Taxonomic review of the Tribe Melaenini (Coleoptera: Carabidae), with observations on morphological, ecological and chorological evolution
Author
Ball, George E.
Author
Shpeley, Danny
text
Zootaxa
2005
2005-12-19
1099
1
1
120
https://biotaxa.org/Zootaxa/article/view/zootaxa.1099.1.1
journal article
10.11646/zootaxa.1099.1.1
11755334
5051674
CA0A0ED3-D49D-4198-A409-407EF86A164E
Cymbionotum
Baudi di Selve 1864
Synonymy.
See following treatment of subgenus
Cymbionotum
.
Recognition.
In general form, adults of
Cymbionotum
(
Fig. 1B
) resemble small adults of
Siagona
Latreille. However
, they exhibit the features of the tribe
Melaenini
(see above), and are recognized by the following combination of character states: overall size small (length ca.
3–7 mm
); body form depressed, broad, pedunculate; integument generally setose; color pale (though elytra of some species bicolored); vertex of head with distinct postocular transverse sulcus; head with two pairs of supraorbital setae, mandibular scrobes each with several setae; pronotum with two pairs of lateral setae; elytral surface virtually smooth, striae absent or very shallowly impressed.
Description
. Adult, with character states of tribe
Melaenini
, and recognition features of
Cymbionotum
, restricted and/or amplified as follows. Color of body black to rufotestaceous, elytra concolorous or bicolored. Body form depressed. Antennae and mouthparts rufotestaceous to testaceous, concolorous. Legs rufotestaceous.
FIGURE 11.
Heads and antennae of
Cymbionotum
species
illustrated with SEM photographs: A–C, head, dorsal aspect of: A,
C. semelederi,
Fabricius
; B,
C. microphthalmum
Chaudoir
; C,
C. helferi
Chaudoir. D
–G, left antenna of: D,
C. semirubricum
Reitter
(antennomeres 1–3, anterior aspect); E,
C. striatum
Reitter
(antennomeres 1–3 and base of 4); F and G,
C. semirubricum
and
C. striatum
,
respectively (antennomeres 6 and 7, posterior aspect). Legend:
A2, A3
,
A6
and
A7
, antennomeres 2, 3, 6 and 7;
ct
, clypeal tooth;
poi
, postocular impression. Scale bars: A–C = 1.0 mm; D–G = 100µm.
Microsculpture. Dorsal surface generally with isodiametric mesh pattern, microlines shallow; or microlines absent, except in postocular transverse impression of dorsal surface of head (
Fig. 11C
,
poi
) with isodiametric to slightly transverse mesh pattern, or smooth, without microlines; or microlines absent, surface without evident mesh pattern. Scutellum with mesh pattern isodiametric to slightly transverse, microlines confined to anterior area, or generally distributed; or completely smooth, without microlines. Middle tibia (
Fig. 13D
,
ifs
)with mesh pattern on inner surface elongate,sculpticells flat,microlines not fimbriate. Luster. Body surface dull or shiny.
Punctation and vestiture (cf.
Figs. 2A–F
,
3A–F
,
11A–C
). Body surface generally punctate and setose, elytral punctures relatively small (
Figs. 2B–E
) to large (
Fig. 2F
), and sparse (
Fig. 3A
), intermediate (
Fig. 3B
), dense (
Figs. 3C–E
), and very dense (
Fig. 2A
,
3F
); elytral setae short (
Figs. 2B–C
), intermediate (
Figs. 2D–E
), or long (
Fig. 2F
). Head posteriad eyes each side with row of anteriorly directed setae (
Fig. 11B
). Proepipleuron with row of upturned setae (
Figs. 11A–C
). All antennomeres setose; all setae relatively long and not especially dense (
Figs. 11D–G
).
Fixed setae (body sclerites, mouthparts, elytra). Head capsule with two pairs of supraorbital setae. Mouthparts: mandible with several setae in scrobe (
Figs. 12A–D
,
pss, sss
); glossal sclerite distally quadrisetose (
Fig. 13A
,
gs
). Pronotum with two pairs of lateral setae. Elytra dorsally without parascutellar or discal setae. Abdominal sterna IV– VII each with pair of setae near posterior margin.
Head (
Figs. 11A–C
). Wide, dorsal surface slightly convex, with distinct transverse impression posteriad eyes (
Fig. 11C
,
poi
); clypeus with anterior margin evenly curved, almost straight (
Figs. 11A–B
), or more markedly concave, with dentiform projection (
Fig. 11C
,
ct
) in both sexes, or only in males; frontoclypeal suture not evident, frontal impressions hardly indicated, or broad and shallow; frontovertex each side laterad eye with slightly elevated longitudinal ridge extended posteriorly to postocular transverse impression; supraorbital ridge sharp, each side sinuously extended anteriorly to mandibular fossa, here joined to apex of frontovertical ridge, and posteriolaterally extended to postocular transverse impression.
