Revision of the Neotropical Brown Lacewing Genus

Nomerobius (Neuroptera: Hemerobiidae)

JOHN D. OSWALD
Department of Entomology, Cornell University,
Ithaca, New York 14853

Ann. Entomol. Soc. Am. 83(1): 18-29 (1990)

ABSTRACT The Neotropical hemerobiid genus Nomerobius is revised. Four valid species
are recognized, two of which, cuspidatus Oswald and spinosus Oswald, are described as
new. The valid species Hemerobius signatus Hagen is transferred to Nomerobius. Sym-
pherobius modestus var. connexus Banks is transferred to Nomerobius, but its species identity
is uncertain. Nomerobius annulicornis Navis is removed from Nomerobius and synonymized
with Sympherohus intervenalis Banks. Lectotypes are designated for Megalomus psycho-
doides Blanchard, Sympherobius modestus Banks, and S. modestus var, connexus Banks.
The males and females of each valid species are keyed. The following interspecific relation-
ships are hypothesized: ((psychodoides + cuspidatus) + (signatus + spinosus)).
KEY WORDS Insecta, brown lacewings, taxonomy, Neotropical Region

Materials and Methods

BROWNLACEWIKGS, family Hemerobiidae, are a
cosmopolitan group of approximately 550 species.
'Sdults and larvae of all studied species are pred-
ators of arthropods associated with emergent vege-
tation. Some species have shown potential value as
biological control agents (New 1975, 1986).
The brown lacewings of the Neotropical Region
are poorly known taxonomically and biologically.
Only t ~oi of the 20 genera found in the Neotropics
have been revised in recent years (Megalomus [see
Gonzales-Olazo 19811 and Sotiobiella [see Mon-
serrat & Penny 1983, hlonserrat 19841). hlost of
the 110 hemerobiid species listed by Penny (1978)
from the Americas south of the contiguous United
States have received little or no attention since their
original descriptions.
I revise here the genus Xomerobius, \vhich is
endemic to the southern two-thirds of South Amer-
ica. In addition to its type species, psychodoides
(Blanchard), three species have heretofore been
referred to Nomerobius: Navis (1915) synony-
mized Sympherobius modestus Banks with psy-
chodoides; Sympherobius marmoratus Navis was
transferred to Nonzerobius by Nakahara (1965)and
was later synonymized with psychodoides b?- Pen-
n) B hlonserrat (1985); and ,\:omerobius annuli-
cornis was described by Navis (1929) and was con-
sidered a valid species b!. Penny & hlonserrat (1985).
In this paper, the validity of the first two synon-
lmies is upheld, but Nornerobius annulicornis is
removed from the genus as a synonym of S y m -
pherobius intervenalis Banks. With the addition
of two new species and the transfer of Hemerobius
signatus Hagen to Nornerobius, four valid species
are recognized in the genus: cuspidatus Oswald,
psychodoides, signatus, and spinosus Oswald.
0013-8746/90/0018-0029$02.00/0 C 1990 Entomological Society
Material. This study is based on the examination
of 250 adult specimens of Xomerobius from the
follohi-ing collections: American h'luseum of Nat-
ural History, New York, N.Y. (AhlNH); Cornell
University Insect Collection, Ithaca, N.Y. (CU);
Florida State Collection of Arthropods, Gainesville,
Fla. (FSCA); James B. Johnson private collection
(JOHKSON); hluseum of Comparative Zoology,
Harvard University, Cambridge, blass. (14CZ);
hlusCum Kational d'Histoire Naturelle, Paris
(hlNHN); Norman D. Penny private collection
(PEKNY); National Museum of Natural History,
LVashington, D.C. (USNhl).
Terminalia Preparation. Terminalia were pre-
pared for examination by removal and immersion
of the abdomen in hot or cold KOH until it was
sufficiently softened and cleared to allo\v dissection
and manipulation.
Illustrations. Drawings were prepared bvith a
camera lucida using compound and dissecting mi-
croscopes. Figures of individual male components
(e.g., Fig. 3-7) are illustrated in standard orien-
tations (i.e.,anterior to left and dorsal or right side
up). Lateral views of the gonarcus (e.g., Fig. 9)
have been rotated 35-90' clock\i~iseto facilitate
plate composition; these views also show the para-
gonarcal membrane unnaturally pulled back to il-
lustrate the line of attachment of this membrane
to the gonarcus. In the species treatments, relative
positional descriptors of gonarcus regions (e.g.,
anterodorsal, ventral) refer to the gonarcus as in-
dividually illustrated, with assumed anterior to left
and dorsal up. The relative in situ positions of com-
ponents of the male terminalia are illustrated in
Fig. 1.
January 1990 OSWALD: OF Nomerobius
REVISION
Measurements. Forewing lengths were mea-
sured from the apex of the wing to the base of the
teeula.
u
Terminology. Terms used for sclerites and
membranes of the male terminalia have been
adapted from Tjeder (1961), with additions pro-
posed by Oswald (1988).
Annotations. The follo\ving annotations are used
in the synonymical listings: A, adult; Bib, bibli-
ography; Dst, distribution; FW, forewing; Lst, list,
listed; MT, male terminalia; NC, new combination;
NS, new synonymy; OD, original description; RD,
redescription; Tax, taxonomy; W, wing(s). An as-
terisk (*) following an annotation indicates a figure
(e.g., FW*, fore\ving figure).
