Wm. Barnes and J. McDunnough.
On the Early Stages of Some Western Catocala Species.
Psyche 20:188-201, 1913.

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188 Psyche [December
n=nucleus.
nc= nerve-cord.
ns = neural septum.
ce = oesophagus.
mv = oesophageal valve.
pe = peritoneal membrane.
ph=pharynx.
pm ~peritrophic membrane.
pp = phwyngeal pump,
pro =prothorax.
R, rec = rectum.
s = set*.
sg= salivary glands.
sgd = salivary gland duct.
see = subcesophageal ganglion.
tg = thoracic ganglion.
tm = t,ransverse muscles.
tr= trachea.
vm = valve muscles.
y = yolk.
1.2, s=come.
ON THE EARLY STAGES OF SOME WESTERN
CATOCALA SPECIES.
BY WM. BARNES M. D. AND J. MCDUNNOUGH PH. D. Decatur, 111.
It was our good fortune in the autumn of 1912 to obtain ova of several species of Catocala whose early stages had never been stud- ied. Most of these we successfully bred through to the adult stage; colored figures of the larvae have been made and will be published later in connection with Beutenmuller's Monograph of the Genus Catocala, which we have been asked by the trustees of the American Museum to revise and complete for publication; in the meantime we offer the following notes on the larval stages. The species in question may be roughly divided into two groups- the oak feeders, comprising zoe, aholibah, ophelia, beutenmuelleri, and desdemona, and the willow and poplar feeders consisting of faustina, californica, irene, pura, and the species going under the name of aspasia Strecker. These two groups may be readily separated in the first larval stage by the fact that the set= arising from the primary tubercles are much longer in the oak feeders than in the willow and poplar feeders, giving the former under a lense quite a spiny appearance, whereas the latter appear almost smooth. Among themselves the larvse of each group are very similar in the first stage; the oak feeders are of a bluish-gray color with more or less strongly developed deep brown lateral blotches on the first four abdominal segments, 5-6 brown lateral lines and at times a similar centro-dorsal line; the presence of this dorsal



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19131 Barnes and McDunnovgh-Western Catocda 189 line in Stage 1 separates zoe and aholibah from the other three species; these two may be distinguished from each other, apart from the ova, which are very dissimilar, by the fact that the brown lateral blotches in aholibah tend to form more or less evident trans- verse banding and the two spiracular lateral lines are more broken than in zoe; in later stages the two species diverge widely from each other. Of the other three desdemona is distinguished by possessing 6 lateral brown lines in Stage 1 instead of 5 as in beutenmuelleri and ophelia; of these latter two species ophelia shows two sub- dorsal lines in Stage I which are not developed in beutenmuelleri until Stage 11. In the later stages of all these species the spaces between these brown lines gradually become paler than the ground color, forming stripes to which the lines act as a border, becoming more wavy in character and finally breaking up into a series of dots scarcely to be distinguished from the general surface of the body which is strongly sprinkled with similar dots; the dorsal stripe with its characteristic diamond-shaped enlargements on the abdominal segments appears usually in Stage 11, either as pale irregular bordering to the centro-dorsal line, or failing this line as a pale stripe between the two subdorsal lines, in which case the centro-dorsal line makes its appearance later. The willow and poplar feeders bear still more resemblance to each other than the oak group and are almost indistinguishable from one another in Stage I. All have a reddish-brown head and a pale greenish body shading into purplish or blackish laterally, this darker portion crossed by three pale rather waved lines; aspasia is the palest, and is almost entirely whitish-green with only faint traces of the lines; caliJornica is the darkest, being laterally almost black in Stage 11; the others are intermediate. The development of the maculation is the direct antithesis of that found in the oak-feeders; the stripes develop directly from the pale lines of Stage I, the darker bordering lines (the only ones visible in the 1st stage of the oak-feeders) appearing only in the 2nd or 3rd stages.
We append a more detailed description of the various stages:- Catocala zoe Behr.
Ovum.