Antennae. Nearly moniliform (
Fig. 1B
), shorter, extended to about basal third of elytra. Antennomere 2 pyriform (
Fig. 11D
) or globose (
Fig. 11E
); antennomeres variously proportioned: 2–4 and 11 slightly longer than wide, antennomeres 5–10 slightly longer than wide, slightly (
Fig. 11F
) or markedly (
Fig. 11G
) asymmetric.
Mouthparts (
Figs. 12A–L
,
13A–C
). Labrum dorsally (
Fig. 12I
) with anterior margin broadly and shallowly concave, or more deeply and angularly so (
Figs. 11A–C
). Epipharynx (
Fig. 12J
): pedium (
ped
) triangular, approximately isosceles, with broad base along distal margin; parapedial projection (
pp
) obtuse; extra parapedial setae (
eps
) mediad parapedial ridge (
pr
); coeloconic sensilla (
cs
) relatively few.
Left mandible, dorsal surface (
Fig. 12B
) with anterior retinacular tooth not evident, from dorsal aspect. Maxilla as in
Figs. 12K–L
. Labium (
Figs. 13A–C
): paraglossae (
pg
), surface with extensive covering of setalike microtrichia (
Fig. 13B
,
mtr
); in dorsal aspect (
Fig. 13C
), distal portion of glossal sclerite (
gsap
) small, relative to medial portion (
gsmp
).
FIGURE 12.
Structural features of
Cymbionotum fernandezi
,
new species
, illustrated with SEM photographs. A–H, mandibles: A, D, lateral aspect, left and right mandible, respectively; B, C, dorsal aspect, left and right mandible, respectively; E, H, occlusal aspect, left and right mandible, respectively; and F, G, ventral aspect, left and right mandible, respectively. I, J, labrum, dorsal and ventral aspect, respectively. K,L, left maxilla, ventral and dorsal aspects, respectively. Legend:
aog
, anterior occlusal groove;
art
, anterior retinacular tooth;
as,
apical seta;
b;
basal portion of mandible;
bb
; basal brush;
bemr,
basal extension of molar ridge;
c
, cardo;
cr,
crepis;
cs
, coeloconic sensillum;
dgs
, dorsal groove of scrobe;
dls
, dorsal lobe of stipes;
eps,
extra parapedial seta;
g1,
galeomere 1;
g2,
galeomere 2;
it,
incisor tooth;
lc
, lacinia;
lct
, lacinial tooth;
lps,
lateral parapedial seta;
mp2,
maxillary palpomere 2;
mp3,
maxillary palpomere 3;
mp4,
maxillary palpomere 4;
mps,
medial parapedial seta;
mr,
molar ridge;
mt,
molar tooth;
mtr,
microtrichia;
omr,
occlusal part of molar ridge;
ped,
pedium;
pf
, palpifer;
pfs,
palpiferal seta;
pmr
, premolar ridge;
pmt
, premolar tooth;
pog,
posterior occlusal groove;
pp,
parapedial projection;
pr,
parapedial ridge;
prt
, posterior retinacular tooth;
pss,
primary scrobal seta;
rr
, retinacular ridge;
s,
scrobe;
sss,
secondary scrobal seta;
svg
, secondary ventral groove;
t
, terebra;
tr
, terebral ridge;
tt,
terebral tooth;
vd,
ventral depression;
vg
, ventral groove;
vls
, ventral lobe of stipes. Scale bars = 100 µm.
FIGURE 13.
Structural features of
Cymbionotum fernandezi
,
new species
, illustrated with SEM photographs. A, labium, ventral aspect; B. paraglossa, dorsal aspect; C, labium, dorsal aspect; D, middle tibia, apical portion, anterior aspect; E, middle tarsomere 5, apical portion, and claws, ventral aspect; F–J, ovipositor, left gonocoxa––F, gonocoxa 1 and 2, lateral aspect; G–I, gonocoxite 1, apical portion and gonocoxite 2, lateral, dorsal and ventral aspects, respectively; J, gonocoxite 2, apical portion, oblique aspect. Legend:
cs,
coeloconic sensillum;
ats
, anterior tibial spur;
avl
, apical ventral lobe of t5;
css
, clamshell sensillum;
dts
, dorsal trichoid seta;
e,
epilobe of mentum;
es
, ensiform seta;
gc1,
gonocoxite 1;
gc2,
gonocoxite 2;
gs
, glossal apical seta;
gsap
, glossal sclerite, apical portion;
gsbp,
glossal sclerite, basal portion;
gsc
, glossal sclerite;
gsmp,
glossal sclerite, medial portion;
ifs,
inner face sculpticell;
ll
, lateral lobe of mentum;
lp1
, labial palpomere 1;
lp2
, labial palpomere 2;
lp3
, labial palpomere 3;
m
, mentum;
ms
, paramedial seta of mentum;
msms
, mentalsubmental suture;
mt
, mental tooth;
mtb
, middle tibial brush;
mtr
, microtrichia;
ns
, nematiform seta;
pg
, paraglossa;
pge,
palpiger;
psms
, primary submental seta;
sm,
submentum;
ssf
, subapical sensory furrow;
ssms
, secondary submental seta;
tc
, tarsal claw;
ts
, trichoid seta;
up
, unguitractor plate;
vls;
ventrolateral seta. Scale bars: A, C– F, and I = 100 µm; B and J = 10 µm.
