ON THE 
Affinities of Aphriza Virgata. 
BY 
R. W. SHUFELDT, M.D., C.M.Z.S. 
[Reprinted from Journal of Morphology, Vol. II., No. 2, November, 1S88.] 
BOSTON: 
GINN AND COMPANY. 
1888. ON THE 
Affinities of Aphriza Virgata. 
BY 
R. W. SHUFELDT, M.D., C.M.Z.S. 
[Reprinted from Journal of Morphology, Vol. II., No. 2, November, 1888.] 
BOSTON: 
GINN AND COMPANY. 
1888. ON THE AFFINITIES OF APHRIZA VIRGATA. 
\_Based upon a comparative study of its skeleton.] 
R. W SHUFELDT, M.D., C.M.Z.S. 
During the latter part of November, 1885, Surgeon Thomas 
H. Streets of the United States Navy, then naturalist of the 
United States Exploring Steamer "Patterson," kindly sent me 
a skeleton of the Surf-bird, the subject of the present memoir; 
and again in August, 1886, the same distinguished officer for- 
warded me a fine pair of skeletons of this species, represent- 
ing both sexes, adult. All of this material was collected by 
Dr. Streets in Alaska, and I am indebted to him also for skele- 
tons of Charadrius squatarola and Arenaria melanocephala, the 
former taken in San Francisco Bay, the latter at Port Townsend, 
Washington Territory, and placed at my disposal for comparison 
with the skeletons of Aphriza. 
As additional material for the osteological comparisons the 
writer here proposes to make, I find I have at my command two 
skeletons of adult specimens of Htzmatopus bachmani (Nos. 
13,636 and 13,637), belonging to the Smithsonian Institution of 
Washington; and finally, in my own cabinet several skeletal 
preparations of the Charadriidce and Tringece, all of which will 
be of assistance in the work now in hand. 
Aphriza virgata constitutes but another one of those forms 
around which centres so much that is of interest to the sys- 
tematic ornithologist, owing to the fact that even after more 
than a mere superficial examination we discover not a little in 
its anatomy that is inclined to puzzle one, when called upon to 
pronounce as to its kinship with more or less nearly related 
groups or types. 
Of the first skin that he ever examined, Audubon wrote: 
" The remarkable bird here represented, which in form and 
size bears a considerable resemblance to the Knot \Tringa ca- 
mitus\, was procured by Mr. Townsend on the shore of Cape 312 
SHUFELDT. 
[Vol. II. 
Disappointment, and proved to be a female."1 And Dr. 
Coues remarks, in defining the genus, - " General character 
of plumage, in its pattern of coloration and seasonal changes, as 
in Tringetz. One species, a remarkable, isolated form, perhaps 
a plover, and connecting this family with the next by close 
relationships with Strepsilas, but with hind toe as well devel- 
oped as usual in Sandpipers, and general appearance rather 
sandpiper-like than plover-like. Aphrizintz might go under 
Hcematopodida next to Strepsilas; or, perhaps better, Aphriza 
and Strepsilas might together constitute a family Aphrizid^e, 
next to, but apart from, Htzmatopodidcz." 2 
Two years after the publication of the doubts here candidly 
expressed by this eminent authority, the American Ornithologi- 
cal Union issued its official Check-List, wherein the arrangement 
proposed was adopted; and we find the Aphrizidtz with its one 
species, A. virgata, including the sub-family Arenariintz, the 
Turnstones, standing between the Plovers on the one hand, and 
the Oyster-catchers on the other,3 an arrangement which proba- 
bly meets with the views of the majority of avian taxonomists of 
the present time. 
Believing that under these circumstances a full account of 
the skeleton of Aphriza will prove an acceptable contribution to 
the literature of the family, as well as a useful work to the better 
understanding of the osteology of the Limicoltz generally, it was 
in view of this that I undertook the task. So far as my mate- 
rial at hand will admit of it, the study will be made thoroughly 
comparative, and in the end it may go to show that, for osteolog- 
ical considerations at least, the position selected in the system 
by ornithologists may be the most natural one to create for our 
Surf-bird, or a somewhat different light may be cast upon the 
problem; in any event we are satisfied that more light in such 
matters is a thing most to be desired, and this alone morphology 
has the power of affording. 
The Skull. - When we come to submit the skulls of adult 
specimens of certain true Tringetz to moderately prolonged ma- 
ceration in water, we invariably find that the premaxillary bone 
detaches itself, as does the dentary of the mandible, leaving in 
1 Birds of America, Vol. V., pl. 228. 
2 Coues, E, Key to North American Birds, 2d. ed., p. 605. 1884. 
8 Check-List of North American Birds, pp. 160-166. New York, 1886. No. 2.] 
APHRIZA VIRGATA. 
313 
the first case the long, thread-like extremities of the palatines and 
median processes of the nasals freely projecting forwards, and 
in the latter instance, a similar condition of the mid-ramal 
elements of the lower jaw. Tringa, Aditis, and others have 
their skulls behave in this manner, under similar circumstances, 
and it is quite characteristic of them; whereas, on the other 
hand, macerate the skulls as long as we may, no such detach- 
ment ever takes place in the Surf-bird, nor in Turnstones and 
Oyster-catchers, nor in such forms as Gallinago and Philohela. 
Indeed, this peculiar feature seems to be restricted to the more 
typical Sandpipers, and it is worthy of our notice that in Aphriza 
it never happens, and I may add that one never observes it to 
be the case in any of the Plovers. 
Aside from this fact, the general facies of the form of the 
superior osseous mandible in Aphriza, carried back as far as the 
rhinal chamber and frontal region, is more as we find it in such 
a species as Aditis, for example, or some of the genus Tringa, 
than it has any semblance to either Arenaria, or much less 
Hamatopus. Even when we come to compare the individual 
parts, this assertion holds true, including in the observation the 
extensive outline of the triangular aperture, on either side, of 
the external nares ; and the superior ends of the fronto-maxil- 
lary processes of the nasals not overlapping, mesiad, the outer 
and juxtaposed margins of their premaxillary processes; which 
latter condition obtains both in the Turnstones and in Oyster- 
catchers. Again, in Aphriza we note that the extremity of the 
superior mandible in the skull is somewhat tumefied, and marked 
over with minute pits, as we find it to be in Gallinago. This 
character is absent in such a form as Aditis and other Sand- 
pipers, nor do we find it to be present in Charadrius, Arenaria, 
and Hcematopus. 
In Bachman's Oyster-catcher (H. bachmani) the superior 
osseous mandible is about seven centimetres long, the remainder 
of the cranium being something over 3.5 cm. long, measured 
along the median line. Taking these same distances in the 
Black Turnstone, we find the first to be 2.5 cm., and the latter 
2.3 cm.; but barring this difference in proportion between beak 
and cranium in these two species, the superior osseous mandible 
in the Turnstone is almost the perfect miniature of that part of 
the skull in an Oyster-catcher. We are to note, however, one 314 
SHUFELDT. 
[Vol. IL 
other difference, and that is, in Hcematopus the anterior extrem- 
ities of the palatines fuse across the median interspace much 
further back than they do in the Turnstone, but essentially, as 
I say, the plan of these parts in the two birds in question is the 
same, and quite different from the corresponding structure in 
Aphriza. Plovers in this particular seem to stand between the 
Surf-bird and Arenaria. 
Taking next these several skulls upon their superior aspects, 
and comparing them with the same view in A. virgata (Fig. i), 
we find that the latter bird again agrees with such a Sandpiper 
as Actitis macularia, or with such a species as T. minutilla. 
The cranium being very narrow in the frontal region between 
the superior orbital margins, while the vault of the brain-case is 
externally rounded, smooth, and ample, being occasionally 
marked in some of the Sandpipers by a median, longitudinal 
furrow. 
In both, the lacrymals jut out about in the same proportion, 
and much in the same manner, as they do in a Turnstone. 
Aphriza and Actitis show scarcely any tilting upwards of the 
superior orbital peripheries, a feature so very conspicuous in 
Charadrius squatarola; less so in Hcematopus, and again quite 
absent in Arenaria. 
