|
Rusicada Walker Gen. rev.
Type
species: nigritarsis
Walker.
Synonym:
Gonopteronia
Bethune-Baker
(type species albopunctata Bethune-Baker, New Guinea to Solomons) syn.
n.
This
account contains much of the material in an unpublished review of the genus by
the author that was referred to by Galsworthy (1997) and Holloway & Nielsen
(1999). Though not fully illustrated, the keys presented should allow all known
Indo-Australian species to be identified.
In
that review, Rusicada
was
retained as a subgenus of Anomis, but is here given full generic status for the reasons presented
on p. 223. The species are relatively large and robust with variegated brick-red
to brown or greyish and pale orange-red hindwings.
The
male genitalia (see also p. 222) have deep valves that extend ventrally to a
shallow saccus that has often a concave ventral margin. The valves have deeply
based, slightly double coremata, and the Indo-Australian species usually have a
slight fold or lobe at three-quarters from the valve base. The juxta is strongly
developed, usually bifid distally (though this is reduced in one lineage), a
feature shared with Cosmophila Boisduval. There are setose projections from where
the anellus fuses with the aedeagus, and the aedeagus vesica has two reversed
cornuti when everted. The male abdomen shares with Cosmophila
and
other members of the Anomis group a corema on the eighth sternite, but this is
much more developed in Rusicada.
The
female genitalia share with Cosmophila presence of two setose lobes arising from the
posterior margin of the lamella postvaginalis. The corpus bursae is scobinate
and corrugated, but lacks a signum.
The
genus contains 17 Oriental and Australasian species of which six occur in as
described later. The other 11 are as follows:
|
Rusicada fulvida Guenée comb.
n. =
inducens
Walker
|
Oriental tropics and subtropics to Japan, Sumatra and Java
|
|
Rusicada mafalui Bethune-Baker comb.
n.
|
Papua
New Guinea
|
|
Rusicada albopunctata Bethune-Baker comb.
n.
|
New
Guinea, Bismarcks, Solomons
|
|
Rusicada schistosema Hampson comb.
n.
|
Seram,
Queensland, New Guinea, Solomons, Vanuatu
|
|
Rusicada privata Walker
comb.
n. =
commoda
Butler
|
China,
Korea, Japan, Vietnam (Zilli, 1999), Thailand (VK) – introduced to
eastern U.S.A.
|
|
Rusicada metaxantha Walker
comb.
n.
|
N.E. Himalaya, Vietnam (Zilli,1999)
|
|
Rusicada lineosa Walker
comb.
n.
|
India
|
|
Rusicada fructusterebrans Bänziger comb.
n.
|
N. Thailand
|
|
Rusicada fulminans Bethune-Baker comb.
n.
|
New
Guinea
|
|
Rusicada vulpina Butler
comb.
n. =
nagalia
Mabille
& Vuillot
|
Fiji,
?Samoa
|
|
Rusicada sp. n. (slide 11946)
|
N. Thailand
|
Keys
to the Indo-Australian species are presented below, based separately on external
appearance, male genitalia and female genitalia. The similarity of the external
appearance of many of the species means that reliable diagnosis is best obtained
from genitalia structures. However, distinction by female genitalia characters
is least reliable for closely related species pairs such as revocans
/
fulvida,
combinans
/
schistosema
and
the group prima
/
leucolopha
/
fulminans
/
vulpina,
and it is recommended that external appearance and geography be used to confirm
any diagnosis so made.
