The Canadian Entomologist, Vol. XII., No. 2, February 1880
Part 1
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The Canadian Entomologist.
VOL. XII. LONDON, ONT., FEBRUARY, 1880. No. 2
ON CERTAIN SPECIES OF SATYRUS.
BY W. H. EDWARDS, COALBURGH, W. VA.
1. NEPHELE.—Kirby, Faun. Bor. Amer., 1837, described this species as follows: “Wings brown; primaries both above and below with a paler submarginal broad band including two eyelets; the upper ones surrounded by a paler atmosphere, with a black iris and white pupil; on the under side the atmosphere of the eyelets is most distinct and forms a kind of glory round them,” etc. Nothing is said of the sex, but apparently this is the description of a female. The wings of the male are blackish-brown, usually of uniform shade throughout—that is, in the typical male, corresponding to the female of Kirby. But there is a frequent departure from this type in the direction of _Alope_, the “pale atmosphere” about the ocelli appearing in the male, and in both sexes gradually widening and becoming less obscure till it culminates in a clear yellow band. When this is reached we have _Alope_, Fabr. So that _Nephele_ intergrades completely with _Alope_. But this is not everywhere and always. The metropolis of the typical _Nephele_ is in Canada and northern New England, that of _Alope_ in the States south of New York. There is a line running about with the southern boundary of New York, or it may be, in Pennsylvania, below which _Alope_ holds sole possession, and no tendency is discoverable towards _Nephele_. In the extreme northern area, if there is any departure from typical _Nephele_, it is the exception, not the rule.
Mr. Scudder, in his essay on The Distribution of Insects in New Hampshire, 1874, says of _Alope_: “This insect is tolerably abundant, sometimes very common, in the southern half of New England. The most northern localities … are Norway, Me., Thornton and Shelburne, N. H., and Sudbury, Vt.” Thornton is just south of the White Mountains, and Shelburne is close by the mountains on the north-east. Of _Nephele_ he says: “It is found over the whole northern half of N. E. in great abundance. The only locality in which I have met with it is in Massachusetts, in the elevated region about Williamstown,” &c. This place is in the north-west corner of the State, next the Vermont line, and the elevated region spoken of is a continuation of the Green Mountains. So it appears that _Nephele_ comes down to the Massachusetts line and _Alope_ flies as far as the White Mountains. In the intervening district the intergrades fly just as in New York.
I made application to Canadian lepidopterists for information about the occurrence of _Alope_, and soon ascertained by examples sent me that _Nephele_ with a pale atmosphere, but not at all indicative of a band, passed by the name of _Alope_. Thereupon I sent a typical _Alope_ to Mr. William Murray, of Hamilton, who kindly offered to make inquiry of his acquaintances in different sections of Ontario. He replies, 31st Dec, 1879: “I now send you my information. Of all my correspondents not one has ever seen an _Alope_ that has been taken in Canada, but _Nephele_ has been taken by all. I begin to think that _Alope_ is not to be found in Canada at any point.”
Mr. H. H. Lyman writes from Montreal: “In July, 1876, I spent a couple of days at a farm near Freligsburg. P. Q., one mile north of the Vermont border, and found _Nephele_ very common. Most of the specimens taken showed a yellow ring about the eye-spots on primaries, but one of them shows on upper side a somewhat faint, but quite discernible, patch corresponding to the yellow band of _Alope_. Was at same place in 1877. _Alope_ was not seen either year.” Mr. Caulfield writes Mr. Lyman: “I have never taken a specimen of _Nephele_ showing any tendency towards _Alope_, nor have I seen any Canadian examples showing it.” Mr. Lyman adds that at Portland, Maine, where he collected several summers, _Alope_ was common as well as _Nephele_ and all intergrades.
