Revision of the genus Parada horvath (Hemiptera: Tingidae) with cladistic analysis

15 Pages·2000·5.9 MB·

by  E Guilbert

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 816-830
REVISION OF THE GENUS PARADA HORVATH (HEMIPTERA: TINGIDAE)
WITH CLADISTIC ANALYSIS
Eric Guilbert
ESA 8043 du CNRS, Museum National d'Histoire Naturelle, Laboratoire
d'Entomologie, 45 rue Buffon, F-75005 Paris, France (e-mail: [email protected])
—
Abstract. Three new species ofParada from New Caledonia are described: P. minuta,
P. paitae and P. majuscula. The monotypic Alloeocysta approba Drake is considered a
junior synonym of Parada and to which its only included species is transferred. The
evolution of certain characters, and the relationships among species of Parada are pre-
sented together with a cladistic analysis. An identification key for these species is also
given.
Key Words: Parada, Tingidae, new species, evolution, phylogeny, key
Horvath (1925) erected Parada as a sub- nus, shares with Parada the form ofthe lat-
genus of Cysteochila Stal for the reception eral carinae, the paranota and the hood, and
of Cysteochila {Parada) taeniophora Hor- is considered ajunior synonym.Alloeocysta
vath (1925). Cysteochila has the paranota approba Drake is transferred to Parada. In
widely reflected, inflated, covering the lat- addition, Cysteochila cubens Guilbert (in
eral carinae, and the internal margins (« mar- press), described from New Caledonia
gine intemo ») rounded, while Parada has shows affinities with the species of Parada
the paranota narrowly reflexed, superficially and could be an intermediate species be-
adpressed onto the pronotum and not cov- tween these two genera. Among these re-
ering the lateral carinae, and the internal lationships, the evolution of some charac-
margins slightly sinuate and never rounded ters getting narrower are discussed through
(Horvath 1925). Drake (1942) raised Par- a cladistic analysis. Deposition of types is
ada to generic level, and add two new spe- specified: MNHN for Museum National
BPBM
cies to the genus {P. torta and P. popla). d'Histoire Naturelle, Paris; for
Later, he added four more new species to Bernice P. Bishop Museum, Hawaii. All
the genus {P. absona, P. darlingtoni and P. measurements are in millimeters.
hackeri Drake 1952; P. solla Drake and Ru-
hoff 1961). Systematics
The genus presently comprises seven
Parada Horvath 1925
Australian species (New South Wales and
Queensland) (Drake and Ruhoff 1965). Cysteochila {Parada) Horvath 1925: 3.
Three new species from New Caledonia are Type species: Cysteochila {Parada) tae-
described here. In addition, Drake (1961) niophora Horvath 1925: 3, by orig. desig.
described a new genus and a new species Parada: Drake 1942: 4.
from a single specimen caught in New Alloeocysta Drake 1961: 109. New Syn-
South Wales: Alloeocysta approba. This onymy. Type species: Alloeocysta appro-
species, the only included species ofthe ge- ba Drake 1961: 109, by orig. desig.
VOLUME 102, NUMBER 4 817
—
Diagnosis. Head with at least two fron- form a subcylindrical areolate cyst that is
tal and two occipital spines. Bucculae wide, closed inward by the internal margins
with apices meeting in front. Antenna long, touching the pronotum, while it is not the
slender, first segment short, second small- case for the species of Parada. The hem-
est, third longest and fourth longerthan first elytra are a little longer than abdomen with
two together, pubescent. a costal area uniseriate, a subcostal areabis-
Pronotum gibbose, deeply punctuate, are- eriate and a discoidal area four to five are-
olate at apex, pubescent near collar, tricar- olae broad. Despite some characters which
inate. Median carina long, slightly erected, distinguishA. approba from species ofPar-
contiguous to hood. Lateral carinae elevat- ada; approba being the unique species of
ed, ending before collar, curved inward, not the genus Alloeocysta, the latter is consid-
resting on pronotum, modified as to form a ered a synonym ofthe genus Parada. Thus,
long inflated areolate cyst. Hood inflated, I propose the new combination Parada ap-
raised. Paranota long, wide, reflexed up- proba (D—rake) n. comb.
right, not resting on pronotum, three to four Type. Holotype: 6, Bogan river. New
areolae broad, almost reaching lateral cari- South Wales, Australia (Australian Muse-
nae. um, Sydney).
