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dominant factor attaching itself to a wild species in recent times. None of the cases are
satisfactorily recorded or analysed as yet, but the evidence is clear that doubledayaria is a
dominant to its type, and in several other dark varieties, though the pigment deposited is
not black, the records show that the increased amount of the pigment almost certainly is
due to a positive factor. Of this, Hemerophila abruptaria is a good example.[24] There are
some irregularities in the results, but taken together they leave little doubt that the dark
brown variety is a dominant and the light, yellowish brown a recessive.
A curious parallel to the rise of the melanic moths in England is provided by the case of
the Honey-creepers or Sugar-birds, in certain West Indian islands.[25] These birds of the
genus Coereba (Certhiola) range from Southern Mexico to the Northern parts of South
America and through the whole chain of the West Indian islands and Bahamas except
Cuba. There are numerous local forms, and many of the islands have types peculiar to
themselves, as is usual in such cases. Some of the types or species range through several
islands, but according to Austin Clark[26] no island has more than one of them. Cory[27]
reckoned twelve such species within the Antillean region. They are small birds about the
size of a nuthatch with a general colouring of black, yellow, and white. From the island of
St. Vincent the Smithsonian Institution received in the late seventies of last century several
completely black specimens in addition to two of the usual type of colouring. The black
were described by W. N. Lawrence as atrata, and those marked with the usual yellow and
white were called saccharina. The collector (Mr. F. A. Ober) reported that the black form
was common, and that the saccharina form was rarer. Lawrence remarks, "Had there been
only a single example (of the black form) I should have considered it as probably a case of
abnormal colouring, but it seems to be a representative form of the genus in this island."
[28] There is of course no doubt of the correctness of the view taken by Austin Clark that
"atrata" is a black variety. The black bird is in every respect, other than colour, identical
with saccharina, and it is even possible to detect a greenish colour in the areas which
would normally be yellow, showing plainly enough the yellow pigment obscured by the
black.
We have next the interesting fact that like our melanic moths the dark form is replacing
the "type." At the time of Ober's visit the type was already in a minority, but now it is
nearly or perhaps actually extinct, though the black form is one of the commonest birds on
the island. Austin Clark found no specimen when he collected there in 1903-4, though
formerly it was not uncommon in the vicinity of Kingston and in the immediate windward
district of St. Vincent.
The Grenadines are geographically just south of St. Vincent, though separated by a
deep channel. In these islands no black forms have yet been taken, but Grenada, the next
island to the south, has both normals and blacks. There are trifling differences of size
between the Grenada birds and those from St. Vincent, the Grenada specimens being
slightly smaller and for this reason they have received distinct names, the form marked
with yellow and white being called Godmani (Cory) and the black, Wellsi (Cory), but this
merely introduces a useless complication. There is evidence that in Grenada, as in St.
Vincent, the black is gradually ousting the original type, but the process has not gone so
far as in St. Vincent. Austin Clark very properly compares this case of the Sugar-birds with
that of Papilio turnus, which as is well-known, has a black female in the southern parts of
its distribution, in addition to a female of the yellow type, but in the Northern States the
black female does not occur.
During the present year P. R. Lowe, who lately studied Coerebas on a large scale in the
West Indies, has published an important paper on the subject.[29] He calls attention to the
fact that Cory recently found a black form of Coereba on Los Roques Islands, and he
himself discovered another on the Testigos Islands. Both localities are on the coast of
Venezuela, far from St. Vincent and Grenada. The whole problem is thus further
complicated by the fact that the black varieties have, as we are almost driven to admit,
arisen independently in remote places. Improbable as this conclusion may be, it is still
more difficult to regard all the black forms as derived from one source. For first, they
present definite small differences from each other; and secondly we have to remember a
consideration of greater importance, that the very fact that each island has its own type
must be accepted as proving that the localities are effectively isolated from each other, and
that migration must be a very rare event.
The rarity of such illustrative cases is, I believe, more apparent than real. It is probably
due to the extreme reluctance of systematists to admit that such things can be, and of
course to the almost complete absence of knowledge as to the genetic behaviour of wild
animals and plants. Only in such examples as this of the Coereba, where colour constitutes
the sole difference, or that of the moths which have been minutely studied by many
collectors, does the significance of the facts appear. The arrangement of catalogues and
collections is such that much practical difficulty of a quite unnecessary kind is introduced.
For example, in this very case of Coereba, I find the British Museum has a fine series from
Grenada including 3 normals and 11 black, and also 16 blacks from St. Vincent. If the black
specimens from Grenada were put with the normals which are almost certainly nothing but
a recessive form of the same bird, the variation would strike the eye on even a superficial
glance at the drawer. But following the notions so naively expressed in the passage quoted
above from W. N. Lawrence, the blacks from Grenada are put apart together with the other
blacks from St. Vincent, though two of them were shot on the same date as one of the
normals.
 CHAPTER VII
Local Differentiation. Continued

Overlapping Forms
The facts of the distribution of local forms on the whole are consistent with the view
that these forms come into existence by the sporadic appearance of varieties in a
population, rather than by transformation of the population as a whole. Of such
sporadically occurring varieties there are examples in great abundance, though by the
nature of the case it can be but rarely that we are able to produce evidence of a previous
type being actually superseded by the variety. When the two forms are found co-existing in
the same area they are usually recorded as one species if intergrades are observed, and as
two species if the intergrades are absent. On the other hand when two forms are found
occupying separate areas, when, that is, the process of replacement is completed in one of
the areas, then forthwith each is named separately either as species or subspecies.
Successive observations carried out through considerable periods of time would be
necessary to establish beyond question that the history proceeds in one way rather than
another. Such continuity of observation has for the most part never been attempted. The
kind of information wanted has indeed only been lately recognized, and really critical
collecting is a thing of only the last few decades. The methods of the older collectors, who
aimed at bringing together a few typical specimens of all distinct forms, are of little service
in this class of inquiry, which is better promoted by the indiscriminate collection of large
numbers of common forms from many localities. When this has been done on a
comprehensive scale we shall be in a position to form much more confident judgments as
to the general theory of evolution.
Some little work of the kind has however been done and the results are already of
great value. Seeing that the differentiation of local forms is only made possible by isolation,
it necessarily happens that the collector finds one form in one locality and another in a
distinct locality, and there is no evidence as to the behaviour which the two representative
species might exhibit if they came into touch with each other. In the most familiar
examples of such distinction each inhabits an island, completely occupying it to the
exclusion of any other similar form. It can only be when the two representative species
occupy parts of a continental area connected with each other by regions habitable for the
organism in question, that there is a chance of seeing the two forms in contact. Often also,
even where this condition is satisfied, the habits, social organisation, or some other special
cause may act as a barrier which prevents the distinguishable forms from ever coming into
such complete contact as to interbreed or to behave as a genetically continuous race.
