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ANNALS
OF THE
CARNEGIE MUSEUM
Volume 31
1948-1951
Published by the Authority of the
Board of Trustees of the Carnegie Institute
Pittsburgh, Pennsylvania
Arranged for publication by
Arthur W. Henn
TABLE OF CONTENTS
Title Page . . . i
Table of Contents iii
List of genera, species, and forms new to science v
ARTICLE
1. Two new Passerine birds from Angola. By James P. Chapin
(Issued February 26, 1948) 1-4
2. Critical remarks on the Wood -hewers. By W. E. Clyde
Todd (Issued July 2, 1948) 5-18
3. A review of the Hydnaceae (Fungi) of Western Pennsyl-
vania. By LeRoy K. Henry (Issued July 30, 1948) 19-32
4. Critical remarks on the Oven-birds. By W. E. Clyde Todd
(Issued October 15, 1948) 33-44
5. Sphoenognathus curviceps (Coleoptera; Lucanidae), a new
species from Bolivia. By Bernard Benesh (Issued
November 3, 1948) 45-48
6. Taxonomy and distribution of the genus PiereUa (Lepi-
doptera). By F. Martin Brown (Issued December 8,
1948) 49-88
7. The land snails of West Virginia. By Gordon K. MacMillan
(Issued February 28, 1949) 89-238
8. A review of the pileate Thelephoraceae (Fungi) of Western
Pennsylvania. By LeRoy K. Henry (Issued Alay 4, 1949) . 239-256
9. Larval development of NectopJn'ynoides tornieri (Roux), with
comments on direct development in frogs. By Grace L.
Orton (Issued June 28, 1949) 257-276
10. Description of a new race of the salamander Pseiidohranchiis
striatus (Le Conte). By Coleman J. Goin and John W.
Crenshaw, Jr. (Issued November 21, 1949) 277-280
11. A review of the Elvellaceae (Fungi) of Western Pennsylvania.
By LeRoy K. Henry (Issued December 19, 1949) 281-288
iii
iwrs-igsf
12. A northern race of Red-tailed Hawk. By W. E. Ch-de Todd
(Issued April 25, 1950). 289-296
13. A new race of Paroquet of the species Aratinga astec from
the Republic of Honduras. By Arthur C. Twomey
(Issued April 28, 1950) 297-298
14. A study of the salamander, Ambystoma cingiilatiim, with the
description of a new subspecies. By Coleman J. Coin
(Issued September 15, 1950) 299-320
\A..
SkSA » 15-
,1
(
16.
17.
18.
An analysis of color and pattern in butterflies of the Asiatic
genus Karanasa. By Audrey Avinoff (Issued October
31, 1950) 321-332
A new race of Hudsonian Chickadee. By W. E. Clyde Todd
(Issued October 27, 1950) 333-334
A new geomyid rodent from the Miocene of Alontan^. By
Albert E. Wood (Issued December 21, 1950) 335-338
The mammals of the Alazinaw Lake region of Ontario; their
reproduction and population dynamics. By John J.
Christian (Issued February 19, 1951 ' 339-386
IV
LIST OF GENERA, SPECIES AND FORMS
NEW TO SCIENCE
Bradypterus mariae boultoni, subsp. nov. Aves. . .
Terpsiphone rufocinerea bannermani, subsp. nov. “ ...
Xiphorhynchus eytoni vicinalis, subsp. nov. “ ...
“ guttatus connectens, subsp. nov. “ ...
“ triangularis distinctus, subsp. nov. “ ...
“ pardalotus caurensis, subsp. nov. “ ...
Dendroplex picus rufescens, subsp. nov. “ . . .
Campylorhamphus procurvoides successor, subsp. nov. “ ...
Glyphorynchus spirurus subrufescens, subsp. nov. “ ...
Sittasomus griseicapillus viridior, subsp. nov. “ ...
“ amazonus sordidus, subsp. nov. “ ...
Dendrocincla merula obidensis, subsp. nov. “ ...
“ meruloides neglecta, subsp. nov. “ ...
Synallaxis albescens littoralis, subsp. nov. “ . . .
“ 'griseonota, subsp. nov. “ ...
“ » cabanisi obscurior, subsp. nov. “ ...
Certhiaxis cinnamomea albescentior, subsp. nov. “ ...
Philydor erythrocercus suboles, subsp. nov. “ ...
Automolus infuscatus.purusianus, subsp. nov. “ ...
Sclerurus rufigularis furfurosus, subsp. nov. “ ...
“ “ brunnescens, subsp. nov. “ ...
Sphoenognathus curvipes, sp. nov. Coleoptera
Pierella hyalinus fusimaculata, subsp. nov. Lepidoptera
“ lena obsoleta forma nova “
“ stollei boliviana, subsp. nov. “
“ luna rubra forma nova “
“ hortona albopunctata forma nova “
“ albofaciata decepta, subsp. nov. “
Pseudobranchus striatus spheniscus, subsp. nov. Amphibia
Buteo jamaicensis abieticola, subsp. nov. Aves
Aratinga astec melloni, subsp. novc , “
Ambystoma cingulatum bishopi, subsp. nov. Amphibia
Penthestes hudsonicus labradorius, subsp. nov. Aves
Gregorymys kayi, sp. nov. ’ Rodentia
1
3
7
8
8
9
11
13
14
15
15
16
16
36
37
37
38
39
40
41
42
45
64
67
74
79
81
83
277
291
297
300
334
335
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ART. 1. TWO NEW PASSERINE BIRDS FROM ANGOLA
By James P. Chapin
The American Museum of Natural History
During the years when Mr. Rudyerd Boulton was devoting special
attention to the birds of Angola, he made several valuable collections
there, for the American Museum of Natural History and for the Carnegie
Museum. A number of the new forms he discovered were described by
Mr. Boulton, but two more which deserved new names still remained in
the Carnegie Museum collection. At the request of Mr. W. E. Clyde Todd,
I have prepared the following diagnoses.
The Representative of Bradypterus mariae in Angola.
In the revisions of the genus Bradypterus by Mackworth-Praed and
Grant {This, 1941, pp. 441-455) and by J. Delacour {Ihis, 1943, pp. 31-40),
there is no mention of any race of B. marice or B. cinnamomeus from the
highlands of Angola. Yet the Carnegie Museum has long had three skins
from that region which clearly represent Bradypterus marice. Two of
them are immature birds, retaining a yellowish wash on the underparts,
from 6,200-6,300 feet on Mount Moco, February, 1931 ; and the other is an
adult female from Mombolo, around 5,000 feet, taken on March 7, 1931.
All were collected by Rudyerd Boulton.
It might seem unwise to describe a new race with but one adult speci-
men, and I do it only because the Angola highland is so far removed from
any district where a race of B. marice has hitherto been known to live.
The new race is named in honor of my friend Rudyerd Boulton.
Bradypterus marise boultoni, subsp. nov.
Type, No. 109,511, Collection Carnegie Museum, adult female; Mom-
bolo, in the northwestern part of the Benguela highland, March 7, 1931;
Rudyerd Boulton.
Suh specific characters. — Rather similar to Bradypterus marice usamharce
Reichenow of eastern Africa, but a little lighter brown on crown, back,
and rump, somewhat lighter beneath, and more tinged with buff on breast
and flanks. Somewhat larger than B. m. cameruneitsis Alexander of
Mount Cameroon.
Measurements. — The type has the wing 62 mm. ; tail 61 ; culmen to base
15.5; tarsus 25. Two immature females from Alount Moco have wings
63, 64; tails 59, 60; culmen to base 15, 16; tarsi 23, 24.5.
1
sued February 26, 1948.
2
Annals of the Carnegie Museum
VOL. 31
Range. — This probably includes many more districts of Angola in ad-
dition to Mombolo and Mount Moco, at elevations above 5,000 feet,
where suitable scrubby woods exist.
Remarks. — Both young individuals have 10 rectrices, but the tail of
the type is incomplete. There is good reason to suppose that the Bradypter-
us marice group is conspecific with B. barratti Sharpe of South Africa,
and the one specimen of barratti I have recently examined also had a tail
of only 10 quills.
A New Race of Terpsiphone from Northwestern Angola.
The Lower Congo district and the adjacent Loango Coast are the home
of a Paradise Flycatcher with rather short crown-feathers, blackish head,
underparts more grayish, and back, wings, and tail rufous, the median
rectrices lengthened in adult males. It w^as named Terpsiphone rufocinerea
by Cabanis in 1875, the type locality being Chinchoxo, a little north of
Landana.^
In 1857, Verreaux had described Tchitrea melampyra from the Gaboon,^
and that name has often been supposed to antedate rufocinerea. But in
1938, Professor Erwin Stresemann informed me that he had examined
the type of melampyra in the British Museum and found it to be a dull-
colored example of some form of T. viridis.
Terpsiphone rufocinerea batesi ChapiiP is a well-marked race living in
the forests of the Upper Congo and Cameroon, lighter and grayer on head
and breast, with median rectrices never much longer than the rest of the
tail. In 1921, I examined four specimens of nominate rufocinerea from
Chinchoxo in the Berlin Museum, and compared with them an adult
male I had myself collected at Boma on the lower Congo River. The
range of this race seems to extend northward along the coast to the base
of Mount Cameroon, and examples of intermediate nature are not un-
common in forested southern Cameroon.
To the southward, rufocinerea has been reported as extending to Ndala
Tandoand even Novo Redondo in Angola. Doctor D. A. Bannerman, in
1 Terpsiphone rufocinerea Cabanis, 1875, Jour. Ornith., p. 236 (Chinchoxo,
Loango Coast).
^ Tchitrea melampyra Verreaux, 1857, in Hartlaub, System der Ornithologie
Westafrica’s, p. 90 (Gaboon).
^ Terpsiphone batesi Chapin, 1921, Amer. Mus. Novitates, no. 7, p. 6, fig. 3
(Medje, northeast Belgian Congo).
1948
Chapin: New Passerine Birds from Angola
3
1936,“^ discussed the possibility that specimens from Ndala Tando were
different from those of the Loango Coast, but he regarded the name
rufocinerea as a synonym of melampyra, and came to no final conclusion.
In March, 1931, at Ngara, some 30 kilometers northeast of Novo
Redondo, Rudyerd Boulton collected two adult males which are plainly
conspecific with T. rufocinerea, but distinctly grayer on head and throat
than the nominate form, although retaining the bright rufous under tail-
coverts. At the same locality, on September 29, 1939, he had also secured
an adult male of T. viridis plumbeiceps. This does not prove that both
birds nest there, for plumbeiceps is known to be migratory between April
and October.
The two males of T. rufocinerea, taken in March, were noted as ready
to breed, and they show that an unnamed race of that species does live in
northwestern Angola. Doctor Bannerman found that eight specimens
from Ndala Tando were lighter in color than all but one of those he had
seen from farther north; and in the Rothschild Collection there is a female
from Canhoca, Angola, which plainly represents the race I propose to
name in honor of my friend David Bannerman.
Terpsiphone rufocinerea bannermani, subsp. nov.
Type, No. 109,533, Collection Carnegie Museum, adult male; Ngara,
northwestern Angola, Alarch 10, 1931; Rudyerd Boulton.
Subspecific characters. — Similar to Terpsiphone r .rufocinerea Cabanis of
the Loango Coast, but with head, throat, and breast lighter and grayer,
much less glossy on throat. There is a slight crest in bannermani, and the
whole crown is very dark gray, glossed with blue-green. The longest
feathers in the crest measure 14-14.5 mm. The rufous coloration of back
and tail is usually a little lighter than in nominate rufocinerea, the rufous
of under tail-coverts not quite so deep. The under wing-coverts are whit-
ish, washed with pale rufous, and becoming more rufous toward the outer
edge of the wing.
Measurements. — The type has the wing 84 mm.; long median tail-
feathers 252, next longest pair 122, and outermost rectrices 79 mm.; cul-
men to base 19.5; tarsus 15.5. The second male from Ngara has wing 84;
median rectrices 227, next pair 94, outermost 75; culmen to base 19;
tarsus 16. The female from Canhoca, wing 79; tail 84; culmen to base 18;
tarsus 15.5.
^ D. A. Bannerman, 1936, The Birds of Tropical West Africa, vol. 4, p. 301.
4
Annals of the Carnegie Museum
VOL. 31
Range. — Northwestern Angola, from the vicinity of Novo Redondo at
least to Ndala Tando, probably to be found only in the more heavily
wooded situations.
Remarks. — In this same region Terpsiphone viridis plumbeiceps is
widely distributed; but it is readily distinguished by its better-developed
crest, with feathers measuring 15 to 19 mm. in males, and by its pale under
tail-coverts, which are either whitish or tinged only lightly with rufous.
In more ways than one, this new race, hannermani, is intermediate
between T. r. rufocinerea and T. v. plumbeiceps. Yet it would be difficult
to treat all three as conspecific, because from the interior of southern
Cameroon to the forested Upper Congo T. rufocinerea batesi lives side by
side with T. viridis speciosa, the former keeping to the heavy shade of
primary forest, while the latter haunts second growth and leafy trees in
clearings. A similar difference in haunts may well prevail in Angola, espe-
cially during the breeding season.
Oiro
ART. 2. CRITICAL REMARKS ON THE WOOD-HEWERS
' By W. E. Clyde Todd
The Wood-hewers or Tree-creepers (Dendrocolaptidae) are represented
in the collection of the Carnegie Museum by 2,544 specimens,^ belonging
to 155 species and subspecies. The Wood-hewers are wholly neotropical
ij in their distribution, and their center of abundance is in the equatorial
I region of South America. Studies made some years ago resulted in the
[ description of eight new forms of this family, and a more recent critical
i study has revealed the existence of eleven additional subspecies which
I appear to be new to science. In connection with the descriptions of these
1 new forms there are included herewith some taxonomic discussion of other
I known forms which seem to be in need of further elucidation. Certain of
I . .
these studies were made possible only through the loan of specimens from
other collections. I am deeply grateful to Mr. John T. Zimmer of the
x\merican Museum of Natural History and to Dr. Herbert Friedmann of
the U. S. National Museum for the loan of key specimens of Wood-
hewers from the collections of their respective institutions,
j All measurements are in millimeters, and that of the bill is for the ex-
posed culmen. The names of colors are taken mostly from Ridgway’s
Ij “Color Standards and Color Nomenclature.” Most of the localities ap-
j pearing in the present paper were listed by the author in an earlier paper,
I on Hummingbirds (Annals Carnegie Museum, Vol. 29, 1942, pp. 365-370).