Hemelytra wider and longer than abdo-
men, slightly wider than pronotum width. Parada majuscula Guilbert, new species
Costal area uni- to biseriate, outer margins —(Figs. 1-3)
bent upward. Subcostal areabi- to triseriate, Description. Head and body black, legs
outer margins bent downward. Discoidal and antenna yellowish to brown fuscous,
area five to seven areolae broad. Sutural distal part of fourth antennal segment and
area large, with larger areolae than subcos- tarsi brown to black. Pronotum clear brown
tal and discoidal areas. Hypocostal laminae to fuscous. Hemelytra yellowish with three
uniseriate. dark brown spots, one near base that may
occupy just basal third of costal area or be
Parada approba (Drake), new expanded to cover all of the basal third of
combination the hemelytron, plus clear brown spots at
(Figs. 7-8) apical third and apex of costal area. Length
Alloeocysta approba Drake 1961: 109. 3.05; width 1.17.
— Head with two frontal and two small,
Comments. According to Drake (1961) slender occipital spines. Bucculae wide,
Alloeocysta approba is different "from oth- triseriate, with apices meeting in front. La-
er Australian genera by having the hood bium reaching hind coxae and labial sulcus
distinctly wider than long and the lateral ca- sinuate. Sternal laminae widely separated
rinae inflated, cyst-like on disc of prono- on metastemum. Antennal measurements: I,
tum" (Holotype measurements, body length 0.19: II, 0.12: III, 1.01: IV, 0.45), last an-
2.99; body width 1.26; hood length 0.37; tennal segment pubescent.
hood width 0.55). All species of Parada Pronotum gibbose, deeply punctuate, are-
have lateral carinae inflated likeA. approba olate on hind process, pubescent nearcollar,
and some of them have a hood wider than tricarinate. Median carina long, slightly el-
long. This is the case of P. majuscula and evated, contiguous to hood. Lateral carinae
P. paitae. Alloeocysta approba has anten- elevated, ending before collar, curved in-
nae as long as that ofthe species ofParada ward, not resting on pronotum, modified as
(I, 0.15: II, 0.12: III, 0.85: IV, 0.3), a pron- to form a long inflated open cyst, four are-
otum gibbose, deeply punctuate and areo- olae broad. Hood inflated, raised, slightly
late at apex, a hood forming a vesicule wid- wider than long (length 0.33; width 0.38).
er than long, lateral carinae modified as to Paranota long, wide, reflexed and partially
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
818
Fig. 1. Parada majuscula, habitus. Scale = 1 mm.
VOLUME 102, NUMBER 4 819
Figs. 2-3. Parada majuscula. 2, Profile. 3, Ventral face. Scale = 1 mm.
recurved above pronotum, not resting on CM. Yoshimoto coll. (BPBM); 1 c?. New
pronotum, four small areolae broad, almost Caledonia, Mt Koghis, 400-600 m, 11.1980,
reaching lateral carinae. N.L.H. Krauss coll.. Ace. #1980.128
Hemelytra wider, longer than abdomen, (BPBM). —
slightly wider than pronotum width. Costal Comments. The body length of this
area bent upward, mostly biseriate but un- species may be sexually dimorphic (Table
iseriate at apex, outer areolae larger than 1), but this characteristic cannot be assessed
inner. Subcostal area bent downward, tris- here because of the small number of spec-
eriate. Discoidal area pentaseriate. Sutural imens.
area large, with larger areolae than subcos-
Parada paitae Guilbert, new species
tal and discoidal areas at apex. Hypocostal
(Figs. 4-5)
laminae u—niseriate. —
Types. Holotype: c?. New Caledonia, Description. Head and body brown.
