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Psyche 8:315-322, 1897.

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Pscvhe, Vol. 8. Plate 8.
- ~~ - -
Isothermal Lines and Faunal Areas of New England.



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Isothenns by W. M. Davis; Faiinal Zones by S. H. ScudderÌÔ(fro Scudder's Butterflies of the Northeastern States).
- - Mean annual isotherms.
Vertical Ruling - Canadian fauna, restricted limits (= Boreal). Unrulcd - Canadian fauna, ordinary limits (= Transition). Oblique Ruling-iIllegl1anian fauna, ordinary limits (= Transition and Carolinian). Horizontal Ruling - Alleghanian restricted (= Carolinian). Oblique Broken Ruling-Dilute Carolinian locust fauna. Ml. Ktaadn, Me.
Jackman, Me.
Hudson, Me.
Aft. Desert Id., Me.
Fryeburg, Me.
Norway, Me.
York, Me.
White Mountains. N. H.
Jackson, N. 11.
North Conway, N. H.
Jaffrey, N. 11.
Newport, Vt.
Montgomery, Vt.
Woodstock, Vt.
Ascutney Mountain, Vt.
Boston, Mass.
Cambridge, Mass.
Faneiiil, Mass.
Newtonville, Mass.
Wellesley, Mass.
Lynn, Mass.
Winchendon, Mass.
Amherst, Mass.
Palmer, Mass.
Springfield, Mass.
Greylock Mtn Mass.
Dedham, Mass.
Walpole, Mass.
Provincetown, Mass.
Martha's Vineyard, Mass
Wickford, R, I.
Block Id., R I.
Thompson, Ct.
Deep River, Ct.
No. Madison, Ct.
No. Haven, Ct.
New Haven, Ct.
Canaan, Ct.
So. Kent, Ct.
Stamford, Ct.
Greenwich, Ct.
Ravenswood, L. 1.




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PSYCHE.
THE DISTRIBUTION OF THE NEW ENGLAND LOCUSTS. BY ALBERT P. MORSE, WELLESLEY, MASS.
[Annual address of the retiving president of the Cambridge Entomological Club, Jamra-ry 13, i8qq.l The Acridiidae, short-horned grass-
hoppers or true locusts, are a family of insects of especial value for the study of geographical distribution, 'I life zones," and kindred topics,--.probably more
valuable than any other group of insects, and perhaps as important as the birds :
they are, in consequence, deserving' of
more attention from biological surveys
than they have yet received.
The reasons -for ascribing to them
so great value in connection with this
subject are as follows :-They are
terrestrial (as distinguisliecl from aqua- tic) in all stages of their existence. They are almost universzilly distributed -
with the exception of dense forests-
from desert to jungle, from sea-side to
mountain-top, from the arctic zone to
the equator. They are conspicuous in
size and habits: being from one-half
nch to four inches in length, one inch
to nine inches in expanse, diurnal,
active and alert, flying or leaping freely when approached, often adorned with
striking colors or producing loud sounds either in flight or at rest. They are
voracious, their food being general in
character rather than special, a question of quantity rather than quality. They
are numerous in individuals, whether
local or widespread occurring in suffi-
cient numbers to make it possible with
reasonable effort to procure series snffi- ciently large to permit of critical study. They require a relatively small amount
of care and delicacy of handling, in&-
ing it possible to devote a proportion-
ally large amount of time during field-
work to the securing' of material rather than to its preparation. Finally, while
the majority are winged and active in
habit they are, with few exceptions,
relatively stationary during life, for not only is the range of individuals com-
paratively restricted but the same is true also of the range of a species in a given locality, which seems to be dependent
less upon the presence or absence of
particular food-plants than upon physi-
cal conditions often of extremely local
character, such as the quality of the
soil, exposure, drainage, humidity, etc. So true is this last statement that in a rcgion where the locust fauna is fairly
well-known, e. g., New England, given