Oyster-catchers exhibit their strong larine affinities in their 
prominent out-jutting lacrymal bones, and still more in their 
deep and sharply defined glandular fossae over the orbital roofs. 
These latter depressions are but fairly well marked in Turn- 
stones, while in A. virgata, again agreeing in this respect with 
most, if not all true Tringece, they are entirely absent. 
Passing to the basi-posterior aspect of the cranium, the Surf- 
bird is seen to possess, in common with all North American 
Limicolce, a well-marked supraoccipital prominence, which shows 
one on either side, the supraoccipital vacuities being larger in 
this species than they chance to be in any Sandpiper at my 
hand. Plovers have them next largest, they being very minute 
in Arenaria, and not present in any of the skulls of the genus 
Hcematopus before me, although these latter have large supra- 
occipital prominences. 
In this last-named genus, too, the "occipital area" is sharply 
defined by a very strong and raised crest, of a subcircular out- 
line. This crest is very much mellowed down in Charadrius; No. 2.] 
APHRIZA VIRGATA. 
315 
more manifest again in the Turnstones, in which genus the 
occipital area is proportionately much smaller; and finally, in 
Sandpipers and Aphriza the crest is present only as a raised 
line, which in them also bounds an occipital area of a subcircu- 
lar outline, the extremities of the boundary line terminating, on 
either side, at the apex of a paroccipital process. 
Most limicoline types have the plane of the foramen magnum 
nearly coincident with the horizontal plane; and so it is in all 
the skulls of the species now before me. But the outline of 
this great foramen of the occiput differs considerably in the 
several genera, being perfectly cordate in the Surf-bird, with 
the prominent posterior apex standing well in between the pair 
of vacuities of the supraoccipital prominence, which condition 
is enjoyed by most Tringe<z\ while in Arenaria and the Plov- 
ers there is a slight tendency for this aperture to become 
rounder, its circularity becoming quite evident in Hcematopus. 
In all these species, the condyle of the occiput is proportion- 
ately very small in each and every species, it being of a hemi- 
spherical form, and unnotched. 
So in general pattern, taken as a whole, the basi-posterior 
aspect of the cranium is more nearly alike in Aphriza and any 
true tringine form; while in Turnstones the approach is 
towards the Oyster-catchers. And Plovers in this particular 
seem to maintain sort of a mid-position between the two latter 
groups. Hamatopus is notorious for the restricted confines of 
its basi-temporal area at the base of the cranium,1 and this space 
is never very wide and deep in any of the Limicola. 
Figure 3 of my drawings in the plate shows the skull of A. 
virgata upon direct lateral view, and here we have a number of 
points to examine. First, it will be observed that thepars plana 
is of a quadrilateral outline, thoroughly ossified, and with its 
superior and inferior borders not in contact with either the 
orbital roof above or the maxillary bar below. The out-jutting 
lacrymal sends down a spicula of bone, which diminutive process 
turns backwards to have its lower end anchylose with the pars 
1 Had not the writer already fully figured the skeleton of the Black Oyster-catcher 
(which figure and its description has been accepted for publication by the Journal of 
Anatomy of London), I should have taken pleasure in presenting drawings of it in 
the present connection, as it constitutes a wonderfully interesting form for comparison 
in this way with species now under consideration. 316 
SHUFELDT. 
[Vol. IL 
plana, on its anterior aspect at a point below, and to the inner 
side of its externo-superior angle. I am thus particular in de- 
scribing in detail the point of attachment of this extremity of 
the descending process of the lacrymal, for the reason that in 
some Sandpipers it is found to be attached to the very angle 
above referred to, while just before arriving there it sends for- 
wards another small, pointed spine, the extremity of which 
latter is free, and directed somewhat inwards towards the 
rhinal chamber. No little variety, however, is to be met with 
in this particular among the Tringea, and in such a form as the 
solitary Sandpiper (R. solitaries') the free end of the lacrymal is 
attached at a point corresponding to the site of its attachment 
in Aphriza. With the true Plovers the case is still different, for 
in them Xhe pars plana is quite small and with rounded angles, 
while the lacrymal has a long, slender, descending process ter- 
minating in a free point below, and likewise has the anterior- 
projecting spine coming off from a mid-point of its continuity.1 
Arcnaria has a large square pars plana, and in this form the 
lacrymal soon fuses at its lower end with the supero-external 
angle of the osseous lamina in question; while in Hcematopus 
the pars plana is again found to be small, and the lacrymals 
very large, with their descending processes thick and strong, 
being fully anchylosed with the outer border of the pars plana, 
and projecting a short distance below that plate, though not arriv- 
ing at the maxillary bar. 
From all this I should say, then, that in Aphriza, Arenaria, 
and most true Sandpipers the pars plana and the lacrymal bones 
were more or less alike, having much the same relation to each 
other; but in Plovers and Oyster-catchers these structures are 
essentially very different; not only do they differ with each 
other, but with the aforesaid mentioned types, and in a manner 
set forth in the last paragraph. 
Limicoline birds are notorious for the large vacuities that occur 
in their interorbital septa, rendering that osseous partition in the 
vast majority, if in not all the species, a very deficient bulwark 
between the orbits. Hcematopus has it most entire, while in 
Aphriza, the Turnstones, Sandpipers, and Plovers, it commonly 
1 For an excellent figure of this arrangement of the lacrymal, see Huxley's figure 
of the enlarged skull of C. pluvialis in his paper, " On the Classification of Birds." 
P.Z.S., p. 427, Fig. 7, L. No. 2.1 
APHRIZA VIRGATA. 
317 
presents two large deficiencies, or often one imperfectly divided 
by a backward-extending remnant of the septum springing from 
the mesethmoid (Fig. 3). 
In the Sandpipers {Aditis) the entire anterior wall of the 
brain-case is, mesiad, wonderfully lacking in bony support; on 
the other hand, in the black Oyster-catcher the foramina for the 
first pair, as well as for the optic nerves, are entire and com- 
pletely encircled by bone. All the species that we have thus far 
mentioned in this memoir possess a straight and slender infra- 
orbito maxillary bar of nearly uniform calibre throughout. 
Figure 1 of the plate will show the peculiar orbital processes 
of the qziadrates in the Surf-bird, and indeed they are nothing 
less than unusually enlarged, thin, and compressed lamina of 
bone, which in no small degree contribute to the floor of the 
orbit on either side. We also find the two remaining apophyses 
of a quadrate to be more than commonly large and stout, and 
each of these bones has a double mastoidal facet; two mandib- 
ular facets, the inner one being as good as two in Aphriza, 
though less markedly divided in Turnstones, Oyster-catchers, 
or Sandpipers; and finally, a proportionately large pterygoidal 
facet situated upon the internal aspect of the bone, just above 
mandibular articulation. Few departures are to be detected 
from this almost common pattern of the quadrate among the 
Limicolce; where, too, I may add, this bone is always found to 
be perfectly pneumatic, as is the greater share of the skull itself 
among these shore-birds. Still confining ourselves to the lateral 
aspect of the skull, we are to note the long, spicula-form, sphe- 
notic process possessed by Aphriza, and the less conspicuous and 
bifurcated squamosal apophysis; both of which features are far 
better marked in this species than they are in any of its allies 
now before me. Turnstones show quite deeply marked temporal 
fossce that lack something less than a centimetre of meeting 
posteriorly over the supraoccipital prominence. Hcematopus also 
show these, but here they are strictly confined on either side to 
the squamosal region, and do not encroach in the slightest 
degree upon the posterior aspect of the cranium. Charadrius 
likewise faintly displays the same feature, a feature which is 
so prominently seen in any of the skulls of the Larida and 
their kin. 
Careful search through my material fails to detect the presence 318 
SHUFELDT. 
[Vol. II. 
of such fossae, either in the Surf-bird or among the Sandpipers. 
