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Article
ZOOTAXA
Programa de Ps-Graduao em Entomologia, Depto. de Ecologia, Zoologia e Gentica, IB, Universidade Federal de Pelotas,
Capo do Leo s/n., 96010-900, Brazil
2
Instituto Nacional de Pesquisas da Amaznia (INPA), Coordenao de Biodiversidade, av. Andr Arajo, 2936, 69060-001, Manaus,
Amazonas, Brazil
3
Pontifcia Universidade Catlica do Rio Grande do SulFaculdade de Biocincias - Depto. de Biodiversidade e Ecologia - Lab. de
Entomologia. Av. Ipiranga, 6681, 90619-900, Porto Alegre, Brazil
4
Programa de Ps-Graduao em Ecologia, Depto. de Biologia Geral, Universidade Federal de Viosa, Campus Universitrio, Av.PH
Rolfs s/n, Viosa, Brazil
5
Programa de Ps-Graduao em Biologia Animal, Instituto de Biocincias, UNESPSo Jos do Rio Preto, SP, Brazil
6
Corresponding author. E-mail: edzefa@gmail.com
Abstract
We provide herein a description of a new species of Oecanthus collected from the tobacco plantation in southern Brazil,
municipality of So Loureno do Sul, State of Rio Grande do Sul. Description focused metanotal gland features, phallic
sclerites, and calling song. A large sampling of individuals was distributed into four groups according to body and appendages color and dotted. We also rank all kind of marks present in the scape and pedicel. We compare linear morphometric
variables of the metanotal glands and tegmina, as well as calling song parameters between groups in order do define all
of them as the same species. Photographs and measurements are provided.
Key words: Insect, Ensifera, taxonomy, bioacoustics, metanotal gland
Introduction
The species of Oecanthus Serville, 1831 present in vivo pale and homogeneous color, usually in shades of green or
brown, often translucent; some have pigmented marks on the ventral region of the scape and pedicel, whose shape
varies according to species (Fulton 1925; Walker 1963). The body color is lost in individuals preserved in alcohol,
but the dotted and pigmented spots around the body and appendages are usually preserved.
Intraspecific variation in body color, as well as scape and pedicel marks in Oecanthus species have been
recorded for a long time. In 1926, Fulton documented geographic variation of these traits in populations of O.
nigricornis Walker, 1869, O. quadripunctatus Beutenmller, 1894, and O. argentinus Saussure, 1874 from United
States, as well as the overlap of these characteristics between these species, which generated doubts about the
identity of each. Walker (1963) has shown that inter and intraspecific shades of brown and green in O. laricis
Walker, 1963 and O. pinus Beutenmller, 1894 vary according to color tree they inhabit. This author also shows
variations in O. nigricornis body color, and scape and pedicel black marking of the O. nigricornis, O. celerinictus
Walker, 1963, O. argentinus, and O. quadripunctatus.
We found a large amount of individuals of Oecanthus in a tobacco plantation, and native bushes in the
municipality of So Loureno do Sul, state of Rio Grande do Sul, Brazil. Two species were previously identified as
O. pallidus Zefa, 2012 and O. lineolatus Saussure, 1897, both with light green body color, and a black rod-shaped
mark in scape and pedicel (Zefa et al. 2012). The other individuals are smaller when compared with both species,
and shown color ranging from yellow-green and light dotts to green-brown with darker dots, which initially led us
to believe dealing with several species.
In this work we separate individuals into four groups according to body color and pigmentation shades, and
compare between groups conventional diagnostic traits as metanotal gland features, phallic sclerites, as well as
calling song, in order to describe a new species of Oecanthus.
267
FIGURES 14. Male habitus of Oecanthus pictus n. sp. paratypes representing the groups according color pattern. 1Male of
the group 1. 2Male of the group 2. 3Male of the group 3. 4Male of the group 4.
Description, male holotype (Figs 512). Body slender, light green, punctulated; Head glabrous; clypeus and
labrum whitish; gena with pigmentar strips; frons with bristles; ecdysial suture pigmented, with short bristles;
group of bristles above the eyes; occiput with a pair of parallel stripes posterior each eyes; green eyes in vivo, but
black eyes outlined of depigmented ommatidia when fixed in alcohol, depigmented ommatidia where lies the
scape; ocelli absent; J-shaped marked scape, L-shaped marked pedicel, antennae flagellum alternating four times
two thinner pingmentar ring with a thicker one, pattern is lost toward apex; maxillary palpi moderately elongate
with five joints, first and second subequal, smaller than the other three, marked with an incomplete pigmentar ring,
third to fifth subequal, third with ventral strip, fourth with incomplete basal ring, and ventral strip, fifth fusiform,
depressed at inner, with incomplete basal ring, and an apical diagonal complete ring; labial palpi three-jointed,
gradual increase in size toward apex, pigmentar mark at apex, third claviform, incomplete pigmentar ring at base
and apex. Thorax: pronotum slightly pubescent, pigmented, translucent, longer as wider, bristles on the border;
ZEFA ET AL.