Large, echinus-shaped, liver-brown with irregular ring of yellow around widest part; strongly ribbed, about 16 ribs diverging from the micropylar area and branching into 2 or 3 almost immediately, these ribs crossed at right angles by



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190 Psyche [December
very fine ribbing; micropylar area a rosette of minute cells. Width, 1.5 mm.
Height, .8 mm.
Stage I.
Head flat, black; body blue-gray; a dark dorsal line; a lateral band paler than the ground color, bordered on each side by a fine dark line and containing a similar central line; two further dark subspiracular lines; on abdominal segments I-IV small round deep brown patches laterally; primary tubercles black, prom- inent especially those situated ventrad to the pale lateral stripe, each with a long black seta Beneath pale gray with central dark blotches. Length, 6 mm. Width of head, .5 mm.
Stage II. Head whitish, strongly marbled with black, with two black lines in front, bending away from each other as they approach apex of cheeks, a central dark line in clypeus and a curved line around lateral portion of the cheeks; body gray with dark brown dorsal line irregularly bordered with whitish, this color tend- ing to spread out into diamond-shaped patches towards rear of each segment and defined outwardly by a blackish line; lateral whitish band as before with dark borders and central line; below this a black line on a level with tubercle IV; a 5th subspiracular line bordered dorsally with whitish; dark blotches on abdominal segments 1-4 as in previous stage; tubercles large, black, increasing in size towards posterior end; prolegs with a strong black chitinous lateral plate. Beneath whitish with dark central blotches.
Length, 12 mm.
Stage III.
Head as in previous stage; on the body the light stripes of the previous stage have so broadened that the general ground color appears light gray slightly mottled with flesh color, the darker portion being confined to a subdorsal irregular band and a similar lateral band in the region of tubercle 111; a light gray, very irregular dorsal band defined outwardly by black lines and broadening into diamond patches at the rear of each segment with a dark centro-dorsal line; the subdorsal dark areas are more or less shaded with gray anterior to tubercle 11; lateral gray area and dark lines as before; ventrad to this is the lateral dark area on a level with tubercle IV, paler on thoracic segments, with mere traces of dark lateral patches of previous stages, bounded ventrally by dark line; the light area ventrad to this contains two dark lines, the lower forming the outer margin; dorsal tubercles conical, shaded black and whitrf, especially large on 8th abdominal segment; a small transverse dark wart dnrsally on 5th abdominal segment. Beneath whitish with black central patches.
Length, 18 mm.
Stage IV.
Head gray strongly striate with black; a black upright dash in central front portion of cheek and a broad curved lateral line of same color; body pale gray, marbled, rough and tubercular in appearance; markings essentially as in pre- ceding stage; the pale gray dorsal stripe with diamond-shaped enlargements is usually quite prominent owing to its being bordered by a darker gray subdorsal stripe, deepest in color on the lst, 2nd and 5th abdominal segments; dark supras- piracular stripe distinct, paler in anterior portion and on 4th abdominal segment; dorsal tubercles pale, ochreous, raised, with more or less of a transverse ridge between the two tubercles 11, especially marked on lst, 2nd, 5th and 8th ab- dominal segments; on 5th abdominal segment a small dark transverse dorsal wart, shaded posteriorly with blackish; tubercle I1 on 8th abdominal segment large and conical; tubercle IV situated on a rather raised wart just behind spiracle; below



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19131 Barnes and McDunnough-Western Catocala 191 this a dark line in a paler field is distinguishable. Beneath pinkish white with the
usual dark blotches. Length, 30 mm.
Stage V.
Head pale ochreous, the upper portion of the lobes tinged with orange, marbled with dark purple-gray with black lateral line descending as far as the ocelli; slight central black line to clypeus. Body very warty both dorsally and laterally, with deep lateral indentations above and below the spiracular area, pale gray, marbled with olive brown, maculation quite similar to that of the pre- ceding stage; dorsal stripe pale gray, irregular, broadening at rear of each segment into diamond-patches which are tinged with ochreous on abdominal segments 1-4; subdorsal stripe dark brownish, broad, waved, prominent behind tubercle I1 on abdominal segments 1-3, 5 and 8, paler and more diffuse on the other segments; dark spiracular stripe, paler on the thoracic segments, abdominal segment 4, the anterior portion of 5 and posterior portion of 6; abdominal segments more or less humped dorsally between tubercules 11, most so on 5 and 8; tubercles situated on a raised hump, tinged with ochreous. Lateral filaments whitish. Beneath deep purple-pink with blackish central patches. Length mature, 40 mm.