When genetic continuity is ensured by a constant diffusion of the population over the
whole area which they inhabit there will manifestly be no formation of local races. The
practical uniformity, for example, of so many species of birds which inhabit widely
extended ranges of Western Europe is doubtless maintained by such constant diffusion.
When, as in the case of the Falcons, many localities have peculiar forms, the fact may be
taken as conclusive evidence that there is little or no diffusion; and when we find in such a
species as the Goldfinch that in spite of migratory fluctuations there are nevertheless
geographical races fairly well differentiated, it may similarly be inferred that these
fluctuations habitually move up and down on paths which do not intermingle. There are
however a few examples of animals, not given to much irregular wandering, which occupy
a wide and continuous range of diversified country and are differentiated as local races in
two or more districts, though the distinct races meet in intervening areas. Of these the
most notorious illustration which has been investigated with any thoroughness is that of
the species of Colaptes (Woodpeckers) known in the United States as Flickers. The study of
the variations of these forms, made by J. A. Allen[1] is an admirable piece of work, with
which every student of variation and evolutionary problems should make himself familiar.
The two forms with which we are most concerned are known as C. auratus and C. cafer,
and are very strikingly different in appearance. In size, proportions, general pattern of
colouration, habits, and notes, the two are alike, but they differ in the following seven
respects as stated by Allen.
Auratus Cafer

1. Quills yellow. 1. Quills red.

2. Male with a black malar stripe. 2. Male with a red malar stripe.
Adult female with usually a brown malar
3. Adult female with no malar stripe. 3.
stripe.
4. A scarlet nuchal crescent in both sexes. 4. No nuchal crescent in either sex.
5. Throat and fore neck brown. 5. Throat and fore neck grey.
6. Whole top of head and hind neck grey. 6. Whole top of neck and hind neck brown.
General plumage with an olivaceous
7. 7. General plumage with a rufescent cast.
cast.
These differences are illustrated in the accompanying coloured plate, which has been most
kindly prepared for me under the instructions of Dr. F. M. Chapman of the American
Museum of Natural History. Before going further it is worth considering the nature of these
differences a little more closely. All but the last are large differences which no one would
overlook even in a hasty glance at the birds. If the only distinction lay in the colour of the
quills we might feel fairly sure that auratus was a recessive form of cafer, and so probably
it is in this respect. Similarly the black malar stripe of auratus is in all probability recessive
to the red malar stripe of cafer and I imagine the pigments concerned are comparable with
those in the Gouldian Finch (Poephila gouldiae) of Australia. Both sexes in that species may
have the head black, red, or, less often, yellow, and though it is not any longer in question
that birds may breed in either plumage, I believe that the young are always black-headed
and I imagine that those which become red-headed possess a dominant factor absent from
the permanently black-headed birds.[2] Yellow as a recessive form of a red is certainly very
common, but red and black as variants of the same pigment are less usual. In the Gouldian
Finch we seem to have a case where a pigment can assume all three forms. It would be
interesting to know whether the red of the malar stripes in Colaptes is a pigment of the
same nature as the red of the quills. Both in Colaptes and in Poephila gouldiae I have seen
specimens intermediate between the black and the red, and the appearance of the part
affected was exactly alike in the two cases, red feathers coming up among the black ones,
and many feathers containing both red and black pigments mixed together.
The development of the scarlet nuchal crescent in auratus and the absence of this
conspicuous mark in cafer constitute from the physiological point of view the most
remarkable pair of differences. When the red crescent is not formed, the feathers which
would bear it are exactly like the rest, and no special pigment is visible in them which one
can regard as ready to be modified into red. If the crescent is due to a factor it must
therefore be supposed that this factor has the power of modifying the pigment of the neck
in one special place alone. Dr. W. D. Miller called my attention to the fact that a similar
variation occurs in another American woodpecker, the Sapsucker, Sphyropicus varius.[3]
I do not suggest that such variations are without parallel: indeed in P. gouldiae the
factor which turns the black of the head into scarlet affects one special region of the black
only, being sharply distinct from the unmodified black of the throat. These regions of the
head are however often the seat of special colours in birds.[4] So also may be instanced
the variety of the Common Guillemot (Uria troile) which has a white line round the eyes
and at the sides of the head where the normal has no such mark; but this line is formed in
a very special place, the groove joining the eye to the ear, whereas the feathers of the
nuchal crescent are not ostensibly distinguished from those adjacent.[5]
The transposition of the brown and the grey on the back and front of the neck also
constitutes a very remarkable difference. If either grey or brown depends on a factor then
it must be supposed that auratus has one of these factors and cafer the other.
From these several considerations it is quite clear that if auratus and cafer are
modifications of the same type produced by presence or absence of factors, several
independent elements must be concerned, and to unravel their inter-relations would be
most difficult even if it were possible to breed the types under observation, which is of
course quite beyond present possibilities.
The distribution of the two is as follows. On the east side of the Continent C. auratus,
relatively pure, occupies the whole of Canada and the States from the North to Galveston.
Westward it extends across the whole continent in the more northern region to Alaska, but
in its pure form it only reaches down the Pacific coast to about the northern border of
British Columbia. Its southern and western limit is thus roughly a line drawn from north of
Vancouver, southeast to North Dakota and then south to Galveston. C. cafer in the
comparatively pure form inhabits Mexico, Arizona, California (except Lower California and
the opposite coast), central and western Nevada, Utah, Oregon, and is bounded on the
east by a line drawn from the Pacific south of Washington, south and eastward through
Colorado to the mouth of the Rio Grande or the Gulf of Mexico. Between the two lines thus
roughly defined is a band of country about 1,200-1,300 miles long and 300-400 miles
wide, which contains some normal birds of each type, but chiefly birds exhibiting the
characters of both, mixed together in various and irregular ways. Even in the areas
occupied by the pure forms occasional birds are recorded with more or less indication of
characteristics of the other form, but within the area in which the two forms are
conterminous, the mixed birds are in the majority. The condition of these birds of mixed
character is described by Allen as follows:
 "As has been long known—indeed, as shown by Baird in 1858—the
'intermediates' or 'hybrids' present ever-varying combinations of the
characters of the two birds, from individuals of C. auratus presenting only
the slightest traces of the characters of C. cafer, or, conversely—
individuals of C. cafer presenting only the slightest traces of the
characters of C. auratus—to birds in which the characters of the two are
about equally blended. Thus we may have C. auratus with merely a few
red feathers in the black malar stripe, or with the quills merely slightly
flushed with orange, or C. cafer with either merely a few black feathers in
the red malar stripe, or a few red feathers at the sides of the nape, or an
incipient, barely traceable scarlet nuchal crescent. Where the blending of
the characters is more strongly marked, the quills may be orange-yellow
or orange-red, or of any shade between yellow and red, with the other
features of the two birds about equally blended. But such examples are
exceptional, an unsymmetrical blending being the rule, the two sides of
the same bird being often unlike. The quills of the tail, for example, may
be part red and part yellow, the number of yellow or red feathers varying
in different individuals, and very often in the opposite sides of the tail in
the same bird. The same irregularity occurs also, but apparently less
frequently, in the quills of the wings. In such cases the quills may be
mostly yellow with a few red or orange quills intermixed, or red with a
similar mixture of yellow. A bird may have the general colouration of true
cafer combined with a well-developed nuchal crescent, or nearly pure
auratus with the red malar stripes of a cafer. Sometimes the body
plumage is that of C. auratus with the head nearly as in pure cafer, or
exactly the reverse may occur. Or we may have the general plumage as in
cafer with the throat and crown as in auratus, and the malar stripe either
red or black, or mixed red and black, and so on in almost endless
variations, it being rare to find, even in birds of the same nest, two
individuals alike in all their features of colouration. Usually the first trace
of cafer seen in auratus manifests itself as a mixture of red in the black
malar stripe, either as a few red feathers, or as a tipping of the black
feathers with red, or with merely the basal portion of the feathers red.