tegmina translucent, longer than the abdome; hindwings short, reaching first two abdominal tergites; femur I dotted
in dorsal face; tibia I proximal to distal marked with three dots, followed by an incomplete ring, plus three dots and
four pairs of dots, unarmed, swollen near base and mounted each side with an elongated-oval tympanum, inner
longer (0.44mm) than the outer (0.35mm); tarsus I three-jointed, basitarsus lighter and longer than two others
together; leg II with the same pattern of leg I, without tympanum; femur III dotted in dorsal face; tibia III same
marks pattern of tibia I, serrulated, armed with three pairs of subapical spurs tipped with black; there are black
spines between subapical spurs; three apical spurs at outer and four at inner face of the tibia III; basitarsus longer
than the two others together, armed with two subequal, apical, tipped with black spurs; the median tarsus the
smallest. Abdomen: tergites with a pair of transversal strips reducing in intensity backward; cerci slender, shorter
(2.25mm) than the abdomen (5.86mm), marked with a subapical light brown ring; supranal plate translucent,
rounded posteriorly, margin with long bristles; subgenital plate rounded posteriorly, with short lateral protuberance.
Metanotal Gland (Figs 9, 1314) pit wide, scutum with a pair of inflated protuberance, each including a tuft of
bristles projected medio-anteriorly (Fig. 13, a), a dense tuft of long bristles projected medio-posteriorly (Fig. 13, b),
and a tuft of short bristles projected dorso-posteriorly (Fig. 13, c); posterior median lobe with median indentation,
pubescent (Fig. 14, pml); scuto-scutellar suture slightly sinuous, without bristles (Fig. 13, ss); scutellum with
latero-posterior bristles. Measurements cavity area 0.47mm; scutum area 0.89mm; scutellum area 0.34mm;
total gland length 1mm; total gland width 1.43mm; scutum lenght 0.75mm; scutum width 1mm; scutellum length
0.25mm; scutellum width 1.28mm.
Right Tegmen (Fig. 10). Lateral field width, 3.42mm; lateral field length, 6.83mm; 12 acessory veins; dorsal
field projected beyond abdomen; basal area reduced, including veins 1A, 2A, 3A and Cu2, last forming the file with
47 teeth (Fig. 7); file length, 0.86mm; chordal area bearing the veins 2A, 1A e Cu2; harpa area, 3.42mm, harp with
three cross veins slightly sinuous; mirror area, 8.65mm, mirror with two straight cross veins; apical area reduced.
Calling song (Figs 1517). Field recorded in 19.iii.2011, at 19h30, ambient temperature at 14C; trill with
dominant frequency of 2422 Hz, pulse period of 0.037s, and pulse rate of 28 pulses/second.
Observations in Paratypes. There were four groups of body color pattern in O. pictus n. sp.: group 1, light
yellow-green, and very light dotts (Fig. 1); group 2, light green, and dark dotts (Fig. 2); group 3, green, and dotts
darker than group 2 (Fig. 3); and group 4, green-brown, and dotts more intense than the group 3 (Fig. 4).
Legs I, II and III punctulated (Figs 2325); tibiae I and II proximal to distal (dorsal view) with three marks
forming, followed by an incomplete ring, plus three marks, and then three or four pairs of marks (Fig. 25, ag),
some individuals showed failures in the pair marks formation.
Antennae. Scape and pedicel marks varying in the shape and pigmentation marks (Fig. 27).
Phallic sclerites (Figs 2838). Pseudepiphallic main lobe slightly inflated with V-median indentation;
pseudepiphallic apodemes present; rami prolonged laterally, and fused posteriorly; lateral endophallic sclerite
posteriorly bilobed, anteriorly bifurcated, with outer branch truncated (Figs 29, 30, 37, 38, arrow); ectophallic
sclerite posteriorly V-bifurcate with inner branches forming an ectophallic fold, and outer placed between
endophallic lobes.
Male and Female body Measurements. Tab. 1.
TABLE 1. Oecanthus pictus n. sp. Male and female measurements (mm).
