expanded and serrated; no notch
on its border.

Penis (Fig. ID) short, extending back one or at most two
abdominal segments when everted. Armed with c. 9 large
asymmetrical teeth on the lateral (convex) face and with
numerous long papillae on the side and medial (concave)
face. These papillae shorter near the distal end. Distal end lobed
on the inner curvature and bears a harder asymmetrical tooth
on the outer apex. Base of penis bare of teeth or papillae.
No swelling lateral to the penes. Cercopods of telson (Fig.
IE) heavily setulated on both margins and subequal in length
to the last two abdominal segments.

Description of female

Length 13.9 mm. Antenna I almost twice the length of the
eye and eye stalk (Fig. 1G). Antenna II (Fig. 1G) equal in
length to antenna I. Foliaceous, about twice as long as wide
and ending in a narrow, sharp protrusion from a shouldered
distal portion. Numerous small thin setae on the distal
margin. Brood pouch (Fig. 1H) extending posteriorly over
a little more than four abdominal segments. Distal part
curved away from abdomen and with a small neck before
the ovipore. Thoracopods similar to those of male, but knobs
on the bases of endopodite setae not as well developed.
Cercopods as in the male.

Description of other material

Length variable but around 10-15 mm in males and slightly
smaller in females (Table 1).

The frontal appendage of the male in life can be opened
into a large sheet considerably wider than the body and
about the length of the thorax. Trunk of frontal appendage
with 2-5 (usually 3-4) papillae on lateral margins. Outer
distal corners of each branch of frontal appendage with 5-

Table 1 . Length (mean+SD) of Branchinella budjiti from different localities. Ten males and ten females from each
locality measured, except entry marked ( a ): only 7 specimens measured.

locality

length of 66

length of ??

RA Pan, Currawinya NP (Dec 96)

9.7+1.0 mm

8.5+0.8 mm

RC Pan, Currawinya NP (Jun 99)

14.6+0.9 mm

13.0+1.8 mm

Sues Pan, Bloodwood (May 98)

15.3+1.2 mm

14.7+0.8 mm a

Goat Pan, Muella (Jun 99)

14.2+1.0 mm

12.6+0.9 mm

Timms: new Branchinella species 249

Figure 1. Branchinella budjiti n.sp. Male holotype A-F, female allotype G and H. A, lateral view of whole animal;
B, dorsal view of antenna II with frontal appendage removed; C, ventral view of frontal appendage; D, penis; E,
telson and cercopods; F, anterior view of right 5th phyllopod; G, dorsal view of head, antennae I and II; H, lateral
view brood pouch and associated abdominal segments. Each scale bar represents 1 mm.

250 Records of the Australian Museum (2001) Vol. 53

8 longer papillae. Papillae on lateral margins and lower
surface variable in density between specimens, though they
are always sparcest on the ventral areas of the fused trunk
and inner area of branches and most numerous towards the
outer distal corners.

Endites 3-5 of thoracopods normally bear 2 setae but
sometimes only one is present and this occurs mostly on
endite 5.

In females the size and shape of the brood pouch is
variable, with the length varying between four and five
abdominal segments and the small neck before the ovipore
sometimes absent.

Differential diagnosis

In Geddes’ (1981) key to Australian species of Branchinella,
B. budjiti straddles both halves of couplet 9, so it needs to
be separated the step beforehand—after couplet 7 part 2
(basal segments of antenna II with medial swellings or
outgrowths elsewhere). This can be achieved by adding
a couplet:

- Trunk of frontal appendage shorter than

branches, branches flattened, large and with
many papillae, especially on posterolateral
margin. B. budjiti

—— Trunk of frontal appendage longer than
branches and if branches expanded distally,
then relatively small and no concentration
of papillae on posterolateral margin.9

The only species likely to be confused with B. budjiti is
B. denticulata from Kalgoorlie, Western Australia. Both
have a lamellar outgrowth from the posterodistal corner of
the basal segment of antenna II (not the anterodistal corner
in B. denticulata as shown in Linder [1941] and Geddes
[1981]), and both have the two branches of the frontal
appendage expanded distally. However, the lamellar
outgrowth in B. budjiti always has a markedly serrate
anterior margin (with 4-6 serrations), but in B. denticulata
the anterior margin is weakly serrate (with c. 2 serrations)
as shown in Linder (1941) and repeated in Geddes (1981)
and as seen in the paratypes (C. Rogers, pers. comm.).
Lurthermore, the lamellar outgrowth is carried parallel to
the axis of the basal segment in B. budjiti, but in B.
denticulata its position is variable, either at right angles (as
reported by Linder [1941] and repeated by Geddes [1981]),
or parallel with the axis of the basal segment (C. Rogers,
pers. comm.). It is possible that this variation in position of
the lamellar outgrowth is an artifact due to preservation;
how the lamellar outgrowth is carried in life is unknown.
The frontal appendages of the two species are also
superficially similar, but in B. budjiti the trunk is short and
the two branches much longer, while the reverse applies in
B. denticulata. The diagram of B. denticulata in Linder
(1941) and repeated by Geddes (1981) is unrepresentative,
showing perhaps a juvenile specimen, since adults among
the paratypes examined by C. Rogers (pers. comm.), have
the branches more expanded and with papillae on the edges

and some denticles or spines scattered over the dorsal
surface. The difference between the two species in the
branches of the frontal appendage lies in much greater
relative size and the larger number of posterolateral papillae
in B. budjiti. Lurther differences between the two species
are in antenna II in the adornment of the basal segment and
in the shape of the second segment. In B. denticulata the
basal segment has a few papillae scattered on its medial
upper surface and the second segment is evenly arched and
lacks any thickening along its length. By contrast in B.
budjiti there are two medial distal swellings bearing hairs
on the basal segment and the second segment has a medial
swelling and is unevenly bent at this point. In conclusion
the two species seem closely related, but are distinctly
different in many key features.

Distribution and habitat

Widespread and common in temporary turbid waters in the
middle portion of the Paroo catchment in northwestern NSW
and southwestern Qld (Sanders, 1999). The most common
habitat is a very turbid (Secchi disc depth <5 cm) claypan
that holds water for a month or two after filling (Sanders,
1999; Timms, 1997).