Sometimes, however, there is a mixture of orange or reddish quills, while
the malar stripe remains normal. In C. cafer the traces of auratus are
usually shown by a tendency to an incipient nuchal crescent, represented
often by merely a few red-tipped feathers on the sides of the nape; at
other times by a slight mixture of black in the red malar stripe."
Such a state of things accords very imperfectly with expectations under any received
theory of Evolution. As in some of the instances discussed in the first chapter we have here
two fairly definite forms, nearly allied, which on any evolutionary hypothesis must have
been evolved either the one from the other, or both from a third form at a time not very
remote from the present, as time must be measured in evolution. Yet though intermediates
exist in some quantity, no one can for a moment suggest that they are that definite
intermediate from which auratus and cafer descend in common. One cannot imagine that
the immediate ancestor of these birds was a mosaic, made up of asymmetrical patches of
each sort: but that is what many of the intermediates are. It is not much easier to suppose
the ancestor to have been a nondescript, with a compromise between the developed
characters of each, with quills buff, malar stripes neither black nor red, with a trace of
nuchal crescent, and so on. Such Frankenstein-monsters have played, a considerable part
in the imaginations of evolutionary philosophers, but if it were true that there was once a
population of these monsters capable of successful existence, surely they should now be
found as a population occupying the neutral zone between the two modern forms. Yet,
though much remains to be done in clearing up the facts, one thing is certain, namely that
the neutral zone has not a definite and normally intermediate population, but on the
contrary it is peopled by fragments of the two definite types and miscellaneous mongrels
between them.
On the other hand, one cannot readily suppose that either form was the parent of the
other. The process must have involved both addition and loss of factors, for whatever
hypothesis be adopted, such changes must be supposed to have occurred. A careful
statistical tabulation of the way in which the characters are distributed in the population of
the mixed zone would be of great value, and till that has been done there is little that can
be said with certainty as to the genetics of these characters. In the collection of Dr. Bishop
of New Haven I was very kindly allowed to examine a sample, all taken at random, near
together, in Saskatchewan. There were females 4 adult, 2 young; males 4 adult and 5
young. This number, though of course insufficient, is enough to give some guide as to the
degree of definiteness which the characters generally show in their variations. Of the 15
birds, 8 had simply yellow quills; 2 had red; 1 was almost red but had one yellow tail-quill;
3 were intermediate and 1 was buff. As regards the malar patch, which can only be
determined properly in the adult males, 1 was red, 1 was approximately red, 2
intermediate. As to nuchal crescent 4 females had none, 2 females very slight; 7 males had
it, 1 had only a slight crescent, and 1 had none. In point of quills therefore 10 were
definite out of 15; in point of crescent, 11 were definite out of 15; and in point of malar
patch 1 only was definite out of 4. The last is a feature directly dependent on age and so
counts for less, but as regards the other two features there is some indication that the
factors show definiteness in their behaviour. It must be remembered that we have no
knowledge what the heterozygous form may be, and in the case of red and yellow it is
probably a reddish buff. The patch-works are no doubt to be compared with other well-
known pied forms, and in these we must suppose the active factor broken up, which it
probably can be very easily. The asymmetry, which Allen notices as so marked a feature, in
the distribution of the red and yellow quills of the tail especially, recalls that of the black
markings in the pied Canaries. As is well known to students of variations some pigment-
factors in some animals are apparently uncontrolled by symmetry, while in other specific
cases symmetry is the rule. On the other hand the blackness or redness of the malar
patches is, I think, as a rule nearly symmetrical. It should be mentioned that two of Dr.
Bishop's young birds belonged to the same nest, one a female with red quills, the other a
male with yellow. Both are without crescent.
As to the question whether certain combinations of characters occur with special
frequency, the evidence is insufficient to give a definite answer. Among all the birds I have
seen in America or in England I have not yet found one having the malar patches black
without any nuchal crescent. Of Dr. Bishop's 8 adults not one, however, showed the
combination of the three chief features normal for auratus or for cafer.
 Besides the two forms that we have hitherto considered, several other local types exist,
and these throw some further light on the problem. Of these the most important in this
connexion is chrysoides, which inhabits the whole of southern California and the mainland
opposite. This remarkable form is as Allen says, very different from auratus except that it
has the quills yellow like auratus, not red like cafer. So that we find here in the extreme
west of the whole distribution a type agreeing in one of its chief features with the eastern
type. Between this and cafer intergrades have, according to Allen, not been found. The
relations of this chrysoides are, Allen thinks, rather with mexicanoides, a southern, smaller
race with colours more intense, which inhabits Guatemala, but however that may be, it
must be regarded as a cafer which has lost its red quills. The island of Guadeloupe off
Lower California has an island form. Beyond the other side of the continent there is also an
island form of auratus, inhabiting Cuba, so that clearly the yellow quills can extend into the
tropics.
The above account is in many respects incomplete, but it suffices to give an outline of
the chief facts. The whole problem is complicated by the undoubted effects of an uncertain
amount of migration, and in many, perhaps all, districts, the winter population differs from
the summer population of the same localities. The existence of these seasonal ebbs and
flows is now well known to ornithologists, and most of the bird species of temperate
regions are subject to them.
Difficult as it may be to conceive the actual process of origin of the two types auratus
and cafer, it is I think still harder to suggest any possible circumstance which can have
determined their development as distinct races, or which can maintain that distinctness
when created. Some will no doubt be disposed to appeal once more to our ignorance and
suggest that if we only knew more we should see that the yellow quills, the black
"moustache" and the red crescent, specially qualify auratus for the north and eastern
region, and the red quills, red "moustache" and absence of crescent fit cafer to the
conditions of its homes. Each can judge for himself, but my own view is that this is a vain
delusion, and that to cherish it merely blunts the receptivity of the mind, which if
unoccupied with such fancies would be more ready to perceive the truth when at last it
shall appear. Think of the range of conditions prevailing in the country occupied by auratus
—a triangle with its apex in Florida and its base the whole Arctic region of North America.
