A remarkable new genus and six new species of fern-inhabiting plant bugs endemic to the Society Islands (Insecta: Heteroptera: Miridae: Mirinae: Filicicoris gen. nov.)
Author
Anna Namyatova
Author
Gerasimos Cassis
text
Insect Systematics & Evolution
2016
47
285
312
journal article
10.1163/1876312X-47032144
f37ece11-478d-4950-907f-a05410a22f4f
269854
Filicicoris
gen. nov.
Etymology
The genus is named for its affiliation with ferns, from the Latin, ‘
filix
’, and the suffix ‘
coris
’, from the Latin, meaning bug. The gender is masculine.
Type
species
Filicicoris meyeri
Namyatova and Cassis
, by original designation.
Diagnosis
Filicicoris
is recognized by the following characters: macropterous; body straight, not declivous, smooth, impunctate; body length
3–5 mm
; body covered with simple short pale and sometimes dark setae, head transverse (
Fig.
2
A, D); wings translucent; posterior margin of head carinate; eye not pedunculate or protruding laterally, contiguous with pronotum (
Fig.
2
A, D); in lateral view head shorter than high, distance from eye to inferior margin of head subequal to half of eye height (
Fig.
2
B); antennal segment I cylindrical (
Fig.
1
), not widened, longer than half of head width, usually subequal to or longer than head width; antennal segment II narrow and cylindrical; collar narrow; labium slightly surpassing posterior margin of metasternum; pronotum not constrict- ed behind calli (
Fig.
2
A, D); lateral margin of pronotum rounded, not angulate or carinate; calli shallow, scutellum flat, shorter than claval commissure; R+M reaching cuneus; cuneus twice as long as wide; femora slender, not modified; tarsal segment I shorter than segment II, segments II and III subequal in length (
Fig.
2
J); right paramere digitiform with small apical hook (
Figs.
3
B, E, I, L, O,
4
C); left paramere L-shaped (
Figs.
3
C, F, J, M, P,
4
D); vesica bearing
3–6
lobes, with small sclerotized spicules, but without long spicules (
Figs.
3
G,
4
A); female posterior wall with interramal lobes, without dorsal or lateral lobes or sigmoid process (
Fig.
5
B, D, F, H, J).
Table 1.
Morphological measurements of the species described in this study.
| Species Length |
Width |
| Body Cun-Clyp |
Pronotum |
AntSeg |
AntSeg |
Head Pronotum |
InterOcDi |
|
F. huahine
|
|
♂ (
N
= 5) Mean 3.28 2.38
|
0.58 |
0.66 |
1.75 |
0.72 1.01 |
0.36 |
| SD 0.07 0.04 |
0.03 |
0.03 |
0.11 |
0.00 0.04 |
0.01 |
| Range 0.17 0.11 |
0.07 |
0.06 |
0.24 |
0.01 0.10 |
0.02 |
| Min 3.22 2.33 |
0.53 |
0.63 |
1.67 |
0.72 0.95 |
0.35 |
| Max 3.39 2.44 |
0.60 |
0.69 |
1.91 |
0.73 1.05 |
0.37 |
|
♀ (
N
= 5) Mean 3.71 2.69
|
0.64 |
0.57 |
1.61 |
0.76 1.14 |
0.36 |
| SD 0.08 0.08 |
0.03 |
0.04 |
0.10 |
0.01 0.02 |
0.03 |
| Range 0.19 0.23 |