Genus -Vomerobius Navas
Nomerobius Nayis, 1915: 130 (type species: Mega-
l o n ~ u psychodoides
s Blanchard in Gay 1851: 127,
by monotypy); Nakahara 1960: 21 (RD, Tax);
Penny & hlonserrat 1985: 894 (Tax).
Diagnosis. Xomerobius is distinguished from
other Neotropical hemerobiid genera by the fol-
lowing combination of forewing venational char-
acters (Fig. 31): radial stem with only 2 oblique
branches, intra-MP area (Fig. 31, right-slanted
hatching) traversed by 2 crossveins, and mediocu-
bital area (Fig. 31, left-slanted hatching) traversed
by 4 crossveins. Forewings of the similar genus
Synzpherobius possess only 1 intra-MP crossvein
and only 3 mediocubital (m-cu) crossveins.
Description. Small hemerobiids, forewing lengths
4.0-6.5 mm. Head. Temporal sutures distinct; epi-
cranial suture absent; frons with a transverse row
of four brown spots which are sometimes fused into
a pair of elongate lateral maculations or a single
transverse brown band; vertex spotted with brown;
maxillary palp 5-segmented, ultimate segment with
an apical subsegment; labial palp 3-segmented, ul-
timate segment with an apical subsegment. Thorax.
Pronotum and mesonotum with brown spotting
medially, brown laterally. Forewing (Fig. 31). Hu-
meral vein recurrent and pectinately branched;
longitudinal veins and costal crossveins with brown
spotting; radial stem with 2 oblique branches; 2
subcostal crossveins, one near origin of first oblique
radial branch, a second (frequently indistinct) sub-
tending distal end of stigma1 region; 2 intra-MP
crossveins (the distal one occasionally absent); 4
mediocubital crossveins (the distal one occasionally
absent); 6 outer gradate crossveins (including distal
intra-hlP and distal mediocubital crossveins); 1cul-
cu2 crossvein; crossveins frequently brown and
margined with brown. Hindwing. As in Fig. 31.
Male Terminalia (Fig. 1). Tergite 9. Postero-
ventral angles produced as narrow processes which
subtend ectoprocts; posterolateral margins each
with a distinct angular cusp; antecosta prominent.
Sternite 9. Attenuate and abruptly upturned dis-
19
tally; acute apex with or without setae; each side
of sternite below attenuation with an elongate mac-
rotrichia borne on a prominently raised base. Ec-
toproct. Bearing a short subapical tubercle termi-
nating in an elongate thickened seta; posterior apex
acute, inwardly curved and terminating in a peg-
like modified seta. Gonarcus. Gonopons narrow,
with an acute medioventral process; intragonarcus
narrow; extragonarcus broad, extrahemigonarcus
joined posteroventrally by a second transverse
sclerotized bridge (here termed the secondary gon-
opons); lateral surface of each hemigonarcus with
an anterodorsal carinate process, around the mar-
gin of which the paragonarcal membrane attaches
(in spinosus, Fig. 24 and 25, carinate processes
much enlarged, forming quadrate lateral plates);
pseudomediuncus and mediuncus present (Fig. 1);
pseudomediuncus continuous with gonarcus at
gonopons, developed as a narrow sclerotized strip
along the longitudinal midline of the paragonarcal
membrane, with or without one or two large distal
teeth; mediuncus composed of a pair of slender,
symmetrical processes which are associated proxi-
mally with the gonopons. Parameres. Paired distal
lobes small, lateral margins prominently toothed
in some species; apophysis proxima narrow and
elongate; apophysis proxima cavity present.
Female Terminalia (Fig. 2). Tergite 8. Lateral
lobes enclosing and extending below eighth spi-
racles, sclerite margin usually pale or indented just
above spiracles. Tergite 9. Expanded ventral lobes
continuous with dorsal arch. Gonapophyses lat-
erales. Semicircular, styli present. Gonapophyses
posteriores. Absent. Subgenitale (e.g., Fig. 33 and
42). Present, apex rounded or slightly emarginate.
Sperrnatheca. A long tubular structure, convoluted
(Fig. 40) or arranged in a series of arched loops
(Fig. 34), attached proximally to anteroventral sur-
face of bursa copulatrix. Bursa copulatrix. A flat-
tened dorsal diverticulum of the vagina bearing a
pair of accessory glands which insert directly above
insertion of spermatheca; middorsal surface of va-
gina near junction with gonapophyses laterales with
a short papilla which serves as the point of attach-
ment for the ducts of two additional glands.
Natural History and Immature Stages. Un-
known. Ward et al. (1977) recorded adult "psy-
chodoides" from mesquite (Prosopis).
Distribution. I have verified distribution from
Argentina, Bolivia, Chile, and Peru between ap-
proximately 10°S and 40"s. Unverified but reliable
records of Nomerobius spp. also exist for P a r i state,
northern Brazil (Penny & Monserrat 1985,897)and
Uruguay (Nakahara 1960, 23), both cited as "psy-
chodoides." However, the restricted species con-
cept of psychodoides used here renders previous
identifications and geographic records of this species
unreliable. Unless otherwise noted, distributional
information given in this work reflects only per-
sonally examined material.
Vertical distribution is from near sea level to at
least 2,800 m.
 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Val. 83, no. 1