Actitus has a skull as free from any such depressions in its tem- 
poral region as the veriest Thrush that ever lived, and the same 
remark fully applies to our least Sandpiper and others of its 
genus. Another interesting point to notice in the cranium of 
A. virgata is the marvellous degree to which a tympanic cavity is 
exposed, as well as the external opening of the Eustachian tube 
within it. These parts seem to depend upon the large quadrate 
anteriorly, and the somewhat arching paroccipital process be- 
hind, for their protection, in so far as any osseous surrounding 
defence is concerned. 
This condition obtains in varying degree in the allied forms 
we have under consideration ; it being notably the case in Turn- 
stones and in Hcematopus, though Plovers and Tringece are but 
little better off in this respect, perhaps the former of these last 
being the better provided of them all. 
And now, finally, turning to the nether aspect of the skull in 
Aphriza, we are at first struck with the diminutive size of its 
pair of pterygoid bones. These are short, compressed from 
above downwards, sharpened all along their anterior margins ; 
while behind they are characterized by strong basipterygoid 
processes, which articulate in the usual fashion with the facettes 
designed for them at the base of the sphenoidal rostrum. 
In the black Oyster-catcher, a pterygoid is no longer than the 
mandibular aspect of the corresponding quadrate is wide, and 
this statement holds true for the other skulls we have been ex- 
amining in connection with Aphriza. 
For the Plovers the pterygoids are well shown in Professor 
Huxley's figure of Charadriuspluvialis^ although they are com- 
paratively much smaller in such a species, for instance, as C. 
squatarola, or others of our North American Charadriidce. 
Upon comparing the palatines of A. virgata with those bones 
in the allied forms before us, we meet with several excellent dis- 
tinctive characters, especially when we include in this compari- 
son the maxillo-palatines. In Plovers, Sandpipers, Turnstones, 
and Oyster-catchers, the postero-internal angles or heads of the 
1 Ibid. P. Z. S., 1867, p. 427, Fig. 6. These bones are also represented in my figure 
of the base of the skull in the Mountain Plover montanaD Jour, of Anat, and 
Phys. London, October, 1883, Pl. V., Fig. 3,7^., where their relatively small size can 
at once be appreciated. No. 2.] 
APHRIZA VIRGATA. 
319 
palatines, each turns outward as it articulates with the anterior 
end of the corresponding pterygoid ; nor do these palatine heads 
meet each other in the medium plane. With this common fea- 
ture, Aphriza agrees ; but when we come to examine the postero- 
external angles of these bones, we find them obliquely truncated 
in our subject, or at least the hinder ends of the palatines them- 
selves thus obliquely truncated, forming obtuse postero-external 
angles, which angles are rounded off in Actitus, Charadrius, and 
in Arenaria, while in Hcematopus a moderate, backward-extend- 
ing and rounded process is found at their site. In this latter 
species, too, the internal and external laminae of a palatine bone 
are very conspicuous, as they are, though less so, in the Surf- 
bird, Spotted Sandpiper, and the Black Turnstone. Plovers have 
this character much subdued. Mesiad, the superior and at the 
same time, internal borders of the palatines, beyond the "ptery- 
goidal processes," along the rostrum, come in contact with each 
other in all of the enumerated species ; and in all completely 
fuse anteriorly with the hinder limbs of the vomerine bifurca- 
tion. Coalescence, again, is most thorough in all these forms 
among the anterior ends of the palatines and those parts of the 
maxillaries and premaxillaries with which they come in contact, 
upon either side (Fig. 2). From this description, then, of the 
palatines, it is clear that A. virgata possesses them in a form 
essentially its own, and further that the pattern varies in all the 
allied groups or species. 
Not so, however, is this altogether the case with the maxillo- 
palatines. One of these in Aphriza virgata is seen to be a 
shell-like process; extending backwards, concave externally; 
the reverse being the case upon its mesial aspect; perforated by 
a few foramina; and finally, not united along its inferior border 
with the palatine of the same side which extends beneath it. 
Charadrius also possesses a maxillo-palatine almost identical 
with this, and in the last-named feature, the true Sandpipers 
also agree, but in these latter birds the bone is much narrower 
and more curved. 
Passing to the Oyster-catchers and Turnstones, a very different 
state of affairs is to be met with ; for in Hcematopus, for instance, 
we find a maxillo-palatine to be quite a thick, laterally compressed 
and unperforated lamina of bone, which completely fuses with 
the corresponding palatine for the entire length of its inferior SHUFELDT. 
[Vol. II. 
320 
margin, appearing in the adult to be a part of the palatine itself. 
A similar arrangement is to be found in Arenaria, - so that in 
this character, Turnstones and Oyster-catchers agree, while the 
Surf-bird, Plovers, and Tringcce are fundamentally alike; in 
the first, the maxillo-palatines have, in the adult individual, 
coalesced with the palatines, each on its own side; while in the 
three last-mentioned families the maxillo-palatines are indepen- 
dent and freely projecting processes. 
Aphriza agrees with all the allied types before us, in having a 
long, slender vomer, its posterior limbs anchylosing with the 
palatines behind, and a median carination extending for its en- 
tire length along its nether aspect. With the exception of 
Hcematopus, the bone in all is pointed anteriorly, and in the ex- 
cepted genus it is at that end distinctly bifurcated.1 
In the form of its mandible, A. virgata substantially agrees 
with the Plovers and Sandpipers. The upturned, slender 
process at the angle is well developed, and the inturned ends 
of the articular cups are quite prominent, showing the usual 
single pneumatic foramen at each extremity. Two vacuities 
occur in either ramus, a small surangular one, and a long, semi- 
closed, splenial one, the sides of the jaw in this region being verti- 
cal and rather deep. The symphysis is rounded beneath, concaved 
above, and equals in length about one-fifth the length of a ramus. 
In the Black Oyster-catcher the symphysis equals more than a 
third of the ramus, and is scarcely at all excavated above, while 
it is decidedly wedge-shaped below. Arenaria also has a deep 
symphysis to its mandible, but here again it is concaved above 
and rounded beneath, and the larger of the two ramal vacuities 
is represented by a very open, subelliptical foramen. 
Among the more ordinary types of existing birds, the hyoidean 
apparatus, as we know, presents but few differences when we 
come to compare it for the various families. Our subject forms 
no exception to this statement, and little need be said here in 
reference to this part of the skeleton in Aphriza. We find the 
arches alluded to to be very delicately constructed, and have 
the basibranchials in two separate pieces, the second one being 
very short and finished off behind in cartilage. The cerato- 
1 Here we have a departure from the rule laid down by Huxley, who has said 
that in Schizognathous birds, " the vomer, sometimes large and sometimes very small, 
always tapers to a point anteriorly." P. Z. S., 1867, p. 426. No. 2.] 
APHRIZA VIRGATA. 
321 
hyals largely coalesce, but the ossification proceeds but a lim- 
ited distance into the glosso-hyal, which latter element in the 
adult is found to be in cartilage only. Thyro-hyals are notably 
slender and of considerable length; they, too, being posteriorly 
tipped with cartilage throughout the life of the individual. 
Most Sandpipers possess a hyoidean apparatus similar to the 
one I have just described for Aphriza ; and, indeed, no material 
difference can be detected for the allied species from other 
groups.1 
Upon carefully examining the special ossifications of the ear 
in a skull of A. virgata, I fail to detect anything worthy of 
detailed record ; the form of the stapedial plate and the coossi- 
fied shaft of the columella have much the same pattern as 
we find these parts in the Fowl, and so clearly drawn for us 
by Parker.2 Dr. Streets, in preparing my skeletons of Aphriza 
for me, threw away the eye-balls, so I am unable to make any 
observations here upon the sclerotal plates in this species ; they 
have also been lost from my skeletons of Hcematopus, the Turn- 
stones, and Plovers. 
This completes what I have to say here in reference to the 
comparative osteology of the skull and its associate parts of the 
skeleton in Aphriza virgata, and such deductions as may be 
drawn from the study the writer proposes to defer making until 
the closing paragraphs of the memoir are arrived at, when the 
skeleton as a whole, in this species, will be passed in review, and 
the proper comparisons made with the others we have been 
considering. 