Food-plant: Buds and catkins of burr-oak. The young larvae emerge with the first warmth of spring, all the ova hatching within a short period of each other; the growth is very rapid, maturity being reached in less than a month; the pupal stage is about three weeks.
The species is very easy to raise, the critical period being just before attaining full growth, when the larvse seem liable to intes- tinal troubles which are apparently more or less contagious. In our brood there was also a noticeable difficulty in properly shed- ding the larval skin in transforming into the pupa. Catocala aholibah Stkr.
Ovum. Dark dirty green, surface of egg strongly granulate but without ribs; micropylar area scarcely visible as a rosette of minute hexagonal cells. Diameter,
2 mm.
Stage I.
Head deep brown, prothoracic plate black; body bluish-gray with brown dorsal line, three brown lateral lines of which the middle one is most distinct; two similar subspiracular lines the upper one rather curved and much broken; abdominal segments 1-4 crossed by broken broad transverse brownish bands, heavi- est laterally; tubercles large, black, with long setae. Beneath pale with central dark blotches. Length, 6 mm.
Stage II. Head pale brown, strongly veined with black-brown, forming a line on each side arising from the mouth and curving outwards and upwards, broadest towards apex of lobes; central dark line to clypeus; body pale brown, slightly banded with brown on abdominal segments as before; a brown dorsal line broad- ening slightly towards rear of each segment and irregularly bordered with creamy, forming more or less distinct diamond-shaped patches as usual; two pale slightly



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192 Psyche [December
waved lateral lines and a third subspiracular one, much curved; tubercles large, black; slight dorsal transverse wart on the 5th abdominal segment and more prom- inent conical protuberance on dorsal portion of 8th; prolegs pale with black stripes. Beneath whitish with prominent black patches. Length, 10 mm. Stage III. Head pale, strongly lined with black, apex of lobes tipped with orange behind which is a heavy black curved line; body gray-purple with distinct black tubercles which at times are orange-tipped; a prominent transverse blackish wart on 5th abdominal segment and a conical protuberance tipped by tubercle I1 on 8th; similar lesser prominence on the 9th; lines and stripes as in preceding stage but broader and less distinct; lateral black shading below transverse wart of 5th abdominal segment. Length. 18 mm.
Stage IV. Head pale creamy strongly lined with black in front and with apex of lobes tipped with orange behind which is heavy black marking. Body purple- brown, the 1st abdominal segment with pale creamy suffusion between and lateral to tubercles I and 11, often very faint; stripes of previous stage not distinguishable but the black border lines have broken up into a series of irregular dotted lines on a more or less whitish ground, giving a general mottled appearance; tubercles orange, tipped with black and ringed at basewithwhite, the black color often predominating; transverse wart of 5th abdominal segment black, tipped with white at apex and defined laterally by the orange tubercles 11; laterally to this is more or less oblique dark suffusion; prominent hump on 8th abdominal segment tipped by large conical tubercles 11, orange in front,- black posteriorly. Beneath whitish with black central blotches. Length, 30 mm.
Stage V. Head pale ochreous, pitted, marbled with purplish gray with tips of lobes tinged with umber-brown, beneath which is a blackish dot and behind which on the dorsal portion of the lobe is a broad black stripe slightly curved at each end but not descending downwards below the level of the umber-brown area. Body pale liver-brown with numerous black dots encircled with white the remnant of the lines of earlier stages; tubercles coral or deep brown ringed at base with white; transverse wart of 5th abdominal segment whitish strongly marbled with black; 6th and 7th abdominal segments laterally and posteriorly slightly discolored with brownish; 8th abdominal segment with strong dorsal hump on which are situated tubercles 11, large, conical, directed backwards, umber tipped with black; spiracles situated in pale area, black-rimmed; legs pink, shields of prolegs, prothoracic and anal plates ochreous; filaments white; first abdominal segment between tubercle I1 and spiracle slightly paler, the remains of the patch of the previous stage. Beneath white with black and brown central blotches. Length, 60 mm.