Branchinella campbelli n.sp.

Fig. 2

Type material. Holotype: S, AM P58547, length 12.4
mm; allotype: 1$, AM P58548, length 15.4 mm;
PARATYPES: KMc? (length 12.2+0.6 mm), 10?? (length
13.0+1.2 mm), AM P58549.

Type locality. Lake Muella, Tredega Station, 140 km NW
Bourke, NSW, 29°3FS 144°53’E, altitude c. 119 m a.s.l.
Collected by the author 17 November 1999.

Other material. Live males and three females from Carters
swamp, 12 km N of homestead, Muella Station, 130 km
NW of Bourke, NSW, AM P58550. Collections by the
author from five localities, some on multiple occasions,
during 1995-2000 on Bloodwood, Muella and Tredega
Stations, 130-150 km NW Bourke.

Etymology. Named for Steve Campbell, Manager of Muella
Station for many years.

Description of male

Length 12.4 mm. Cephalothorax slightly longer than the
abdomen minus the cercopods. Antenna I a little longer than
the basal segment of antenna II (Lig. 2A). Antenna II (Lig.
2A) of normal relative size and angle of repose (c. 20-40°
to body axis) for Branchinella. Bases fused for almost half
their length. Basal segment with a well defined mound about
one-third of the way along the free inner surface. Mound
clothed with numerous very short thin setae. Apical segment
about equal in length to the basal joint, curved and tapering
to a wide point. Lor much of its length the inner surface
bears numerous weakly developed transverse ridges.

Timms: new Branchinella species 251

Figure 2. Branchinella campbelli n.sp. Male holotype A-D, female allotype E and F. A, dorsal view of head,
antennae I and II with frontal appendage removed; B, dorsal view of frontal appendage; C, penis; D, anterior view
of right 5th phyllopod; E, dorsal view of head, antennae I and II; F, lateral view of brood pouch and associated
abdominal segments. Each scale bar represents 1 mm.

Frontal appendage (Fig. 2B) moderately large and
elaborate, approximately equal in bulk to antennae II proper
and reaching almost half way along the cephalothorax. Basal
section fused into an unadorned trunk, though the trunk is

ringed. Terminal branch with an unadorned flange
protruding medially and attached by a wide stem to the inner
margin about two-fifths along its length; and expanded
basolaterally into a roughly equilateral triangle tapering

252 Records of the Australian Museum (2001) Vol. 53

distally into a long flagellum-like process. The outer margin
of the expanded base and flagellum bordered with finger¬
like papillae, each terminating in a spine. These papillae
number c. 35. The first 20 papillae of similar size, but those
towards the tip of the flagellum decrease in size until the
last few are swellings rather than papillae.

Fifth thoracopod (Fig. 2D) with endopodite broadly
rounded and bearing numerous setae. No knobs on the bases
of endopodite setae. Endites 1 and 2 heavily setulated and
each with a stout anterior seta, the first about half the length
of the second. Endite 3 and 4 with 2 anterior setae and endite
5 with one anterior setae. Exite much longer than
endopodite, narrow and with many more and longer setae
than endopodite. Epipodite sausage-shaped and smooth.
Preepipodite expanded and with a few small sharp denticles
on its distal border; no notch on this border. Thoracopods 1
and 11 essentially similar to 5th thoracopod, but smaller
and slightly different in proportions.

Penis (Fig. 2C) short, extending back one abdominal
segment when everted. Armed on the outer lateral (convex)
side with about five broadly based asymmetrical teeth and
on the medial (concave) side with numerous (c. 15-21)
sharp teeth crowded distally. Innerside teeth more
symmetrical, but curved, particularly those near the tip of
the penis. Area between the two sets of teeth with thin
papillae and some weakly developed, broad-based teeth.
No swelling lateral to the penes.

Telson with cercopods heavily setulated on both margins.
Cercopods equal in length to the last two abdominal
segments.

Description of female

Length 15.4 mm. Antenna I (Fig. 2E) about one and three-
quarters the length of the eye and eye stalk. Antenna II (Fig.
2E) equal in length to antenna I, foliaceous, almost three
times as long as wide and tapering to a sharp protrusion
distally. Numerous small thin setae on the distal margin.
Brood pouch (Fig. 2F) deep-bodied and extending
backwards over almost four abdominal segments.
Thoracopods and cercopods as in the male.

Description of other material

Length variable between 8 and 15 mm (Table 2).

In live males the long apical flagellum of the frontal
appendage reaches three quarters the length of the
cephalothorax and the basolateral part of the side branches
reach somewhat beyond (by a factor of one and a half) the
cephalothorax. In many mature specimens this basolateral
expansion of the side branches protrudes at its distal corner
to make another flagellum-like process, but not as long as
the apical process.

Differential diagnosis

In Geddes’ (1981) key to Australian species of Branchinella,
B. campbelli keys to couplet 11, but neither alternative
offered in this couplet is applicable. An additional couplet
is needed after couplet 10:

-Frontal appendage with two simple

branches. 11

- Frontal appendage with each branch having

medially directed flange and a large
triangular expansion basolaterally. B. campbelli

Branchinella pinnata and to a lesser extent B. frondosa
and B. arborea are the species most like B. campbelli, all
bearing ramification (fine branches of each main branch)
of the frontal appendages. In B. frondosa and B. arborea
many of these sub-branches of the frontal appendage are
further branched (secondary ramification) whereas in B.
pinnata and B. campbelli all sub-branches are whole. In B.
pinnata most sub-branches of each branch are long and
some are placed on the upper inner surface of each main
branch. However in B. campbelli each sub-branch is very
short and papilla-like (so much so that they do not seem to
be the main structure of the frontal appendage as when they
are long in B. pinnata, B. arborea and B. frondosa ).
Branchinella campbelli therefore should not key to these
species in couplet 5 of Geddes’ key. Also none of the sub-

Table 2. Length (mean+SD) of Branchinella campbelli from different localities. Number of specimens measured in
brackets.

locality and date

length of 33

length of 22

Lower Crescent Pool (Bloodwood) (Oct 96)

15.0 mm (1)

18.0 mm (1)

Carters Swamp (Dec 97)

8.0+0.0 mm (2)

8.7+0.4 mm (6)

Lake Muella (Jun 98)

11.9+1.6 mm (9)

14.0+0.2 mm (4)

Carrols Swamp (Jun 99)

9.2+0.8 mm (6)

10.2+1.0 mm (5)

Timms: new Branchinella species 253

branches/papillae is placed on the inner surface of the two
main branches in B. campbelli. Furthermore, this species
has an inwardly directed flange along the medial surface of
each branch and a large triangular expansion basolaterally
on each branch. A further distinguishing feature between
the two species is the presence of a spine at the distal end of
the first segment of antenna II in B. pinnata.