0.06 |
0.10 |
0.25 |
0.03 0.04 |
0.07 |
| Min 3.63 2.56 |
0.60 |
0.52 |
1.54 |
0.74 1.12 |
0.33 |
| Max 3.82 2.79 |
0.67 |
0.62 |
1.79 |
0.77 1.16 |
0.40 |
|
F. mayeri
|
|
♂ (
N
= 5) Mean 3.31 2.34
|
0.64 |
0.39 |
1.26 |
0.67 1.03 |
0.34 |
| SD 0.12 0.09 |
0.05 |
0.24 |
0.10 |
0.02 0.03 |
0.01 |
| Range 0.34 0.25 |
0.15 |
0.57 |
0.25 |
0.04 0.09 |
0.03 |
| Min 3.14 2.19 |
0.57 |
0.00 |
1.14 |
0.65 0.99 |
0.33 |
| Max 3.48 2.44 |
0.72 |
0.57 |
1.39 |
0.69 1.08 |
0.36 |
|
♀ (
N
= 5) Mean 3.58 2.50
|
0.69 |
0.53 |
1.18 |
0.71 1.10 |
0.36 |
| SD 0.09 0.05 |
0.01 |
0.04 |
0.08 |
0.01 0.02 |
0.02 |
| Range 0.24 0.14 |
0.01 |
0.11 |
0.20 |
0.03 0.05 |
0.05 |
| Min 3.48 2.44 |
0.68 |
0.46 |
1.10 |
0.69 1.08 |
0.35 |
| Max 3.72 2.58 |
0.69 |
0.57 |
1.30 |
0.72 1.13 |
0.41 |
|
F. rubramaculus
|
|
♂ (
N
= 5) Mean 3.90 2.79
|
0.55 |
0.73 |
1.88 |
0.77 1.05 |
0.35 |
| SD 0.09 0.07 |
0.02 |
0.06 |
0.11 |
0.01 0.04 |
0.02 |
| Range 0.25 0.18 |
0.06 |
0.17 |
0.29 |
0.03 0.10 |
0.04 |
| Min 3.77 2.72 |
0.52 |
0.64 |
1.73 |
0.76 1.02 |
0.33 |
| Max 4.02 2.90 |
0.57 |
0.81 |
2.02 |
0.79 1.11 |
0.37 |
|
♀ (
N
= 2)
N
= 1 4.27 3.07
|
0.30 |
0.78 |
1.86 |
0.81 1.17 |
0.37 |
|
N
= 2 4.24 3.21
|
0.29 |
0.77 |
1.94 |
0.81 1.27 |
0.36 |
|
F. tatarnici
|
|
♂ (
N
= 1)
N
= 1 4.18 2.90
|
0.70 |
0.85 |
2.15 |
0.79 1.17 |
0.37 |
|
F. viridis
|
|
♂ (
N
= 5) Mean 4.58 3.27
|
0.72 |
0.91 |
2.11 |
0.81 1.23 |
0.40 |
| SD 0.33 0.22 |
0.05 |
0.07 |
0.09 |
0.03 0.15 |
0.03 |
| Range 0.77 0.49 |
0.12 |
0.17 |
0.23 |
0.07 0.34 |
0.07 |
| Min 4.25 3.04 |
0.65 |
0.82 |
2.04 |
0.76 1.09 |
0.37 |
| Max 5.02 3.53 |
0.77 |
0.99 |
2.27 |
0.84 1.43 |
0.43 |
|
♀ (
N
= 5) Mean 4.73 3.34
|
0.76 |
0.86 |
1.99 |
0.86 1.27 |
0.42 |
| SD 0.34 0.24 |
0.04 |
0.06 |
0.10 |
0.09 0.11 |
0.02 |
| Range 0.76 0.57 |
0.11 |
0.15 |
0.25 |
0.22 0.29 |
0.05 |
| Min 4.26 2.98 |
0.68 |
0.80 |
1.82 |
0.79 1.14 |
0.40 |
| Max 5.02 3.55 |
0.79 |
0.95 |
2.07 |
1.01 1.43 |
0.45 |
1 2
Fig. 1.
Habitus of
Filicicoris huahine
(male and female),
F. meyeri
(male and female),
F. raiatea
(male and female),
F. tatarnici
(male),
F. rubramaculus
(male and female) and
F. viridis
(male and female). This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.
Description
Male.
Macropterous. Coloration (
Fig.
1
): Body usually pale, either yellow or green, sometimes with dark brown or red markings, with one species having dorsum mostly dark brown to black, wings translucent. Surface and vestiture: Mostly smooth and shiny, impunctate, pronotum weakly rugose; hemelytron translucent, covered with pale short simple setae, sometimes with dark setae on head and pronotum; tibiae with pale brown spine-likesetae.Structureandmeasurements:Head:Indorsalviewheadshort,transverse, with rounded frons (
Fig.
