Larvae of : Generic Characteristics and a Description of Micromus subanticus (: )

ALAN H. KRAKAUER AND CATHERINE A. TAUBER Department of Entomology, Comstock Hall, Cornell University, Ithaca NY 14853-0901

Ann. Entomol. Soc. Am. 89(2): 203-211 (1996) ABSTRACT Micromus and Hernerobius are the most common and agriculturally important genera of hemerobiids in North America. Twelve morphological traits (8 cephalic, 4 thoracic) differentiate the larvae of these genera. Additional structural, chaetotaxic, and color traits distinguish M. subanticus (Walker) and M. posticus (Walker) larvae. The larval stages of M. subanticus are described.

KEY WORDS Hemerobiidae, Micromus subanticus, larvae

THE BROWN LACEWING family Hemerobiidae is a Our article focuses on the comparative mor- cosmopolitan neuropteran group of ~575species; phology of North American hemerobiid larvae, worldwide, it currently contains 10 subfamilies and with emphasis on the Micromus. It has the 27 genera (Monserrat 1993; Oswald 1993a, b, following 3 goals: (1)identification of generic-level 1994). In North America the hemerobiid fauna larval characters, (2) delineation of -specific consists of 6 genera and ~60species (Kevan and characters for Micromus larvae, and (3) description Klimaszewski 1987, Klimaszewski and Kevan 1988, of the instars of M. subanticus. Oswald 1993a). Most s~eciesof hemerobiids are Ideally, for a phylogenetically based analysis, we predaceous ii both iAaginal and preimaginal would have contrasted Micromus larvae with those stages, and many are of considerable value as bi- of its sister-group (Megalomina, according to Os- ological control agents (Dunn 1954, New 1975a, wald 1993a); unfortunately larvae of this and other Miller and Cave 1987). genera in the subfamily are unknown. Systematic studies of the family have focused on Therefore, we emphasized Hemerobius in the the adults; there has been very little consideration comparison for several reasons. First, along with of hemerobiid larvae. Oswald's (1993a) treatment Micromus, Hemerobius is a large and common ge- of the higher phylogeny and worldwide nus for which we had larval specimens. Second, larvae of the 2 genera share similar body plans, life of the family considered only adult characters. styles, and habitats. Third, the 2 genera frequently Keys to the adults of North American genera and occur syrnpatrically in agricultural situations, and species can be found in Carpenter (1940), Kevan biological control practitioners as well as other pest and Klimaszewski (1987), Klimaszewski and Kevan management personnel would benefit from relia- (1987a, b; 1988; 1992), and Oswald (1993a). Lar- ble means for identifying the larvae. vae of some North American species are described Worldwide, descriptions of various quality exist (for example, Smith 1923; Withycombe 1924; Kil- for the immatures of 7 species of Micromus, 3 of lington 1936, 1937; MacLeod 1960; Miller and which inhabit North America: M, angulatus (Ste- Cave 1987: Miller and Lambdin 1984). and a brief phens), M, posticus, and M. vaviegatus (F.).In ad- key exists for the larvae of some genera in Europe dition, preimaginal stages of 12 species of Heme- (Veenstra et al. 1990). These studies, together with robius have been described, including the several systematic work on the larvae of other Neuropter- species that occur in North America: H. humulinus an groups (e.g., Tauber 1969, Stange and Miller L., H. pacijicus Banks, H. simulans Walker, and H. 1990, Hoffman and Brushwein 1992), indicate that stigma Stephens. larval morphology may be very useful in the sys- tematic analysis of the family (Tauber and Adams 1990). Moreover, improvements to the systematics Materials and Methods of hemerobiid larvae would be very useful for in- Material Examined. We examined larvae of Mi- creasing the effiecient use of hemerobiids in bio- cromus subanticus, M. posticus, Hemerobius hu- logical control programs, especially during moni- mulinus, and H. conjunctus (Fitch). All specimens toring and evaluation. were F1 offspring of adults collected in Ithaca,

46/96/0203-0211$02.00/0 O 1996 Entomological Society of America 204 ANNALSOF THE ENTOMOLO(XCAL SOCIETYOF AMERICA Vol. 89, no. 2

