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SYSTEMATICS
Descriptions of Immature Stages of Three Species of Laccocorinae
(Hemiptera: Heteroptera: Naucoridae) From India
S. SUNDAR,
1,2
ROBERT W. SITES,
3,4
AND M. ARUNACHALAM
1
Ann. Entomol. Soc. Am. 107(6): 1056Ð1065 (2014); DOI: http://dx.doi.org/10.1603/AN14051
ABSTRACT Adults and Þrst through Þfth instars of three species of Naucoridae in the subfamily
Laccocorinae were collected from localities in Tamil Nadu and Karnataka states in southern India.
SpeciÞcally, Heleocoris indicus (Montandon) and Heleocoris vicinus (Montandon) were collected from
the vegetated margins of streams that contained no congeners. Pogonocaudina indica Sites & Zettel
was collected from vertical rock surfaces at the type locality. All instars of these three species are
described here. First instars of both species of Heleocoris are immaculate, whereas subsequent instars
are densely punctate. The Þrst instar of P. indica has pronounced dorsal tubercles with stout spines,
whereas subsequent instars lack these structures. All instars of P. indica have a deep terminal cleft and
dense fringe of hairs on the perimeter of the posterior abdominal segments. Second through Þfth
instars of these three species can be distinguished based on relative length of the mesonotal wing pad
compared with the exposed part of the metanotal lateral margin.
KEY WORDS Nepomorpha, Naucoridae, Laccocorinae, India, nymph
The Indian subcontinent is host to a diversity of spe-
cies of creeping water bugs in the subfamily Lacco-
corinae. Recently, in a treatment of two Indian genera
that occur in waterfalls, a new genus was described
and another revised (Sites et al. 2011). More speciÞ-
cally, Diaphorocoris is represented by three species in
India and a fourth in Sri Lanka; the monotypic Pogono-
caudina indica Sites & Zettel is known only from
southern India (Sites et al. 2011). The most speciose
genus of Laccocorinae in India is Heleocoris, with 11 of
the 28 described species recorded from that country
(Thirumalai 2007). An undescribed species of Cteni-
pocoris also occurs in India. Species of Heleocoris occur
in rivers and streams, while those of Diaphorocoris and
Pogonocaudina occur on the vertical rock faces of
waterfalls; Ctenipocoris is lentic.
Although the family is represented worldwide by
⬇380 described species, the nymphal stages of only a
few of these have been studied including Ambrysus
attenuatus Montandon (Lopez Ruf and Hernandez
2007), Ambrysus bergi Montandon, Ambrysus ochra-
ceus Montandon (Lopez Ruf 1990), Ambrysus funebris
La Rivers (Whiteman and Sites 2008), Ambrysus lu-
natus lunatus Usinger (Sites and Nichols 1990), Am-
brysus mormon Montandon (Usinger 1946), Cry-
phocricos barozzii Signoret (Lopez Ruf 1996a),
Cryphocricos hungerfordi Usinger (Sites and Nichols
1993), Ctenipocoris schadei (De Carlo) (Lopez Ruf
2004), Limnocoris lutzi La Rivers (Sites and Nichols
2001), Limnocoris ovatulus Montandon (Lopez Ruf
1993), Pelocoris binotulatus nigriculus Berg (Estevez et
al. 1989), Pelocoris femoratus femoratus (Palisot de
Beauvois) (de la Torre-Bueno 1903, Hungerford 1927 [as
P. carolinensis, see La Rivers 1948], McPherson et al.
1987), and Placomerus micans La Rivers (Lopez Ruf
1996b).
Heleocoris indicus (Montandon) and Heleocoris vici-
nus (Montandon) are among the most common spe-
cies in the genus in southern India. Little information
exists in the literature concerning identiÞcation of the
species of Heleocoris in the adult stage, and the
nymphal stages are unknown. Adults of H. indicus and
H. vicinus can be distinguished most easily by differ-
ences in an abdominal characteristic of the male. Spe-
ciÞcally, the left sixth ventral laterotergite is com-
pletely ßat in H. vicinus and partially recessed in H.
indicus. Diagonostic information for adults of Pogono-
caudina and a key to the species of Naucoridae oc-
curring in waterfalls in southern India were presented
by Sites et al. (2011); however, nymphal stages are
unknown. Presented here are descriptions of all Þve
nymphal instars of three species of Laccocorinae: H.
indicus, H. vicinus, and P. indica.
Materials and Methods
All specimens of the three species were collected in
various water bodies of Tamil Nadu state in southern
India. H. vicinus also was collected in Karnataka, which
borders Tamil Nadu to the northwest. Descriptions
1
Sri Paramakalyani Centre for Environmental Sciences, Manon-
maniam Sundaranar University, Alwarkurichi, Tamilnadu, India.
2
Present address: Department of Ecology and Environmental Sci-
ences, School of Life Sciences, Pondicherry University, Puducherry
605014, India.
3
Enns Entomology Museum, Division of Plant Sciences, University
of Missouri, Columbia, MO 65211.
4
Corresponding author, e-mail: bugs@missouri.edu.
