GENERA OF ICHNEUTINAE
JAN. 7, 1993
TITLE: A Revision of the Genera of Ichneutinae (Hymenoptera: Braconidae)
BY: Michael J. Sharkey1 and Robert A. Wharton2
RUNNING TITLE: Genera of Ichneutinae
1. Biological Resources Division, Centre for Land and Biological Resources Research, Agriculture Canada, Ottawa,
Ontario, Canada, K1A 0C6.
2. Department of Entomology, Texas A&M University, College Station, Texas 77843, United States of America.
1
GENERA OF ICHNEUTINAE
JAN. 7, 1993
ABSTRACT. The genera of the subfamily Ichneutinae are revised using cladistic analysis. The following new genera
are proposed: Hebichneutes, Helconichia, Masonbeckia, and Muesonia. The following new synonymies are proposed:
Ciliosa Mason and Pulchaukia Mason as junior subjective synonyms of Oligoneurus Szépligeti; Muesebeckia Mason
and Anaprixia Mason as junior subjective synonyms of Paroligoneurus Muesebeck; Ichneutidea Ashmead and
Proteropoides Viereck as junior subjective synonyms of Proterops. The subgenus Proterops is proposed for those
species of Proterops formerly restricted to the genus Proterops auct. and all other species of the genus Proterops, i.e.,
those formerly placed in Ichneutidea, are considered incertae sedis in the genus Proterops. A total of 15 new species are
described viz.: three species of Hebichneutes, three new species of Helconichia, six new species of Masonbeckia, and
three new species of Muesonia. A key to the world genera and a key to all of the newly described species are included.
Cladistic relationships among the world genera and among the newly described species are discussed.
2
GENERA OF ICHNEUTINAE
JAN. 7, 1993
TABLE OF CONTENTS
INTRODUCTION
PHYLOGENETIC CONSIDERATIONS
MONOPHYLY OF THE ICHNEUTINAE
RELATIONSHIPS AMONG ICHNEUTINAE GENERA
Character List
Taxonomic Units
Results
KEY TO THE WORLD GENERA OF ICHNEUTINAE
SUBFAMILY TREATMENT
DIAGNOSIS
DESCRIPTION
DISTRIBUTION
HOSTS
GENERIC TREATMENTS
Hebichneutes new genus
Helconichia new genus
Ichneutes Nees von Esenbeck
Lispixys Mason
Masonbeckia new genus
Muesonia new genus
Oligoneurus Szépligeti
Paroligoneurus Muesebeck
Proterops Wesmael
SPECIES TREATMENTS
Hebichneutes new genus
Key to species
Relationships of the species of Hebichneutes
Hebichneutes bicolor n. sp.
Hebichneutes citrinus n. sp.
Hebichneutes tricolor n. sp.
Hebichneutes wasbaueri n. sp.
Helconichia new genus
Key to species
Relationships of the species of Helconichia
Helconichia areolata n. sp.
Helconichia trichiops n. sp.
Helconichia trichopteryx n. sp.
Masonbeckia new genus
Key to species
Relationships of the species of Helconichia
Masonbeckia brunneigaster n. sp.
Masonbeckia compressicornis n. sp.
Masonbeckia flavofemorata n. sp.
Masonbeckia hansoni n. sp.
Masonbeckia pauxipalpus n. sp.
Masonbeckia townesi n. sp.
Muesonia new genus
Key to species
Relationships of the species of Muesonia
3
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Muesonia sigalphoides n. sp.
Muesonia straminea n. sp.
Muesonia venus n. sp.
REFERENCES
4
GENERA OF ICHNEUTINAE
JAN. 7, 1993
INTRODUCTION
Förster (1862) established the subfamily Ichneutinae for the genera Ichneutes Nees and Proterops Wesmael
and characterized it largely on the basis of the shape of the 2R1 (marginal) cell of the fore wing. Mason (1969)
described the tribe Muesebeckiini for a group of six genera with reduced venation whose relationships to other
braconids are problematic. He assigned this tribe to the Ichneutinae, citing a number of morphological features shared
by Ichneutes and muesebeckiines. Achterberg (1976) used the tribal names Proteropini and Ichneutini to segregate
Proterops from Ichneutes in his key to braconid subfamilies. Although Mason (1969) presented a fairly complete
description and diagnosis of the Muesebeckiini, comparable descriptions for the Proteropini and Ichneutini (sensu
Achterberg 1976, 1984; Quicke and Achterberg 1990; Shaw and Huddleston 1991) have never been assembled.
Several authors have commented on the problem of grouping the Ichneutinae genera together in a single
subfamily (Mason 1969; Achterberg 1976, 1984; Quicke and Achterberg 1990; Wharton et al. 1992). Tobias and
Belokobylskij (1981), Tobias (1986a,b), and Belokobylskij (1989) rejected the putative relationship between the
Ichneutinae s.s and the genera Oligoneurus and Paroligoneurus, (their Muesebeckiini), and placed these with the
Miracinae.
The Ichneutinae is a relatively small subfamily with only 60 valid species, including the 16 described in this
revision (Shenefelt 1973; Tobias and Belokobylskij, 1981; Belokobylskij 1986, 1990; Papp 1987; Austin and Wharton,
1992). Fifteen generic names have been applied to these species, including four introduced in this revision (Nees, 1816;
Wesmael, 1835; Ashmead, 1900; Szépligeti, 1902; Viereck, 1909; Muesebeck, 1931; Mason, 1957, 1969, 1991). The
paucity of described species reflects a Holarctic bias, since there are many undescribed species from the tropics,
especially in the genera Oligoneurus, Paroligoneurus and Proterops.
Abbreviations for specimen depositories are as follows:
AEI: American Entomological Institute, Gainesville, Florida, U.S.A.
CNC: Canadian National Collection, Ottawa, Ontario, Canada.
INBio: Instituto Nacional de Biodiversidad de Costa Rica, 3100 Santo Domingo, Heredia, Costa Rica.
TAMU: Entomology Department, Texas A&M University, College Station, Texas, U.S.A.
PHYLOGENETIC CONSIDERATIONS
MONOPHYLY OF THE ICHNEUTINAE.
Perhaps the most convincing synapomorphy for members of the Ichneutinae is the presence of one or more
spines at the apex of the fore tibia (Fig. 16). Some members of the Agathidinae (most notably members of the Earinini)
have spines on the lateral surface of the fore tibia but these are not restricted to the apex as in the Ichneutinae. A possible
synapomorphy for members of the Ichneutinae is that the ground plan hosts appear to be members of the Symphyta
(Hymenoptera). Somewhat mitigating the persuasiveness of this character is the fact that nothing is known of the
biology of two of the basal lineages, i.e., Helconichia, and Muesonia (Figs 48, 49). Four additional synapomorphies
are: 1) presence of simple tarsal claws, a character with only one instance of homoplasy, i.e., a reversal in Ichneutes; 2)
absence of a sculptured sternaulus; 3) presence of an enlarged subgenual organ in the hind tibia (Figs 1, 14, 15); 4) setae
of the ovipositor sheath restricted to the sheath apex (Fig. 27).
PLACEMENT OF THE ICHNEUTINAE WITHIN THE BRACONIDAE
Quicke and Achterberg (1990) place the Ichneutinae either basal to the non-cyclostomes (their Figs 3-5) or as a
member of the microgastroid complex (their Figs 1, 6). Either placement is suspect due to numerous errors in their
analysis (Wharton et al., 1993).
We hypothesize that the Agathidinae, Sigalphinae (including Pselaphanus) and Ichneutinae form a
monophyletic group. All have subpronopes (Figs 6, 17), which are dorsolateral pits near the dorsum of the pronotum.
Subpronopes do not occur in other braconid subfamilies except Microtypinae. The latter taxon may be the sister group
of the aforementioned subfamilies or distantly related with a convergent appearance of subpronopes. Within the
Agathidinae, Sigalphinae, and Ichneutinae, subpronopes are widespread, occurring in the great majority of the genera.
Subpronopes are missing from most species of Proterops but these have lost almost all trace of sculpture of any kind on
the mesosoma. Subpronopes are also missing from members of Oligoneurus, Paroligoneurus and Lispixys, but the very
5
GENERA OF ICHNEUTINAE
JAN. 7, 1993
reduced size of the pronotum dorsally and their convincing placement as derived ichneutines (Figs. 48, 49) strongly
suggest a secondary loss of the pits.
The Sigalphinae appears most likely to be the sister-group of the Ichneutinae as suggested by the presence of a
short, sharp, ovipositor combined with the presence of subpronopes. The biology of the Sigalphinae is not known,
however, egg-larval parasitism is a possibility. If this were the case, the hypothesis of a sister-group relationship
between the two subfamilies would be strengthened.
Another candidate for the sister group to the Ichneutinae is the clade (Cheloninae + Adeliinae). Members of
this clade are known to be egg-larval parasites. Possibly all five subfamilies (Ichneutinae, Agathidinae, Chaeloninae +
Adeliinae, Sigalphinae) are closely related. The derived position of the last abscissa of Rs of the fore wing (ending well
basad the wing apex) suggests this.
6
GENERA OF ICHNEUTINAE
JAN. 7, 1993
RELATIONSHIPS AMONG ICHNEUTINE GENERA.
Character List
To polarize character states the Sigalphinae and Agathidinae were used as the primary outgroups, a
hypothetical ancestor was also included which is basically a ground plan for the endoparasitic (non-cyclostome) lineage
of the Braconidae. The hypothetical ancestor was necessary to polarize some characters that were ambiguous in the
Agathidinae and Sigalphinae and to polarize character 34 (see character list that follows) which is otherwise
plesiomorphic only in the Agathidinae. For most characters almost any braconid subfamily could be used as an
outgroup with unequivocal effect. Most characters in the following list are polarized with `0' representing the
plesiomorphic state. Due to ambiguity in the outgroups and homoplasy, however, the reader should refer to the data set
of Table 1 and the cladograms of Figures 48 and 49 to get a complete understanding of character polarity decisions.
Notes on a few of the characters follow. Both states of character 15 are found in the Agathidinae however it is coded as
state 0 since this is by far the most widespread state. Likewise, both states of character 34 are found in the Agathidinae
but it is coded as state 0 (ovipositor at least as long as the metasoma) since Sharkey (1992) demonstrated that a short
ovipositor is derived within the Agathidinae, i.e., if one codes the length of the ovipositor character as polarity unknown
(? in the outgroup) using the data set of Sharkey (1992:431) the short ovipositor state always appears as a derived
character and a autapomorphy of the tribe Disophrini (Agathidinae).
HEAD
1
Number of maxillary palpomeres.
0: 6,
1: 5 (one segment lost),
2: 4 (two segments lost).
Ordered transformation series 0
2
Labial palpal segment 3.
0: 0.5x length of segment 4 or longer,
1: less than 0.5x length of segment 4,
2: absent.
3
4
5
6
7
Occipital carina.