Remainder of the Axial Skeleton. - Since the days that 
anatomists first took to counting the total number of vertebrae 
in the spinal column of any of our existing Carinatce, there 
have been difficulties and differences upon that point; and 
as to the differences, these have been but multiplied by the 
many attempts that have been made to state exactly the num- 
1 Unfortunately the hyoid arches of both my specimens of Hamatopus bachmani 
have been lost, and I am unable to add, by way of comparison, anything about them 
here. I am inclined to believe, however, that if any essential difference is to be 
hereafter detected in these arches for adult specimens of various representations of 
the Limicolce, it may be that some form will show a fusion of the first and second 
basibranchials ; but I look for nothing more than some such a difference as this, and 
even it may never be found to exist. 
2 Parker, W. K., Morphology of the Skull, p. 258, Fig. 74, m.st. and st. 322 
SHUFELDT. 
[Vol. II. 
ber of cervical vertebrae, and the number of true dorsals, in any 
particular species. In the case of the total count for the verte- 
brae in any adult bird of the kind alluded to, the trouble is with 
the pelvic sacrum, and the skeleton of the tail. In nearly 
every example it is almost impossible to decide with certainty 
as to the number of vertebrae that have been incorporated in 
its fusion ; while in the case of the tail, the pygostyle is the 
stumbling-block, for sometimes in representatives of the same 
species this bone in one individual will thoroughly appropriate 
to itself a terminal caudal vertebra, that will perhaps remain 
free in the other, thus making an additional segment for a bird 
of the same species. Then, again, it is pretty well agreed that 
when we come to define the line between cervical and dorsal 
divisions of the column, we look chiefly to the ribs for assist- 
ance ; yet these are by no means to be always relied upon ; as 
sometimes, in the same species, an additional pair may remain 
free at the further end of the cervical region, or an additional 
pair (always at the anterior part of the dorsal division) may 
connect with the sternum by a pair of haemapophyses. So 
that one observer might state that the first pair of free ribs 
in a certain species of bird were to be found upon the 13th 
cervical vertebrae, while another observer, equally careful, finds 
an individual of the same species wherein it is the 14th cer- 
vical that has the leading pair of free ribs. Thus it is that 
confusion and doubt creeps into the work. 
After having paid no little attention to the osteology of birds 
for more than ten years, now, I am fairly convinced that there 
is but one safe or very nearly safe method of arriving at the 
total number of vertebrae in the spinal column of any particular 
species of the class, and to do it we should have the prepared 
skeletons of the young of the species, two, or, better, three, of 
them, made just at the time when fusion is about to take place 
among the vertebrae, which eventually go to form the pygostyle 
in the adult, and when the vertebrae composing the pelvic 
sacrum are all still individualized. Then the count can be made 
with certainty, and we should not fail to have by us at the time 
several skeletons of the adult of the same species for compari- 
son. Finally, our remaining difficulty, it is evident, cannot be 
settled in any such manner, and the only course left to pursue 
here, is to state, if possible, the general rule and the noted No. 2.] 
APHRIZA VIRGATA. 
323 
exceptions. For instance, we may say in a certain species, it 
has been found as a general rule that free ribs occur upon the 
13th and 14th vertebrae, and that upon the 15th, the ribs connect 
with the sternum by costal ribs, but examples have been found 
in the same species where free ribs were found upon the 12th, 
13th, and 14th vertebrae ; or again, a specimen of the same 
species showed free ribs on the 12th and 13th vertebrae, while 
the ribs on the 14th connected with the sternum. This is by 
no means a fanciful description, as I have studied cases quite 
like it, and it only goes to show the utter hopelessness of laying 
down hard and fast lines in describing the cervical and dorsal 
divisions of the vertebral column in existing birds. On the 
other hand, I am inclined to believe that if the total number of 
vertebrae be counted in the skeleton of the chick of any par- 
ticular species, the number will be found to be quite constant. 
Now I believe, after carefully re-counting the vertebrae in the 
skeleton of the tail in the Mountain Plover (^. montanus), 
there should be a quaere placed after the number I have given 
in my memoir upon the osteology of that species,1 as I believe 
it should be seven, not counting the pygostyle ; further, I am 
inclined to think I made another miscount, for some reason, in 
my specimen of Hcematopus, and too many vertebrae have been 
mentioned as forming the skeleton of its tail in another paper 
of mine.2 
At the present writing I have not at hand the immature skel- 
etons of any of the species, we have here under consideration, 
and consequently am unable to carry out the suggestions made 
in the foregoing paragraphs, so I can candidly admit that the 
number of vertebrae given below as found in the sacra of the 
birds there enumerated must be taken cum grano salts ; although 
I will add that I counted them to the best of my ability: the 
process can often be assisted by holding the pelvis up to a strong 
light, with the dorsal aspect towards you. 
In my female specimen of A. virgata, the " epipleural appen- 
dages " are present upon the pair of free ribs on the 15th 
vertebra, the reverse being the case in the skeleton of two males 
1 Ibid., p. 97. 
2 Shufeldt, R. W., "Osteology of Numenius longirostris, with notes upon the 
Skeletons of other American LimicolaP Journ. of Anat, and Phys. London. Vol. 
XIX., October, 1884, pp. 71, 72. 324 
SHUFELDT. 
[Vol. II. 
of the same species. Charadrius sqtiatarola seems to have the 
usual cervical vertebrae (6th to 9th inclusive) modified beneath for 
the carotid canal; while in a Killdeer before me, the series so 
modified includes the fifth cervical. Only the free caudal verte- 
brae have been counted in the subjoined "Table," omitting in 
every case to include the pygostyle; a bone that may include 
several more, and perhaps vary for the species here given. 
TABLE. 
Aphriza virgata 
Charadrius squatarola . 
Actitis macularia .... 
Haematopus bachmani . 
Arenaria melanocephala 
Rhyacophilus solitarius 
Species. 
G) GJ W GJ GJ GJ 
Number of cervical vertebrae without 
free ribs. 
The first cervical vertebra that has 
free ribs. 
In CH 4- Ln in Ln 
The second cervical vertebra that has 
free ribs. 
.. 16 
.. 16 
.. 16 
15 16 
. . 16 
. . 16 
The third cervical vertebra that has 
free ribs. 
The first vertebra that has ribs which 
reach the sternum by costal ribs. 
CN ON Os O' 0N 
Number of vertebrae that possess ribs 
which reach the sternum by costal ribs. 
« „ « HH hl 
Pairs of sacral ribs (they do not reach 
sternum). 
U GJ 4- to b) GJ 
Number of vertebrae in pelvic sacrum. 
*4 CO *4 -4 ^4 CO 
Number of free caudal vertebrae. 
21 42 
21 40 
21 40 
21 42 
21 42 
21 40 
Total number of free prepelvic verte- 
brae. 
Total number of vertebrae in column 
exclusive of pygostyle. 
6th to 9th 
6th to 9th 
6th to 9th 
6th to 9th 
6th to 9th 
6th to 9th 
Carotid canal in following cervical ver- 
tebral (inclusive). 
We find nothing especially noteworthy in the vertebrae of 
Aphriza virgata, as they in a general way are fashioned upon 
the usual ornithic pattern of those bones among ordinary birds. 
The axis, and the three cervical vertebrae that follow it, are 
especially conspicuous for their prominent neural and haemal 
spines, and their postzygapophyses, which latter extend upwards, 
outwards, and backwards, as strong processes in the second and 
third cervicals. This last feature is not so manifest in Aphriza 
but becomes more so in Arenaria, and in Hamatopus arrives at 
its maximum development, for in this species the postzyga- 
pophyses of the axis and the vertebra next behind it are nearly 
as lofty as the great neural spines upon these vertebrae, giving 
them the appearance of being tricornuted upon their dorsal 
aspects. Oyster-catchers have comparatively short parapophy- No. 2.] 
APHRIZA VIRGATA. 