0013-8746/14/1056Ð1065$04.00/0 䉷2014 Entomological Society of America
were prepared using only Þeld-collected specimens.
Adults and nymphs of Heleocoris were collected from
vegetated margins of streams in which no congeners
occurred, thereby ensuring Þdelity of nymphal asso-
ciation with adults. More speciÞcally, adult and im-
mature specimens of H. indicus were collected at
Tamil Nadu, Illuppiar, 08⬚47⬘54.6⬙N, 77⬚19⬘37.7⬙Eon
27 March 2010. Adult and immature specimens of
H. vicinus were collected at Tamil Nadu, Kurangani,
10⬚05⬘24.7⬙N, 77⬚14⬘50.7⬙E on 26 October 2010;
Karnataka, Meena Kundi, Nethravathi, 12⬚55⬘16.5⬙N,
75⬚30⬘26.1⬙E on 20 April 2010; and Karnataka, Thin-
galae (Commerce Graduate Farm), Seethanathi, 13⬚
27⬘17.9⬙N, 75⬚3⬘33.2⬙E on 19 April 2010. Adult and
immature specimens of P. indica were collected in
Tamil Nadu at the type locality of Kiliyur Waterfall on
11 March 2010 and 25 October 2010. Samples of H.
indicus and H. vicinus were taken using an aquatic
D-net by kick-sampling the rocky substratum and mar-
ginal vegetation. Samples of P. indica were collected
by hand picking the insects from the surface of rocks
of the waterfall with either Þngers or forceps. A soft-
bristled brush was used to sweep over the rocks, which
did not dislodge the insects but disturbed them suf-
Þciently that they moved, thereby enabling detection
and collection. All samples were preserved in 70Ð80%
ethyl alcohol.
Measurements are based on 1Ð6 individuals of each
stage and given in millimeters. All specimens have
been deposited as vouchers in the Enns Entomology
Museum, University of Missouri, Columbia, MO. Pho-
tographs were prepared using the Leica Application
Suite v.4.4 Extended Depth of Focus module with a
Leica MZ16 stereomicroscope. Descriptive measure-
ments are given in Tables 1Ð3. Length of body was
measured from the tip of the labrum to the tip of the
abdomen; width was measured across the widest part
of the body, usually the metathorax.
Systematics
The Þrst instar is described in detail, but only
changes unrelated to those associated with allometric
development in subsequent instars are described.
Typically, the nymphal coloration in live specimens is
greenish yellow but this fades to yellow after preser-
vation in alcohol. Thus, coloration reported here may
appear slightly greener in live specimens. Patterns and
extent of maculation vary among individuals and typ-
ically become more pronounced in older individuals
within an instar and among instars.
A notable, previously unknown difference among
subfamilies of Naucoridae is the number of meso- and
metatarsal segments of the nymphs. In Cryphocricinae,
A. lunatus and C. hungerfordi have two-segmented tarsi
(Sites and Nichols 1990, 1993, respectively); in Limno-
corinae, L. lutzi La Rivers has two tarsal segments (Sites
and Nichols 2001); and in Naucorinae, P. femoratus Pal-
isot de Beauvois has two tarsal segments (McPherson et
al. 1987). Although the nymphal stages have been de-
scribed for one species in Laccocorinae, C. schadei (Lo-
pez Ruf 2004), no mention was made of the number of
tarsal segments. Nymphal stages have not been de-
scribed for any species of Cheirochelinae. Only a single
meso- and metatarsal segment occur in the three species
of Laccocorinae reported here.
Heleocoris indicus Montandon
Fig. 1AÐE
Heleocoris indicus Montandon: in Montandon 1897:
450Ð451.
Descriptions of Immature Stages. First Instar (Fig.