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$16 PSYCHE,. [February 1899.
a field of any particular character it is possible to predict with a high degree
of certainty what species of locusts will be found therein.
This susceptibility to local physical
conditions is a very important attribute of the group in this connection. Most
cultivated crops and many wild plants
are equally susceptible to the same con- ditions, --a fact especially noticeable
in parts of California where exposure
to sea-wind or direct sun-rays produces
a marked difference in the amount and
character of the vegetation on contiguous hill-slopes, and also in parts of New
England, where fields a few acres in
extent are seldom homogeneous in chai-
acter.
Now the successful cultivation of
crops by the individual agriculturist is dependent on the factor here involved :
a close adaptation of a particular species or variety to its environment,-the en-
vironment being constituted by condi-
tions, as has been said, often extremely local in character. For this reason it
seems probable that close study of the
distribution of this group of animals
would prove of exceptional value in
delimiting the smaller details of life
zones, especially in the more valuable
narrow extensions of such areas, whose
importance has been indicated by Dr.
Merriam.
It is perhaps worth while to point
out in this connection that species be-
longing to a southern zone and adapted
to a sandy soil may find their northern
limit considerably higher up, both lati- tudinally and veitically, than other spe- cies equally characteristic of the same
zone but restricted to a damp and heavy
soil. More rarely the reverse arrange-
ment would occur. Consequently, that
the final limit of a given zone would
vary according to the species selected
to delimit it and the adaptation of these species to the physical conditions of the locality in question.
So much by way of preface and in
regard to details. It remains to be seen whether the distribution of the group is of value in determining the broader
features of life zones. For this purpose let us turn our attention to a district
sufficiently well-known to allow a corn- arson to be made. Such a district
we have in New England, whose locust-
fauna, with the exception of noithern
and eastern Maine and western Ver-
mont, is as well known as that of any
part of the Union of equal extent and
whose faunal areas also have been the
subject of study. While it is true that
a great deal remains to be done in
working out the details there is suffi-
cient available to enable a close ap-
proximation to the truth to be secured.
Geographically considered the locusts
of New England fall into three groups :
- [st, species believed to be distributed over practically the entire district ; 2nd, speciesknown to be distributed over but
a part of the district ; yd, species whose distribution - either from local char-
acter or positive rarity-is insufficiently known. Of these groups the first is of
interest in showing the relation of New
England to the rest of the country ; the third may be ignored for the present;




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February 1S99,1 PSYCHE. 31 7
to the second we must look for the divi- sion of the district into fannal areas. It we include Long Island as a part of
New England, whicli wc may reason-
ably do, we have actual record of 48
species from this district.
Of these 19
belong to the first group, 3 to the third, and 26 to the second.
The second group - those distributed
over but a part of New England - falls
naturally into two divisions, a boreal
d an anstrat.
The former, consisting
of several species of boreal character,
extends from tile north over varying
prtions of the district ; the latter, ans- tral in character, extends similarly from the south over vnrying poi liuns. These
two divisions overlap each other widely, a consequence clue in part to the genernl north and south trend of the more i~n-
portant physical features, -- mountain
chains, river valleys and sea-coast,-.-
with its consequent effect on tcnnpcr-
atures, but chiefly to other causes, par- ticularly that of specific adaptability, as will be shown.
The northern division contains six
species. The first of these, Podisma
glacialis, is of a truly boreal and sub- alpine character, and is found more or
less commonly in the vicinity of tree-
line on the higher mountains of New
England (Ktaadn, I, White Mts., 8,
Ascutney, 15, Greylock, 26, 3000 to
4500 ft.) anil at a lower level in north- ern Maine (Jackm~n, 2).
Closely related to it in distribution
and associated with it in several of the same stations is Melanopl-ns mancus,
which seems, however, to be able to
withstand a higher temperature. This
species hcis been found on Mt. Desert
Id., 4, Hie Wliite Mts., I, Ascutney
Mt., 15, and occurs, probably in an
outlying colony, as far south as southern Connecticut, 34. This southward ex-
tension of its range is probiibly due to the influence of the ice-age.
r ,
lhese two species are apterous or
sub-apterous and in consequence of their boreal character, sire markedly discon-
tinuous in distrib~if.ion, except, possibly, in the extreme north. Thc remaining
four are winged, in consequence are
able to range more widely and occur,
ill the adult stage at least, on the alpine si~i~m~its of thc White Mis. as well as at lower levels.
Two of them prefer moist situations
and apparently have n-iiicli the sjime
distribution. These are MecostethniS
grad& (Norway, Me., 6, Wliite Mts.,
8, and Jafirey, H, N. H., northern
Vermont, 12, 13, Greylock Mt., 26,
Mass.) ;I 116 Afelano/Z?u extrems
(Hudson, 3, and Norway, 6, Me.,
White Mts., 8, Jiickson, 9, and No.
Conway, 10, N. H., Asci~tne~ Mt., 15,
Woodstock, 14, and northern Vermont,
12, 13, Mt. Grcylock, 26, and Winch-
enclon, 32, Mass). The fi~st of these
is met by its congener, lineatus, which
takes its place in southern New Eng-
land.
The remaining two boreal species
extend even further to the so~ith, reach- ing northern Connecticut. These arc
Circotettix verni-cnlatus and Camnula
/eUzidda. The former frequents rocky
ground and is met (or overlapped) at