Riviere Bleue P7, dense evergreen rainfo- Pronotum, hemelytra, legs and antenna
rest, 24.x.1992, L. Bonnet de Larbogne, J. clear to yellowish, apex of last antennal
Chazeau & E. Guilbert colls (fogging) segment brown fuscous. Length 2.98; width
(MNHN). Paratypes: 1 $, same data as ho- 1.17; hood length 0.31; hood width 0.31.
lotype (MNHN); 3 d and 1 ?, New Cale- Same characters as P. majuscula, except
donia, Yahoue III.1978, N.L.H. Krauss rostrum extends between median coxae but
coll.. Ace. #1978.114 (BPBM); 1 9, New does not reach hind coxae, costal area is
Caledonia, 6 km N of Paita, 25.1.1963, uniseriate, areolae are rather large, except
820 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Means of body, hood and antennal segments measurements for 5 males and 3 females of P. ma-
jiiscula in mm.
VOLUME 102, NUMBER 4 821
Table 2. Body length, body width, hood length, hood width and antennal segments measurements i the
New Caledonian species and P. approba. Last antennal segment measurements on P. approha is missing on the
type.
Body AntennalSegments
Species
822 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-8. Parada approba. 7, Profile. 8, Ventral face. Scale = 1 mm.
Cladistic Analysis active and ten inactive. These ten characters
—
Choice of the taxa. The eleven species define the size ofvarious structures in terms
of the genus Parada are included in this of number of areolae. They are not used in
analysis as the ingroup. Three species are the analysis because the homology between
included as outgroups: Cysteochila cubens the areolae cannot be established with cer-
Guilbert, Physatocheila dissimilis Guilbert tainty, but they figure as attributes. When a
and Nobamus pilosus Guilbert. Cysteochila structure has two areolae, does it mean that
cubens shows some affinities with the genus an areola divided into two areolae, or does
Parada, while A^. pilosus is clearly different. a second areola appears near the existing
Physatocheila dissimilis could be considered one? The results of the two analyses are
as intermediate between these two generaby compared. Inactive characters in the second
its general morphology. Therefore, 14 spe- analysis are mentioned. Characters of the
cies were included—in this analysis. genitalia are not taken into account because
Character states. Sixty-one morpholog- they do not vary within the genus. Multi-
ical characters are analysed. They are ana- state characters are treated unordered. The
lysed in two differents ways. A first anal- matrix of characters is given in Table 3.
ysis concern all the characters. A second Question marks representcharacterslacking
analysis concern only fifty-one characters or uncertainly stated.
.
VOLUME NUMBER
102, 4 823
Figs. 9-15. Ptirada spp. left costal area, paranotaand lateral carinae. 9, P. popla. 10, P. absona. 11, Ptorta.
12, P. hackeri. 13, P. taeniophora. 14, P. darlingtoni. 15, P. sollo. Scale = 1 mm.
0. Head black (0), brown (1), yellowish to 12. Occipital spines stout (0), slender (1).
white (2). 13. Median spine long (0), short (1), tuber-
1. Pronotum black (0), brown (1), yellow- cle-like (2), absent (3).
ish (2). 14 Median spine erect (0), prostrate (1).