1A). Length, 2.44 ⫾0.02 mm; width, 1.46 ⫾0.02 mm
(n⫽2). Body elongate-oval, general appearance dor-
soventrally ßattened, greatest width at meso- and met-
anota; overall convex dorsally, concave ventrally; im-
Table 1. Descriptive measurements (mean mm ⴞSE) of instars of H. indicus
Character
a
First instar Second instar
b
Third instar Fourth instar 4 Fifth instar
Body length 2.44 ⫾0.02 3.29 ⫾0.05 4.50 ⫾0.06 5.68 7.73 ⫾0.17
Body width 1.46 ⫾0.02 2.08 2.78 ⫾0.06 3.64 4.99 ⫾0.03
Head length 0.49 ⫾0.03 0.67 ⫾0.01 0.83 ⫾0.01 1.08 1.31 ⫾0.01
Head width 1.03 ⫾0.01 1.36 1.74 ⫾0.02 2.32 2.96 ⫾0.02
Synthlipsis 0.48 ⫾0.03 0.58 ⫾0.02 0.73 ⫾0.01 1.04 1.15 ⫾0.03
Pronotum length 0.29 ⫾0.01 0.40 ⫾0.01 0.55 ⫾0.01 0.70 1.02 ⫾0.01
Mesonotum length 0.23 ⫾0.01 0.35 ⫾0.01 0.54 ⫾0.02 0.76 1.13 ⫾0.02
Metanotum length 0.40 ⫾0.00 0.52 ⫾0.02 0.73 ⫾0.03 0.94 1.15 ⫾0.02
Leg length
Profemur 0.56 ⫾0.02 0.77 ⫾0.02 0.98 ⫾0.00 1.24 1.69 ⫾0.03
Protibia 0.36 ⫾0.00 0.49 ⫾0.01 0.62 ⫾0.00 0.86 1.18 ⫾0.01
Protarsus 0.18 ⫾0.00 0.21 ⫾0.01 0.27 ⫾0.01 0.31 0.42 ⫾0.01
Mesofemur 0.53 ⫾0.01 0.69 ⫾0.01 0.94 ⫾0.02 1.28 1.68 ⫾0.01
Mesotibia 0.37 ⫾0.01 0.49 ⫾0.01 0.67 ⫾0.01 0.88 1.21 ⫾0.01
Mesotarsus 0.22 ⫾0.00 0.27 ⫾0.02 0.35 ⫾0.01 0.44 0.57 ⫾0.02
Metafemur 0.58 ⫾0.08 0.90 ⫾0.00 1.17 ⫾0.03 1.60 2.13 ⫾0.02
Metatibia 0.69 ⫾0.01 1.01 ⫾0.01 1.28 ⫾0.02 1.74 2.31 ⫾0.02
Metatarsus 0.43 ⫾0.01 0.55 ⫾0.02 0.67 ⫾0.03 0.92 1.09 ⫾0.02
SE values ⬍0.005 are listed as 0.00. Number of specimens measured for each instar: Þrst, 2; second, 3; third, 2; fourth, 1; Þfth, 6.
a
Measurements of body, head, and notal lengths taken at midline.
b
Two specimens unnaturally decurved laterally and most widths not measurable.
November 2014 SUNDAR ET AL.: IMMATURE STAGES OF LACCOCORINAE FROM INDIA 1057
maculate yellowish brown dorsally and ventrally;
dorsally with pale margins.
Head transverse, anterior margin convex, continu-
ous with lateral margins of pronotum, posterior margin
convex, immaculate yellowish brown dorsally and
ventrally. Compound eyes dark red, ßattened, con-
vergent anteriorly. Synthlipsis ⬇2.2 times the width of
one eye. Lateral hyperoche slightly widening ante-
riorly. Antenna light yellow, apparently four-seg-
mented, basal segment recessed within head. Labrum
originates caudad from anterior margin of head. La-
bium yellowish white, elongate-conical, with three
visible segments, overall length ⬇1.6 times the width
at base, segment 1 mostly within head, segment 2
subequal in length to 3.
Nota yellowish brown with pale lateral margins.
Spines arising from tubercles lacking. Pronotum with
anterior margin concave; lateral margins evenly
rounded; posterior margin straight medially, slightly
produced caudad laterally; overlaps ⬇20% of mesono-
tum. Mesonotum with posterior margin mostly
straight medially, with slight indication of wing pads
posterolaterally. Metanotum longest of nota at mid-
line, 1.4 times the length of pronotum, 1.7 times the
length of mesonotum. Metanotal wing pads not evi-
dent. Metanotum completely overlaps abdominal ter-
gum I and 80Ð90% of tergum II at midline.
Prothoracic leg retentorial (see Sites and Nichols
1990), generally milky white with yellowish brown
tibia, tarsus, and pretarsal claws. Procoxa ⬇2.0 times
the length of trochanter, ⬇0.3 times the length of
femur. Profemur compressed, two dense brushes of
setae along anterior margin above and below row
of stout spines. Protibia and tarsus with sulcus extend-
Table 2. Descriptive measurements (mean mm ⴞSE) of instars of H. vicinus
Character
a
First instar
b
Second instar Third instar 3 Fourth instar Fifth instar
Body length 2.88 ⫾0.04 3.90 4.80 ⫾0.12 6.00 ⫾0.08 7.79 ⫾0.12
Body width 1.72 ⫾0.02 2.32 2.91 ⫾0.05 3.71 ⫾0.03 5.12 ⫾0.08
Head length 0.68 ⫾0.02 0.80 0.96 ⫾0.02 1.10 ⫾0.02 1.35 ⫾0.01
Head width 1.18 ⫾0.02 1.54 1.92 ⫾0.04 2.39 ⫾0.03 3.05 ⫾0.06
Synthlipsis 0.48 ⫾0.02 0.66 0.75 ⫾0.05 0.94 ⫾0.03 1.13 ⫾0.04
Pronotum length 0.34 ⫾0.02 0.46 0.60 ⫾0.00 0.78 ⫾0.01 1.07 ⫾0.01
Mesonotum length 0.27 ⫾0.01 0.40 0.59 ⫾0.01 0.84 ⫾0.04 1.20 ⫾0.02
Metanotum length 0.50 ⫾0.02 0.60 0.77 ⫾0.01 0.96 ⫾0.02 1.12 ⫾0.02
Leg length
Profemur 0.69 ⫾0.01 0.90 1.09 ⫾0.01 1.37 ⫾0.01 1.78 ⫾0.01
Protibia 0.48 ⫾0.00 0.62 0.78 ⫾0.04 1.01 ⫾0.04 1.31 ⫾0.06
Protarsus 0.20 ⫾0.00 0.26 0.31 ⫾0.01 0.37 ⫾0.01 0.45 ⫾0.03
Mesofemur 0.62 0.84 1.06 ⫾0.02 1.32 ⫾0.02 1.80 ⫾0.02
Mesotibia 0.40 0.56 0.73 ⫾0.02 0.91 ⫾0.01 1.25 ⫾0.03
Mesotarsus 0.26 0.30 0.34 ⫾0.01 0.46 ⫾0.03 0.55 ⫾0.03
Metafemur 0.79 ⫾0.01 1.04 1.29 ⫾0.04 1.70 ⫾0.02 2.24 ⫾0.01
Metatibia 0.83 ⫾0.01 1.14 1.42 ⫾0.02 1.85 ⫾0.02 2.41 ⫾0.02
Metatarsus 0.50 ⫾0.02 0.58 0.69 ⫾0.01 0.87 ⫾0.01 1.13 ⫾0.01
SE values ⬍0.005 are listed as 0.00. Number of specimens measured for each instar: Þrst, 2; second, 1; third, 3; fourth, 5; Þfth, 3.