Fig. 1 and 2. N.psychodoidtcs. (1) Apex of male abdomen (some\vhat expanded to show components), lateral.
(2) Apex of female abdomen, lateral. Abbreviations: cc, calus cercus; ect, ectoproct; gl, gonapophysis lateralis; gon,
gonarcus; med, mediu~~cus; par, parameres; pmd, pseudomediuncus; pmdt, pseudomediuncus teeth; 7s, 8s, 9s,
sternites 7, 8, 9; sg, subgenitale; sty, stylus; 7t, 8t, St, tergites 7, 8, 9.

Etymology and Gender. T h e etymology of
Yomerobius was not ueiven b~ NavAs and is un-
known. T h e gender is masculine.
Excluded Species. N. annulicornis \\.as de-
scribed by Navis (1929) from material originally
contained in the Hamburg hluseum. T h e type ma-
terial of this species is presumed to have been de-
stroyed in the bombing of that museum during
World W a r I1 (Weidner 1972, 124-125).
There is, however, good reason to suspect that
this species was not a true i\iomerobius. T h e fol-
lowing characters, illustrated in Navis's original
figure of the forewing of annulicornis (see Navis
1929, 320, Fig. 2), strongly support the contention
that annulicornis was actually a Sympherobius.
The forewing of annulicornis is shown to b e short
and broad; all known Nomerobius species have the
forewing narrower a n d more elongate (Fig. 31). In
all Yomerobius specimens that I have examined,
the four crossveins of the outer gradate series in
the radial area are stepped in one direction (Fig.
31); Navis's figure of annulicornis shows these
crossveins to be alternating, as in most Symphero-
&us. A;. annulicornis is shown to possess only one
intra-MP crossvein as in Sympherobius; all No-
merobius species (excluding aberrant individuals)
have two intra-MP crossveins. N. annulicornis is
shown to possess only three mediocubital cross-
veins. as in Sunzvherobius:
- , all Nomerobius species
(excluding aberrant individuals) possess four me-
diocubital crossveins.
The following additional evidence supports the
synonymy of ankulicornis with S , interuezlis. T h e
short broad form of the forewing figured by Navis
is characteristic of 5'. intervenalis. Navis (1929,
820) stated that the forewing membrane of an-
nulicornis is liyaline, a distinctive feature of S.
intervenalis, the forewing of which is entirely hya-
line except for the fuscous margining of some fore-
wing crossveins, Kavis's (1929, 319) description of
the apex of the male abdomen, "cercis superioribus
6 triangularibus, elongatus, interne arcuatis, acu-
minatus, apice fusco," agrees very well with the
form of the ectoproct of S . interuenalis, which is
subtriangular in lateral view a n d bears distally a
single inwardly curved, attenuate process which
terminates in a dark peglike modified seta. Both
S . annulicornis and S. intercenalis were described
from Colombia, where no true Xomerobius species
a r e presently known. T h e northernmost western
South l m e r i c a n specimens of Nomerobius exam-
ined for this stud>-were collected in central Peru.
Based on this evidence, I propose the follo\ving
synonymy:
Sympherobius intervenalis Banks
Sympherobius intervenalis Banks [1915] 1914-
1913: 630 (OD, A) (female holotype examined,
b1CZ).
Nomerobius annulicornis Navis 1929: 319 (OD,
A, FW*) (type material previously in the Ham-
burg Museum, now presumed destroyed). New
syriorlymy.
Key to Adult Male and Female
RTomerobius Species
1. Males (Fig. 1)-sternite 9 prominent, nar-
rowed and abruptly upturned distally; gon-
apophyses laterales absent . . . . . . . . . . . . . 2
January 1990 OSWALD:REVISIONOF Nomerobius
Females (Fig. 2)-subgenitale present; gon-
apophyses laterales present, each bearing a
prominent stylus . . . . . . . . . . . . . . . . . . . . . 5
2. Apex of sternite 9 with several dark, spinelike
macrotrichia (Fig. 4) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . .psychodoides (Blanchard)
hpex of sternite 9 without macrotrichia (Fig.
18 and 27), or with a field of minute peglike
macrotrichia (Fig. 13) . . . . . . . . . . . . . . . . 3
3. hpex of sternite 9 markedly attenuate (Fig.
18 and 27), without minute peglike macro-
trichia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Apex of sternite 9 relatively short and stout,
with a field of minute peglike macrotrichia
(Fig. 13) . . . . . . . cuspidatus Oswald, n. sp.
4. Anteroventral margin of ectoproct with a spi-
nose process terminating in a peglike mod-
ified seta (Fig. 26) . . spinosus Oswald, n. sp.
Anteroventral margin of ectoproct without a
spinose process (Fig. 17) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . .signatus (Hagen), n comb.
5. Posterior margin of sternite 7 produced into
a cusp at its ventral midline (Fig. 33) . . .
. . . . . . . . . . . . . . . . cuspidatus Oswald, n. sp.
Posterior margin of sternite 7 not cuspate
midventrally (Fig. 32) . . . . . . . . . . . . . . . . 6
6. Spermatheca short and convoluted but not
arranged in a series of arched loops (Fig.
40); subgenitale broadl) T-shaped (Fig. 42)
. . . . . . . . . . . . . . . . . . spinosus Oswald, n, sp.
Spermatheca longer and arranged in a series
of arched loops (Fig. 34 and 38); subgeni-
tale not T-shaped (Fig. 35 and 39) . . . . . 7
7. First elongate loop proximal to bursa with
closely parallel arms which are not strongly
arched, spermatheca in this loop distinctly
wider than in more distal loops (Fig. 38)
. . . . . . . . . . . . . . .signatus (Hagen), n. comb.
First elongate loop proximal to bursa with
arms not closely parallel nor much wider
than in more distal loops; distal loops long
and strongly arched, normally encompass-
ing 270 or more degrees (Fig. 34) . . . . . . .
. . . . . . . . . . . . . . . .ps ychodoides (Blanchard)
Nomerobius psychodoides (Blanchard)
(Fig. 1-9, 31, 32, 34, 35)
Megalomus psychodoides Blanchard i n Gay 1851:
127 (OD, A).
Hemerobius psychodoides: Hagen 1888: 566 (RD,
A).
Sympherobius modestus Banks 1910: 158 (OD, A);
Navis 1915: 131 (NS).
Sympherobius marmoratus Navis 1910a: 70 (OD,
A); Kimmins 1929: 188 (RD, A, W*); Penny &
Xlonserrat 1985: 895 (NS).
Sympherobius psychodoides: Navis 1910b: 237
(NC, Dst); Stange 1984: 52, 57, 68, 71 (Dst).
Nomerobius psychodoides: Iiavhs 1915: 131 (NC,
Dst); Navis 1917: 196 (Dst);Navis 1920: 62 (Dst);
21
Navis 1924: 363 (Dst); Kavis 19.2613: 108 (Dst);
Navis 1930: 19 (Dst); Nav6s 1933: 96 (Dst);
Nakahara 1960: 22 (RD, A, MT*, W*, Dst);
Nakahara 1965: 111 (Dst); Stange 1967: 27 (Bib,
Dst); Penny 1978: 32 (Lst); Penny & Monserrat
1983: 895 (RD, A, hlT*, W*).
Nomerobius marmoratus: Nakahara 1965: 111(NC,
Dst); Stange 1967: 27 (Bib, Dst); Penny 1978: 32
(Lst).
Nomerobium [sic] psychodoides: Ward et al. 1977:
61 (Lst, Dst).
Diagnosis. Males. Distinguished by the presence
of several dark spiniform setae at the apex of the
ninth sternite (Fig. 4). Females. Spermatheca very
similar to that of cuspidatus (compare Fig. 34 and
36), but posterior margin of sternite 7 without a
prominent midventral cusp (compare Fig. 32 and
33).
Description. Forewing length 3.99-5.21 mm
(mean, 1.75, n = 10). ,Vale. Sternite 9 (Fig. 4 and
5): attenuate apical region with several (usually 4-
6) spinose, black, apical and subapical setae; an-
teroventral concavity of sternite margin shallow in
dorsal view. Ectoproct (Fig. 3): without a spinose
anteroventral process. Gonarcus (Fig. 8 and 9):
anterodorsal carinate processes small and rounded;
ventral border of extragonarcus without prominent
bulges or cusps. Parameres (Fig. 6 and 7): apices
of distal lobes rounded in dorsal view, lateral mar-
gins without prominent teeth. Female. Sternite 7
(Fig. 32): posterior margin without a prominent
midventral cusp. Subgenitale (Fig. 35): small, apex
rounded or slightly emarginate. Spermatheca (Fig.
34): arranged in a series of long arched loops; loops
regular, parallel and normally arched 270 or more
degrees; first elongate loop proximal to bursa with
arms strongly arched, not closely parallel nor much
wider than in more distal loops.
Remarks. The species concept of psychodoides
used here is significantly restricted relative to ear-
lier authors, hence previous identifications and dis-
tributional records for this species must now be
considered unreliable. For the sake of complete-
ness, the synonymical listing presented for psycho-
doides contains earlier records of this species pub-
lished under its previous species concepts (excluding
verified misidentifications). Some of these records
may actually apply to one or more of the succeed-
ing species.
Etymology. Unexplained but likely derived from
the Greek "psyche," butterfly, and "-oides," having
the form of.
Distribution. Recorded here from Argentina,
Chile, and Peru. Reports of this species from Serra
Norte, southern Par; state, Brazil (Penny & Mon-
serrat 1985, 897) and Uruguay (Nakahara 1960,
23) have not been confirmed.
Type Material. Megalomus psychodoides. Fe-
male lectotype (MNHN), by present designation.
Verbatim label data: "Museum Paris / Chili / Gay
13-43" [tan rectangular label], "15 / 43" [pale (re-
22 ANNALSOF THE ENTOMOLOGICAL
SOCIETYOF AMERICA V O ~83.
. no. 1