0: present,
1: absent.
Mandible.
0: not twisted (Fig. 11),
1: twisted (Fig. 10).
Mandible.
0: basal area not separated by transverse carina (Figs 10, 11),
1: basal area separated by transverse carina (Fig. 12).
Specialized sensillae on flagellomere 1.
0: absent,
1: present (Fig. 4).
Apical
0: without fields of specialized sensillae and without pair of widely separated longitudinal placodes,
1: with fields of specialized sensillae between pair of widely separated longitudinal placodes (Figs 2, 3).
WINGS
8
1M of fore wing.
0: evenly curved (Fig. 30),
1: abruptly curved at mid-length (Fig. 28),
2: abruptly curved anteriorly (Fig. 32).
Unordered transformation series.
9
1Rs vein of fore wing.
7
GENERA OF ICHNEUTINAE
10
11
12
13
14
15
JAN. 7, 1993
0: present (Figs 28),
1: absent (Figs 29, 35).
Venation.
0: complete (Fig 28),
1: reduced (Fig. 29).
Rs of hind wing.
0: pigmented (Figs 41, 42),
1: unpigmented (Figs 38, 47).
R vein of hind wing.
0: not sharply curved posteriad near distal end (Fig. 47),
1: sharply curved posteriad near distal end (Fig.45).
Sensory setae at junction of R and r-m of hind wing.
0: absent (Figs 40, 41),
1: present (Figs 38, 39).
1A vein of hind wing.
0: complete or almost so (Fig. 46),
1: reduced (Figs. 38, 39).
Hamuli.
0: in one cluster (Fig. 45),
1: scattered (Fig. 44).
MESOSOMA
16
Longitudinal carina of propleuron.
0: present (Fig. 6),
1: absent (Fig. 7).
17
Subpronope.
0: present (Figs 6, 17),
1: absent.
18
Epicnemial carina.
0: present (Fig 7),
1: absent (Fig. 6).
19
Sternaulus.
0: sculptured (Fig. 7),
1: unsculptured (Fig. 6) .
20
21
22
23
24
Notaulus.
0: sculptured,
1: smooth.
Posterior scutellar depression.
0: present (Fig. 18),
1: absent (Fig. 19).
Propodeum.
0: with areolae (Fig. 18),
1: lacking areolae (Fig. 19).
Apex of fore tibia.
0: without one or more spines,
1: with one or more spines (Fig. 16).
Hind claw.
0: basal tooth present.
1: basal tooth absent.
8
GENERA OF ICHNEUTINAE
25
JAN. 7, 1993
Subgenual organ of hind tibia.
0: of normal proportions,
1: large (Figs 1, 14, 15).
METASOMA
26
Pair of median longitudinal carinae on tergum 1.
0: present (Figs 20, 22),
1: absent (Figs 21, 23).
27
Unmargined median longitudinal groove on tergum 1.
0: present (Fig. 21),
1: absent (Figs 20, 22, 23)
28
Spiracle on metasomal tergum 7.
0: present,
1: absent.
29
Setose lateral pits on tergum 7 of males.
0: absent,
1: present (Fig. 26).
30
Cercus.
0: palpiform,
1: plate-like.
31
Male tergal glands of metasonal segment 6.
0: absent,
1: present (Fig. 24).
32
Male tergal glands of metasomal segment 7.
0: absent,
1: present (Figs 24, 25).
33
Female hypopygium.
0: patch of apical setae present,
1: patch of apical setae absent.
34
Ovipositor.
0: at least as long as metasoma,
1: shorter than metasoma.
35
Setae of ovipositor sheath.
0: scattered over entire surface,
1: restricted to apex (Fig. 27).
36 Host.
0: Lepidoptera,
1: Hymenoptera, Argidae,
2: Hymenoptera, Tenthredinidae.
Partially ordered transformation series, i.e., 0
since the transformation to Hymenoptera is presumed to have occurred once.
Taxonomic Units
In this treatment of the Ichneutinae four new genera are proposed. The autapomorphies that define each of
these are enumerated in the data matrix (Table 1) and more fully elucidated in the Remarks section following the
description of each genus. There are also several new synonyms in this revision. Since the descriptions of Mason
(1957, 1969), it has become increasingly difficult to arrange the species under the available generic names in what was
formerly the tribe Muesebeckiini. We have retained Oligoneurus, Paroligoneurus and Lispixys but have been unable to
find adequate supporting characters for the remaining nominate genera. Pulchaukia and Ciliosa are treated as junior
synonyms of Oligoneurus, and Muesebeckia and Anaprixia are treated as junior synonyms of Paroligoneurus. It is
9
GENERA OF ICHNEUTINAE
JAN. 7, 1993
further proposed that Ichneutidea and Proteropoides are junior subjective synonyms of Proterops. The rationales for all
synonymies are found in the Remarks section following the respective generic descriptions.
Results
The ie* option of Hennig86 (Farris, 1988), which is designed to execute an exhaustive search for all minimum
length trees, was used. The result was two minimum length trees (Fig. 48, 49). Excluding the seven autapomorphic
characters, there are 27 informative characters in the matrix. The consistency and retention indices of these are 0.66 and
0.79 respectively.
Due to the pectinate nature of the phylogeny (Fig. 48), a tribal classification is not used. For example, if one
wanted to maintain the tribe Muesebeckiini, a separate tribe would be necessary for every other genus in the subfamily.
Based on the phylogeny here proposed (Figs 48, 49) sawflies, probably argids, appear to be the groundplan
hosts for ichneutines. This infers a switch to Tenthredinoidea in the genus Ichneutes and a switch to lepidopterous
leafminers in the ancestral lineage leading to the clade ((Oligoneurus) + (Paroligoneurus + Lispixys)). The switch to
lepidopteran hosts is a reversal and the first reversal to lepidopteran hosts documented in the Braconidae. It is not clear
from a strict parsimony argument whether the switch to Lepidoptera came from an ancestor attacking tenthredinids or
argids however I find the tenthredinids a more likely ancestor since the majority of these hosts are gall-makers of the
genera Pontania and Euura. A host switch from gall-makers to leaf-miner (Lepidoptera) seems simpler than leaf tier
(Argidae) to leaf-miner.
Members of the four basal clades (Figs 48, 49) are restricted to the Neotropical region. The next clade,
Proterops, is found in the Holarctic, Neotropical, and Oriental Regions. The next clade, Ichneutes, is restricted to a
Holarctic distribution. Paroligoneurus is found in the Nearctic, Neotropical, Palaearctic and its sister-group,
Oligoneurus, is cosmopolitan. The fact that the more derived clades are more widely distributed than the basal clades
deserves explanation. The limited distributions of the basal clades may be due to a lack of collecting in some areas,
especially tropical Africa and Southeast Asia, but if the distributional data are not artifacts of collecting then there has
been large scale dispersal, or extinction, or both. There are many plausible scenarios, but since the putative sister-group
of the Ichneutinae, the Sigalphinae, is cosmopolitan (as are most braconid subfamilies and more than a hundred genera)
a Pangean origin of the Ichneutinae is suggested. This infers extinctions in the ranges of most of the basal clades of the
Ichneutinae.
10
GENERA OF ICHNEUTINAE
1
8
JAN. 7, 1993
KEY TO WORLD GENERA OF ICHNEUTINAE
Venation reduced, Rs vein of fore wing not reaching wing margin. (Figs 29, 32, 36)...............................2
Venation complete, or nearly so, Rs vein of fore wing reaching wing margin. (Figs 28, 30)................................
4
2
Vein 1A of hind wing incomplete, not nearly meeting cu-a (Figs 38, 39).....................................................3
2'
Vein 1A of hind wing complete, meeting or nearly meeting cu-a (Fig. 46).........................................Oligoneurus
3
Ventral side of female flagellomeres in apical 3rd quarter of antenna with two widely spaced longitudinal
placodes and with fields of modified sensillae between them (Figs 2, 3); metasomal tergum 1 often raised
medially, always without narrow median longitudinal depression as in Fig. 21 ...... ............................................
Paroligoneurus
3'
Female flagellomeres with evenly spaced longitudinal placodes and without distinct fields of modified
sensillae;
metasomal
tergum
1
with
narrow
median
longitudinal
depression
(Fig.
21)...........................................Lispixys
4
Pigmented portion of hind wing vein Rs absent or represented by short stub (shorter than r-m) on R. (Fig.
47)...Ichneutes
4'
Pigmented portion of hind wing vein Rs ending near wing margin (Figs 40, 44).............................................5
5
Occipital carina present. .........................Helconichia
5'
Occipital carina absent. ....................................6
6
Epicnemial carina present. (cf. Fig.7)............Muesonia
6'
Epicnemial carina absent. (Fig. 6)...........................7
7
Notaulus smooth; hind claw with rounded basal tooth........ .....................................................Proterops
7'
Notaulus sculptured, at least some crenulae anteriorly; hind tarsal claw simple, lacking basal tooth.
....................8
Longitudinal carina on propleuron present; metasomal tergum 1 without pair of median longitudinal carinae (Fig. 23);
without patch of setae at apex of hypopygium. .....Masonbeckia
8'
Longitudinal carina of propleuron absent; metasomal tergum 1 with pair of median longitudinal carinae (Fig.
Hebichneutes
1
1'
SUBFAMILY TREATMENT
Ichneutinae Förster
Ichneutoidae Förster, 1862: 255.
Ichneutides: Marshall, 1885: 10.
Ichneutinae: Cresson, 1887: 61.
Ichneutidae: Marshall, 1888: 66.
Ichneutini: de Gaulle, 1907: 187.
DIAGNOSIS. Apex of fore tibia, and usually middle and hind tibia, with one or more spines laterally (Fig. 16); 1M
vein of fore wing curved (Figs 28-37); ovipositor short, from barely exerted to as
long as metasoma (Fig. 27); labrum not concave; clypeus and labrum not forming a cyclostome depression; fore wing
with last abscissa of Rs, if complete, meeting margin of wing well before wing apex (Figs 28-37); fore wing lacking
second anal cross-vein; hind wing lacking r crossvein; hind wing lacking 2Cu (last abscissa of cubitus).
DESCRIPTION. For characters that are variable within the subfamily, the supposed ground plan is presented in italics.
Head: Occipital carina either present and meeting hypostomal carina well above base of mandible, or, much
more commonly, completely absent; number of maxillary palpomeres 6 or 4; number of labial palpomeres 4 or 3;
placodes (=longitudinal plate sensilla) generally short and either scattered or in multiple ranks; compound eyes with or
without long setae.
