Corresponding author: Catherine A. Tauber (
Academic editor: A. Contreras-Ramos
An expanded list of generic level larval characteristics is presented for
Uma lista expandida das características larvais em nível de gênero é apresentada para
Although most of the taxonomic work on the genus
With the above in mind, we describe and provide images of the larvae of five additional species of
The specimens used in our study were reared from field-collected females. The rearing, preservation, descriptive procedures, and terminology are identical to those published previously (
The earlier study of
In our previous work, we have used two terms “submedian setae” (e.g.,
It is noteworthy that bilateral asymmetry in setal numbers is common, and that specimens occasionally exhibit variation in the numbers and sizes of setae (especially in the dorsal thoracic setae and the submedian abdominal setae) of all instars. The numbers presented here reflect this asymmetry. Also, in our descriptions, unless stated otherwise, all setae, other than the submedian setae, are smooth and pointed (not thorny, hooked or blunt).
The chrysopid life cycle includes a larval stage with three instars. The first instar differs markedly in structure, setation and often coloration from the other two instars, which differ from each other only in minor ways, largely related to size. Thus, for taxonomic purposes, the first instar constitutes Semaphoront A, whereas Semaphoront B includes both the second and third instars and Semaphoront C encompasses all instars (= the larva) (See
Chrysopid genera fall into two general categories: those with “naked” larvae and those with “trash-carrying” larvae.
In general, the species studied here exhibit all of the larval characteristics proposed earlier to typify
Examples of dorsal setae that typify
Note: To identify cephalic setae, see Fig. 5 on page 477 of
1 | Head predominantly brown; epicranial marking entire (with mesal and lateral arms wholly or partly confluent) and fused mesally ( |
2 |
– | Head predominantly white to cream-colored; epicranial marking consisting of two longitudinally elongate brown stripes, with mesal and lateral arms completely separate ( |
3 |
2 | Cranium with two dorsal setae (S1 and S11) thorny (Fig. 5A in |
|
– | Cranium with four dorsal setae (S1, S4, S6 and S11) thorny; dorsum of abdominal segments A1-A5 each with total of 16–20 elongate, smooth, hooked setae (submedian and spiracular setae) ( |
|
3 | Paired frontal marking with posterior ends straight, not meeting mesally ( |
|
– | Paired frontal marking with posterior ends curved inward, meeting mesally ( |
4 |
4 | More than three (usually six) cranial setae (S1, S3, S4, S5, S6, S11) thorny (Fig. 5A in |
|
– | Only two or three cranial setae (S1, S11, sometimes S4) thorny; large lateral tubercles on thorax and abdomen with setae (LS) dark brown ( |
|
1 | Epicranial mark broad, with two arms wholly or partly confluent; paired frontal markings broadly fused mesally ( |
2 |
– | Epicranial mark consisting of two paired longitudinal stripes (= mesal and lateral arms); paired frontal markings close to each other, but separate, except sometimes posterior tips curve and meet mesally ( |
3 |
2 | Mesal and lateral arms of epicranial marking fully confluent throughout ( |
|
– | Mesal and lateral arms of epicranial marking confluent basally, separate distally ( |
|
3 | Paired frontal marking with posterior ends straight, not meeting mesally ( |
|
– | Paired frontal marking with posterior ends curving and meeting mesally ( |
4 |
4 | Cranium with three to four pairs of small secondary setae between S1 and S4 (Fig. 5B in |
|
– | Cranium without secondary setae (Fig. 5A in |
|
Because the Appendix lists the generic-level characteristics that are shared by all
Adults of
Argentina, Brazil, British Guiana, Colombia, Cuba, Jamaica, Paraguay, Peru, Uruguay, and Venezuela (
The first instar of
(Semaphoront A).
Head and thorax, dorsal, first instar
Gena, ventral margin of cranium brown; genal marking with small white spot behind eye. Labial palpus tinged with light brown, slightly darker distally. Mandibles amber, with brown basolateral spot. Antenna with scape brownish, pedicel white basally, brown distally, flagellum light brown.
T1: Row of three very small setae (R1) at anteromesal base of LTs. Sc1 brown mesally, basolaterally, with cream colored interior; S2Sc1 small, immediately above S1Sc1. S1 long, S3 intermediate-length. T2: Spiracle with lips of atrium protruding above integumental surface. Sc1, Sc2, transparent; Sc3 marked with light brown; S2Sc3 variable, medium-length to long; S2 smaller. T3: S1Sc1, S2Sc1 present, S1Sc2 very small, sometimes absent, S2Sc2 usually present. Posterior fold with row (R1) of six (sometimes seven) long, thorny, pointed setae on chalazae with ovate, light brown marks anteriorly; lateral two chalazae on each side juxtaposed.
A1: Dorsum with ten to twelve SMS in single anterior row, with 34–42 SMS in double to triple row posteriorly; spiracles at end of posterior row, without distinguishable SSp. A2: Dorsum with ~10 SMS in single anterior row, with ~44 SMS in double to triple posterior row; anterior row bending posteriorly at each end, coalescing with setae in posterior row; spiracle at end of posterior row, with SSp near anterodistal margin. A3-A5: Dorsum with ~8 (A3, A4) or 6–7 (A5) SMS in single anterior row, with ~36 (A3), ~32 (A4), ~26 (A5) SMS in double to triple posterior row; anterior row bending posteriorly at each end, coalescing with posterior row; spiracle with SSp anteroventrally. A6: Anterior region with four SMS; spiracle with SSp mesally. A7: Anterior region with pair of microsetae; spiracle with SSp mesally. A8: Venter with two pairs of medium-length setae posteriorly, one pair of short setae slightly anteriorly.
