Article beginning on page 219.
Psyche 7:219-228, 1894.

Full text (searchable PDF)
Durable link: http://psyche.entclub.org/7/7-219.html

The following unprocessed text is extracted from the PDF file, and is likely to be both incomplete and full of errors. Please consult the PDF file for the complete article.

PSYCHE.
A COMPARISON OF COLIAS HECLA WITH C. MEADXI AND c. ELIS.
BY THOMAS E. BEAN, T.AUGAN, ALBERTA PROVINCE, CANADA. [Annual address of the retiring president of die Cambridge Entomological Club, 8 March, 1895.j As the males of Meadti and &is ai c
but narrowly separate, I have made it
series of measurements from which to
derive numerical averages in the two
species, as to total expanse of the front pair of wings, as to breadth of dark
border of primary at middle oS outer
margin, and particularly as to the pro-
portion between this breadth of dark
border and the expanse of the corres-
ponding wing. These tests were also
applied to Hecla, althoi~g'h there
scarcely of equal significance on
account of the small number of
examples at my command. Mr. Davici
Bruce very kindly sent me a large
series of f&%adii for cxaiviination, so
that I have been en;ibled to compare
42 (f , 23 9 yfcadti with 7 8, 5 9
Hecia and 56 (f, 75 9 å£'/is The
males were individually measured, and
the data reduced to general averages,
with following results.
Average total expanse (sum of the
length of the two primaries plus breadth of body) : MeadZ, nearly 47 inm. ;
Hecla, nearly 4'; mm, ; Elis, neaily
50 mm.
Average breadth of clink border of
primary, at middle of outer margin:
Mdii, 4; mm. ; Hecia, å´1s inn). ;
mm.
Elis. 3;
Proportion between average breadth
of c1.u-k border of primary and average
length of primary (measured from apex
to centet of base of wing) : in ,!k%ad/z' the breadth of dark border proved to be
slightly over 19& per cent of length of
primary, in Heda 13 per cent. in Elis
slightly over 14 per cent.
These are significant averages. The
individual measurements from which
they were obtained show that in regard
to expanse of wing, as in other respects, Mead// is in considerable degree less
subject to variation than Elis. Meadii,
in its extreme terms of expanse, only
varies to 4 rnm. below its average figure and to 3 tnm. above its average. I3li.r
ranges to 10 min. below and to 6 mm.
above its own average term. The
extremes in Meadii are only 7 mm.
apart, but in HecZa they are 10 mm..
apart, and EZis has a range of 16 mrn.
of variation in this dimension, or
11 mm. if one extremely abnormal
example be omitted from the com-
parison. The uniformity of expanse in