325 
ses in the cervicals, while in both the Surf-bird and in Turn- 
stones these are quite long and spicula-form in the fourth, fifth, 
sixth, and seventh vertebrae; and these last species differ with 
Hcematopus, in that in this bird the vertebrae of the entire col- 
umn are unusually large in proportion to the size of their owner. 
Free ribs in all the species before us possess both capitula and 
tubercula; the leading pair being of diminutive size, and gradu- 
ally increasing as they near the ribs of the dorsal region. The 
last pair usually do not develop unciform appendages, though 
they may do so, as in the skeleton of my female Surf-bird, and 
an Oyster-catcher before me. In the dorsal region only the first 
two, or at the most, three, leading vertebrae possess haema- 
pophyses, while ossified metapophyses link their transverse pro- 
cesses together above. Aphriza has the neural spines of its 
dorsals very low, they being more lofty in the Turnstone, and 
still more so in Hcematopus. Plovers and Tringece have them of 
a medium height. We find few or no especial distinctive differ- 
ences in the true dorsal ribs of the several species before us, to 
render any aid as pointing to near or remote kinship in the case 
of any two species compared. Agreeing with the majority of 
limicoline birds, these parts are for the most slender, curved as 
usual, and rather long, with uncinate processes that do not 
completely anchylose with the several ribs to which they belong. 
Elsewhere, as I have already said above, I have presented 
figures of the pelvis of the Black Oyster-catcher, and it can be 
said here now that this bone in Hcematopus very markedly dif- 
fers from the pelvis either of Aphriza or of Arenaria. Indeed, 
in some respects, the pelvis of an Oyster-catcher reminds us not 
a little of the pelvis in Gallus bankiva; it lacks, however, the 
propubic spine, and the antero-median borders of the ilia do not 
fuse with the sacral crista. Further, it is noted for having a 
certain solidity and thickness of the several bones that com- 
pose it, quite in keeping with the other parts of the skeleton of 
this powerfully built bird. It differs with the general plan of 
the pelvis in Aphriza (Fig. 21), and in the Turnstones, in that 
in Hcematopus the ilium forms a ledge, overhanging to some 
extent the ischium and the ischiadic foramen; the postpubis is 
far separated from the lower ischiadic margin, and is truncated 
almost immediately after passing the ischium behind; finally, 
anchylosis is very strong between all the several bones making 326 
SHUFELDT. 
[Vol. II. 
up the pelvis in the Oyster-catcher, so that prolonged macera- 
tion does not tend to separate them. 
One skeleton of Hcematopus at my hand shows a curious dila- 
tation of the latter third, or less, of the postpubis of the left side, 
not present in any other specimen, nor upon the opposite side 
of the same individual. All these points and characters are 
very different in the pelvis of the Surf-bird, wherein this bone 
is remarkable for its breadth, and for being, as it were, flattened 
out in the vertical direction, - shallowed ; its sides narrow, and 
ilia more or less spread. As among Plovers, and most or all 
Sandpipers, it shows a double row of interapophysial foramina 
down the sacrum (Fig. 21); and the anterior ends of the ilia are 
subtruncated and embellished with a delicately raised rim, 
nearly always to be seen in limicoline birds. Between the 
hinder ends of ilium and ischium, a rounded notch occupies the 
entire margin of the bone, which notch in Hcematopus is angu- 
lated. Postpubis is in contact, more or less, for its entire 
length with the lower border of the ischium above it; and is 
carried beyond for some little distance to a pointed extremity, 
much the same being the case in the Turnstones. 
The ischiadic foramen is of considerable size, and of a subel- 
liptical outline; this vacuity being unusually large in an Oyster- 
catcher, and such a species as the Killdeer Plover. 
Without, then, further entering upon a fuller discussion of 
minor details, although they have all been carefully noted and 
weighed by the writer, I would state that in so far as the Hcema- 
topodidce are concerned, the pattern of the pelvis in that group, 
as represented by the bone from the skeleton of H. bachmani, 
is of quite a different style from the pelvis of either Aphriza or 
Arenaria. That when we come to study the pelvis of A. vir- 
gata, and compare it with the bone in allied types, we discover 
that notwithstanding the fact that it exhibits many characters 
held in common by the limicoline forms generally, still it is 
impressed with a character of its own, being proportionately 
broader and shallower than the generality of either charadrine 
or tringine pelves. It is most nearly approached by the pelvis 
of a Turnstone, while this latter bird shows in its pelvis a marked 
shading towards the style of the bone in the Plovers. 
Again, Actitis and PEgialitis vocifera possess pelves almost 
identically alike in their general form ; and in both these No. 2.] 
APHRIZA VIRGATA. 
327 
species the postpubis is separated by more or less of an interval 
from the lower margin of the ischium of the same side, along 
the middle of its continuity. 
Figure 20 of the Plate shows the form upon lateral view of 
the pygostyle in A. virgata, and that is not far departed from 
by the representatives of nearly affined Limicolce. In Oyster- 
catchers it varies, because in them the ultimate caudal vertebra 
usually is coalesced with the bone, giving it the different shape. 
The Surf-bird and its allies show the diapophyses of the caudal 
vertebrae to be spreading in the first few leading ones; then in 
mid-course they become shorter and bend downwards; then long 
and more flaring again, to terminate by an aborted one just 
before coming to the pygostyle. The ultimate two or three 
may develop bifid haemapophyses, which anteriorly stand be- 
tween the joints of the centra as chevron bones ; this feature is 
well marked in H. bachmani. 
Oyster-catchers have the neural spines of the caudal verte- 
brae notched in front, while Aphriza, Charadrius, and the major- 
ity of the Tringece possess these apophyses as plain points. 
That time-honored standby of avian skeletologists, the ster- 
num, presents among the birds we have been noticing in this 
memoir but few insignificant departures from a common pat- 
tern, except in the cases of R. solitarius and Actitis. 
Aphriza has the bone (Figs 12 and 15) doubly notched on 
each side posteriorly, the outer notches being fully twice the 
dimensions of the inner ones, - the outer pair of xiphoidal pro- 
cesses thus formed having a tendency to flare outwards; 
otherwise, the outline of the sternal body is quadrilateral, being 
much concaved above and correspondingly convexed upon its 
pectoral aspect. Either costal border shows six haemapophysi- 
cal facettes for the costal ribs, while extensive " costal pro- 
cesses " of a triangular outline rear above these in front. 
The manubrium is small and sessile, being sharp beneath and 
blunt above, and directly between coracoidal grooves, thus pre- 
venting the coracoids in the articulated skeleton from infringing 
upon each other; the reverse being the case in Oyster-catchers 
and Arenaria, wherein these bones are in contact posterior to 
the manubrium, when the former are in situ. Returning to the 
sternum of Aphriza, we are to note its ample carina, which ex- 
tends the entire length of the sternal body, being thickened for SHUFELDT. 
[Vol. II. 
328 
the upper two-thirds of its anterior border, sharpened below, and 
concaved throughout. Along its lower margin a thickened rim 
defines its edge, which is outlined by a long, gentle convexity 
from carinal angle in front to mid-xiphoidal process behind 
(Fig. 12). 
Viewing the bone upon its pectoral aspect, a muscular line - 
the interpectoral muscular line we will call it here - extends, 
on either side, from the boundary of the costal fossa beneath, 
to the inner of the pair of xiphoidal notches behind, to the 
mesial edge of the same. This line occurs sharply defined 
in both Plovers and Oyster-catchers, where it is similarly drawn, 
as it is likewise in a Turnstone. 
Arenaria has a sternum, which, excepting the fact of its being 
a size smaller, is the perfect counterpart of the bone in A. vir- 
gata, though in the latter species the sternal keel seems to be 
comparatively a trifle deeper in proportion. 
