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I1172 The "Collar Bone," in theMaminalia. [December,

THE SIGNIFICANCE OF THE "COLLAR BONE" IN

THE MAMMALIA.
BY SPENCER TROTTER, M.D.

WHEN running through a series of forms in animal life a

structureis found fullydeveloped in some and in others
rudimentaryor entirelywanting,we are led to consider the
causes acting upon the structurethrough the lifeof the animal
which has brought about the conditionof developmentor non-
developmentpresent. Every fullydeveloped tissue in an organ-
ism is needed or it would not be there; and just so soon as by
increasing change in life and habits,it becomes a factorof less
and less importanceto the animal; it fails more and more to
attainits formerstandardof development,and in timefallsback
to the primitivecondition fromwhich it arose and finallydis-
appears.
The "collar bone," or clavicle is an unstable factorin the
shoulder girdle; by this I mean an element not always found
presentthroughoutvertebrates,and its presence,absence or rudi-
mentarycondition,is in relationto the lifeof the animal.
In this articleit is my intentionto note,principallyamong the
mammals,the relation which the clavicle bears to the various
modes of life,but beforetaking up the considerationof individ-
ual forms,reviewingbrieflythe general anatomyof the part.
The shouldergirdle consistsfirstof the scapula, or " shoulder
blade," a more or less irregularplate of bone preformedin car-
tilage and ossifyingfromtwo centers,the dorsal or scapular,the
ventralor coracoid,in positionagainst the anteriorthoracicribs,
its long axis varyingin inclination.
Its ventralend terminatesin the " glenoid cavity," a ridge and
process,spinousand acromion, on its outer surface are more or
less developed in differentforms,and in all mammals above
the Ornithodelphiathe coracoid is reduced to a mere process.
Second, the clavicle,when present,preformedin fibroustissue
extends as a bar of bone from the acromion process above the
glenoid cavityto the manubriumi sterni,forminga strongsupport
to the girdleand an extended surface for ligamentousand mus-
cular attachments. Mechanicallyconsidered the shouldergirdle
(by using the term" girdle" both sides of the body are implied)
is nothing more than the fulcrumof which the fore limb is the
lever of the thirdkind,and its specialization is in directrelation
1885.] The "Collar Bone" in theMcuninalia. 1173

to the amount and character of the work done. The clavicle

firstappears in the ganoid fishes as a secondaryapparatusdevel-
oped in connectionwiththe primarycartilaginousscapular arch.
In the teleost or bonyfishesit becomes a more importantelement
in the shouldergirdle,having connectionwiththe skull and the
opposite sides joining in the ventral medianline.
It undergoesa reduction and becomes of less significancein
Amphibia and Reptilia, but in birds it assumes an important
positionin relation to flight,the entire shoulder girdle in fact
being specialized to meet the conditions incidentto aerial loco-
motion. The coracoids-large, strongbones-act as braces; the
clavicles,peculiarlymodified,are unitedat theirsternalends into
one bone, the furculum,or " merrythought1"which,as Owen has
pointedout, acts as an elastic,bony arch opposing the forceinci-
dent upon the downwardstrokeof the wing in flight, thus aiding
the humeri,or arm bones, to regain their formerpositionin the
succeeding upwardor counter stroke with as little loss of time
and energyas possible. In terrestrial forms(grouse, fowls,etc.),
where flightis sustainedforshortintervalsonly,the arch is nar-
rowerand the structuremore slenderand delicate.
Taking up the Mammalia witha view to ascertainingthe rela-
tionshipexisting between the developmentof the clavicles and
the work done by the forelimbs,let us start with the following
general propositionas a basis for our observations,namely,that
those animals which have the forelimbsspecialized over the hind
limbs in relationto work,possess a clavicle,and where the hind
limbsare the most highlyspecialized,the clavicles are rudiment-
ary or entirelywanting.
Leaving the ornithodelphmammals (the Australian duck-bill,
Echidna, etc.),with their bird-like shoulder girdle,we note the
presenceof more or less well developed clavicles in all the mar-
supials,.with the single exception of the " bandicoots" (Pera-
rnelidac),where it is wanting,and its absence may be accounted
forfromthe factthatthe bandicootsfromtheir terrestrialmode
of life, nesting in hollow places and feeding on insects, roots
etc.,have fora long time had their fore limbs subjectedto less
complex conditionsthanthe arboreal and predatoryfamilies,the
.opossums(Didelphidme), the Dasyuridw,the phalangers(Phalang-
istidx) and the kangaroos (Macropodidaz),which constitutethe
restof the order. In the kangaroos the clavicles,though pres-i
1174 The "Collar Bone" in theMammalia. [December,