Mesosoma: Longitudinal carina of propleuron present (Fig. 6) or, more often, absent (Fig. 7); epicnemial
carina present (Fig. 7) or, more often, absent (Fig. 6); sternaulus unsculptured or rarely sculptured; notaulus varying
from absent to present, with or without sculpture; postscutellar depression present (Fig. 18) or absent (Fig. 19), usually
11
GENERA OF ICHNEUTINAE
JAN. 7, 1993
consisting of two shallow pits when present. Wing venation reduced in Oligoneurus, Lispixys and Paroligoneurus (Figs
29, 32, 36); otherwise with relatively complete venation (Figs 28, 30, 31, 32, 33, 35, 37). For those genera with
relatively complete venation the following characters apply: 1M vein of fore wing gently to abruptly curved (Figs 2837); fore wing with last abscissa of Rs meeting margin of wing well before wing apex; 1Rs of fore wing present (Fig. 28)
or absent (Fig. 35); 2Rs+M vein of fore wing present (Fig. 30) or absent (Fig. 31); m-cu crossvein of fore wing usually
entering first R cell (Fig. 30), rarely interstitial or entering first Rs cell; 2Cu cell closed; first anal cross-vein (a´) present
(Fig. 28) or absent (Fig. 36), second anal crossvein absent. Hind wing lacking 2Cu (last abscissa of cubitus) and without
the following secondary veins m-cu´, r´, 2r-m´, a´. Hind tibia broad, with subgenual organ (Figs 14, 15). Tarsal claws
simple or with a weak basal lobe; apex of fore tibia and usually middle and hind tibia with one or more spines laterally
(Fig. 16).
Metasoma: Metasomal terga usually densely and irregularly setose; ovipositor short, from barely exerted to as
long as metasoma; ovipositor sheath glabrous basally with long setae apically (Fig. 27).
DISTRIBUTION. Worldwide, though only Oligoneurus is truly cosmopolitan.
cladisticly and in terms of the number of species, is in the Neotropical Region.
The greatest known diversity,
HOSTS. Sawflies of the families Tenthredinidae and Argidae, and leafmining Lepidoptera of the taxa Nepticulidae and
Bucculatrix (Lyonetiidae = Bucculatrigidae); Neodiprion sertifer Geoffroy, of the family Diprionidae, has been recorded
as a host of Ichneutes reunitor Nees von Esenbeck only once (Kolubajiv, 1962) and needs verification.
Biological information for the genera Oligoneurus and Paroligoneurus is summarized in Whitfield and Wagner
(1991).
GENERIC TREATMENTS
Hebichneutes NEW GENUS
(Figs 22, 37, 45)
Type species: Hebichneutes citrinus n. sp.
DIAGNOSIS: female hypopygium with dense patch of setae apically, epicnemial carina absent; notaulus crenulate.
DESCRIPTION
Length: 2.6 to 4.6 mm.
Head: occipital carina absent; antenna with 26 to 31 flagellomeres; first flagellomere without patch of basiconic
sensillae; ventral side of female flagellomeres in apical 3rd quarter of antenna without two, widely spaced, longitudinal
placodes and without fields of modified sensillae; maxillary palpi with 6 segments; labial palpi with 4 segments; labial
palpal segment 3 longer than 0.5x segment 4; mandible not twisted for scissor-like function (cf. Fig. 11); base of
mandible depressed but without microsculptured area set off by acute transverse ridge.
Wings (Figs 37, 45): vein 1M of fore wing evenly curved; vein 1Rs of fore wing present; venation complete; R
of hind wing sharply curved posteriad near apex; Rs of hind wing extending near wing apex, much longer than r-m; hind
wing without long sensory setae at junction of veins R and r-m; 3 hamuli in one cluster; hind wing with vein 1A
complete to cu-a.
Mesosoma: propleuron without longitudinal carina; pronotum with dorsolateral pit (subpronope); epicnemial
carina absent; sternaulus smooth; notaulus crenulate and complete, approaching transscutal articulation; posterior
scutellar depression with two pits; propodeum areolate; hind tarsal claw simple, without basal tooth.
Metasoma (Fig. 22): tergum 1 with pair of median longitudinal carinae and lateral longitudinal carinae, median
carinae convergent in basal half, forming deep depressions at conjunctions with lateral carinae; tergum 1 without
unmargined median longitudinal depression; spiracle of metasomal segment 7 absent; tergum 7 of male setose and
without lateral pits (tergal gland openings); male without large medial excavations on terga 6 and 7; cerci palpiform;
female hypopygium with dense patch of setae apically.
12
GENERA OF ICHNEUTINAE
JAN. 7, 1993
DISTRIBUTION: Neotropical, known from Mexico south to Brazil.
ETYMOLOGY: A conjunction of Ichneutes and hebe which is Greek for puberty and pubic hair. The name refers to
the tuft of setae on the female hypopygium. The name is masculine.
Helconichia NEW GENUS
(Figs 31, 40)
Type species: Helconichia trichiops n. sp.
DIAGNOSIS: Occipital carina present; R vein of hind wing sharply curved posteriad near apex.
DESCRIPTION
Length: 2.7 to 3.3 mm.
Head: occipital carina present meeting hypostomal carina well above base of mandible; antenna with 25 to 29
flagellomeres; first flagellomere without patch of basiconic sensillae; ventral side of female flagellomeres in apical 3rd
quarter of antenna without two widely spaced longitudinal placodes and without fields of modified sensillae; maxillary
palpi with 6 segments; labial palpi with 4 segments; third labial palpal segment 0.5x length of segment 4; mandible not
twisted for scissor-like function; base of mandible not depressed and without microsculptured area set off by acute
transverse ridge.
Wings (Figs 31, 40): vein 1M of fore wing evenly curved; vein 1Rs of fore wing present; venation complete; R
of hind wing sharply curved posteriad near apex; Rs of hind wing extending near wing
apex, much longer than r-m; hind wing without long sensory setae at junction of veins R and r-m; 3 or 4 hamuli in one
cluster; hind wing with vein 1A complete to cu-a.
Mesosoma: propleuron with longitudinal carina; pronotum with weak dorsolateral pit (subpronope) (cf. Fig.
17); epicnemial carina present; sternaulus smooth; notaulus crenulate and complete, approaching transscutal articulation;
posterior scutellar depression with two pits; propodeum areolate; hind tarsal claw simple, without basal tooth.
Metasoma: tergum 1 with pair of median longitudinal carinae and lateral longitudinal carinae; tergum 1 without
unmargined median longitudinal depression; spiracle of metasomal segment 7 absent; tergum 7 of male setose and
without lateral pits (tergal gland openings); male without large medial excavations on terga 6 and 7; cercus palpiform;
female hypopygium without dense patch of setae apically.
DISTRIBUTION: South America.
ETYMOLOGY. Helconichia means helcon-like in Greek and refers to the superficial likeness this species has to the
genus Helcon, especially in the presence of a complete occipital carina. The name is feminine.
Ichneutes Nees von Esenbeck
(Figs 1, 7, 16, 20, 26, 35, 47)
Ichneutes Nees von Esenbeck 1816 (1813): 275. Type species Ichneutes reunitor Nees von Esenbeck.
Ichneustes: Szépligeti 1896: 195. Typographical error.
DIAGNOSIS: anterior portion of vein 1M of fore wing curved abruptly apically (Fig. 35); males with pair of pits on
metasomal tergum 7 (Fig. 26).
DESCRIPTION
Length: 3.0 to 8.0 mm.
Head: occipital carina absent; antenna with 23 to 38 flagellomeres; first flagellomere without patch of basiconic
sensillae; ventral side of female flagellomeres in apical 3rd quarter of antenna without two widely spaced longitudinal
placodes and without fields of modified sensillae; maxillary palpi with 5 segments (since no segments are particularly
13
GENERA OF ICHNEUTINAE
JAN. 7, 1993
long one segment is presumed lost rather that two segments fused); labial palpi with 4 segments; labial palpal segment 3
less than 0.5x length of segment 4; mandible not twisted for scissor-like function; base of mandible not depressed and
without sculptured area set off by acute transverse ridge.
Wings (Figs 35, 47): vein 1M of fore wing mostly straight but anterior portion curved abruptly apicad; vein
1Rs of fore wing absent; venation complete; R of hind wing not sharply curved posteriad near apex; Rs of hind wing
unpigmented or indicated only by small stub much shorter than r-m; hind wing without long sensory setae at junction of
veins R and r-m; 3 or rarely 4 hamuli in one cluster; hind wing with vein 1A complete to cu-a.
Mesosoma (Figs 7, 16): propleuron without longitudinal carina; pronotum with or without weak dorsolateral
pit (subpronope); epicnemial carina present; sternaulus crenulate; notaulus crenulate and complete, approaching
transscutal articulation; posterior scutellar depression with numerous small crenulae or rarely with two pits; propodeum
areolate rugose; hind tarsal claw with basal tooth.
Metasoma (Figs 20, 26): tergum 1 with pair of median longitudinal carinae and lateral longitudinal carinae
(Fig. 20) (Often the carinae are weak but the medial region of the tergum is raised giving the appearance of carinae.);
tergum 1 without unmargined median longitudinal depression; spiracle of metasomal segment 7 present; male with
small, setose lateral pits on otherwise glabrous tergum 7 (Fig. 26) (tergal gland openings, Buckingham and Sharkey
1988); male without large medial excavations on terga 6 and 7; cercus plate-like; female hypopygium without dense
patch of setae apically.
HOSTS: Tenthredinid sawflies.
DISTRIBUTION: Holarctic.
Lispixys Mason
(Fig. 21)
Lispixys Mason 1969: 274. Type species, Lispixys avispa Mason
DIAGNOSIS: Flagellomeres of female without fields of basiconic sensillae that sit between longitudinal placodes that
are unusually widely separated; fore wing venation reduced (cf. Figs 29, 36); 1A vein of hind wing not complete to cu-a
crossvein (cf. Figs 38, 39); propodeum areolate; unmargined medial groove on tergum 1.
DESCRIPTION
Length: 1.8 to 3.5 mm.
Head: occipital carina absent; antenna with 20 to 24 flagellomeres; first flagellomere without patch of basiconic
sensillae; ventral side of female flagellomeres in apical 3rd quarter of antenna without two widely spaced longitudinal
placodes and without fields of modified sensillae; maxillary palpi with 5 segments (since no segments are particularly
long one segment is presumed lost rather that two segments fused); labial palpi with 4 segments, third segment very
small; labial palpal segment 3 less than 0.5x length of segment 4; mandible twisted for scissor-like function; base of
mandible not depressed and without microsculptured area set off by acute transverse ridge.
Wings (cf. Figs 29, 36, 38, 39): vein 1M of fore wing mostly straight but anterior portion curved abruptly
apicad; vein 1Rs of fore wing absent; venation reduced; R of hind wing not sharply curved posteriad near apex; Rs of
hind wing unpigmented or indicated only by small stub much shorter than r-m; hind wing with two or more long sensory
setae at junction of veins R and r-m; 3 hamuli in one cluster; hind wing with vein 1A incomplete, extending less than 3/4
length to cu-a.