When I published my Osteology of Numenius longirostris, I 
therein stated that the sternum to a skeleton of the Spotted 
Sandpiper (A. macularia) possessed two notches upon either 
side, a large outer pair and a very small inner pair. Those 
skeletons were prepared and the species diagnosed by a very 
incompetent man, and at the present writing I am not prepared 
to vouch for the correctness of the statement. I have had to 
regret many times since, the confidence I placed in the identifi- 
cation of the skeletons which went to make up the collection 
of the vertebrate series in the Museum and Library of the 
Surgeon General's Office of the Army at Washington; they 
were the cause of a number of oversights creeping into my 
avian osteology in early days, which now are being corrected 
in my more recent papers. So I am inclined to believe that the 
forir-notched sternum of Actitis may be a specimen from the 
same category; it was "on file" in the same institution. At 
any rate, we have before us now an excellent skeleton of Actitis 
of my own collecting, and it possesses but a pair of notches in 
its sternum, otherwise the bone has the general pattern of the 
Limicolce at large. 
In a few days The Auk for July (1888) will be issued, and in 
there, in a letter to its distinguished editors, I point out the 
fact that the Solitary Sandpiper is another species which has 
but a pair of notches in its sternum, and suggest that its old No. 2.] 
APHRIZA VIRGATA. 
329 
genus, Ryacophilus, be restored for it (at the present writing it is 
a subgenus of Totanus). But the majority of Tringece, as for 
example, all the true representatives of the genus Tringa, have 
the "four-notched" sternum, and the fact that Aditis and Rya- 
cophilus have each only a pair of notches, may be of more signi- 
ficance in so far as the true affinity of those two species is 
concerned, than some of their external characters (that are so 
very subject to change from minor causes), but which have alone 
been relied upon thus far by the systematist, to classify them. 
Finally, then, the fact may be broadly stated, that with respect 
to the sterna of Aphriza and Arenaria, they are to all intents 
and purposes the miniatures of that bone as we find it in Hcema- 
topus bachmani ; and although this is very interesting, the cir- 
cumstance goes to prove, in the light of other skeletal characters 
of these species, we would utterly fail were we to rely upon this 
bone alone as any indication of family, or much less, generic, 
affinities. 
All the sterna that the writer has examined from the skele- 
tons of North American Limicolce were non-pneumatic bones, 
and if it is ever found to be otherwise, we must believe that it 
is the exception. In a specimen of Tringa minutilla in my col- 
lection, the bone is so thin that a large perforation occurs on 
either side of the keel in the sternal body, and one through the 
keel itself, high up in front. 
Speaking of the pneumaticity of the skeletons of the several 
species we are here considering, I am strongly inclined to be- 
lieve that the skull is the. only part in any of them where air 
gains access to the interior of the bones, and as a rule to no 
very great extent, comparatively, there. 
Turning our attention next to the shoulder girdle (Figs, n, 
16, and 17) in Aphriza virgata, we meet with a scapitla having 
much the form that that bone assumes among the Shore Birds, 
generally, being shaped a good deal like the blade of a miniature 
cimeter, and when in situ in the articulated skeleton bearing the 
usual relations to the clavicle and coracoid of its own side. A 
coracoid is conspicuous for its short, sub-cylindrical shaft; its 
head being tuberous and much crooked over towards the median 
plane, while the sternal extremity of the bone is not only thick- 
ened from before, backwards, but much expanded lateral-wise, 
showing at its outer angle a prominent lamelliform process, com- 330 
SHUFELDT. 
[Vol. II. 
pressed in the antero-posterior direction, and curved somewhat 
upwards. Os furcula typifies the U-shape form of the bone, and 
has in all Limicolce, that the writer has ever studied, the merest 
apology for a hypocleidium, below. Its limbs are of nearly 
uniform calibre throughout, being slightly heavier above than 
below, and moderately compressed transversely for their entire 
lengths. Their heads, that is, the free ends of the clavicular 
limbs, are much elongated, and when articulated in situ, reach 
far back to the heads of the scapulae. Viewed upon direct lat- 
eral aspect, it will be noticed that the os furcula is very con- 
siderably, though gracefully, curved in the antero-posterior 
direction, the convexity of the curvature being to the front. 
In Hcematopus the shaft of a coracoid is pierced by a small 
oval foramen, from before, backwards, and just below the scap- 
ular process, that does not occur in either the Surf-bird or among 
Turnstones. 
Now, aside from the exceptions of this foramen, and differen- 
ces as to the method of articulation of the coracoids with the 
sternum, already alluded to, and finally, the mere matter of size, 
- the description of the shoulder girdle here rendered for 
Aphriza, will apply almost equally well to the shoulder girdles of 
the Tringece, the Turnstones, and the Plovers, and no doubt to 
many other Shore Birds. Were we to depend for characters, 
upon which to base distinctive differences, on the shoulder gir- 
dle alone in Aphriza, Arenaria, and a true Plover of a correspond- 
ing size, we should surely be disappointed, for the bones of this 
arch do not offer them in the genera enumerated. 
Of the Appendicular Skeleton. 
The Pectoral Limb.-The majority of the bones of this part 
of the skeleton in Aphriza are shown by my drawings in the 
Plate, as already stated above. The corresponding parts for 
Hcematopus and Plovers have been given in other connections. 
It will be seen that in the humerus of our Surf-bird, both 
"radial crest " and "ulnar tuberosity" are strikingly well devel- 
oped, while the articular humeral head between them is rather 
small (Figs. 18 and 19). The pneumatic fossa is an extensive 
excavation, although the bone itself is not pneumatic. As to 
the shaft, it shows but little curvature, is smooth, and subcylin- 
drical in form. At the distal extremity of the bone, a conspicu- No. 2.] 
APHRIZA V/RGATA. 
331 
ous " ectocondyloid process" is to be observed; and at the 
extreme end, the oblique and radial tubercles, or condyles, stand 
out with more than their usual boldness, a condition which is 
enhanced by the fact that just above them there exists a fossa 
of some considerable depth. When the bones of this limb are 
articulated, and the arm tightly closed, the head of the radius 
accommodates itself to this latter depression at the distal end of 
humerus. Upon the anconal aspect of this extremity of the 
bone, we also find strong and powerfully impressed grooves for 
the passage of the tendons of muscles inserted upon it, and at 
the shoulder joint, during the life of the individual. 
Neither radius nor ttlna exhibit any very great degree of cur- 
vature, and as a consequence the interosseous space in the 
articulated arm is not very wide. The latter bone develops 
down its shaft a row of some eight or nine papillae, for the quill 
butts of the secondaries of the wing, a feature common to so 
many of our Shore Birds. Comparatively speaking, the distal 
extremities of these bones of the antibrachium in Aphriza, as in 
all Limicola which the writer has examined, are small, though 
to this fact the Oyster-catcher seems to offer an exception. 
The carpal segments are two in number, as usual, and present 
nothing, either in their size, form, or method of articulation, in 
any way different from what we usually find. 
In the pinion (Figs. 13 and 14), the carpo-metacarpal bone is 
long, large-headed, and with its coalesced shafts straight, and 
more than commonly close together. As a negative character, 
we miss the overlapping process at the proximal end of the mid- 
metacarpal, - an apophysis which characterizes this bone in all 
true Gallina. Prominent and upturned, the poll ex metacarpal 
supports a phalangeal joint that is clawless; the same being 
the case with the distal phalanx of the digit of the succeeding 
finger. 
Regarding the expanded and proximal phalanx of this last 
digit, it is to be noted that its somewhat narrow blade, although 
more or less scooped out upon either aspect, is never perforated 
as we find it in some of the Larida. At its further end a little 
process protrudes, against which, in life, the head of its follow- 
ing phalanx plays ; this latter joint being very long and slender, 
as in some Plovers and Sandpipers. The little phalangeal joint 
of the remaining digit is crooked so as to accommodate itself to 332 
SHUFELDT. 
[Vol. II. 
the outline of the posterior border of the blade of the proximal 
joint of index finger. 
Equal correspondence of characters in the pectoral limbs of 
Plovers, Sandpipers, and Turnstones, as compared with the 
bones of the skeleton of the arm, in Aphriza seem to prevail, 
as they were found to prevail among the shoulder girdles and 
sterna of the same species. Practically, the description I have 
just given for the arm bones of the Surf-bird will answer all 
purposes of exactness for the representatives of the genera 
Tringa, Arenaria, and most, if not all, true pluvialine types. 