a
Measurements of body, head, and notal lengths taken at midline.
b
One specimen missing both mesothoracic legs.
Table 3. Descriptive measurements (mean mm ⴞSE) of instars of P. indica
Character
a
First instar Second instar Third instar 3 Fourth instar Fifth instar
b
Body length 2.77 ⫾0.04 3.74 ⫾0.05 4.97 ⫾0.01 6.72 ⫾0.00 8.38 ⫾0.10
Body width 1.80 ⫾0.02 2.52 ⫾0.05 3.27 ⫾0.04 4.32 ⫾0.16 5.36 ⫾0.08
Head length 0.67 ⫾0.01 0.87 ⫾0.02 1.07 ⫾0.01 1.38 ⫾0.02 1.54 ⫾0.06
Head width 1.17 ⫾0.02 1.60 ⫾0.01 2.05 ⫾0.01 2.58 ⫾0.10 3.20 ⫾0.04
Synthlipsis 0.67 ⫾0.02 0.88 ⫾0.02 1.06 ⫾0.03 1.36 ⫾0.00 1.62 ⫾0.02
Pronotal length 0.40 ⫾0.00 0.57 ⫾0.00 0.77 ⫾0.01 1.02 ⫾0.02 1.33 ⫾0.01
Mesonotum length 0.36 ⫾0.02 0.53 ⫾0.01 0.77 ⫾0.02 1.12 ⫾0.04 1.62 ⫾0.02
Metanotum length 0.40 ⫾0.00 0.52 ⫾0.01 0.64 ⫾0.01 0.86 ⫾0.06 0.88 ⫾0.00
Leg length
Profemur 0.67 ⫾0.01 0.87 ⫾0.01 1.12 ⫾0.01 1.46 ⫾0.04 1.88
Protibia 0.38 ⫾0.01 0.52 ⫾0.00 0.71 ⫾0.01 0.95 ⫾0.03
Protarsus 0.23 ⫾0.01 0.28 ⫾0.01 0.34 ⫾0.00 0.41 ⫾0.01
Mesofemur 0.75 ⫾0.02 1.01 ⫾0.00 1.36 ⫾0.02 1.80 ⫾0.06 2.34 ⫾0.02
Mesotibia 0.56 ⫾0.01 0.77 ⫾0.01 1.06 ⫾0.03 1.38 ⫾0.04 1.90 ⫾0.02
Mesotarsus 0.35 ⫾0.01 0.41 ⫾0.01 0.51 ⫾0.01 0.63 ⫾0.01 0.84 ⫾0.04
Metafemur 0.90 ⫾0.01 1.24 ⫾0.02 1.66 ⫾0.02 2.27 ⫾0.03 2.92 ⫾0.00
Metatibia 0.95 ⫾0.01 1.34 ⫾0.02 1.81 ⫾0.02 2.49 ⫾0.05 3.26 ⫾0.02
Metatarsus 0.53 ⫾0.01 0.66 ⫾0.01 0.82 ⫾0.01 1.07 ⫾0.01 1.32 ⫾0.00
SE values ⬍0.005 are listed as 0.00. Number of specimens measured for each instar: Þrst, 3; second, 5; third, 3; fourth, 2; Þfth, 2.
a
Measurements of body, head, and notal lengths taken at midline.
b
Prothoracic legs represented only by femora on one specimen.
1058 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 6
ing length of occlusal surface, profemoral spines em-
bracing anterodorsal margin of tibia and tarsus when
appressed; tarsus one-segmented. Pair of short pretar-
sal claws with pronounced basal tooth.
Meso- and metathoracic legs generally yellowish.
All segments of metathoracic leg longer than corre-
sponding segments of mesothoracic leg. Mesocoxa
conical, ⬇1.4 times the length of trochanter and 0.7
times the length of femur. Meso- and metacoxal mesal
margins each with sparse row of stout spines. Mesofe-
mur with two rows of spines along basal two-thirds of
posterior margin, one row on ventral surface, one row
of about eight spines in basal half of anterior margin.