Structure
Male
Female (n=10)
body length
110.6 (1011.6)
pronoto length
1.70.06 (1.61.8)
pronoto width
1.70.06 (1.61.8)
head width
1.50.05 (1.41.5)
interocular distance
0.590.03 (0.530.62)
4.20.23 (44.4)
5.70.34 (5.36.7)
ovipositor length
4.20.14 (44.4)
269
FIGURES 512. Oecanthus pictus n. sp. holotype male. 5habitus in lateral view; 6habitus in dorsal view; 7stridulatory
file; 8head and pronotum, dorsal view; 9Metanotal gland; 10right tegmen; 11subgenital plate; 12supranal plate.
ZEFA ET AL.
FIGURES 1314. Metanotal gland of Oecanthus pictus n. sp paratype. 13whole gland, dorsal view; 14posterior median
lobe. Conventions: Scscutum; Stscutellum; ssscutoscutellar suture; pmlposterior median lobe; atuft of bristles
projected medio-anteriorly; bbristles projected medio-posteriorly; cbristles projected dorso-posteriorly.
Right Tegmen Measurements. Dorsal field length 7.8mm0.33 (7.18.7, n=47); dorsal field width
3.6mm0.17 (3.34, n=47); lateral field acessory veins 11.20.97 veins (913, n=36); mirror area 10.1mm0.93
(8.212.2, n=47); harp area 4.3mm0.39 (3.25 n=41); file length 1.1mm0.06 (0.82.6, n=50); file teeth number
463.64 (3949, n=45). The table 2 presents right tegmen measurements comparing the four groups of O. pictus n.
sp. We carried out a Principal Component Analysis comparing of the tegmen measurements in the four groups
deffined by the body colour of specimens, and no differences were seen (component one 90.7%, and component
two 7.6%) (Fig. 39).
TABLE 2. Oecanthus pictus n. sp. rigth tegmen measurements.
Structure/Groups
1, n=12
2, n=12
3, n=11
4, n=6
Average
7.80.23
(7.488.25)
7.80.35
(78.26)
7.60.34
(7.088.21)
7.60.28
(7.227.94)
7.80.33
(7.18.7)
3.690.15
(3.483.94)
3.70.18
(3.273.97)
3.610.17
(3.323.96)
3.570.13
(3.403.78)
3.60.17
(3.34)
11.41.1
(913)
10.90.76
(1012)
11.730.75
(1013)
11.670.94
(1013)
11.20.97
(913)
10.50.87
(9.112.2)
10.20.89
(7.9311.2)
9.540.82
(8.1911.3)
9.800.71
(8.5710.7)
10.10.93
(8.212.2)
4.50.26
(4.124.99)
4.40.38
(3.684.99)
4.040.37
(3.234.81)
4.270.29
(3.904.65)
4.30.39
(3.25)
1.00.04
(0.931.09)
1.00.06
(0.81.05)
0.930.06
(0.791.01)
0.950.04
(0.901)
1.10.06
(0.82.6)
47.22.44
(4451)
45.73.32
(4149)
412.06
(3945)
48.21.67
(4651)
463.64
(3949).
Metanotal Gland Measurements (n=86). Metanotal gland length, 1.2mm0.07 (1.11.4); metanotal gland
width, 1.5mm0,08 (1.21.7); cavity area 0.57mm0,15 (0.450.65); scutum area 1.1mm0.08 (0.91.3mm);
scutellum area 0.42mm0.04 (0.310.52); scutum length 0.88mm0.06 (0.761.11); scutum width 1.5mm0.08
(1.21.7); scutellum length 0.36mm0.03 (0.280.45); scutellum width 1.3mm0.06 (1.11.5). The table 3
presents metanotal gland measurements comparing the four groups of O. pictus n. sp. The comparison of the
tegmen measurements shows no difference in the four groups (Principal Component Analysis: component one
68.8%, and componente two 19.7%) (Fig. 40).
271
FIGURES 1519. Calling songs of Oecanthus pictus n. sp. 15Holotype spectrogram; 16oscillogram of two pulses; 17
oscillogram of the trill; 18regression lines showing the effect of temperature on pulse rate; 19relationship between
dominant frequency and pulse rate.
ZEFA ET AL.
FIGURES 2025. Head, pronotum and legs of Oecanthus pictus n. sp. paratypes. 20, 21, 22head and pronotum, lateral view,
increasing pigmentation; 23legs I and II, respectively, lateral view, inner and outer faces; 24legs III, lateral view; 25
Tibia III, lateral view. Conventions: Lleft leg outer side; Rright leg inner side; a,cset of three marks; bring; dg
pairs of marks.