I
j Xiphocolaptes orenocensis paraensis Pinto
j Three specimens: Santarem, Colonia do Mojuy, and Apacy, Brazil,
jl This form was described by Sr. Oliveira Pinto (Papeis Avulsos De-
I partmento de Zoologia, etc., Sao Paulo, 5, No. 15, 1945, 135) as a race of
Xiphocolaptes promeropirhynchus, which according to Hellmayr’s views
is not conspecific with X. orenocensis. With this disposition I agree, after
examination of the several forms involved. Pinto’s type-series (all but
one specimen) came from the right bank of the Rio Madeira, but his de-
scription applies well enough to our birds from the Rio Tapajoz. They
resemble Xiphocolaptes orenocensis berlepschi Snethlage of the Rio Purus
and upper Amazon, but the pileum is more blackish, less brownish, with
I 1 This number includes 249 specimens alienated from the collection by ex-
change.
I
jlssued July 2, 1948.
JULS /- 'k
6
Annals of the Carnegie Museum
VOL. 31
narrower shaft-streaks, and the bill is much darker-colored. According
to the collector’s notes, the maxilla in paraensis is “blackish,” and the
mandible is “intensely dark horn gray with blackish tip.” In the Rio
Purus specimens, however, the maxilla is “dark horn gray,” and the mandi-
ble is “light horn gray with a faint bluish pearly gloss on the basal half.”
Hellmayr (1925, 285) also mentions these differences, as well as Griscom
and Greenway (1941, 210), but they made no formal separation.
Compared with two specimens of true orenocensis (American Museum),
our Rio Tapajoz birds have the bill darker-colored, while the striping on
the underparts is much heavier, and the pileum is blacker. One of our
specimens is more rufescent than the other two.
Xiphocolaptes orenocensis orenocensis von Berlepsch and Hartert.
Except for its deeper coloration, a single specimen from Manacapuru
agrees well enough with two Orinoco skins, and should doubtless be re-
ferred to orenocensis, the range of which will have to be extended accord-
ingly.
Dendrocolaptes certhia ridgwayi Hellmayr.
Messrs. Griscom and Greenway (1941, 208-9) remark that this series
“supports Zimmer’s comments [1934b, pp. 2-3] on the variability of this
species.” They add that “three Santarem birds are the concolor type,”
but I find myself unable to pick them out, and I unhesitatingly refer all
our fifteen specimens from the east bank of the Rio Tapajoz to one and
the same form, which, as these authors show, must bear the name ridgwayi.
Dendrocolaptes certhia juruanus von Ihering.
Twenty-three specimens: Hyutanahan, Nova Olinda, Arima, Sao
Paulo de Olivenga, and Caviana.
Caviana birds are almost certainly subspecifically separable by reason
of their paler coloration and narrower barring, but inasmuch as they are
closely approached in these respects by certain of the Rio Purus series I
do not care to take this step with only two specimens available. Birds
from Sao Paulo de Olivenga, on the other hand, in their heavier barring
and their tendency to more rufescent coloration, approach polyzonus of
Bolivia. If we discard the supposed record from Marabitanas on the Rio
Negro, as suggested by Zimmer (1934b, 1), juruanus emerges as a race
confined to the region south of the middle and upper Amazon.
1948
Todd: Critical Remarks on Wood-Hewers
7
Dendrocolaptes concolor von Pelzeln.
Nine specimens: Villa Braga and Itaituba, Brazil.
To my eye these birds are perfectly distinct from the east-bank birds,
for which the name ridgwayi will have to be restored, as already said. The
very fact that the two forms behave as they do, as described by Zimmer
(1934b, 2-3), indicates that they are really distinct species. I find no
specimens in our series which cannot certainly be referred to one or to
the other.
Xiphorhynchus eytoni vicinalis, subsp. nov.
Type, No. 78,061, Collection Carnegie Museum, adult male; Apacy,
Rio Tapajoz, Brazil, April 30, 1920; Samuel M. Klages.
Subspecific characters. — Similar to Xiphorhynchus eytoni eytoni (Sclater)
of the Para region of Brazil, but underparts with more buffy suffusion,
and the abdomen usually tinged with clay-color; streaking of the upper-
and underparts averaging heavier.
Range. — Rio Tapajbz region of Brazil, and eastward for an undeter-
mined distance.
Remarks. — Mr. Zimmer (1934c, 1-4) has fully discussed the variations
in the series of this and related forms examined by him. In spite of his
dictum that eytoni is best left undivided, I agree with Griscom and
Greenway (1941, 212) in believing that another form should be recognized.
They call attention to the fact that Zimmer had no topotypes of eytoni.
Certainly there is a definite line of demarcation between our twelve
Benevides birds on the one hand and those from the Rio Tapajoz (twenty-
five) on the other, as above indicated. I have examined also ten speci-
mens (from Villa Bella Imperatriz, Borba, and Faro) which Zimmer calls
eytoni, but which are not eytoni at ail, but the new form I am describing
herewith. Some of them come from the same places as undoubted speci-
mens of guttatoides, from which they can at once be distinguished by the
differently colored bill. X. guttatus guttatoides, be it noted, has a pale bill,
while eytoni and the new form both have dark-colored bills (at least the
maxilla is dark). X. g. guttatoides and the present form may be subspecies,
separated ecologically, as Zimmer suggests, but would not this be a new
kind of subspecies? I am inclined under the circumstances to rank them
as species. Two specimens from Faro show that this new form ranges
north of the Amazon.
8
Annals of the Carnegie Museum
VOL. 31
Xiphorhynchus guttatus connectens, subsp. nov.
Type, No. 83,523, Collection Carnegie Museum, adult male; Obidos,
Brazil, January 29, 1921; Samuel M. Klages.
Subspecific characters. — Most nearly resembling Xiphorhynchus gut-
tatus polystictus (Salvin and Godman) of the Guianas and eastern Vene-
zuela, but the underparts are more deeply suffused with buff, and the light
stripes on the pileum, nape, and upperparts average heavier, and deeper
buff. (Thirteen specimens examined.)
Range. — Known only from the type-locality, but probably ranging
westward to the Rio Negro and eastward for an indefinite distance, on the
north bank of the Amazon.
Remarks. — Messrs. Griscom and Greenway, who handled these speci-
mens, referred them without comment to polystictus, but more careful
comparison shows that they differ appreciably from that form, as it is
represented by a series from Venezuela and French Guiana, in the respects
pointed out above. They form a connecting link, in fact, between poly-
stictus and guttatoides, and come from the region between the respective
ranges of these forms. As a rule, however, French Guiana forms usually
range unchanged to the Amazon at Obidos.
Xiphorhynchus triangularis triangularis (Lafresnaye).
Of our series of twenty-seven specimens, all but four come from Vene-
zuela, and should therefore represent hylodromus Wetmore (1939, 2).
Even with his type-series before me, I have great difficulty making out
this form. Four specimens from Las Ventanas, Santander, Colombia,
which may be considered to represent true triangularis, are scarcely
separable, in my opinion. The light spotting of the underparts is perhaps
a little less dense, but the difference is very slight, and I would question
the validity of hylodromus. However, in the Western Andes of Colombia
there lives a population about whose distinctness there can be no question.
Let it be called, therefore,
Xiphorhynchus triangularis distinctus, subsp. nov.
Ten specimens: Heights of Caldas, Bitaco Valley, and La Cumbre,
Colombia.
Type, No. 67,543, Collection Carnegie Museum, adult male; Bitaco
Valley, Colombia, July 6, 1918; M. A. Carriker, Jr.
Subspecific characters. — Similar to Xiphorhynchus triangularis triangu-
1948
Todd: Critical Remarks on Wood-Hewers
9
laris (Lafresnaye) of the Eastern Andes of Colombia, but general colora-
tion darker; upperparts between orange citrine and medal bronze (in-
stead of citrine) ; rump and upper tail-coverts heavily shaded with argus
brown; wings externally much darker (more rufescent, less olivaceous);
tail bay (instead of auburn) ; underparts darker olive, the throat deeper
buffy; and under tail-coverts less rufescent.
Range. — Subtropical Zone, Western (and Central ?) Andes of Colom-
bia.
Remarks. — Chapman had some specimens of this species from the
Western Andes, but could “detect practically no racial variation,” except
that birds from the “Central and Western Andes may average slightly
more rufescent above” (1917, 420). But with smoothly and uniformly
made-up skins available for comparison, the difference between birds from
the Eastern Andes on the one hand and those from the Western Andes
on the other is obvious, and it is too great to ignore, as Hellmayr and other
authors have done.
The range of this form approximates that of cequatorialis of the lower
levels (Tropical Zone) in this region, but there is no reason to suppose
that they intergrade, and I consider them specifically distinct.
Xiphorhynchus pardalotus caurensis, subsp. nov.
Sixteen specimens: La Lajita, Rio Mocho, El Llagual, and Rio Yuruan,
Venezuela.
Type, No. 32,306, Collection Carnegie Museum, adult male; Rio Mocho,
Rio Caura, Venezuela, November 23, 1909; M. A. Carriker, Jr.
Subspecific characters. — Similar to Xiphorhynchus pardalotus pardalotus
(Vieillot) of French Guiana, but general color of upperparts more rufes-
cent, less olivaceous, and buffy streaks of upper- and underparts nar-
rower and less pronounced.
Range.- — Southern Venezuela, from the Rio Caura east to the Guiana
frontier.
Remarks. — It is not strange that the geographical variation in this
species has been overlooked, since satisfactory series for comparison are
wanting in most collections. But with the specimens above listed, and a
fine series (thirty-two skins) from French Guiana as well, it is at once
obvious that they represent two different races. In the nominate form the
upperparts (in fresh plumage) may be described as medal bronze of Ridg-
way, while in the Venezuelan form it is nearer Dresden brown. The
underparts differ in a corresponding manner; the streaking on both the
10
Annals of the Carnegie Museum
VOL. 31
upper- and underparts differs as aforesaid; it is less strongly marked in
the Venezuelan race.
Xiphorhynchus elegans ornatus Zimmer.
Four specimens: Sao Paulo de Olivenga, Brazil.
These were handled and listed by Zimmer (1934c, 7-10). They belong
to a form closely allied to elegans, but in my opinion not conspecific with
spixii, which form occurs, however, on the east bank of the Rio Tapajoz.
That ornatus is not conspecific 'with juruanus, as it is given by Zimmer, is
proved by the fact that they both occur at the same place — Sao Paulo de
Olivenga. Zimmer actually lists them thus, but suggests that one of them
must have been taken “near?” that place. This is not the case. Speci-
mens of both forms were actually taken on the same date, and Mr. Klages
assures me that they came from precisely the same collecting-station.
This circumstance definitely disposes of the case of ornatus and juruanus,
and by implication that of elegans and spixii also. Even Hellmayr, who
always tried to unite related forms as conspecies wherever possible, could
not see his way clear to do so in these cases, and neither can I.
Xiphorhynchus obsoletus multiguttatus (Lafresnaye).
Twenty-three specimens: Hyutanahan, Nova Olinda, and Arima,
Brazil.
Rio Purus specimens are referred to typical obsoletus by Count Gylden-
stolpe (1945, 141), but I do not agree to this disposition. Since our ten
specimens from Sao Paulo de Olivenga and Tonantins compare favorably
with Ecuadorean specimens of X. obsoletus palliatus, I would refer the
Rio Purus birds to multiguttatus, described from Fonteboa. The two races
are easily separable in series from each other, and from obsoletus.
Xiphorhynchus necopinus (Zimmer).
Of this species we have thirty-one specimens, from the following
localities in Brazil: Santarem, Villa Braga (Rio Tapajoz), Hyutanahan,
Nova Olinda, Arima (Rio Purus), Sao Paulo de Olivenga, Caviana,
Islands near Manacapuru, and Rio Manacapuru (Rio Solimoes). In addi-
tion I have examined six specimens from the American Museum collection
which were listed by the original describer. Our specimens indicate that the
range of this form is actually more extensive than he suspected. Moreover,
I am convinced that the resemblance, close as it is, between necopinus and
Dendroplex picus is superficial and fortuitous, so to speak, and not indica-
1948
Todd: Critical Remarks on Wood-Hewers
11
tive of congeneric affinity. Zimmer suggested that it might become neces-
sary to submerge Dendroplex under Xiphorhynchus. I propose instead to
retain Dendroplex for D. picus and its allies, and to transfer necopinus to
Xiphorhynchus, where it undoubtedly belongs. Santarem specimens of
necopinus appear to be slightly smaller and duller than the others; how-
ever, I do not venture to separate them. We have two young birds (April
2), with dark-colored bills.
Dendroplex picus rufescens, subsp. nov.
Sixteen specimens: Santarem, Villa Braga, Itaituba, and Apacy, Brazil.
Type, No. 75,252, Collection Carnegie Museum, adult male; Villa
Braga, Rio Tapajoz, Brazil, December 1, 1919; Samuel M. Klages.
Subspecific characters. — Similar to Dendroplex picus picus (Gmelin) of
the Guianas and adjacent parts of Brazil, but underparts more rufescent,
less brownish, and light streaks on upper- and underparts wider and more
extended posteriorly. Similar also to D. p. kienerii (Des Murs) of the
middle Amazon, Rio Purus, etc., but averaging less rufescent below and
rather more rufescent above; markings on the underparts finer and less
extensive; size smaller. Wing (type), 101; tail, 86; bill, 30.
Range. — Certainly known only from the Rio Tapajoz (both banks),
but probably ranging eastward to the island of Marajo and westward to-
wards the Rio Madeira.
Remarks. — With the series before me I have no alternative but to
describe this race, although neither Zimmer nor Griscom and Greenway
considered it tenable. The characters it shows are in part those given by
Zimmer for his race duidce, but direct comparison with the latter elimi-
nates the possibility of identity therewith. The general color of the pos-
terior underparts in the present form is tawny olive; in true picus they are
considerably duller and more brownish in tone (near Saccardo’s umber).
In the color of the underparts in general this race is intermediate between
picus on the one hand and kienerii on the other. There is some variation
it is true, but the general difference is too great to overlook. In any event,
the form is as readily recognizable as are the races of Xiphorhynchus
ohsoletus. The exact limits of its range, however, must be determined from
examination of specimens. In this connection it may be noted that speci-
mens from Caviana (opposite Manacapuru on the middle Amazon) are
clearly referable to kienerii. From this we may infer that rufescens does
not reach the Rio Madeira on the west; the form occurring on that
stream is probably kienerii.
12
Annals of the Carnegie Museum
VOL. 31
Dendroplex picus dugandi Wetmore and Phelps.