Food-plani: Buds and catkins of burr-oak. The emergence and habits of the young larvae coincide with those of zoe; in all its stages the species shows a close resemblance to sponsa of Europe. Ova from Truckee, Calif. and Provo, Utah gave rise to similar larvse, the resulting imagines being also identi- cal except that the Utah specimens were generally larger. About
1-4 of the Utah larvae gave the form coloradensis Beut. which is a



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19131 Barnes and McDunnough-We3tern Catocala 193, mere aberration in which the primaries are distinctly paler and the" brown band beyond the t. p. line more marked in consequence. Catocala beutenmuelleri B. & McD.
Ovum. Flat, strongly ribbed; micropylar area forms a slightly raised button with central depression containing the micropyle itself; from the rim of this area 17-20 broad unbranched ribs arise, crossed at right angles by numerous fine wavy cross-ribs. Color purple with yellow ring a short distance above the base. Diam-
eter, 1 mm.
Sfage I. Head black. Body pale.gray with 3 brown lateral stripes; diffuse brown patches laterally on 1st four abdominal segments with traces of a 4th brown line posterior to them; below these a 5th subspiracular line; tubercles black, small, with long setse. Beneath pale with central dark blotches. Length, 4 mm.
Stage II. Head pale gray, heavily mottled with brown except in front and with two heavy dark curved lines below apex of lobes not reaching lower than apex of clypeus; five equidistant lateral and subspiracular lines as before, the 3rd line rather waved, the lower line distinct; dark lateral abdominal patches much fainter than in preceding stage; dorsal region with two brown lines narrowing between tubercles I and spreading out towards rear of segments tending to form diamond-shaped patches of the enclosed space; traces of a centro-dorsal line between tubercles I on abdominal segments; on 5th abdominal segment a slight raised dorsal wart; on 8th abdominal a sma!l conical projection; tubercles black with setae as before. Length, 9 mm.
Stage III.
Head as before but more striate with black. Body and markings.
essentially as before but pale dorsal diamond-shaped patches more noticeable and the dark lines appearing more as border lines to enclosed yellowish stripes; small transverse wart on 5th abdominal segment; the rear portion of this segment and anterior portion of 6th shaded with blackish; 8th segment raised dorsa!ly with conical reddish tubercles and a small black lunate mark posterior to these; pro- legs with black stripe.
Length, 15 mm.
Stage IV. Head pale, strongly marbled, apex of lobes slightly orange, below which is a small blackish curved mark; behind the apex of lobes a curved double black line extending laterally down the sides of the cheeks. Body gray, somewhat ochreous at the incisions of -the first four abdominal segments; lines and stripes much as before but fainter, the former being broken up into a series of dots; tub- ercle on 5th abdominal segment black, prominent, with dirty brown oblique patch extending laterally downwards to base of prolegs; tubercles deep orange, those on 8th abdominal segment situated on a strong hump behind which is a curved orange mark; prolegs pale, striped with black;
beneath whitish with the usual black
patches. Length, 25 mm.
Stage V. Head purplish gray marbled in the front of the lobes with darker; the two tubercles at the apex of the lobes tipped with orange with dark blackish, curved mark behind the apex. Body light gray, fairly smooth, with small oblique ridges on the anterior abdominal segments extending laterally forward from tuber- cle 11; pale dorsal stripe with diamond-shaped enlargements; remainder of body heavily marbled with black dots, representing the dark border lines of previous



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194 Psyche [December
stages; a very prominent fleshy beak-like wart on 5th abdominal segment shaded with black and with black posterior markings extending obliquely downward over the anterior portion of the following segment; laterally the 5th segment is shaded with light brown extending between the prolegs on to the 6th segment; prom- inent conical dorsal tubercles on the 8th abdominal segment with Iunate brown mark behind extending to above spiracle; filaments whitish; beneath white with central black blotches.