(Semaphoront B).
Head and thorax, dorsal, second instar
Head and thorax, dorsal, third instar
All cephalic setae smooth, pointed; S1 slightly robust, medium length, S11 long, S2–10, S12 short to medium length; Vx setae fairly long, robust; with one to two pairs of secondary setae. Anterior margin of head protruding, straight with angled lateral margins; mesal pair of anterior setae much longer than two lateral pairs.
Head width across eyes, 0.53–0.59 mm (L2), 0.78–0.83 mm (L3); mandible length, 0.52–0.56 mm (L2), 0.81–0.90 mm (L3); ratio mandible length to head width = 0.92–1.0 : 1 (L2), 1.0–1.1 : 1 (L3). Tip of mandible with six teeth mesally.
Cervix brownish dorsally, slightly darker laterally; venter cream-colored mesally, light brown laterally; white ventrally.
T1: LT with 14–17 (L2), 17–19 (L3) LS. Sc1 large, rhomboid, extending around posterior base of LT, darker brown laterally than mesally. Sc2 triangular, appearing as paired brown marks, with two small, irregularly shaped sclerites (brown) above; sometimes small brown spots anteriorly. S2, S3 smooth. T2: Anterior sclerite (Sc1) light brown to brown; spiracles on prominent protuberances. Posterior subsegment with Sc2 light brown; Sc3 pronounced, brown. LT with 12–13 (L2), 16–19 (L3) LS. T3: Sc1 transparent. LT with 11–13 (L2), 13–18 (L3) LS. Posterior fold with 14 to 15 robust, thorny setae; L3 sometimes with additional pair of long, smooth, pointed setae laterally, arising from smaller chalazae.
A1: Dorsum with ~78–88 (L2), 142–172 (L3) SMS in two double-triple transverse bands between spiracles. A2-A5: Dorsum with 64–112 (L2), 120–196 (L3) SMS in two broad transverse bands. LTs each with 11–21 (L2), 29–40 (L3) LS: one to five robust, thorny, blunt to spatulate LS on distal surface; remaining LS long, smooth, hooked, in large patch on dorsal surface. A6: Dorsum with transverse band of 20–30 (L2), 56–66 (L3) SMS across anterior of segment; midsection with one to two pairs of smooth setae, mesal pair hooked (similar to SMS), lateral pair short, pointed. LT with nine to eleven (L2), ten to eleven (L3) LS of various sizes. A7: Dorsum with one to two pairs of very small setae (S1, S2) anteriorly, between spiracles; LDTs each with one medium-length, robust, thorny, blunt to spatulate LDS, one to two smaller, smooth, pointed LDS; pair of very small setae between LDTs. LT with ten to 13 (L2), ten to 14 (L3) LS of various sizes. A8: Anterior region with one to two pairs of very small setae (S1, S2). Venter with pair of medium-length setae between LTs, two to three smaller setae slightly anteroventral to LTs. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14–16 short to medium-length setae, several in each ring robust. A10: Dorsum with several pairs of small setae on V-shaped anterior sclerites, one slightly anterior to terminus. Two pairs of robust lateral setae. Venter with ~five pairs of small setae in V-shaped pattern, posterior row of microsetae anterior to terminus.
At oviposition, light yellowish green to green, with white micropyle; ovoid, 0.82 to 1.06 mm long, 0.36 to 0.43 mm wide. Stalk smooth, hyaline, 2.89 to 6.80 mm long.
Numerous lots, each originating from a single gravid female collected in
Adults and larvae of this species were collected on shrubs in disturbed, dry forest habitats. Adults are agile; they exhibit fast, evasive flight, usually inward toward the interior of the bush or tree.
Based on the following observations, we think that
In the lab, eggs from all the lots listed above were deposited separately (with isolated stalks), in small groups with no particular pattern; the stalks were sticky, but without droplets. During the first 24 hours after oviposition, the eggs were bright yellowish green to green, without spots. On the second day, they began to develop a bluish brown color, with grey or brownish mottling which became more pronounced as hatching approached (
In one case, a small proportion (n = 2 of 28) of the eggs laid by a female from Minas Gerais had a prolonged incubation period (approximately one week to ten days longer than the usual six- to eight-day incubation period). They were a dark bluish brown color during the period of delay. The cause of the prolongation is unknown; however, the resulting larvae developed normally and appeared healthy.
Larvae of
Adults of this recently described species are readily distinguished from other
Brazil, Venezuela (
Like the larvae of
(Semaphoront A).
Gena, ventral region of head capsule brown, with white spot in anterior region of genal mark. Labial palpus white, marked with light brown on basal segment, distal two annuli of middle segment, base of terminal segment. Mandibles amber to light brown. Antenna with light brown scape, pedicel white basally, light brown distally; flagellum light brown.