Tompkins County, New York. We also examined palpi; the distal segment is the longest. Three-seg- larvae of M. subanticus that had been reared from mented antennae arise above and behind the man- adults collected in Chandler. Mari~osaL Countv. i, Ar- dibles. The scapes are small, approximately as long izona. Representative specimens are in the Cornell as wide, and the pedicelli and flagella are thin, an- University Collection (Lot #1205). In com- nulated, and long (combined, usually longer than piling generic and species-specific traits, we con- the jaws). The eyes consist of 3 transparent, ovoid sidered previous descriptions of hemerobiid larvae stemmata raised over a variable number of pig- wherever possible. mented areas (New and Boros [I9831 report 4 Rearing. Field-collected adults were kept in stemmata in M, tasmaniae). glass vials under a photoperiod of 16:8 (L:D) h at The thorax and abdomen are of approximately 21 f- 1°C. They were provided green peach , equal length. The prothorax consists of 2 subseg- Myzus persicae (Sultzer), as prey; cotton plugs in ments; the anterior one, the cervix, is unscleroti- the vials served as the substrate for oviposition. zed, whereas the posterior subsegment bears a pair

Larvae were individually reared on a diet of green of I~rominent dorsal sclerites. The meso- and meta- peach aphids and, occasionally pea aphids, Acyr- thoraces each have very short anterior subseg- thosiphon pisum (Harris); they were examined dai- ments and a pair of small dorsal sclerites. A pair ly for molting and cocoon spinning. To increase the of large spiracles occurs laterally on the mesotho- humidity for the prepupae and pupae, we held the rax. (There is a singular reference to metathoracic cocoons in a plastic bag containing a moist paper spiracles in LM. tasrnuniae [New and Boros 19831.) towel. The cursorial larvae have well-developed, 3-seg- Specimen Preparation. Larvae were killed in a mented legs (with pulvilli arising apically on the solution of kerosene, ethanol, glacial acetic acid, tarsi of 1st instars). The abdomen consists of 10 and dioxane (KAAD solution; Stehr 1987), and segments; the 1st segment is the widest and short- stored in 95% ethanol. Samples of each instar were est; subsequent segments taper. Spiracles occur cleared in ~10%KOH solution for 2 d. Larger laterally on the 1st 8 abdominal segments, a dorsal instars were sectioned transversely through the sclerite on the 9th, and dorsal, ventral, and lateral thorax or abdomen to facilitate the removal of the sclerites on the 10th. Overall, the body is sparsely internal viscera. Subsequently, some larvae were covered with simple setae; and there are no pro- placed in clearing fluid (a mixture of lactic acid, tuberances, tubercules, or modified setae (with the liquid phenol, and glacial acetic acid) with 1 drop possible exception of spatulate abdominal setae on of Double stain for 3-6 h, then transferred to un- M. tasrnuniae [see New and Boros 19831). stained clearing solution for an additional 3 d. All larvae were taken through a series of alcohol so- Comparison of Micromus and Hemerobius lutions (70% [l min], 80% [2 min], and 95% EtOH [3 rnin]). Finally, specimens were transferred to a Hemerobius and ~Micromuslarvae share several mixture of glycerin and 95% EtOH, and, after sev- traits that distinguish them from the 6 other gen- eral minutes, mounted in glycerin on glass slides. era for which larvae have been described. Unlike Measurements. We measured the following the larvae of Psectra, Megalomus (including Bor- structures: total length of body (tip of mandibles iomyia), and Sympherobius, whose antennae are to tip of abdomen), width of head (at the level of relatively short, the antennae of Hemerobius and the eyes, including the eyes), length of head cap- Micromus project beyond the jaws and are longer sule (at the midline), length of mandibles (straight- than the width of the head capsule (Killington line distance from tip to medial margin of the 1936, 1937, 1946; MacLeod 1960; Veenstra et al. base), length of flagellum and pedicel, length of 1990). Also, unlike Megalomus, Psectra, Drepana- scape (medial), length of labial palp (base of men- cra, or Sympherobius, the head capsule is not re- tum to tip of terminal segment), length of distal cessed into the cervix (references above and New segment of labial palp (base to tip), width and [1975b]). Finally, with the possible exception of length of cervix, prothorax and mesothorax (me- (New and Boros 1983), the dial), length of prothoracic tibia (at the midline), terminal segment of the palps is elongate and fu- and length of prothoracic tarsus (excluding pulvil- siform in Microm~zusand Hemm~vbius,not swollen . \ as in Megalomus, Psectra, Drepanacra, and Sym- pherobius. \Vesmaelius larvae have jaws that are -1.5 times as long as the width of the head cap- General Morphological Characteristics of sule; in Micromus and Hemerobius the jaw length Hemerobiid Larvae is approximately equal to the head width (Killing- Hemerobiid larvae are fusiform and flattened ton 1936, 1937; Veenstra et al. 1990; however, see dorsoventrally (Smith 1923, Withycombe 1924, plate XI, figure 5, Killington 1936). Drepanepteryx Killington 1936, Tauber 1991). The head is prog- larvae are distinguished by anterior legs almost nathus, cordate, and depressed. The jaws-man- twice as long as the head, including the jaws (Mor- dibles above and the maxillae below-are long and ton 1910, Killington 1937); in Micromus and Hem- medially curved; the mandible tips are serrated. erobius the prothoracic legs are only 1-1.5 times Below the maxillae originate %segmented labial as long as the head and jaws. March 1996 KRAKAUERAND TAUBER:HEMEROBIID LARVAE 205