I had identified all our specimens of Dendroplex from the Santa Marta
region of Colombia as D. picirostris, but without seeing the type or topo-
typesof that form. It was no surprise, therefore, when Messrs. Wetmore
and Phelps described the bird of northern Colombia as a new race,
dugandi. The surprise came when I discovered that while they considered
specimens from the southwestern part of the Santa Marta region to be
dugandi, they referred our birds from Bonda, Mamatoco, etc., to pici-
rostris. I have again gone over our series very carefully, and can find no
justification for dividing it up in such fashion. As I see it, all birds from
the Santa Marta region proper belong to one and the same race, which
I would call dugandi. They have been compared directly with eight speci-
mens of picirostris from Riohacha (the type-locality) and vicinity, in the
Goajira Peninsula. The difference is striking, and dugandi is an excellent
race, but the describers failed to state that their comparisons were made
with topotypical material. In all our specimens the white of the under-
parts is much more restricted than in picirostris; the bill, too, is shorter.
Two specimens from Dibulla show intermediate tendencies.
Lepidocolaptes layardi duidae Zimmer.
One specimen from Tonantins, Rio Solimoes, belongs to this race,
which is known to reach the upper Rio Negro. The specimen in question
compares favorably with six of duidcc in the American Museum.
Lepidocolaptes layardi madeiras (Chapman).
Four specimens: Santarem, Villa Braga, Miritituba, and Arima, Brazil.
Hellmayr handled these specimens some years ago, and noted his
identifications on the labels; they are duly listed by him (1925, 327).
Griscom and Greenway, however (1941, 214), set his work aside, and
argued that the Rio Tapajoz must be the demarcation line between the
respective ranges of layardi and madeircc. The specimens before me do
not at all support this view. They show that madeircc is found on both
banks of the river, and not just on the west bank alone. These specimens
agree with each other, and differ from five skins of typical layardi from
Benevides (near Para) in just the way that Chapman claims. Incidentally,
I am of the opinion that layardi should be kept specifically separate from
both fuscicapillus and alholineatus, from both of which it differs by valid
characters.
1948
Todd: Critical Remarks on Wood-Hewers
13
Lepidocolaptes angustirostris certhiolus (Todd).
In describing this form some years ago I compared it with hivittatus,
from which it differs as I said. It is actually a form intermediate between
hivittatus and angustirostris — rather nearer the latter, but lighter rufous
above, and not so heavily streaked below. The indications are that it has a
rather restricted range — a consideration that may account for the
difficulty some authors find in recognizing it.
Campylorhamphus procurvoides successor, subsp. nov.
Nineteen specimens: Islands near Obidos, Hyutanahan, Nova Olinda,
Arima, Sao Paulo de Olivenga, and Caviana (opposite Manacapuru),
Brazil.
Type, No. 92,371, Collection Carnegie Museum, adult male; Nova
Olinda, Rio Purus, Brazil, August 3, 1922; Samuel M. Klages.
Subspecific characters. — Similar to Campylorhamphus procurvoides
probatus Zimmer of the Rio Tapajoz region of Brazil, but general colora-
tion more rufescent, less brownish, and pale streaking of the upper- and
underparts heavier. Similar also to C. procurvoides multostriatus (Sneth-
lage), but general coloration more rufescent, pale streaking less pro-
nounced, and pileum brownish rather than blackish.
Ra7ige. — Brazil, south of the Amazon, and west of the Rio Madeira;
reappearing on the islands near Obidos.
Remarks. — This is the undescribed race referred to by Zimmer (1934a,
11) in discussing a single specimen from Teffe. This specimen, courteously
sent me for examination, fits right in with the above series. Count Gylden-
stolpe (1945, 142) also had but a single specimen from the Rio Jurua,
which he was unable to place. With the good series of specimens now
available it is obvious that they represent a race which is fully as well
differentiated as any of its conspecies. Much to my surprise I find that
six specimens from islands in the Amazon near Obidos belong to this same
form, although we have one specimen of true procurvoides from Obidos
itself.
Glyphorynchus spirurus, subsp.?
Four specimens: Manacapuru and Rio Manacapuru.
These resemble G. spirurus castelnaudii, but are paler above and rather
more extensively streaked below. They do not exactly fit in any of the
14
Annals of the Carnegie Museum
VOL. 31
described races. I sent them to Mr. John T. Zimmer for his opinion, and
he writes as follows:
“The Glyphorynchus offer a problem. In the abdominal streaking they
are most like spirurus and some of my specimens from Faro are very like
them in other respects, although the average Faro bird agrees too well with
the Guianan to suggest any real separation there. Your birds undoubtedly
are paler-backed than almost all the series of spirurus.
“I have extremes of castelnaudii with about as much streaking on the
belly as your 98,522 and the lighest extreme of that form at hand is lighter
than your darkest specimen, but again the average is darker than in your
birds. Your 105,489 and 98,522 could be lost in the series of castelnaudii,
but the other two would stand out by reason of their ventral streaks, not
by other characters.
“I would expect my birds from Muirapinima (west bank of the lower
Negro) to agree with your Manacapurii birds, but two of them are darker
above and one of these darker ones is the only one of the four that has any
noticeable belly streaks and not very much at that. On the whole they
probably are best kept in rufigularis, where I first placed them.
“It looks possible, therefore, that the Manacapuru population is not
to be matched exactly by any of the named forms, unless you have more
from there less well marked than these.”
However, I do not venture to describe the Manacapuru bird at this
time; it would only complicate the definition of the ranges of the allied
forms. In this connection I might call attention to G. spirurus inornatus,
described by Zimmer and restricted by him to the west bank of the Rio
Tapajoz. We have four skins of this race from the west bank of this stream
(Villa Braga, Apacy), but these can be matched by others from the east
bank. To my eye the Rio Tapajoz series of Glyphorynchus cannot be
divided on this basis at all; it is all one race. It was compared primarily
with castelnaudii, from which it differs as was said. As a matter of fact,
however, it is much more closely related to cuneatus, from which it differs
only in the slightly warmer brown shade of the underparts, apparent in
series.
Glyphorynchus spirurus subrufescens, subsp. nov.
Twelve specimens: Murindo, El Tambo, Andagoya, Potedo, and
Malagita, Colombia.
Type, No. 66,470, Collection Carnegie Museum, adult male; Poted6,
Choco, Colombia, May 2, 1918; M. A. Carriker, Jr.
1948
Todd: Critical Remarks on Wood-Hewers
15
Subspecific characters. — Similar to Glyphorynchus spirurus sublestus
Peters, but smaller; general coloration more rufescent, less brownish; and
streaked area on breast less extensive.
Range. — Valleys of the Rio Atrato (except lower part) and of the Rio
San Juan, western Colombia.
Remarks. — These birds would be referred to sublestus by Peters (1929,
443) and by Zimmer (1934d, 5), but they certainly differ from a specimen
from the lower Atrato (Soatata), which in its turn closely resembles a
series examined from eastern Panama (American Museum). The pe-
culiarities of the western Colombia bird would scarcely be remarked unless
a series was available — a circumstance which probably accounts for the
fact that thus far they have gone undetected. The richer general colora-
tion of this population and the restriction of the breast markings are
obvious when series are compared. Seven adult males average: wing,
68 mm. ; tail, 62.7 ; bill, 12. Ten males of sublestus from Panama and Costa
Rica: wing, 74; tail, 70; bill, 13.4. Costa Rican birds may be regarded
as intermediate between sublestus and pectoralis. Birds from the region
around the Gulf of Maracaibo in Venezuela may be separable as still
another subspecies.
Sittasomus griseicapillus viridior, subsp. nov.
Six specimens: Palmarito and Rio Quiser, Bolivia.
Type, No. 80,141, Collection Carnegie Museum, adult male; Palmarito,
Rio San Julian, Chiquitos, Bolivia, May 22, 1918; Jose Steinbach.
Subspecific characters. — Similar to Sittasomus griseicapillus viridis Car-
riker of the montane region of Bolivia, but general coloration much lighter,
more yellowish green, and the remiges more rufescent.
Range. — Chiquitos region of eastern Bolivia.
Remarks. — This remarkable new race adds another to those hailing from
this particular region. It is well marked, and is perfectly distinct from
viridis, of which we have nine specimens from various localities in the
mountains and foothills of Bolivia. The upperparts are buffy citrine, and
the underparts are similar but paler; the primaries externally are Sudan
brown, and the secondaries are Sanford’s brown. All the characters of
viridis are carried a step further in the present race.
Sittasomus amazonus sordidus, subsp. nov.
Five specimens: El Llagual and Rio Yuruan, Venezuela.
Type, No. 33,807, Collection Carnegie Museum, adult male; Rio Yuruan,
Venezuela, March 25, 1910; M. A. Carriker, Jr.
16
Annals of the Carnegie Museum
VOL. 31
Suhspecific characters. — Similar to Sittasomus amazonus amazonus
Lafresnaye, but somewhat smaller, and general coloration duller; upper-
parts with a wash of antique brown; primaries edged externally with
Dresden brown (instead of buckthorn brown or dull grayish brown) ;
underparts (near) deep olive gray (instead of deep grayish olive) ; under
tail-coverts amber brown (instead of ochraceous tawny). Wing (type),
82; tail, 69; bill, 16; tarsus, 17.
Range. — Venezuela, from the Rio Caura east to British Guiana.
Remarks. — The characters of this race, although perfectly obvious
upon comparison, are not easy to formulate. Hellmayr (1925, 259) re-
ferred birds from Venezuela to amazonus without hesitation, but our series
from that region, although small, is definitely distinct. The underparts
almost entirely lack the greenish shade so characteristic of amazonus;
instead they are dingy grayish brown. Comparison has been made with
specimens from the upper Amazon, which are believed to correctly repre-
sent amazonus (described from Peru).
Dendrocincla merula obidensis, subsp. nov.
Type, No. 83,250, Collection Carnegie Museum, adult female; Obidos,
Brazil, December 31, 1930; Samuel M. Klages.
Suhspecific characters. — Similar to Dendrocincla merula merula of
French Guiana, but larger, and chin-spot paler buffy.
Measurements.— One male: wing, 108; tail, 76; bill, 25; tarsus, 25. Two
females: wing, 102, 113; tail, 78, 80; bill, 24.5, 28; tarsus, 25.
Range. — Brazil, north of the Amazon (Obidos; Faro).
Remarks. — With only one specimen at the time, Zimmer (1934a, 15)
was unwilling to describe this race, but now that we have two more (both
from Obidos), closely matching the first specimen, there can remain no
good reason for refusing to recognize it by name. It is very distinct by
reason of its larger size. The wings of French Guiana specimens of true
merula run 92, 99, 100; the tails, 65, 66, 67. Except for the slightly paler
chin-spot, there is no difference in coloration.
Dendrocincla meruloides neglecta, subsp. nov.
Twenty-one specimens: Nova Olinda, Arima, Sao Paulo de Olivenga,
Tonantins, opposite Tonantins, Manacapuru, and Rio Manacapuru,
Brazil.
Type, No. 96,388, Collection Carnegie Museum, adult male; Sao Paulo
de Olivenga, Brazil, April 10, 1923; Samuel M. Klages.
1948
Todd: Critical Remarks on Wood-Hewers
17
Subspecific characters. — Similar in general to Dendrocincla meruloides
phceochroa von Berlepsch and Hartert of the Orinoco Valley, Venezuela,
but decidedly larger; upperparts slightly darker, more olivaceous; and
underparts slightly paler (less rufescent) brown.
Measurements. — Ten males: wing, 108-114 (average, 111.6); tail, 86-96
(91.5); bill, 28-31 (30.2); tarsus, 22.5-27 (25). Eight males of D. m.
phceochroa: wing, 100-110 (103); tail, 78-95 (84); bill, 27-30 (28); tarsus,
22-25 (23.5).
Range. — Valley of the Amazon (both banks), above the Rio Negro,
and south to include the lower Rio Purus. Rio Madeira and upper Ama-
zon records probably belong here.
Remarks. — In working out this form it became necessary to examine and
compare the other members of the fuliginosa-meruloides group, of all of
which we have an ample series. My conclusions differ from those of Zim-
mer (1934a, 18-20), in that I recognize two species instead of uniting them
all under one. With the nominate race fuliginosa I would range rujo-
olivacea and atrirostris as conspecies. These agree in having the bill dark-
colored (the maxilla at least), the throat with distinct mottling or barring,
and the breast with fine buffy streaks. The buffy postocular stripe is
usually well marked. These three races are readily distinguishable from
each other by their different general coloration. D. fuliginosa atrirostris
was originally described from Bolivia, but its range was extended to the
west bank of the Rio Tapajbz by Zimmer (1934a, 19) and by Griscom and
Greenway (1941, 219). There are indications that the birds of these two
respective regions are not precisely the same, but into this point I do not
care to go further at present.
Under the second species, meruloides, I would range, besides the nomi-
nate race, phceochroa, lafresnayei, christiani, ridgwayi, and the new form
I am describing herewith. In these forms the bill tends to be lighter-
colored, although in the northern races, christiani and ridgwayi, it is al-
most as dark as in atrirostris. The underparts are almost unicolored, only
the throat being paler than the rest. The breast is without pale streaks,
and the throat is either plain or else with very faint and narrow indications
of cross-barring — not at all like it is in the fuliginosa group. The pale
postocular stripe is restricted and indistinct, and sometimes wanting. I
am satisfied that these two groups represent two specific types, even
though at present I have no proof that they occur together at any one
place. It is hard to believe that the new race has not been taken by other
collectors, but if so it must have been misidentified, or confused with
18
Annals of the Carnegie Museum
VOL. 31
phceochroa. It is nevertheless perfectly distinct from the latter, as shown
by a comparison in series.
Incidentally, I find that D. meruloides christiani of western Colombia
is an easily recognizable race when smoothly and uniformly made-up
specimens are used for comparison; in coloration it is intermediate be-
tween lafresnayei and ridgwayi, but separable from both.
Literature Cited
Chapman, Frank M.
1917. The Distribution of Bird-Life in Colombia. — Bulletin American
Museum of Natural History, 36, 1917, pp. x -f 729, 41 pis.
Griscom, Ludlow, and Green way, James C., Jr.
1941. Birds of Lower Amazonia. — Bulletin Museum Comparative
Zoology, 88, No. 3, June, 1941, 83-344.
Gyldenstolpe, Nils.
1945. The Bird Fauna of Rio Jurua in western Brazil. — Kungliga Sven-
ska Vetenskaps-Akademiens Handlingar, (3), 22, No. 3,
August 14, 1945, pp. 338, 1 map.
Hellmayr, C. E.
1925. Catalogue of Birds of the Americas (etc.). Furnariidae-Den-
drocolaptidae. — Field Museum of Natural History Zoological
Series, 13, Part 4, December 29, 1925, pp. iv -f 390.
Peters, James L.
1929. An Ornithological Survey of the Caribbean Lowlands of Hon-
duras.— Bulletin Museum Comparative Zoology, 69, October,
1929, 397-478.