Length, 40 mm.
Food-plant: Buds and catkins of oak.
The larvse emerged about a week later than zoe and aholibah maturing rather more slowly than these two species; a number of the full grown larvae refused to pupate but gradually shrivelled up and died; it is probable that better results would be obtained by isolating the larvae in the last stage. The prominent dorsal wart on the fifth abdominal segment would place the species in the ultronia group. In our opinion beutenmuelleri will prove to be a local race of verrilliana but we can find no larval description of this species. We cannot agree with Hampson who makes this species and werneri Bied. synonymous; this latter species, of which the type is in Coll. Barnes, shows much more affinity to violenta Edw. and we should not be surprised if it proved to be a mere aberrant form of the same; ophelia Hy. Edw. we have proved by breeding to be a species very distinct from beutenmuelleri and presumably therefore from verrilliana; breeding alone will show how closely it is related to violenta Edw. Catocala ophelia Edw.
Ovum.
Hemispherical, much higher than beutenmuelleri; ribbed, about 20 ribs attaining the edge of the raised circular micropylar area, every second rib branch- ing dicotomously in contradistinction to the preceding species where the ribs always remain single; color liver-brown with yellow ring near the base. Diameter, 1.2 mm. Stage I. Head blackish; body blue-gray more or less completely banded with brown on the first four abdominal segments, tending to form lateral and dorsal blotches; tubercles black with long black setae; geminate rather waved dorsal brown lines situated in the area between tubercles I and I1 and tending to separate towards rear of each segment,; three lateral brown lines and two further subspirac- ular ones. Beneath whitish with brown central patches. Length, 6 mm.
Stage II. Head pale gray, marbled with black with inverted V shaped black mark on front of each lobe and central black dash on clypeus. Body pale yellowish
gray with black tubercles and traces of lateral purplish patches of previous stage; markings as before with the addition of a centro-dorsal broken stripe, thickened slightly at intervals. Length, 11 mm.
Stage III. Head same as before.
The lines on the body of the previous stage



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19131 Barnes and McDunnough-Western Catocala 195 now appear as border lines to three pale ochreous stripes; dorsal stripe broadening to the usual diamond shaped patches and with central dark line, thickening in these same patches; small dark transverse tubercle on 5th abdominal segment; dorsal protuberance on the 8th abdominal segment with large conical flesh-colored tuber- cles; 9th segment similar but smaller. Length, 18 mm. Stage IV. Head pale with V mark and central dash as before; apex of lobes tinged with orange; cheeks bordered by a double black curved line. Body pale gray, the markings essentially as before but fainter, the black border-lines broken up into dots, the lateral stripes most marked between prolegs of 5th and 6th abdom- inal segments, the ground color being here somewhat darker; dorsal wart on 5th abdominal segment small, shaded with blackish; tubercles orange, I1 on the 8th abdominal segment large, extended backward and situated on a prominence. Beneath white with black central patches. Length, 30 mm.
Stage V. Head pale purple-gray, apex of lobes white, slightly lined with orange- red and purple, the white color extending down cheeks frontally in irregular lines; a black lateral border line to cheeks shaded inwardly with orange. Body deep gray, smooth, heavily dotted with black; stripes of the previous stages fairly dis- tinct; tubercles small, brick-red, tubercle I1 on 8th abdominal segment large, situated on prominence; wart on 5th abdominal segment small dusky; an och- reous subspiracular stripe between prolegs of 5th and 6th abdominal segments, shaded dorsally with deep smoky gray; filaments whitish. Beneath white with the usual dark central patches. Length, 40 mm. Food-plant: Buds and catkins of burr-oak. The species may at once be separated from the preceding by the lack of the prominent hump on the 5th abdominal segment and the presence of an ochreous patch between prolegs of 5th and 6th abdominal segments. It coincides with beutenmuelleri in its general habits. The parent moth was captured at Provo, Utah. Catocala desdemona Edw.
Ovum. Pale yellowish, very finely ribbed with numerous branching ribs, hemi- spherical; micropylar area slightly raised, circular. Stage I.