1 long segment, when viewed at magnifications up to 400X. Antennal Length. Antennal lengtwjaw length ra- tio: Micromus, >2.0; Hemerobius, <2.0 (usually) (Table 1).Values for Micromus species are consis- tently >2.0; Hemerobius larvae show more varia- tion, but values are usually <2.0. The ratio of an- tennal lengthhead width has also been used to differentiate Micromus and Hemerobius larvae (for instance, Veenstra et al. 1990); the same values and variation apply to this ratio. Frontal Width. Frontal widthhead width ratio (at the level of the eyes): Micromus, >0.5; Heme- robius, <0.5. In the species we studied and in all published figures with sufficient detail, the frons occupies >1/2 the width of the head capsule in Micromus, and generally <1/2 the width of the head in Hemerobius. Frontal Suture. Frontal suture: Micromus, U- shaped (Fig. 1);Hemerobius, V-shaped. Typically, in Micromus the arms of the epicranial suture are broadly joined and the frons is rounded posteri- orly; in Hemerobius the epicranial arms join acute- ly and the frons is angular posteriorly. The frontal suture of Micromus 1st instars is similar to that of 2nd and 3rd instars; however, in Hemerobius, 1st instars often lack dorsal pigmentation, making the structure difficult to identify. Genae. Left and right genae: ~Micromus,meeting dorsally; Hemerobius, without dorsal contact. In Micromus the frontal sutures extend only to the middle of the head and the genae occupy the dor- sum of the head posteriorly; a coronal suture oc- curs between the genae. In Hemerobius the frons extends to (or near to) the posterior margin of the