This, of course, does not apply to the mere question of the 
difference of size, but to the salient features only. Our engag- 
ing little T. mimitilla has a pectoral limb, the skeleton of which 
corresponds, in so far as its available taxonomic points are con- 
cerned, with the same parts in the Surf-bird, to a degree of 
nearness most exasperating to the comparative anatomist, and 
discouraging to one eagerly searching for strong diagnostic dis- 
tinctions in the structure of such types. Turning to the skele- 
ton of the black Oyster-catcher we find but one sure point of 
difference that can be relied upon, apart from its size, as distin- 
guishing the fundamental plan of the structure of the skeleton 
of its arm from the above species. In this bird we find a claw 
upon the end of the pollex phalanx, which, by the way, is also 
found in some Curlews. 
Aside from this character, then, we might, were it possible, 
reduce the arm bones of this Oyster-catcher until they equalled 
in size those of a Surf-bird, and it would puzzle the best of us 
to correctly decide to which species either belonged, after such 
a hypothetical reduction. 
These limicoline birds all seem to lack the os humero scapu- 
lare at the shoulder joint, and I have yet to find in any of 
them the sesmoidal ossicles at the elbow, such as we do find, 
for instance, in some of the Puffins. 
Of the Skeleton of the Pelvic Limb. - In Aphriza virgata the 
femur possesses only a moderately sized, semi-globular head, 
with barely any depression on the top of it for the insertion of 
the teres ligament. The trochanter is lofty and thick, reaching 
conspicuously above the articular summit of the bone, which 
latter is broad and smooth. Little or no curvature is met with 
as we descend the shaft, and this is quite cylindrical for its No. 2.] 
APHRIZA VIRGATA. 
333 
middle third. As to the condyles, we find nothing worthy of 
special note, as they are fashioned much as we generally find 
them in ordinary birds; the external one being the larger and 
lower, while it shows behind the usual vertical groove for the 
head of the fibula. 
In the leg the fibula is free, not anchylosing with the tibio- 
tarsus, being but a trifle longer than the half of this latter bone. 
Its lower two-thirds are almost hair-like in proportion, and as 
for that matter it would seem that this delicate osseous fila- 
ment can add next to nothing to the skeletal support for the 
structures of the limb (Fig. 8). 
Tibio-tarsus also has very nearly a straight shaft, but com- 
monly shows, as in Hcematopus, a curvature in the transverse 
plane, being gently bowed outwards towards the fibular side. 
Its procnemial process is much the larger of the two laminar 
crests in front of the head of the bone, and it is pointed almost 
directly forwards, while the ectocnemial process is turned out- 
wards, and curves downwards, as a pointed hook. 
They both rear slightly above the articular summit of the 
proximal end of the shaft. Occupying the middle part of the 
upper third, the fibular ridge stands out quite boldly at the 
outer aspect, as in all birds. Proceeding to the distal ex- 
tremity of the shaft we find a marked difference in the form of 
the two condyles, the outer one being decidedly uniform in out- 
line, and the inner one more elongated and showing compres- 
sion in the vertical direction. Between, and at the same time 
above them, in front, we detect the usual little osseous bar for 
tendinal confinement. A patella is absent in the Surf-bird, the 
case, I believe, in all Limicolce-at least the writer is not aware 
of any exception to the rule. 
Having a length just equal to that of the femur, the tarso- 
metatarsus exhibits at its usual site at the upper end and back 
part of its shaft a stumpy hypotarsus, that appears to be both 
pierced and grooved, for the guidance of tendons. Two minute 
foramina in this locality also perforate the bone in the antero- 
posterior direction, which is also the case in an Oyster-catcher. 
A mid-section of the shaft of tarso-metatarsus would show a 
quadrilateral outline, but the calibre increases, and this form 
changes as we pass to its distal extremity, where we find three 
large and prominent trochleae, the middle one of which is the 334 
SHUFELDT. 
[Vol. II. 
biggest and placed the lowest on the bone. The ones upon 
either side of it are directed to the rear. At its usual site, 
above the innermost one, is faintly to be discerned the facette 
for the articulation of the os metatarsale accessorius, of hallux 
phalanx. Already the ornithologist has placed at our command 
the fact that the arrangement of the joints of pes in this spe- 
cies, as in Areuaria and many Tringece, is upon the most com- 
mon plan of 2, 3, 4, and 5 bones from hind toe to outermost 
digit, respectively, and a study of these in the skeleton reveals 
nothing peculiar in them. 
The Black Turnstone being a somewhat smaller bird than 
Aphriza, it has a correspondingly smaller pelvic limb and skele- 
ton ; aside from this fact, however, the several bones of that 
extremity are fashioned quite as they are in our Surf-bird, and 
present no decided differences in any particular. This state- 
ment applies with equal truth to various Sandpipers, and would 
to Charadrius were it not that in the latter genus a dispropor- 
tion, and a disproportion merely, exists in the hallux, - that 
toe being much smaller in the Plover, though all the other 
bones of the limb are remarkably similar in all their essential 
features, bone for bone, as we have described and figured them 
for Aphriza virgata. Hallux is also missing in Hcematopus, 
where we sometimes find a process of bone anchylosed to the 
shaft of tarso-metatarsus, at a point where the accessory meta- 
tarsal is commonly attached by ligament. In a specimen of 
H. bachmani before me, such a process is to be found upon the 
shaft of the tarso-metatarsus of the right pelvic limb, but is 
missing from the corresponding bone in the opposite limb of 
the same individual. Strange to say, this is precisely the state 
of affairs in the skeleton of a second specimen at my hand, - 
the process is present in the right, absent in the left. It may 
prove of interest to look further into this point with additional 
specimens. Returning to our comparison between the pelvic 
limb of this Oyster-catcher, then, with the same part of the 
skeleton in Aphriza, we may say in brief that beyond the few 
exceptions mentioned the limb in the former is simply an ampli- 
fication of the limb in the latter species, - its bone for bone, 
character for character, - the same thing over again, only the 
mould happened to be several sizes larger wherein the skeleton 
of these parts in our Oyster-catcher were cast. No. 2.] 
APHRIZA VIRGATA. 
335 
Having now briefly passed over the more salient characters 
to be found in the skeletons of Aphriza, Arenaria, Hcematopus, 
and several representative Plovers and Sandpipers, we are, I 
think, in a position to offer our conclusions upon the affinities 
of the Surf-bird, in so far as they seem to be indicated through 
this study of its osteology. 
Conclusions. 
Taking the skull as the part of the skeleton, I would invite 
attention to the peculiar form of the superior osseous mandible 
in Hcematopus; and in the cranium of the same bird to the deeply 
sculpt, supra-orbital, glandular depressions ; to its bold, outstand- 
ing lacrymal bones, and their mode of meeting the pars plana on 
either side; to its vomer, which is forked at its anterior ex- 
tremity ; to the absence of the supra-occipital foramina; to the 
complete coalescence of the maxillo-palatine, with the palatine 
of the same side; to its cranial foramina in front for exit of 
the first and second pair, they being completely surrounded by 
bone; and, finally, to the subelliptical outline of its foramen 
magnum. All these points, and many minor ones in its skull, 
are totally different in Aphriza, and alone are sufficient to war- 
rant us in at least drawing strong lines of family distinc- 
tion between these two types. Equally clear is the differ- 
ence between the skull in Hcematopus and Charadrius, but 
when we come to compare it in the former with the skull 
of Arenaria, a faint indication of affinity may be made out, 
which indication is principally seen in the Turnstone, having 
similarly formed nasals; feebly developed supra-orbital glandu- 
lar depressions ; a somewhat similar superior osseous mandible; 
and, lastly, in that its maxillo-palatines unite along their entire 
lower margins with the corresponding palatine on either side 
below them. 
The differences presented to us, however, in the skulls of 
Hcematopus and Arenaria, amount at least to family distinction. 
So, too, when we come to compare the skulls of Arenaria and 
Ckaradruts, they are very different, and undoubtedly belong to 
representatives of distinct families. 