T1: Row of three very small setae (R1) at anteromesal base of LTs not observed. Sc1 with edges brown, center white; S2Sc1 very small, immediately above S1Sc1. S1, S3 intermediate-length, robust. T2: Spiracle with lip of atrium protruding above integumental surface. Sc1, Sc2, Sc3 light brown; S2Sc3, S2 long, robust, thorny, of approximately equal size. T3: S1Sc1, S2Sc1, S1Sc2 very small; S2Sc2 absent. Raised posterior fold with row (R1) of four very long, robust, thorny setae on chalazae with ovate, light brown marks anteriorly.
A1-A5: Dorsum with 12 SMS in two rows, with four mesal pairs equally spaced, two lateral-most pairs juxtaposed; spiracle with SSp elongate, robust, mesal to spiracle; chalazae of all SMS and SSp large, robust. A6: Anterior region with two SMS. Spiracle with very small SSp mesally. A7: Anterior region without setae or microsetae. Spiracle without apparent SSp. A8: Venter with one pair of medium-length setae posteriorly.
(Semaphoront B).
All cephalic setae smooth, pointed; S1-10, S12 of medium length, S11 long; Vx setae fairly long, robust.
Head width across eyes, 0.57–0.58 mm (L2), 0.77–0.85 mm (L3); mandible length 0.51–0.57 mm (L2), 0.82–0.85 mm (L3); ratio mandible length to head width = 1.0–1.1 : 1 (L2), 0.97–1.1 :1 (L3). Tip of mandible with four teeth mesally.
Cervix cream-colored, tinged with light brown dorsally; sides with pair of broad brown patches; venter cream-colored mesally, light brown laterally.
T1: LT with 14–18 (L2), 16–20 (L3) LS. Sc1 large, rhomboid, extending around posterior base of LT, darker brown laterally than mesally. Sc2 triangular, mostly brown, with small secondary sclerite above. Notum with four brown spots in longitudinal row along midline. S2, S3 thorny. T2: Anterior sclerite (Sc1) light brown; spiracles on small protuberances. Posterior subsegment with Sc2 light brown; Sc3 pronounced, brown. LT with 13–17 (L2), 16–20 (L3) LS. T3: LT with 11–15 (L2), 14–18 (L3) LS. Posterior fold with row (R1) of twelve robust, thorny setae.
A1: Dorsum with ~54–59 (L2), ~90–94 (L3) SMS in two double-triple transverse bands between spiracles. A2-A5: Dorsum with 42–86 (L2), 100–166 (L3) SMS in two broad transverse bands. LTs each with 9–13 (L2), 13–20 (L3) LS; apical four to eight LS long, robust, thorny, pointed to blunt; remaining LS less robust, smooth, hooked, in patch on dorsal surface. A6: Dorsum with transverse band of ~26 (L2), ~45 (L3) SMS across anterior of segment; midsection with two pairs of smooth setae, mesal pair hooked, lateral pair pointed. LTs with ~9 (L2), ~15 (L3) LS of various sizes. A7: Dorsum with two pairs of very short setae (S1, S2) anteriorly, between spiracles. LDTs each with one medium-length, robust, thorny, blunt to spatulate LDS, one to two shorter, thorny, robust LDS, one to three small, smooth, pointed LDS. LTs with ~8 (L2), ~11 (L3) LS of various sizes. A8: Dorsum with two to three pairs of very small setae anteriorly; two to three pairs of small setae posteromesal to spiracles; four pairs of small, posterior setae in transverse row mesal to LTs. Venter with three transverse rows of setae, each with three to four smooth, pointed setae of increasing size posteriorly. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~16–18 short to medium-length setae, several in each ring robust. A10: Dorsum with two pairs of small setae: one posterior to V-shaped anterior sclerites, one slightly anterior to terminus. Two pairs of lateral setae, robust. Venter with ~five pairs of small setae, posterior row of microsetae anterior to terminus.
At oviposition, light green, with white micropyle; ovoid, 0.99 to 1.08 mm long, 0.40 to 0.41 mm wide. Stalk smooth, hyaline, 4.39 to 5.42 mm long.
Several lots, each originating from a single gravid female collected in
Adults and larvae of
Among
Brazil (
(Semaphoront A).
Gena brown with small central white spot, ventral region of head capsule white to cream-colored, with ventral margin of head capsule brown. Labial palpus white, with tinge of light brown on basal segment, lateral side of middle segment, darker brown on distal annulation, terminal segment. Mandibles amber to light brown, with dark brown basally. Antenna with light brown scape, pedicel light brown basally, brown distally; flagellum brown. Cervix with pair of light brown sublateral spots.
T1: Row of three very small setae (R1) at anteromesal base of LTs present. Sc1 with distal section light brown, center white; S2Sc1 very small, immediately above S1Sc1. S1 intermediate-length; S3 intermediate-length to short, thorny. T2: Spiracle with lip of atrium protruding slightly beyond level of integumental protuberance. Sc1, Sc2, Sc3 light brown; S2Sc3, S2 small to intermediate length, thorny, of approximately equal size. T3: S1Sc1, S2Sc1, S1Sc2 very small; S2Sc2 absent. S1, S2 absent. Raised posterior fold with row (R1) of four very long, robust, thorny, pointed setae on chalazae with ovate, light brown marks anteriorly.