================================================================================

Meadii is remarkable. Omitting two
examples, the largest (TO mm.) and
the smallest (43 mm.), the remaining
forty show a variation in expanse of
only 4 mm., From 45 to 49 mm. In
El6 leaving" out the lowest term (one
specimen of 40 mm.) and the highest
term (two of 56 mm.), the pendulum
of variation stillswings between 45 and
54 mm., a range more than double that
of Meadii. Hecia, also, is evidently
far more variable in expanse than
Meadii, as shown even by this small
series.
A comparison of the shape of dark
border of fore-wing furnishes a valid
distinction, parting Hecia from Elis
and even more emphatically from
Meadti, while it separates less deci-
sively Blis from Meadii. In a very
large majority of the Afeadii, the dark
border of fore-wing extends a consider-
able projection toward base of wing,
both at internal angle and at apex.
Owing to this salient projection the
fore-wing is rendered extremely broad
on costa. The curvature or inner edge
of border, however, is so great that the costal excess of breadth is rapidly parted with in the backward course of the
border. From a point of fore-wing
nearly opposite the cell-spot, 1.0 a point. :I little back of posterior median
nervule, the dark border of Meadii 3
in a large majority of the specimens
examined maintains closely an equal
breadth. This is plainly the case in 39
of the 42. In the other 3 the border
narrows very slightly from the front to
the back of this median part; in one of
them on both fore wings, in the other
two on right wing only. The rule
then in MeadU-, and a rule of almost
universal application, is that the dark
border offore-wing is dispi'oportionatciy broad on costa, loses this excess of
breadth anterior to a point nearly oppo- site the discal spot, and thro~igho~it the median portion of the wing maintains
closely an equal breadth. In a con-
siderable proportion of individuals the
border abruptly narrows more or less a
little anterior to the submedian nerve,
the disk color encroaching upon the
marginal border in a broad irregular
sinus whose deepest extension usually
occurs at the s~ibmeclian nerve.
The seven males of Hecia present a
radically difl'criiig pattern of lore-wing border from that displayed by Meadti.
The excess of breadth at costa, instead
of being quickly dissipated in the back- wan1 progress of the border (as in
Meadii), is parted with very gradually
and evenly, so that Hecla's dark border
becomes progressively narrower from
costa to internal angle. This general
method is plain in all the seven, though in two of them a part of the median
extent of the border shows but a slight
narrowing within itself. In all the
seven a gradual reduction of breadth
from costa to internal angle is a sys-
tematic detail, and it would effectively part them from all these Meadii males
were there no other distinction. The
differing shape of the fore-wing dark
bonier is a far more clecisive distinction between the two species than is the
difference merely in breadth of border.




================================================================================

A ~ ~ ~ I 1a95.1 PS 2THE. 221
.Hecia in these specimens differs from
Meadii by exhibiting less tendency to
extension of the dark border inward at
costa and at internal angle : in Meadii, the color-pattern is salient and aggres- sive ; in Hecla, stationary or retrograde. r,
lhe color on veins crossing dark
border of primary seems to be a fluctu-
ating and indecisive character. A ten-
dency to show yellow scales on the
veins of anterior part of border is some- what prevalent in Meadii. while in
Hecla more commonly than in Meadii
the posterior part of wing also exhibits yellow veining on the border. The
present, material indicates that when
the yellow veining occurs in Hecia it
will generally be somewhat uniformly
presented throughout the border, but in
Meadii usually predominant toward
apex.
The male of Elis approaches that of
=adii so closely that individusils of
the former can be found which scarcely
seem to dil'Tcr tangibly from the latter: by a vague contrast in general appear-
ance they part from Meadii and iden-
tify themselves with Ell's. This some-
what elusive difference between closely
.
similar individuals of contiguous species is difficult of analysis. Adjoining spe- cies are most readily discriminated as
unities, by the opposition or unlikeness of the entire systems or coml>inations of pattern, containing in part closely
similar or identical elements, and in
part alien elements. A large series of
each tends to manifest the entire
amount of alienation separating each
species from its nearest allies. Single
individuals are at best imperfectly rep- resentative ; being themselves special-
ized fragments, they merely represent
what the species would be if similarly
specialized tl~roughout. In estimating
the relation between adjacent species
adequate series should be examined.
Safe conclusions cannot be drawn from
comparison of isolated cxamples until
the representative value of those exani- pies has been ascertained. Individuals
are typical or divergent: in the latter
case, if divergent in the direction of the proximate species they express in
relation to that species less than a
typical amount of alienation ; if diver- gent in a direction leading away from
the proximate species, they exhibit rela- tivcly to it a degree of alienation- (bio- logical estrangement) which is greater
than the typical contrast. From this it
follows that individuals which have a
more tlian typical degree of divergence
from each other, representing adjacent
species, will over-contrast those species, and on the other hand, unusually
approximate examples in contiguous
species will under-contrast the two
species. In species so near of kin as
Elis and Meadii, the most closely
resemblant examples are exactly those
which are least representative of the
characters peculiar to their respective
species, and most highly representative
of characters common to the several
closely allied species. Elis being
greatly more variable than Meadit, the
closcst approximation between them is
on the part of exceptional males of
Elis. It may be said that Aladii as