ent,are slenderand delicate,the weak forelimbs cominginto use

in manipulatingthe "pouch," etc.,thus throwinga variety of
motion into the shoulder joint, which explains the presenceof
clavicles in an animal whose locomotion is almost entirelyper-
formedby the hind limbs.
Among the Insectivora,a large orderof wide geographical
range and ratheruniformlife,the majorityof species being ter-
restrial and fossorial or burrowing,we have the clavicles well
developed withbut one exception,thatof Potamogalevelox,a rare
formfromWesternAfrica,and this is the only insectivorewhich
is almost entirelyaquatic. It measures about two feetin length
witha long cylindricalbody, tapering continuouslyinto a thin,
laterally compressed tail,which is the main propulsive power
when swimming,the shortlegs withtheir unwebbedfeetdrifting
back against the body. Anotherform,Myogale,fromthestreams
of SoutheasternRussia, is natatorialand possesses a clavicle,but
the feetare all webbed and come into play along withthe tail as
organs of propulsion.
The moles, Talpidae,are eminentlyfossorial,their excavations
being everywhereknown; an East Indian squirrel-likeformis
arboreal; all the restare terrestrial and fossorial.
The order Edentata is divided into two primary groups,the
"leaf-eaters" (Phytophaga) and the " insect-eaters " (Entomoph-
aga). The Phytophagacomprisethe "sloths," curious,arboreal
formsinhabitingthe South Americanforests,representedby only
two livinggenera; they are entirelyarboreal,making character-
isticprogressionamong the tree-topson the leaves of which they
feed. In the two-toed sloth,or "unau" (Cholcepus), the clavi-
cles are well developed; in the " ai," or three-toedvariety(Brady-
pus), theyare small,rudimentary, having lost theirsternalattach-
ments and evidently undergoing a retrogressivechange, due
probablyto some variationin the animal's life and habits which
bringsthe part into less active use.
In the Entomophaga (ant-eaters,armadillos,etc.) we findthe
clavicles fullydeveloped in the climbing two-toed ant-eaterCy-
clothurus,a small South Americanspecies; also in the Cape ant-
eater,or "Caard-vark,"Orycteropus,a burrowing form fromthe
Cape of Good Hope; while in the great ant-eater,Myrme-
cophaga, a strictlyterrestrialformbut not fossorial,the clavicles
are absent,the long claws tearingopen the ant hills and termites'
I885.] The "Collar Bone" in tkeMcnmalui. I 175

mounds,and the narrow,flexibletongue securingthe food,stand

in place of the burrowinghabit,thus bringingthe shoulderjoint
under much less active conditions. In the South American
armadillos,which are more omnivorousand are " diggers,"the
clavicles are developed. The " pangolins," scaly covered ani-
mals, mainlyterrestrialand rolling themselves in a ball on the
approach of danger,have no clavicles.
In the orderRodentia we findall the Sciuromorphgroup with
fullydeveloped clavicles; the squirrelsare eitherarborealor bur-
rowers; the marmots,Arctomyina,are typical burrowers,while
the beaver,though eminentlyaquatic, brings the forelimbs into
active use in the constructionof its dams and lodges.
The Myomorpha (rats, mice, gophers, etc., typical gnawers
and diggers) have well developed clavicles,the gnawing habit
bringingthe forelimbs and shoulder girdle activelyinto use to
steady the head and neck and the object which is being worked
upon.
In the Hystricomorpha(porcupines,cavies, etc.) clavicles are
founddevelopedin the climbingtree-porcupines (Sphingurinac),of
which our Canadian species (Erethizon) is a member,and absent
in the large Africanporcupine and all othersof the Hystricine
group,which is strictlyterrestrialbut not fossorial.
The chinchillas have well developed clavicles,the horse-trip-
ping burrowsof the " Pampas Viscacha " being too well known
to the ridersof thatregion. The agoutis (Dasyproctidae),inhab-
itingthe banks of streams in the South Americanforests,semi-
aquatic and sub-ungulate(i. e., having the feet partiallyencased
in a horny sheath or hoof) have rudimentaryclavicles. The
Caviidoe (guinea pigs) and the " capybara," the largest living
rodent,semi-aquatic,withwebbed feet,have imperfect clavicles.
The Lagomnorphaconsistsof two livingfamilies,th'eLagomyi-
do, small burrowinganimals called "picas," inhabitingAlpine
regions,and the Leporidze(hares and rabbits); the formerfamily
having well developed, the latter rudimentaryclavicles. The
hares are typical runners,solitary,never burrowing,depending
forsafetyin speed and making their characteristic"form"; the
species known as the rabbit,however, differslargely in habits
from the other membersof the family,being more or less gre-
garious,livingin "warrens" which it burrows out, and having
much less capacityforspeed. Clavicles are developed in both;
VOL. XIX..TN. :KTo 77
1176 TSe "Collar Bone" in theMamnalia. [December,