Is it seriously suggested that there is some element common to the "conditions" of such an
area which demands a nuchal crescent in the Flickers, though the birds of the cafer area,
almost equally varied, can dispense with the same character? Curiously enough, the
geographical variation of Sphyropicus varius, another though a very different
Woodpecker[6] shows that conversely the nuchal crescent can be dispensed with in the
Eastern form though it is assumed by the Western.[7]
Allen points out the interesting additional fact that superposed upon each of the two
distinct forms, auratus and cafer, are many geographical variations which can very
naturally be regarded as climatic. Each decreases in size from the North southward, as so
many species do.[8] They become paler in the arid plains, and show the ordinary phases
which are seen in other birds having the same distribution. Such differences we may well
suppose to be determined directly or indirectly, by environment, and we may anticipate
with fuller knowledge it will be possible to distinguish variations of this nature as in the
broad sense environmental, from the larger differences separating the two main types of
Colaptes, which I surmise are altogether independent of such influences.
It is generally supposed that phenomena like those now so well established in the case
of Colaptes are very exceptional, and as has already been stated a number of
circumstances must combine in order that they may be produced. I suspect however that
the examples are more numerous than is commonly thought. In all likelihood the three
forms Sphyropicus varius, nuchalis and ruber are in a very similar condition though the
details have not, so far as I know, been worked out. A complex example which is closely
parallel to the case of Colaptes was described by F. M. Chapman[9] at the same date as
Allen's work. This is the case of Quiscalus, the Grackles, which in the North American
Continent have three fairly distinct forms which Chapman speaks of as Q. aeneus, Q.
quiscula, and Q. quiscula aglaeus. The birds are all, so far as pigment is concerned, dark
blackish brown, but the head and mantle have superposed a metallic sheen of
interference-colours which in the various forms take different tints, bluish green, bronze
green, or bronze purple. The details are complicated and difficult to appreciate without
actual specimens, but the two common types are sufficiently distinct. The birds inhabit the
whole area east of the Rockies, quiscula aglaeus occupying Florida and the Southern States
southwest of a band of country about a hundred miles broad extending roughly from
Connecticut to the mouth of the Mississippi; and aeneus taking the area north and west of
this band. In discussing this case Chapman expresses the same view as Allen does in the
Colaptes case, that there are two distinct populations, substantially fixed, and that the
band of country in which they meet each other has a mongrel population, with no
consistent type, but showing miscellaneous combinations of the character of the two chief
types.
The warblers of the genus Helminthophila provide another illustration which has points
of special interest. The two chief species are H. pinus, which has a yellow mantle and
lower parts, white bars on the wings, a black patch behind the eyes and a broad black
mark on the throat; and H. chrysoptera with dark grey mantle and pale whitish grey lower
parts, yellow bars on the wings, and grey marks on cheeks and throat where pinus has
black. These two birds are exceeding distinct, and in addition their songs are quite unlike.
H. pinus ranges through the eastern United States up to Connecticut and Iowa. H.
chrysoptera is a northern form extending down to Connecticut and New Jersey. Both are
migrants.
In these two States, where the two types overlap, certain forms have been repeatedly
found which have been described as two distinct species, Lawrencei and leucobronchialis.
Dr. L. B. Bishop and Mr. Brewster showed me two long series of Helminthophila containing
various intergrades between the four named kinds, and details regarding these may be
found in Chapman's North American Warblers and in Dr. Bishop's paper in Auk, 1905, XXII.
Though the characters evidently break up to some extent, the series can be represented as
due to recombinations of definite factors more easily than the others which I have
described. The differentiating characters are:

Pinus Chrysoptera
 1. Mantle and lower parts yellow (Y1). 1. Mantle and lower parts grey (y1).
2. Wing-bars white (y2). 2. Wing-bars yellow (Y2).
3. Cheek and throat not black (b). 3. Cheek and throat black (B).

The grey pigment of the mantle is common to both, but is masked by the yellow in
pinus, the net result being an olive-green.[10]
I am much indebted to Dr. F. M. Chapman for the loan of the coloured plate in which
these distinctions are shown. It first appeared in his book, North American Warblers.
We cannot tell whether yellow or not-yellow is due to the presence of a factor, but we
may suppose that one or other gives the special colour to the parts. The black of character
3 is no doubt a dominant. Thus pinus becomes Y1y2b and chrysoptera in y1Y2B. The
Lawrencei which has the underparts yellow, wing-bars white, and black patches is Y1y2B
and leucobronchialis which has mantle and underparts not-yellow, wing-bars yellow and no
black patches is y1Y2b. This representation, it should be clearly understood, is tentative
and approximate only. The characters are not really sharp, for there is much grading; but
allowing for the effects of heterozygosis and for some actual breaking-up of factors I
believe it gives a fairly correct view of the case. In particular we can see how it meets the
difficulty which Chapman felt in accepting leucobronchialis as in any sense derived from
pinus which has a yellow breast, and chrysoptera which has a black throat, seeing that
leucobronchialis has neither. We now recognize at once that this form could be produced
by ordinary re-combination of the absence of Y1 with the absence of B.
I note also with great interest that the modern observers agree that the so-called
hybrids may have the song either of the one species, or of the other, or a song
intermediate between the two. It may also be added that these two types have several
times been seen, in the breeding season, paired with each other or with one of the other
combinations.

Fig. 1. Helminthophila pinus, male.

Fig. 2. Helminthophila pinus, female.
Fig. 3. "Lawrence's Warbler," male; one of the integrading forms.
Fig. 4. "Brewster's Warbler," male; another of the integrading forms.
Fig. 5. Helminthophila chrysoptera, male.
Fig. 6. Helminthophila chrysoptera, female.
 Allen[11] has described another excellent American example, the Tits of the group
Baeolophus bicolor-atricristatus. The form bicolor belongs to the eastern States and ranges
from the Atlantic coast to the Great Plains, and atricristatus, of east Mexico, extends from
Vera Cruz to central Texas. In southern and central Texas the breeding ranges adjoin, and
in this country various intermediates occur. The chief types differ in two main points.
 B. bicolor B. atricristatus

Forehead varies from deep black Forehead white to buffish white.

to dull black, suffused with rusty brown.
Crown and crest grey, Crown and crest black, abruptly
slightly darker than the back. contrasting with the back.

The intergrades between the two have, as usual, received specific names. A detailed
description is given by Allen, from which it appears that the gradation is very complete. In
one case a series of 16 adults were all intermediates. It is not stated whether the collector
took these at random, but from the local lists it is clear that the types are found not far
away from the place where the intergrades were shot.