Distribution and habitat

Restricted to a small area about 50 km 2 on Bloodwood,
Muella and Tredega Stations, 130-150 km NW of Bourke,
NSW. It is generally found in temporary clear-water
wetlands with few macrophytes in the water column, but
occasionally occurs in limited numbers in creek pools
where it is probably washed in from wetlands elsewhere
(Sanders, 1999).

Discussion

These two species are distinctive and easily distinguished
from other Australian species of Branchinella. In B. budjiti
the placement of the basal segments of antennae II at almost
right angles to the body axis is characteristic, though a few
other species, particularly B. affinis, approach this
arrangement. Furthermore the lamellar outgrowth at the
posterobasal corner of the basal joint is an uncommon
feature, as is the large sheet-like frontal appendage.
Branchinella campbelli is equally remarkable by virtue
of its elaborate frontal organ, which has no close
counterpart in other species. Certainly the flange
protruding medially along the inner margin of each branch
and the plate-like basal expansion with all its finger-like
papillae are unique. Females of both species, like most in
Branchinella , are unremarkable.

It is difficult to accommodate either species into any of
the three groups of Branchinella erected by Geddes (1981).
Certainly neither fit into his group I ( B. australiensis, B.
occidentalis, B. compacta, B. nichollsi, B. buchananensis,
B. hattahensis ) which are large, sturdy bodied, and males
have lateral swellings to the penes and females have a long
antenna II. Thoracopods have elongated distal endites only
sparsely setulated medially and there are large numbers (10-
20) of anterior setae on endites 3-5 of thoracopod 1. Both
of the new species are of small to medium size (<15 mm),
have a rounded distal endite, endites 3-5 with few (<3)
anterior setae, no lateral swelling to the penes in males, and
short antenna II in females.

Group II ( B. affinis, B. apophysata, B. denticulata, B.
latzi, B. longirostris, B. probiscida) animals are small to
medium in size, with short distal endites, 2-4 anterior setae
on endites 3-5 of thoracopod I, a frontal appendage
consisting of a long trunk and two (usually simple) branches,
and swellings lateral to the penes (Geddes, 1981).
Branchinella budjiti has some affinities with this group by
reason of small size, the gross structure of the frontal
appendage, short distal endite and presence of 2-3 anterior
setae on endites 3-5 of the cercopods. However it lacks
lateral swellings to the penes, and the frontal appendage
has a short trunk and greatly expanded branches.

Likewise B. campbelli shares many features with Group
III ( B. arborea, B. basispina, B. dubia, B. frondosa, B.
pinnata, B. wellardi) such as frontal appendage complexly
branched, no swellings lateral to the penes, absence of spines
on the basal part of the penis, and endites 3-5 of thoracopod
I with 2-7 anterior setae. Even so, endites 3-5 are small
and not particularly bubble-like, and the animal is small
compared to many other species in the group.

Two species ( B. lyrifera and B. simplex ) do not fit Geddes’
groupings. Probably B. budjiti also should be ungrouped
but it has some affinities with Group II, especially with B.
denticulata , while B. campbelli is perhaps best considered
a non-core member of Group III.

Branchinella budjiti is the sixth Australian Branchinella
with an outgrowth from the distal end of the basal joint of
antennae II. In B. nichollsi, B. hattahensis and B.
buchananensis the outgrowth is from the inner corner and
is about as long as the second segment (see Geddes, 1981:
fig. 5a,b,c). Relative development of lateral papillae varies
between the three species, from none in B. hattahensis, to a
few papillae on the medial side in B. nichollsi, to many
papillae on the lateral side in B. buchananensis (Geddes,
1981). In B. budjiti and B. denticulata the lamellar
outgrowth is smaller and protrudes from the posterodistal
comer. There are minor differences between the two species
in its structure and angle of repose (see “Differential
Diagnosis”). Branchinella apophysata also has a lamellar
outgrowth of moderate length from the distal end of the
first segment (Geddes, 1981: fig. 8f) but insufficient is
known on its detailed structure for further comments to be
made. In summary it seems that the outgrowths in B.
nichollsi, B. hattahensis and B. buchananensis may be
homologous (which is supported by Geddes considering
them as subspecies within B. nichollsi ), and those in B.
denticulata and B. budjiti are quite different but perhaps
also homologous. The outgrowth of B. apophysata may
have some affinities with those in B. budjiti and B.
denticulata. Interestingly, the first three species are placed
in Group I and the second three in or near to Group II,
showing that lamellar outgrowths are not basic features in
the classification of Branchinella into groups, though the
detail of the outgrowth may be of some value. Neither are
lamellar outgrowths important generic determinants, for
besides being found in some Branchinella, they occur in
the related genus Thamnocephalus (Moore & Young, 1964).

The relationships among Australian Branchinella are
further clouded by another character, a medial swelling on
the basal joint of the second antennae, shared between
otherwise unrelated species. Such a swelling occurs in both
of the new species and in five other species shared between
all three groups ( B. australiensis, B. compacta, B.
probiscida, B. affinis and B. arborea ) as well as one (B.
lyrifera ) that does not fit into Geddes’ groups. The swellings
take different forms and are probably not homologous—
those in B. campbelli are large and bulbous and look like
the swellings in B. affinis and B. lyrifera, while those in B.
budjiti are simply raised pads with thin setae, perhaps like
those of B. arborea. These similarities cross Geddes’
groupings (B. campbelli is similar to a Group II species
and an ungrouped species, while B. budjiti is included with

254 Records of the Australian Museum (2001) Vol. 53

a Group III species). Because of these problems and others
mentioned by Geddes (1981) meaningful grouping and
phylogeny of Australian Branchinella probably cannot
be achieved by conventional analysis, but DNA
sequencing should be useful and is in progress (Hebert
& Timms, in prep.)