================================================================================

222 PSYCHE. [April 1895.
a species is very close to certain excep- tional males of Elis, and conversely
that some Elis males are very near to
the species Me&. The approxi-
mation is on the part of occasional
conservatively divergent Elis males.
These divergent individuals of Bit's,
wlicn isolated from their fellows and
brought into comparison with Meadii,
seem closer to the latter than they
really are: the cause of this is, that the resemblance to Meadii is made con-
spicuous on account of the attention
being attracted to it by the presence of examples of ^lead% the greater reseti'i- blance to the species Elis being over-
looked by lack of a proper representst-
tion of Elis for comparison.
In these three species the females
compare very differently from the
males. Elis and Mead< 'which some-
times :ippr~~ach closely in the male,
manifest in the female only a moderate
general affinity, offset by a striking
antagonism of patteri~ rarely found
between such close allies in this genus. flecia and ^lea&/, l~owever, whose
males differ widely, are parted in the
female sex chiefly by fluctuating avcr-
ages, so that their contrast in that sex is comparatively vague and informal,
and in occasional individuals is found
resemblance to the allied species in a
degree which has sometimes proved
confusing.
In the border of primary in Meadii
the yellow spots are ordinarily irregular in the extreme in size and outline; and
they are sometimes hazy or nebulous,
and extremely pallid, - two such
examples are in this series. The vari-
ance between individuals as to degree
of development of the yellow spots is
great. There is a decided tendency to
total eclipse of the spots, a tendency so influential that in seven specimens of
this small lot the border is almost
entirely solid black. This species
exhibits great eccentricity in the pre-
sentation of the maculae, which are
frequently of irregular shapes, often
indeed peculiarly erratic in outline. It is the far rarer instances, with tolerably neat and conip;iratively even-sized
spots, which closely resemble some
Hecia; but such Meadii are not at all
representative examples. Thc cccen-
tricity of . maculation spoken of is
foreign to the method of Elis, and it is a peculiarity of which my small series
of IlccZa contains no suggestion. In
the maci~lation of h'ecln., as comparcd
with that of Mead-it, the spots are more equal in the same individual, and more
uniform, taking one example will1
another.
In both Meadii and Hecla the yellow
spots on dark border of primary are
iineqii~illy developed, the mid-wing
spot being often obsolete, and almost
invariably at least feeble in dcvclop-
merit. This is the established method
in both species.
Elis is remarkable for its symmetry
of pattern, the maculation being in
general conspicuously harmonic in the
individual, and the middle spot having




================================================================================

approximate equality nit11 the others.
Examples of Elis in which the border
spots of primary are not pretty uniform
in size and shape. and those in which
the inid-wing spot is not at least mod-
erately well developed. are exceptional. The most informal in this series of Jilts have the spots more neatly regular than
in the most formal Meadii in the lot.
Those Meadii with the spots largest
are not ncarci- to /?//.Y on that account, but rather show stronger contrast, as
tlie diverse system of maculiition in the two species is thereby rendered more
obvious.
An occasional Me&, having the
spots on primary border more symmet-
rical than is usual, closely resembles
some Hecla, so that in a large scries of both a small percentage of the Meadii
might prove difficult to part from a
few of the Hecla by this test solely.
In a large majority of cases, however,
the two females difl'er widely in chsir- acter of dark border of hind-wing : 18
out of 23 of these Meadii have a prac-
tically solid black hind-wing border,
and 3 of these 5 Hecia have a most
distinctly maculated border with tlie
yellow spots large and conspicuous.
yfcadii with hind-wing border partly
maculated, and Hecia with niaculation
of liincl-wing border partly obscured,
might not readily separate 011 this
character alone. It follows then, that
among the sni;ill number not distin-
guishablc by characters of Core-wing
aclei : that is to say, a percentage of a percentage occurs wherein the rnacu-
lation of dark border of both primary
and secondary is liable to prove too
/ipp~-oxiin~te or too obscure to rely
ipon in determining the position of the
individ~~iil. Even these rare instances
of close approximation should not
defeat a student familiar with the two
species. Other specialties of pattern
are in some degree indicative, and it
rarely indeed occurs that i-ill significant distinctions lapse in a single individual. There remains also that indefinite but
obvions Fact known as " ttic general
appearance," which fnrnishes an expe-
rienced observer with the most reliable
and satisfactory guide in distinguishing one species from anullier.
In a general view of the three species,
the most obvious fact is that fJda
male is distinct from the respective
males of ATeadii and Bit. Also,
properly rcprcscnt~itive material makes
conspicuously evident the divergence
of the female of EHs from the females
of Hecla and Meadii. The close
approximation occurs between Hecia
and Meadii in the fcrnalc, between
Etis and Meadii in the male. This
complex of relationship is perhaps
partly capable of explanation. &la
may safely be considered the oldest of
the tlirec species, zinc1 Elis the
youngest. In Colias the law of the
agency of sex in race progress appears
to be that the male is the conservative
border, one individual in many may and tlie female the progressive sex. also fall in the small proportion with Accordingly, in the alliance under con- hind-wing border of ambiguous char- sideration, IlecZa being the oldest