A1-A5: Dorsum with 12 SMS in two rows, with lateral-most two pairs juxtaposed. Spiracle with SSp elongate, robust, mesal to spiracle. A6: Anterior region with two SMS. Spiracle with SSp mesally. A7: Anterior region without setae or microsetae. Spiracle anterior to LT, without apparent SSp. A8: Venter with two pairs of medium-length setae posteriorly.
(Semaphoront B).
Cephalic seta S1 moderately long, thorny, S2-S12 smooth, S11 long; Vx setae relatively long.
Head width across eyes, 0.64–0.66 mm (L2), 0.98–1.05 mm (L3); mandible length, 0.51–0.58 mm (L2), 0.91–0.98 mm (L3); ratio mandible length to head width = 0.82–0.87 : 1 (L2), 0.93–0.99 : 1 (L3). Tip of mandible with four teeth mesally.
Cervix: dorsum cream-colored, tinged with light brown; sides with pair of broad brown patches; venter brown laterally, becoming cream-colored mesally.
T1: LT with 15–19 (L2), 18–22 (L3) LS. Sc1 with elongate brown marks laterally, mesally, cream-colored streak centrally; two heavily sclerotized rods extending from base of sclerite, mesal one stretching along mesal margin of Sc1 to mesal base of LT, lateral one forked distally, with mesal fork extending onto posterolateral margin of LT, lateral fork extending laterally below posterior base of LT. Sc2 triangular, included in elongate mesal brown mark, with two small, brown, secondary sclerites anteriorly. S2, S3 thorny. T2: Anterior sclerite (Sc1) brown; spiracles on small protuberances. Posterior subsegment with Sc2 light brown; Sc3 pronounced, brown. LT with 15–19 (L2, L3) LS. T3: LT with 14–18 (L2, L3) LS. Posterior fold with row (R1) of twelve robust, thorny setae.
A1: Dorsum with 62–76 (L2), ~ 156–188 (L3) SMS in two double-triple transverse bands between spiracles. A2-A5: Dorsum with 62–114 (L2), 176–268 (L3) SMS in two broad transverse bands. LTs each with 9–14 (L2), 12–34 (L3) LS: four to eight long, robust, thorny, pointed to spatulate LS on distal surface; remaining LS less robust, smooth, hooked, in patch on dorsal surface. A6: Dorsum with transverse band of 22–32 (L2), 42–56 (L3) SMS across anterior of segment; midsection with two pairs of smooth setae, mesal pair hooked, lateral pair pointed. LT with 8–10 (L2), ~11–14 (L3) LS of various sizes. A7: Dorsum with three pairs of very short setae anteriorly, between spiracles. LT with ~8–10 (L2), 11–14 (L3) LS of various sizes. A8: Dorsum with two pairs of very small setae between spiracles; four pairs of small setae in transverse row between LTs. Venter with two transverse rows of setae, each with three to four smooth, small, pointed setae. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14–16 short to medium-length setae, several in each ring robust. A10: Dorsum with two pairs of small setae: one posterior to V-shaped anterior sclerites, one slightly anterior to terminus. Two pairs of lateral setae, robust. Venter with ~five pairs of small setae, posterior row of microsetae anterior to terminus.
Green, ovoid, 0.95–1.00 mm long; 0.43–0.45 mm wide; stalk hyaline, 10.7–12.6 mm long.
Several lots, each originating from a single gravid female collected in
Eggs of
Argentina (
The dorsal head markings of
(Semaphoront A).
Gena, ventral margin of head capsule brown, with small clear patch within genal mark. Labial palpus tinged with light brown, slightly darker distally. Mandibles light brown. Antenna with light brown scape, pedicel white basally, brown distally; flagellum brown.
T1: Row of three small setae (R1) near base of LT not observed. Sc1 brown throughout, but slightly lighter mesally; S2Sc1 small, immediately above S1Sc1. S1 long; S3 intermediate-length, slightly thorny. T2: Spiracle with lip of atrium raised above integumental surface. Sc1, Sc2 transparent: Sc3 marked with light brown; S2Sc3 variable, from medium-length to long, S2 smaller than S2Sc3. T3: S1Sc1 present, S2Sc1 (sometimes absent), S1Sc2 very small. S2Sc2 absent. Raised posterior fold with row (R1) of four long, thorny, pointed setae on chalazae with ovate, light brown marks anteriorly.
A1-A2: Dorsum with four to six SMS in anterior row, with 12 to 16 SMS in posterior row. Spiracles far lateral to anterior row, with SSp mesally. A3-A5: Anterior and posterior rows of SMS largely coalesced, with four to six SMS in anterior row, ten to 14 SMS in curved, posterior row. Spiracle with SSp near anteromesal margin. A6: Anterior region with row of two to six SMS, pair of small, straight setae lateral to SMS. Spiracle with pair of small SSp mesally. A7: Anterior region without setae or microsetae. Spiracle with SSp mesally. A8: Venter with two pairs of medium-length setae posteriorly, one pair of short setae anteriorly.
(Semaphoront B).