Finally it is pertinent to note that while B. budjiti has a
wide distribution within the Paroo, B. campbelli occurs in
a very limited area (c. 50 km 2 ) despite searching over 3,000
collections over 13 years in the Paroo and adjacent areas.
Given that nine species are known only from their type
locality ( B. apophysata, B. basispina, B. hattahensis ) or
from a very limited area ( B. buchananensis, B. campbelli,
B. denticulata, B. nichollsi, B. simplex, B. wellardi ), all
in the arid zone (Geddes, 1981), it is possible that more
species of Branchinella await discovery in remote areas
of Australia.

Acknowledgments. I wish to thank the owners of Bloodwood,
Muella and Tredega Stations for access to their properties and
Steve Campbell, former manager of Muella Station, for hospitality
and assistance in the field. I am also grateful to Christopher Rogers
for information on B. denticulata and for comments on the
manuscript.

References

Belk, D., & J. Brtek, 1995. Checklist of the Anostraca.
Hydrobiologia 298: 315-353.

Geddes, M.C., 1981. Revision of Australian species of
Branchinella (Crustacea: Anostraca). Australian Journal of
Marine and Freshwater Research 32: 253-295.

Hancock, M., & B.V. Timms, in review. Ecology of turbid clay
pans in the Paroo, semi-arid Australia, with special reference
to invertebrate succession. Hydrobiologia.

Linder, F., 1941. Contributions to the morphology and the
taxonomy of the Branchiopoda Anostraca. Zoologiska Bidrag
fran Uppsala 20: 102-303.

Moore, W.G., & J.B. Young, 1964. Fairy shrimps of the genus
Thamnocephalus (Branchiopoda, Anostraca) in the United
States and Mexico. Southwestern Naturalist 9: 68-77.

Sanders, P.R., 1999. Biogeography of Fairy Shrimps (Crustacea:
Anostraca) in the Paroo, Northwestern Murray-Darling Basin.
Honours Thesis, University of Newcastle.

Sayce, O.A., 1903. The Phyllopoda of Australia, including
descriptions of some new genera and species. Proceedings of
the Royal Society of Victoria 15(2): 224-261, pis. 27-36.

Timms, B.V., 1993. Saline lakes of the Paroo, inland New South
Wales, Australia. Hydrobiologia 267: 269-289.

Timms, B.V., 1997. A Study of the Wetlands of Cur raw iny a
National Park. Report to the Queensland Department of
Environment, University of Newcastle, Newcastle.

Manuscript received 12 June 2000, revised 19 October 2000 and accepted

1 December 2000.

Associate Editor: D.J. Bickel.

© Copyright Australian Museum, 2001

Records of the Australian Museum (2001) Vol. 53: 255-262. ISSN 0067-1975

Scaptodrosophila aclinata :

A New Hibiscus Flower-breeding Species
Related to S. hibisci (Diptera: Drosophilidae)

Shane F. McEvey 1 and J.S.F. Barker 2

1 Australian Museum, 6 College Street, Sydney NSW 2010, Australia
shanem @ austmus. gov. au

2 School of Rural Science & Natural Resources, University of New England, Armidale NSW 2351, Australia

sb arker @ metz. une. edu. au

Abstract. Physiological, ecological and evolutionary studies of Scaptodrosophila hibisci have led to
recognition of a second species in the Northern Territory (Australia) which is described here as
Scaptodrosophila aclinata n.sp. The new species is readily distinguishable by reference to the first
orbital: it is large and proclinate in S. hibisci and small and reclinate in S. aclinata. Scaptodrosophila
hibisci has been collected from the flowers of five Hibiscus species in eastern Australia and S. aclinata
uses eleven Hibiscus species in the Northern Territory. Only H. meraukensis is a host for both, and there
is no evidence of narrow host-specialization. The distributions are apparently disjunct. The two species
can be reared in the laboratory on cultured plants. Hybridization studies showed the two species to be
partially interfertile; S. aclinata has delayed sexual maturation and extended copulation latency when
compared to S. hibisci. This species pair is already the subject of various eco-physiological and
reproductive-biological studies because of so many useful experimental attributes: they are interfertile
and can be laboratory-cultured, their hosts and reproductive biology are known, they are abundant and
easy to find, and research is underpinned by extensive genetic information already available for
Drosophila.

McEvey, Shane F., & J.S.F. Barker, 2001. Scaptodrosophila aclinata: a new Hibiscus flower-breeding species
related to S. hibisci (Diptera: Drosophilidae). Records of the Australian Museum 53(2): 255-262.

There are about 300 drosophilid species recorded from
Australia, with some 90% of them described. The genus
Scaptodrosophila Duda, 1923 (for many years treated as a
subgenus of Drosophila but see Grimaldi [1990] for revised
status) has 81 named species and is by far the largest. The
predominance of Scaptodrosophila among the 36 genera
represented, is striking and distinguishes the Australasian

fauna from major drosophilid radiations in other regions—
Afrotropical, Neotropical and Hawaiian. In Australia, the
other large genera Drosophila (35 species), Hirtodrosophila
(31 species ),Leucophenga (25 species) and My codrosophila
(24 species) are much smaller by comparison. In general,
Drosophila species are attracted to fermenting fruit and may
be reared easily in the laboratory; whereas Scaptodrosophila

256 Records of the Australian Museum (2001) Vol. 53

species have, in most cases, unknown resource requirements
(van Klinken & Walter, 2001) and are difficult to rear in
the laboratory. Only 10 of the 35 Drosophila species
recorded in Australia are endemic and of these 10 only D.
birchii and D. serrata have provided useful research
opportunities. In contrast, Drosophila species that occur in
natural habitats in North America and Africa have provided
many important models in the study of evolution, behaviour,
physiology and ecology, with field observations being
further elaborated by genetic and controlled-laboratory
experimentation. The opportunity to explore evolutionary
and ecological aspects of the Australian Scaptodrosophila
radiation, has until recently, been severely hampered by the
lack of an amenable model for field and laboratory studies.