================================================================================

224 PS 2'CHE. [April 1895.
species, its inale should be the most
conservative element in the allianw ;
ncl Bits being the youngest species,
its female should prove to be the most
progressive element. Hecia male and
Blis female occupy outposts as to race
progress, iincl are thus of necessity more completely ditTere~itiated than the other elements of the alliance.
The comparison so far is from notes
made several yearsap. Having lately
re-exainincd all descriptions and fig~ires of Hecliz available to me here in the
wilderness, I am inclined to think my
analysis of the figure-pattern accurate. In order to amplify my knowledge of
Hecla, I applied to Dr. Strecker for
details in regard to the fiecia males in ]is collection. making very specific
enquiry as to shape of dark border of
primary. In response three pen
sketches were sent me, representing the
range of variation observed in the
Lapland and Greenland examples.
These drawings show a method of
border corresponding to that found in
ny own series and described in the
present analysis. Another sketch rep-
resented the border of primary of
Meadii, delincatii~g precisely the style of dark border I have described its
practically the constant fashion in that species.
In print, as also in correspondence,
Mr. W. H. Edwards refers to the
general ground color of some ^fc&
as indistinguishable froin thsit of Hecla, or of some Hec?a! ancl liales this rescni- hlance into court as EI witness to prove yieadii and I'lccia one species. Hec7a
was already well known when Mr.
Edwards named Meadiz', and if there
is now no valid distinction between
Meadii and FIecla there was none
then, and in- that case A-Ieadii should
not have been described. In Bntter-
flies N. A., 1st series, Mr. Edwards
closes thus his A-feadii text : *. Tins
Colias bears close resemblance to Hecla, but may be readily distinguished by
the glandular spot before spoken of
(see Plate), a character not found in
Hecla, and so decisive that it is not
necessary to point unt minor points of
difference." At this late day Mr.
Edwards seems to find but one item of
difference between the two species, -
the å ´ g1andul:ir spot "-and it is as
evident an embarrassment to him at
present as it was convenient formerly.
Probably it is not unfair to say that
Meadii when named was concluded
distinct from tiecla simply on account
of two circumstances: one being the
presence of the '-glandular spot," the
other the fact that the original catch of Meadii consisted of specimens colored
a redder orange lliau Hech ordinarily
displays. Mr. Edwds is open to
criticism in both his earlier and later
attitudes 111 regard to Meadii. Con-
sidering Xeadi-i distinct froni Hecla
bccausc the original matcriiil of fVfeadii was of a redder orange thm that cus-
toinary in Hecia was the initial error.
The acceptance of this color F act as a
proof of dis~inctness was incorrect in
two ways: first, because dependent on
the violent assumption that all other
Meadii would prove of as red an orange