Immature stages of Calycopis bellera (Hewitson) and C. janeirica (Felder) (Lepidoptera, Lycaenidae, Theclinae, Eumaeini): Taxonomic significance and new evidence for detritivory
Author
Duarte, Marcelo
Author
Robbins, Robert K.
text
Zootaxa
2009
2325
39
61
journal article
10.5281/zenodo.192050
1b056045-15f1-4e10-9b30-31e6c8ae56c9
1175-5326
192050
Calycopis janeirica
(Felder, [1863]) (
Figs. 39–83
)
Egg
(
Figs. 39, 40
,
60–65
). With regard to color pattern, shape, and chorionic ornamentation, the egg of
C. janeirica
looks like the eggs of
C. bellera
,
C. caulonia
,
C. cecrops
, and
C. isobeon
. Chorionic “islands” (
sensu
Downey & Allyn 1981
) protrude from floor of rosette cells. There are three micropylar openings in
C. janeirica
(
Fig. 62
), six in
C. caulonia
, and three or four in
C. bellera
.
Diameter
0.72–0.80 mm
(X =
0.78 mm
, SD = 0.02, N = 35); height
0.40–0.52 mm
(X =
0.43 mm
, SD = 0.03, N = 35). Duration 6–11 days (X = 9.14 days, SD = 1.01, N = 55).
First instar
(
Figs. 41–44
,
66–71
,
75, 76
). Morphologically similar to that of
C. bellera
and
C
.
caulonia
, but distinguished by the presence of whitish maculae at base of dorsal and lateral setae (
Fig. 42
). Mandibles of first instar of
C. janeirica
resemble those of
C. bellera
with six teeth and no tooth on oral surface (
Figs. 69, 70
).
Head capsule width
0.24–0.32 mm
(X =
0.28 mm
, SD = 0.03, N = 35). Body length ranging from
1.20 mm
to
3.52 mm
(X =
2.33 mm
, SD = 0.85, N = 30). Duration 4–8 days (X = 6.05 days, SD = 0.80, N = 55).
Head chaetotaxy (
Figs. 66, 67
). Same number of primary setae and pores as
C. bellera
and
C. caulonia
, and as in
C. bellera
, setae AF2, CD3, F1, F2, L1, and P2 are absent in
C. janeirica
. A1, A2, AF1, C1, and C2 usually shorter than those of
C. bellera
(
Fig. 66
), but size of setae vary intra- and interspecifically (
Duarte
et al
. 2005
).
Labrum chaetotaxy. Identical to that of
C. caulonia
(
Duarte
et al
. 2005
: 12, fig. 29) and
C. bellera
.
Body chaetotaxy (
Fig. 68
). Same number of primary setae and pore cupola organs as in
C. bellera
(this paper), distributed as follows:
Prothorax (
Fig. 68
). Identical to that of
C. bellera
and
C. caulonia
first instar (see
Duarte
et al
. 2005
: 13, figs. 30, 32).
Mesothorax (
Fig. 68
). Identical to that of
C. bellera
and differing from
C. caulonia
in having MD1 much shorter, about a tenth length of D1.
Metathorax (
Fig. 68
). Identical to that of
C. bellera
and
C. caulonia
first instar (see
Duarte
et al
. 2005
: 12, 15, fig. 30), including a conspicuous subdorsal pore cupola organ (SDL) of almost half the width of metathorax in all species.
Abdominal segment 1 (
Fig. 68
).
As
in
C. bellera
, with 10 pairs of setae and one pair of subdorsal PCOs (SDL of
Ballmer & Pratt 1992
). D2 slightly shorter than D1. Unlike
C. caulonia
, subventral PCOs (SVL of
Ballmer & Pratt 1992
) are lacking in
C. janeirica
as well as in
C. bellera
.
Abdominal segment 2 (
Fig. 68
). Similar to that of
C. bellera
and
C. caulonia
(
Duarte
et al
. 2005
: 15, fig. 30). D1 and D2 as in previous abdominal segment.