In this paper we report the discovery of a sibling species
of Scaptodrosophila hibisci that offers many of the same—
and some new—research opportunities as do some of the
important and well-documented Drosophila models. This
new species, Scaptodrosophila aclinata, is readily
distinguishable morphologically, has a very specific host-
plant relationship, can occur in very large numbers, can be
reared under laboratory conditions and can be induced to
hybridize (with some negative heterosis) with its sibling
species S. hibisci.

Scaptodrosophila hibisci (Bock in Cook etal., 1977) was
found to breed in flowers of Hibiscus splendens and H.
heterophyllus. Both these plant species have been recorded
from central Queensland to the Wollongong district in
southern New South Wales (Wilson, 1974). Collections of
S. hibisci have since been made from H. diversifolius in
New South Wales and Queensland, and from H. divaricatus
and H. meraukensis in Queensland (Starmer et al., 1997;
Wolf et al., 2000; Barker unpubl.). With its widespread
distribution in eastern Australia, and utilization of a number
of Hibiscus species as breeding sites, S. hibisci has already
become a model for the study of population structure and
genetic variation, and possible host-plant specialization.
Completed studies of this species include ecological aspects,
quantitative genetic analyses and reproductive biology
(Starmer et al., 1997, 1998, 2000; Polak etal., 1998,2001;
Wolf et al., 2000, 2001).

A number of Hibiscus species occur in the Northern
Territory and not in eastern Australia. Collections were made
in the Northern Territory from 11 Hibiscus species (H.
aneuthe, H. arnhemensis, H. byrnesii, H. cf. byrnesii, H.
fallax, H. menzeliae, H. meraukensis, H. petherickii, H.
riceae, H. symonii, H. zonatus) at 22 locations in May, 1998.
Differences between S. hibisci and the flies collected in the
Northern Territory were noted in terms of the ovariole-
number body-size relationship (Wolf et al., 2000), and in
microsatellite allele frequencies (Barker unpubl.). Here we
describe the Northern Territory fly as a new species, and
present results of host-plant specialization and its laboratory
hybridization with S. hibisci. Given the diverse Hibiscus
flora in northern Australia and the discovery of cryptic
flower-breeding Scaptodrosophila species in a variety of
Hibiscus species throughout the Afrotropical Region,
Lachaise & Tsacas (1984) predicted that sibling species of
S. hibisci would be found in northern Australia.

Taxonomy

Morphological terms and morphometric formulae have been
given previously (Grimaldi, 1987; McEvey, 1990). Material
has been lodged in the following museums:

AM Australian Museum, Sydney
ANIC Australian National Insect Collection, Canberra
NSMT National Science Museum, Tokyo
NTM Museum and Art Gallery of the Northern
Territory, Darwin

QMB Queensland Museum, Brisbane.

Specimens used for SEM images are preserved on stubs in the
Australian Museum SEM Unit. Wing-length was measured
from the humeral to the wing apex (W) cf. axillary area to
apex (L). Specimens have been individually numbered by
McEvey, this information is abbreviated “Reg.” below.

Scaptodrosophila aclinata n.sp.

Figs. 2, 6-8, 9-12

Type material. Holotype 6 , Nitmiluk NP, Northern Territory,
14°18.77'S 132°27.00'E, ex Hibiscus menzeliae flowers, March
2000, Rick Hope & J.S.F. Barker; Reg. 15345, Australian Museum
K118208. Paratypes (24 66, 40 $9, all Northern Territory): same
data as holotype but Reg. 15308-15310dd, QMB; Reg. 15311-
153169$, NTM; Reg. 15317-15318c?d, NTM; Reg. 15332-3 &
15335-8 99 , QMB; Reg. 15334 9 (AM K118230, SEM Unit); Reg.
15339-15344 (AM K118202-K118207, Reg.15342 in SEM
Unit)dd; Bardedjilidji Walk, nr Cahill’s Crossing [c. 12°26'S
132°58'E], Kakadu NP, Hibiscus flowers, 23 Feb. 1996, D.K.
McAlpine & G.R. Brown, Reg. 15368-15383 (AM K118209-
K118224) 99 and Reg. 15384-15388 (AM K118225-K118229)
66, AM; Bukalara Plateau, 46 km SSW of Borroloola [c. 16°26'S
136°04'E], 23 Apr. 1976, D.H. Colless, on Hibiscus flowers, Reg.
15389-15398 99 , Reg. 15399-15405 66, ANIC; McArthur River,
48 km SSW of Borroloola [c. 16°26'S 136°04'E], 14 Apr. 1976,
D.H. Colless, malaise trap, Reg. 154069 and 15407 6, ANIC.

Distinguishing features. All three orbital setae are reclinate,
and foretarsi are unmodified.

Description. Holotype measurements given with paratype
range between parentheses where appropriate.

Body length. 2.0 mm (2.0-2.2 mm).

Head. Arista with 3 short, straight rays above and 2
below, plus a small terminal fork. Frons slightly longer than
wide (fw:fl = 0.9); with numerous frontal hairs; blackish
brown, paler anteriorly (Figs. 2, 8). Ocellar-triangle also
blackish brown. Ocellars subequal in length to the
postocellar and first orbital setae. Pedicel and first
flagellomere yellowish brown. Carina prominent, narrow
between pedicels, broad and square below, upper surface
flat (Figs. 7, 8). Face yellowish brown. Palpus tan, rounded
with 68 setae apically and subapically and about 4 ventrally.
Gena curved, slightly broader anteriorly, about one tenth
greatest diameter of eye, o:j = 13 (10-16), o:ch =11 (10-
14). Vibrissa single. Eye dark reddish brown with dense
pile (Fig. 2). Orbitals short, barely distinguishable from
frontal hairs (especially or2), anterior most orbital (orl)
reclinate, or2 and or3 also reclinate, in approximate ratio
6:6:7, orl:or3 = 0.9 (0.8-0.9), orl:or2 = 1.0 (1.0-1.2) (Fig.
2). Ocellars (oc) short and pointing posterolaterally,