Wetmore, Alexander.
1939. Five New Races of Birds from Venezuela. — Smithsonian Miscel-
laneous Collections, 98, No. 4, March 10, 1939, 1-7.
Zimmer, John T.
1934a. Studies of Peruvian Birds. XIII. — American Museum Novitates
No. 728, May 31, 1934, 1-20.
1934b. Studies of Peruvian Birds. XIV. — American Museum Novitates
No. 753, November 10, 1934, 1-26.
1934c. Studies of Peruvian Birds. XV. — American Museum Novitates
No. 756, November 30, 1934, 1-20.
1934d. Studies of Peruvian Birds. XVI. — American Museum Novitates
No. 757, November 30, 1934, 1-22.
ART. 3. A REVIEW OF THE HYDNACE^ (FUNGI) OF
WESTERN PENNSYLVANIA
By LeRoy K. Henry
(Plates 1-2)
The family Hydnaceae or tooth-bearing fungi belong to that class of
basidiomycetes which bear their spores upon downward directed teeth,
spines, warts, or folds. They vary in habit from resupinate, through later-
ally sessile to upright forms with distinct stem and cap. The resupinate
plants are often inconspicuous and this may account for their scarcity in
herbarium collections. The upright plants with fleshy or leathery stems
and caps and the large, laterally sessile, fleshy forms are more readily seen
and recognized. These fungi grow in a variety of habitats such as, on the
ground in leaf mold, on decaying wood, and on injured areas of living trees.
I have included here the intermediate genera Irpex and Phlebia which
grade into the Polyporaceae by having the hymenium more or less poroid in
the young stage of development.
This review is based upon the specimens in the Herbarium of the Car-
negie Museum which have been collected in western Pennsylvania.
As the eastern boundary of western Pennsylvania, I have arbitrarily
chosen the eastern borders of Potter, Clinton, Centre, Huntingdon, and
Fulton counties. The initials following the collection data are to be in-
terpreted as follows: l.k.h. — L. K. Henry ; o.e.j. — O. E. Jennings; d.r.s. —
D. R. Sumstine; m.b.k. — Marie B. Knauz; H.s.w. — H. S. Wieand. The
numerals following the author’s initials are his field numbers.
Synopsis of the Genera of Hydnace^
I. Fruiting-body absent.
A. Clavate spines attached directly to the substratum; on wood
Mucronella
and bark.
H. Fruiting-body present
A. Fruiting-body resupinate.
1. Hymenium borne on variable spines, conical, awl-shaped, or
cylindrical, typically divided; sporophore membranaceous,
floccose, crustaceous, or rarely waxy; cystidia present; on
wood
Odontia
19
of Issue July 30, 1948.
20
Annals of the Carnegie Museum
VoL. 31
2. Hymenium borne on long awl-shaped spines; sporophore
floccose, fleshy or waxy ; no cystidia ; on wood Mycoacia
3. Hymenium borne on short, blunt, hemispherical or cylindrical
warts, sometimes awl-shaped and fragile; sporophore mem-
branaceous, soft crustaceous or waxy Grandinia
4. Hymenium borne on spines, soft, conical to awl-shaped;
sporophore soft floccose, dark colored; on wood. . . . Caldesiella
B. Fruiting-body resupinate to reflexed to pileate.
1. Hymenium borne on low, elongated folds; sporophore fleshy-
waxy Phlehia
2. Hymenium borne on blunt, often flattened and branched teeth ;
sporophore fleshy-tough Radulum
3. Hymenium borne on spines; sporophore consisting of branched
processes in a coarse brownish tomentum Gloiodon
4. Hymenium borne on long, pendant, awl-shaped spines;
sporophore tuberculiform or much branched, fleshy or sub-
fleshy ; on wood Hericium
C. Fruiting-body substipitate or sessile, reflexed and broadly at-
tached; coriaceous to subfleshy; on wood.
1. Hymenium borne on round or flattened spines. . Steccherinum
2. Hymenium at first poroid, breaking up into irregular, flattened
teeth Irpex
D. Fruiting-body generally pileate and centrally stipitate.
1. Sporophore fibrous, tough, often composed of two layers of
dilTerent texture; dark colored; terrestrial Calodon
2. Sporophore fleshy, white to pinkish-cinnamon; terrestrial.
Dentium
3. Sporophore fleshy to subfleshy, generally dark colored; on
wood Hydnum
Key to Calodon
I. Pileus single or confluent, up to 12 cm. in diameter, brownish in center and
depressed, tomentos, azonate to subzonate, with a broad whitish marginal
zone; texture duplex, the outer layer soft and felty, the inner hard, compact
and woody, extending down into the stipe as a core; odor fragrant in drying;
spines awl-shaped, round, 2-4 mm. long, white when fresh, becoming grayish-
brown when dry C. amicus
1948
Henry: Hydnace^ of Western Pennsylvania
21
I. Pileus and spines darker in color.
A. Pileus convex to plane, azonate, finely tomentose, subrotund to irregular,
2-12 cm. in diameter; texture duplex, consisting of two thick layers, a
soft, spongy outer layer and a hard, compact inner layer continuing into
the stipe as a core; stipe bulbous at base; spines decurrent, 1-6 mm. long,
awl-shaped, round, acute, color of pileus C. velutinus
B. Pileus thin, centrally depressed to infundibuliform, distinctly zoned,
usually light colored at margin to pecan brown at center; texture more or
less homogeneous, tough, fibrous; stipe short and broad with a spongy,
bulbous base, colored like the pileus; spines decurrent, 1-3 mm. long,
slender, round, color of pileus C. zonatus
C. Pileus more rigid, thicker, less zonate and rougher than in C. zonatus; not
exuding a reddish liquid if bruised when fresh C. scrobiculatus
Key to Hericium
I. Sporophore a solid or porous body, laterally attached, bearing on the lower
part soft, fleshy, white spines, 1-4 cm. long and more or less coalesced at base.
H. Erinaceus
I. Sporophore soft, fleshy, whitish, consisting of a mass of branched processes.
A. Intricately branched and anastomosing; spines 1-6 mm. long, uniformly
distributed on the under side of the branches H. laciniatum
B. Several main branches, relatively short and stout, from which arise short,
slender, terminal branches; spines 5-15 mm. long, arising in terminal
clusters or on short lateral spurs H. coralloides
Key to Hydnum
I. Pileus inherently scaly, light brown to fawn; stipe tapering to a small white
root; teeth up to 2 mm. long H. Underwoodii
II. Pileus not scaly, smooth to pubescent or felted, young teeth violet.
H. fuligineo-violaceum
III. Pileus distinctly scaly, dark brown, thick; stipe thick, stout, base not taper-
ing and of same color as pileus; teeth up to 1 cm. long when mature.
H. imbricatum
IV. Pileus densely tomentose or strigose-hairy, tan or yellow H. cristatum
V. Pileus smooth, blackish-green; stipe of same color H. atroviride
Key to Irpex
I. Sporophore resupinate, cinnamon to dark brown I. cinnamomeus
II. Sporophore reflexed, whitish I. mollis
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Annals of the Carnegie Museum
VoL. 31
Key to Mycoacia
I. Sporophore adnate to the substratum, waxy, bright yellow, with a strong
odor of bitter almonds when fresh; spines 4-10 mm. long, in nodules or singly.
M. setosa
II. Sporophore separable from the substratum, fleshy-membranaceous, yellow
to chrome orange when fresh, margin fibrillose with orange, rhizomorphic
strands; spines 1-4 mm. long, occasionally confluent M . fragilissima
III. Sporophore adnexed to the substratum by a loose, white, floccose layer;
hymenial layer waxy and drab to brownish in color, not completely covering
the floccose layer, its spines 6 mm. long .M. Himantia
Key to Odontia
I. Teeth very short, granular, confluent; on conifers O. ahieticola
II. Teeth fine, awl-shaped; often with 2-3 points, scarcely distinct. .0. fusco-atra
Key to Phlebia
I. Sporophore usually resupinate, waxy-soft, flesh colored to bright red, margin
fimbriate to dentate-radiate; hymenium red to reddish-brown, with radiating
folds P. radiata
II. Sporophore usually reflexed, coriaceous, rust-brown, drying darker, somewhat
tomentose, zonate; hymenium smoky-purplish, undulate, rugose, subradiat-
ing P. strigoso-zonata
Key to Radulum
I. Sporophore waxy-fleshy, edge white, membranaceous-cottony. .R. orhiculare
II. Sporophore crustaceous-waxy, edge white, villose-floccose R. quercinum
Key to Steccherinum
I. Fruiting-body resupinate to reflexed or entirely pileate and laterally sessile,
fibrous, tough; upper surface of pileus soft, tomentose, zoned, buff; hymenial
surface pinkish-cinnamon to vinaceous-cinnamon, with a whitish, or pubes-
cent margin; spines up to 2 mm. long, slender, awl-shaped, round, angular or
flattened *S. ochraceum
II. Fruiting-body distinctly pileate or rarely reflexed.
A. Sporophore tough, fibrous, consisting of many sessile, horizontal, im-
bricated and confluent pilei; single pilei 3-15 cm. wide, 2-15 cm. long;
fibrous and pubescent on upper surface, white at first, drying wrinkled
and ochraceous-buff ; spines 6-18 mm. long, slender, found, pointed,
white, becoming reddish to brown on drying. . .' 5. septenfrionale
1948
Henry: Hydnace^ of Western Pennsylvania
23
B. Sporophore soft, fibrous- fleshy, juicy when fresh and containing a milky,
sticky sap; pilei sessile, solitary to imbricate, often arising from a re-
supinate base; single pilei up to 15 cm. wide and 8 cm. long, densely
fibrous or hairy on upper surface, whitish at first, becoming light buff
to liver brown in drying; spines 1-6 mm. long, slender, round, pointed,
tawny S. pulcherrimum
C. Sporophore fibrous, subfleshy to tough, usually laterally stipitate, but
occasionally eccentric or sessile with abortive or fully developed, kidney-
shaped pilei arising from the upper surface of the lower ones; surface of
pilei finely pubescent, dull cinnamon-buff, faintly zoned toward the mar-
gin; spines 1-3 mm. long, slender, round, angular or flattened, whitish at
first, becoming pinkish in drying and finally russet S. adustum
Enumeration of Species
Caldesiella ferruginosa (Fries) Sacc.
Armstrong County: near Leechburg, d.r.s. Allegheny County: Frick
Park, Pittsburgh, d.r.s. Mercer County: Transfer, d.r.s. Westmoreland
County: Idlewild Park, Ligonier, d.r.s.
Calodon amicus Quel. (Plate 1, fig. 1)
Armstrong County: Kittanning, d.r.s. Bedford County: Sulphur Springs,
D.R.S. Fayette County: Ohiopyle, d.r.s.; near Somerfield, d.r.s. Venango
County: 1 mi. n. of Lisbon, h.s.w.
Calodon scrobiculatus (Fries) Quel. (Plate 1, fig. 2)
Beaver County: Temple Hollow woods, west edge of Aliquippa, l.k.h.,
2915. Centre County: Woodward, d.r.s.; Poe Paddy Park, m.b.k. Clarion
County: near Clarion, d.r.s. ; Cook Forest, d.r.s. Fayette County: Ohiopyle,
D.R.S.; near Somerfield, d.r.s. Venango County: near Emlenton, m.b.k.
Westmoreland County: Forbes Forest, 3 mi. s.e. of Rector, m.b.k.
Calodon velutinus (Fries) Quel.
Armstrong County: Kittanning, d.r.s. Clarion County: near Clarion,
D.R.S. Crawford County: French Creek near Cochranton, h.s.w. Fayette
County: Ohiopyle, d.r.s. Westmoreland County: near Mellon Estate, 6 mi.
s.e. of New Florence, d.r.s. Venango County: 1 mi. n.e. of Lisbon, h.s.w.
Calodon zonatus (Fries) Quel.
Armstrong County: Kittanning, d.r.s. Erie County: Weiss Library
woods, near Erie, o.E.j. Fayette County: Ohiopyle, o.e.j.; near Somerfield,
D.R.S.
24
Annals of the Carnegie Museum
VoL. 31
Dentium repandum (Fries) S. F. Gray
Allegheny County: 1 mi. west of Mt. Nebo. Armstrong County: Kit-
tanning; across Big Buffalo Creek from W. Winfield. Bedford County:
Sulphur Springs. Butler Comity: Little Buffalo Creek near Monroe Sta-
tion; Watson’s Run, 2 mi. s. of Leasuresville; Little Connoquenessing
Creek, 3 mi. n.e. of Harmony; near Slippery Rock. Cameron County: along
Route 872, 10 mi. s. of Sinnemahoning. Centre County: near Rebersburg;
Woodward. Clarion County: Cook Forest. Clearfield County: State Game
Lands, No. 34, s. of Medix Run. Erie County: Weiss Library w'oods, s.w.
of Erie. Fayette County: Ohiopyle. Forest County: Cook Forest. Somerset
County: near Buckstown; 3 mi. w. of Berlin. Venango County: 1 mi. n. of
Lisbon. Westmoreland County: Shades Ravine, 2 mi. e. of Trafford; Forbes
Forest, 3 mi. s.e. of Rector; Mellon Estate, 6 mi. s.e. of New Florence.
Gloiodon strigosus (Fries) Karst. (Plate 2, fig. 2)
Armstrong County: near Leechburg, d.r.s. Centre County: in Poe Paddy
Valley, m.b.k.
Grandinia Brinkmanni (Bres.) Bourd. & Galz.
Allegheny County: Frick Park, Pittsburgh, d.r.s.
Hericium coralloides Scop. (Plate 1, fig. 3)
Allegheny County: Frick Park; near Sewickley. Armstrong County:
Kittanning. Butler County: Cook Forest; near Clarion. Elk County: south
of Kane. Lawrence County: Muddy Creek Falls, near mouth of Muddy
Creek. Mercer County: 2 mi. n.e. of Grove City. Potter County: near Carter
Camp. Venango County: 1 mi. n.e. of Lisbon; Little Scrubgrass Creek,
n.e. of Sutton’s Mills. Westmoreland County: near Kiski campus, Salts-
burg.
Hericium erinaceus (Fries) Pers. (Plate 1, fig. 4)
Allegheny County: Tom’s Run back of Dixmont; Frick Park; Allegheny
Cemetery; Thornburg; along Washington Blvd., East Liberty district,
Pittsburgh; Herron Hill, Pittsburgh. Beaver County: 2 mi. west of Beaver.