Mesotibia with four rows of stout spines (three lateral,
Fig. 1. Nymphs of H. indicus. (A) First instar. (B) Second instar. (C) Third instar. (D) Fourth instar. (E) Fifth instar.
(Online Þgure in color.)
November 2014 SUNDAR ET AL.: IMMATURE STAGES OF LACCOCORINAE FROM INDIA 1059
one medial). Meso- and metatarsi one-segmented.
Meso- and metapretarsal claws paired, equal, basal
tooth lacking, 0.5 and 0.3 times the length of respective
tarsus. Metacoxa and trochanter resembling those of
mesothoracic leg in shape and size. Metafemur with
four rows of spines extending entire length: two pos-
terior, one posteroventral, one anteroventral. Meta-
tarsus with two rows of short spines. Natatorial hairs
sparse on mesotibia and tarsus, abundant on metatibia
and tarsus.
Abdomen yellowish brown, pale in anterior two-thirds
of lateral margin of each segment giving connexivum
checkered appearance; paired transversely elongate se-
ries of midlateral punctures on terga IIIÐVII; spines aris-
ing from tubercles lacking; paired ostioles of scent glands
near midline of posterior margin of tergum III, con-
nected by arcuate groove, not set on separate sclerite.
Lateral margins of IIIÐVIII with row of short spines,
posterolateral margins of IIIÐVI with single elongate
spine and sparse elongate hairs. Abdominal apex entire,
without cleft. Ventrally with middle one-third convex,
covered with long setae; spiracles one-third to half the
distance from lateral margin to midline on segments
IIÐVIII.
Second Instar (Fig. 1B). Length, 3.29 ⫾0.05 mm;
width, 2.08 mm (n⫽3). Greatest body width generally
at metanotum. General body coloration yellowish
brown, no longer immaculate. Head, nota, and ab-
dominal terga with scattered brown punctures; brown
coloration associated with punctation conßuent in ar-
eas on vertex and near middle of each notum. Nota
with punctation absent near lateral margins. Mesono-
tum with posterior margin convex at midline. Mesono-
tal wing pads more pronounced. Metanotum with dark
transverse band along posterior margin, especially
dark at midline. Metanotum longest of nota at midline,
1.3 times the length of pronotum, 1.5 times the length
of mesonotum. Checkered appearance of connexiva
progressively less evident in each instar.
Third Instar (Fig. 1C). Length, 4.50 ⫾0.06 mm;
width, 2.78 ⫾0.06 mm (n⫽2). Metanotum longest of
nota at midline; pro- and mesonotum subequal, each
1.3 times the length of metanotum. Lateral margin of
mesonotum, including wing pad, subequal in length to
exposed part of lateral margin of metanotum. Postero-
lateral margins of abdominal segments IIIÐVII with
group of 3Ð4 long spines and elongate hairs.
Fourth Instar (Fig. 1D). Length, 5.68 mm; width,
3.64 mm (n⫽1). Metanotal transverse band consis-
tently dark across width. Metanotum longest of nota at
midline, 1.3 times the length of pronotum, 1.2 times the
length of mesonotum. Lateral margin of mesonotum,
including wing pad, 1.6 times the length of exposed
part of lateral margin of metanotum.
Fifth Instar (Fig. 1E). Length, 7.73 ⫾0.17 mm; width,
4.99 ⫾0.03 mm (n⫽6). Meso- and metanota subequal
in length at midline, each 1.1 times the length of prono-
tum. Lateral margin of mesonotum, including wing pad,
completely overlapping lateral margins of metanotum
and abdominal terga I, II, and 20Ð30% of III.
Notes. This species was known only from the Indian
states of Andhra Pradesh, Maharashtra (Thirumalai
2007), and Tamil Nadu (as Madras; Distant 1906).
Brief diagnostic information including comparative
notes to distinguish adults from those of Heleocoris
elongatus Montandon were provided by Distant
(1906). Recently, the adult H. indicus was redescribed
by Ismail et al. (2009).
Heleocoris vicinus Montandon
Fig. 2AÐE
Heleocoris vicinus Montandon: in Montandon 1910:
655Ð656.
Descriptions of Immature Stages. First Instar (Fig.
2A). Length, 2.88 ⫾0.04 mm; width, 1.72 ⫾0.02 mm
(n⫽2). Body elongate-oval, general appearance dor-
soventrally ßattened, greatest width at meso- and met-
anota; overall convex dorsally, concave ventrally; im-
maculate yellowish dorsally and ventrally; dorsally
with pale margins.
Head transverse, anterior margin convex, continu-
ous with lateral margins of pronotum, posterior margin
convex, immaculate yellow dorsally and ventrally.
Compound eyes dark red, ßattened, convergent an-
teriorly. Synthlipsis ⬇2.2 times the width of one eye.