1, n=17
2, n=43
3, n=14
4, n=12
Average
1.240.06
(1.131.36)
1.240.06
(1.11.35)
1.220.105
(1.091.5)
1.210.063
(1.111.34)
1.20.07
(1.11.4)
Gland width
(mm)
1.480.07
(1.251.56)
1.500.08
(1.31.69)
1.480.068
(1.351.59)
1.450.05
(1.361.53)
1.50.08
(1.21.7)
cavity area
(mm)
0.580.05
(0.50.68)
0.540.034
(0.490.62)
0.540.04
(0.450.64)
0.550.04
(0.490.64)
0.570.15
(0.450.65)
scutum area
(mm)
1.090.07
(11.23)
1.050.08
(0.931.24)
1.060.09
(0.891.17)
1.040.08
(0.951.21)
1.10.08
(0.91.3)
0.440.03
(0.390.5)
0.410.039
(0.330.47)
0.400.04
(0.350.46)
0.410.04
(0.350.48)
0.420.04
(0.310.52)
0.880.042
(0.80.97)
0.880.047
(0.770.97)
0.890.091
(0.761.11)
0.850.045
(0.780.96)
0.880.06
(0.761.11)
1.480.073
(1.251.56)
1.500.084
(1.31.69)
1.480.069
(1.351.59)
1.450.05
(1.361.53)
1.50.08
(1.21.7)
0.350.028
(0.320.41)
0.360.032
(0.310.45)
0.30.038
(0.280.39)
0.350.03
(0.30.41)
0.360.03
(0.280.45)
1.330.048
(1.241.42)
1.300.066
(1.141.46)
1.290.067
(1.21.41)
1.260.041
(1.211.35)
1.30.06
(1.11.5).
Calling song (Figs 1819). Individuals recorded: Group 1: R04 and R05; Group 2: SP04, GG06, GG10, GG11,
GG12, R01, R11, R13, R18, R21, R25, R29, W02, W07, W08, W11, W12 and W14; Group 3: GG05, GG07,
GG09, R06, R09, R12, R15, R17, R19, R20, R22, R26, W01, W03, W04, W06, W09 and W10; Group 4: SP06,
GG14, R08 and R14. There were significant positive effect of temperature on pulse rate (F1,39=717.58, p<0.001,
R=0.95), and of pulse rate on dominant frequency (F1,39=627.88, p<0.001, R=0.94). For both models we did not
detect any difference of pulse rate and dominant frequency between groups (pulse rate x temperature: F3,36=2.32, p
= 0.07; dominant frequency x pulse rate: F3,36=0.16; p=0.92).
A NEW SPECIES OF OECANTHUS FROM BRAZIL
273
Female (Figs 4149). Body color and dotts pattern similar to males; supranal and subgenital plates as Figures
48 and 49, respectively; ovipositor apex denticulated (Figs 4247); Measurements, Tab. 1.
Habitat. Same as O. lineolatus and O. pallidus (Zefa et al. 2012).
FIGURE 27. Oecanthus pictus n. sp. Variation found in scape and pedicel marks, in ventral view. Conventions: apedicel;
bscape.
ZEFA ET AL.
FIGURES 2838. Male genitalia of Oecanthus pictus n. sp. 28ventral view with membranes; 29phallic sclerites, dorsal
view; 30phallic sclerites, ventral view; 31main lobe of pseudepiphallus, dosal view; 32pseudepiphallus, ventral view;
33 pseudepiphallus, dorsal view; 34phallic sclerites, diagonal view; 35pseudepiphallus, diagonal view; 36ectophallic
sclerite; 37ectophallic and endophallic sclerites; 38endophallic sclerite. Conventions: MLPsmain lobe of
pseudepiphallus; MEnd.Sc-Median endophallic sclerite; LEnd.Sclateral endophallic sclerite; End.Sc.pendophallic
sclerite posterior lobes; Ect.Scectophallic sclerite; Ect.Fectophallic fold; Ecendophallic cavity; Rrami.
Discussion
Thirty four of the sixty nine species of Oecanthus have the scape and pedicel features documented, four of them
without marks, and 30 with pigmented marks varying since a simple dot, as occurs in O. alexanderi Walker, 2010,
to elaborated marks as occurs in O. argentinus and O. walkeri Collins & Symes, 2012. It is usual intraspecific
variations in the intensity of the marks, as occurs in O. rileyi Baker, 1905, O. fultoni Walker, 1962, O. forbesi Titus,
1903, O. nigricornis, O. celerinictus, O. argentinus, O. quadripunctatus, and O. californicus Saussure, 1874
(Fulton 1926b; Walker 1962a, 1963). Here we found over forty different scape marks patterns in O. pictus n. sp.,
all of them differing of the marks present in species in which this characteristic has been described.