Another very striking case is that of the Tanagers, of the genus Rhamphocoelus. In this
group there are several local forms which are related to each other in remarkable ways.
The forms known as passerinii and icteronotus exhibit the clearest phenomena of
intergradation. The species passerinii has a brilliant scarlet and black male, and it inhabits
Honduras and Nicaragua. Proceeding southwards along the isthmus we find next
costaricensis which has a male like that of passerinii (but a female with more orange than
the olive-grey female of passerinii). Next we come to Panama which is occupied by
icteronotus, sharply distinguished from passerinii by the fact that the scarlet is replaced by
lemon-yellow. This same icteronotus occurs again as a pure type in Ecuador and many
other parts of South America; but Colombia, between Panama and Ecuador, contains
scarlets like passerinii, yellows like icteronotus, and various intergrades of several shades
of orange. The passerinii males from Nicaragua are indistinguishable from those of
Colombia, and the icteronotus of Ecuador are the same as those in Panama. The orange
intergrades, doubtless heterozygous forms, though collected at the same locality (Medellin
in Colombia) as several pure yellows and pure scarlets, are in the British Museum series
sorted out as a separate species under the name chrysonotus! Complications are
introduced by the relations of these forms to another named type, flammigerus, but we
may for our purpose leave that out of consideration, and say that the order of geographical
sequence from Honduras to Ecuador is (1) scarlet, (2) yellow, (3) mixture of types, scarlet,
yellow, orange, (4)yellow.
Similar examples exist in the birds of the old world, but I do not know of any that have
been studied so fully as those of America. The best known is that of the two Rollers,
Coracias indicus which spreads from Asia Minor through Persia, Baluchistan, the Indian
Peninsula and Ceylon, and affinis which ranges from Nepal, through Assam, Tenasserim
and the Indo-Chinese countries. The two types are very different and may be distinguished
as follows:

C. indicus C. affinis
 Mantle drab brown-chestnut. Dark olive-green.
Breast chestnut. Dull purple brown.
Throat purplish, streaked with white. Purple, streaked with blue.
Upper tail-coverts indigo. Turquoise.

The wings are the same in both. In the provinces of Nepal, Sikhim, and Darjiling the
two species coexist, with the result that intergrades have been frequently recorded. The
line of intergradation extends to the coast, and birds showing various combinations of the
two types from the Calcutta district exist in collections.[12] The case is interesting inasmuch
as like that of Quiscalus it shows a series of combinations of various metallic colours. Some
of these are probably evoked by the development of pigment behind striations or other
interferences already existing, but in the present state of knowledge it would be quite
impossible to suggest what the actual factors producing these appearances may be.
There are, naturally, many other cases among birds which are suspected of being in
reality comparable, but in most of them the evidence is still inadequate. Among
Lepidoptera also there are a few of these; perhaps the most striking is that of Basilarchia
"proserpina."[13] The genus is well known to European collectors under the name
Limenitis, of which we in England have one species, L. sibylla, the "White Admiral." A
species very like sibylla in general appearance is common in the northern parts of the
United States, ranging through Canada and Northern New England, but rarely south of
Boston. This species has the conspicuous white bands across both wings like our sibylla.
There is also a more Southern type known as astyanax, which is very different in its
appearance, being without the white bands and having a broad irroration of blue scales on
the posterior border of the hind wings. The two are so distinct that one would not be
tempted to suspect any very close relation between them. In its distribution astyanax is
described by Field as replacing arthemis south of latitude 42°. About Boston it is much
more common than arthemis.
The two forms encroach but little on each other's territory, but where they do coexist, a
third form, known as proserpina, is found which is almost intermediate, with the white
bands much reduced. There is now no doubt that this proserpina is a heterozygous form,
resulting from a combination of the characters of arthemis and astyanax. Field succeeded
in rearing a brood of 16 from a proserpina mother caught wild which laid 31 eggs, and of
these, nine (five males, four females) resembled the mother, being proserpina, and seven
(four males, three females) were arthemis. There can be no question therefore that the
mother had been fertilised by a male arthemis and that no-white-band is a factor partially
dominant over the white band. Another point of interest which Field observed was that the
proserpina female refused to lay on birch, poplar or willow, but accepted wild cherry
(Prunus serotina) a species on which astyanax can live, though that tree is not known to
be eaten by arthemis. Incidentally also the observations show that sterility cannot be
supposed to be the bar which maintains the distinctness of arthemis and astyanax.
In this connection Papilio oregonia and bairdii should be mentioned.[14] P. oregonia is
one of the numerous forms like machaon, but rather paler. It is a northern insect,
inhabiting British Colombia east of the Cascade Range, and reaching to Colorado. P. bairdii
is a much darker butterfly, representing the asterias group of the genus Papilio. Like
asterias it has the abdomen spotted at the sides, not banded as in the machaon group. It
belongs to Arizona and Utah extending into Colorado. From Colorado the form brucei is
described, more or less intermediate, like bairdii but with the abdomen banded as in
oregonia. W. H. Edwards records the results of rearing the offspring of the bairdii-like and
of the oregonia-like mothers. Each was found able to have offspring of both kinds, that is
to say, bairdii females gave both forms, and oregonia females gave both forms. It is not
possible to say which is dominant, since the fathers were unknown. On general grounds
one may expect that the bairdii form will be found to dominate, but this is quite doubtful.
From this particular discussion I omit reference to those examples in which the
permanently established types are obviously associated with special conditions of life.
Where considerable climatic differences exist between localities, or when we pass from
South to North, or from the plains into Alpine levels we often find that in correspondence
with the change of climate there is a change in the characteristics of a species common to
both. When I say "species" in such a connection I am obviously using the term in the
inclusive sense. Some would prefer to say that in the two sets of conditions two
representative species exist. Whichever expression be preferred it is plain that such
examples present another phase of the problem we have been just considering, and in
them also we have an opportunity of observing the consequences of the overlap of two
closely related types, but there are advantages in considering them separately. In the
examples hitherto given, with the possible exception of the Papilios,[15] the two fixed types
severally range over so extensive a region that it may fairly be supposed that in the
different parts they are subject to considerable diversities of climate. There is no
outstanding difference that we know distinguishing the habitats of the two forms; but in
comparing Alpine with Lowland forms, or essentially northern with essentially southern
forms we do know an external circumstance, temperature, that may reasonably be
supposed to have an influence, direct or indirect, on the population.
 CHAPTER VIII
LOCALLY DIFFERENTIATED FORMS. Continued.