Fig. 3-16. (3-9) N.psychodoides, male terminalia. (3) Tergite 9 and ectoproct, lateral. (4) Sternite 9, lateral.
(5) Sternite 9, dorsal. (6) Parameres, dorsal. (7) Parameres, lateral. (8) Gonarcus and mediuncus, dorsal (pseudo-
mediuncus and membranes omitted). (9) Gonarcus, mediuncus and pseudomediuncus, lateral. (10-16) N . cuspidatus
(holotype), male terminalia. (10) Gonarcus and mediuncus, dorsal (pseudomediuncus and membranes omitted). (11)
Gonarcus, mediuncus and pseudomediuncus, lateral. (12) Tergite 9 and ectoproct, lateral. (13) Sternite 9, lateral.
(14) Sternite 9, dorsal. (15) Parameres, dorsal. (16) Parameres, lateral.

verse green) circular label], "Sympherobius / psy-
chodoides / Blanch." [white, ruled, rectangular la-
bel with clipped corners, written in an unknown
handwriting (h'avhs?)], "Lectotype j Megalomus /
psychodoides / Blanchard in Gay, 1851 / Det. J.
D. Oswald 1988" [handwritten red rectangular la-
bel]. Condition: Good, several wing fragments, ter-
minal antenna1 segments, and distal segments of
some legs missing, otherwise complete. Glued flat
on a transparent (mica?) card. Terminalia cleared,
stained with chlorazol black and placed in a glyc-
erin-filled microvial pinned below specimen.
The labels associated with this specimen indicate
that it was accessioned in the MNHN as part of
Gay's collection of Chilean insects, the h'europtera
of which were described by Blanchard in 1851.
The specimen fits Blanchard's brief original de-
scription of Megalomus psychodoides and is the
only specimen of Nonzerobius in the MIVHN dating
from the mid-1800s (J. Legrand, personal com-
munication). Based on this information, it seems
certain that this specimen was a member of the
type series of M . psychodoides, and I here desig-
nate it as the lectotype of psychodoides. This spec-
imen is apparently the same as that cited by Penny
& hlonserrat (1985, 897) as "the possible type" of
psychodoides. The precise collection locality of the
lectotype within Chile is unknown. One Chilean
locality, "cordilleras de Elequi," was specifically
mentioned in Blanchard's original description, but
no such locality label is present on the pin of the
lectotype.
Sympherobius marmoratus. Male lectotype
( M N H N ) , designated by Kimmins (1929, 189).
January 1990 OSWALD:REVISIONOF Nomerobius
Verbatim label data: "Type," "hluseum Paris /
Chaco d e Santiago del Estero / Bords du Rio
Salado / env. d'Icafio / E.-R. Wagner 1904,"
"Sympherobius / marmoratus / Nav.," "Symphero-
bius / marmoratus / 8 Navhs / det. D. E. Kimmins.
/ 9-1-1928, / Type.," "abdomen in / glicerine /
V. J. Monserrat 84," "Nomerobius / psychodoides
/ Det JDOswald." Condition: Excellent, minuten
pin mounted, part of left antenna missing, other-
wise complete. Terminalia cleared and placed in
a glycerin-filled microvial pinned below specimen.
This is the only specimen cited by Navis in the
original description of marmoratus and may be
the sole specimen upon which marmoratus was
based. However, because Navis (1910a) gave no
definite holotype designation nor any explicit in-
dication of the number of specimens in the type
series, I here accept Kimmins's (1929, 189) "type"
citation as a lectotype designation.
Sympherohius m o d e s t u s . F e m a l e lectotype
(?.lCZ),by present desig~lation.Verbatim label data:
"type," "htlendoza / Argentina," "Type / 11931,"
"Sympherobius / modestus / type Bks," "Lecto-
type / Sympherobius / modestus / Banks, 1910 /
J. D. Oswald 1987." Condition: Excellent, pinned,
tips of antennae missing, otherwise complete. Ab-
domen cleared and placed in a glycerin-filled mi-
crovial pinned below specimen.
This specimen bears type labels in Banks's hand-
writing and is almost certainly the specimen con-
sidered by him to be the holotype of modestus;
however, Banks (1910) gave no definite holotype
designation nor any explicit indication of the num-
ber of specimens in the type series. Rather than
assume this specimen to be the unique holotype, I
here designate it, in accordance with Recommen-
dation 73F of the Code, as a lectotype.
Other Material Examined (155 specimens). AR-
GENTINA: Catamarca Prov.: 388, Los Nacimen-
tos, Punta Balasto, 28-XI-1983, Peiia (PENSY); 18,
l a , Bunta Balasto, 26-XI-1983, Peiia (PENNY). La
Rioja Prov.: 2288, 2599. Guandacol (and Guadacol
[sic]), 1,000 m, 1-3-XII-1983, Peiia (PENNY).
Xlendoza Prov.: 18, 499, San Rafael, 7-XII-1983,
Peiia (PENNY); 288, 399, Uspallata, 1,600 m,
3-6-XII-1983, Peiia (PENNY); lP, Va. El Salto, 12
km SW Potrerillos, 20-XII-1973, Flint (USKM).
Neuquen Prov.: 18, Rio Alumine, 9 km N Alumine,
27-11-1978, Flint (USNM); 18, Kio Litran, 28-XII-
1969, 1,500 m, Gentili (FSCA); 19, Cochico (Aroyo
Dumuvo), N Neuquen, 1,500 m, 5-11-1969, Gentili
(FSCA). San Juan Prov.: 18, 299, Iglesia, 31-XII-
1983, Peiia (PEKNY). CHILE: Aconcagua (Acon-
agua [sic]) Reg.: 1188, 1999, El Tartaro, N. Putaen-
do, 4-6-11-198.1, Peiia (PENNY); 19, Rio Blanco,
10-111-1968, Flint & Peiia (USKl4). Atacama Reg.:
1088, 1199. Finca Chanaral, Inca d e Oro, 17-18-
X-1980, Peiia (PENNY); 18, Finca Chanaral, 200
m, 7-8-X-1980, Peiia (PENKY); 399, El Transito,
E. Vallenar, 25-X-1980, Peiia (PENNY). Liberta-
dor General Bernardo O'Higgins Reg.: 19, Bajo Los
Xlaitenes, K W Rancagua, 1.600 m , 1-11-1982, Pefia
23
(USNM). h4aule Reg.: 18, Curico, Rio Teno, 2 5 x 1 -
1981, Peiia (PENNY); 19, Kio Ancoa, 35 km E
Linares, 320 m, 23-1-1978, Flint (CSNh4). Santiago
Reg.: 368, 299, Colina, 111-1980 and 10-111-1980,
Peiia (PENNY); 18, Conchali, XI-1978, Peiia
(PENKY); 18. hlaipa, Rio S. Jose, 1,500 m , 12-11-
1984, Peiia (PENNY); 19, 17 km W Maipu nr.
Rinconada, 530 m, 10-X-1981, Schuh & Platnick
(AhtINH); 388, Quilicura, X-1979 and 16-XII-1979,
Peiia (PENNY); 388, 499, El Alfalfal, 2-11-1968,
Flint & Peiia (USK14); 18, 19, El Portezuelo, 7 km
N Santiago, 500 m , 22-25-X-1981, Davis (USNM).
Region unknown: 19, 25-1'-1927, Kisliuk? (USNM);
299, Curico El Coigo, E of Potrero Grande, 950 m,
6-1-1988, Peiia (JOHSSON). PERU: Arequipa
Dept.: 299, Yura, 28 km N Arequipa, 2,500 m, 23-
24-1-1972, Schuh (AMKH). Lima Dept.: 299,
Xlatucana, 2,374 m (7,788 ft), 14-VI, Parish (hlCZ).
Nomerobius cuspidatus Oswald, new species
(Fig. 10-16, 33, 36, 37)
Nomerobius psychodoides: Navis 1926a: 96 (mis-
identification, female from near Icatio is cuspi-
datus).
Diagnosis. Males. hlost similar to psychodoides
but apical setae of ninth sternite modified into mi-
nute pegs (compare Fig. 4 and 13). Females. Dis-
tinguished from all other species by the prominent
midventral cusp on the posterior margin of sternite
7 (Fig. 33).
Description. Forewing length 4.80-5.66 m m
\ -
irnean. 5.38. n = 10). ,llale. Sternite 9 (Fig. 1 3 and
14): apical region bearing a transverse field of mi-
nute toothlike pegs; anteroventral concavity of ster-