Mesosoma: propleuron without longitudinal carina; pronotum without weak dorsolateral pit (subpronope);
epicnemial carina absent; sternaulus rarely smooth (2%), when present sculptured and usually crenulate, at least
anteriorly; notaulus crenulate but absent in posterior half; posterior scutellar depression absent; propodeum areolate to
areolate rugose; hind tarsal claw simple, without basal tooth.
Metasoma (Fig. 21): tergum 1 with weak lateral longitudinal carina, but lacking pair of median longitudinal
carinae; tergum 1 with unmargined median longitudinal depression (Fig. 21); spiracle of metasomal segment 7 absent;
14
GENERA OF ICHNEUTINAE
JAN. 7, 1993
tergum 7 of male setose and without lateral pits (tergal gland openings); male without large medial excavations on terga
6 and 7; cercus plate-like; female hypopygium without dense patch of setae apically.
HOSTS: Unknown, but based on its phylogenetic position, lepidopteran leafminers of the families Nepticulidae and/or
Bucculatrigidae are likely hosts.
DISTRIBUTION: Published records are from Peru and from Sinaloa, Mexico. An undescribed species from Arkansas,
United States of America is in the Canadian National Collection.
REMARKS. We do not feel confident that this clade should be recognized as a genus because there is equally
parsimonious evidence suggesting that it may be a derived lineage of Oligoneurus. Members of a group of Oligoneurus
have an unmargined median longitudinal depression on the first metasomal tergum as do members of Lispixys.
Additionally, members of Lispixys superficially resemble members of this clade of Oligoneurus. Despite these
arguments, we choose to agree with Mason (1969) and maintain the equally parsimonious alternative choice of treating
Lispixys as a genus on the basis of a reduction in the anal vein in the hind wing which is a consistent character within the
Ichneutinae.
Masonbeckia NEW GENUS
(Figs 4, 5, 6, 12, 15, 17, 18, 23, 33, 34, 42, 43)
Type species: Masonbeckia compressicornis n.sp.
DIAGNOSIS: base of mandible with depressed area set off by transverse ridge (Fig. 12); patch of basiconic sensillae
present on first flagellomere (Fig. 4).
DESCRIPTION
Length: 2.3 to 4.0 mm.
Head (Figs 4, 5, 12) : occipital carina absent; antenna with 24 to 31 flagellomeres; first flagellomere with patch
of basiconic sensillae on lateral surface (Fig. 4); ventral side of female flagellomeres in apical 3rd quarter of antenna
without two widely spaced longitudinal placodes and without fields of modified sensillae; maxillary palpi with 6 or 4
segments (in the latter case two segments are considered lost since no segments are long enough to suggest fusion);
labial palpi with 4 or 3 segments (segment 3, when present, longer than 0.5x segment 4); thus palpal formula 6/4 or 4/3;
mandible not twisted for scissor-like function; base of mandible with depressed microsculptured area set off by acute
transverse ridge (Fig. 12).
Wings (Figs 33, 34, 42, 43): vein 1M of fore wing evenly curved; vein 1Rs of fore wing present; venation
complete; R of hind wing not sharply curved posteriad near apex; Rs of hind wing extending near wing apex, much
longer than r-m; hind wing without long sensory setae at junction of veins R and r-m; 3 hamuli in one cluster; hind wing
with vein 1A complete to cu-a.
Mesosoma (Figs 6, 15, 17, 18): propleuron with longitudinal carina (Fig. 6); pronotum with dorsolateral pit
(subpronope) (Figs 6, 17); epicnemial carina absent (Fig. 6); sternaulus smooth; notaulus crenulate and complete, almost
meeting transscutal articulation; posterior scutellar depression with two pits (Fig. 18); propodeum areolate (Fig. 18);
hind tarsal claw simple, without basal tooth.
Metasoma (Fig. 23): tergum 1 mostly smooth, lacking median longitudinal carinae and with or without lateral
longitudinal carinae (Fig. 23); tergum 1 without unmargined median longitudinal depression; spiracle of metasomal
segment 7 present or absent; tergum 7 of male setose and without lateral pits (tergal gland openings); male without large
medial excavations on terga 6 and 7; cercus plate-like; female hypopygium without dense patch of setae apically.
DISTRIBUTION: Neotropical, known from Mexico south to Argentina.
REMARKS: This genus is rather remarkable in that some species have a spiracle on metasomal segment 7 and others
do not, the same species that have the reduction in spiracle number also have the maxillary palpi reduced to 4 segments
15
GENERA OF ICHNEUTINAE
JAN. 7, 1993
and the labial palpi reduced to 3 segments. In a preliminary phylogenetic analysis of the ichneutine genera I (MJS)
entered these two groups separately and obtained the same results as presented in the cladogram of Figure 50. The
entries for Masonbeckia in the data set reflect the ground plan based on outgroup analysis.
ETYMOLOGY: This genus is named in the honour and memory of W.R.M Mason and Cf.W. Muesebeck. The name is
feminine.
Muesonia NEW GENUS
(Figs 28, 44)
Type species: Muesonia straminea n. sp.
DIAGNOSIS: Vein 1M of fore wing vein abruptly curved at mid-length (Fig. 28); hamuli scattered and unevenly
spaced (Fig. 44).
DESCRIPTION
Length: 3.9 to 7.5 mm.
Head: occipital carina absent; antenna with 26 to 31 flagellomeres; first flagellomere without patch of basiconic
sensillae; ventral side of female flagellomeres in apical 3rd quarter of antenna without two widely spaced longitudinal
placodes and without fields of modified sensillae; maxillary palpi with 6 segments; labial palpi with 4 segments; labial
palpal segment 3 more than 0.5x length of segment 4; mandible not twisted for scissor-like function; base of mandible
not depressed and without microsculptured area set off by acute transverse ridge.
Wings (Figs 28, 44): vein 1M of fore wing rather abruptly curved at mid-length (Fig. 28); vein 1Rs of fore
wing present; venation complete; R of hind wing not sharply curved posteriad near apex (Fig. 44); Rs of hind wing
extending near wing apex, much longer than r-m; hind wing without long sensory setae at junction of veins R and r-m; 4
to 6 hamuli scattered and unevenly spaced; hind wing with vein 1A complete to cu-a.
Mesosoma: propleuron with longitudinal carina absent or weakly indicated anteriorly; pronotum with weak
dorsolateral pit (subpronope; epicnemial carina present; sternaulus smooth; notaulus crenulate and complete,
approaching transscutal articulation; posterior scutellar depression with two pits; propodeum areolate; hind tarsal claw
simple, without basal tooth.
Metasoma: tergum 1 with pair of median longitudinal carinae and lateral longitudinal carina; tergum 1 without
unmargined median longitudinal depression; spiracle often (60%) at apex of lateral flange; spiracle of metasomal
segment 7 present; tergum 7 of male setose and without lateral pits (tergal gland openings); male without large medial
excavations on terga 6 and 7; cercus palpiform; female hypopygium without dense patch of setae apically.
DISTRIBUTION: Neotropical, Mexico south to Venezuela and Colombia.
ETYMOLOGY: This genus is named in the honour and memory of Cf.W. Muesebeck and W.R.M Mason. The name
is feminine.
Oligoneurus Szépligeti
(Figs 24, 29, 46)
Oligoneurus Szépligeti 1902. Type species: Oligoneurus concolor
Pulchaukia Mason 1969: 277. NEW SYNONYM.
Ciliosa Mason 1969: 277. NEW SYNONYM.
Szépligeti.
DIAGNOSIS: Venation greatly reduced (Figs 29,); 1A of hind wing complete to cu-a crossvein (Fig. 46); males with
large tergal glands on metasomal terga 6 and 7 (Fig. 24).i.e
DESCRIPTION
16
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Length: 1.2 to 5.0 mm.
Head: occipital carina absent; antenna with 12 to 28 flagellomeres; first flagellomere without patch of basiconic
sensillae; ventral side of female flagellomeres in apical 3rd quarter of antenna without two widely spaced longitudinal
placodes and without fields of modified sensillae; maxillary palpi with 5 segments; maxillary palpi with 5 segments
(since no segments are particularly long a segment is presumed lost rather that two segments fused); labial palpi with 4
or rarely 3 segments (third segment sometimes lost); labial palpal segment 3 (when present) less than 0.5x length of
segment 4; most (75%) species with mandible twisted for scissor-like function; base of mandible not depressed and
without microsculptured area set off by acute transverse ridge.
Wings (Figs 29, 36, 46): vein 1M of fore wing mostly straight but anterior portion curved abruptly apicad;
vein 1Rs of fore wing absent; venation reduced; R of hind wing not sharply curved posteriad near apex; Rs of hind wing
unpigmented or indicated only by small stub much shorter than r-m; hind wing with two or more long sensory setae at
junction of veins R and r-m; 3 hamuli in one cluster; hind wing with vein 1A complete to cu-a.
Mesosoma: propleuron without longitudinal carina; pronotum without weak dorsolateral pit (subpronope);
epicnemial carina absent; sternaulus crenulate or smooth; notaulus often absent, if present crenulate or smooth, always
absent in posterior two thirds; posterior scutellar depression absent; propodeum areolate, to areolate rugose, to smooth;
hind tarsal claw simple, without basal tooth.
Metasoma (Fig. 24): tergum 1 with or without lateral longitudinal carinae, usually mostly smooth; tergum 1
without pair of median longitudinal carinae; tergum 1 with or without unmargined median longitudinal depression (cf.
Fig. 21); spiracle of metasomal segment 7 absent, in one apparently monophyletic group spiracles are absent on terga
1,3,5 and 7); tergum 7 of male setose and without lateral pits (tergal gland openings); male with large medial
excavations on terga 6 and 7 (Fig. 24); cercus plate-like; female hypopygium without dense patch of setae apically.
HOSTS: Whitfield and Wagner (1991) report Nepticulidae and Bucculatrigidae (both lepidopteran leafminers) as hosts.
Because their identification of these genera was not based on the criteria we define in his paper it is not certain that they
refer to Oligoneurus or Paroligoneurus. I have checked some of their specimens and verified Bucculatrix sp. on
Artemisia douglasiana as a host of Oligoneurus sp.
DISTRIBUTION: Worldwide.
REMARKS. Oligoneurus and the clade Paroligoneurus + Lispixys are readily identified by the presence and absence,
respectively, of the apical portion of the hind wing anal vein. Oligoneurus is defined by the presence of a median pit on
tergum 6 of males. Although the species of Oligoneurus generally have more setose eyes than those of Paroligoneurus
and Lispixys, we have been unable to define discrete states for this character, and assignment of some species on this
basis alone has been equivocal. The difficulty of defining Paroligoneurus relative to Oligoneurus has been discussed
elsewhere (Tobias and Belokobylskij, 1981; Austin and Wharton, 1992).
The type species of Ciliosa and Pulchaukia differ little from that of Oligoneurus. Each possesses a few derived
features as enumerated by Mason (1969) but there are no characters that define Oligoneurus with the exclusion of these
two clades.