2. Abdomen black (0), brown (1), yellow- 15 Median spine stout (0), slender (1).
ish (2). 16. Antennae densely pubescent (0), not
3. Hemelytra black (0), brown (1), yel- pubescent (1).
lowish to white (2), brown with colored 17. Last antennal segment longer than first
spots (3). and second together (0), shorter than
4. Body glabrous (0), pilose (1). first and second together (1).
5. Head convex (0), flat (1). 18. Last antennal segment longer than first
6. Frontal spines long (0), short (1). (0), shorter than first (1).
7. Frontal spines erect (0), prostrate (1). 19. Third antennal segment 1 time as long
8. Frontal spines stout (0), slender (1). as the last one (0), 2 times (1), 3 times
9. Frontal spines in contact or crossed (0), (2), 4 times (3).
not in contact (1). 20. Bucculae biseriate (0), triseriate (1),
10. Occipital spines long (0), short (1). quadriseriate (2), pentaseriate (3). In-
1 1 Occipital spines adpressed (0), pros- active character.
trate (1). The occipital spines are pros- 21. Bucculae open in front (0), closed in
trate when the spines do not rest onto front (1).
the head, otherwise they are adpressed. 22. Labium almost extending beyond the
824 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 3. Matrix of states ofthe 61 characters for 14 species.
Characterstates
Nobamus pilosiis 00001100000003??0111012100100?00?01??0004200010010030??202133
Physatncheila dissimilis 01110100000100001102101110101210701??0111011111111001??11042
CysteochUa cubens 222300011000120?1?0221200101073051127032001111111110102010512
Parada darlingUmi 1013000010001011100211107001121122032111411110110112110111220
Parada hackeri 1113010100000111100101401011122121117111311110110110110111110
Parada torta 101200111000111110022110100110214212111141111011011210011112
Parada absona 0003000111001111100211100001122152110111211111110111110111220
Paradapopla 0013001110001111100211300001122132042111411110110111120111320
Parada solla 1113010110001111100211201001112162121121311111110111100111221
Parada taeniophora 10130011100011011??21111100111212202112131111011011211011112
Parada majuscida 01030011101013??100121010101112102132121311111110111122111111
Parada paitae 11110011111013??100111210001122112111121411111110111101111221
Parada mimita 22230011101013??10011131000112210211112131111111011110011112
Parada approba 101300011?00?00?100311310101??212214211141111011011111011102
metastemum (0), extending to the mid- 14 areolae long (5), 15 areolae long (6).
dle of the metastemum (1), extending Inactive character.
little beyond the mesosternum (2), 33. Hood absent (0), present but flat (1),
reaching the posterior margins of the cyst-like and partly covering the head
mesosternum (3), extending to the mid- (2). When the hood is present, it covers
dle of the mesosternum (4). most of the head but generally not the
23. Mesostemal laminae subparallel (0), eyes.
not subparallel (1). 34. Hood higher than posterior pronotal
24. Metasternal laminae wide (0), narrow lobe (0), not higherthan posteriorpron-
(1). otal lobe (1).
25. Sternal laminae open behind (0), closed 35. Hood 3 areolae long (0), 4 areolae long
behind (1). (1), 5 areolae long (2), 6 areolae long
26 Pronotum strongly gibbose (0), slightly (3), 7 areolae long (4). Inactive char-
gibbose (1). acter.
27 Pronotum pubescent (0), glabrous (1). 36. Hood 1 areola high (0), 2 areolae high
28 Median carina not areolate (0), uniser- (1), 3 areolae high (2). Inactive char-
iate (1). acter.
29. Median carina with large areolae (0), 37. Collar triseriate (0), biseriate (1). In-
small areolae (1), minute areolae (2). active character.
30. Lateral carinae ridge like (0), with one 38. Paranota 2 areolae broad (0), 3 areolae
or two rows of areolae (1), with 3 to 5 broad (1), 4 areolae broad (2), 5 areolae
rows of areolae (2), with 6 or more broad (3). Inactive character.
than 6 rows of areolae (3). Inactive 39. Paranota with large areolae (0), small
character. areolae (1), minute areolae (2).