(To the west of New York, in the latitude of the belt spoken of, it is believed that the two forms fly together at least as far as Wisconsin. Prof. A. J. Cook writes that both are common in Michigan, south of the latitude of Grand Rapids. At Toledo, Mr. John Wilson writes that _Nephele_ is rare, and _Alope_ unknown, so far as appears. At Cleveland, O., Dr. J. F. Isom informs me that _Alope_ is very rare, but that _Nephele_ is abundant in some seasons. In south-west Ohio, Dr. H. K. Landis, of Columbus, writes that he cannot learn that either form has ever been taken. They are not mentioned in Mr. Dury’s list of butterflies found about Cincinnati. But in northern Illinois _Nephele_ is abundant and _Alope_ not found at all. So that somewhere between New York and Illinois, in Ohio and Indiana, _Alope_ seems to disappear, while _Nephele_ becomes the sole form; but whether the separation is abrupt or gradual is not ascertained. As the information which I have been able to gather is so meagre as regards the States west of New York. I shall confine my remarks to that State and New England.[1])
We have therefore in these separated districts two apparently good species, answering to any definition of that name. But between, there is a belt of latitude passing through New York and southern New England, where in one section or other both types are found and the whole series of intergrades. In this belt _Alope_ and _Nephele_ are found to be dimorphic forms of one and the same species. I formerly was of the opinion that they were distinct species, though in some districts there were intergrades. I thought these approaches of one to the other did not bridge the whole space between. In a paper printed in Proc. Ent. Soc. Phil, 1866, I gave my reasons therefor. But some observations made in July, 1876, at Martha’s Vineyard, led me to suspect a closer relationship between the two species or forms. In the open country back of Oak Bluffs, I found these butterflies fresh from chrysalis, and in considerable numbers. They were all very black, diminutive, and there was every grade from what I had been in the habit of calling _Nephele_ to unquestionable _Alope_, with a broad clear-colored band. The band was not yellow, however, as in the typical _Alope_, but reddish-yellow like that of _Pegala_, which Fabricius called _rufa_ in distinction from _flava_, applied to _Alope_. Mr. Scudder took the same small reddish-banded form on Nantucket, which island is about 30 miles from the mainland, Martha’s Vineyard being about 7. I call this variety _Maritima_, but whether it is restricted to the islands, or appears on the adjacent coast, I am not yet advised. Mr. Mead obtained for me a large number of eggs of this butterfly, while at the Bluffs shortly after my departure. They were laid by the broad-banded females in confinement and mailed to Coalburgh. There the larvæ hatched out, and these as well as the eggs were found to be precisely like the same stages of _Nephele_ from Catskills. But none of the larvæ survived the winter.
In the belt spoken of, _Nephele_ rather keeps to the highlands. It is the prevailing form in the Catskills, if with it are classed the intergrades, but full-banded _Alope_ may be taken in small numbers every season. Along the Hudson River, _Alope_ is the common form, but I have received intergrades very near to _Nephele_ from Mr. Hulst, taken at Hoboken, N. J.; and a black _Nephele_ ♂ from Mr. H. Laitloff, which he writes me was taken some five years since near Greenville, Jersey City. It was so unusual a form that Mr. Laitloff sent it to me for name. At Coalburgh, W. Va., _Nephele_ is never seen, but _Alope_ is the only form; and so on southward.
2.—_Alope_ was described by Fabricius, Ent. Syst., 1793, as fuscous (_fusca_) with a yellow (_flava_) band; with two ocelli on fore wings; on hind wing one ocellus above, six below. The band is very broad in the female, usually narrower in the male, pale yellow in both sexes. The ocelli resemble those of _Nephele_ and vary in same manner. Usually they are round, but sometimes oval; are either small or large, often equal, but sometimes the upper is larger, at others the lower. Now and then a third pupilled ocellus appears, and individuals have been taken with but one ocellus (the upper). It is not very unusual to find examples in which a black point, or what may be considered as a rudimentary ocellus, presents itself. On the upper side of hind wing is often a small but complete ocellus near inner angle, but in many cases it is partly or wholly wanting; and occasionally there are one or two black spots in addition. The males in the majority of examples have six small ocelli on the under side of the hind wings; the females rarely have six, and often none at all. At the north, _Alope_ is blackish-brown, more brown in the female; but to the southward brown prevails in both sexes; and it is of a lighter shade, while the under side has a tint of yellow more or less decided over whole surface, often mixed with gray. The band is of yellow, or with a slight ochrey tint. This is a description of the extreme southern type, and to distinguish I call it var. _Texana_. All examples from Texas which I have seen have a complete anal ocellus, and six ocelli beneath, of pretty large size—larger than in northern _Alope_—in distinct ochrey rings; the pupils white points with a few blue scales about them in the larger ocelli. Of 70 _Nephele_ ♂ examined, 50 have 6 ocelli, 11 have 5, 3 have 4, 3 have 3, 2 have 1, 1 has 0.
Of 55 _Nephele_ ♀, 6 have 6, 1 has 5, 4 have 4, 13 have 3, 7 have 2, 13 have 1, 11 have 0.