Butler County: Little Buffalo Creek at Monroe Station; Iron Bridge, Big
Buffalo Creek. Bedford County: near Bedford. Fayette County: Ohiopyle;
Cranberry Glade Run, n. end of Wharton Township. Lawrence County:
Muddy Creek Falls at mouth of Muddy Creek. Potter County: near Carter
Camp. Washington County: along Buffalo Creek near junction with Buck
Run. Westmoreland County: near Edgecliff; Idlewild Park near Ligonier ;
near Waterford; n.w. of Saunders.
1948 Henry: Hydnace^ of Western Pennsylvania 25
Hericium laciniatum (Leers) Banker
Allegheny County: 2 mi. due east of Upper Talleycavey; Turtle Creek;
Frick Park; 4 mi. east of Monongahela; near Sewickley; Fallen Timber
Hollow opposite Sutersville. Armstrong County: near Leechburg. Beaver
County: Beaver Falls. Bedford County: Sulphur Springs. Butler County:
near Saxonburg; 2 mi. s. of Leasuresville ; 4 mi. n.e. of Harmony. Crauford
County: Conneaut Lake. Mercer County: 2 mi. n.e. of Grove City. Potter
County: near Carter Camp. Somerset County: near Jennerstown; near
Buckstown.
Hydnum atroviride Morgan
Fayette County: Ohiopyle, d.r.s.
Hydnum cristatum Bres.
Fayette County: Ohiopyle. d.r.s.
Hydnum fuligineo-violaceum Kalch.
Fayette County: Ohiopyle; fits description except for small size.
Hydnum imbricatum (L.) Fries
Centre County: Woodward; Pleasant Gap, d.r.s. Fayette County: Ohio-
pyle, O.E.J.; near Somerfield, d.r.s.
Hydnum Underwoodii Banker
Erie County: Weiss Library woods, 8 mi. s.w. of Erie, o.E.j. Fayette
County: Ohiopyle, o.E.j.
Irpex cinnamomeus Fries
Allegheny County: Coraopolis; North Park. Armstrong County: Kit-
tanning. Bedford County: Sulphur Springs. Centre County: Woodward;
Reitz Gap. Clearfield County: State Game Lands, No. 34, s. of Medix Run.
Erie County: Presque Isle. Fayette County: Killarney Park, s. of Normal-
ville. Somerset County: near Jennerstown. Westmoreland County: Forbes
Forest, 3 mi. s.e. of Rector; Seward; Youngstown; Jones’ Mills; Derry;
Laurel Ridge, 1 mi. e. of Kregar.
Irpex mollis B. & C. (Plate 2, fig. 1)
Allegheny County: Frick Park, Pittsburgh. Armstrong County: Kit-
tanning. Butler County: Nixon Station on Butler Short Line, 4 mi. n.e. of
Harmony. Erie County: Presque Isle. Fayette County: Ohiopyle. Mercer
County: Blacktown. Westmoreland County: Ligonier; Derry; Mellon
Estate, 6 mi. s.e. of New Florence; near South Greensburg.
26
Annals of the Carnegie Museum
VoL. 31
Mucronella calvum A. & S.
Allegheny County: Frick Park, Pittsburgh, d.r.s.
Mycoacia fragillissima (B. & C.) Miller
Allegheny County: near Sardis; Sandy Creek. Armstrong County: Kit-
tanning; near Ford City. Bedford County: Sulphur Springs. Butler County:
near Zelienople; near Butler. Centre County: Penn’s View. Washington
County: along Raccoon Creek 2 mi. above Murdocksville. Westmoreland
County: Latrobe; Jones’ Mills.
Mycoacia Himantia (Schw.) Miller
Allegheny County: Sandy Creek; Coraopolis; along Route 910, 3 mi. s.e.
of Bakerstown. Armstrong County: Kittanning. Butler County: Saxonburg;
near Culmerville. Crawford County: Pymatuning Swamp at Hartstown.
Cambria County: Cresson. Fayette County: Ohiopyle. Somerset County:
near Jennerstown. Washington County: along Raccoon Creek above
Murdocksville. Westmoreland County: Kingstown; near Saltsburg.
Mycoacia setosa (Pers.) Donk
Armstrong County: Kittanning, d.r.s. Westmoreland County: Idlewild
Park near Ligonier. d.r.s.
Odontia abieticola B. & C.
Allegheny County: Nadine, d.r.s., determined by S. H. Jackson. Clarion
County: near Clarion, d.r.s.
Odontia fusco-atra (Fries) Bres.
Allegheny County: Frick Park, Pittsburgh, d.r.s.
Phlebia radiata Fries
Allegheny County: near Sewickley; near Bellevue; Frick Park, Pitts-
burgh. Armstrong County: Kittanning; near Leechburg. Butler County:
near Saxonburg; near Slippery Rock; near Butler; Nixon Station on Butler
Short Line; near Zelienople; 4 mi. n.e. of Harmony. Clarion County: near
Clarion ; Cook Forest. Crawford County: near Hartstown. Lawrence County:
at mouth of Muddy Creek. Somerset County: Kooser State Park. Potter
County: near Carter Camp. Venango County: near Lisbon. Westmoreland
County: Waterford; along Conemaugh River opposite Saltsburg.
Phlebia strigosa-zonata Schw.
Clarion County: near Clarion, d.r.s. Elk County: near Kane, d.r.s.
1948
Henry: Hydnace^ of Western Pennsylvania
27
Radulum orbiculare Fries
Butler County: near Slippery Rock, d.r.s. C/anow near Clarion,
D.R.s. Centre County: Woodward, d.r.s.
Radulum quercinum Fries
Centre County: Woodward, d.r.s.; State Game Lands, No. 33, along
Port Matilda-Philipsburg Road, l.k.h., 5034.
Steccherinum adustum (Schw.) Banker (Plate 2, fig. 3)
Allegheny County: Frick Park, Pittsburgh; Power’s Run opposite
Verona; along Beaver Grade Road near Montour Run. Armstrong County:
Kittanning. Beaver County: across Ohio River from Ambridge. Bedford
County: Sulphur Springs. Butler County: Slippery Rock; 4 mi. n.e. of
Harmony. Centre County: 10 mi. s. of State College. Clarion County: near
Leeper. Elk County: near Kane. Fayette County: Ohiopyle; Killarney
Park, s. of Normalville. Lawrence County: Slippery Rock Creek above
Wurtemberg. Mercer County: Transfer. Somerset County: near Somerset ;
Haines; near Trent. Venango County: 1 mi. n. of Lisbon. Westmoreland
County: Mellon Estate, 6 mi. s.e. of New Florence; Forbes Forest, 3 mi.
s.e. of Rector; Shades Ravine, 2 mi. e. of Trafford.
Steccherinum ochraceum (Fries) S. F. Gray
Allegheny County: Frick Park, Pittsburgh; Guyasuta Hollow at Aspin-
wall; near Wildwood; Y2 i^ii. s. of Smithdale; along Route 910, 3 mi. s.e. of
Bakerstown ; Powers Run opposite Verona. Armstrong County: Kittanning;
near Leechburg. Bedford County: Sulphur Springs. Butler County: near
Zelienople; along Watson’s Run, 2 mi. s. of Leasuresville. Cambria County:
Cresson. Cameron County: along Route 872, 10 mi. n.e. of Sinnemahoning.
Centre County: State Game Lands, No. 33, 3 mi. s.e. of Philipsburg.
Lawrence County: Slippery Rock Creek at Kennedy’s Mills; near Wes-
minster College, New Wilmington. McKean County: Kane; Tionesta
Tract near Brookston. Somerset County: near Buckstown on Fomes
near Somerset. Venango County: 1 mi. n.e. of Lisbon. Wash-
ington County: near New Eagle; above Raccoon Creek IJ^ mi. s. of
Murdocksville. Westmoreland County: Shades Ravine, 2 mi. e. of Trafford;
Idlewild Park near Ligonier; near New Florence; along Conemaugh River
across from Saltsburg; Lynn Run, 3 mi. s.e. of Rector; along Route 80,
10, mi. w. of Saltsburg.
28
Annals of the Carnegie Museum
VoL. 31
Steccherinum pulcherrimum (B. & C.) Banker
Allegheny County: Frick Park, d.r.s.; Pittsburgh; near Sandy Creek,'
L.K.H. Armstrong County: Kittanning, d.r.s. Westmoreland County: near
New Kensington, m.b.k.
Steccherinum septentrionale (Fries) Banker (Plate 2, fig. 4) •
Cambria County: Cresson, d.r.s. Erie County: near Erie, J. H. Miller.
Mercer County: no locality given, h.s.w. Somerset County: near Bucks-
town, o.E.j.
Summary
A generic key and descriptive keys to thirty-two species of Hydnaceae
have been provided. Grandinia Brinkmanni, Hydnum atroviride, H.
cristatum, H. fuligineo-violaceum, Mticronella calvum, and Odontia fusco-
atra are represented by one collection of each.
Gloiodon strigosus, Hydnum Underwoodii, Mycoacia setosa, Odontia
abieticola, Phlebia strigosa-zonata and Radidum quercinum were each
collected in two different localities.
All species collected from three or more localities, and yet not plentiful
enough to be considered as common, have been designated by including the
collector’s name or initials after the locality.
Out of a total of thirty-two counties in western Pennsylvania, no col-
lections of Hydnaceae have been obtained from Blair, Clinton, Fulton,
Huntingdon, Indiana, Jefferson, and Warren counties.
In order to obtain a more complete record of the distribution of the
Hydnaceae in western Pennsylvania collections are needed from these
counties and additional collections from the southeastern and north-
eastern counties of western Pennsylvania.
1948
Henry: Hydnace^ of Western Pennsylvania
29
BIBLIOGRAPHY
Miller, L. W. and Boyle, J. S.
The Hydnaceae of Iowa. University of Iowa Studies in Natural History,
Vol. 18, No. 2, 1943.
Banker, H. J.
A contribution to a revision of the North American Hydnaceae. Memoirs
of the Torrey Botanical Club, Vol. 12, No. 2, 1906.
Coker, W. C.
The Hydnums of North Carolina. Journal of the Elisha Mitchell Sci-
entific Society, Vol. 34, No. 4, 163-197, 1919.
Bourdet, H. and Galzin, A.
Hymenomycetes de France. Societe Mycologique de France, 1927.
Rea, Carleton.
British Basidiomycetes. The British Mycological Society, Cambridge
University Press, London, 1922.
30
Annals of the Carnegie Museum
VoL. 31
EXPLANATION OF PLATE 1
Fig. 1. Calodon amicum Quel. X 3/7.
Fig. 2. Calodon scrobiculatus (Fries) Quel. X 3/7.
Fig. 3. Hericium coralloides Scop. X l/3.
Fig. 4. Hericium erinaceus (Fries) Pers. X 5/12.
ANNALS, CARNEGIE MUSEUM, Vol. XXXI
Art. 3
Plate 1
32
Annals of the Carnegie Museum
VoL. 31
EXPLANATION OF PLATE 2
Fig. 1. Irpex mollis B. & C. X 2/5.
Fig. 2. Gloiodon strigosus (Fries) Karst. X 3/5.
Fig. 3. Steccherinum adusium (Scho.) Banker X 1/2.
Fig. 4. Steccherinum se plenty ion ale (Fries) Banker. X 2/5.
ANNALS, CARNEGIE MUSEUM, Vol. XXXI
Art. 3
Plate 2
v.^
$01 .1^
.f Af>b
ART. 4. CRITICAL REMARKS ON THE OVEN-BIRDS
By W. E. Clyde Todd
Like the Wood-hewers or Dendrocolaptidae, to which they are closely
allied, the Oven-birds or Furnariidae are wholly neotropical and largely
South American in their distribution. As yet their classification cannot be
said to be absolutely settled, and some genera, notably Asthenes and
Cranioleuca, are composed of diverse elements. The group as a whole is
much more difficult to study than are the Tree-creepers; racial differences,
when present, are often obscured by seasonal, sexual, and age variations,
and large series are indispensable in working out the taxonomy. The
Carnegie Museum collection has 3,286 specimens of Oven-birds, repre-
senting 245 species and subspecies. For the loan of additional specimens of
certain forms needed for comparison I am deeply indebted to the authori-
ties of the U. S. National Museum, the Academy of Natural Sciences of
Philadelphia, the Museum of Comparative Zoology, and in particular
the American Museum of Natural History. In the present paper descrip-
tions of eight apparently new races are presented, together with some
critical notes and remarks on the taxonomy of certain other forms.
As in other papers by the writer, all measurements are in millimeters,
and that of the bill is for the exposed culnien. The names of colors are taken
mostly from Ridgway’s “Color Standards and Color Nomenclature.”
Most of the localities appearing in the text were listed in an earlier paper
by the writer, on Hummingbirds (Annals Carnegie Museum, Vol. 29,
1942, pp. 365-370).
Furnarius torridus Sclater and Salvin.
Mr. Zimmer (1936a, 4) has discriminated this form from F. tricolor,
with which Hellmayr (1925, 20) had confused it. I would call the upper-
parts of torridus Sanford’s brown (not chestnut), and those of tricolor raw
sienna to Mars yellow. We have specimens of both forms from Sao Paulo
de Olivenga.
Furnarius leucopus exilis Todd.
Hellmayr (1925, 22) after examination of von Pelzeln’s type of F.
longirostris, found that F. agnatus of Sclater and Salvin is a synonym of
the other — a wholly unexpected development. He doubted if two races
occurred in the region, and relegated my exilis to synonymy. After corn-
sued October 15, 1948
34
Annals of the Carnegie Museum
VOL. 31
paring our series of the latter (twenty-four specimens) with eight speci-
mens from the Goajira Peninsula (U. S. National Museum), I find that
exilis is a barely recognizable race, distinguishable only by its slightly
deeper general coloration and (average) shorter bill. Our single specimen
of longirostris from Rio Hacha is exceptionally pale. But if, as Mr. Peters
suggests (in litt.), birds from Rio Aurare, Venezuela, are the same as those
from the Santa Marta region, then Cory’s name venezuelensis, based on
the former, would naturally have priority. The easiest way out of the
dilemma is to follow Hellmayr in recognizing only one form from northern
Colombia and Venezuela.
Cinclodes fuscus oreobates Scott.
Although in 1922 I took a different view, I am now inclined to follow
Hellmayr in reducing oreobates to a subspecies of fuscus. Greatly to my
surprise 1 find no difference between Santa Marta specimens and those
from the Eastern Andes of Colombia (Lagunillas, Rio Negro), whereas the
bird from the Venezuelan Andes is clearly distinct {heterurus von Ma-
darasz).
The Northern Races of Synallaxis albescens.
Hellmayr, writing in 1925, recognized four races of Synallaxis albescens
in northern South America and one additional race in Costa Rica. To
S. a. albigularis Sclater he assigned an immense range, from Guiana, north-
ern Brazil, and most of Venezuela, west to Peru, Ecuador, and Colombia.