McEvey & Barker: a new Hibiscus- breeding drosophilid 257

Figures 1,2. Frontal setation of Scaptodmsophila hibisci, left (1), and S. aclinata n.sp., right (2). Note the prominent
proclinate first orbital (orl) in S. hibisci (broken off on left side) and its diminutive reclinate form in S. aclinata ; see
text for abbreviations. (Specimens: S. hibisci, Reg. 15327, coll. Bellingen NSW, JSFB; S. aclinata, Reg. 15334,
same data as holotype, head on stub in AM SEM Unit, rest of body in Collection). Scale bar = 100 pm.

postocellars (poc) as short as first orbitals, oc:orl = 1.0 (1.0-
1.3), poc:oc = 0.9 (0.9-1.3). Inner (iv) and outer (ov) vertical
setae longer than the orbitals, or3:iv = 0.6 (0.6-0.8), iv:ov
= 1.0 (0.8-1.1) (Figs. 6, 8).

Thorax. Mesoscutum subshining blackish brown.
Dorsocentrals in two pairs; posterior dorsocentrals about
twice the length of the anterior setae, and slightly shorter
than the anterior scutellar setae, adc:pdc = 0.5 (0.5-0.7),
pdc:asc = 0.8 (0.7-0.9). Scutellum and mesoscutum
concolorous. Acrostichals in 8 rows, 6 between dorso¬
centrals. Prescutellar setae developed, adc:pre.sc =1.0 but
less well developed and shorter (0.6) in some paratypes.
Halter yellowish brown. Fine propleural seta present.
Anepisternum bare. Katepisternal setae barely distinguish¬
able from hairs and all arising near upper edge of sternite,
sterno-index = 1.0, m:a kepst = 0.9 (0.7-0.9), p.kepst:pdc
= 0.3. Two short humerals; anterior supra-alar about twice
as long. Legs and halters concolorous and paler than
mesoscutum; forelegs with unmodified tarsi and with tarsal
hairs strongly curved; mid tibia with 3-4 apical bristles,
hind tibia with 2 short ventroapical bristles. Pre-apical
bristles absent or not differentiated.

Wing. Length from axillary area to apex 1.56 mm
(paratype range 1.45-1.78), length from humeral crossvein
to apex 1.36 mm; C-index 1.44 (1.25-1.88), 4v-index 2.19
(2.00-2.70), 4c-index 1.50 (1.29-1.67), 5x-index 1.63
(1.25-1.80), M-index 0.65 (0.52-0.78), ac-index 4.80
(3.60-5.71), C3fringe 0.60 (0.56-0.67). Third and fourth
longitudinal veins slightly convergent apically.

Abdomen. Uniformly dark brown, slightly paler than thorax.

Male terminalia (Figs. 9-12). Epandrium narrow, without
lateral or ventral broadening, pale tan, with a single large
seta ventrally and pubescent hairs restricted to small areas
posterodorsally, posterolaterally and narrowly along
posterior border in between. Cercus not indented, covered
entirely with short hairs and with long setae becoming
smaller and shorter ventrally (Figs. 9-10). Surstylus with

row of c. 12 short stout prensisetae along inner margin and
6-7 longer setae arranged irregularly behind them.
Hypandrium with two long submedian spines; aedeagus
expanded apically, with curved apodeme slightly bulbous
distally (Figs. 11-12); parameres rounded with cluster of
fine sensilla apically.

Female. Forelegs with tarsal hairs only slightly curved (cf.
strongly curved in males), otherwise external morphology
similar to male.

Female terminalia. Egg guide sclerotized with large
marginal teeth.

Distribution (Fig. 13). Northern Territory north of 17°S.

In January 2001 no Hibiscus plants were found west of
Charters Towers on the Barkly Highway, south of
17°11.70'S 133°28.08'E on the Sturt Highway (Northern
Territory) or southeast of Halls Creek in the Tanamai Desert
(Western Australia-Northern Territory). Mr Terry A.
Woodger (Richmond-based botanist, pers. comm.) reports
Hibiscus from the Selwyn Ranges (c. 21.5°S 140.5°E) and
further collecting in that region would be important in order
to determine the extent to which populations of S. aclinata
and S. hibisci are geographically isolated.

Other specimens examined. Specimens from eastern
Australia in the AM and previously determined as
Scaptodrosophila hibisci by Bock or McEvey were re¬
examined and found to be correctly identified. Mt Cahill
specimens (ANIC, see paratype series above) were found
to be incorrectly identified as hibisci. A series of
Scaptodrosophila aclinata flies from Tolmer Falls,
13°11.60’S 130°42.32'E, Litchfield NP, Northern Territory,
1998, J.S.F. Barker, were dissected and discarded—this
represents an additional locality for the new species.

Remarks. Scaptodrosophila aclinata n.sp. is closely related
to S. hibisci (Bock in Cook et al., 1977) because it has very

258 Records of the Australian Museum (2001) Vol. 53

Figures 3-8. Comparative views of the head of Scaptodrosophila hibisci (left) and S. aclinata n.sp. (right). View
of the back of the head showing supracervical setae: Fig. 3, S. hibisci (Reg. 15327); Fig. 6, S. aclinata n.sp. (Reg.
15334). Frontal setation and facial morphology, Figs. 4-5, S. hibisci (Reg. 15322, coll. Bellingen NSW, JSFB);
Figs. 7-8, S. aclinata n.sp. (Reg. 15342). Note the complete lack of proclinate setae (arrowed in hibisci Fig. 5) in
the anterior frontal half of S. aclinata n.sp. (Fig. 8).

McEvey & Barker: a new Hibiscus -breeding drosophilid 259

similar morphology (Figs. 1-8) and habitat preference, and
it produces progeny—albeit with reduced fertility—when
hybridized (Table 1). However, it is distinctly different by
virtue of the first orbital being proclinate and relatively
large in hibisci and reclinate and relatively small in
aclinata. Of less significance is that the humeral setae
are larger and the overall coloration darker in S. hibisci.
Other differences have been noted in ovariole-number to
body-size relationship (see “ Drosophila hibisci —Northern
Territory flies” in Wolf et al., 2000) and microsatellite allelic
frequencies (Barker unpubl.).