head ca~sule:L ' as a result the 0genae do not make contact dorsally and the coronal suture is absent or greatly reduced. Pleurostomal Suture. Pleurostomal suture: Mi- Fig. 1. Head of M. subanticus 3rd instar, dorsal, left cromus, unbranched; Hemerobius, branched (Fig. and ventral, right. Setal nomenclature follows the scheme 2). Micromus is characterized bvi a I~leurostomal for chrysopid larvae (Rousset 1966, Tauber et al. 1992). suture that extends between the eye and the base Setal designations indicate position on the head and are of the scape. In Hemerobius this suture branches not meant to imply homology with chrysopid setae; ec- around the eye (2nd and 3rd instars), or it has a dysial suture, es. Scale = 0.4 mm. small bud-like protrusion below the eye (1st in- stars). This character is not mentioned in any pre- Below we present 12 morphological character- vious descriptions, but it appears in illustrations istics of 2nd and 3rd instars that distinguish the (for instance, Killington 1936, p. 98). Micromus from the Hemerobius species we exam- Genal Marking. Triangular genal marking at the ined. Eight of these characteristics also apply to lateral base of the mandibles: Micromus, dark; 1st instars. Because many of the distinguishing Hemerobius, pale (Fig. 2). The genal marking in traits are noted or figured in descriptions of other Micromus typically is medium to dark brown (al- Micromus and Hemerobius species, we propose most as dark as the eyes); in Hemerobius it is only that they generally differentiate the larvae of these outlined or lightly tinted (often pale in 1st instars). 2 genera. This character was consistent in all the specimens Head Characters. Antennal Pedicel. Distal we examined and in published illustrations (for ex- width of antenna1 pedicel: Micromus, subequal to ample, Killington 1936, p. 98). adjacent flagellar width (Fig. 1);Hemerobius, wid- Frontal Markings. Frontal region: Micromus, er than adjacent flagellar width. Hemerobius is uniformly colored (pale yellow or tan to dark characterized by a distinct narrowing at the pedi- brown), without a medial marking; Hemerobius, cel-flagellum junction; this narrowing is absent in pale with dark brown or black medial marking. Micromus so that the 2 segments appear fused into This character was visible only on our 2nd- and 206 ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA Vol. 89, no. 2

Table 1. Characters that distinguish Micromus from Hemerobius larvae

Species Character Instar M. subanticus M. posticus H. humulinus H. conjunctus Junction of pedicel and flagellum All Indistinct Indistinct Distinct Distinct Ratio: antenna lengtha/jaw length 3rd 2.5-3.3 2.0-2.3 1.61.8 1.61.7 2nd 2.4-3.5 2.1-2.3 1.5-1.9 1.5-1.7 1st 2.44.1 2.03.6 1.2-2.1 1.8-1.9 Ratio: antenna lengthahead width 3rd 2.1-2.2 1.9-2.3 1.3-1.5 1.0-1.1 2nd 2.1-2.4 1.8-2.4 1.2-1.3 0.9-1.0 1st 2.1-2.9 2.62.5 1.41.8 1.0-1.1 Color of head (dorsal) 3rd, 2nd Brown with a pale frontal structure White with dark longitudinal mark- ing medially on frons Epicranial suture (Fig. 1) All Y-shaped, branching centrally on V-shaped, branching at posterior head capsule margin of head capsule Pleurostomal suture (Fig. 2) All Unbranched Unbranched Branchedb Branchedb Triangular genal marking (Fig. 2) All Medium to dark brown As light or lighter than dorsum of head capsule Dorsal body markings 3rd, 2nd Diffuse purple or brown dorsolateral Well-defined dorsolateral bands on pigmentation on buff or beige white body body Ratio: cervical lengthwidth 3rd 0.7-1.2 0.9-1.5 0.7-0.8 0.60.8 2nd 0.61.3 0.61.3 0.60.8 0.4-0.6 1st 0.5-0.9 0.7-0.9 0.40.6 0.5-0.7 Prothoracic sclerites 3rd, 2nd Sigmoid or crescentric; only anterior Crescentric, with anterior and poste- end curved medially nor ends curved laterally 1st Indisctinct, triangular, or elliptical Distinct, often with lateral edge straight Ratio: tarsial length/tibial length 3rd 0.20-0.25 0.20-0.24 0.46-0.58 0.67-0.72 2nd 0.31-0.33 0.28-0.33 0.56-0.76 0.62-0.75 1st 0.41-0.50 0.33-0.60 0.50-1 .OO 0.75-0.83

In addition ot our specimens, we considered published descriptions and figures of Micromus and Hemerobious larvae (Terry 1980; Smith 1923; Withycombe 1924; Killington 1936, 1937; Dunn 1964; Nakahara 1954; Kawashima 1958; New and Boros 1983; Miller and Lambdin 1984; Miller and Cave 1987). Antenna length includes the pedicel and flagellum, but not the scape. Branch may be small, bud-like in 1st instar.