275
FIGURES 3940. Comparison of the metanotal glands and tegmina by Principal Component Analysis (PCA) using linear
variables. 39Metanotal gland; 40Tegmina. Group 1, red; Group 2, green; Group 3, blue; Group 4, yellow.
Phallic sclerites of nineteen species of Oecanthus had been studied, generally with little detailed descriptions,
focusing on the pseudepiphallic main lobes, which in fact is the most divergent structure among species (Toms &
Otte 1988). Here we break apart O. pictus n. sp. phallic sclerites and found close relationship with O. pallidus and
O. lineolatus (Zefa et al. 2012), but differing in the pseudepiphallic main lobe, and anterior outer branch of the
endophallic sclerite truncated.
The file teeth number were described in fifty species, ranging from 17 in O. exclamationis Davis, 1907 to 65 in
O. rufopictus Chopard, 1932 (Toms & Otte 1988). The file with 463.64 teeth (3949, n=45) separated O. pictus n.
sp. from O. henryi Chopard, 1936 (32.4), O. capensis Saussure, 1878 (5057), O. galpini Toms & Otte, 1988 (63),
O. pseudosimilis Otte, 1988 (37), O. similis Chopard, 1932 (53), O. macer Karsch, 1893 (30), O. sycomorus Toms
& Otte, 1988 (28), O. quadripunctatus (56,7), O. exclamationis (1723), O. leptogrammus Walker, 1962 (2132),
O. niveus (De Geer, 1773) (2230), O. varicornis Walker, 1869 (2636), O. bilineatus Chopard & Chatterjee, 1937
(33), O. indicus Saussure, 1878 (36), O. immaculatus Bruner, 1906 (24), O. jamaicensis Walker, 1969 (31), O.
minutus Saussure, 1878 (37), and O. prolatus Walker, 1967 (28) (Metrani & Balakrishnan 2005; Toms & Otte
1988; Walker 1962a, 1963, 1969).
We compared metanotal gland morphology of O. pictus n. sp. with thirty seven species, and found: i) the
posterior median lobe rounded, as well as scutum bristles similar to O. bakeri Collins & van den Berghe, 2014 and
O. belti Collins & van den Berghe, 2014 (Collins et al. 2014; Eades et al. 2015); ii) bristles tufts position similar to
O. californicus and O. allardi Walker & Gurney, 1960, and scutum and scutellum features similar to O. nigricornis
and O. henryi (Collins 2015; Collins et al. 2014; Metrani & Balakrishnan 2005; Toms & Otte 1988; Walker &
Gurney 1967).
Oecanthus pictus n. sp. shared with O. pictipes the occiput paired postocular lines, punctulate limbs, scape
marks similar to one of the patterns here described, annulate antennae flagellum, as well as the aborted hindwings
(Rehn 1917). But we refuse to consider both as same species since O. pictipes have the body color antimony
yellow, and head vinaceous-rufous (Rehn 1917). Taxonomic status of O. pictipes will be solved after further
specimens collections in the municipality of Natal, State of Rio Grande do Norte, Brazil, followed by detailed
characterization on large amount of individuals. Nothing is known of calling song, metanotal glands, and phallic
sclerites for this species.
ZEFA ET AL.
FIGURES 4149. Oecanthus pictus n. sp. female. 41habitus; 42, 44, 46cerci and ovipositor, lateral, dorsal and ventral
views, respectively; 43, 45, 47ovipositor apex, lateral, dorsal and ventral views, respectively; 48supranal plate; 49
subgenital plate.
277
Acknowledgment
This paper has benefited from grants received from Conselho Nacional de Desenvolvimento Cientfico e
Tecnolgico (CNPq, Brazil) for the scholarship granted to Luciano de Pinho Martins (350059/2010-1 and 350059/
2012-8), as well as financial support received from CNPq (Conselho Nacional de Desenvolvimento Cientfico e
Tecnolgico) and FAPEMIG (Fundao de Amparo Pesquisa do Estado de Minas Gerais)Process number
563360/2010-0EditalMCT/CNPq/MMA/MEC/ CAPES/FNDCTAo Transversal/FAPs N 47/ 2010
Sistema Nacional de Pesquisa em BiodiversidadeSISBIOTA Brasil.
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