31. Lateral carinae erect (0), reflexed but 40. Paranota more than 14 areolae long (0),
not resting onto the pronotum (1), re- 13-14 areolae long (1), 11-12 areolae
flexed and resting onto the pronotum long (2), 10 areolae long (3), 8-9 are-
(2). olae long (4). Inactive character.
32. Lateral carinae 8 areolae long (0), 9 41. Paranota reflexed with free margins in
areolae long (1), 10 areolae long (2), contact with pronotum (0), reflexed
11 areolae long (3), 12 areolae long (4), with free margins not in contact with
VOLUME 102, NUMBER 4 825
pronotum (1), not reflexed with free areolae wide (1), 12 areolae wide (2),
margins not in contact with pronotum less than 6 areolae wide (3). Inactive
(2). character.
42. Paranota not covering pronotum (0),
covering part of pronotum (1). The phylogenetic analysis was performed
43. Hemelytra wide (0), narrow (1). Hem- with Hennig86 (Farris 1988). The algorithm
elytra are considered narrow when not ie* was used to build tree (s), using Fitch
extending far beyond the abdomen. parsimony.
44. Hemelytra sharply widened at base (0),
Results
not sharply widened at base (1). —
45. Junction of RM and Cu veins raised First analysis. Three trees of 119
and forming a little swollen zone (0), steps, with ci = 61 and ri = 50 were gen-
not raised and forming a little swollen erated. They differ by the relative position
zone (1). of the three New Caledonian species
46. Hemelytra areolae hyaline (0), not hy- among themselves. Seventy-seven percent
aline (1). of the active characters are not informa-
47. Hemelytra pubescent (0), glabrous (1). tive for the resolution of these conflicting
48. Hemelytral veins raised, forming ridges branches. Here is presented one of the
(0), not raised and forming ridges (1). trees which corresponds to the following
49. Costal area wide (0), narrow (1). interpretation (Fig. 16). For the 7 infor-
50. Costal area plane (0), raised (1). mative characters, Parada minuta and P.
51. Costal area areolae very small (0), paitae share the same state for four of
small (1), large (2), very large (3). them, while P. majuscula and P. minuta
52. Costal area 6 areolae broad (0), uni- share the same state for one of them, and
seriate (1). P. majuscula and P. paitae for any of
53. Costal area uniseriate at base (0), bi- them. Then, Parada minuta may be more
seriate at base along 3-5 areolae (1), closely related to P. paitae than to P. ma-
biseriate at base along 7-10 areolae (2). juscula. The coefficients and number of
The costal area is mostly uniseriate and steps for each characters is given Table 4.
shows in part two areolae for the same The genus Parada (including the species
width like if the original areolae were approba) is monophyletic. The monophyly
divided in two smaller areolae (Figs. of the genus is supported by the raised
9-15). veins on the hemelytra, the lateral carinae
54. Costal area uniseriate at the middle (0), and the paranota which are curved inwards
biseriate at the middle along 2-3 are- but not resting onto the pronotum.
olae (1), biseriate at the middle along The genus is divided in two groups, the
5-6 areolae (2). New Caledonian group which comprise
55. Subcostal area areolae very small (0), the three New Caledonian species, and the
small (1), large (2), very large (3). Australian group comprising the other
56. Subcostal area large (0), small (1). eight species. The Australian species have
57. Discoidal area areolae very small (0), five cephalic spines, while New Caledon-
small (1), large (2), very large (3). ian species have four. No unambigous
58. Discoidal area 4 areolae wide (0), 5 synapomorphy support the monophyly of
areolae wide (1), 6 areolae wide (2), 7 the Australian group, while the short oc-
areolae wide (3), 8 areolae wide (4), 9 cipital spine and the lack of a median
areolae wide (5). Inactive character. spine are the unambiguous synapomor-
59. Sutural area areolae very small (0), phies of the New Caledonian group. Par-
small (1), large (2), very large (3). ada approba is completely integrated in
60. Sutural area 6-7 areolae wide (0), 8-9 the genus; however, it is the basal species