3rd-instar specimens. It is consistent in most larval crescentic with the anterior end arched laterally. descriptions; its reliability as a diagnostic character The sclerites of 1st instars are: Micromus, indis- needs study. tinct, wedge-shaped, or elliptical; Hemerobius usu- Body Characters. Thoracic Pigmentation. Tho- ally distinct, rectangular. rax: Micromus, with broad transverse bands ante- Prothoracic Tarsi. Ratio of prothoracic tarsavtib- riorly on the meso- and metathorax; Hemerobius, ial length: Micromus, <0.25; Hemerobius, >0.3 with 2 well-defined dark purple or brown longi- (2nd and 3rd instars); Micromus, <0.5; Hemero- tudinal bands. Both species have longitudinal dor- bius, >0.5 (1st instars). This trait is consistent in solateral stripes on the cervix and longitudinal all previously published illustrations that show suf- bands on the dorsum of the abdomen. In Micro- ficient detail (for example, New and Boros 1983, mus the abdominal bands are diffuse; in Hemero- Kawashima 1958, Miller and Lambdin 1984). bius they are well defined. This trait is visible in (Measurements of the tarsi do not include the pul- 2nd and 3rd instars only and it may be variable villi.) among Hemerobius species. Ceruical Length. Ratio of cervical length /width Distinguishing Characteristics of M. posticus (at widest point): 3rd instar Micromus, >0.9; Hem- and M. subanticus erobius, < 0.8; 2nd instar Micromus, >0.8; Hem- erobius, <0.65. Obtaining consistent measure- To the naked eye, M. posticus and M. subanticus ments of membranous regious, such as the cervix, appear nearly identical. Nevertheless, several char- is difficult, and there was some overlap in the ratios acteristics differentiate the species. of the 2nd instars of the 2 genera (Table 1).This Structural Characters. Cervical Length. Ratio character is generally consistent in the descriptions of cervix length/width (2nd and 3rd instars only): and illustrations of Micromus and Hemerobius lar- M. posticus, >1; M. subanticus <1. Because the vae, except M. tasmaniae, which appears to resem- cervix is membranous, this character may vary ble Hemerobius (New and Boros 1983). among preserved specimens (see Table 1). Prothoracic Sclerite. Dorsal prothoracic sclerites Labial Palpus. Angle of attachment below a hor- of 2nd and 3rd instars: Micromus, sigmoid or cres- izontal plane through the longitudinal axis of body centic, with the anterior end parallel to the longi- (2nd and 3rd instars): M. posticus, =90°; M. sub- tudinal body axis or curved medially; Hemerobius, anticus, <45'. March 1996 KRAKAUERAND TAUBER:HEMEROBIID LARVAE

Fig. 2. Lateral view of 3rd instar head: H. humulinus (above), M. subanticus (below). Pleurostomal suture, pl; Genal marking, g. Scale = 0.5 mm.

Coloration. Dorsal Head Markings. Markings Tibia1 Markings. Ratio of tibial marking length/ anterior to ecdysial suture (2nd and 3rd instars: M. tibial length (all instars): M. posticus, >0.5; M, sub- posticus, obscure or lacking, not contrasting with anticus, <0.4. ecdysial area; M, subanticus, light brown, contrast- Chaetotaxy. The head capsule and abdominal ing with ecdysial area. chaetotaxy of M. subanticus and LM.posticus are