Climatic Varieties
In this chapter we will examine certain cases which illustrate phenomena comparable
with those just considered, though as I have already indicated, they form to some extent a
special group. The outstanding fact that emerges prominently from the study of the local
forms is that when two definite types, nearly allied, and capable of interbreeding with
production of fertile offspring, meet together in the region where their distributions
overlap, though intergrades are habitually found, there is no normally or uniformly
intermediate population occupying the area of intergradation. Such phenomena as these
must, I think, be admitted to have great weight in any attempt to construct a theory of
evolution. True we must hesitate in asserting their positive significance, but I see no
escape from the conclusion that they throw grave doubt on conventional views. Again and
again the same question presents itself. If A and B lately emerged from a common form
why is that common form so utterly lost that it does not even maintain itself in the region
of overlapping? Almost equally difficult is it, in the cases which I have numerated, to apply
concrete suggestions based on any factorial scheme. We may see that in Heliconius erato
the type with the red mark on the hind wing probably contains a dominant factor, and that
where the red mark is absent the metallic colours are exposed; and that similarly the green
metallic colour may have another factor which distinguishes it from the blue. In this way
we can fairly easily represent the various types of erato on a factorial system as the result
of the various possible combinations of two pairs of factors. But there we stop, and we are
quite unable to suggest any reason why one area should have the red and the green type
while another should have the blue also. So again with Colaptes or the Warblers. By
application of a factorial system, admittedly in a somewhat lax fashion, the genetic
interrelations of the types can be represented; but how it comes about that each type
maintains a high degree of integrity in its own region we can only imagine. Each has in
actual fact a stability which the intermediate forms have not, but we cannot yet analyse
the nature of that stability. Mendelian conceptions show us how by segregation the
integrity of the factors can be in some degree maintained, but not why certain
combinations of factors should be exceptionally stable. All that is left us to fall back on is
the old unsatisfying suggestions that some combinations may have greater viability than
others, that there may be a tendency for like to mate with like, and so forth.
These difficulties acquire more than ordinary force in those cases in which the two fixed
types inhabit regions differing in some respect so obvious and definite that we are
compelled to regard each type as climatic and as specially adapted to the conditions. When
for example an animal has a distinct type never met with except in Arctic or Alpine
conditions, and another type proper to the plains and temperate regions, what are the
characteristics of the population of intermediate latitudes or at intermediate levels? Some
of the examples discussed in the last chapter may be instances of this very nature, but
even if they are not, others are forthcoming which certainly are. The evidence of these
cases leads to the suspicion that with further knowledge they will be found to consist of
two classes, some in which the observer as he passes from the one climate to the other
will find the intermediate area actually occupied by a population of intermediate character,
and others in which, though we may presume the maintenance of intermediate conditions
in the transitional area, there is no definite transitional population. This interrupted or
discontinuous distribution seems, so far as I have means of judging, to be by far the more
common of the two. I do not doubt that by sufficient search individuals representing every
or almost every transitional form can be found, but it is apparently rare that populations
corresponding to these several grades can be seen. The question has in few if any cases
been studied with precision sufficient to provide a positive answer; but I suspect that real
and complete continuity, in the sense thus defined, will only be found where the character
of the local populations depends directly on the conditions of life, and shows an immediate
response to changes in them apart from that postponed response which we suppose to be
achieved by selection. Obviously the character must be one, like size for instance, capable
of sensibly complete gradation.
The only example I have met with of the phenomenon of anything like a complete
intergradation between local types really distinct in kind is that provided by the butterfly
Pararge egeria. It is well known to entomologists that this insect exists in two very
different types, a northern one, the "Speckled Wood" of England, in which the spots are a
pale whitish yellow, and a southern type having the full fulvous colour that we know as
characteristic of megaera, the "Gatekeeper." It appears that Linnaeus gave the name
egeria to the southern type,[1] and our own is now called egerides. Broadly speaking, so
far as Great Britain, France, and the Spanish Peninsula are concerned, the tawny-coloured
egeria occupies Spain and western France up to the latitude of Poitiers and the pale yellow
egerides extends from Scotland, where it has a scanty distribution, through southern
England, where in suitable localities it is common, and the north of France to Paris.[2] The
two types when placed side by side are strikingly different from each other, and are an
excellent illustration of what is meant by climatic variation. The insect is not a great
traveller and probably scarcely ever wanders far from its home. It should therefore be
possible by collecting from north to south to find out how the transition is effected,
whether suddenly or gradually. This at various times I have endeavoured to do, but I am
still without exact information as to the population in certain critical areas. In addition to
the information derived from specimens which I have collected or seen in the collections of
others there is a good account of the general distribution in Europe given by the Speyers,
[3] who evidently paid more attention to the subject than most lepidopterists have done,

and many more recent records. In particular Oberthür[4] has published many details as to
the distribution in western France and I am especially indebted to Mr. H. Rowland-Brown
for a long series of notes as to the distribution in France generally, and to Mr. H. E. Page
and Dr. T. A. Chapman, Mr. Oberthür Prof. Arrigoni degli Oddi, Mr. H. Williams and other
correspondents, for showing me forms from many localities. The butterfly is attached for
the most part to woods of deciduous trees and to country abounding in tall hedges or
rough scrub. It is not usually to be found in highly cultivated districts or in very dry
regions. Hence there is necessarily some want of continuity in the distribution at the
present time and I should think a mile or two of arable land without big hedges would
constitute a barrier hardly ever passed. The larva feeds on several coarse grasses,
especially Dactylis glomerata. Barrett mentions also Triticum repens. In this country the
winter is usually passed in the larval stage, but I have found that in captivity, at least,
there is much irregularity. The larvæ feed whenever the weather is not very cold and may
pupate, but if sharp cold comes on when they are pupating or nearly full-grown they often
get killed unless protected.
Some writers speak of a difference between the early and later broods, but I have
never noticed this, and I do not think that the general tone of the yellow is affected by the
seasons (see Tutt, Ent. Rec., IX, 1897, p. 37).[5]
Beginning at the south of Spain the thoroughly fulvous type egeria is common at
Gibraltar in the Cork woods, at Granada, and doubtless generally. Lederer is said to have
found only this type in Spain (Speyer), and though I have no precise information as to
other places in the Peninsula north of Jaen I feel tolerably sure that there is no change
from south to north.[6] Immediately north of the Pyrenees we still meet egeria exclusively,
and up to Poitiers at least there is no noticeable change. But somewhere between Poitiers
and the bottom of the Loire valley at Tours, the genuine southern type comes to an end,
and the whole population begins at the Loire to be of an intermediate type, easy to
distinguish both from egeria and from egerides. As to the exact condition of the species in
the fifty miles separating St. Savin on the Vienne from places on the Loire I have no
adequate information. I have only one small sample from there, but it does contain insects
both of the southern and intermediate types taken on the same day, in a wood near
Preuilly. Oberthür also states that at Nantes the true southern form exists in company with
the northern. From this I infer that the southern form extends up the coast further than it
does inland, but I imagine the representative spoken of as northern would be of usual
Brittany or intermediate type.