S. a. josephince Chubb (1919) and 5. occipitalis von Madarasz (1903) were
considered to be synonyms. S. a. nesiotis Clark (1902), a “rather unsatis-
factory race,” was restricted to Margarita Island, Trinidad, and the
coastal region of northeastern Venezuela. S. a. perpallida Todd (1916),
originally described from the Goajira Peninsula, was extended to cover the
region south to Maracaibo. Birds from western Colombia and Panama
were assigned to S. a. hypoleuca Ridgway (1909), and those from south-
western Costa Rica to 5. a. latitabunda Bangs.
Chapman was the next author to make a critical study of this group
(1931, 88-91). He showed very clearly that Sclater’s albigidaris had noth-
ing to do with S. albescens, but belonged to a perfectly distinct species — a
conclusion which I have independently verified after examination of our
series. He revived Chubb’ s josephince for the Roraima bird, and suggested
that the bird from northwest Venezuela would have to be called occipitalis,
1948
Todd: Critical Remarks on Oven-Birds
35
instead of alhigularis. Also, he intimated that my perpallida might be the
same as the latter.
Now comes Zimmer (1936b, 12-18) with further additions and refine-
ments. He describes a new race, trinitatis, with a range extending from
Trinidad through northeastern Venezuela to the Orinoco Valley. The bird
from eastern and western Colombia he calls insignis, subsp. nov. He
restricts nesiotis to Margarita Island and the mainland opposite. He takes
occipitalis for the bird of the Merida region (the type-locality), but in-
cludes specimens from Caracas under the same name. He identifies speci-
mens from an intermediate locality in Venezuela, however, as perpallida,
and he questions the pertinence of Ridgway’s name hypoleuca for the
Panama bird.
Examination of our series of specimens has led me to somewhat dif-
ferent conclusions in certain respects. Insofar disjosephince is concerned, I
have nothing to offer, as this form is unrepresented in our collection.
Otherwise, however, we have a good series from Venezuela and Trinidad.
Comparing birds from the latter locality with three specimens in fresh
plumage (U. S. National Museum) of nesiotis from Margarita Island, I
cannot verify any of the distinctions which Zimmer seeks to make for his
proposed new race trinitatis, although he claims that it is “very distinct.”
Moreover, birds from La Cumbre de Valencia, Tocuyo, and Guarico,
Venezuela, so far as I can see, are not definitely separable. The last two
localities are in the same region from which Zimmer records specimens of
perpallida. Obviously, Zimmer could not have seen true perpallida, or he
would not have so identified them. Five specimens from the Santa
Marta region of Colombia, which at one time (following other authors) I
called alhigularis, are close counterparts of these Venezuelan specimens.
A series from the north coast of Venezuela (from Carabobo to the Cumana
region) is not different either. In short, I can find no valid grounds for
recognizing more than one race from Trinidad, Margarita Island, the
Orinoco Valley and north coast of Venezuela, and the Santa Marta high-
lands of Colombia. For this form I adopt the oldest name, nesiotis Clark —
type-locality Margarita Island.
Synallaxis albescens occipitalis von Madarasz, from the Merida region
of Venezuela, is not represented in our collection, but four topotypical
specimens have been available for examination through the courtesy of the
American Museum of Natural History. It proves to be an easily recogniza-
ble race by reason of its darker general coloration. The forehead is black-
ish, with no grayish shade. Its range appears to be restricted to the higher
36
Annals of the Carnegie Museum
VOL. 31
elevations of the Andes of Merida. At lower elevations it is represented by
other races, nesiotis and insignis. The latter is the form of the Andean
region of Colombia, and inhabits the Tropical Zone of that country on
either side of the Andean ranges and in the Magdalena Valley. Specimens
from western Colombia are slightly different, as Zimmer points out, and
approach Central American birds in some respects. Zimmer refers a speci-
men from Calamar, on the Caribbean coast of Colombia, to insignis. I
have examined this specimen; it is grayish above like the interior form,
but our specimens from Calamar and from the north coast in general
obviously represent a different race, which I propose to call
Synallaxis albescens littoralis, subsp. nov.
Twenty-five specimens: Fundacibn, Calamar, Turbaco, Cartagena,
Puerto Zapote, Lorica, Monteria, and Soatata, Colombia.
Type, No. 52,654, Collection Carnegie Museum, adult male; Lorica,
Rio Sinu, Colombia, February 23, 1916; M. A. Carriker, Jr.
Subspecific characters. — Similar in general to Synallaxis albescens insignis
Zimmer of the Andean region of Colombia, but tail shorter, upperparts more
brownish, less grayish ; and underparts averaging paler, with less grayish
wash. Wing (type), 57; tail, 70; bill, 11.5; tarsus, 19.
Range. — Caribbean littoral of Colombia.
Remarks. — In describing his new race insignis Zimmer (1935b, 3 ; 1936b,
16-18) noted that “birds from the lower Magdalena river are somewhat
more grayish on the back.” 1 have seen ten specimens from this region; to
my eye they are inseparable from the rest of the series of insignis; at any
rate, they seem distinct from the Caribbean coast race. The latter is a
short- tailed, brownish-backed bird by comparison. It differs from
of Panama in its darker-colored underparts, paler wing-coverts, and par-
ticularly in its less developed (more restricted) crown-patch. Mr. de-
Schauensee (1945, 8) suspected the distinctness of Soatata birds.
Synallaxis albescens perpallida Todd
This form is the one that inhabits the Arid Tropical Zone of the Goajira
Peninsula, and ranges southward to Maracaibo and across to the eastern
shore of the Gulf of the same name. It is a small, pale race, markedly
whiter below than either nesiotis or littoralis. As already noted, Zimmer’s
reference of certain Venezuelan specimens to perpallida is questionable.
1948
Todd: Critical Remarks on Oven-Birds
37
Synallaxis albescens hypoleuca Ridgway.
Zimmer discounts hypoleuca, and states that “skins from Santiago, only
a little southwest of Nata [the type-locality], are inseparable from Costa
Rican examples.” I am at a loss to understand this dictum, since I find
that these same specimens, agreeing as they do with the type, are definitely
distinct from six Costa Rican specimens in our collection by reason of their
whiter underparts, with paler flanks. Our Costa Rican birds came from
Boruca (the type-locality of latitabunda Bangs), Paso Real de Terraba, and
Buenos Aires. Such being the case, I cannot at all follow Zimmer in con-
sidering the two one and the same form. Just where hypoleuca and littoralis
meet remains to be determined.
Synallaxis albescens griseonota, subsp. nov.
Two specimens: Santarem, Brazil.
Type, No. 72,311, Collection Carnegie Museum, adult male; Santarem,
Brazil, April 25, 1919; Samuel M. Klages.
Subspecific characters. — Similar to Synallaxis albescens incsqualis Zim-
mer, but upperparts more grayish, less brownish, and pileum and wing-
coverts paler rufous.
Range. — Rio Tapajoz region of Brazil.
Remarks. — In addition to the above I have examined two specimens from
Lago Grande in the collection of the Museum of Comparative Zoology.
These four specimens as a group are easily separable from a series of
incequalis from French Guiana by the characters above specified. It so
happens that incequalis was described from the south bank of the Amazon
(Villa Bella Imperatriz), but French Guiana birds handled at the same time
were considered to be identical.
Synallaxis cabanisi obscurior, subsp. nov.
Six specimens: Tamanoir, French Guiana.
Type, No. 61,152, Collection Carnegie Museum, adult male; Tamanoir,
Mana River, French Guiana, May 2, 1917; Samuel M. Klages.
Subspecific characters. — Resembling Synallaxis cabanisi macconnelli
Chubb of Roraima, British Guiana, but general coloration darker and
grayer; underparts dull neutral gray, the flanks with a slight brownish
wash (instead of hair brown to drab) ; upperparts more grayish, less brown-
ish; and size smaller.
38
Annals of the Carnegie Museum
VOL. 31
Measurements. — Three males: wing, 56, 58, 59; tail, 61, 67, 68. One
female: wing, 56; tail, 61.
Range. — French Guiana.
Remarks. — This new race has been compared only with macconnelli of
Roraima, which form is said to be very close to cabanisi (fide Hellmayr).
The smaller size and grayish general coloration set it off as an easily separa-
ble race. No form of this species, so far as I know, has ever been attributed
to French Guiana, so that the present record involves a considerable ex-
tension of range.
Synallaxis propinqua von Pelzeln.
A single specimen from Pied Saut, Oyapock River, French Guiana,
March 18, 1918, agrees well with specimens from eastern Ecuador in the
collection of the American Museum of Natural History. This constitutes a
new record for French Guiana.
Synallaxis cinnamomea Lafresnaye.
Eight specimens: El Cauca, La Palmita, and Pueblo Nuevo, Colombia.
These appear to be the first definite locality records for this species,
heretofore known only from “Bogota” skins. Two young birds are included
(August 2 and 19). Further comparisons convince me that this form dif-
fers enough from the Venezuelan bolivari to merit rank as a distinct species.
Nowhere, so far as known, do these two forms approach each other
geographically ; there is a wide gap between their respective ranges.
Certhiaxis cinnamomea albescentior, subsp. nov.
Twenty-four specimens: El Trompillo, Venezuela.
Type, No. 46,905, Collection Carnegie Museum, adult male; El Trom-
pillo, Carabobo, Venezuela, May 12, 1914; Samuel M. Klages.
Subspecific characters. — Similar to Certhiaxis cinnamomea fuscifrons
(von Madarasz) of northern Colombia and adjacent parts of Venezuela,
but underparts whiter, with less buffy wash; upperparts and wings ex-
ternally deeper rufous; and brown frontal areas averaging more re-
stricted.
Range. — Known at present only from the type-locality.
Remarks. — In this series, taken from May 11 to 22, there is great uni-
formity of coloration. As a series it is readily separable from another of
1948 Todd: Critical Remarks on Oven-Birds 39
fuscifrons by the above characters, although occasional odd specimens
might be hard to place, taken by themselves. The deeper rufous coloration
of the upper parts and the general whiteness of the underparts stand out
well by comparison. So far as I am aware this species has not been re-
corded from the north coast proper, but Sclater cites a record from the
“Plains of Valencia” (Proc. Zool. Soc. London, 1869, 252) ; doubtless it per-
tains to the present form.
This form has not been compared with orenocensis Zimmer, in which,
however, the forehead is said to be merely a little paler rusty than the
crown — not the case with the present series.
Pseudocolaptes boissonneautii boissonneautii (Lafresnaye).
Twenty-seven specimens: Paramo de Rosas, La Cuchilla, and Heights
of Tabay, Venezuela; Las Ven tanas, Ramirez, La Pica, Pena Blanca, and
Bitaco Valley, Colombia.
A remarkable feature about this species is the sexual difference in the
size and shape of the bill. In the male the bill runs 19-21 mm., and the
culmen is more or less curved downwards. In the female the bill is from
24.5 to 26.5 mm., and the culmen is straighter. What is the significance of
this difference?
After examining and comparing the above series I fail utterly to make
out the proposed race meridce of Hartert and Goodson. Hellmayr (1925,
177) admits it without question, but I cannot verify a single one of the
characters he specifies. The streaking on the crown, the mottling on the
underparts, and the yellow tinge on the throat and malar region are all
variable quantities both in Eastern Andean and in Venezuelan birds. A
single specimen from Bitaco Valley has the squamation below more dis-
tinct, and the tail darker rufous. It would represent oberholseri of Cor}^ —
if this is a valid race.
Philydor erythrocercus suboles, subsp. nov.
Nine specimens: Tonantins, Brazil.
Type, No. 96,926, Collection Carnegie Museum, adult male; Tonantins,
Rio Solimoes, Brazil, June 23, 1923; Samuel M. Klages.
Subspecific characters. — Similar to Philydor erythrocercus erythrocercus
(von Pelzeln) of the Guianas and northern Brazil, but upperparts more
rufescent, less olivaceous; underparts paler, with less yellowish wash.
Range. — Presumably the north bank of the upper Amazon in Brazil.
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Annals of the Carnegie Museum
VOL. 31
Remarks. — In Brazil, south of the Amazon, there is only one race of
Philydor erythrocercus, P. e. lyra Cherrie, which ranges from Maranhao
westward into Peru. On the north bank of the Amazon we have typical
erythrocercus coming in from the Guianas, and extending as far up the
river as Faro (at least). Still farther up, at Tonantins, we find the species
represented by a perfectly distinct race, characterized by rufescent
(medal bronze) upperparts and Dresden brown wing-edgings, while the
yellowish wash on the underparts is restricted mostly to the throat. It is
readily distinguished from lyra by the duller and paler color of the under-
parts, with no buffy suffusion whatever, and by the more olivaceous, less
brownish wing-edgings. Some of our specimens of lyra from Sao Paulo de
Olivenga and the Rio Purus are almost as pale below, but they show a
slightly buffy wash, and the wing-edgings are not at all the same. The
new race has nothing to do with P. subfulvus of eastern Ecuador, with
which it has been directly compared. I doubt if the two are even con-
specific.
Automolus infuscatus purusianus, subsp. nov.
Thirty-eight specimens: Hyutanahan, Nova Olinda, and Arima (Rio
Purus), Sao Paulo de Oliveng-a and Caviana (Rio Solimoes), Brazil.
Type, No. 87,395, Collection Carnegie Museum, adult male; Hyutana-
han, Rio Purus, Brazil, January 19, 1922; Samuel M. Klages.
Sub specific characters. — Similar to Automolus infuscatus paraensis
Hartert of the region south of the Amazon and east of the Rio Madeira,
but larger; upperparts more rufescent, less olivaceous by comparison; and
forehead more decidedly tinged with rufescent. Similar also to A . infuscatus
infuscatus (Sclater) of the north bank of the middle and upper Amazon,
but upperparts not quite so rufescent; and pileum more distinctly streaked
or squamate. Wing (type), 96; tail, 76; bill, 22; tarsus, 20.5.
Range. — Brazil, south of the Amazon, and west of the Rio Madeira.
Remarks. — Hellmayr (1925, 213) referred birds from the Rio Purus to
infuscatus, as also did Gyldenstolpe (1945, 158). Zimmer (1935a, 15) re-
marks that in Peru the birds from both banks of the Amazon are alike, so
that the question of the exact type-locality of infuscatus is of no great
importance. This is fortunate, since in western Brazil the birds from op-
posite banks of the Amazon are certainly not alike. Zimmer refers birds
from Teffe to infuscatus, which would by implication fix that name on our
birds from Manacapuru and Sao Paulo de Olivenga. With good series from
both banks available, there is no difficulty in discriminating between
1948
Todd: Critical Remarks on Oven-Birds
41
them. The new race is intermediate in its color characters between in-
fuscatus and paraensis. It has the striated pileum of the latter, but ap-
proaches the former in the rufescence of the upperparts. I find that our
Caviana birds (from east of the Rio Madeira) are somewhat intermediate,
but are perhaps best referred here.