2
D) or slightly elongated and acute (
Fig.
2
A), vertex carinate; eye round, not pedunculate or protruding, contiguous with pronotum (
Fig.
2
A, D); in anterior view head triangular, only slightly elongated, clypeus length somewhat shorter than eye height, not delimited basally, antennal fossa placed slightly above inferior margin of eye, tubercles around antennal fossae slightly upraised, inferior margins of antennal fossa and eyes placed at the same level; eye higher than wide, its inner margin concave (
Fig.
2
C); in lateral view head shorter than high; distance between eye and inferior margin of head subequal to half of eye height, antennal fossa placed above suture between mandibular and maxillary plates, buccula nearly twice as short as labial segment I, distance between buccula and pronotum distinct, but shorter than buccula length (
Fig.
2
B). Labium (
Fig.
2
E): Slightly surpassing posterior margin of metasternum; segment I reaching forecoxa, approximately twice wider than other segments; segment II subequal to or slightly longer than segment I, segment III ca.
0.6–0.7
× as long as segment II, segment IV slightly longer than segment III (
Fig.
2
B). Antenna (
Fig.
1
): Subequal to body length; segment I cylindrical, longer than vertex width, subequal to or shorter than head width, not widened; segment II narrow and cylindrical,
2–3
× as long as segment I and slightly thinner than segment I, segment III slightly longer than half of segment II and as wide as segment II; segment IV half length of segment III and slightly thinner than segment III. Pronotum (
Fig.
2
A, D): Trapeziform, with lateral margins rounded, not angulate or carinate and posterior margin almost straight; collar distinctly delimited, narrow; calli shallow. Scutellum and mesoscutum: Scutellum flat, almost as long as wide, shorter than calval somissure; mesoscutum not exposed or slightly exposed. Scent gland evaporative area (
Fig.
2
F): Triangular, with anterior margin slightly shorter than posterior margin. Hemelytron: Costal margin rounded; clavus wide, subequal to eye width basally, indistinctly divided into inner and outer parts with upraising; claval commissure slightly longer than scutellum length; embolium wide, thinner basally; R+M reaching cuneus; cuneus triangular, twice as long as width at base, longer than scutellum, its inner margin straight; cuneal fracture directed anteriorly, slightly notched; membrane cell slightly longer than distance from apex of cell to apex of membrane (
Fig.
1
). Legs: Femora not modified, not widened; hind tarsal segment I short, tarsal segments II and III subequal in length, each of them ca.
1.5
× longer than segment I (
Fig.
2
J); pulvilli present, short (
Fig.
2
G), parempodia wide, longer than claw (
Fig.
2
G–I). Male genitalia: Genital capsule. Not rotated relatively to other segments (
Fig.
2
K), trapeziform, longer than width, its ventral wall longer than dorsal wall, acute posteriorly (
Fig.
4
E, F). Right paramere (
Figs.
3
B, E, I, L, O,
4
C): Digitiform, often folded dorsally (
Figs.
3
B, E, I, L), with apical process more or less hook-shaped, directed right or posteriorly. Left paramere (
Figs.
3
C, F, G, M, P,
4
D). Twice longer than right paramere, distinctly shorter than aedeagus, L-shaped, widened basally, with apical process narrow. Aedeagus (
Figs.
3
A, D, G, H, K, N,
4
A, B). Theca moderately sclerotized, transparent; endosoma subdivided into vesica and conjunctiva, vesica with
3–5
lobes (
Figs.
3
G,
4
A); vesica without long spicules, only with weekly sclerotized areas, sometimes lobes with small tubercles or small dense sclerotized spicules apically; conjunctiva entirely membranous; secondary gonopore large, oval; basal part of ductus seminis sclerotized, medial part short, coiled apical part of ductus seminis sclerotized, bowlshaped, secondary gonopore surrounded with loop bearing distinct sculpture.
Female.
Coloration (
Fig.
1
): As in male. Surface and vestiture: As in male. Structure and measurements: As in male, larger than males. Female genitalia (
Fig.
5
): Dorsal labiate plate (
Fig.
5
A, C, E, G, I) transverse, with anterior margin sclerotized (
Fig.