r~itemargin deep and broad in dorsal view. Ecto-
proct (Fig. 12): without a spinose anteroventral
process. Gonarcus (Fig. 10 and 11): anterodorsal
carinate processes rounded, similar to that of psy-
chodoides; ventral border of extragonarcus without
prominent bulges or cusps. Parameres (Fig. 15 and
16):apices of distal lobes broadly rounded in dorsal
vie\\,, lateral margins without large teeth. Female.
Sterrlite 7 (Fig. 33): posterior margin with a prom-
inent midventral cusp. Subgenitale (Fig. 37): small,
rectangular; apex slightly emarginate. Spermathe-
ca (Fig. 36): arranged in a series of long arched
loops; loops regular, parallel, and normally arched
270 or more degrees; very similar to those of psy-
chodoides but loops usually somewhat shorter; first
elongate loop proximal to bursa with arms strongly
arched, not closely parallel or much wider than
more distal loops.
Remarks. Some earlier records of "psycho-
doides" may refer to this species.
Etymology. An adjective in the nominative sin-
gular from the Latin "cuspidatus," pointed, in ref-
erence to the cusp on the midventral posterior mar-
gin of female sternite 7.
Distribution. Recorded here from Argentina,
Bolivia, and Chile.
24 ANNALSOF THE ENTOMOLOGICAL
SOCIETYOF AMERICA
Type Material. hlale holotype (USNM),by pres-
ent designation. Verbatim label data: "ARGEN-
TINA: Bs. As. / Rio Santiago, Palo / Blanco, Berisso
, 19 Dec. 1979 / C . h4. & 0.S. Flint, Jr.," "USNPVI,"
"HOLOTYPE / Nomerobius / cuspidatus Oswald
1 J . D. Oswald 1987." Condition: excellent, tips of
antennae missing, otherwise complete. Terminalia
cleared and placed in a glycerin-filled microvial
pinned below specimen.
Other Material Examined (23 specimens). AR-
GENTINA: Catamarca Prov.: 19, 82 km SW Santa
Maria, Campo Arenal, 2,150 m, 1-1-1982, Schuh &
\lassie (r\hlNH). Santiago de Estero Prov.: 19, Bords
d u Rio Salado, env. d'Icaiio, 1909, Wagner
Jl'lNHN). BOLIVIA: La Paz Prov.: 288, Yungas
hlts., nr. Sorata, 2,800 m, 15-XII-1984, Pefia
(JOHNSON). CHILE: Aconcagua Reg.: 18, 599,
"Yalpo." [=Vaparaiso?], La Cruz, IX-1966, Rojas
(USNM). Atacama Reg.: 19, Finca Chanaral, Inca
d e Oro, 17-18-X-1980, Peiia (PENKY). Santiago
Reg.: 18, 899, Portezuelo, 7 km N Santiago, 22-25-
X-1981, Davis (USNM);288, 19, Quilicura, X-1979
and 16-XII-1979, Peiia (PENNY).
Nomerobius signatus (Hagen),
new combination
(Fig. 17-23, 38, 39)
Hemerobius signatus Hagen 1861: 322; 1866: 420.
Nomen nudum (see Remarks below).
Hemerobius signatus Hagen 1888: 565 (OD, A).
Somerobius psychodoides: Nakahara 1965: 111
(misidentification, male from Bariloche is sig-
natus).
Diagnosis. Males. Similar to spinosus, particu-
larly in the shape of the apex of the ninth sternite,
but without a n anteroventral ectoproct spine (com-
pare Fig. 17 and 261, anterodorsal carinate pro-
cesses of hemigonarcus much smaller (compare Fig.
22 and 24), and each extrahemigonarcus with a
ventral oval convexity (Fig. 23). Females. Distin-
guished by the weakly arched loops of the sper-
matheca (Fig. 38).
Description. Fore\ving length 4.92-6.51 m m
(mean, 5.71, n = 10). Male. Sternite 9 (Fig. 18 and
19): tip of attenuate apical region without setae;
anteroventral concavity of sternite margin shallow
in dorsal view. Ectoproct (Fig. 17): without a spi-
nose anteroventral process. Gonarcus (Fig. 22 and
23): anterodorsal carinate processes rounded but
longer than in psychodoides; ventral border of each
extrahemigonarcus with a prominent oval convex-
ity or bulge but without posteroventral cusps. Par-
ameres (Fig. 20 and 21): apices of distal lobes
rounded in dorsal view, anterolateral margins with
several large teeth. Female. Sternite 7: posterior
margin without a midventral cusp (as in Fig. 32).
Subgenitale (Fig. 39): small, apex rounded. Sper-
matheca (Fig. 38): roughly arranged in a series of
short arched loops, loops irregular and usually not
arched more than 180"; first elongate loop proximal
Val. 83, no. 1
to bursa with arms closely parallel, distinctly wider
than more distal loops and not strongly arched.
Remarks. Some earlier records of "psycho-
doides" may refer to this species.
Hagen (1861, 1866) published the combination
Hemerobius signatus without description, defini-
tion, or indication. These uses of the specific name
signatus are nomina nuda under Article 12 of the
International Code of Zoological Nomenclature
(International Commission on Zoological Nomen-
clature 1985). The name signatus was apparently
first made available in 1888 when Hagen provided
a description for this species.
Etymology. From the Latin "signatus," mark,
likely in reference to the facial spots found in this
genus.
Distribution. Recorded here from Argentina,
Chile, and Peru.
Type Material. Female holotype (MCZ), by
monotypy (Hagen [1888, 5661 states: "the only
specimen is perhaps a female" [emphasis added]).
Verbatim label data: "Chile," "Type / 10458," "H.
signatus / Hag." [first and last labels handwritten,
probably by Hagen]. Condition: Fair, right fore-
wing, pedicels and flagella of antennae and some
leg segments missing. Thorax broken and reglued
to pin between pro- and mesothorax. Terminaiia
cleared and placed in a glycerin-filled microvial
pinned below specimen.
The labels on this specimen conform precisely
to Hagen's brief original (1861) citation of signa-
tus: "signatus! Hemerobius signatus Hagen, Col-
lect. / HAB.Chili." The specimen agrees extremely
well with Hagen's 1888 description and possesses
a "type" label applied by a previous worker (prob-
ably Nathan Banks, not Hagen).
Other Material Examined (62 specimens). AR-
GENTINA: Mendoza Prov.: 19, 4 km SW Potre-
rillos, 18-XII-1973, Flint (USNM). Rio Negro Prov.:
19, El Bolson, 11-11-1962, Kovacs (USNM); 18, Bari-
loche, XI-1926, Shannon (USNM). CHILE: Acon-
cagua Reg.: 588, 599, El Tartaro, N. Putaendo, 4-
6-11-1984, Peiia (PENNY); 299, Rio Blanco, 10-111-
1968, Flint & Peiia (USNM); 18, 19, Huaquen, XII-
1986, Pefia (JOHNSON). Atacama Reg.: 299, Finca
Chanaral, Inca d e Oro, 17-18-X-1980, Pefia (PEN-
KY); 288, 399, El Transito, E . 17allenar,25-X-1980,
Peiia (PENNY); 288, Juntas (Cohiapo), 1,600 m,
2-X-1980, Peiia (PENNY); 29P, S. Huasco, 13-X-
1980, Pefia (PENNY). Bio Bio Reg.: 288, 499, El
Albanico, 1,000 m, 18-111-1984, Pefia (PENNY).
Coquimbo Reg.: 18, 19, 15 km E Choros Bajos, Rio
Los Choros, 300 m, 10-11-XI-1981, Davis (USNM);
19, Tres Cruces, 29-30-IX-1980, Peiia (PENNY);
299, Tongoy, 3-5-11-1984, Peiia (PENNY). Maule
Reg.: 19, Curico, Rio Teno, 25-XI-1981, Peiia
(PENNY). Santiago Reg.: 18, nr. Pta. Yeso, ca. 70
km SE Santiago, 1,250 m , 27-28-X-1981, Davis
(USNM); 288, 299, El Alfalfal, 29-11-1968, Flint &
Peiia (USNM); l P , Pilay, Rio Peuco, ca. 45 km S
Santiago, 800 m, 23-24-XI-1981, Davis (USNM);
888, 3PP, El Portezuelo, 7 km N Santiago, 500 m ,
January 1990