Prothorax. Pronotum (
Figs. 1B
,
14
) cuplike in outline from dorsal perspective, lateral margins not beaded, more or less sinuate posteriorly; posteriolateral angles rectangular to acute, prominently projected; lateral grooves narrow; posterior margin beaded or not; proepipleuron evident (at least posteriorly) in dorsal aspect; disc almost flat, lateral declivity each side slightly sloped.
FIGURE 14.
Pronotum, dorsal aspect, of
Cymbionotum
species
, illustrating subgeneric differences with outline drawings. A,
C. (Procoscinia) fernandezi
,
new species
. B,
C. (Cymbionotum) semelederi
Chaudoir. Legend
:
pep
, proepipleuron. Scale bars = 1.0 mm.
Elytra (
Figs. 1B
,
2A
). Dorsal surface plane for most of length, apical declivity gradually sloped; basal ridge narrow, not extended to edge of scutellum, anteriorly smooth, not angulate; stria 1 shallow, striae 2–8 not or hardly evident; intervals flat.
Hind wings (
Fig. 15A
). Macropterous. Wedge cell absent. Veins
RP 3 + 4
and
AA 3 + 4
absent. Vein
MP 4
less than half length of
MP 3.
Legs. Middle tibia with distinct distal brush of setae (
Fig. 13D
,
mtb
). Unguitractor plate (
Fig. 13E
,
up
) narrow.
Male genitalia (
Figs. 15B–D, 15F
, 17D–E, 17I–K, 19C–D, 19G–H, 19K, 19O, 22D). Endophallus with sclerite x much shorter than median lobe, or absent; microtrichial field various, from very small to large; parameres (
Figs. 15C–D
) distally glabrous.
Ovipositor (
Figs. 13F–J
). Ramus about 0.5–0.8 length of gonocoxite 1. Laterotergite IX (
Fig. 15E
,
lt
) without anteriorly directed apophysis. Gonocoxa dimerous, gonocoxites 1 and 2 subequal (
Fig. 13F
,
gc1, 2
), gonocoxite 2 dorsally with pair of trichoid setae in lateral aspect (
Fig. 13G
,
dts)
, dorsal margins curved; narrow in outline, distal margin narrowly rounded (
Figs. 13H–I
), ventral surface smooth, each side with row of coeloconic sensilla (
Fig. 13G
,
cs
).
Female internal genitalia (
Fig. 15E
). Spermathecal duct much longer than spermathecal reservoir (
sp
), latter reniform.
FIGURE 15.
Structural features of
Cymbionotum fernandezi
,
new species
(A–E) and
C. negrei
Perrault
(F) illustrated with line drawings. A, left hind wing, dorsal aspect. B, male genitalia: phallus, left lateral aspect, with endophallus partially everted; C–D left and right paramere; respectively, ventral aspect. E, female genitalia, ventral aspect. F, male genitalia, left lateral aspect. Legend: wing veins and cells
AA1+ 2,
anal vein 1 + 2;
MP3
and
MP4,
posterior median veins 3 and 4, respectively;
o
, oblongum cell;
RA4
, anterior radial vein 4;
RP2
, posterior radial vein. Male genitalia:
eph
, endophallus;
ph
, phallus;
x,
sclerite x of endophallus. Female genitalia:
bc
, bursa copulatrix;
co
, common oviduct;
di
, spermathecal diverticulum;
gc 1
and
2,
gonocoxite 1 and 2, respectively;
lt
, laterotergite of abdominal segment IX;
sp,
spermatheca;
spd
, spermathecal duct. Scale bars: A and E = 1.0 mm; and B–D and F, = 0.5 mm.
Included taxa
. This genus includes two subgenera:
Procoscinia
,
n. subg.
, and
Cymbionotum
(
sensu stricto
)
.
Habitat.
The members of
Cymbionotum
occupy the five vegetation zones occupied by the tribe
Melaenini
, with concentration in temperate grassland and tropical deciduous forest and savannah vegetation zones (Table 17)
Geographical distribution
(
Figs. 16
, 18, 20, 21, 23). The range of this genus, including both Western and Eastern Hemispheres, is coextensive with the range of tribe
Melaenini
.