The Vienne river joins the Loire, so the true southern type reaches over into the basin
of the Loire. From the Loire (Tours, Corméry) north to Calvados (Balleroy) only the
intermediate is found, so far as I know, and the same type extends over Brittany.[7] In
general, however, the woods near Paris have the thoroughly northern type egerides, but at
St. Germain-en-Laye and at Etampes (Oberthür) the population approaches the
intermediate type.
On the whole the intermediate type is certainly less homogeneous than either of the
extremes, and females with the two central spots either paler or more fulvous than the rest
are not uncommon, but I have never taken one on the Loire or in Brittany which I should
class with either of the extreme types.
Before speaking of the distribution in other parts of France and in Europe generally I
will briefly state the results of my breeding experiments. The work was done many years
ago before we had the Mendelian clue, and it is greatly to be hoped that some one will find
opportunities of repeating it. Crossing the English and the thoroughly southern type the
families produced agree entirely with the intermediates of Brittany and the Loire.
Reciprocals are alike. Of F2 I only succeeded in raising very few and of those that I had
(about 30) nearly all were intermediate in character, though perhaps rather less uniform
than F1. One family alone, containing only 4 specimens, had one egerides, and three
fulvous intermediates. As the case stands alone I hesitate whether or not to suppose it due
to some mistake. Moreover from F1 crossed back with the respective parental types I had
fairly long series, especially from F1 × the southern type, and looking at these families I
cannot see any clear evidence of segregation. On the contrary, I think that though there
are slight irregularities, they would, taken as a whole, be classed as coming between the
intermediate type and the extreme form used as the second parent. This at least is true
when the second parent was of the southern type.
On this evidence I have regarded the case as one in which there is no good evidence of
segregation and as conforming most nearly with the conventional view of gradual
transition in response to climatic influences. Such influence must however be indirect; for I
reared five generations of the northern type in England, and these, though they included
several abnormal-looking specimens in the last generation and then died out, did not show
any noticeable change from the fulvous colour of the wild type. Merrifield[8] also found that
heat applied to pupae of the northern type produced no approach to the southern type.
Looking at the facts now in the light of more experience it seems to me just possible
that the case may be one in which, as in Nilson-Ehle's Wheats, the dominant differs from
the recessive in having two pairs of factors with similar effects. The fulvous type for
example may have two or more elements in separate pairs which together produce the full
effect, and the intermediate may have one of these. If this were so, some segregation
should of course eventually be observable, but the proportion of the various fulvous and
fulvous-intermediate individuals would be large, and the reappearance of actual
representatives of the northern type might be rare. I admit that this is a somewhat
strained interpretation of the facts, and as yet it is not entitled to serious consideration.
Nevertheless I am led to form some such expectation partly from the great difficulty in the
way of any other, partly from the evidence of the small mixed sample found at Preuilly and
partly from the statements given by Oberthür. There are moreover other features in the
general distribution of the species which make it improbable that the dependence on
climate can after all be so close. Published lists are unfortunately of little use in deciding
which form occurs at a particular place, because, since the name Meone has ceased to be
used for the southern form, there is no complete unanimity among authors as to the
application of the names egeria and egerides, and unless more particulars are given, either
name may be used for either form. Besides this, difficulty arises from the fact that the
intermediate type is not generally distinguished at all, and English collectors finding it, may
easily record it as the southern type. From Staudinger's note on the distribution, I gather
that he, on the contrary, reckoned the intermediate with the northern type, as do the
Speyers also. The late Mr. J. W. Tutt was careful to distinguish the three forms and has left
several useful records. Easy therefore as it might seem to be to make out the distribution
of such a familiar insect in its various modifications, there are serious practical difficulties,
and until long series are brought together with this special object in view many obscurities
will remain.
With only the series from England, the west of France, and Spain before one it would
be easy to regard the successive series of tones as a fair measure of climate; the brighter
the colour, the hotter might one expect the locality to be. Such rough correspondence is
often to be observed in butterflies and birds. It becomes impossible to take these simple
views in the light of more complete knowledge. Beginning with France the fulvous egeria
occupies the lower valley of the Rhone, probably from well above Lyon, though I have no
exact information respecting the country above Avignon. According to Speyer it also takes
the department of Lozère. The same authority says that Puy-de-Dôme has "egeria,"
meaning perhaps the intermediate form, with the fulvous form much less commonly. Next
comes the curious fact that though the Lower Rhone (Avignon, Tarascon, Nîmes) has the
true fulvous form, Hyères, Cannes, Grasse, Nice, Digne, and Alassio have the intermediate.
Savoy has the intermediate (Chambéry) and even egerides perhaps, though in the same
latitude on the west of France there is nothing but the fulvous type. At Chalseul and
Besançon (Doubs) the ordinary northern type is found. Switzerland generally, I believe, has
the northern type, but Staudinger gives egeria for Valais and the intermediate occurs in
Vaud.[9] The south side of the Alps has probably colonies of the pale egerides, and of
intermediates. Orta, with a very hot summer, has the English type (Tutt, Ent. Rec., XII,
1900, p. 328). Locarno has the intermediate (ibid., XV, 1903, p. 321). North Italy in general
and western Piedmont have the intermediate; but further south egeria begins, at what
region I do not know. Speyer gives on his own authority the remarkable statement that at
Florence both extremes occur, but chiefly intermediates between the two. Mr. R. Verity
however kindly informs me that in his experience this is not so, and that neither the real
southern type nor the northern occur there. Sardinia, Sicily, Crete all have the southern
type. Greece probably has various types. Staudinger (Hor. Ross., VII, 1870, p. 78) says
intermediates resembling Nice types common everywhere, but from "Greece" the British
Museum has a series that would pass for English specimens; and the same type occurs
near Constantinople. The island of Corfu has a pale intermediate, distinct from egerides
but approaching it. In Roumania all three forms are recorded from various places: egeria in
the Dobrutscha; not quite typical (presumably an intermediate) at Bukharest; intermediate
in various mountainous localities as well as in Macedonia and Dalmatia; but egerides in
Azuga at about 3,000 feet.[10] Hungary has the true egerides also. (Cf. Caradja, Deut. Ent.
Zt., IX, p. 58.) Mathew records the same from Gallipoli (E. M. M., 1881, p. 95). Staudinger
does not distinguish the intermediates from the northern, but he gives "egerides" for
Armenia and Fergana (Central Asia). As against the mere proximity of a great mountain
chain being the influence which keeps the Riviera population intermediate may be
mentioned the fact that the northern foothills of the Pyrenees have the pure southern type,
and the climate of Cambo must surely be far cooler than that of Nice. The exact locality of
the Greek specimens is not given, but there can be no part of Greece which is not much
hotter in summer than Brittany, or Calvados, which have the intermediate, not the English
type.