Paroligoneurus Muesebeck
(Figs 2, 3, 13, 25, 27, 32, 36, 38, 39)
Paroligoneurus Muesebeck 1931: 7. Type species: Paroligoneurus
johnsoni Muesebeck .
Muesebeckia Mason 1957: 355. NEW SYNONYM.
Anaprixia Mason: 1991: 95. NEW SYNONYM.
DIAGNOSIS: Middle and sometimes apical flagellomeres of female with fields of modified sensillae that sit between
longitudinal placodes that are unusually widely separated (Figs 2, 3); fore wing venation reduced (Fig 32); 1A vein of
hind wing not complete to cu-a crossvein (Figs 38, 39).
DESCRIPTION
17
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Length: 1.8 to 3.5 mm.
Head (Figs 2, 3, 13): occipital carina absent; antenna with 15 to 25 flagellomeres; first flagellomere without
patch of basiconic sensillae; female with field of modified sensillae ventrally on middle and usually on distal
flagellomeres (Figs 2, 3); maxillary palpi with 5 segments (since no segments are particularly long one segment is
presumed lost rather that two segments fused); labial palpi with 4 or rarely 3 segments (i.e., third segment sometimes
lost); labial palpal segment 3 (when present) less than 0.5x length of segment 4; mandible twisted for scissor-like
function ( cf. Fig. 10) or rarely exodont (Fig. 13); base of mandible not depressed and without microsculptured area set
off by acute transverse ridge.
Wings (Figs 32, 38, 39): vein 1M of fore wing mostly straight but anterior portion curved abruptly apicad (Fig.
32); vein 1Rs of fore wing absent; venation reduced; R of hind wing not sharply curved posteriad near apex; Rs of hind
wing unpigmented or indicated only by small stub much shorter than r-m (Figs 38, 39); hind wing with two or more long
sensory setae at junction of veins R and r-m; 3 hamuli in one cluster; hind wing with vein 1A incomplete, extending less
than 3/4 length to cu-a.
Mesosoma: propleuron without longitudinal carina; pronotum without weak dorsolateral pit (subpronope);
epicnemial carina absent; sternaulus rarely completely smooth (2%), usually crenulate, at least anteriorly; notaulus
usually (90%) present and if so then crenulate but always absent in posterior half or more; posterior scutellar depression
absent; propodeum varying from smooth to rugose, often with median longitudinal carina or rugosities, not areolate;
hind tarsal claw simple, without basal tooth.
Metasoma (Figs 25, 27): tergum 1 with weak lateral longitudinal carina, usually (90%) without pair of median
longitudinal carinae, rarely with weak median longitudinal carina; tergum 1 without unmargined median longitudinal
depression; spiracle of metasomal segment 7 absent; tergum 7 of male setose and without lateral pits (tergal gland
openings); male without large medial excavation on tergum 6, rarely (2%) with large medial excavation on tergum 7
(Fig. 25); cercus plate-like; female hypopygium without dense patch of setae apically.
HOSTS. P. johnsoni has been reported from leafmining species of Nepticula (Lepidoptera: Nepticulidae). (Muesebeck
and Walkley, 1951). Whitfield and Wagner (1991) report Nepticulidae and Bucculatrigidae as hosts. From their
collection I have verified only Stigmella sp. (Nepticulidae) on Crataegus. There are two unidentified specimens of
Paroligoneurus in the Canadian National collection on Nepticula species reared from Ulmus rubra and Betula sp.
(probably papyrifera).
DISTRIBUTION: Nearctic, Palaearctic, Neotropical, there are also some records from border areas between the
Oriental and Palaearctic (e.g., Taiwan and Nepal) but we presume these to be Palaearctic elements.
REMARKS. Autapomorphies for Paroligoneurus are the reduction in hind wing venation and the presence of a field of
modified sensillae on the middle and usually the apical flagellomeres of the female.
The species formerly assigned to Anaprixia (Mason 1991) are quite distinctive, having exodont mandibles and
a reduced number of abdominal spiracles (missing from segments 2,4,6, and 8). They are otherwise inseparable from
Paroligoneurus, some members of which also have a reduced number of abdominal spiracles. Since recognition of
Anaprixia as a genus renders Paroligoneurus paraphyletic Anaprixia is here considered to be a junior synonym of
Paroligoneurus.
Proterops Wesmael
(Figs 8-11, 14, 19, 30, 41)
Proterops Wesmael 1835: 201. Type species: Proterops nigripennis
Ichneutidea Ashmead 1900: 133. NEW SYNONYM.
Proteropoides Viereck 1909: 42. NEW SYNONYM.
Wesmael.
DIAGNOSIS: Maxillary palpi with 4 segments; venation complete, notaulus smoothly impressed.
18
GENERA OF ICHNEUTINAE
JAN. 7, 1993
DESCRIPTION
Length: 2.5 to 9.0 mm.
Head (Figs 8 -11): occipital carina absent; antenna with 25 to 34 flagellomeres; first flagellomere without patch
of basiconic sensillae; ventral side of female flagellomeres in apical 3rd quarter of antenna without two widely spaced
longitudinal placodes and without fields of modified sensillae; maxillary palpi with 4 segments (since no segments are
particularly long two segments are presumed lost rather than three segments fused); labial palpi with 3 segments (third
segment presumably lost); mandible variable, twisted for scissor-like function (Fig. 10) or not (Fig. 11); base of
mandible not depressed and without microsculptured area set off by acute transverse ridge (Figs 10, 11).
Wings (Figs 30, 41): vein 1M of fore wing evenly curved (Fig. 30); vein 1Rs of fore wing present; venation
complete; R of hind wing not sharply curved posteriad near apex (Fig. 41); Rs of hind wing extending to or nearly to
wing apex, much longer than r-m; hind wing without long sensory setae at junction of veins R and r-m; 3-5 hamuli in
one cluster; hind wing with vein 1A complete to cu-a.
Mesosoma (Figs 14, 19): propleuron without longitudinal carina; pronotum usually (90%) without weak
dorsolateral pit (subpronope); epicnemial carina absent; sternaulus smooth; notaulus smooth, lacking crenulae but
usually complete, approaching transscutal articulation, rarely absent; posterior scutellar depression absent (Fig. 19);
propodeum usually smooth (Fig. 19), rarely with two prominent longitudinal carinae; hind tarsal claw simple, without
basal tooth.
Metasoma: tergum 1 usually smooth, rarely with two prominent median longitudinal carinae or two lateral
carinae; tergum 1 without unmargined median longitudinal depression; spiracle of metasomal segment 7 present; tergum
7 of male setose and without lateral pits (tergal gland openings); male without large medial excavations on terga 6 and 7;
cercus palpiform to almost plate-like; female hypopygium without dense patch of setae apically.
DISTRIBUTION: Nearctic, Neotropical, Oriental, Palaearctic.
REMARKS: We propose the subgenus P. Proterops NEW SUBGENUS to include those species that were formerly
placed in Proterops. The remainder of the species that we include in Proterops (those formerly in the genus Ichneutidea)
are now considered incertae sedis in Proterops. It is our aim to retain as many monophyletic groups (that are reasonably
diverse and speciose) as possible in the classification. The monophyly of P. Proterops is supported by the apomorphic
arrangement of the ocelli, such that the median ocellus is closer to the antennal pit than to the lateral ocellus (Fig. 8), the
plesiomorphic state shared by the remainder of Proterops is that the median ocellus is closer to the lateral ocellus than to
the antennal pit (Fig. 9). In the data set used for phylogenetic analysis Proterops is coded as having the mandibles not
twisted (Fig. 11). Those species with twisted mandibles (Fig. 10) constitute a derived lineage with the subgenus P.
Proterops.
SPECIES TREATMENTS AND KEYS TO NEW SPECIES
Hebichneutes NEW GENUS
Key to species
1
Terminal flagellomeres yellow, contrasting sharply with darker basal flagellomeres; body of mesosoma entirely
yellow....... ................................................. H. citrinus.
1'
Terminal flagellomeres brown, concolorous with basal flagellomeres; body of mesosoma partly brown..............
2
2
2'
Middle and hind femora brown. ..................H. tricolor.
Middle and hind femora dull yellow. .......................3
3
3'
Hind tibia entirely brown. ......................H. wasbaueri
Hind tibia not entirely brown. ....................H. bicolor
Relationships of the species of Hebichneutes
19
GENERA OF ICHNEUTINAE
JAN. 7, 1993
There are very few characters on which to base a phylogenetic hypothesis for the group, and these are
continuous or meristic characters that are difficult to code. No grouping is proposed for the three included species.
Hebichneutes bicolor n. sp.
FEMALE DESCRIPTION.
Length: 2.6 to 4.0 mm.
Colour: Dull yellow except as follows: flagellum, scape and pedicel light brown, lateral lobes of mesoscutum,
hind tibia apically and hind tarsi all brown, metasomal terga 4 to 8 dull yellow or brown; all coxae somewhat paler than
mesosoma and femora; wings infuscate.
Structural features: antenna with 26 to 29 flagellomeres; ratio of vein 2Rs:3Rs of fore wing 1.3 - 1.5.
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY: The specific name refers to the two colours, brown and dull yellow, possessed by members of this
species.
TYPE MATERIAL.
, Estancia do Santo Castello, Nov. 1976, M. Alvarenga, (CNC).
Hebichneutes citrinus n. sp.
FEMALE DESCRIPTION.
Length: 4.1 to 4.6 mm.
Colour: Yellow except as follows: basal 2/3 of flagellum brown, scape and pedicel light brown, hind tarsus
and apex of hind tibia brown, wings infuscate.
Structural features: antenna with 31 flagellomeres; ratio of vein 2Rs:3Rs of fore wing 1.3 - 1.4.
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY. The specific name citrinus is Latin meaning "of citron" and is a reference to the yellow colour of this
species.
TYPE MATERIAL.
Alvarenga, (CNC).
Hebichneutes tricolor n. sp.
(Figs 22, 37, 45)
FEMALE DESCRIPTION.
Length: 3.0 to 3.6 mm.
Colour: Brown, dull yellow and ivory, dull yellow except as follows: head brown dorsally, laterally and
20
GENERA OF ICHNEUTINAE
JAN. 7, 1993
posteriorly, antenna brown, scape and pedicel light brown, metapleuron and propodeum brown; middle and hind femora,
tibiae and tarsi brown, most of metasoma except anterolaterally and first tergum apically brown, first tergum apically
and metasoma anterolaterally ivory, all coxae and trochanters ivory to ivory and brown; wings infuscate.
Structural features: antenna with 26 to 29 flagellomeres; ratio of vein 2Rs:3Rs of fore wing 1.3-1.4 (Fig. 37).
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY: The specific name refers to the three colours, brown, ivory and dull yellow, possessed by members of
this species.
TYPE MATERIAL.
D.H. Janzen, (AEI).
June 16, 1976; June 5, 1977; Sept. 27, 1977; Sept. 28, 1977; and
la, June 6, 1969, W.R.M.