Of 24 _Alope_ ♂, 15 have 6 ocelli, 3 have 5, 4 have 1, 2 have 0.
Of 25 _Alope_ ♀, 12 have 6, 1 has 4, 4 have 2, 4 have 1, 4 have 0.
Therefore of _Nephele_ ♂, 71 per cent. have 6 ocelli, 4 per cent. have under 3; 1.4 per cent. have 0.
Of _Nephele_ ♀, 11 per cent. have 6, 56 per cent. have under 3, 20 per cent. have 0.
Of _Alope_ ♂, 62 per cent. have 6 ocelli, 25 per cent. under 3, 8 per cent. 0.
Of _Alope_ ♀, 24 per cent. have 6, 48 per cent. have under 3, 16 per cent. have 0.
3.—The dark Satyrus which inhabits Illinois and westward has gone by the name of _Nephele_, though differing somewhat from _Nephele_ of the east. I was struck by the difference between a series sent me by the late Mr. Walsh from Galena, years ago and when I first began collecting butterflies, and a series of _Nephele_ taken in the Catskills, and I have always kept the two apart in my cases, considering the Illinois form as at least a well marked variety. Mr. Worthington has recently written me: “I have received a lot of _Nephele_ from New Hampshire and am surprised at the difference between them and the Illinois _Nephele_.”
The males of this last are almost black, the ocelli are very small and without rings. But in some examples there is a faint russet or yellowish tint about the ocelli, and perhaps on the space between them. On the under side the rings are russet or ochraceous, on both wings. The females are almost invariably and uniformly dark, and only occasionally is there a paler shade over the extra discal area of fore wings. Out of a number of females I find but one in which there is a clouded yellow space about the ocelli, and only three on which there are yellow, though hazy, ocellar rings. Of 16 ♂, 14 have 6 small ocelli beneath, 1 has 5, 1 has 2. Of 19 ♀, 2 have 6, 2 have 5, 6 have 4, 2 have 3, 6 have 2, 1 has 1. This form prevails exclusively to the Rocky Mountains. I have received it from Nebraska, Montana, Colorado and New Mexico, but _Alope_ is unknown to me from that region.
In CAN. ENT., ix., 141, 1877, I gave the history of _Nephele_, bred from eggs laid by a typical female from the Catskill Mountains, Hunter, N. Y. In fall of 1878, I wrote to several correspondents for eggs, and by their good will obtained many. Prof. Lintner and Dr. Bailey sent eggs of _Alope_ from Albany, N. Y. Rev. Mr. Hulst, with the zeal and kindness which distinguishes him, crossed the rivers from Brooklyn to Hoboken, and brought away females of _Alope_, from which he obtained eggs for me. I got _Alope_ eggs here at Coalburgh from three females. A friend at Hunter sent eggs of _Nephele_, and Mr. Worthington sent many of the Illinois form from Chicago. In each case the parent was sent with the eggs that the type might be noted. From Albany, Hoboken is 150 miles south; Coalburgh 800 miles southwest; Hunter is 35 miles southwest of Albany and of about 2,000 feet greater elevation. Chicago is about 800 miles northwest of Coalburgh and 1,000 west of Albany. So that the five localities are separated by considerable distances, and there has probably been no intercommunication at any time so far as these insects are concerned.
The eggs of the six lots were kept apart and as the larvæ hatched (at from 14 to 28 days from deposition, depending on the temperature), they were placed on sods in separate pots and left in the coolest room in my house. But some of the Illinois eggs were sent to Mr. C. P. Whitney, at Milford, N. H., who offered to put them on ice. I wished to try the effect of cold in retarding the hatching. Early in February I received the boxes again and found a number of healthy larvæ, with a few unbroken eggs. These last proved to be dead. The eggs had been sent in a paper pill box which was within a flat tin box, and this was set directly on the ice. The young larvæ when I received them were fixed to the rough sides of their box and had not been attacked by mould, the enemy most to be dreaded. Mr. Whitney wrote that he was notified in December that the ice-house was empty, and he thereupon removed the tin box without opening it, and placed it in a snow bank, where it remained till I sent for it. The larvæ may have been emerging from the eggs when he first received them, or perhaps did so in the interval between ice-house and snow. This method of keeping larvæ which become lethargic immediately upon leaving the egg will probably be found successful with all species of butterflies which have that habit—as the large Argynnids—and make it possible to breed them in numbers. I have been unable to find any other mode of wintering such larvæ without a certain loss of most of them.