All cephalic setae present; S1, S11 long; S2-S10, S12 medium length to short; S1 thorny; S4, S6, S11 lightly thorny (thorniness difficult to see except under high magnification, especially on L2); other setae smooth. Vx setae relatively short. Anterior margin of head straight, with angled lateral margins; mesal pair of anterior setae medium-length, lateral two pairs short or very short.
Head width across eyes, 0.55–0.61 mm (L2), 0.91–0.96 mm (L3); mandible length, 0.49–0.55 mm (L2), 0.96–1.00 mm (L3); ratio mandible length to head width = 0.87–0.93 (L2), 1.02–1.07 (L3). Tip of mandible with six teeth mesally.
Cervix brownish, with dense covering of spinules; sides with pair of broad brown patches; venter brown throughout, darker laterally.
T1: LT with 15–19 (L2), 18–24 (L3) LS. Sc1 extending around posterior base of LT, dark brown laterally, mesally, with cream-colored to light brown, elongate streak with broad base. Three sclerites on midline: Sc2 triangular, cream-colored, with two brown spots anteriorly, mesal brown spot posteriorly, two smaller, light brown sclerites anterior to Sc2 [not all apparent on L2]. S2, S3 thorny. T2: Anterior sclerite (Sc1) brown; spiracle on small protuberance. Posterior subsegment with Sc2 transparent; Sc3 pronounced, dark brown. LT with 12–16 (L2), 14–18 (L3) thorny, robust LS, most long, several basal ones shorter. T3: LT with 12–16 LS (L2, L3). Posterior fold with 12–13 robust, thorny setae.
A1: Dorsum with 40–76 (L2), ~128–150 (L3) SMS in two double-triple transverse bands between spiracles; bands not extending laterally beyond spiracles. A2-A5: Dorsum with 41–102 (L2), 119–173 (L3) SMS in two broad transverse bands. LT with 10–15 (L2), 18–30 (L3) LS: five to seven robust, thorny, blunt LS of various lengths, on distal surface; remaining LS less robust, smooth, hooked in patch on dorsal surface. A6: Dorsum with transverse band of 16–28 (L2), 28–38 (L3) SMS across anterior of segment; midsection with two pairs of smooth setae, mesal pair hooked, long, lateral pair pointed, shorter. LT with 9–11 (L2), 11–17 (L3) robust, thorny, blunt LS of various sizes. A7: Dorsum with two pairs of short setae between spiracles. LT with 8–10 (L2), 10–13 (L3) LS of various sizes. Venter with setal number and size variable, usually with two pairs of setae anteriorly (both small, smooth), three pairs of sublateral setae posteriorly (two lateral-most pairs robust, thorny, mesal pair small, smooth). A8: Dorsum with two pairs of small setae between spiracles; four pairs of small setae in transverse row between LTs. Venter with scattered, small setae anteriorly, two robust, thorny setae on small chalazae between LTs. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14–16 setae of various sizes, several in each ring robust. A10: Dorsum with one pair of setae anteriorly, two pairs mesally, patch of several setae distally; one pair of small setae posterior to V-shaped anterior sclerites. Lateral region with two pairs of robust setae, two to three pairs of smaller setae. Venter with ~two pairs of robust setae, five pairs of small setae.
At oviposition, green, with white micropyle; ovoid, 0.92 to 0.99 mm long, 0.40 to 0.44 mm wide. Stalk smooth, hyaline, 3.1 to 6.2 mm long.
Several lots, each originating from a single gravid female collected in
Adults of
In the lab, eggs were deposited separately (with isolated stalks), in small groups with no particular pattern. During the first 24 hours of oviposition, the eggs were bright green, with dark green blotches. On the second day, they began to develop a bluish brown tone, with brownish mottling; by the third day the eggs were greyish blue to pinkish, with brown mottling. At 24 ± 1°C, hatching occurred within six days (n = 12).
Larvae of
Experimental life history studies of
Although the female and male genitalia of
Brazil (
Like the larvae of
(Semaphoront A).
Gena, ventral margin of head capsule mostly white, with light brown genal mark posterior to eye, with clear spot near anterior margin. Labial palpus mostly white, middle segment tinged with light brown laterally, distal segment mostly tinged with brown. Mandibles amber to light brown, with dark brown basolateral spot. Antenna with scape light brown, pedicel white basally, brown distally; flagellum light brown.
T1: Row of three very small setae (R1) at anteromesal base of LTs. Sc1 with scattered brown areas, especially laterally; S2Sc1 small, immediately above S1Sc1. S1, S3 intermediate-length. S2, S3 thorny. T2: Spiracle with lip of atrium flush with level of integumental surface. Sc1, Sc2 transparent; Sc3 light brown; S2Sc3 medium-length to long, slender, S2 shorter. T3: S1Sc1, S2Sc1 usually present, S1Sc2 very small; S2Sc2 absent. Raised posterior fold with row of four robust, thorny, pointed setae on chalazae with ovate, light brown marks anteriorly.
A1-A5: Dorsum with 12 SMS in two rows, with four mesal pairs equally spaced, two lateral-most pairs juxtaposed; spiracle with SSp elongate, robust, mesal to spiracle; chalazae of all SMS and SSp not large. A6: Anterior row with two SMS, pair of small, straight setae lateral to SMS; spiracle at anterior base of LT, without apparent SSp. A7: Anterior region without setae or microsetae. Spiracle without apparent SSp. A8: Venter with two pairs of medium-length setae posteriorly.