Abdominal segments 3–6 (
Fig. 68
). Identical to those of
C. bellera
,
but missing a pair of subdorsal PCOs on each segment (SDL of
Ballmer & Pratt 1992
). Distinguished from those of
C. caulonia
first instar by absence of
V2
(
Duarte
et al
. 2005
: 12, 15, fig. 30).
Abdominal segment 7 (
Fig. 68
).
As
in
C. bellera
,
C. cecrops
,
C. isobeon
and
C. caulonia
, with a dorsal PCO (DL of
Ballmer & Pratt 1992
) joined to chalaza of D1. Differs from
C. bellera
by lacking SV1 and subventral PCOs (SVL of
Ballmer & Pratt 1992
). These PCOs also absent in first instar of
C. caulonia
(
Duarte
et al
. 2005
: 12, 15, fig. 30).
Abdominal segment 8 (
Fig. 68
). Differs from
C. bellera
in having L3 almost as long as L2; SV1 much shorter, nearly equal to length of
V1
as in
C. caulonia
. Unlike both
C. bellera
and
C. caulonia
, subventral PCOs missing.
Abdominal segments 9+10 (
Fig. 68
). Differs from
C. bellera
by the presence of SD2 on ninth segment. Differs from first instar of other
Calycopis
with chaetotaxy available by presence of SV1 on A9. Length and position of setae similar to those of
C. caulonia
.
FIGURES 39–51.
C. janeirica
, egg and larva. 39. Newly laid eggs, dorsal view. 40. Mature egg, dorsal view. 41. Newly hatched larva lateral, dorsal view. 42. Idem, lateral view. 43. First instar after feeding on artificial diet, dorsal view. 44. First instar preparing to molting, lateral view. 45. Second instar, dorsal view. 46. Third instar, dorsal view. 47. Idem, lateral view. 48. Fourth instar, dorsal view. 49. Idem, dorso-lateral view. 50. Idem, preparing to pupate, dorso-lateral view. 51. Idem, lateral view.
FIGURES 52–59.
C. janeirica
, pupa. 52. After molting, dorsal view. 53. Idem, lateral view. 54. After 24 hours, dorsal view. 55. Idem, lateral view. 56. At middle of pupal stage (specimen with few dark brown maculae), dorsal view. 57. Idem, lateral view. 58. Idem, ventral view (it is already possible to see the imago’s eyes). 59. At end of pupal stage (one day after of this picture adult emerged), ventral view.
Second instar
(
Figs. 45
,
72
,
77, 78
). Cephalic color pattern resembling that of
C. bellera
and
C. caulonia
. Mandibles identical to those of
C. bellera
with six teeth visible externally as well with a toothlike process located ventrally on oral surface (
Fig. 72
); same number of setae as first instar. Integument coloration identical to
C. bellera
, including the two longitudinal reddish stripes extending dorsally and laterally from mesothorax to last abdominal segment. Unlike
C. bellera
,
C. caulonia
,
C. isobeon
, and
C. vitruvia
(
Duarte
et al
. 2005
; Duarte unpubl.), dendritic setae (
sensu
Ballmer & Pratt 1992
) missing on this and later instars. Prothoracic shield pentagonal with number and position of setae and pore cupola organs variable. Other characteristics identical to
C. caulonia
and
C. bellera
: prothorax wider and shorter than other thoracic segments; integument highly sculptured and distinctive, with uniformly-spaced oval depressions over epicuticular surface; pattern of oval depression similar in remaining instars; prolegs with uniordinal crochets in uniserial mesoseries, interrupted near center by a conspicuous fleshy pad (
Figs. 77, 78
), separating the crochets into anterior and posterior groups.
FIGURES 60–65.
C. janeirica
, scanning electron microscopy of egg. 60. Dorsal view. 61. Dorso-lateral view. 62. Micropylar area (rosette with three cells). 63. Cup-shaped cell of the exochorion. 64. Spine-like protuberance (=tubercle) arising from intersected ribs of the exochorion. 65. Aeropyle on top of tubercle.
FIGURES 66–67.
C. janeirica
, head chaetotaxy of first instar. 66. Anterior view. 67. Lateral view. Abbreviations (see text for setal nomenclature): as, adfrontal suture; en, epicranial notch; es, epicranial suture; tp, tentorial pit.