On 23rd Jan., 1879, I transferred such of the Satyrid larvæ as were living (and this included some of each lot) to fresh sods, and 28th Jan. noticed that several were feeding. One Hunter _Nephele_ passed 1st moult 23rd Feb’y, and before 4th March several of the same lot had passed the moult. But the Illinois _Nephele_ and all _Alope_ lingered. One Coalburgh _Alope_ and one from Hoboken passed 1st moult 7th March, by which date the Hunter _Nephele_ spoken of was swollen for 2nd moult, which it passed two days later. Two Illinois _Nephele_ passed 1st moult 8th March. To the end some of the Hunter _Nephele_ were in advance of all, and some of the Illinois examples lingered behind all. The stages of Coalburgh _Alope_ were as follows:
1st moult passed 7th March. 2nd ” ” 21st ” 1st to 2nd—14 days. 3rd ” ” 14th April. 2nd to 3rd—24 ” 4th ” ” 2nd May. 3rd to 4th—18 ” In chrysalis 26th ” 4th to chrys.—24 ” Imago issued 9th June. chr. to imago—14 ”
Of Hunter _Nephele_ I find no notes, but in 1877 the stages were
1st moult to 2nd—23 days. 2nd ” to 3rd—14 ” 3rd ” to 4th—14 ” 4th ” to chrys.—28 ” Chrys. to imago—14 ”
Of Illinois _Nephele_ the stages were:
1st moult passed 8th March. 2nd ” ” 21st ” 1st to 2nd—13 days. 3rd ” ” 9th April. 2nd to 3rd—19 ” 4th ” ” 26th ” 3rd to 4th—17 ” In chrysalis 17th May. 4th to chry.—21 ” Imago issued 30th ” chry. to im.—13 ”
The eggs of all these forms are alike, not to be distinguished from each other. They are conoidal, truncated at top and slightly arched; marked by about 18 vertical ridges running from base to top, the spaces between excavated roundly, and crossed by fine striæ; the top is covered with shallow cells, the outer ones irregularly hexagonal, the inner long and narrow about a central oval cell.
Nor can the larvæ of these forms be distinguished from each other up to second moult. The young, of first stage, are very peculiar and quite unlike what they become after first moult, as well as unlike all other Satyrid larvæ which I have bred. Under the microscope they look like the vertebræ of a fish, by reason of the many rows of long hooked bristles, those of upper and lower rows being curved back, the middle row forward. General color carnation, with a medio-dorsal line, and three lines on each side, all of crimson. After 1st moult the color is green, and the stripes, which are the same in number and position as the lines of first stage, are dark green; the hairs short and straight.
After 2nd moult the color becomes yellow-green and the stripes are changed. There is now a dark green one on middle of dorsum and a yellow one covering the ridge over the feet. On 24th March, I compared Hoboken _Alope_, Hunter and Illinois _Nephele_, of same age, all lately past second moult, and could see no difference whatever between them. Some _Alope_ and Hunter _Nephele_ were much covered with long hairs which were bent to the surface and gave them quite a shaggy appearance. But others did not show this peculiarity.
On 18th April, I compared Coalburgh _Alope_ and _Nephele_ from both localities, all past 3rd moult; length from .68 to .75 inch. All were yellow-green and in general alike; all had the yellow basal ridge, but in addition to this, the Illinois _Nephele_ had a distinct longitudinal yellow stripe on upper part of side, and on either edge of the green dorsal stripe was a fine yellow line. The Hunter _Nephele_ showed very faint traces of the yellow side stripe; the _Alope_ none at all.
Comparing another Coalburgh _Alope_ and Hunter _Nephele_ a few days later, both past 3rd moult, neither showed traces of these lines and I could see no difference between the two.
At 4th moult all the Illinois _Nephele_, now .7 inch long, showed same peculiarities as at last stage. No other larva of the several lots presented the yellow lines so plainly at the same age, that is, just after the moult; but there were one or more _Alope_ and Hunter _Nephele_ which gave indications of the side line, and this came out more distinctly as the stage progressed. But most were without the side line.
Comparing mature larvæ:
One Albany _Alope_, length 1.25 inch, greatest breadth .16 inch; color very yellow-green, no yellow side or dorsal stripes or lines.
One Coalburgh _Alope_, length 1.6, gr. br. .2 inch; color yellow-green, the side more green than dorsum; a yellow side line, quite indistinct.