(Semaphoront B).
Cephalic seta S1 moderately long, thorny, S2-S12 smooth, only S11 long; Vx setae moderately long; three to four pairs of small secondary setae between S1 and S4.
Head width across eyes, 0.5–0.6 mm (L2), 0.84–0.86 mm (L3); mandible length, 0.54–0.57 mm (L2), 0.86–0.90 mm (L3); ratio mandible length to head width = 0.91–0.99 : 1 (L2), 1.00–1.05 : 1 (L3). Tip of mandible with six teeth mesally.
Cervix cream-colored, tinged with light brown; sides with pair of broad brown patches; venter brown laterally, becoming cream-colored mesally; with three pairs of small setae ventrally.
T1: LT with 16–17 (L2), 17–19 (L3) LS; five to six short, smooth setae anterobasally. Sc1 large, extending up mesal base of LT, light brown mesally, transparent laterally. Sc2 triangular, light brown; without secondary sclerites. S2, S3 thorny. T2: Sc1 light brown; spiracle on small protuberance. Posterior subsegment with Sc2 transparent to very light brown; Sc3 pronounced, brown. LT with 12–13 (L2), 17–19 (L3) LS. T3: LT with 11–13 (L2), 16–18 (L3) LS. Posterior fold with ten to twelve robust, thorny setae.
A1: Dorsum with 40–56 (L2), ~116–124 (L3) SMS in two double-triple transverse bands between spiracles. A2-A5: Dorsum with 66–84 (L2), 134–174 (L3) SMS in two broad transverse bands. LTs each with 8–11 (L2), 11–21 (L3) LS: four to nine long, robust, thorny, usually pointed LS on distal surface; remaining LS less robust, smooth, hooked in patch on dorsal surface. A6: Dorsum with transverse band of 16–18 (L2), 44–58 (L3) SMS across anterior of segment; midsection with two pairs of smooth setae, mesal pair long, hooked, lateral pair short, pointed. LT with 7–8 (L2), ~14 (L3) LS of various sizes. A7: Dorsum with three pairs of very short setae anteriorly, between spiracles. LT with 6–7 (L2), 9–12 (L3) LS of various sizes. A8: Dorsum with three pairs of very small setae between spiracles; three pairs of small setae in transverse row between LTs. Venter with four transverse rows of setae, each with three to four smooth, small to medium-length, pointed setae. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14–16 short to medium-length setae, several in each ring robust. A10: Dorsum with one pair of small setae posterior to V-shaped anterior sclerites. Several pairs of lateral setae. Venter with ~five pairs of small setae, posterior row of microsetae anterior to terminus.
At oviposition, green, with white micropyle; ovoid, 0.92 to 0.97 mm long, 0.42 to 0.44 mm wide. Stalk smooth, hyaline, 8.8 to 10.1 mm long.
Several lots, each originating from a single gravid female collected in
The thermal influence on rates of development and reproduction in
We thank the following who assisted with obtaining specimens: V. Becker, E. M. G. Fontes, F. Franca, S. L. Lapointe, J. S. Multani, A. Nascimento, C. S. S. Pires, E. A. Silva, B. Souza, E. R. Sujii, A. J. Tauber, and P. J. Tauber. CAT and MJT acknowledge L. E. Ehler and M. Parella for their cooperation in a variety of ways.
Our project is long-standing; it is a pleasure to acknowledge the support from the following agencies and institutions: the USDA/NRI (Competitive Grant 9802447, MJT, CAT), the National Geographic Society (MJT, CAT, GSA), the National Science Foundation (Grants INT-9817231, DEB-0542373, MJT, CAT), the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil – Grants 300504/96-9, 466439/00-8, 475848/04-7, 484497/07-3, GSA), Regional Project W-1385, Cornell University, and the Universidade Estadual do Norte Fluminense.