Head brownish; body pale gray with fairly large black tubercles and brown lateral patches on abdominal segments 1-4 tending to broaden into trans- verse bands; three brown lateral lines and traces of three further lines below these the upper two of which are only visible behind the lateral brown patches. Beneath pale with central brown blotches. Length, 6 mm. Stage II. Head gray with curved black line in frontal portion of cheek curving outward toward apex of lobes; central black line on clypeus. Body gray with
pale germinate dorsal stripes bordered with dark brown and tending to coalesce; two pale lateral stripes similarly bordered; a third subspiracular one; a faint brown line defines the pale ventral area; traces of the dark lateral patches still present. Length, 13 mm.
Stage III. Head pale gray slightly tipped with orange; dark W mark in frontal



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196 Psyche [December
portion of cheeks and dark curved lateral border to same; clypeus with central dark line. Body pale gray; dorsal and three lateral creamy stripes prominent, bordered by brown lines; 5th abdominal segment with small dorsal wart tipped with reddish at apex; from this wart a dusky lateral shade extends downward to the prolegs, crossed by the pale lateral stripes; tubercles reddish, tubercle I1 of 8th abdominal segment conical, with dark lunate reddish line posterior to it; - prolegs striped with black. Length, 18 mm. Stage IV.
Head gray, apex of lobes tinged with orange-red; lobes strongly marbled vertically forming W mark frontally; geminate lateral black curved line and central line to clypeus. Body gray with paler creamy stripes; dorsal stripe, as usual, tending to diamond-shaped enlargements, with centro-dorsal dark line and similar colored border-line, most prominent at rear of 1st four abdominal segments; three pale lateral stripes bordered with blackish lines which tend to break up into dots; on 1st abdominal segment a dark oblique lateral shade extends forward from tubercle I1 toward the 3rd pair of legs; dorsal wart on 5th abdominal segment flesh-colored with diffuse darkish shade extending from it laterally to prolegs; tubercles ,orange; curved line behind the conical dorsal tubercles on 8th abdominal segment brownish. Beneath whitish with the usual dark blotches. Length, 20-30 mm.
Stage V.
Head whitish tinged with orange in front and along sides of cheeks; pale purple gray marbling in front forming a W mark, black border stripe to cheeks laterally. Body pale to dark gray caused by numerous dark dots on a whitish ground; dorsal and lateral stripes of previous stage distinct, latter waved and irreg- ular, the dark border lines broken up into dots, most distinct behind tubercle I1 on first 5 abdominal segments as lower border of the dorsal stripe and upper border of the first lateral one; on first abdominal segment an oblique pale shade running from tubercle I1 to beyond spiracle and bordered posteriorly slightly with dark gray; wart on 5th abdominal segment fairly large, tipped with white, with brown marbling at base, lateral to this wart an oblique brownish shade crossed by the pale stripes and with the ground color showing on a level with the spiracle as a dark gray cross- stripe; reddish lunate dorsal line on 8th abdominal segment posterior to the large conical tubercles 11; tubercles orange with white base; prolegs pinkish marbled with brown; filaments white. Beneath white shaded with pink with the usual dark central blotches. Length, 50 mm.
Food-plant: Buds and catkins of burr-oak. This species was the latest of the oak-feeders to emerge; after reaching its full size the larvae seem to find difficulty in pupating, a number of ours dying at this stage. The larva is considerably larger and more variegated than that of either ophelia or beuten- muelleri.
Catocala pura Hlst.
Ovum. Hemispherical, deep purplish-red banded with yellowish near base, ribbed, the ribs branching into two or three shortly after leaving micropylar area



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19131 Barnes and McDunnough-Western Catocala 197 which is slightly raised.
Very similar to the egg of C. favstina.
Diameter, 1.25
mm.
Stage I. Head red-brown. Body when first emerged blackish, later dirty green shaded with purplish anteriorly and posteriorly, with three pale lateral lines; tubercles small, black, with short setae. Beneath paler with faint central dark blotches. Length, 5-6 mm.
Stage II.