veryrudimentary in the hares, and in the rabbits existing as a

bone incompleteat the ends, a conditionwhich we would expect
to findfromthe variationin lifehabits.
Taking up the large orderof " hoofed" animals,the Ungulata,
we note the completeabsence of the clavicles in all the forms,
not a vestige of the bony structurebeing found in any one of
them. They are the " runners" par excellence,and if we may
use the expression,live on theirlegs,the capacityforspeed and
endurance being one of the chief factorsaimed at by natural
selectionin maintainingand perfectingthe species.
Here we findthe forelimbs subservingthe hind limbs in rela-
tion to work; the latterare the main motor power in running,
while the forelimbs act chieflyas supportersand guiders; hence
the entireabsence of clavicles. The marineMammalia,withtheir
rudimentarylimbs,have no clavicles.
In the flesh-eatinganimals,Carnivora,the clavicles are always
rudimentary, and in many cases entirelywanting. They are bet-
ter developed in the cats, Felidaz,than in any otherfamilyof the
order(though the articularextremitiesof the bone are lost), for
the clutch whichfollowsthe springin securingtheirprey,brings
the forelimbs into more active use than in the dogs and wolves,
Canidae,which run theirquarryto " the death,"and then pull it
down by force of numbers. Many 'of the-cats, too, frequent
trees,but theyspring fromplace to place and do not properly
climb. Several species of bears also climb, but the habit is
more acquired than natural,the forelimbs clasp the trunkwhile
the hind ones secure a footholdand, so to speak, shove the ani-
mal upward,verydifferent fromthe active movementsof a nat-
ural climber.1
In the bats, Chiroptera,as we would expect fromtheirmode
of life,the anteriorlimbs being the main factorsin their flight,
clavicles are developed,long, curved and strong.
The arboreal Lemurs also have them developed.
In the order Primateswe have the clavicles presentin all, from
the arboreal monkeysup through the anthropoid apes to man,
where the anteriorlimbs attain such perfection,with ball and
1 The badger (Taxidea) is a strikingexceptionto the general rule,being an emi-
nentlyfossorialanimalyetdevoid of clavicles. The fore feet are armedwithex-
ceedinglylong, slightlycurved claws, probablybringinggreaterleverage on the
wristand elbowjoints and thuspartiallysupplementing the absence of clavicles.
I1885.] Pear Bligkhand its Cause. 1177

socket shoulderjoint capable of such variedand extensivemotion,

witha high degree of pronationand supinationof the forearm,
and last, but not least, the wonderfullyspecialized hand withits
thumbopposable to each of the fourdigits.
Afterthisbriefsurvey,and withthe forementioned proposition
in view,viz.,the correlationexistingbetweenthe developmentof
the clavicle and the work done by the fore limbs,we are left to
draw the rationalconclusionthatthe subject underconsideration
is one of uiseand disuseof parts,as Darwin has so clearlypointed
out in his chapter on rudimentary organs in the Originof Spe-
cies. The factswe have noted in our hasty glance at the Mam-
malia confirmthis,in the more or less perfectdevelopmentof
clavicles in arboreal,fossorial,aerial and all other formswhere
the forelimbs are the active,aggressive pair in the life of the
animal,and theirabsence or rudimentary conditionin the hoofed
animals,the marinespecies and all otherswherethe anteriorpair
take a secondaryplace in the work done by the limbs.
As there is, of course,no actual disuse of a part as a whole
(the nearestapproach to this being in marine forms),a simple,
uncomplicatedmotion existing,with little strainat the shoulder
joint, the parts requireless supportand fewerpointsforligament-
ous and muscular attachmentthan where the movementsare
more complicatedand the strain more severe. Consequentlywe
have a greateror less differentiation in the elementsof the shoul-
der girdleas the case may be, and the clavicle,holding as it does
a positionof secondaryimportance,is the unstable,variable ele-
ment.

PEAR BLIGHT AND ITS CAUSE.

BY J. C. ARTHUR.

DEAR trees in this countryare subject to an endemic disease

I that,owing to its malignancyand frequentoccurrence,is well
knownto cultivatorsand fairlywell discriminatedby them. It is
knownboth as pear blightandfire blzokzt, and the same disease'in
the apple and quince is also called twigbligit. The term blight
is applied to manykinds of plant diseases,and especiallyto those
thateventuallykill withoutrenderingthe cause conspicuous; it
is also the name of a class of disease-producingfungi. The pear
maladybearingthis ulameis, however,a specificdisease,although