Characteristics of
(1) setaeof four types
(a) smooth, pointed, short to long [most cephalic setae, some setae on abdominal segments A6-A10, some ventral setae, all small setae] (Fig. 1)
(b) thorny, pointed to blunt or slightly spatulate, medium length to long [some cephalic setae, most setae on thoracic lateral tubercles (LS), some primary thoracic setae, some LS on A2-A6, some setae on laterodorsal tubercles of A6, A7 (LDS), some ventral setae] (Fig. 1)
(c) smooth, hooked, long (some LS on A2, A3 of L1, some LDS on A6, A7 of L3)
(d) smooth, arising from brown chalazae, with elongate, narrow stem, laterally flattened terminal hook [abdominal submedian setae (SMS) on A1-A5, anterior region of A6] (Fig. 1)
(2) setal bases usually brown
(3) spiracles simple (with cylindrical sides, atrium not enlarged), with brownish margins
(4) mandible with four to six small terminal “teeth” (serrations)
(5) mandibular base with one seta and dorsal row of four placoid organs (Rousset 1966)
(6) stipes elongate, narrow, folded inward on longitudinal axis
(7) stemmata black
(8) epicranial (mesal and lateral), postfrontal, frontal markings present
(9) genal marking present, forked anteriorly or, more usually, with clear spot near anterior margin
(10) labrum with three pairs of setae (mesal pair long, lateral pairs shorter)
(11) all primary setae, including S12 present, S1, S11 long
(12) short, stout seta laterally on tip of pedicel
(13) long terminal seta on tip of flagellum
(14) lateral tubercles (LTs) about two times longer than broad
(15) primary setae straight, with pointed tips
(16) secondary setae absent
(17) paired lateral sclerites (Sc1) large, rhomboid, marked with brown, with associated seta S1Sc1 relatively long, thorny, stemming from brown chalaza
(18) primary seta S1 thorny, stemming from brown chalaza, S5 absent
(19) venter with one pair of very small setae mesal to coxae (minute on L1)
(20) *segment divided into two distinct subsegments: anterior subsegment with single fold (= “lobe” of Snodgrass 1935), posterior subsegment consisting of two transverse folds, separated by subtle transverse groove (folds on posterior subsegment difficult to discern on L1)
(21) anterior subsegment bearing pair of small sclerites (Sc1) anteriorly, pair of spiracles (Sp) laterally
(22) *posterior subsegment: anterior fold bearing large tubercles (LTs) laterally, pair of small sclerites (Sc2) anteriorly; posterior fold bearing large, usually brown sclerite (Sc3) on anterolateral margin (posteromesal to LTs); Sc3 with one small seta (S1Sc3) and one longer, thorny seta (S2Sc3)
(23) spiracles located on rounded, lateral protuberances, without associated setae (SSp)
(24) venter with one pair of very small setae on anteromesal margin
(25) *segment with three transverse folds; anterior two folds separated by subtle, transverse groove; dorsal surface of both folds level with mesothorax; dorsum of posterior fold prominently raised above anterior two folds, separated from anterior folds by subtle groove
(26) S1, S2 absent
(27) posterior fold with transverse row (R1) of long, robust, thorny setae arising from large chalazae, each with brown, teardrop-shaped mark on anterior surface
(28) *each transverse fold with pair of small, transparent sclerites (Sc1, Sc2, Sc3); Sc1, Sc2 (when visible) located on anterior margin of fold; Sc3 (when visible) nestled in anterolateral corner of groove above R1; Sc2, Sc3 sometimes difficult to discern (L1, L2)
(29) venter with pair of very small setae on anteromesal margin
(30) overall shape gibbous
(31) depth of A2-A5 greater than depth of thorax, width not greater than thorax
(32) A6-10 becoming narrow and tubular distally; segments not withdrawn
(33) *terminus with pair of eversible, membranous pouches (small on L1)
(34) *A1-A5 each with two, ill-defined, transverse folds separated by small grooves; each fold bearing transverse row of SMS
(35) *A6-A10 without folding; A6 with row of SMS anteriorly, without SMS posteriorly; A7-A10 without SMS
(36) A1 without pleural or ventral region; without lateral tubercles (LTs) or laterodorsal tubercles (LDTs)
(37) A2-A7 with LTs robust
(38) A8 with LTs round, fleshy
(39) A2-A5, A8-A10 without LDTs
(40) A6, A7 each with pair of LDTs
(41) A1-A8: spiracles (Sp) sessile
(42) A1, A6, A7: Spiracular setae (SSp) small, smooth, pointed (when present); *A2-A5 with SSp indistinguishable from SMS; A8 without SSp
(43) dorsal setae S1, S2 absent from all segments
(44) *venter with following setae: A2-A7 each with pair of sublateral setae and pair of submedian setae. Sublateral setae short, smooth, extending from small chalazae on segments A2, A3, becoming larger, smooth or thorny, stemming from larger chalazae or small tubercles on segments A4, A5, stemming from small tubercles with at least one other small seta on segments A6, A7. Submedian setae long, smooth, slender on A2, A3, short on A4-A7. Note: Occasionally, one or more of the above setae may be absent, or additional ones may be present.