In face of these facts it can scarcely be maintained that average temperature is the
efficient cause of the particular tone of colour which the butterfly shows in a given region.
Nevertheless it is clear that climate counts for much in determining the distribution. It is
noticeable that though the pale egerides can be established in a warm climate we never
find egeria in cold climates, and even the intermediate is not found in places that have a
hard winter. I suspect that the distribution of the broods through the year and the
condition of the animal at the onset of hard frost are features which really determine
whether a strain can live in a particular place or not. Though the truth of the suggestion
cannot be tested by experiments in captivity, which at once introduce disturbances, I
incline to the idea that egeria has not got the right periodicity for northern climates. If it
could arrange its life so that the population consisted either of young larvae, or perhaps of
thoroughly formed pupae[11] at the onset of winter, it might, for any obvious reason to the
contrary, be able to live in England. It is irregularly "polyvoltine," as the silk-worm breeders
say, and as soon as a little warmth encourages it, a new generation starts into being,
which if the frost comes at an untimely moment, is immediately destroyed. Many species
are continually throwing off individuals which feed up fast[12] and emerge at once if the
temperature permits, and I imagine a species of Satyrid wholly or largely represented by
such individuals could scarcely survive in a country which had a hard winter. For such a
climate some definite periodicity in the appearance of the broods may well be
indispensable. But assuming that egeria is cut off from cold climates for such a reason,
there is nothing yet to connect these habits with the fulvous colour, and until breeding can
be carried out on a satisfactory scale there is no more to be said.
From time to time records appear of individual specimens more or less fulvous being
caught in southern England, especially in the New Forest.[13] It would be interesting to
know what offspring such individuals might produce. From the evidence now given some
notion both of the strength and the weakness of the case considered as one of continuous
climatic variation can be formed. I know no other equally satisfactory. Whether or not
definite mixture of the intermediates with either of the extremes will be proved to occur,
the case differs materially from those considered in the last chapter in the fact that at all
events there is no general overlapping of forms. In a species so little given to wandering,
overlapping could indeed scarcely be expected to occur. It is this circumstance which
makes the species preeminently suitable as a subject for the study of climatic influences,
and I trust that entomologists with the right opportunities may be disposed to explore the
facts further.
Just as many species, like egeria, have varieties which can be regarded as adapted to
northern and southern regions, so there are also several which have lowland and Alpine
forms quite distinct from each other. Every such case presents an example of the problem
we have been considering. As the collector passes from the plains to the Alpine region,
how will he find the transition from one form to the other effected? Does the lowland form
give place to the Alpine form suddenly, with a region in which the two are mixed, or will he
find a zone inhabited by an intermediate population? I have spent a good deal of time
examining the facts in the case of Pieris napi and its Alpine female variety bryoniae, and
though there are many complications which still have to be cleared up, no doubt is possible
as to the main lines of the answer. If in any valley in the Alps inhabited by both napi and
bryoniae the collector catches every specimen he can, beginning at the bottom and
working up to 7,000 feet, he will at first get nothing but napi. At about 2,500 feet, he may
catch an occasional bryoniae flying with the napi. After 3,000 feet napi usually ceases, and
only bryoniae are found. As an exception a colony of napi may be met with at much
greater heights. I once found them in numbers at about 6,000 feet.[14] Not only were they
free from any trace of modification in the direction of bryoniae, but they were of the
thoroughly southern type of napi, being a late brood of that large and very pale kind
(meridionalis) almost destitute both of dark veining above and of green veining below,
which are common on the shores of Lago Maggiore and in other hot southern localities.
Not far off at the same level were typical bryoniae in fair abundance. Occasionally an
intermediate may be met with. I have taken a few, for example, at Macugnaga and at
Fobello. These, however, in my experience are rarities in the Alps. Fleck[15] gives notes on
the distribution in Roumania which shows the same state of things. The lowland form is
not transformed though found at great heights, and at Azuga (nearly 3,000 feet) bryoniae
occurs with only occasional "flavescens," viz., intermediates of the second brood.
If this were all the evidence we should be satisfied that the lowland and Alpine types
keep practically distinct, overlapping occasionally, but rarely interbreeding. The problem
would remain, how is the distinctness of the two types maintained in the region of
overlapping? Nowadays, I suppose, we should incline to answer this question by reference
to segregation, and perhaps by an appeal to selective mating. The suggestion that
segregation does take place is certainly true to some extent. There are, however,
difficulties in the way, and the whole subject is one of great complexity. My own
experiments were made in pre-Mendelian times and were not arranged with the simplicity
which we now know to be essential. The results are neither extensive enough nor clear
enough to settle the many collateral questions which have to be considered, and the work
ought to be done again. Nevertheless, some notes of the observations may have a
suggestive value.
When I began, I did not sufficiently appreciate that the "napi" group, omitting the
North American forms, and the Asiatic representatives, has at least three chief types in
western Europe. The differences we have to deal with are manifested by the females only,
so in this account particulars as to the males are omitted for the most part. These are (1)
our own British napi; (2) the form found in the south, from the Loire downwards, and in
the Italian Alps, which I think may be spoken of as meridionalis; (3) bryoniae, which is a
form clearly recognizable in the female only, and is found only in the arctic regions and in
the Alps above 2,500 feet. The first two have several broods, two, three, or more,
according to opportunity, and the first brood is different from the later ones. In napi the
markings on the upper surface are a dark grey but in meridionalis they are a pale silvery
grey and much less extensive. In the later broods of napi there is much less general
irroration of the veins, and the spots stand out as more defined and blacker. These
differences vary greatly in degree of emphasis. In meridionalis the later broods are entirely
different from the first. Instead of having silvery markings they have the ground colour
quite white, with the spots large and a full black. On the under side of the hind wings the
usual green veins are almost absent, and I have seen individuals which could scarcely be
distinguished from rapae. To these later broods the term napaeae is sometimes applied,
but I here use meridionalis for the southern race in general as applicable to all broods.
The female bryoniae is totally unlike the others. The ground colour is a full yellow, and
each nervure is thickly irrorated with a brown pigment often spreading so far as to hide the
ground almost entirely in the fore-wings. The males corresponding with these females are
not certainly distinguishable from those of our own napi. Both sexes have the green
veining of the underside of the hind wing fully developed, rather more than is usual in the
lowland races, but this is not really diagnostic of the variety. The first serious difficulty
arises in regard to the second brood of bryoniae. It is stated that there is only one brood,
[16] but I feel fairly sure that a second brood is sometimes produced, and that the females
with a yellow ground and diminished irroration of the veins, not very uncommon in the
Italian Alps in July to August, are generally representatives of it. Such insects would of
course be classed with bryoniae in collections.
My experiments began with eggs of true bryoniae females caught at about 2,500 feet
early in July. These emerged in August-September as intermediates with yellow ground
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