Hunter _Nephele_, 3 examples; length of one 1.2 inch, gr. br. .15 inch; of another 1.15, br. .16 inch. Two were yellow-green, of same shade as nearly all the Coalburgh larvæ. One was more decidedly yellow, with less green; but in none was there a yellow side stripe or the fine dorsal lines.
Illinois _Nephele_, length 1.36, gr. br. .14 inch. Color bright yellow-green, the dorsum more yellow than side; on the side as broad a yellow stripe as the one along basal ridge, and the green dorsal stripe edged by yellow.
Summary as to larvæ:
The five lots could not be separated before 2nd moult. After that, through the stages to maturity, the _Alope_ from different localities and the Hunter _Nephele_ varied somewhat in the shade of green, being more or less yellow; in some yellow prevailing on dorsum, green on sides; all had the yellow band on basal ridge, either pale or deep colored. If the yellow side line was present, as in some examples it was, it was indistinct, or obsolescent. They varied also in the hairy surface, some having the hairs short and upright, others long and bent down. The Hunter _Nephele_ could not be distinguished from _Alope_ by any permanent character.
The Illinois larvæ were deep yellow-green after second moult, and the side stripe was always present and distinct. The hairs were never long and bent. The larvæ were distinguishable from all the others.
Comparing chrysalids:
One Albany _Alope_, A; length .56 inch, greatest breadth .21 inch; color deep green, covered with smooth specks and patches of a lighter color, but which scarcely affect the general green hue; top of head case, ridge of mesonotum and ventral edges of wing cases cream color. This was the only one I obtained, and it produced a male butterfly.
One Hoboken _Alope_, same size and color, and produced a male.
One Hunter _Nephele_, length .6, br. .2 inch; was precisely like the Albany _Alope_ in appearance, and produced a male.
Another Hoboken _Alope_; color yellow-green, and on the dorsum were three longitudinal yellow bands, one on middle of abdomen, ending at base of mesonotum, the others sub-dorsal, extending from last segment to head. This died before imago.
One Coalburgh _Alope_; length .8, br. .24 inch; bright yellow-green, covered with the lighter specks and patches, but not so as to obscure the ground; the wing cases clouded with darker green in long stripes; the three yellow dorsal bands as in the _Alope_ last mentioned; edges of head, wing cases and mesonotum cream color. This produced a female butterfly, with broad yellow band and like the parent.
Another Coalburgh _Alope_, length .6, br. .22 inch; like the foregoing, being both banded and clouded. Produced a female butterfly, with broad yellow band.
One Hunter _Nephele_, B; color yellow-green, bands and clouding of wings present but indistinct. Produced a female.
Three Hunter _Nephele_; all yellow-green, with no bands or clouding; the edgings cream color. These all gave males. The chrysalis described CAN. ENT., ix., 143, produced a female, but showed no band or clouds; the edgings cream color.
Two Illinois examples; length .6, br. .22 inch; color a pale blue-green, the powdery covering giving a whitish hue to the whole; no bands or clouds; the edges of mesonotum, head and wing cases white. Both these gave males. I obtained no females from this lot.
Summary as to chrysalids:
The largest _Alope_ and Hunter _Nephele_ were alike in color and in dorsal stripes, clouds on wings, and edgings of head case, etc.; but the bands and clouds were most distinct in _Alope_. All these large chrysalids produced female butterflies. The plainer and smaller chrysalids were male. But one female _Nephele_ chrysalis is recorded as without clouds or bands.
The Illinois chrysalids were of same shape as the rest, but were small and plain colored, and were blue or whitish-green; the edgings white instead of cream color. They were readily to be distinguished from any other.
Results in butterflies:
The chrysalis A, Albany _Alope_, gave a male not differing from many males taken at Hunter, and which there I always regarded as true _Nephele_, though off type, being without band, but with a narrow yellow nimbus about the ocelli and connecting them, the edges everywhere fading into the black ground.
On the other hand, the chrysalis B, Hunter _Nephele_, gave a typical female _Alope_, with a broad and clear yellow band. The female which emerged in 1877 from the Hunter _Nephele_ before spoken of had both ocelli surrounded and connected by yellow, and stood midway between the types of the two forms.
The two chrysalids from Illinois, as I have said, gave males; one wholly dark, the irides without rings; the other had a faint russet nimbus about them, and over the intervening space was a tint of russet.