Automolus melanopezus (Sclater).
Four specimens: Hyutanahan (Rio Purus), Brazil.
These specimens agree with the description of this species ; I have found
no others for comparison. The record involves a considerable extension of
the known range.
Automolus pallidigularis pallidigularis Lawrence.
Eleven specimens: El Cauca, El Tambor, Soatata, and Murindo,
Colombia.
Chapman says that this form is unknown in western Colombia, so that
our specimens are the first recorded from this region. The specimens have
been compared with a series from Panama (the type-locality) in the Bio-
logical Survey collection, and found to be indistinguishable therefrom.
I hold that this form should be kept specifically separate from A.
ochrolcemus.
Sclerurus rufigularis furfurosus, subsp. nov.
Four specimens: Obidos, Brazil.
Type, No. 83,826, Collection Carnegie Museum, adult female; Obidos,
Brazil, December 30, 1920; Samuel M. Klages.
Subspecific characters. — Similar to Sclerurus rufigularis fulvigularis
Todd of the Guianas and eastern Venezuela, but general coloration lighter
in tone ; upperparts Brussels brown (instead of argus brown) ; upper tail-
coverts argus brown (instead of auburn) ; buffy color of throat and breast
less intense; and posterior underparts Dresden brown (instead of Front’s
brown).
Range. — Known at present only from the type-locality, but probably
ranging farther west on the north bank of the Amazon (to the Rio Negro?).
Remarks. — Our Obidos specimens above listed were identified by Hell-
mayr as fulvigularis, but they differ consistently from a series of eleven
specimens of that form in the respects pointed out above, and in my
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Annals of the Carnegie Museum
VOL. 31
judgment they represent a valid race of somewhat restricted distribution
The Rio Negro probably separates the range of the present form from
that of the one now to be described as
Sclerurus rufigularis brunnescens, subsp. nov.
Eight specimens: Tonantins, Manacapuru, and Rio Manacapuru,
Brazil.
Type, No. 97,394, Collection Carnegie Museum, adult male; Tonantins,
Rio Solimoes, Brazil, July 21, 1923; Samuel M. Klages.
Subspecific characters. — Similar to Sclerurus rufigularis fulvigularis
Todd, but general coloration deeper, more brownish, less rufescent.
North bank of the Amazon, west of the Rio Negro.
Remarks. — This race varies away from fulvigularis in a direction op-
posite to that of furfurosus. It is deeper brown than the former; this is
particularly pronounced on the underparts and wing-edgings. The rusty
buff wash on the throat and breast is about the same in both. The wing-
edgings may be described as raw umber in the present form. Front’s
brown in fulvigularis . S. r. brunnescens is much duller- and darker-colored
than true rufigularis from the south bank of the Amazon.
1948
Todd: Critical Remarks on Oven-Birds
43
Literature Cited
Chapman, Frank M.
1931. The Upper Zonal Bird-Life of Mts. Roraima and Duida. — Bulletin
American Museum Natural History, 63, November 20, 1931,
1-135.
Gyldenstolpe, Nils.
1945. The Bird Fauna of Rio Jurua in western Brazil. — Kungliga Svenska
Vetenskaps-Akademiens Handlingar, (3), 22, No. 3, August 14,
1945, pp. 338, 1 map.
Hellmayr, C. E.
1925. Catalogue of Birds of the Americas (etc.). Furnariidae-Dendro-
colaptidae. — Field Museum of Natural History Zoological Series,
13, Part 4, December 29, 1925, pp. iv + 390.
deSchauensee, Rodolphe M.
1945. Notes on Colombian Antbirds, Ovenbirds, and Woodhewers, with
the Description of a New Form from Peru. — Notulae Naturae
No. 153, May 23, 1945, 1-15.
Zimmer, John T.
1935a. Studies of Peruvian Birds. XVII. — American Museum Novitates
No. 785, March 18, 1935, 1-24.
1935b. Studies of Peruvian Birds. NAJWl.— American Museum Novitates
No. 819, September 17, 1935, 1-8.
1936a. Studies of Peruvian Birds. XIX. — American Museum Novitates
No. 860, June 19, 1936, 1-17.
1936b. Studies of Peruvian Birds. XX. — American Museum Novitates
No. 861, June 22, 1936, 1-26.
ART. 5. SPHOENOGNATHUS CURVIPES (COLEOPTERA;
LUCANIDAE), A NEW SPECIES FROM BOLIVIA.
By Bernard Benesh
North Chicago, Illinois
During a visit to the Carnegie Museum, a small series of a Lucanid of
doubtful status attracted my attention. After further study, I am con-
I vinced that the insect is new to science, which I propose to describe in the
I subjoined short diagnosis, under the specific name
[ Sphoenognathus curvipes, sp. nov.
Male: subparallel, depressed, golden-brown, aeneous; head, pronotum,
and venter, with dense, fulvous pubescence.
I Head transverse-quadrate, broader than long, produced in front and
truncate, rugose, antero-laterad angles oblique, subacute, sides emargi-
I nate and narrowing gently to base; anteocular area elevated; from base
of the mandibles to vertex of the head, a broad V-shaped depression; eyes
large, convex, completely divided by the canthus. Mandibles longer than
the head, porrect, trigonate, incurved in apical third, acute, inner margin
with several unequal ill-defined teeth; upper ridge bending outward in
basal third, thence diagonally to exterior margin; inner margin and the
ridge darker (nearly black) ; inner area rugose, lateral granulate; basal half
(dorsad and ventrad) covered with long fulvous pubescence. Antennae
slender, piceous; scape club-shaped, slightly bent, nitid, posterad pube-
scent, distinctly shorter than the clava and twice as long as the funicle;
funicle of three segments, first segment subglobose (pear-shaped), second
similar and one quarter longer than the first, third regularly dilated from
base to apex and one-eighth shorter than the second; clava of six seg-
ments, rufous, subopaque, first five segments of equal length, anterad
lobate, terminal of irregular shape and twice as long as the preceding,
with anterior margin feebly lobed by a longitudinal impression; funicle
and bases of first three segments of clava nitid, sparsely setose.
Pronotum twice as broad as long, anterad sinuate, antero-laterad angles
; obtuse, sides diverging diagonally to center, center rounded to posterior,
I basal angles oblique-truncate, base nearly straight, margin elevated;
disc with a median longitudinal canaliculation, subrugose, glabrous,
antero-laterad area irregularly depressed and densely clothed with whorls
of fulvous pubescence; sides with a deep marginal pit.
I 45
ssued November 3, 1948.
46
Annals of the Carnegie Museum
VOL. 31
Scutellum broad, rounded. Elytra nearly parallel, broadest beyond the
middle, posterior regularly rounded, alutaceous, strongly aeneous, minutely
punctured with remote punctures (not discernible by naked eye), humeri
rounded, suture and declivity with sparse, short, grayish pubescence.
The space between the thorax and elytra covered by dense pubescence,
partially hiding the scutellum, which is destitute of clothing (in lindeni
pubescent.)
Beneath, rufous, with head, pro-, meso-, and meta-sternum covered
with long fulvous pubescence; abdominal segments granulose, with short
prostrate grayish pubescence.
Legs: femora stout, remotely punctured, posterior and lower areas
pubescent; tibiae strongly curved, anterior nude, intermediate and pos-
terior pubescent; anterior tibiae distad with broad furcation and an ill-
defined denticle near the center; intermediate with one small spine near
the apical fourth, posterior spineless; tarsi piceous, setose beneath; claws
large, simple.
Female: robust, convex, broader than the male, darker (chestnut-
brown to piceous), nitid, without the aeneous luster typical to the male;
antennae stouter, clava more compact than in the male, and as long as the
funicle and scape combined; elytra covered throughout with sparse gray-
ish pile; tibiae less arcuate and more massive, anteriors with a broad furca-
tion and two lateral subequal spines, intermediate with a single spine in
apical third (more pronounced than in the male), posterior unarmed.
Beneath, darker (nearly black), with pro-, meta-, and mesosternum
strongly tinted with green, aureous, femora purplish; pubescence less
dense and shorter.
Principal Dimensions
(In millimeters)
Length X Width
9
Head ' 3 x 5.9 2.0 x 5.75
Mandibles 3.5 2.25
Prothorax 5 xlO.O 5.25x11.0
Elytra 16 xl2.0 17.0 xl3.0
Types: cT and 9, Incachaca, Bolivia, alt. 2,500 M., Collector Jose
Steinbach, Acc. 6373, Collection of the Carnegie Museum.
Paratopotypes: cf* and 9 , in the writers collection and that of the Car-
negie Museum.
1948 Benesh: Sphoenognathus curvipes, a new species 47
Allied to 5. Undent Murray^ from which it may be readily separated
by the peculiar antennae, arcuate tibiae (in lindeni apparently doubly bent,
e.g., in-, ex-, and incurved), uniformly rounded elytra (in lindeni each
elytron is rounded) and other characters.
My thanks are due to Dr. A. Avinoff, and to Dr. George Wallace of the
Carnegie Museum, for the privilege of examining and reporting on these
insects, as well for the generous contribution to my reference collection of
the paratopotypes, which greatly enhance my meager series of neotropical
Lucanidae.
J Murray, Andrew: New Coleoptera from the Andes (Edinburgh New Phi-
losophical Journal), 1857, 5: 221-226, pi. 3, figs. 1, d^; 2, 9.
48
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 1
Sphoenognalhus curvipes, sp. ncv.
Fig. 1. Male. Head. 1. a antenna, 1. b anterior tibia, 1. c intermediate tibia, 1. d
posterioi tibia.
Fig. 2. Female. Head. 2. a antenna, 2. b anterior tibia, 2. c intermediate tibia,
2. d posterior tibia.
Fig. 3. Sphoenognalhus lindeni Murray. Tracing of Murray’s original figure,
enlarged 2 X. Anterior tibia (3), and intermediate tibia (3. a), d”.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 5
Plate 1
V y ART. 6. TAXONOMY AND DISTRIBUTION OF THE
^ : GENUS PIERELLA (LEPIDOPTERA)
r By F. Martin Brown^
Colorado Springs, Colorado
These large dark-winged satyrids are found in the deep tropical rain
forest. A few of the species stray into the subtropical forest but their true
home is hot and humid. There they flit along the trails and slip into the
protection of the thickets at the slightest alarm. Their lazy flight would
make them easy to capture in any other environment. When they settle,
the cryptic markings on the underside of the wings afford them ample
protection from all but the keenest eyes.
Genus Pierella, Westwood, 1851
The first member of the genus to be described was lena, by Linneaus in
1767. Since then over thirty forms have been named. The authors of the
earliest species included them in the genus Papilio. Huebner used Oreas
in 1807 and Godart used Satyrus in 1823. Neither of these names are
available. In the Genera of Diurnal Lepidoptera, 2:365, 1851, Westwood
proposed Pierella as a subgenus, “Section 2, Haetera Fabricius.” In this
he placed the species then known. Herrich-Schaeffer {Prodromus 1:55,
1864) was the first to use Pierella as the name of a full genus. He has been
followed by all except Hewitson, Erichson and Felder who rigidly adhered
to Haetera.
In 1868, Butler {Ent. Mo. Mag. 4:195) specified nereis Drury as the
type of Pierella. This was a particularly happy selection since that species
was figured by its author and thus there can be no question as to what the
name stands for.
^ During 1940 and 1941 I prepared several papers based upon my collections
made in Ecuador during 1938 and 1939. This series, of which this will be the sixth
paper, was interrupted by the war. Upon returning to my laboratory in the sum-
mer of 1946, I decided to continue with the Neotropical Satyrids but to enlarge the
scope of the papers. This is the first of the enlarged papers.
During the preparation of this paper the following men and museums have
given me advice and loaned me material: Mr. William P. Comstock and Dr.
Charles D. Michener at the American Museum of Natural History, Mr. Richard D.
Fox formerly at the Reading Public Museum and now at the Carnegie Museum,
the late Capt. Roswell C. William of the Academy of Natural Sciences, Phila-
delphia, and Dr. William C. Field at the United States National Museum. Without
their aid this study could not have been made.
49
Issued December 8, 1948.
50
Annals of the Carnegie Museum
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The genus Pierella may be described as follows:
Head: The antennae are rather stout with clubs only slightly heavier
than the shafts. They range in length from about two-thirds to the same
length as the cell of the forewing, and consist of 48 {rubecola) to 66 {nereis)
joints.
The eyes are glabrous.
The palpi are large: the distal joint the shortest, the middle joint the
longest {see fig. 1650 a).* In tzerm the proportions are about 23:110:40;
in hyalinus 20:108:37 ; and in lesbia 30:110:40.
Thorax: The male prothoracic legs have a single tarsal joint, a little
longer than either the tibia or femur {see fig. 1650 b). The female pro-
thoracic legs have a tarsus made up of five joints, the basal one being
longer than the combined length of the others {see fig. “9 lena”)- The
second most distad joint is the shortest. The mesothoracic and meta-
thoracic legs are perfectly normal in both sexes. The legs bear no spurs.
The tegulae are large. There is a long spur sweeping backward from the
body {see fig. 1651 c).
FEMALE
PROTHORACIC LEGS
Fig. 1. Appendages of head and thorax.
The venation of the forewings: The basal portions of Sc, Cu, and A2
are inflated. The radial is five branched; Ri and R2 arise before the
end of the cell. The origin of R2 is closer to that of Ri than to the end of
the cell. Ml, R3+4+5 and the UDC are conate or nearly so. The origin
of R3 is variable but usually about half-way from the end of the cell to the
apex. There is a spur on A2 that arises just beyond the end of the swelling
*The figure numbers used refer to the serial numbers of my dissections.
1948
Brown: Taxonomy of Pierella
51
and is sharply recurved. There are three strong recurrent veins in the cell.
Another trace lies between Cu2 and A2 {see fig. 1651 a).
The venation of the hindwings: The Sc and Rs are free at the base.
The Sc is sharply angled at the origin of H. The H is a short straight
spur. M3 and Cui branch from a common vein well beyond the end of the
cell. The UDC is shorter than the MDC which is about equal to the
weak LDC in length. The LDC meets the Cug just beyond the origin of
the Cu2 {see fig. 1651 b).
PIERELLA
VENATION
Fig. 2. Venation in Pierella,
The scalation of nereis is weak. In all other species it is normal.
Male Genitalia: The tegumen is subtriangular in shape, usually about
half the total dorso-ventral dimension of the genitalia in depth and vary-
ing from less than to more than that in anterior-posterior dimension.