Mason, (CNC).
Hebichneutes wasbaueri n. sp.
MALE DESCRIPTION.
Length: 4.0 mm.
Colour: Yellow except as follows: scape and pedicel brown, flagellum black, temple with small patch of
brown, mid and hind tibiae and tarsi brown, metasomal terga 6-8 pale brown, wings infuscate.
Structural features: antenna with 28 flagellomeres; ratio of vein 2Rs:3Rs of fore wing 1.4.
REMARKS. Female unknown.
ETYMOLOGY. Named in honour of the collectors of the type specimen.
TYPE MATERIAL.
H
Wasbauer, (CNC).
-4, 1992, 300m, M.&J.
Helconichia NEW GENUS
Key to species
1
Compound eyes with short setae, about equal to diameter of ommatidium ................................... H. areolata
1'
Compound
eyes
with
long
setae,
about
4X
diameter
of
ommatidium
...........................................................2
2
2'
Hind wing with 3 hamuli.......................H. trichopteryx
Hind wing with 4 hamuli...................... H. trichiops
Relationships of the species of Helconichia
On the basis of the following synapomorphies H. trichiopteryx is proposed as the sister species of H. trichiops:
1. reduced number of flagellomeres. 2. compound eyes (and body generally) with long setae. 3. posterior median areola
of propodeum lost and replaced with median longitudinal carina. 4. median longitudinal carinae of metasomal median
tergite 1 almost meeting medially.
Helconichia areolata n. sp.
21
GENERA OF ICHNEUTINAE
JAN. 7, 1993
(Figs 31, 40)
FEMALE DESCRIPTION.
Length: 3.0 to 3.2 mm.
Colour: Dull yellow and brown; brown as follows: antenna (scape and pedicel somewhat lighter), ocellar
triangle, hind tibia and tarsus, most of terga 3-7; wings lightly infuscate.
Structural features: antenna with 28 to 29 flagellomeres; setae of eyes short, about equal to diameter of
ommatidium; ratio of vein 2Rs:3Rs of fore wing 1.65 (Fig. 31); hind wing with 3 hamuli (Fig. 40); propodeum without
posterior median longitudinal carina; metasomal median tergite 1 ratio of length from base of medial carina / apical
width, 0.72; median tergite 1 with longitudinal carinae widely separated medially, distance between carinae about equal
to distance from carinae to lateral margin of tergite.
REMARKS. Male unknown.
ETYMOLOGY. Areolata is Latin meaning with small spaces and refers to the relatively well areolated propodeum of
this species.
TYPE MATERIAL.
-31, 1976, S. and J. Peck, (CNC).
C).
Helconichia trichiops n. sp.
FEMALE DESCRIPTION.
Length: 2.7 to 3.3 mm.
Colour: Dull yellow and brown; brown as follows: antenna (scape and pedicel somewhat lighter), ocellar
triangle, mesoscutum laterally and anteriorly (mesoscutum may be entirely dull yellow), mesopleuron, apex of hind tibia,
hind tarsus, metasomal terga 2 to 8 light brown (reduced to terga 4 to 8 in some specimens); wings lightly infuscate.
Structural features: antenna with 24 to 25 flagellomeres; setae of eyes long, at least 4 X diameter of an
ommatidium; ratio of vein 2Rs:3Rs of fore wing 1.2; hind wing with 3 hamuli; propodeum with posterior median
longitudinal carina; metasomal median tergite 1 ratio of length from base of medial carina / apical width, 0.79; median
tergite 1 with longitudinal carinae almost meeting medially, distance between carinae much smaller than distance from
carinae to lateral margin of tergite.
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY. Trichiops is Greek meaning `hairy eyes' and refers to the long setae on the eyes of this species.
TYPE MATERIAL.
Prov., 47 km S. Santo Domingo, Rio Palanque Res. Sta., July 22-31, 1976, S&J Peck, (CNC).
Helconichia trichopteryx n. sp.
MALE DESCRIPTION.
Length: 2.8 mm.
Colour: Dull yellow and brown; brown as follows: flagellum; scape and pedicel light brown, ocellar triangle,
mesoscutum except posteromedially, apex of hind tibia, hind tarsus, metasomal terga 4-8; wings lightly infuscate.
Structural features: antenna with 24 flagellomeres; setae of eyes long, at least 4 X diameter of an ommatidium;
22
GENERA OF ICHNEUTINAE
JAN. 7, 1993
ratio of vein 2Rs:3Rs of fore wing 1.0; hind wing with 4 hamuli; propodeum with posterior median longitudinal carina;
metasomal median tergite 1 ratio of length from base of medial carina / apical width, 0.96; median tergite 1 with
longitudinal carinae almost meeting medially, distance between carinae much smaller than distance from carinae to
lateral margin of tergite.
REMARKS. Female unknown.
ETYMOLOGY. Trichopteryx is `of hairy wings' in Greek and refers to the long setae on the wings of this species.
TYPE MATERIAL.
Holotype:
Masonbeckia NEW GENUS
Key to species
1
1'
2
2'
3
3'
4
4
5
5'
r crossvein of fore wing shorter than 3Rs (Fig. 34) ____________________________________M.
compressicornis n. sp.
r
crossvein
of
fore
wing
equal
to
or
longer
than
3Rs
(Fig.
33).________________________________________________ 2
Maxillary palpi with 4 segments, labial palpi with 3 segments.
_____________________________________________________3
Maxillary palpi with 6 segments, labial palpi with 4 segments.
_____________________________________________________ 5
Metasoma predominantly yellow _____________M. hansoni n. sp.
Metasoma predominantly brown _______________________4
Flagellomere 6 with perpendicular setae ventrally at mid length and at apex (cf. Fig. 5). ______ M.
pauxipalpus n. sp.
Flagellomere 6 with perpendicular setae ventrally restricted to apex. _____________________________M.
brunneigaster n. sp.
Hind femur brown, at least basally __________M. townesi n. sp.
Hind femur yellow ____________________M. flavofemorata n. sp.
Relationships of Species of Masonbeckia
Characters used in analysis
0 = plesiomorphic 1 = apomorphic
1.
Flagellomeres. 0: not laterally compressed, 1: laterally compressed.
2.
Number of maxillary palpomeres. 0: 6, 1: 4.
3.
Number of labial palpomeres. 0: 4, 1: 3.
4.
Base of mandible. 0: unsculptured, 1. sculptured (Fig. 12).
5.
Base of mandible. 0: without transverse carina. 0: with transverse carina (Fig. 12),
6
7.
Perpendicular setae on flagellomere 6. O: absent, 1: at apex,
2: at mid-length (unordered transformation series).
Spiracle on metasomal segment 7. 0: present, 1: absent.
The data set of Table 2 was run on Hennig86 (Farris 1988) and one tree, that of Figure 50 resulted.
Masonbeckia brunneigaster n. sp.
23
GENERA OF ICHNEUTINAE
JAN. 7, 1993
MALE DESCRIPTION.
Length: 2.9 mm.
Colour: brown, mouthparts and legs (except hind tarsus and tibia in distal 0.7) slightly paler brown; wings
infumate.
Structural features: antenna with 28 flagellomeres; flagellomeres not laterally compressed, flagellomere 6 as
wide as high; flagellomere 6 with perpendicular setae ventrally at apex and at mid-length (cf. Fig. 5); maxillary palpi
with 4 segments; labial palpi with 3 segments; basal area of mandible sculptured; ratio of vein 2Rs:3Rs of fore wing 2.0;
2Rs+M present; presence of spiracle of metasomal segment 7 unknown due to folding on metasomal terga of the sole
specimen known.
REMARKS. Female unknown.
ETYMOLOGY. Brunneigaster means brown belly in Latin and refers to the brown metasoma of this species.
TYPE MATERIAL.
Holotype:
Mason, (CNC).
Masonbeckia compressicornis n. sp.
(Figs 4, 5, 6, 12, 15, 17, 18, 23, 34, 43)
FEMALE DESCRIPTION.
Length: 2.4 to 3.8 mm.
Colour: yellow and brown; from almost entirely yellow except for brown flagellomeres, hind tarsus, and apical
0.7 of hind tibia, to mostly brown as follows: flagellomeres, mesosoma dorsally and metasoma dorsally, hind tarsus and
tibia in distal 0.7; wings infumate.
Structural features: antenna with 21 to 27 flagellomeres; flagellomeres laterally compressed, flagellomere 6 1.4
X higher than wide; flagellomere 6 with perpendicular setae ventrally at apex and at mid-length (Fig. 5); maxillary palpi
with 4 segments; labial palpi with 3 segments; basal area of mandible sculptured (Fig. 12); ratio of vein 2Rs:3Rs of fore
wing 1.2 to 1.6 (Fig. 34); 2Rs+M present; spiracle of metasomal segment 7 absent.
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY. Compressicornis means compressed horns in Latin and refers to the laterally compressed antennae of
this species.
Host Data. One specimen from Costa Rica collected by Dan Janzen has some biological data as follows: Host plant,
Cydista heterophylla, Bignoniaceae. Host insect, sawfly. Dan Janzen sent us the following note regarding the biology
of the parasite, "larvae (of host) eating young leaves, 3 larvae (of host) spun tight brown cocoons between leaves about 8
June, 1 parasite eclosed 14 July 1984 and I killed the other 2 cocoons by accident (parasite was mistaken at that time for
another sawfly); determination of sawfly must await further rearing." David Smith (expert on Symphyta from the
U.S.D.A. in Washington) suggested (personal communication) that the host is probably an argid sawfly but that a
confident identification at the family level is not possible with the limited information.
TYPE MATERIAL.
April, 1972, (CNC).
Paratypes: ARGENTINA: Tucumán, Villa Nogués, Jan. 1, 1966, H. & M. Townes, (AEI). BRAZIL:
M. Alvarenga, (AEI). Guanaba
24
GENERA OF ICHNEUTINAE
JAN. 7, 1993
-500 m, Nov. 1968, Fritz
-78, D.H. Janzen, (AEI).
Rosa Park, ex. cocoon of sawfly, host plant Cydista heterophyla, July 14, 1984, D.H. Janzen, (INBio).
ECUADOR: Pinchincha: Rio palanque Res. Station, July 1Grande N.P., cloud forest, maxinet, 1,100 m, A
-13,
Townes, (AEI).
Masonbeckia flavofemorata n. sp.
(Figs 33, 42)
FEMALE DESCRIPTION.
Length: 2.5 to 3.6 mm.
Colour: body yellow; flagellomeres brown; scape and pedicel from yellow to pale brown; head brown dorsally
at least between ocelli; mid and hind tarsi brown; hind tibia partly brown; metasoma posteriad segment 1 yellow to
brown; wings infumate.