Lateral hyperoche slightly widening anteriorly. An-
tenna light yellow, apparently four-segmented, basal
segment recessed within head. Labrum originates cau-
dad from anterior margin of head. Labium yellowish
white, elongate-conical, with three visible segments,
overall length ⬇0.9 times the width at base, segment
1 mostly within head, segment 2 subequal in length
to 3.
Nota yellowish brown with pale lateral margins.
Spines arising from tubercles lacking. Pronotum with
anterior margin concave; lateral margins evenly
rounded; posterior margin straight medially, slightly pro-
duced caudad laterally; overlapping ⬇10Ð20% of me-
sonotum. Mesonotum with posterior margin straight
medially, with slight indication of wing pads postero-
laterally. Metanotum longest of nota at midline, 1.5
times the length of pronotum, 1.8 times the length of
mesonotum. Metanotal wing pads not evident. Met-
anotum completely overlaps abdominal tergum I and
⬇50% of tergum II at midline.
Prothoracic leg retentorial, generally milky white
with yellowish brown tibia, tarsus, and pretarsal claws.
Procoxa ⬇1.5 times the length of trochanter, ⬇0.5
times the length of femur. Profemur compressed, two
dense brushes of setae along anterior margin above
and below row of stout spines. Protibia and tarsus
with sulcus extending length of occlusal surface, fem-
oral spines embracing anterodorsal margin of tibia and
tarsus when appressed; tarsus one-segmented. Pair of
short pretarsal claws with pronounced basal tooth.
Meso- and metathoracic legs generally yellowish, all
segments of metathoracic leg longer than correspond-
ing segments of mesothoracic leg. Mesocoxa conical,
⬇2 times the length of trochanter, half the length of
femur. Meso- and metacoxal mesal margins each with
sparse row of stout spines. Mesofemur with two rows
of spines along basal two-thirds of posterior margin,
1060 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 6
one row on ventral surface, one row of about eight
pegs in basal half of anterior margin. Mesotibia with
four rows of stout spines (three lateral, one medial).
Meso- and metatarsi one-segmented. Meso- and
metapretarsal claws paired, equal, basal tooth lacking,
0.5 and 0.3 times the length of respective tarsus. Meta-
coxa and trochanter resembling those of mesothoracic
leg in shape and size. Metafemur with four rows of
spines extending entire length: two posterior, one pos-
teroventral, one anteroventral. Metatarsus with two
rows of short spines. Natatorial hairs sparse on me-
sotibia and tarsus, abundant on metatibia and tarsus.
Fig. 2. Nymphs of H. vicinus. (A) First instar. (B) Second instar. (C) Third instar. (D) Fourth instar. (E) Fifth instar.
(Online Þgure in color.)
November 2014 SUNDAR ET AL.: IMMATURE STAGES OF LACCOCORINAE FROM INDIA 1061
Abdomen yellowish brown, pale in anterior two-
thirds of lateral margin of each segment giving con-
nexivum checkered appearance; paired transversely
elongate series of midlateral punctures on IIIÐVIII;
spines arising from tubercles lacking; paired ostioles of
scent glands near midline at posterior margin of III,
connected by arcuate groove, not set on separate scle-
rite. Lateral margins of IIIÐVIII with row of short
spines, posterolateral margins of IIIÐVI with single
elongate spine and sparse elongate hairs. Abdominal
apex entire, without cleft. Ventrally with middle one-
third convex, covered with long setae; spiracles one
third to half distance from lateral margin to midline on
segments IIÐVIII.
Second Instar (Fig. 2B). Length, 3.90 mm; width, 2.32
mm (n⫽1). Greatest body width at metanotum. General
body coloration yellowish brown, no longer immaculate.
Head, nota, and abdominal terga with scattered brown
punctures; brown coloration associated with punctation
becoming conßuent in areas on vertex and near middle
of each notum. Nota with punctation absent near lateral
margins. Mesonotum with posterior margin convex at
midline. Mesonotal wing pads progressively more pro-
nounced in each instar. Metanotum with dark transverse
band across posterior margin, especially dark at midline.
Metanotum longest of nota at midline, 1.3 times the
length of pronotum, 1.5 times the length of mesonotum.
Checkered appearance of connexiva progressively less
evident in each instar.
Third Instar (Fig. 2C). Length, 4.80 ⫾0.12 mm;
width, 2.91 ⫾0.05 mm (n⫽3). Metanotum longest of
nota at midline; pro- and mesonota subequal, each 0.8
times the length of metanotum. Lateral margin of
mesonotum, including wing pad, 1.1 times the length
of exposed part of lateral margin of metanotum. Pos-
terolateral margins of abdominal segments IIIÐVII
with group of 3Ð4 long spines and elongate hairs.
Fourth Instar (Fig. 2D). Length, 6.00 ⫾0.08 mm;
width, 3.71 ⫾0.03 mm (n⫽5). Metanotum longest of
nota at midline, 1.2 times the length of pronotum, 1.1
times the length of mesonotum. Lateral margin of
mesonotum, including wing pad, twice the length of
exposed part of lateral margin of metanotum.