Fig. 17-30. (17-23) S . signatus, male terminalia. (17)Tergite 9 and ectoproct, lateral. (18) Sternite 9, lateral.
(19) Stertlite 9, dorsal. (20) Parameres, dorsal. (21) Parameres, lateral. (22) Gonarcus and mediuncus, dorsal (pseu-
domediuncus and membranes omitted). (23) Gonarcus, mediuncus and pseudomediuncus, lateral. (24-30) N. s p i -
nosus (holotype), male terminalia. (24) Gonarcus and mediuncus, dorsal (pseudomediuncus and membranes omitted).
( 2 3 ) Gonarcus, mediuncus and pseudomediuncus, lateral. (26) Tergite 9 and ectoproct, lateral. (27) Sternite 9,
lateral. (28) Sternite 9, dorsal. (29) Parameres, dorsal. (30) Parameres, lateral.

22-25-X-1981, Davis (USNbl); 19, La Vilama, SE
of hlelipilla, 350 m, 16-XII-1987, Peiia (JOHN-
SON). Kegion unknown: 18, "Chili Central," 1912,
Lucet (;LlKHN) [misidentified by Navis as psy-
chodoides]; 299, Curico El Coigo, E of Potrero
Grande, 950 m, 6-1-1988, Pefia (JOHNSON). PERU:
Arequipa Dept.: 18, Arequipa, 2,700 m, 6-XI-1983,
Peiia (PENNY).
Nomerobius spinosus Oswald, new species
(Fig. 24-30, 40-42)
Diagnosis. Males. Distinguished from all other
species by the anteroventral spine of the ectoproct
(Fig. 26) and the large, quadrate, anterodorsal car-
inate processes of the hemigonarcus (Fig. 23). Fe-
males. Distinguished by the large, T-shaped
subgenitale (Fig. 32) and a spermatheca which is
convoluted but not arranged in a series of arched
loops (Fig. 40).
Description. Forewing: length 4.84-6.53 mm
(mean, 5.63, n = 5 ) ;with an irregular hyaline streak
beginning between the inner and outer gradate
series in the cubital and medial area, running dis-
tally along the radiomedial area before curving
anteriorly beyond the outer radial gradate cross-
veins; hyaline streak strongly contrasting with ad-
jacent dark brown mottled membrane; cells m2
and m-cu4 (Fig. 31) hyaline except for brown mar-
gining of enclosing crossveins. Male. Sternite 9 (Fig.
27 and 28): tip of attenuate apical region without
setae; anteroventral concavity of sternite margin a
deep semicircle in dorsal view. Ectoproct (Fig. 26):
lvith a spinose anteroventral process terminating
in a dark peglike modified seta. Gonarcus (Fig. 24
and 25): anterodorsal carinate processes large and
quadrate; posteroventral margin of extragonarcus
26 ANNALSOF THE SOCIETYOF
ENTOMOLOGICAL AMERICA Val. 83, no. 1

Fig. 31-42. (31) 3'.psychodoides, forewing and hindwing (right hatching: intra-MP area, note 2 crossveins;
left hatching: mediocubital area, note 4 crossveins). (32 and 33) Sternite 7, female, ventral. (32) N. psychodoides.
i33) S , crcspidatu.~.(34-39) Spermatheca and subgenitale, both ventral. (34 and 35) N. psychodoides. (36 and 37)
S . ctcspidatt~s.(38 and 39) S . signatus. (40-42) N.spinosus. (40) Spermatheca, lateral. (41) Subgenitale, lateral.
(42) Subgenitale, central. Abbreviations: m2, second cell of intra-hlP area; m-cu4, fourth cell of mediocubital area.