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PSYCHE.
[February 1899.
Canaan, Ct., 38, by S-pharagemon
saxdile, a species having precisely
similar habits which covers the south-
ern part of New England. C. veryucu-
latus occurs at Palmer, 24, Mass., and
is recorded from Cambridge, 17, but
seems not to be found there now, while
saxatile is plentiful. C. fellucida
reaches So. Kent, 39, in western and
Thompson, 33. in northeastern Con-
necticut on high hills, outposts of the
highlands of western and central Mas-
sachusetts.
So much for boreal species. Passing
to the austral we find a group of three, Tryxalis ifevicornis, Clinocephalus
elegans, Orfihula olivacea, occurring
in swamps and salt-marshes in the
vicinity of New York (Ravenswood,
L. I., 42, Stamford, 40, and Greenwich,
41, Ct.) but perhaps extending further
to the northeast. A fourth, Scktsto-
cerca americana, is found not infrc-
quently about New York, and has been
taken once near Boston. This is,
however, an insect of exceptionally
powerful flight am1 strong migratory
tendency and is entirely adventitious in New ~ngland.
The remaining species inhabit a
much larger portion of the district.
Sckistocerca alutacea is common in
swamps in southern Connecticut and
reaches Martha's Vineyard. 30. Mela-
noplus scudderi is found in western
(So. Kent, 39), southern (New &ven
37), and southeastern (Deep River,
31) Connecticut, and is reported from
*The reporie4 occurrence of this species in liruiiswick Springfield, Mass., 25.* Dichromor-
$ha viridis is common in all quarters
of Connecticut, 33, 36, 38, 40, and at
Amherst, 23, Mass., and is reported
from New Hampshire which it probably
reaches in the Connecticut valley. It
is found at Wickford, 31, R. I., and
probably throughout most of the State
but is absent from the vicinity of
Wellesley, 20, and Boston, 16. Par-
oxya floridana, common in southern
Connecticut, reaches Boston, 18, and
Cambridge, I 7. Scirtetica marmorata
is found in southern Connecticut, 36,
on Martha's Vineyard, 30, and at
Provincetown, 29, Mass. Schistocerca
yuliiginosa is found in northeastern
Connecticut, 33, at Provincetown, 29,
Dedham, 28, and Wellesley, 20, Mass.
Oy$hula macul@ennis, also common
11 northeastern Connecticut, is found
along- the coast at least as far as Lynn, 21, Mass., hut for some reason is practi- cally absent from the vicinity of Wel-
lcsley, 20. Paratettix cucullatus
occurs in extreme northwestern, 38,
southern, '57, and northeastern, 33,
Connecticut and has been reported
from the vicinity of Boston. Arfihia
xanthoftera, Spharagemon saxatsle,
and Sph. collars scudderi cover much
or all of Connecticut, a large part of
central Massachusetts, are common
about Boston, and probably extend
considerably farther to the noitli and
east, probably reaching southwestern
Me. (Sadder, Rev. Melanopli, p. 214) is earoneous, the specimen proving to be another species-MeIawplits mi, not previoidy known to occur in New England,