Fifth Instar (Fig. 2E). Length, 7.79 ⫾0.12 mm; width,
5.12 ⫾0.08 mm (n⫽3). Mesonotum longest of nota at
midline, 1.1 ⫻length each of pro- and metanota. Lateral
margin of mesonotum, including wing pad, completely
overlapping lateral margins of metanotum and often ab-
dominal terga I, II, and 20Ð30% of III.
Notes. This species was known only from Assam,
Bihar, Kerala, Maharashtra, Uttar Pradesh, and West
Bengal states in India (Distant 1910, Thirumalai 2007).
These are the Þrst records of H. vicinus from Karnataka
and Tamil Nadu states. Distant (1910) provided a
supplemental description of the adult.
Pogonocaudina indica Sites & Zettel
Fig. 3AÐE
Pogonocaudina indica Sites & Zettel: in Sites et al. 2011:
6Ð8.
Descriptions of Immature Stages. First Instar (Fig.
3A). Length, 2.77 ⫾0.04 mm; width, 1.80 ⫾0.02 mm
(n⫽3). Body elongate-oval, general appearance dor-
soventrally ßattened, greatest width at pronotum to
metanotum; slightly convex dorsally and ventromedi-
ally, slightly concave ventrolaterally; uniformly brown
dorsally with pale margins, yellowish brown ventrally.
Head transverse, anterior margin convex, continu-
ous with lateral margins of pronotum, posterior margin
more or less straight medially, brown dorsally, yellow-
ish brown ventrally, strongly curved longitudinally,
strongly declivent anteriorly. Compound eye dark red,
bulging. Synthlipsis ⬇2.6 times the width of one eye.
Lateral hyperoche slightly widening anteriorly. An-
tenna whitish, apparently four-segmented, basal seg-
ment recessed within head. Labrum originates caudad
from anterior margin of head. Labium yellowish white,
elongate-conical, with three visible segments, overall
length beyond labrum subequal to width at base; seg-
ment 1 mostly within head, segment 2 subequal in
length to 3.
Nota brown in middle three-fourths with pale lateral
margins. Scattered stout spines arising from tubercles in
middle three-fourths. Pronotum with anterior margin
slightly concave between eyes; posterior margin
nearly straight, slightly produced caudad laterally;
pronotum overlaps ⬇20% of mesonotum. Mesonotum
with posterior margin slightly convex medially, with
slight indication of wing pads posterolaterally. Pro-
and metanota longest of nota, equal in length at mid-
line, each 1.1 times the length of mesonotum. Met-
anotal wing pads not evident. Metanotum completely
overlaps abdominal tergum I and ⬇75% of tergum II at
midline. Row of strong spines along lateral margins of
meso- and metanota, spines continuing along abdom-
inal terga IIIÐV. Mesosternum with strong median
tubercle near posterior margin.
Prothoracic leg retentorial, generally milky white
with yellowish brown tibia, tarsus, and pretarsal claws.
Procoxa ⬇1.5 times the length of trochanter, and ⬇0.6
times the length of femur. Profemur compressed,
brush of stout setae along anterior margin. Tarsus
one-segmented. Pair of short, widely divergent, artic-
ulated pretarsal claws with pronounced basal tooth.
Meso- and metathoracic legs milky white basally;
meso- and metatibiae and -tarsi yellowish brown,
darkening distally. All segments of metathoracic leg
longer than corresponding segments of mesothoracic
leg. Mesocoxa conical, about twice the length of tro-
chanter and 0.6 times the length of femur. Meso- and
metacoxal mesal margins each with sparse row of stout
spines. Meso- and metafemora with two rows of sparse
pegs along posterior margin, one row along anterior
margin. Mesotibia with four rows of stout spines (two
ventral, one lateral, one medial). Meso- and metatarsi
one-segmented, several stout spines ventrally. Meso-
and metapretarsal claws paired, large, widely diver-
gent, strongly curved, with basal tooth, 0.5 and 0.4
times the length of respective tarsus. Metacoxa and
trochanter resembling those of mesothoracic leg in
shape and size. Natatorial hairs sparse on mesotibia
and tarsus, abundant on metatibia and tarsus.
1062 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 6
Abdomen mostly dark brown with yellowish brown
anterolateral corners giving connexivum checkered
appearance. Paired ostioles of scent glands between
segments IIIÐIV, nearly contiguous, connected by
short groove, not set on separate sclerite. Stout spines
arising from tubercles on anterior abdominal terga:
6 Ð8 on III, 4 Ð6 on IV. Ventrally with middle one-third
convex and covered with long setae; spiracles one-
third distance from lateral margin to midline on seg-
ments IIÐVIII. Deep terminal cleft at midline, rim of
cleft reßexed dorsally. Fringe of long hairs on perim-
eter of segments VÐVIII.
Second Instar (Fig. 3B). Length, 3.74 ⫾0.05 mm;
width, 2.52 ⫾0.05 mm (n⫽5). Brown coloration no
longer uniform, now interspersed with yellow, dense
brown punctation throughout, loss of scattered stout
spines arising from tubercles. Metanotum with brown
band along posterior margin. Lateral hyperoche with
Fig. 3. Nymphs of P. indica. (A) First instar. (B) Second instar. (C) Third instar. (D) Fourth instar. (E) Fifth instar.