\vith a prominent cusp on either side of the sec-
ondar) gonoporis. Parameres (Fig. 29 and 30): dis-
tal lobes elongate i11 dorsal view, posterolateral
rnargiri of each \\.ith a rob\ of large teeth. Female.
Sternite 7: posterior margin without a midventral
cusp (as in Fig. 32). Subgenitale (Fig. 31 and 42):
large, broadl?, T-shaped, apex slightly emarginate.
Spermatheca (Fig. 40): short; convoluted but not
arranged in a series of arched loops; with a short
bulge riear bursa.
Remarks. Some earlier records of "psycho-
doides" may refer to this species.
Etymology. .4n atljecti\.e in the ~lomi~lative sin-
gular from the Latin "spinosus," thorny, in refer-
ence to the elongate spine on the anteroventral
margin of the male ectoproct
Distribution. Kno\\ 11 o111\ from Chile
Type ?laterial. 2lale holot) pe ( C L ) , b\ present
designatio~l \ e r b a t ~ mlabel data "El hlanzano /
(Santiago) CHILE / Feb. 9 '31, 1200 hl. / L. E.
Peiia-G.." "Cornell University / Insect Collec-
,
tions," "HOLOTYPE Nomerobius / spinosus Os-
\vald J. D. Oslvald 1987." Condition: excellent,
no parts missing. Terminalia cleared and placed in
a glycerin-filled microvial pinned below specimen.
Other Material Examined ( 4 specimens).
CHILE: Antofagasta Reg.: 19, Antofagasta, San Pe-
dro, 18-XI-1983, Pefia (PENNY). Bio Bio Keg.: 2PP,
El Albanico, 1,000 m, 18-111-1984, Pefia (PENNY).
Santiago Keg.: 19, El Ilanzano, 1,200 m, 9-11-1951,
Peiia (CU).
Nomen Dubium
ATomerobiusconnexus (Banks),
new7 combination
Sympherohus modestus \a1 connexua Banl\s [I9151
1914-1915 630 (OD, A)
January 1990 OSWALD:REVISIONOF Nomerobius
Fig. 43. Cladogram showing hypothesized phylo-
genetic relationships among species of Nomerobius.
Numbers to left of parentheses refer to characters in
Character List. Numbers within parentheses indicate
character state transitions.
Type Material. Female lectotype (MCZ), by
present designation. Verbatim label data: "type,"
"Chosica Peru / 2800 ft 9-1'1,'' "Parish / Coll,"
"Type / 11932," "Sympherobius / modestus / con-
nexus / type Bks," "Lectotype / Sympherobius /
modestus / var. connexus Banks, 1915 / J. D. Os-
wald 1987." Condition: poor, minuten pin-mount-
ed, parts of antennae and legs missing, head and
metathorax disarticulated and placed dry in a mi-
crovial pinned below specimen. Terminalia cleared
and placed in a glycerin-filled microvial pinned
below specimen (spermatheca missing).
This specimen bears type labels in Banks's hand-
writing and is in all probability the specimen con-
sidered by him to be the holotype of S. n~odestus
var, c o ~ ~ n e x zhmvever,
~s; Ranks (1914-1915) gave
no definite holotype designation for this variety nor
any explicit indication of the number of specimens
present in the type series. Rather than assume this
specimen to be the unique holotype, I here des-
ignate it, in accordance with Recommendation 73F
of the Code, as a lectotype.
Remarks. Banks's varietal name connexus is
a\,ailable in the species-group under the rules of
the Code (International Commission on Zoological
Nornenclature 1983); reference to its lectotype,
27
Table 1 . Character-state matrix f o r Nomerobius and
outgroups
CharacteF
Tawon
1 2 3 4 5 6 7 h 8
Sympherohus spp 0 0 0 0 0 0 2 0 C
Neosympherobluscanereus 0 0 0 0 0 0 0 0
Somerohuspsychodozdes 1 1 1 1 1 1 1 0
Vomerohuscuspadatus 1 1 1 1 1 1 1 0
Nomerohus sagnatus 1 2 2 1 1 1 0 1
Nomerohus sp~noszis 1 2 2 1 1 1 0 1
" Characters are defined in the Character List in the text. Matrix
symbols: 0, p1e.ciomorphic state; 1, 2, apomorphic states.
Absence of cuticular pseudomediuncus teeth is assumed to be
plesiomorphic within the Nomerobius genus group. No hemero-
biid genus lacking a pseudomediuncus is known to have teeth
developed in the homologous region of the paragonarcal rnem-
brane.
Absence of prominent lateral teeth on the paramere lobes is
apparently plesiornorphic within Sympherobius, but similar teeth
are developed in some species (Oswald 1988).
designated above, has shown it to be a Nomerobius
species. Hoa~ever,when the abdomen of the female
lectotype was cleared and dissected, the sperma-
theca was inexplicably found to be absent. On the
basis of its noncuspate sternite 7 and simple subgen-
ital, the lectotype is not cuspidatus or spinosus;
with the spermathem missing it has not been pos-
sible to positively associate this specimen w-ith either
psychodoidcs or signatus. Because both of the lat-
ter species are known from Peru, the type locality
of connexus, it seems preferable to treat connexus
as a nomen dubium until additional characters are
available to distinguish among females of signatus
and psychodoides.
Phylogeny
Based on venational similarities of the forewing,
particularly the presence of four radial area outer
gradate crossveins and only two oblique branches
of the radial stem, previous authors (e.g., Navls
1915, Nakahara 1960, Penny & hlonserrat 1985)
have suspected a close phylogenetic relationship
between the genera lVomerobius and Symphero-
bius. Os\\,ald (1988) proposed the designation
"Somerobiz~sgenus group" for the three genera
Somerobius, Sympherobius, and Neosymphero-
bius, the holophyly of this group being supported
by two synapomorphic characters of the male ter-
minalia: presence of a pseudomediuncus and pres-
ence of a spinose process with a terminal peglike
seta positioned ~ e n t r o m e d i a l lon
~ the ectoproct.
Oslvald further hypothesized Somerobius to be the
sister group of the lineage Neosympherobius +
Sympherobius, based on the synapomorphic loss in
the latter two genera of the fore\ving distal radial
crossvein and the mediuncus of the male genitalia.
Based on this work, Sympherobius and Xeosym-
pherobius have been used here as outgroup taxa.
The eight male terminalic characters used in the
present phylogenetic analysis are given in the fol-
28 ANNALSOF THE ENTOMOLOGICAL
SOCIETYOF AMERICA
lo^ ing character list. The matrix of observed char-
acter states is giben in Table 1 .
Character List, Male Terminalia
1. Posterolateral margin of tergite 9: (0) cusp ab-