Table 2. Size of M. subanticus larval structures

Character 3rd instar (n = 2nd instar (n = 17)a 1st instar (n = 14)a Total lengthb 6.7 t 1.1 (5.1-9.6) 4.0 t 0.57 (3.2-5.3) 2.7 t 0.62 (1.8-3.35) Width of headC 0.57 t 0.02 (0.54-0.59) 0.41 t 0.02 (0.38-0.43) 0.31 i 0.01 (0.28-0.33) (n = 5) (n = 5) (n = 8) Length of headd O.SOt0.04(0.460.56) 0.37t0.03(0.33-0.41) 0.28i0.02(0.260.31) (n = 5) (n = 5) (n = 8) Length of mandiblese 0.38 t 0.04 (0.33-0.46) 0.30 t 0.02 (0.26-0.33) 0.22 t 0.03 (0.18-0.26) Length of flagellum and pedicel 1.1 t 0.10 (0.89-1.22) 0.88 i: 0.07 (0.77-0.99) 0.68 t 0.09 (0.54-0.82) Length of scapef 0.10 i: 0.01 (0.08-0.10) 0.07 t 0.01 (0.06-0.08) 0.05 t 0.01 (0.04-0.06) Length of labial pal$ 0.61 t 0.05 (0.54-0.71) 0.46 ? 0.03 (0.41-0.52) 0.37 t 0.04 (0.31-0.41) Length of distal segment of labial palph 0.39 t 0.04 (0.33-0.48) 0.29 t 0.03 (0.260.36) 0.24 t 0.02 (0.20-0.26) Width of cervixf 0.53 t 0.08 (0.41-0.61) 0.34 t 0.03 (0.29-0.41) 0.21 t 0.05 (0.12-0.29) Length of cervixf 0.48 t 0.12 (0.31-0.66) 0.27 t 0.07 (0.20-0.46) 0.15 t 0.05 (0.07-0.23) Width of prothoraxf 0.84 t 0.14 (0.64-1.1) 0.49 t 0.05 (0.460.39) 0.34 t 0.05 (0.260.43) Width of mesothoraxf 1.2 t 0.24 (0.84-1.3) 0.68 t 0.08 (0.56-0.83) 0.43 t 0.10 (0.27-0.59) Width of metathorad 1.3 tO.26(0.89-1.33) 0.73t0.08(0.59-0.89) 0.45tO.10(0.28-0.60) Length of prothoracic tibia 0.67i:0.04(0.560.71) 0.44t0.03(0.34-0.48) 0.31t0.02(0.28-0.34) Length of prothoracic tarsusi 0.16 t 0.02 (0.14-0.18) 0.15 t 0.01 (0.13-0.17) 0.13 t 0.01 (0.11-0.15)

Measurements are given as mean t SD mm (range). a Number of larvae measured unless indicated below. Tip of mandibles to tip of abdomen. At the level of the eyes, including the eyes. d At the midline. Straight-line distance from tip to medial base. f Medial. g Base of mentum to tip of terminal segment. Base to tip. "ncluding pretarsus, but not pulvillus. ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA Vol. 89, no. 2

Fig. 3. Diagram of setation on M. subanticus 3rd-instar cervix (Cer) and thorax (TlLT3). Spiracle, spr; sclerite, scl; coxa, L. very similar (as in Figs. 1, 3, and 4); M, tasmniae Description of M. subanticus Larvae also appears to share this pattern (New and Boros 1983). However, as stated above, the chaetotaxy of Third Instar. Specimens examined n = 11; the mandibles, legs, and thoracic segments differs measurements in Table 2. Body cream-colored to between species; in general, M. subanticus larvae tan, with head, legs, and posterior region darker. have fewer setae on these structures than do M. Integument covered with minute spinules through- posticus larvae. out. Maxillae. M. subanticus, with 7-8 small setae Head (Figs. 1 and 2). Genae dark tan, marked ventrally; M. posticus, with =12 setae. with purplish pigment posteriorly and posterola- Mesothorax. M. subanticus, with 8 dorsal setae terally in some specimens. Frons tan; clypeus and between spiracles; M. posticus, with 10-12. labrum light tan; epicranial suture white. Venter Prothoracic Tibia. M, subanticus, with =24 se- white medially. Jaws light brown, pale at tips; distal tae; M. posticus, with =36 setae. segment of labial palpi, and distal segments of an- March 1996 KRAKAUERAND