External
appearance
| 1 |
Forewing ground colour orange |
2 |
|
— |
Forewing ground colour mainly brown or vinous, though perhaps with orange or
reddish patches or tinge |
11 |
|
|
|
| 2 |
Posterior lobe of forewing reniform smaller than the anterior |
3 |
|
— |
Posterior lobe of reniform equal to or larger than the anterior |
5 |
|
|
|
| 3
|
Average line of zigzag anterior part of forewing postmedial oblique to costa
|
nigritarsis
|
| — |
Average line of anterior part of postmedial more or less at right angles to the
costa |
4 |
|
|
|
| 4 |
Posterior lobe of forewing reniform markedly darker and forewing ground dull
yellowish orange (Australian specimens of revocans could key out here)
|
fulvida
|
| — |
Posterior lobe of forewing reniform only slightly darker and forewing ground
reddish orange |
revocans |
|
|
|
|
|
|
| 5 |
Forewing reniform with small lobes separated by a marked constriction; forewing
margin rounded; pale orbicular large, often almost 1mm in diameter, cream-coloured |
vulpina
|
| — |
Forewing reniform with moderate lobes separated by only a slight constriction;
forewing margin angular; orbicular a minute white dot |
6 |
|
|
|
| 6 |
Forewing reniform with posterior lobe larger, filled with paler grey or dark
brown |
7 |
|
— |
Forewing reniform with lobes roughly equal, filled with ground colour |
8 |
|
|
|
| 7
|
Forewing antemedial more or less evenly curved; hindwings reddish; Papuan
Subregion
|
schistosema
|
| — |
Forewing antemedial irregular; hindwing brown; Oriental |
combinans |
|
|
|
| 8 |
Hindwing margin produced slightly centrally |
prima
|
| — |
Hindwing margin more or less evenly rounded |
9 |
|
|
|
| 9 |
Posterior half of forewing postmedial more or less straight, evenly concave
distad or only slightly wavy; E. China, Japan, Vietnam |
privata
|
| — |
Posterior half of postmedial distinctly wavy |
10 |
|
|
|
| 10 |
Length of forewing costa greater than 20mm |
leucolopha
|
| — |
Length of forewing costa less than 20mm |
metaxantha |
|
|
|
| 11 |
Forewing margin excavate between apex and central angle; orbicular white dot
prominent |
albopunctata
|
| — |
Forewing margin straight or only slightly excavate between apex and central
angle; orbicular white dot minute |
12 |
|
|
|
| 12
|
Posterior part of postmedial evenly curved, pale, colinear with basal edge of
reniform
|
lineosa
|
| — |
Postmedial not as above |
13 |
|
|
|
| 13 |
Forewing marginal angle rounded; reniform with black dots distally; New Guinea
(if Fiji, then vulpina) |
mafalui
|
| — |
Forewing marginal angle square, reniform obscure |
14 |
|
|
|
| 14 |
Line of anterior half of forewing postmedial meets costa obliquely |
15 |
|
— |
Line of anterior half of postmedial meets costa more or less at right angles |
16 |
|
|
|
| 15 |
Reddish brown; orbicular white dot present |
nigritarsis
|
| — |
Dark brown; orbicular not white |
fructusterebrans |
|
|
|
| 16 |
Forewing markings obscure; rich reddish deep brown |
fulminans
|
| — |
Forewing markings obvious; vinous greyish brown or dark brown, sometimes with
orange patch centrally by posterior half of postmedial |
17 |
|
|
|
| 17
|
Forewings dark brown; line of posterior part of postmedial not contiguous with
basal edge of reniform but well basal to it
|
sp.
11946
|
|
—
|
Forewings vinous greyish brown, sometimes with orange suffusion; line of
posterior part of postmedial more or less contiguous with basal edge of reniform
|
albitibia |
Male
genitalia (aedeagus
vesica must be everted).
| 1 |
Setose projections from anellar attachment to aedeagus asymmetrical, usually
only one large one; valve coremata with setal bases pale; juxta robust, strongly
bifid |
2 |
|
— |
Setose projections on aedeagus two, more or less symmetrical; valve coremata
with setal bases dark-centred; juxta weakly bifid or entire, planar. |
6 |
|
|
3 |
| 2 |
Valve with subapical lobes or projections |
4 |
|
— |
Valve without subapical
ornamentation |
|
|
|
|
| 3 |
Valve heavily sclerotised, ventral angle acute, curved; two subapical
projections on valve: a large subdorsal rounded one and a smaller elongate
angled subventral one; uncus square apically, constricted subapically; eighth
sternal corema double |
albopunctata
|
| — |
Valve only lightly sclerotised, ventral apical angle blunt central subapical
projection to valve a transverse flange; uncus tapering apically; eighth sternal
corema single |
mafalui |
|
|
|
| 4 |
Juxta bifurcations extend almost to apex of valve, apically curved; basal
projections from vesica rugose, directed basad |
nigritarsis
|
| — |
Juxta bifurcations straight, terminating well short of apex of valve; one
projection from base of vesica a robust, distally directed spine |
5 |
|
|
|
| 5 |
Anellus projection long, slender, only heavily setose at apex |
revocans
|
| — |
Anellus projection broad, moderate, evenly and extensively setose |
fulvida |
|
|
|
| 6 |
Juxta with lobes lateral to a longer, central, notched process |
sp. n. 11946 |
|
— |