5
C, G, I) or membranous (
Fig.
5
A, E), sclerotized rings often indistinct, their diameter varying from very small (
Fig.
5
A, E) to relatively large (
Fig.
5
C, G, I); posterior wall of bursa copulatrix with pocketlike interramal lobes, distance between them varies from very small,
3
× as short as interramal lobe, to as long as interramal lobe (
Fig.
5
B, D, F, H, J).
Distribution
The genus is known only from Society Islands.
Plant associations
Plant records exist for three
Filicicoris
species. Adults and nymphs of
F. meyeri
are known from the fern
Nephroplepis
hirsulata
(
Lomariopsidaceae
).
Filicicoris viridis
specimens were collected from the ferns
Angiopteris evecta
and
Marattia
sp., (Mariattiaceae), and a few specimens were collected from the angiosperm
Metrosideros collina
(
Myrtaceae
), with the latter likely sitting records. All material of
F. rubromaculus
was collected from
Piper latifolia
(
Piperaceae
).
Very few groups of insects are known to live on ferns, including the
Heteroptera (
Konstantinov and Knyshov
2015
)
. Within the
Heteroptera
, there are a handful of clades belonging to the subfamily
Bryocorinae
(
Namyatova et al.
2016
) that are closely associated with ferns, in particular the tribes
Bryocorini
(
Konstantinov and Knyshov
2015
) and Felisacini (
Namyatova et al.
2016
). Additionally, a few species of the brocorine tribe
Dicyphini
are also associated with ferns (
Knight
1938
;
Wheeler et al.
1979
).
Here we provide the first record of the subfamily
Mirinae
with ferns.
Filicicoris
species are known from two phylogenetically distantly fern families, the
Lomariopsidaceae
and Mariattiaceae (
Christenhusz and Chase
2014
), which supports the contention of
Konstantinov and Knyshov (
2015
)
that mirids can be associated with distantly related fern clades.
Discussion
Based on the divergent, flattened parempodia and ring-like sclerotization of the secondary gonopore,
Filicicoris
belongs to the subfamily
Mirini
(
Schwartz
2008
;
Cassis and Schuh
2012
). According to the descriptions and keys provided in
Carvalho (
1955
)
,
Schwartz (
2008
)
and
Cassis and Schuh (
2012
)
,
Filicicoris
belongs to the nominotypical tribe
Mirini
. Although
Filicicoris
has a glassy appearance, akin to that found amongst most members of the tribe
Hyalopeplini
, it does not have the facies of members of this latter tribe. Various authors have reported on the inadequacy of glassy wings in defining hyalopeplines exclusively, with variation within the tribe, and presence in other mirine tribes (
Schwartz 1987
,
2008
;
Chérot
2001
). The
Hyalopeplini
comprises
17
genera, and all are readily separated from
Filicicoris
by the ecarinate vertex and/or punctate pronotum (
Carvalho and Gross
1979
).
There are
298
genera of
Mirini
known worldwide (Schuh
2002–2013
;
Carpintero and Chérot
2008
;
Cho and Kwon
2008
;
Gapon
2014
). We made an extensive survey of
mirini
diagnoses, and consider
Filicicoris
to be distinctive, based on the characters given in the diagnosis.
Filicicoris
is likely related to
Lygocoris
Reuter,
1875
and allied genera, as they also have a vertical head, carinate pronotum, impunctate or faintly punctate body and green coloration.
Filicicoris
is similar to
Lygocoris
as it also has a translucent hemelytron, but it differs in the posterior carina on the vertex being often indistinct medially, and the theca having an apical process ventrally and the vesica has distinct sclerites (
Yasunaga,
1991
a
). Other similar genera include
Neolygus
Knight,
1917
,
Lygocorias
Yasunaga,
1992
and
Ryukulygus
Yasunaga,
1996
.
Neolygus
differs by having the body covered with golden pubescence, the pronotum is irregularly punctate, the right parameres are L-shaped in most species and the left paramere has a well-developed sensory lobe (
Yasunaga
1991
b
).