Fig. 4. Diagram of setation on M. subanticus 3rd-instar abdomen (Al-A10). Spiracle, spr; sclerite, scl. tennae light brown, annulated. Stipites and cardi- n = 5). Setation in 3 rows (Fig. 3). Tibiae with 27- nes with dark tan margin. Setation as in Fig. 1. 28 setae, femora with 12-14. Cervix. Measurements in Table 2. Buff with 2 Mesothorax. Dorsum of anterior subsegment pig- longitudinal brown bands. Setae in 2 rows as in mented throughout, darker laterally; posteriol; sub- Fig. 3. segment pale anteriorly, pigmented posterior to Prothorax. Pigmentation limited dorsally to sclerites, except for pale marking along midline and small continuation of lateral bands on cervix; pair ventrally. Dorsal sclerites pale yellow to light tan, of posterolateral markings extending ventrally to sometimes with dark tissue beneath. Setae in 4 ill- legs and anteriorly to dorsal sclerite; pale ventrally. defined transverse rows (Fig. 3). Tibia1 and femoral Dorsal sclerites pale yellow to tan. Basal 3rd of markings and setation as on prothoracic legs. tibia dark (length of tinted portion/total length of Metathorax. Anterior subsegment pigmented as prothoracic tibiae: x = 0.375; range, 0.346-0.414; in mesothorax; posterior subsegment pale. Dorsal 210 ANNALSOF THE ENTOMOLOGICAL SOCIETY OF AMERICA VO~.89, no. 2 sclerites similar to mesothoracic sclerites, with Kawashima, K. 1958. Bionomics and earlier stages of dark tissue beneath. Setation in 4 ill-defined trans- some Japanese Neuroptera (11) Eumicromus nume- verse rows (Fig. 3). Tibia1 and femoral markings rosus (Navis) (Hemerobiidae). Mushi 32: 4346. and setation as on prothoracic legs. Kevan, D. K. McE., and J. Klimaszewski. 1987. The Hemerobiidae of Canada and Alaska. Genus Heme- Al: Dorsum pigmented throughout, Abdomen. robius L. G. Ital. Entomol. 16: 305-369. except pale medially; venter lightly pigmented; Killington, F. J. 1936. A monograph of the British midsegmental white patch laterally. Spiracular per- Neuroptera, vol. 1. Ray Society, London. itremes slightly raised, with basal margins dark 1937. A monograph of the British Neuroptera, vol. 2. brown. Setation in 2 rows (Fig. 4). Ray Society, London. A2-A7: Pigmentation similar to Al, except A3- 1946. On Psectra diptera (Burm.) (Neur., Hemerobi- A7 pale laterally and dorsomedially. Spiracular idae), including an account of its life-history Entomol. peritremes not raised. Setation in 2 transverse rows Mon. Mag. 82: 161-176. (Fig. 4). Posterior row on A7 with 2 pairs long Klimaszewski, J., and D. K. McE. Kevan. 1987a. The pinaculate setae laterally, 1 pair pinaculate setae brown lacewings flies of Canada and Alaska (Neurop- tera: Hemerobiidae). Part II(a). The genus Wesmae- ventrally. lius Kruger. Neuroptera Int. 4: 153-204. A8: Pigmented medially, posteriorly; posterodor- 198713. The brown lacewings flies of Canada and Alas- sal sclerites round, lighc tan to pale yellow. Setation ka (Neuroptera: Hemerobiidae). Part IIb. The genus in 2 transverse rows (Fig. 4). Wesmaelius Kruger. Neuroptera Int. 4: 245-274. A9: Anterior posterodorsal sclerites 1988. The brown lacewing flies of Canada and Alaska tan. Two rows of setae (Fig. 4). (Neuroptera: Hemerobiidae). Part 111. The genus Mi- A10: Pale; dorsal, ventral, dorsolateral sclerites cromus Rambur. G. Ital. Entomol. 19: 31-76. tan. Setation in clusters (Fig. 4). 