The new species keys to couplet 80 in Bock’s (1982)
key to the Australian species of Drosophila. Formation of a

Figures 9-12. Male terminalia of Scaptodrosophila aclinata n.sp.
(Reg. 15344, AM K118207) 9-10, hypandrium, caudal and lateral
views; 11-12, epandrium, ventral and lateral views.

triplet at that level with the addition of: “Frontal
macrochaetae greatly reduced, first orbital not proclinate...
aclinata” would lead to a correct identification.

Three other anthophilic drosophilids from northern
Queensland and New Guinea are superficially similar:
Scaptodrosophila moana (McEvey) from Torres Strait
and Cape York Peninsula, and S. aproclinata (Okada &
Carson) and S. paraguma (Okada & Carson) from Wau.
Scaptodrosophila moana has a very distinctive arista with
a single upper ray quite unlike the three rays above and
two below arrangement in aclinata n.sp.; moana also has
a well-differentiated and proclinate first orbital seta.
Scaptodrosophila aproclinata and S. paraguma have not
been examined but they are described as having only two
reclinate orbitals, a condition that would make them very
hard to separate from aclinata n.sp. However, aproclinata
is also described as having extraordinary tarsal modification
and finely pubescent arista (tarsi are unmodified and aristae
are not finely pubescent in aclinata n.sp.); while paraguma
is described as having an arista pubescent in the distal half,
a mesopleural (= anepisternal) seta, and a deeply constricted
cercus (the anepisternum is bare and the cercus is not
constricted in aclinata n.sp.). The prensisetae of the aclinata
surstylus are most unlike the arrangement in S. paraguma.

The unusually short rays of the arista and the overall
reduction in cephalochaetae appears to be characteristic of
a number of drosophilids associated with flowers.

Etymology. The specific name refers to the unusual
inclination of the first orbital seta—proclinate in most other
drosophilids including Scaptodrosophila hibisci but
reclinate in this species.

260 Records of the Australian Museum (2001) Vol. 53

Figure 13. Distribution of Scaptodrosophila aclinata (a) and S. hibisci (•) in Australia (Papua New Guinea record
for S. hibisci not shown). Hibiscus flowers examined for Scaptodrosophila flies (January 2001) without result (O).

Host-plant specialization

Two populations of S. hibisci, each derived from a locality
in nature that has only one of the two Hibiscus species, H.
heterophyllus or H. diversifolius, were used to test
preferences for oviposition of each population on each
species. Wild caught flies (50 c3(3, 70 $9) from Bellingen ( H .
heterophyllus 30°25.155'S 152°49.425'E) were set in a pop¬
ulation cage and maintained breeding on H. heterophyllus
flowers for six weeks. Wild caught flies (250(3(3, 340$$)
from Tyagarah {H. diversifolius 28°34.933'S 153°32.258'E)
were held at 20°C in sugar-agar vials for three days, and
then a population cage was set up for each population with
50 males and 50 females. One H. heterophyllus and one H.
diversifolius flower were added to each cage, each day. Two
days after addition to a cage, flowers were removed to sand
bottles (Starmer et al., 1998), and all emerging progeny
scored daily until there were no further emergences. After
28 days, all remaining flies in the cages were collected and
counted.

Results. Over the 28 days, the Tyagarah population derived
from H. diversifolius produced more progeny than the
Bellingen population from H. heterophyllus (mean progeny/
day = 14.0 and 9.6 respectively, P = 0.07), and survived

better (mean numbers at end of test period = 33(3(3, 29 $$
and 12(5<3, 15 $$ respectively). Hibiscus heterophyllus
flowers were preferred by flies from both populations (mean
progeny/day = 16.0 and 7.4 respectively, P < 0.001). The
regressions of proportion of progeny from//, heterophyllus
on day were not significant for either cage. Thus all two
way interactions were tested in ANOVA against population
of origin x Hibiscus species x day as error. None were
significant.

Discussion. For two species ( H . heterophyllus and H.
diversifolius ) which it does utilize in nature, S. hibisci
laboratory populations from each of these species in nature
produced more progeny on the former. However, as the
population of origin x Hibiscus species interaction was not
significant, there is no evidence for host plant specialization.

Both S. hibisci and S. aclinata n.sp. have been found
breeding only in flowers of the Furcaria section of the genus
Hibiscus in Australia. However, S. hibisci has been recorded
breeding in flowers of okra [Abelmoschus (= Hibiscus)
esculentus] in New Guinea (Okada & Carson, 1982), and
we have bred it on okra flowers in the laboratory.

Scaptodrosophila hibisci has been collected from flowers
of five Hibiscus species and S. aclinata from 11 species.
Only one of these Hibiscus species, H. meraukensis, is

McEvey & Barker: a new Hibiscus -breeding drosophilid 261

Table 1. Results of test crosses for hybridization, copulation latency and duration and interfertility between Scaptodrosophila hibisci

(h) and Scaptodrosophila aclinata n.sp. (a); ft. =

fertile.

mating

mating

number

number

%

copulation

copulation

number

progeny

no. pairs

mean

type

6 x $

pairs

mating

mated

latency 0

duration

pairs

number

tested

day

tested

<lh

<3h

<3h

(min)

(min)

tested for

to first

first

mean+sd

mean+sd

progeny

mean+sd

ft- egg

ft- egg

parent

hxh

18

16

nt b

0.89

19.3+19.1

5.6+3.9

8

66.0+33.1

12

1.83

ax a

11

7

10

0.91

37.7+35.0

4.8+1.6

6

11.7+9.5

5

5.00

F,

axh

23

7

12

0.52

81.4+69.0

6.1+3.0

11

10.0+8.7

8

4.88

hx a

20

8

13

0.65

43.6+33.3

2.9+3.5

4

0.4+0.5

3

4.67

F 2

(a x h) x (a x h )

11

7

9

0.82

26.8+34.3

5.6+2.7

2

0

—

—

(h x a)x (axh)

2

0

0

0

—

—

—

—

—

—

backcross

ax (axh)