Juxta otherwise |
7 |
|
|
|
| 7 |
Juxta distinctly bifid |
8 |
|
— |
Juxta entire or slightly lobed apically |
9 |
|
|
|
| 8
|
Uncus apically excavate; aedeagus with two smaller anellus projections in
addition to two normal ones and two prominent long spines arising from within
the anellus structure
|
schistosema
|
| — |
Uncus tapered apically; aedeagus with only two anellus projections, with two or
three slender spines arising from within the anellar structure |
combinans |
|
|
|
| 9 |
Juxta apically squarish, broadest at apex |
10 |
|
— |
Juxta apically rounded, narrower at apex |
12 |
|
|
|
| 10 |
Cornutus in aedeagus vesica a lobe bearing several spines |
privata
|
| — |
Cornutus simple, unspined |
11 |
|
|
|
| 11
|
Long spines associated with anellar structure as in 7; largest vesica cornutus
acute
|
metaxantha
|
| —
|
No long spines in anellar structure; largest vesica cornutus apically bulbous
|
prima |
|
|
|
| 12 |
Valve ornamentation a sclerotised band extending to the ventral margin
subapically, where it is coarsely scobinate |
lineosa
|
| — |
Valve ornamentation not marginal if present |
13 |
|
|
|
| 13 |
Valve with central subapical projection oblique |
14 |
|
— |
Valve with central subapical projection vestigial or transverse |
16 |
|
|
|
| 14
|
Central projection of valve moderately oblique; uncus long, tapering
|
fructusterebrans
|
| — |
Central projection more longitudinal than transverse; uncus short, somewhat
swollen subapically |
15 |
|
|
|
| 15 |
Aedeagus vesica with one cornutus long, curved, almost reaching exterior angle
of everted vesica |
leucolopha
|
| — |
Aedeagus vesica with both cornuti short, blunt, directed more towards interior
angle of everted vesica |
fulminans |
|
|
|
| 16
|
Valve with central subapical projection reduced, merely a more sclerotised area
|
vulpina
|
| — |
Central subapical projection of valve transverse |
albitibia
|
Female
genitalia
| 1 |
Two posterior, finely setose lobes of lamella postvaginalis as long as, or
longer than broad, separated by a deep, usually
narrow cleft; sclerotisation at base of ductus bursae moderate to weak, separate
from lamella vaginalis |
2 |
|
— |
Lobes of lamella postvaginalis broader than long at base, separated by only
a shallow, angular cleft; sclerotisation at base of ductus bursae extensive,
continuous with that of lamella vaginalis |
12 |
|
|
|
| 2 |
Cleft of posterior lobes terminating in a posteriorly directed pocket |
3 |
|
— |
Cleft of posterior lobe equal to exterior margin, terminating simply |
11 |
|
|
|
| 3 |
Ductus bursae with sclerotisation centrally |
4 |
|
— |
Ductus bursae with sclerotisation only basally or not at all |
5 |
|
|
|
| 4 |
Ductus bursae with central sclerotisation associated with an angle |
privata
|
| — |
Ductus bursae not angled |
metaxantha |
|
|
|
| 5 |
Base of ductus distinctly swollen, connected to pocket of lobes (2) by a broad
band of sclerotisation |
lineosa
|
| — |
Base of ductus unswollen or only slightly so, connected to pocket of lobes by
only a weak band of sclerotisation |
6 |
|
|
|
| 6 |
Pocket of lobes much larger (deeper) than lobes |
albitibia |
|
— |
Pocket of lobes smaller than lobes |
7 |
|
|
|
| 7 |
Sclerotisation present in ductus bursae distal to basal constriction |
8 |
|
— |
No sclerotisation so present |
10 |
|
|
|
| 8 |
Pocket and lobes flanked by a pair of semicircular flanges |
sp. n. 11946 |
|
— |
Pocket and lobes not flanked by flanges |
9 |
|
|
|
| 9 |
Lobe pocket much smaller than lobes; anterior margin square, posterior margin
shallowly cleft |
prima
|
| — |
Lobe pocket almost as large as lobes; anterior margin hastate, posterior
margin rounded, deeply excavate |
leucolopha |
|
|
|
| 10
|
Depth of pocket at sides less than one third length of interior cleft, its lip
only slightly concave
|
vulpina
|
| —
|
Depth of pocket at sides about half length of interior cleft, its lip deeply
concave
|
fulminans |
|
|
|
| 11 |
Lobes mostly parallel-sided, set close together |
combinans
|
| — |
Lobes tapering, set slightly apart |
schistosema |
|
|
|
| 12 |
Lamella postvaginalis with posteriorly directed sclerotised angle between the
lobes; diverticulum on right of base of ductus bursae |
13 |
|
— |
Lamella postvaginalis without such an angle; no diverticulum on right of
base of ductus bursae (but possibly on left) |
14 |
|
|
|
| 13 |
Angle between lobes obtuse, somewhat rounded |
fulvida
|
| — |
Angle between lobes acute |
revocans |
|
|
|
| 14
|
Sclerotised base of ductus straight, simple
|
nigritarsis
|
| — |
Sclerotised base of ductus expanded, convolute |
15 |
|
|
|
| 15
|
Distal diverticulum from left of basal swelling on ductus; lamella postvaginalis
lobes only weakly cleft (to less than half overall depth)
|
mafalui
|
| — |
No distal diverticulum on ductus swelling; lamella postvaginalis lobes cleft to
more than half overall depth |
albopunctata |
Fig
2. Cladogram
of relationships of species of Rusicada, with summaries of distribution.