1992. Review of Canadian and Alaskan brown lace- Second Instar. Measurements on Table 2. Sim- wing flies (Neuroptera: Hemerobiidae) with a key to ilar to 3rd instar, but some small setae on A8 and the genera. Part IV: the genera Megalomus Rambur, Boriomyia Banks, Psectra Hagen and Symperobius A9 absent. Banks. An. Transvaal Mus. 35: 435-457. First Instar. Measurements on Table 2. Bodv MacLeod, E. G. 1960. The immature stages of Bor- white with faint purple or tan transverse bands iomyia jidelis (Banks) with taxonomic notes on the along the posterior margin of each segment, oc- affinities of the genus Boriomyia (Neuroptera: Hem- casionally with pinkish or other colored gut con- erobiidae). Psyche 67: 26-40. tents visible through integument. Setation on head Miller, G. L., and R. D. Cave. 1987. Bionomics of similar to that of 3rd instar, except maxilla with (Walker) (Neuroptera: Hemerobi- only 34 lateral setae, palpiger and palpus each idae) with descriptions of the immature stages. Proc. Entomol. Soc. Wash. 89: 776-789. lacking.0 one seta. A2-A9 similar to 3rd instars. ex- cept some short setae missing; A10 with setae scat- Miller, G. L., and P. L. Lamhdin. 1984. Redescrip- tions of the larval stages of Hemerobius stigma Ste- tered irregularly, not in clusters. The setation on phens (Neuroptera: Hemerobiidae). Fla. Entomol. 67: the thorax and A1 of our 1st-instar specimens was 377-382. difficult to interpret. Monserrat, V. J. 1993. New data on some species of the genus Micromus Rambur, 1842 (Insecta: Neurop- tera: Hemerobiidae). Ann. Mus. Civ. Stori. Nat. Cia- Acknowledgments coma Doria 89: 477-516. Morton, K. J. 1910. Life history of Drepanepteyx We thank J. D. Oswald (University of Texas), Q. D. phalaenoides, Linn. Entomol. Mon. Mag. 8: 54-62. Wheeler and M. J. Tauber (Cornell University) for their Nakahara, W. 1954. Early stages of some Japanese comments on the manuscript, F. Fawcett for help with Hemerobiidae including two new species. Kontyu 21: the drawings, and the National Science Foundation 41-46. (Grant BSR 88-17822), Regional Project 185, and the New, T. R. 1975a. The biology of Chrysopidae and Pew Undergraduate Fellowship Program of the Pew Hemerobiidae (Neuroptera), with reference to their Charitable Foundation, for support. usage as biocontrol agents: a review. Trans. R. Ento- mol. Soc. Lond. 127: 115-140. 197513. The immature stages of Drepanacra binocula References Cited (Neuroptera: Hemerobiidae), with notes on the rela- Carpenter, F. M. 1940. A revision of the Nearctic Hem- tionship of the genus. J. Aust. Entomol. Soc. 14: 247- erobiidae, Berothidae, Sisyridae, Polystoechotidae and 250. Dilaridae (Neuroptera). Proc. Am. Acad. Arts Sci. 4: New, T. R., and C. Boros. 1983. The early stages of 193-280. Micromus tasmaniae (Neuroptera : Hemerobiidae). Dunn, J. A. 1954. Micromus variegatus Fabricus Neuroptera Int. 2: 213-217. (Neuroptera) as a predator of the pea . Proc. R. Oswald, J. D. 1993a. Revision and cladistic analysis of Entomol. Soc. Lond. (Ser. A) 29: 76-80. the world genera of the family Hemerobiidae (Insec- Hoffman, K. M., and J. R. Brushwein. 1992. De- ta: Neuroptera). J. N.Y. Entomol. Soc. 101: 143-299. scriptions of the larvae and pupae of some North 199313. A new genus and species of brown lacewing American Mantispinae (Neuroptera: Mantispidae) from Venezuela (Neuroptera: Hemerobiiidae), with and development of a system of larval chaetotaq for comments on the evolution of the hemerobiid fore- Neuroptera. Trans. Am. Entomol. Soc. 18: 159-196. wing radial vein. Syst. Entomol. 18: 363-370. March 1996 KRAKAUERAND TAUBER:HEMEROBIID LARVAE 211

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Bibliography of the Neuropterida Reference number (r#): 8496

Reference Citation: Krakauer, A. H.; Tauber, C. A. 1996 [1996.??.??]. Larvae of Micromus: generic characteristics and a description of Micromus subanticus (Neuroptera: Hemerobiidae). Annals of the Entomological Society of America 89:203-211.

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