Second instar was unnaturally deßexed anteroposteriorly. (Online Þgure in color.)
November 2014 SUNDAR ET AL.: IMMATURE STAGES OF LACCOCORINAE FROM INDIA 1063
lateral margin clearly divergent. Meso- and metanota
subequal in length at midline, each 0.9 times the length
of pronotum. Mesonotal wing pads progressively more
pronounced in each instar. Ostioles of scent glands
connected by short groove set in small, dark brown,
oval plate. Length of lateral spines shorter, progres-
sively decreasing in each instar. Mesosternal tubercle
with scattered stout spines. Abdomen with paired,
transversely elongate series of midlateral punctures on
IIIÐVIII; terga with anterior margins lined with dark
brown. Fringe of hairs on perimeter of posterior ab-
dominal segments denser, progressively increasing in
each instar. Relative depth of terminal cleft reduced,
progressively decreasing in each instar. Checkered
appearance of connexiva progressively less evident in
each instar.
Third Instar (Fig. 3C). Length, 4.97 ⫾0.01 mm;
width, 3.27 ⫾0.04 mm (n⫽3). Pro- and mesonota
subequal in length at midline, each 1.2 times the length
of metanotum. Lateral margin of mesonotum, includ-
ing wing pad, ⬇1.4 times the length of exposed part of
lateral margin of metanotum.
Fourth Instar (Fig. 3D). Length, 6.72 ⫾0.00 mm;
width, 4.32 ⫾0.16 mm (n⫽2). Brown punctation
more distinct and evenly distributed. Mesonotum lon-
gest of nota, 1.1 times the length of pronotum, 1.3 times
the length of metanotum. Lateral margin of mesono-
tum, including wing pad, ⬇1.8 times the length of
exposed part of lateral margin of metanotum.
Fifth Instar (Fig. 3E). Length, 8.38 ⫾0.10 mm;
width, 5.36 ⫾0.08 mm (n⫽2). Row of spines obsolete
on lateral margins of nota and abdominal segments
IIIÐV. Mesonotum longest of nota at midline, 1.2 times
the length of pronotum, 1.8 times the length of met-
anotum. Mesonotal wing pad completely overlapping
lateral margins of metanotum, abdominal terga I, II,
and 25Ð30% of III.
Notes. This insect was described recently as the
monotypic representative of the genus (Sites et al.
2011). Adults and nymphs are known from the Þlm of
water ßowing over the rock face of Kiliyur Waterfall,
where they often occur in association with fallen
leaves and other debris that adhere to the surface. It
can be found syntopically with Diaphorocoris arunach-
alami Sites & Zettel and Diaphorocoris kiliyur Sites &
Zettel (Sites et al. 2011). Previously, this species was
known only from the type locality. A second locality
is represented by a female specimen misidentiÞed by
John T. Polhemus as Diaphorocoris dubreuili Montan-
don in the Snow Museum, University of Kansas; the
specimen bears the collection data label: S. India,
Paluey Hills, IV-53, P.S. Nathan, Kodaikanal, Silver
cascade, 5,500 feet.
Diagnosis
Nymphs of these species can be determined to be
members of Laccocorinae by the paired, articulated
prothoracic pretarsal claws; these claws are single and
Þxed in known nymphs of the other subfamilies of
Naucoridae. In addition, members of Laccocorinae
have the front of the head folded posteriorly, such that
the rostrum is set back from the anterior margin of the
head, sometimes resulting in an opisthognathous ap-
pearance. Pogonocaudina nymphs are identiÞed read-
ily by the deep terminal cleft and dense fringe of hairs
on the perimeter of abdominal segments VÐVIII. In
addition, the head of Pogonocaudina curves downward
before folding posteriorly, so the front of the head
appears vertical. Members of Heleocoris have no ter-
minal cleft, only scattered long hairs around the perim-
eter of the abdomen, and ßattened heads. Although
adults can be identiÞed, no discrete diagnostic char-
acters are available to distinguish between nymphs of
H. indicus and H. vicinus.
Acknowledgments
We thank K.G. Sivaramakrishnan (Chennai) for his sup-
port, C. Selva Kumar (Sri Paramakalyani Centre for Envi-
ronmental Sciences at Manonmaniam Sundaranar Univer-
sity) for assistance with specimen collections, and S.S.
Mariappan for assistance with preliminary illustrations. We
thank the Department of Science and Technology-Science &
Engineering Research Board (SERB) for Þnancial support
under Fast Track Young Scientist Scheme, File No.SB/FT/
LS-266/2012 dated 02.05.2013. The Enns Entomology Mu-
seum, University of Missouri, provided optical and electronic
graphics resources. S.S. also thanks staff members of the
Department of Ecology and Environmental Sciences, Pondi-
cherry University, for moral and logistical support. J.E.
McPherson (Southern Illinois University) kindly provided
critical reviews of this manuscript.
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Received 2 April 2014; accepted 5 August 2014.
November 2014 SUNDAR ET AL.: IMMATURE STAGES OF LACCOCORINAE FROM INDIA 1065