sent, (1) cusp present.
2. Apex of sternite 9: ( 0 ) not abruptly upturned,
( 1 ) abruptly upturned, short and broad, (2)
abruptly upturned, elongate and attenuate.
3. Setae of apex of sternite 9: (0) length normal,
(1) length reduced, ( 2 ) absent.
4. Elongate lateral setae of sternite 9: ( 0 ) absent,
( 1) resent.
\ ? .
5 . Anterodorsal carinate process of hemigonarcus:
( 0 ) absent, (1)present.
6. Secoridary gonopons of extragonarcus: ( 0 ) ab-
sent, (1) present.
7 . Pseudomediuncus: (0) cuticular teeth absent, ( 1 )
one or two large teeth, (2) a field of small teeth.
8. llargins of paramere lobes: (0) elongate teeth
absent, (1) elongate teeth present.
Discussion
The unique, most parsimonius cladogram de-
rived fro111 the character-state matrix given in Ta-
ble 1 is shown in Fig. 43. This cladogram exhibits
no honioplasy. The holophyly of Nonlerobius is
strong11 supported by six shared derived charac-
ters: posterolateral margins of male tergite 9 with
angular cusps (character 1 , state transition [O-l]),
apex of male sternite 9 abruptl1- upturned (2[0-I]),
apical setae of male sternite 9 reduced in size (3[0-
I]), male sternite 9 with a pair of elongate lateral
setae (4[0-l]), hemigonarcus with anterodorsal car-
inate processes (3[0-11) and extrahemigonarcus
joined b> a secondary gonopons (6[0-11). N. psy-
chodoides and A'. cuspidatus form a clade based
on the presence of one or two large pseudome-
diu~lcalteeth ('i[O-11). N. signatus and A'. spinosus
form a clade united by the attenuate apex of male
sterrlite 9 (2[1-2]), the loss of setae on the apex of
male sternite 9 (3[1-21) and the presence of prom-
inent teeth on the lateral margins of the paramere
lobes (8[0-11).
Acknowledgment
I thank the following persons for loaning material for
this stud): James Carpenter and Scott Shaw (MCZ); Oli-
\ e r S. Flint, Jr. (USNPvl);J. Legrand ( h l S H N ) ; James K.
Liebherr (CU); Randall T. Schuh (AMNH); and Lionel
A . Stange (FSCA). Special thanks are extended to N. D.
Penn! and James B. Johnson for the opportunity to ex-
amine Sonzerobizis in their private collections. I thank
J. K. Liebherr, \I1. L. Brown, and N. D. Penny for re-
vie\ving the manuscript.
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i~lsects.Proc. Entomol. Soc. Wash. 12: 146-160.
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1 9 1 4 - 1 9 1 5 . New neuropteroid insects, native and
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of Vol. 6e.l
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1 8 6 6 . Hemerobidarum synopsis synonymica. Stett.
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1 8 8 8 . Three species of Hen~erobiusfrom Chili [sic].
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Iiimmins, D. E. 1 9 2 9 . Some new and little known
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1909, I1 (Neuroptera, Planipennia, Hemerobiidae).
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[\-I] serie). 14en1. R. Acad. Cienc. Artes Barc. (3)12:
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1 9 1 7 . Algunos insectos neurbpteros de la Argentina.
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Mens. (Acad. Liter. Plata), B. Aires 22(1922): 358-
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1 9 2 9 . Insectos neurbpteros del rnuseo de Hamburgo.
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January 1990 OSWALD:
REVISIONOF Nomerobius
New, T. R. 1975. The biology of Chrysopidae and
Hemerobiidae (Neuroptera), with reference to their
usage as biocontrol agents: a review. Trans. R. Ento-
mol. Soc. Lond. 127: 115-130.
1986. A review of the biology of Neuroptera Plani-
pennia. Neuroptera Int., Suppl. Ser. 1: 1-37.
Oswald, J. D. 1988. '4 revision of the genus Sym-
pherohus Banks (Neuroptera: Hemerobiidae) of
America north of l4exico with a synonymical list of
the world species. J. N.Y. Entomol. Soc. 96: 390-451.
Penny, N. D. 1978. Lista de Xlegaloptera, Neuroptera
e Raphidioptera do MCxico, Amkrica Central, ilhas
Caraibas e America do Sul. hcta Amazonica 7(4) (sup-
plement) (1977):1-61. [Dating: Date stamps on copies
of the number containing this paper in the Cornell
University and Iowa State University libraries suggest
that this work was not issued until 1978.1
Penny, N. D. & V. J. Monserrat. 1985. Neuroptera
of the Amazon Basin. Part 10. Hemerobiidae. Acta
hmazonica 13(1983):879-909. [Dating: Date stamps
on copies of the number containing this paper in the
Cornell University and Iowa State University libraries
suggest that this work was not issued until 1985.1
29
Stange, L. A. 1967. Catlilogo de Neuroptera de Ar-
gentina y Uruguay. Acta Zool. Lilloana 22: 5-87.
1984. Field studies of the insect order Neuroptera in
South America. Natl. Geogr. Res. 17(1976): 51-76.
[Dating: From the bottom of page 17: iv.]
Tj eder, B. 1961. Neuroptera-Planipennia. The lace-
u-ings of southern Africa. 4. Family Hemerobiidae,
pp. 296-408. In R. Hanstrom, P. Brinck & G. Ru-
debec [eds.], South African animal life, vol. 8. Swedish
Natural Science Research Council, Stockholm.
Ward, C. R., C. W. O'Brien, L. B. O'Brien, D. E. Foster
& E. W. Huddleston. 1977. Annotated checklist
of Ne\v World insects associated with Prosopis (mes-
quite). USDA Technical Bulletin 1557.
Weidner, H. 1972. Die entomologischen Sammlun-
gen des Zoologischen Instituts und Zoologischen h4u-
seums der Universitat Hamburg. VIII Teil. Insecta
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Received for publication 23 January 1989; accepted
26 Atarch 1989.
 Bibliography of the Neuropterida
Bibliography of the Neuropterida Reference number (r#):
6546

Reference Citation:
Oswald, J. D. 1990 [1990.??.??]. Revision of the Neotropical brown lacewing genus
Nomerobius (Neuroptera: Hemerobiidae). Annals of the Entomological Society of
America 83:18-29.

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File:
File produced for the Bibliography of the Neuropterida (BotN) component of the Global
Lacewing Digital Library (GLDL) Project, 2006.