Numbers
refer to presumed synapomorphies of each clade, listed below.
1.
Setose projections from where anellus fuses with aedeagus. Vesica, when everted,
with two reversed cornuti (reduced to one in many species). Sternite eight
corema in male strong (present in Cosmophila and some allied species, but weak).
2.
Single valve corema elongate along axis of valve. Setose projections from
anellus well developed. Male forewing costal margin with one or two angles.
Valve with slight fold or lobe centrally at three-quarters from base (lost,
displaced or modified in several species).
3.
Reversed cornuti reduced to one in vesica (also in privata though).
Anellus projections asymmetric. Juxta stout, robust, broadly bifid (only
slightly so in pair defined by 8, entire (bifurcations fused) in other members
of sister group). Base of ductus bursae heavily sclerotised and sclerotisation
continuous with lamella postvaginalis.
4.
Valve central process lost (seen also in privata, metaxantha and
to some extent in vulpina). Vesica of aedeagus with additional basal
projections.
5.
One basal projection on aedeagus vesica a distally directed spine. Angular
projection from lamella postvaginalis between marginal lobes. Diverticulum
basally on ductus bursae on right.
6.
Dull brown coloration. Black dots distally on forewing reniform, one on anterior
lobe, two on posterior lobe. Heavy sclerotisation and convolution of base of
ductus bursae.
7.
Valve apex bilobed. Setal bases on valve corema dark-centred.
8.
Juxta bilobed rather than bifid. Gnathus apically divided, the bifurcations
splayed. Base of ductus bursae spherically swollen. Reniform of forewing with
posterior lobe larger, filled with pale or dark grey (also seen in the African
pair; 10).
9.
Juxta entire (some indication of fusion of bilobed structure in some species).
Interior margins of lobes of lamella postvaginalis come together within a
posteriorly directed pocket.
10.
Juxta narrow, bifurcations close together. Eighth sternal corema strongly
eversible.
The
larvae are slender, cylindrical semi-loopers with the first of prolegs reduced.
The primary setae are small, giving the body a glabrous appearance; the
chaetotaxy was described by Gardner (1941). The larvae are solitary defoliators
of a variety of plant genera, mainly in the family Malvaceae (Abutilon,
Cissampelos,
Gossypium,
Hibiscus,
Kydia,
Sida,
Thespesia,
and Urena)
and the related Tiliaceae (Grewia) and Sterculiaceae (Pterospermum,
Sterculia,
Waltheria),
all Malvales.
The
adults are primary piercers of soft-skinned fruit and secondary piercers of
fruit with tougher skin (Bänziger, 1982); the development of barbs on the
proboscis is generally more pronounced than in other genera of the Anomis
complex
(Bänziger, 1986).
Two
Afrotropical species, leucosema Hampson and melanosema Berio,
are also referable to Rusicada and probably form a sister-group to the
Indo-Australian species as suggested by a cladogram presented by Holloway &
Nielsen (1999). This is reproduced in Fig 2, with synapomorphies listed in the
legend.
Two
major clades are recognised. The nigritarsis clade is fully resolved and includes three widespread
species (two recorded from Borneo) characteristic of open habitats as
sister-group to a pair of species from New Guinea and neighbouring islands.
The
second, much larger, combinans clade includes an unresolved group of ten species.
Apparently derived traits within the group either also occur in combinans
(long
spines arising from within the anellar structure; valve projection transverse,
almost a pocket) or otherwise lack congruence (obliqueness or loss of the valve
projection; elongation of one vesica cornutus; square apex to juxta).
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