Head capsule width
0.38–0.52 mm
(X =
0.46 mm
, SD = 0.04; N = 30). Body length
3.68–6.16 mm
(X =
4.31 mm
, SD = 0.64, N = 30). Duration 3–5 days (X = 3.96 days, SD = 0.61, N = 54).
Third instar
(
Figs. 46, 47
,
79, 80
). Similar to previous instar. Differing in having greater number of spiculate setae on thorax and abdomen, prothoracic shield sagittiform, and prolegs with biordinal crochets in uniserial mesoseries (
Figs. 79, 80
). The typical velvet aspect produced by numerous dendritic setae and found in mature larvae of
C. bellera
and
C. caulonia
is lacking in
C. janeirica
.
Head capsule width
0.62–0.76 mm
(X =
0.68 mm
, SD = 0.07; N = 30). Body length
7.36–8.16 mm
(X =
7.63 mm
, SD = 0.24, N = 30). Duration 4–8 days (X = 6.87 days, SD = 1.03, N = 52).
Fourth instar
(
Figs. 48–51
,
73, 74
,
81, 82
). Similar to previous instar in shape and color of head, but with a greater number of secondary setae concentrated on antero-medial region of frontoclypeus. Mandibles with three setae rather than two (
Figs. 73, 74
). Integument orange-yellow with numerous golden secondary setae (
Figs. 48–51
). Prothoracic shield in two symmetrical parts. Prolegs with triordinal crochets in uniserial mesoseries (
Figs. 81, 82
).
Head capsule width
0.92–1.16 mm
(X =
1.02 mm
, SD = 0.11; N = 30). Body length
8.96–17.60 mm
(X =
13.96 mm
, SD = 4.13, N = 30). Duration 9–15 days (X = 12.23 days, SD = 1.511, N = 47).
Pupa
(
Figs. 52–59
,
83
). Similar to those of
C. bellera
and
C. caulonia
. Head and thorax initially translucent light yellow (
Figs. 52, 53
). Abdomen dorsally with intersegmental pale red maculae extending laterally, but not reaching spiracles (
Figs. 52, 53
). Integument darkening gradually to light brown, with dark brown maculae and small golden prominent setae scattered on dorsal and lateral body (
Figs. 54–57
). Differing from all other reared
Calycopis
species by the presence of dendritic setae around A6 spiracle (
Fig. 83
).
Pupal width on metathorax
2.88–4.96 mm
(X =
3.86 mm
, SD = 0.54, N = 30), on segment A3
3.68–5.76 mm
(X =
4.63 mm
, SD = 0.57, N = 30). Pupal length
6.72–11.04 mm
(X =
9.33 mm
, SD = 1.15, N = 30). Duration 9–14 days (X = 13.00 days, SD = 1.07, N = 46).
Development on artificial diet.
Caterpillars of
C. janeirica
have four instars rather than five as in
C. bellera
and
C. caulonia
. All larval instars of
C. janeirica
fed on artificial diet, which supported complete development. Data on development times from egg to adult are summarized in
Table 1
. Females of
C. janeirica
emerged from pupae as adults earlier on average (X = 50.06 days, SD = 2.07, N = 18) than males (X = 52.39 days, SD = 1.71, N = 28), as was the case with
C. caulonia
and
C. bellera
.
FIGURE 68.
C. janeirica
, body chaetotaxy of first instar. Dorso-lateral view. Abbreviations (see text for setal nomenclature): A, abdominal segment; cx, coxa; pl, proleg; sp, spiracle; ss, suranal shield; T, thoracic segment.
Cannibalism
.
As
with other
Calycopis
, when mature larvae were confined in containers, cannibalism was frequent.
Taxonomy
. The previous identification of
C. janeirica
was incorrect (
Field 1967a
;
Johnson 1988
,
1991
), and the reasons for our identification of this species were outlined above. In southern
Brazil
, including our reared adults, the male dorsal hindwing lacks blue and the female dorsal hindwing has blue, but the extent of this coloration in the female is variable. In other parts of South
America
, the amount of dorsal hindwing blue is variable in both sexes, and the number of species involved is yet to be determined.