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February iSqq.1 PSYCHE. 31 9
Maine. Pseudopomala brachyptera,
Mecostethus lineatus, Melanoi>Zus
minor^ and Oehula aequalis extend
even farther north, in most cases at
least to Woodstock, 14, Vt., Jatirey,
11, and No. Conway, 10, N. H.,
Fryeburg, 5. and Norway, 6, Me. The
sole remaining species is noteworthy as
illustrating in a high degree the peculiar susceptibility of many of the group to
physiographical conditions, and its dis- tribution may rightly be said to be
almost as much physiopaphical as
climatal. This is Trimerotroi>is mari-
tima, the sea-side locust, which is
restricted to the sandy sea-beaches of
the coast from York, 7, Me., southward.
The only exception known to me is
that of a small colony at North Haven,
'56, Ct., on the sandy plains of the old sea-floor, now a few miles inland but
only a few feet above tidewater.
Examining the positive evidence
given above in connection wit11 the
faunal and clirnatal map of New Eng-
land in Scudder's Butterflies of the
Northeastern States we find a remark-
able conformity in the distribution of
the locusts with the courses of the mean annual isotherms, a conformity most
striking in the western, more inland,
part of their courses. Compare, for
instance, the southward extension into
Massachusetts ofthe ranges of Mecoste-
thusffracilis and ~lfelanoplus extvemu s Or$hula aepalis to that of 4z0, of
DicJzromorpha viridis to the western
part of 46', of Melanoplus scudderi to
the western part of 48'.
The distribution of the locusts is dis-
tinctly closer in agreement with the
annual means than with the winter
means ; perhaps a little closer with them, on the whole, than with the faunal zones drawn from the distribution of the
butterflies. I greatly regret to have
been unable to compare it with the
summer means or with lines based
on the laws given by Dr. Merriam,
viz.- that northward distribution is
governed by the sum of positive tem-
peratures for the entire season of growth and reproduction, and the southward
distribution by the mean temperature of
a brief period during the hottest part
of the year.
Of course discrepancies occur. These
are probably due, in part, to insuffi-
cient dam, but may be due to other
agencies, such as, e. g., some unknown
factor, or the difference between the
annual mean temperatures and the fac-
tors stated above or to the possible con- flict of these two factors between them- selves, resulting in unconformable lim-
its of two adjoining zones, distinguished by the overlapping or absence of char-
acteristic species in certain sections.
As nearly as we can judge, however,
from the data at hand, these limits are
conformable in the case of the New
-
with the line of 44', of Circotettix
verrztculatus and Camnula pellucida
into Connecticut with the line of 46';
the northward range of Mecostethus A -
lineatus, fkfelano@us minor, and consisting in the absence from the England locusts except in eastern Mas-
sachusetts, where, also, the principal
discrepancy with the isotherms is found.



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320 PSYCHE. [February 1899.
neighborhood of Boston and Wellesley,
at least, of the austral species Melano- å´plu sczidderi and Dichromorpha
vi~idis, and the boreal species Circo-
tettix verruczitlattis and Camnula
å´pellucida '*å
Notwithstanding discrepancies, to
whatever agency due, it is clear that
the distribution of locusts in New Eng-
land is primarily and distinctly climatal in character, although strongly influ-
enced by physiography and its attend-
ant conditions.