Structural features: antenna with 24 to 25 flagellomeres; flagellomeres not laterally compressed, flagellomere 6
as wide as high; flagellomere 6 without perpendicular setae ventrally; maxillary palpi with 6 segments; labial palpi with
4 segments; basal area of mandible not sculptured; ratio of vein 2Rs:3Rs of fore wing 2.6; 2Rs+M present (Fig. 33);
spiracle of metasomal segment 7 present.
MALE DESCRIPTION. Essentially as in female, though somewhat more melanic.
REMARKS. Differs from M. townesi in that the latter is more melanic and larger, with relatively thicker hind femora.
ETYMOLOGY. Flavofemorata, is Latin meaning yellow femora and refers to the colour of the hind femora of the
species, one of the key diagnostic characters.
TYPE MATERIAL.
-31, 1976, S. & J. Peck,
(CNC).
Rio Palanque Res. Station, May 18-30, 1975. Peck, (CNC).
Masonbeckia hansoni n.sp.
FEMALE DESCRIPTION.
Length: 2.4 - 3.4 mm.
Colour: body yellow and brown, brown as follows: head except mouthparts, body of mesosoma, hind tibia
distally and hind tarsus, metasomal tergum 3 medially and most of terga 4-7; wings infumate.
Structural features: antenna with 26-28 flagellomeres; flagellomeres not laterally compressed, flagellomere 6
0.78 X as high as wide; flagellomere 6 with perpendicular setae ventrally at apex; maxillary palpi with 4 segments; labial
palpi with 3 segments; basal area of mandible sculptured (cf. Fig. 12); ratio of vein 2Rs:3Rs of fore wing 1.7-1.8;
2Rs+M present; spiracle of metasomal segment 7 present.
25
GENERA OF ICHNEUTINAE
JAN. 7, 1993
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY. The species is named after the collector of the type specimen.
TYPE MATERIAL.
Holotype:
-20, 1988, P. Hanson, (CNC).
-20, 1988, P. Hanson, (TAMU).
ill (CNC).
Masonbeckia pauxipalpus n. sp.
FEMALE DESCRIPTION.
Length: 3.8 mm.
Colour: brown; mouthparts and legs yellow except hind tarsus and tibia in distal 0.7 brown; wings infumate.
Structural features: antenna with 29 flagellomeres; flagellomeres weakly compressed laterally, flagellomere 6
1.12 X higher than wide; flagellomere 6 with perpendicular setae ventrally at apex and at mid-length (cf. Fig. 5);
maxillary palpi with 4 segments; labial palpi with 3 segments; basal area of mandible sculptured (cf. Fig.12); ratio of
vein 2Rs:3Rs of fore wing 2.5; 2Rs+M present; spiracle of metasomal segment 7 present.
REMARKS. Male unknown.
ETYMOLOGY. Pauxipalpus means few palpi and refers to the reduced number of maxillary and labial palpomeres of
this species.
TYPE MATERIAL.
Holotype:
Mason, (CNC).
Masonbeckia townesi n. sp.
MALE DESCRIPTION.
Length: 4 to 4.1 mm.
Colour: body yellow and brown, brown as follows: antenna, ocellar triangle, mesosoma (sometimes); pronotum
(sometimes) mesopleuron in part, metapleuron (sometimes) most or all of mesoscutum, all legs (sometimes); base of
anterior femur in one specimen, entire mid leg (sometimes) or restricted to base of mid femur, mid coxa in one
specimen, part of mid tibia in one specimen, mid tarsus; hind leg in one specimen completely brown, in the other partly
yellow on coxa, femur and tibia and median tergites of metasomal segments 4-7.
Structural features: antenna with 28 to 29 flagellomeres; flagellomeres not laterally compressed; flagellomere 6
as wide as high; flagellomere 6 without perpendicular setae ventrally; maxillary palpi with 6 segments; labial palpi with
4 segments; basal area of mandible not sculptured; ratio of vein 2Rs:3Rs of fore wing 2.6; 2Rs+M absent; spiracle of
metasomal segment 7 present.
REMARKS. Masonbeckia townesi differs from M. flavofemorata in that the former is lacking fore wing vein 2Rs+M, it
is also more melanic and larger, with relatively thicker hind femora. Female unknown.
ETYMOLOGY. Named after the late Ichneumonologist and co-collector of the type series, Henry Townes.
TYPE MATERIAL
-29, 1987,
(CNC).
26
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Tucumán, Jan. 5 and Jan. 18, 1966, H. & M. Townes, (AEI).
Muesonia NEW GENUS
Key to species
1
Metasomal terga 1 to 4 rugose. .......... M. sigalphoides
1
Metasomal terga 1 to 4 smooth. .....................2
2
Metasoma mostly dark reddish brown......... M. venus
2
Metasoma dull yellow........................M. straminea
Relationships of the species of Muesonia
Most traits that vary interspecifically are continuous or meristic characters that are difficult to code. The large
tubercles for the spiracles on metasomal tergum 1 suggest the that M. sigalphoides and M. venus are sister species.
Muesonia sigalphoides n. sp.
FEMALE DESCRIPTION.
Length: 7.5 mm.
Colour: Dark reddish brown and dull yellow. Dull yellow as follows: palpi, fore tarsus, middle tarsus in part,
metasoma; wings infuscate.
Structural features: antennae broken; flagellomeres not compressed laterally, flagellomere 6 as high as wide;
ratio of vein 2Rs:3Rs of fore wing 0.9; hind wing with 5 or 6 hamuli; spiracle of tergum 1 on laterally projecting
tubercle; metasomal terga 1 to 4 rugose, forming carapace; tergum 2 with transverse depression.
REMARKS. Male unknown
ETYMOLOGY. Sigalphoides is a combination referring to the braconid genus Sigalphus and oides which is Latin
meaning likeness. The name refers to the striking though partially superficial resemblance between this species and
certain members of the subfamily Sigalphinae.
TYPE MATERIAL.
ICO, Chiapas, 7,200 ft (2,194 m), San Cristobal de las Casas, June 10, 1969, W.R.M.
Mason, (CNC).
Muesonia straminea n.sp.
(Figs 28, 44)
FEMALE DESCRIPTION.
Length: 3.9 to 4.7 mm.
Colour: Dull yellow except brown as follows: flagellum, hind tibia apically, hind tarsus; wings infuscate.
Structural features: antenna with 26 to 28 flagellomeres; flagellomeres strongly compressed laterally,
flagellomere 6 1.4 X higher than wide; ratio of vein 2Rs:3Rs of fore wing 0.9 - 1.0 (Fig. 28); hind wing with 4 hamuli
(Fig. 44); spiracle of tergum 1 not on laterally projecting tubercle; metasomal terga smooth, not forming carapace;
tergum 2 without transverse depression.
MALE DESCRIPTION. Essentially as in female.
ETYMOLOGY. Straminea is Latin for of straw-like and refers to the colour of this species.
TYPE MATERIAL.
Holotype:
1981.
27
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Muesonia venus n. sp.
FEMALE DESCRIPTION.
Length: 5.0 to 5.1 mm.
Colour: Dull yellow, brown and ivory, brown as follows: antenna, ocellar triangle, metanotum, metapleuron,
propodeum, middle tibia apically, middle tarsus, hind leg, metasomal terga; metasomal sterna partly ivory, more so
anteriorly; wings infuscate.
Structural features: antenna with 31 flagellomeres; flagellomeres weakly compressed laterally, flagellomere 6
1.2 X higher than wide; ratio of vein 2Rs:3Rs of fore wing 0.95.; hind wing with 4 hamuli; spiracle of tergum 1 on
laterally projecting tubercle; metasomal terga smooth, not forming carapace; tergum 2 without transverse depression.
REMARKS. Male unknown
ETYMOLOGY. Venus is the Roman goddess of love, and symbol of beauty and grace. The name is a noun in
opposition.
TYPE MATERIAL.
Peck, (CNC).
Paratype: COLOMBIA:
18-20, 1970, H.F. Howden, (CNC).
28
GENERA OF ICHNEUTINAE
JAN. 7, 1993
REFERENCES
Achterberg, K. van. 1976. A preliminary key to the subfamilies of the
Braconidae (Hymenoptera). Tijdschrift Voor
Entomologie. 119:33-78.
Achterberg, K. van. 1984. Essay on the phylogeny of the Braconidae (Hymenoptera: Ichneumonoidea). Entomologisk
Tidskrift 105: 41-58.
Ashmead, W.H. 1900. Classification of the ichneumon-flies or the superfamily Ichneumonoidea. Proceedings of the
United States National Museum 28:127-158.
Austin, A.D. & R. A. Wharton. 1992. New records of subfamilies, tribes and genera of Braconidae (Insecta:
Hymenoptera) from Australia, with descriptions of seven new species. Transactions of the Royal Society South
Australia 116: 41-63.
Belokobylskij, S. 1986. Five new species of braconids (Hymenoptera: Braconidae) from the Asiatic part of the USSR,
pp 28-40. In: Ler, P.A., S.A. Belokobylskij and N.A. Strorozheva, eds., Hymenoptera of Eastern Siberia and
the Far East, collected works. Academy of Sciences USSR, Far East Science Centre, Vladivostok. 152 pp.
Belokobylskij, S. 1989. East Palaearctic species of the braconid genera Dirrhope and Mirax (Hymenoptera:
Braconidae: Miracinae). Vestnik Zoologii 4:34-46.
Belokobylskij, S. 1990. A contribution to the braconid fauna (Hymenoptera) of the Far East. Vestnik Zoologii 6: 31-39.
Buckingham, G.R. and M.J. Sharkey. 1988. Abdominal exocrine
glands in the Braconidae (Hymenoptera). In
Advances In
Parasitic Hymenoptera Research, V.K. Gupta ed., E.J. Brill,
Leiden, pp. 199-242.
Cresson, E.T. 1887. Synopsis of the families and genera of the Hymenoptera of America north of Mexico with a
catalogue of the described species and bibliography. Transactions of the American entomological Society
Supplement 1-350.
de Gaulle, J. 1907. Catalogue systématique et biologique des Hyménoptères de France. Fam. VIII Braconidae. Feuille
des jeunes Naturalistes 38. 1907.
Farris, J.S. 1988. HENNIG86 reference: documentation for version 1.5. port Jefferson Station, New York.
Förster, A. 1862. Synopsis der Familien und Gattungen der Braconiden. Verhandlungen Des Naturhistorischen Vereins
De Preussischen Rheinlande Und Westfalens Bonn 19: 225-288.
Kolubajiv, N.G., 1962. Die Ergebnisse der Zuchten von Entomophagen (der Parasiten und Rauber) der schädlichen
-73.
Marshall, T.A. 1885. Monograph of British Braconidae. Part I. Transactions of the Royal entomological Society London
pp. 1-280.
Marshall, T.A. 1888. Les Braconides. Species des Hyménoptères d'Europe & d'Algérie. E. André (ed.) 4:1-609.
Beaune (Côte d'or).
Mason, W.R.M. 1957. A new genus and species of Microgastrinae (Hymenoptera: Braconidae). The Canadian
Entomologist 89: 355-357.