Comparing next the skull of a Turnstone with that of the 
Surf-bird, we are at first struck with the radical differences 336 
SHUFELDT. 
[Vol. II. 
exhibited in the superior osseous mandibles of the two species. 
Then in the former the maxillo-palatine fuses, in a manner 
already described, with the palatine, and finally, the " croto- 
phyte fossae" are pretty well defined in Arenaria, whereas in 
Aphriza not a vestige of them is to be seen. The posterior 
external angles of the palatines are obliquely truncated in A. 
virgata, rounded in the Turnstone ; and finally, the mandibles 
of these two types are essentially very different, both as to 
form and structure, as in the Turnstone a large vacuity persists 
between the forks of the dentary, which is nearly absent in the 
Surf-bird. 
Taking it all and all, the skull, mandible, and associated parts 
of the skeleton of the head in Aphriza agree more nearly with 
the corresponding structures in the skeleton of the head of some 
of the larger Tringece than with any other class of limicoline 
birds. I have arrived at this point by carefully comparing this 
part of the skeleton in the numerous species at my hand, and I 
am fully convinced of the truth of the assertion. 
Passing next to the remainder of the axial skeleton, and with- 
out enumerating the differences that have been dwelt upon 
quite at length when under review, it is evident from the ar- 
rangement and number of the ribs that Hamatopus differs es- 
sentially from the several types we have been examining, and 
agrees only with Arenaria in the number of its free caudal 
vertebrae. If we were to be guided in this matter by the total 
number of vertebrae, as shown by these several species of birds, 
providing our count is correct, it would seem that in this respect 
Hcematopus, Arenaria, and Aphriza fall together, while some 
Plovers and some Sandpipers are thus associated. Hcematopns 
has a very different style of pelvis from the other species we 
have enumerated, while the pattern of this bone is not a little 
different when compared in Aphriza and Arenaria. Were we 
allowed the expression, it might be said that in so far as the 
pelvis goes in the genera Aphriza, Charadrius, Actitis, Are- 
naria, and some few others, the pluvialine design seems to be 
impressed with more or less strength upon them all. This 
statement will not apply to the pelvis in an Oyster-catcher. 
The sternum and shoulder girdle do not materially assist us 
in this matter; we found it with but two notches in such No. 2.] 
APHRIZA VIRGATA. 
337 
species as Actitis macularia and R. solitarius, and in the other 
forms it is of a markedly pluvialine stripe throughout. 
As to the appendicular skeleton, a marvellous sameness of 
style - barring the question of mere difference in size - seems 
to pertain to the skeleton of the limbs in Surf-birds, Turnstones, 
Sandpipers, Plovers, and others. Hamatopus, as we saw, pos- 
sesses a claw upon its pollex phalanx, while some of the species 
possess but three toes, as in the last-named genus and in Himan- 
topris and others. 
To sum up, then : my comparative studies of the osteology 
of Aphriza virgata lead me to believe that in the first place 
its affinity with Hamatopus is by no means a close one, the 
Oyster-catchers forming a very well-marked and distinct family 
of limicoline birds by themselves, showing by the structure of 
certain parts of their skeletons strong larine derivation. 
Secondly, the skull in Arenaria is in some respects more upon 
the plan of the skull in Hcematopus than it in any way ap- 
proaches the structure of that part of the skeleton in Aphriza; 
indeed, so far as the skulls are concerned, the skulls in Chara- 
drius, Arenaria, and Aphriza might be represented as each 
occupying the angle of an equilateral triangle, each sharing cer- 
tain characters in common and each removed from the other, 
equally distant by an equal set of others, not possessed by 
either of the other species. Were the skulls of Hcematopus and 
an average Tringa introduced into this hypothetical figure, the 
skull of the Oyster-catcher would be, as far as the characters 
they hold in common are concerned and for no other reason, 
on the line joining the skulls of the Turnstone and Surf-bird, 
and considerably nearer the first species than the last. The 
Sandpiper's skull would be on the line joining the skulls of 
Aphriza and the pluvialine type, and considerably nearer the 
former species than the latter. Aside from the skull, the re- 
mainder of the skeleton of Arenaria would answer fully for the 
skeleton of an equal-sized Plover, but hardly so for A. virgata. 
To conclude: the sum total of the skeletal characters of 
Aphriza virgata place this species nearer the Tringece-say some 
typical large-sized Tringa, with a four-notched sternum - than it 
does to the Plovers ; less so to Arenaria, and far less so to 
Hcematopus. Some pluvialine forms, however, are not so dis- 338 
SHUFELDT. 
tantly removed from Aphriza, if we may be allowed to judge 
from osteological premises. 
There is not the slightest doubt in my mind that had such a 
form as Charadrius squatarola been the sole representative of 
the Charadriidce in the entire world's avifauna, and been found 
taking the place, the habitat, of Aphriza, modern systematists 
of the class would surely have grouped it with the Turnstones 
as forming a family Aphrizid^e. We must believe that the 
fashion is far too prevalent, that when an isolated form like 
ApJtriza is met with, to lay about and endeavor to hit upon a 
family, a corner, where best to stow it away, without creating, 
as in the majority of cases should be done, a separate family 
for it. 
Now Aphriza is no more nearly related to Arenaria than is 
Charadrius related to Arenaria, and as at present classified by 
American Ornithologists, by no means indicates its true posi- 
tion in the system. I would propose then, in view of the ana- 
tomical facts brought out in the present memoir, that the family 
Aphrizidte be made to contain the sole representative of it, 
Aphriza virgata; and that another family, the Arenarid^e, be 
created to contain the Turnstones. 
It would seem that Aphriza in some way connects the Plovers 
with such Tringece as may possess the hallax and a four-notched 
sternum ; while Arenaria links the Plovers with Hcematopus, 
albeit that the Turnstone is much nearer the Plover than the 
far more remotely affined Oyster-catcher would in any way 
appear to be. 340 
SHUFELDT. 
EXPLANATION OF PLATE XXV. 
Note. - The figures in this Plate were all drawn by the author from the skeleton 
of a single individual, an adult male, and are all of natural size. 
Fig. i. The skull of A. virgata, seen directly from above; mandible removed. 
Fig. 2. The skull of A. virgata, seen upon basal aspect; mandible removed. 
Fig. 3. The skull of A. virgata, seen upon direct lateral view ; right side, includ- 
ing the mandible. 
Fig. 4. The mandible of A. virgata, direct superior view. 
Fig. 5. The right tarso-metatarsus of A. virgata, anterior view. 
Fig. 6. The right radius of A. virgata, palmar aspect. 
Fig. 7. The right ulna of A. virgata, palmar aspect. 
Fig. 8. The right fibula of A. virgata, inner side. 
Fig. 9. The right tibio-tarsus of A. virgata, mesial aspect. 
Fig. io. The right femur of A. virgata, anterior surface. 
Fig. 11. The os furcula of A. virgata, seen obliquely from the right side. 
Fig. 12. The sternum of A. virgata, seen upon right lateral aspect. 
Fig. 13. The right carpo-metacarpus of A. virgata, anconal surface; pollex pha- 
lanx in situ. 
Fig. 14. The right proximal phalanx of the index digit of A. virgata, anconal 
side. 
Fig. 15. The sternum of A. virgata, seen upon direct anterior view. 
Fig. 16. The left coracoid and scapula of A. virgata, outer aspect, and articula- 
ted in situ. 
Fig. 17. The right coracoid of A. virgata, seen upon direct anterior view. 
Fig. 18. The left humerus of A. virgata, palmar aspect. 
Fig. 19. The left humerus of A. virgata, anconal aspect. 
Fig. 20. The four last vertebrae of the skeleton of the tail of A. virgata, and the 
pygostyle, seen upon the right lateral aspect. 
Fig. 21. The pelvis of A. virgata, dorsal view. PL XXV. 
Mourn. Morph . VoL.lI. 
Ji W Shufeldt,nddel 
B. Meisel, Ji th Boston. 
Mjihriza virgata., ad&.xl.