7

5

5

0.71

8.5+6.6

3.3+1.0

3

35.0+16.1

3

1.00

hx(axh)

8

8

8

1.00

6.6+8.7

4.4+2.9

3

19.7+12.3

4

4.25

(axh) x a

1

0

0

0

—

—

—

—

—

—

(ax h)x h

3

1

3

1.00

60.7+49.8

8.1+8.0

0

—

—

—

b not tested

c time to first copulation (averaged only for pairs that mated)

known to occur within each of the disjunct distributions of
the two Scaptodrosophila species, and it is utilized by both.
Thus there is no field evidence of host plant specialization for
these Scaptodrosophila species. However, the hybridization
tests (Table 1) were done using H. diversifolius, which is
utilized by S. hibisci in nature, but which is not known to
occur within the distribution of S. aclinata. In both parental
and Fj crosses, S. aclinata females produced fewer progeny
than S. hibisci, indicating poorer adaptation of the former
to this Hibiscus species, to which it is not exposed in nature,
or possibly a lower intrinsic fecundity.

Hybridization studies

Adults of Scaptodrosophila hibisci and S. aclinata n.sp.
were reared from flowers of H. heterophyllus collected at
Bellingen, N.S.W. and flowers of H. menzeliae collected at
Nitmiluk National Park, Northern Territory. Some, where
females were collected as virgins, were used in single pair
matings in both parental and F x crosses (both reciprocals).
The remainder were added to population cages (one for each
species, and one for each reciprocal cross to produce Fj
progeny). For all pair matings, males were generally one
day older than females, and most females were collected
and used within 2 h of eclosion, using very light C0 2
anaesthetization. All flies for crosses were placed singly in
vials with about 7 ml 1.5% agar, and allowed 1 h to recover
from anaesthetization. The predetermined male was then
gently aspirated and added to its paired female, and pairs
observed for copulation for 3 h. Copulation latency and
copulation duration were recorded. All observations were
done between 09h00 and 14h00 at 25°C. At the end of the
observation period, each mated pair was placed in a 200 ml
bottle with moist sand in the base, and a small tube with
water holding a single H. diversifolius flower. The pairs

were transferred to a fresh flower each day for 10 days,
with the previous days flower transferred to a bottle with
sand. Four days later, 10 ml distilled water was added to
each of these bottles. Progeny emerging from these flowers
were collected daily, sexed and counted. From parental
matings, progeny were used in backcrosses or added to the
appropriate parental cage. Some of the ¥ l progeny, plus Fj
flies from the cage crosses, were used in F 2 and backcross
matings, with the remainder stored (sexes separate in agar
vials) for use on subsequent days.

Flies in population cages were maintained by adding
one or two fresh flowers to the cage each day, with the
previous days flowers transferred to a bottle with sand
for progeny collection.

Sufficient flowers were not available on some days to
set up all pairs that copulated. Further, some pairs were not
carried through for 10 days because of death or loss of one
or both of the pair. Thus the number of pairs tested for
progeny production is less than the number that copulated,
while the number of pairs recorded for day of first fertile
egg lay is greater than the number tested for progeny, except
where some pairs copulated, but produced no progeny.

Discussion. The results are summarized in Table 1. These
two species are partially interfertile, and clearly are closely
related. The proportion of pairs mating and average progeny
numbers are less for the Fj crosses than for parentals, while
no progeny were obtained from the F 2 crosses. Two of the
backcrosses appear exceptional, both in proportion of pairs
mating and in progeny numbers. However, this is possibly
a function of the much older males used in these crosses,
viz. average of 9-10 day old versus average of two day old
in all other crosses.

In all crosses, the pairs were kept together for 10 days,
so that further matings may have occurred during this period.

262 Records of the Australian Museum (2001) Vol. 53

Previous study of S. hibisci (Polak et al., 1998) has shown
that mature males prefer young virgins, as compared with
older virgin and non-virgin females, and that a mating plug
fills the entire uterus at copulation. For the S. hibisci parental
matings here, copulation latencies for < 2 h and 2 day old
females were 12.5 and 30.1 min (but not significantly
different). The sexual maturation and copulation dynamics
of S. aclinata seem to be different. Mean copulation latency
was about twice as long as for S. hibisci, while copulation
latencies for < 2 h, 1 and 4 day old females were 49.7, 39.2
and 10.9 min respectively (again not significantly different).
However, male age was highly correlated with female age,
and both copulation duration and progeny numbers
increased with parental age. These observations, together
with the later day of first fertile egg lay, suggest delayed
sexual maturity in this species, as compared with S. hibisci.
For the Fj cross (S. hibisci malexS. aclinata female), mean
copulation duration is shorter than for all other crosses.
However, six of the 13 pairs mated more than once in the 3
h observation period—five twice and one three times. In
all cases, the first copulation was short (< 1 min), and the
overall mean copulation duration, using last copulation for
multiple matings was 4.2+4.1 min, similar to the means of
other crosses.

Hibiscus meraukensis is known (records of the
Queensland Herbarium) from a number of localities in
northwest Queensland—the region between the known
distributions of these two Scaptodrosophila species (Fig.
13). Further field work in this region is needed to
determine if either species is present there, and whether
they ever occur sympatrically under natural conditions.
The form of orbital setation in hybrids is also in need of
further investigation so that any naturally occurring
hybrids may be identified as such.

Acknowledgments. We are indebted to Larry Wolf, Syracuse
University, Michal Polak, University of Cincinnati, and Jane
Bowles, University of Western Ontario, for camping company and
collaboration in collections in the Northern Territory, to Rick Hope,
Nitmiluk National Park, for collection of H. menzeliae flowers,
and to Lyn Craven, Centre for Plant Biodiversity Research,
Australian National Herbarium, for determination of Hibiscus
species. Field work in the Northern Territory was supported by
funds from the National Science Foundation (U.S.A.) International
Programs and Syracuse University, and laboratory work by an
Australian Research Council grant to JSFB. We thank Dr Akihiko
Shinohara, NSMT, for lending us material.

References

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Manuscript received 5 March 2001, revised 12 July 2001 and accepted

19 July 2001.

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