Mason, W.R.M. 1969. Muesebeckiini, a new tribe of Braconidae (Hymenoptera). Proceedings of the entomological
Society of Washington. 71: 263-278.
Mason, W.R.M. 1991. A new genus of exodont Ichneutinae (Hymenoptera: Braconidae). Insecta Mundi 5: 95-99.
Muesebeck, C.F.W. 1931. Descriptions of a new genus and eight new species of ichneumon-flies with taxonomic notes.
Proceedings of the United States national Museum. 79(16): 1-16.
Muesebeck, C.F.W. and L. Walkley. 1951. Braconidae, In Muesebeck, C.F.W. Muesebeck, Krombein, K.V. and
Townes, H.K.. Hymenoptera of America north of Mexico. Synoptic Catalogue. Agriculture Monograph 2: 90184.
Nees von Esenbeck, C.G. 1816 (1813). Ichneumonides adsciti, in genera et familias divisi. Magazin Gesellschaft
Naturforschender Freunde Zu Berlin 7: 243-277.
Nixon, K.C. 1992. Clados reference: documentation for version 1.2. Tromansburg, New York.
Papp, J. 1987. Braconidae (Hymenoptera) from Korea, VIII. Acta Zoologica Hungarica 33: 157-175.
Quicke, D.L.J. and C. van Achterberg. 1990. Phylogeny of the subfamilies of the family Braconidae (Hymenoptera:
29
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Ichneumonoidea). Zoologische Verhandelingen (Leiden) 258: 1-95.
Sharkey, M.J. 1992. Cladistics and tribal classification of the Agathidinae (Hymenoptera: Braconidae). Journal of
Natural History 26: 425-447.
Shaw, M.R. and T. Huddleston. 1991. Classification and biology of braconid wasps (Hymenoptera: Braconidae),
Handbooks for the Identification of British Insects 7(11) 126 pp.
Shenefelt, R.D. 1973. Braconidae 5. Microgasterinae and Ichneutinae. Hymenopterorum Catalogus. Pars 9: 669812.
Szépligeti, G.V. 1896. Beitrage zur Kenntnis der Ungarischen Braconiden. Természetrajzi Füzetek 19:228-242, 359386.
Szépligeti, G.V. 1902. Tropische Cenocoelioniden und Braconiden
aus der Sammlung des Ungarischen NationalMuseums. Természetrajzi Füzetek. 25:39-84.
Tobias, V.I. 1986a. Subfam. Ichneutinae. Handbook to the insects of the European part of the USSR. 3. Hymenoptera.
Part 4. 459. Akademia Nauk, Lenningrad.
Tobias, V.I. 1986b. Subfam. Miracinae. Handbook to the insects of the European part of the USSR. 3. Hymenoptera.
Part 4. 291-293. Akademia Nauk, Lenningrad.
Tobias, V.I. and S. Belokobylskij. 1981. Braconid genera (Hymenoptera: Braconidae) from Primorye Territory new to
science and to USSR fauna. Entomologicheskoe Obozrenie 60(2): 354-363.
Viereck, H.L. 1909. Descriptions of new Hymenoptera. Proceedings of the entomological Society of Washington. 11:
42-51.
Wesmael, C. 1835. Monographie des Braconides de Belgique. Nouveaux Mémoires de l'Acadamie r. des Sciences,
Bruxelles 9:1-252
Wharton, R.A., S.A. Shaw, M.J. Sharkey, D.B. Wahl, J.B. Woolley, J.B. Whitfield, P.M. Marsh, J.W. Johnson. (1992).
Phylogeny of the subfamilies of the family Braconidae (Hymenoptera: Ichneumonoidea): a reassessment.
Cladistics 8: in press.
Whitfield, J.B. and D.L. Wagner. 1991. Annotated key to the genera of Braconidae (Hymenoptera) attacking leafmining
Lepidoptera in the Holarctic region. Journal of Natural History 25: 733-754.
30
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Table 1. Data set for genera of Ichneutinae with outgroups and hypothetical ancestor (Out); ? = polymorphic or (in the
case of character 36) unknown.
TAXA
CHARACTERS
111111111122222222223333333
123456789012345678901234567890123456
Out
000000000000?000000000000?00000000?0
Agathidinae 00110000100000001000?000000000000000
Sigalphinae 000000000000000010000000000000000100
Hebichneutes 00100000000100011110001110010000111?
Muesonia
00100001000000101010001110000000011?
Proterops 221?000000000001011111111?0000000111
Helconichia 0000000000010000?010001110010000011?
Ichneutes 1110000210100001?00000101000110001?2
Paroligoneurus 111100121110110101001?111?01010?0110
Lispixys
11110002111011010100101111110100011?
Oligoneurus 111100021110100101001?1111?101110110
Masonbeckia 001011000000000111100011110001000111
31
GENERA OF ICHNEUTINAE
JAN. 7, 1993
Table 2. Data set for Masonbeckia species; hypothetical ancestor = Out; ? = polymorphic.
TAXA
CHARACTERS
1234567
Out
0000000
M. brunneigaster
011112?
M. compressicornis
1111121
M. flavofemorata
0000100
M. hansoni
0111110
M. pauxipalpus
1111120
M. townesi
0000100
32
GENERA OF ICHNEUTINAE
JAN. 7, 1993
CAPTIONS FOR FIGURES
FIGURE 1. Habitus drawing of Ichneutes sp..
FIGURES 2 - 7.
FIGURE 2. Paroligoneurus eximus (Mason), enlarged from Figure 3, showing truncated setae.
FIGURE 3. Paroligoneurus eximus (Mason), penultimate flagellomere showing pair of widely separated longitudinal
placodes, with field of modified (truncated) setae between.
FIGURE 4. Masonbeckia compressicornis n. sp., lateral view of basal flagellomere showing patch of basiconic sensillae.
FIGURE 5. Masonbeckia compressicornis n. sp., lateral view of 6th flagellomere showing perpendicular setae at mid
length and at apex.
FIGURE 6. Masonbeckia compressicornis n. sp., lateral view of anterior portion of mesosoma showing: subpronope (s),
longitudinal carina of propleuron (lc), and the absence of an epicnemium.
FIGURE 7 . Ichneutes sp., showing the presence of an epicnemium (e) and sternaulus (s) and the lack of a longitudinal
carina on the propleuron.
FIGURES 8 -13.
FIGURE 8. Proterops Proterops sp., dorsal view of head showing alignment of ocelli.
FIGURE 9. Proterops proteroptoides Viereck, dorsal view of head showing alignment of ocelli.
FIGURE 10. Proterops Proterops sp., anterior view of head showing twisted mandibles.
FIGURE 11. Proterops proteroptoides Viereck, anterior view of head showing straight mandibles
FIGURE 12. Masonbeckia compressicornis n. sp., anterior view of lower head showing sculptured area and transverse
carina at base of mandible.
FIGURE 13. Paroligoneurus alachua (Mason), anterior view of head showing exodont mandibles (twisted 180 degrees).
FIGURES 14 - 19.
FIGURE 14. Proterops proteroptoides Viereck, showing swollen hind tibia which houses a large subgenual organ.
FIGURE 15. Masonbeckia compressicornis n. sp., base of hind tibia showing modified area which indicates the site of
basal attachment of the subgenual organ.
FIGURE 16. Ichneutes sp., apex of fore tibia showing blunt spine.
FIGURE 17. Masonbeckia compressicornis n. sp., dorsal aspect of anterior portion of mesosoma, showing subpronopes.
FIGURE 18. Masonbeckia compressicornis n. sp., dorsal aspect of posterior portion of mesosoma, showing posterior
scutellar depression.
FIGURE 19. Proterops proteroptoides Viereck, dorsal view of posterior portion of mesosoma showing lack of sculpture
and lack of posterior scutellar depression.
FIGURES 20 - 27.
FIGURE 20. Ichneutes sp., dorsal view of first metasomal tergum showing two precurrent longitudinal carinae.
FIGURE 21. Lispixys avispa Mason, dorsal view of first metasomal tergum showing unmargined median longitudinal
groove.
FIGURE 22. Hebichneutes tricolor n. sp., dorsal view of first metasomal tergum showing converging median
longitudinal carinae (mc) and lateral longitudinal carinae (lc).
FIGURE 23. Masonbeckia compressicornis n. sp., dorsal view of first metasomal tergum showing smooth surface
unmodified by grooves and carinae.
FIGURE 24. Oligoneurus sp. dorsal view of posterior segments of male metasoma showing openings to tergal glands on
metasomal segments 6 and 7.
FIGURE 25. Paroligoneurus sp. dorsal view of posterior segments of male metasoma showing opening to tergal gland
on metasomal segment 7.
FIGURE 26. Ichneutes sp., dorsal view of posterior segments of male metasoma showing openings to tergal gland on
metasomal segment 7.
FIGURE 27. Paroligoneurus eximus (Mason), lateral view of posterior segments of female metasoma, showing terminal
33
GENERA OF ICHNEUTINAE
JAN. 7, 1993
setae on ovipositor sheath and short sharp ovipositor.
FIGURES 28 - 37.
FIGURE 28. Muesonia straminea n.sp., fore wing.
FIGURE 29. Oligoneurus sp., fore wing.
FIGURE 30. Proterops proteroptoides (Viereck), fore wing.
FIGURE 31. Helconichia areolata n. sp., fore wing.
FIGURE 32. Paroligoneurus eximus (Mason), fore wing.
FIGURE 33. Masonbeckia flavofemorata n. sp., fore wing.
FIGURE 34. Masonbeckia compressicornis n. sp., fore wing.
FIGURE 35. Ichneutes sp. fore wing.
FIGURE 36. Paroligoneurus sp., fore wing.
FIGURE 37. Hebichneutes tricolor n. sp., fore wing.
FIGURES 38 - 47.
FIGURE 38. Paroligoneurus eximus (Mason), hind wing .
FIGURE 39. Paroligoneurus johnsoni, hind wing.
FIGURE 40. Helconichia areolata n. sp., hind wing.
FIGURE 41. Proterops proteroptoides (Viereck), hind wing.
FIGURE 42. Masonbeckia flavofemorata n. sp., hind wing.
FIGURE 43. Masonbeckia compressicornis n. sp., hind wing.
FIGURE 44. Muesonia straminea n.sp., hind wing.
FIGURE 45. Hebichneutes tricolor n. sp., hind wing.
FIGURE 46. Oligoneurus sp., hind wing.
FIGURE 47. Ichneutes sp., hind wing.
FIGURES 48 - 49.
FIGURE 48. First of two minimum length cladograms for ichneutine genera (Table 1). This and following cladograms
were made with the clados program (Nixon, 1992).
FIGURE 49. Second of two minimum length cladograms for ichneutine genera (Table 1).
FIGURE 50. Sole minimum length tree for the data set of Masonbeckia species (Table 2).
34