Lygocorias
differs by the body being uniformly clothed with silky hairs, the right paramere is L-shaped, with an elongate hypophysis and the sensory lobe has a protuberance (
Yasunaga
1992
).
Ryukulygus
differs by having the hemelytra strongly declivous at the cuneal fracture, and the vesica possesses an elongate spicule (
Yasunaga
1996
).
Compared to genera from Australian biogeographic region,
Filicicoris
is similar to
Chinamiris
in external characters and male genitalia. However,
Chinamiris
Woodward,
1959
differs in the eye in lateral view being slightly shorter than the head height, the carinate lateral margins of the pronotum, and the hemelytron is opaque, and neither glassy nor translucent (
Eyles and Carvalho
1991
).
Filicicoris
shares some similarities with
Tinginotum
Kirkaldy,
1902
in structure, but the latter genus has an opaque hemelytron, elongate setae and the vesica has distinct spicules (
Eyles
2000
).
Diomocoris
Eyles,
1999
shares some characteristics with
Filicicoris
in having green coloration and translucent hemelytron, but differs in the body being clothed with the dense silvery setae, the antennal segment I subequals to the vertex width, the right paramere is L-shaped, and the vesica has a large spicule.
Filicicoris
may also be related to the New Guinean genus
Prolygus
Carvalho,
1987
, as the latter has similar genitalia, but differs in the finely puncutate body and the cuneus length being subequal to or slightly longer than the cuneus width at base (
Carvalho,
1987
).
Amongst genera inhabiting the Western Hemisphere, we found only three taxa that are similar to
Filicicoris
, based on their original descriptions. Based on the works of
Carvalho (
1984
)
and
Carvalho and Costa (
1992
a)
,
Ganocapsinus
Carvalho,
1984
and
Ganocapsisca
Carvalho and Costa,
1992
are similar to
Filicicoris
in many diagnostic characters and the parameres of
Ganocapsisca
bear resemblance to those of
Filicicoris
.
Ganocapsinus
differs in having a protruding scutellum, the cuneal fracture is notched, the left paramere has an enlarged sensory lobe and the vesica has a large spicule.
Ganocapsisca
differs in having large eyes, reaching the gula in lateral view, the antenna are inserted near the upper third of the eye, and the apical part of right paramere is straight.
Filicicoris
is also similar to
Xavantinisca
Carvalho and Costa,
1992
in the structure and the shape of parameres, but
Xavantinisca
differs in the presence of small punctures on the pronotum and hemelytron, the left paramere has dentate outgrowth, and the aedeagus has a spicule (
Carvalho and Costa
1992
b
).
Key to species
1
. Mostly yellow to green, uniform in coloration..................................................
2
- Mostly glassy to green, with red or black areas on dorsum................................
5
2
. Body pale green or yellow ................................................................................
3
- Body dark green....................................................................
F. tatarnici
sp. nov.
3
. Body length
4.3
–5.0 in males and females; dorsal labiate plate with sclerotized anterior margin (
Fig.
5
I); posterior wall with distance between interramal lobes subequal to interramal lobe width (
Fig.
5
J); antennal segment I usually longer than head width........................................................................
F. viridis
sp. nov.
- Body length in males
3.2–3.4
and in females
3.6–3.9
; dorsal labiate plate with membranous anterior margin (
Figs.
5
A, E); posterior wall with distance between interramal lobes ca.
0.3
× as long as interramal lobe width (
Figs.
5
B, F); antennal segment I usually shorter than head width.........................................................
4
4
. Vesica with six membranous lobes, one of them with small dense pointed spicules apically and another one with small tubercles apically (
Fig.
3
A)........................... ................................................................................................
F. huahine
sp. nov.
- Vesica with four distinct lobes lacking small sclerotized spicules, one of the lobes with tubercles apically (
Fig.
3
D)................................................
F. raiatea
sp. nov.
5
. Body with head, pronotum, scutellum, clavus and marking on corium black; vesica with three lobes (
Fig.
3
H) .....................................................
F. meyeri
sp. nov.
- Body with head, anterior part of pronotum and marking covering posterior part of clavus and inner part of corium red; vesica with four lobes (
Fig.
3
K)............... ........................................................................................
F. rubramaculus
sp. nov.