(1) *stemmata black, with integument between all stemmata brown to black
(2) *exterior margin of stemmata with thin, brown border
(3) antenna: scape without setae
(4) mandible with four terminal “teeth” (serrations), without basal setae
(5) labial palpus: distal margin of basal segment with two setae: one lateral, one mesoventral; middle segment with two distal setae; distal segment without setae
(6) palpiger with two pairs of distal setae
(7) mentum with two pairs of anterior setae, one pair of posterior setae
(8) lateral tubercle (LT) with two LS; one microseta between LS
(9) paired lateral sclerites (Sc1) extending anteriorly to base of LT
(10) posteromesal sclerite (Sc2) not discernible
(11) primary setae S1, S3 intermediate-length to long, thorny; S2 short, adjacent to S1Sc1, S4 absent
(12) LT with three LS; no microseta
(13) *Sc1 with two small associated setae (S1Sc1, S2Sc1), one on each side; S3Sc1 absent. Sc2 with two small associated setae (S1Sc2, S2Sc2) laterally; S3Sc2 absent
(14) primary setae: S1 short to minute or absent, S2 intermediate-length to long, thorny
(15) *posterior fold with four to seven (perhaps eight) R1 setae
(16) *Sc1 with one to two small associated setae (S1Sc1, S2Sc1) (sometimes absent)
(17) Sc2 difficult to discern, sometimes with small associated seta (S1Sc2)
(18) Sc3 with one small associated seta (S1Sc3), sometimes absent
(19) primary setae S1, S2 not discernible
(20) *A1-A5 each with two primary rows of SMS, sometimes lateral two SMS juxtaposed
(21) A6 with anterior area bearing single transverse row of SMS; posterior area without SMS
(22) A2-A7: LT robust basally, tapering, sclerotized distally, bearing two long, robust setae (LS)
(23) *A2-A3: LT with one thorny, pointed LS, one smooth, hooked LS
(24) A4-A7: LT with two thorny, pointed to blunt LS
(25) A8: LT with one medium-length, thorny, pointed LS, one short, smooth LS
(26) A6, A7: LDT bearing one long, robust, thorny LDS, one short, thorny LDS (A6) or two small LDS (A7)
(27) *A6, A7: with pair of small setae between LDTs
(28) *A1: spiracle with or without SSp
(29) *S2-A5 with single pair of smooth, hooked SSp
(30) *A6, A7 with small, smooth, pointed SSp present or absent
(31) A8 without SSp
(32) *A9: anterior section with two pairs of robust, thorny, pointed to blunt setae dorsally, one pair laterally
(33) A9: posterior section with one pair of robust, thorny setae dorsally, one pair laterally, three pairs ventrally
(34) *A10: with scattered small setae, microsetae
(1) dorsal and ventral surface with rounded, brownish spinules
(2) spinules generally pointed, without microsetae
(3) *integument surrounding three anterior and central stemmata brown
(4) *integument around outer margin of posterior two stemmata unmarked
(5) scape with robust seta on distolateral margin
(6) tip of pedicel with relatively long, stout lateral seta
(7) terminal flagellar seta long
(8) basal segment of labial palpus with one small, dorsal seta, one medium-length, lateral seta, one long, ventral seta
(9) middle segment of labial palpus (L3) with ~five annulations, eight setae (three near apex of segment)
(10) middle segment of labial palpus (L2) with ~four annulations, six setae (three near apex of segment)
(11) distal segment of labial palpus without setae, with small rings throughout; ventral surface with two small, longitudinal sensory organs
(12) palpiger with two intermediate-length setae
(13) mentum (L3) with two pairs of long setae anteriorly below palpiger, one pair of long setae laterally above stipes, one pair of medium-length setae posteriorly
(14) mentum (L2) with two pairs of anterior setae, one pair of posterior setae
(15) LTs well sclerotized, papiliform, with globose terminus
(16) paired lateral sclerites (Sc1) extending well onto the base of LT
(17) posteromesal sclerite (Sc2) present, marked with brown, triangular, smaller than Sc1
(18) *secondary sclerites present or absent
(19) S2Sc1 present, short
(20) primary setae S1 thorny, S2, S3 smooth or thorny, S4 short, smooth
(21) row of three small setae (R1) usually on anterolateral margin of notum (sometimes obscured beneath cervical fold)
(22) Sc1 with two (L2) to three (L3) small associated setae – one mesal (S1Sc1), two lateral (S2Sc1, S3Sc1-L3 only)
(23) Sc2 with two very small associated setae, both lateral (S1Sc2, S2Sc2)
(24) *Sc3 with two associated setae – S1Sc3 short, smooth, S2Sc3 intermediate-length to long, thorny
(25) *primary setae S1, S2 smooth, short to medium-length, anterior to Sc3; S3 robust, thorny, intermediate length to long, usually located in transverse row with S2Sc3; S4 small when present, often absent
(26) *Sc1, Sc2, Sc3 each with pair of sclerites bearing one to two small, associated setae (S1Sc1, S2Sc1, S1Sc2, S2Sc2, S1Sc3 sometimes absent)
(27) posterior fold with ten to 16 robust, thorny R1 setae
(28) *A1-A5: dorsum with transverse folds spinule-covered, separated by shallow groove without spinules
(29) each fold bearing double or triple transverse band of submedian setae (SMS)
(30) A2-A7: LTs hemispherical to slightly flattened apically, with brown marks anteriorly and posteriorly (sometimes light, sometimes absent from L2)
(31) *A2-A5: LTs each with one to eight long, thorny, robust, blunt to spatulate LS apically (size, thorniness, robustness of these LS increasing posteriorly), with subapical dorsal setae long, smooth, hooked, similar to SMS
(32) A6, A7: all LS long, thorny, robust, blunt to spatulate, restricted to apical region of LT
(33) A8: LT bearing one medium-length, thorny, pointed seta (LS), one or more short, smooth LS
(34) A6, A7 with LDTs, each bearing one to two long, robust, pointed or blunt (not hooked), thorny setae (LDS) stemming from large, brown chalazae, sometimes a long, smooth, hooked LDS, and three to four smaller, smooth or thorny LDS
(35) A6, A7 with pair of short, pointed setae between LDTs
(36) A1-A5 with patches of setae near or around each spiracle [These setae are indistinguishable in size and shape from SMS, and the patches often merge with the anterior rows of SMS. Thus, here, they are included with the SMS.]
(37) A6, A7, A8 with single pair of small, pointed SSp