The subuncal processes that are usually present on the Satyridae are
absent.
The uncus is simple, slender and curved, and from just short of to equal
to the anterior-posterior dimension of the tegumen.
The vinculum is slender (rather broad in nereis and hortona) and almost
as long as the depth of the tegumen.
The saccus is slender and about as long as to a little longer than the
vinculum.
The annuli are less than half as long as the vinculum and are lanceolate.
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Annals of the Carnegie Museum
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P LENA GLAUCOLENA R LENA LENA p HYCETA HYCETA
R HORTONA
R HYCETA LATONA
RALBOFASCIATA DECEPTA
Fig. 4. Male genitalia of Pierella,
P. lena glaucolena
P. lena lena
P. hyceta hyceta
P. amalia
P. hortona
P. hyceta latona
P. albofaciata decepta
P. hyalinus hyalinus
P. astyoche
1631 Rio Jondachi, E. Ecuador, 800 m., Nov. 1939, F.M.B.
1652 Bartica, British Guiana, A.M.N.H.
1755 Arima, Rio Purus, Brazil, November 1922, C.M.
1769 Macayacu, Colombia, March 1946, A.M.N .H .
1635 Rio Jondachi, E. Ecuador, Nov. 1939, F.M.B.
1633 Puyo, E. Ecuador, 1000 m., Dec. 1938, F.M.B.
1752 Rio Japacani, E. Bolivia, Sept. 1914, Paratype in C.M.
1653 Cayenne, French Guiana, December 1903, A.M.N. H.
1654 Kamadusa, British Guiana, October 1922, A.M.N.H.
Brown: Taxonomy of Pierella
53
R LUNA LUNA R LUNA RU3ECOLA R LUNA LESBIA
P. luna luna
P. luna rubecola
P. luna lesbia
P. rhea lamia
P. rhea columbina
P. rhea x P. hyceta ?
P. stollei stollei
P. lucia
P. boliviana
Fig. 5. Male genitalia of Pierella.
1656 Barro Colorado, Canal Zone, March 1926, A.M.N.H.
1657 Tezonapa, Vera Cruz, Mexico, A.M.N.H.
1632 No data, A.M.N.H.
1655 Para, Brazil, A.M.N.H.
1756 “Colombia,” A.M.N.H.
1749 Arima, Rio Purus, Brazil, September 1922, C.M.
1754 Munez Freire, Espiritu Santo, Brazil, C.M.
1630 Canelos, Eastern Ecuador, December 1938, F.M.B.
1750 Rio Japacani, Eastern Bolivia, August 1913, C.M.
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Annals of the Carnegie Museum
VOL. 31
Fig. 3. Male genitalia of Pierella.
P. helvina incanescens 1649 “Guatemala,” A.M.N.H,
P. nereis 1651 “Brazil,” A.M.N.H.
Note: The four digit number following the name is the serial number of the
dissection and is to be found on both the specimen and the mounted
dissection. These are deposited at the museum indicated by the initials
following the locality data.
The valves are lobate to subtriangular. They are always armed with a
heavy distal spine (digitate process in helvina). The dorsal margin may
be simple or toothed. A sub-dorsal comb frequently is present (absent in
luna, nereis, and hortona; most highly developed in incanescens). The
valves show good specific characters.
The aedaeagus is more or less straight and heavy. It is of uniform di-
ameter except in hortona where it is markedly tapered. It is variously
armed with cornuti which seem to be torn loose during copulation. This
organ is of some use for separating species.
Remarks
In recent years the named forms of Pierella have been distributed
among twelve or thirteen “species.” I am inclined to believe that there
are fewer biologic species than this in the genus. Until the life histories
of the various named forms that compose the bulk of this genus are known,
the true number of species will remain in doubt. The male genitalia
indicate that there are either several groups of very closely related species
or several species that have developed ecological subspecies. This is
1948
Brown: Taxonomy of Pierella
55
particularly true of those members of the genus found in the Amazon
basin.
The development of the pattern on the wings has been fully studied and
discussed by Schwanwitsch {Zeit. Wiss. Biol., Abt. A., 10:433-532, 1928),
On the basis of these studies those species included in Division B in this
paper probably represent the most primitive forms. The present dis-
tribution of the various forms within the genus does not wholly support
this thesis if primitive forms are considered to be peripheral. The present
distribution seems to indicate that the genus antedates the Andean up-
lift and that at that time the genus ranged across the area affected. Those
species now found west and north of the Andes in South America are ap-
parently derived from the Central American progenitors. There are no
members of the genus found in the West Indies^ (excepting Trinidad).
The following key, based on color pattern refers to acceptable taxo-
nomic species.
A KEY TO THE SPECIES OF PlERELLA
Reference is made in this key only to species. Methods for separating the sub-
species and forms will be found in the discussion of each species in the genus,
la. Underside: a dark-bordered light transverse band on Both Wings
3. Division A (p. 56)
b. : three dark transverse lines on the Hindwing 2.
2a. Upperside: forewing with a blue spot or a white spot on the discocellulars.
12. Division C (p. 80)
b. : no such band or spot 9. Division B (p. 70)
Division A
3a. Upperside:
b. :
4a. Upperside:
b. :
5a. Hindwing:
b. :
6a, Upperside:
b. :
distal portion of the hindwing with an ochraceous area.
nereis (p. 57)
not so 4.
hindwings with red markings helvina (p. 57)
no red markings or at most only a rusty flush on the hindwings. 5.
the margin of the wing deeply excavated anterior to M3.
hyalinus (p. 62)
lacking this excavation 6.
a large white spot at or near the margin of the hindwing 7.
no such spot, at most only small interneural white spots 8.
2 Series of P. hyalinus, lena, and lamia, in the Mengel Collection at the Read-
ing (Pennsylvania) Public Library and Museum, are ticketed “Bath, Jamaica.”
The Mengel Collection contains many specimens with obviously incorrect data so
I do not consider these specimens as West Indian in origin.
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Annals of the Carnegie Museum
VOL. 31
7a. Upperside:
b. :
8a. Upperside:
b. :
white spot on hindwing touches or almost touches the margin.
lucia (p. 69)
the white spot separated from the margin by about K the
diameter of the spot amalia (p. 68)
three transverse rows of blue spots, sometimes white centered or
whitish on the hindwing lena (p. 65)
no such series astyoche (p. 69)
Division B
9a. Upperside: two large black submarginal ocelli at the inner angle of the
hindwing (rarely up to five, then the specimen is from northern
Central America) luna (p. 78)
b. : four or five submarginal ocelli on the hindwing (rarely three,
then from S. Brazil or Bolivia) 10.
10a. Upperside: hindwing with large ochraceous or reddish brown areas.
hyceta (p. 76)
b. : no such areas 11.
11a. Upperside: basic color greyish faun stollei (p. 73)
b. : basic color dark grey brown or blackish rhea (p. 71)
Division C
12a. Underside: transverse light band on the forewings crossed by a similar
band at the discocellulars albofaciata (p. 82)
b. : transverse light band only slightly enlarged at discocellulars.
hortona (p. 80)
Division A
Characteristics: There is a distinct, narrow, light band with dark mar-
ginal lines across the underside of both wings. A third dark line crosses
both wings basad of the light band.
I. The nereis group.
Characteristics: There is a brightly colored, transverse band across the
limbal area of the upperside of the hindwings.
This group contains two species, nereis the genotype, and helvina. The
former has a bright ochraceous patch on the upperside of the hindwings,
the latter a red patch. The two species are strictly congeneric. No
other species in the genus is so closely related to the genotype as is helvina.
These species occupy widely separate ranges. P. nereis is restricted to
southeast Brazil and helvina to Central America and that part of South
America west and north of the Andes.
1948
Brown: Taxonomy of Pierella
57
1. Pierrella nereis (Drury)
Original description: 1782, Illustr. Exot. Ent., pi. 35, f. 2, 3.
Earliest figure: in original description.
Other figures:'^ 1835, Lucas, Lepid. Exot., pi. 8a, f. 1. 1911, Weymer,
Seitz's Macrolepid., 5: pi. 42, line d. ^
Material seen: DcTcf, 39 9.
Range: S.E. Brazil in the rain forest.
This species is distinctive and quite constant in its appearance. No-
where does it seem to be common. Most of the specimens seen came from
Petropolis, Sta. Caterina, and Rio de Janeiro.
2. Pierella helvina
This is a plastic species. It is quite probable that no two local popula-
tions are alike. This poses a taxonomic quandary. To dignify each of
those populations with a varietal name would obscure the apparent de-
velopment that is in progress in the species. If and when the genetics of
the species are studied, there will be time enough for further additions to
the nomenclature. There are three reasonably well established foci of
development, each with a basic form distinguishable from the other two.
These occupy different geographic areas butareonly partially isolated. The
numbers of specimens seen, from a large number of stations, have been
by no means sufficient to do more than sketch the relationships of the
forms. Unfortunately the species seems to center in the worst possible
areas as far as disease and the ease of travel is concerned. At least one
collector has lost his life in search of butterflies in those regions.
The species has not been found in the basins of the Orinoco and Amazon
Rivers. Typical helvina helvina is found in northern Colombia. West of
the Andes, from the Rio Guayas in Ecuador to the Canal Zone, the race
found is helvina ocreata. The Central American form is helvina incanescens.
The latter is considered a full species by Weymer (Seitz's Macrolepid.,
5:176-179, 1911) and Gaede (Lepid. Cat., 29:422, 1931). Of these sub-
species, ocreata is the most distinctive and the most variable.
The subspecies may be separated on the following bases:
a. Those specimens having considerable white on the colored band
across the upperside of the hindwings are ocreata;
b. those specimens with no white in the colored band on the hind-
wings, with the red extending to or about to the costal margin
® All species are figured in Schwanwitsch, Zeit. Biol., Abt. A., 10:433-532.
1928. No specific citations are made to these.
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and with little more than a trace of white submarginal spots in
M2-M3 and M-3-Cui are helvina;
c. those specimens with no white on the colored band on the hind-
wings, with the red usually extending no further toward the costal
margin than Mi and with well-developed submarginal spots in
M2-M3, M3-CU1 and occasionally in Cui-Cu2 are incanescens.
2a. Pierella helvina helvina (Hewitson)
Original description: 1860, Exot. Butt., 2: Haetera, pi. 1, f. 4.
Earliest figure: in original description.
Other figures: 1911, Weymer, Seitz's Macrolepid., 5: pi. 42, line d.
Material seen: 14 cf’cf , 10 9 9.
Range: In tropical rain forests of northern Colombia, especially east
of the Rio Cauca, and west of the Cordillera Oriental.
P. h. helvina is the most stable of the three races of the species. The
forewing generally exhibits an ocellus in M1-M2, and white spot in R5-M1,
M2-M3 and M3-CU1; the latter mark is the least constant. The hindwing
bears an ocellus in M1-M2, and white spots in Rg-Mi and traces of white
spots in M2-M3 and M3-CU1. The extent of the red on the hindwings is
somewhat variable, on the majority of specimens it reaches or almost
reaches the costal margin. On a few specimens it does not quite reach Mi
and in this respect these specimens resemble incanescens.
2b. Pierella helvina incanescens Godman & Salvin
Original description: 1877, Proc. Zool. Soc. London, p. 61.
Earliest figure: 1880'^, Godman & Salvin, Biol. Cent.-Amer., Rhop., 3:
pi. 6, f. 5, 6.
Other figures: 1911, Weymer, Seitz's Macrolepid., 5: pi. 42, line d. 1925,
Schwanwitsch, Entomologist, 58: 266.
Material seen: 2S S' cL , 11 9 9.
Range: In the tropical rain forests of Central America from Guatemala
to Chiriqui, Panama.
This subspecies is a little more variable than h. helvina. As has been
pointed out, some authors take the position that it is a separate species
from helvina. I can find no support for this view. The anatomy of the
^ Dates used for Cramer, Pap. Exot. and Godman & Salvin, Biologia, are ac-
cording to Brown: Ann. Ent. Soc. Amer., 34:127-138, 1941.
1948
Brown; Taxonomy of Pierella
59
two forms clearly indicate a single species and the wing pattern of each
varies toward that of the other.
The forewing of incanescens is marked like that of helvina. The hind-
wing exhibits a tendency toward a well-developed submarginal series of
white spots. Only traces of this series are found on helvina. The red on
the hindwing is expanded toward the inner margin. As a result of this
expansion a black line perpendicular to the inner margin appears in the
red between Cu2 and Ai. This line is rarely absent or obscure on incan-
escens and rarely visible on helvina.
A pair of unusually small specimens of this race was collected by Schaus
on the Rio Sixaola in south-eastern Costa Rica near the Panama border.
The red on the hindwings is limited anteriorly as on incanescens and
posteriorly as on helvina. Such limitation is more like the pattern of
helvina than incanescens. The submarginal row of white spots is well de-
veloped and places the specimens in the subspecies incanescens. Curiously
the male displays a white spot in Cui-Cu2, usually found only on females
and the female lacks this spot!
These two specimens might be included under Niepelt’s form costaricana
{Ink Ent. Zeit., 21:50, 1927) described from Turrialba, C.R. This name
was applied to specimens that resemble helvina in the extent of the red
markings on the hindwings. The types and all specimens that I have
seen come from the valley of the Rio Reventazon where they fly with
typical incanescens.
2 c. Pierella helvina ocreata God man & Salvin
Original description: 1868, Ann. & Mag. Nat. Hist., (4) 2:143.
Earliest figure: 1880, Godman & Salvin, Biologia Cent.-Amer., Rhop.,
3: pi. 6, f. 7-9.
Material seen: 3 cfcT, 6 9 9.
Range: In the tropical rain forests from the Canal Zone eastward
through the Isthmus of Darien and then south along the coast to the Rio
Guayas, Ecuador, west of the Cordillera Occidental.
This race is highly variable. It may be recognized always by the white
area in the colored band on the upper-side of the hindwings. The other
markings are much like those found on helvina and incanescens.
The males that I have seen are all alike, the females are each different
from the others. These represent various degrees and combinations of
three lines of variation :
a. replacement of the ocellus on the forewings by a white spot;
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b. a color change of the submarginal spots from white to red;
c. restriction of the red area in the colored band on the hindwing.
This tendency to vary has lead so far to the proposal of four varietal
names. Without doubt at least as many more will be proposed! The
Panamanian specimens that I have seen are reasonably constant. I am
inclined to believe that the race has invaded the South American part of
its range in recent geologic times and is responding to new environmental
pressures. I believe that every female from the Pacific coast of Colombia
that has reached Germany has been dubbed with a separate varietal
name!
The currently named varieties are these:
9 f. hymettia Staudinger, {Exot. Tagf., 1:220, 2: pi. 77, 1888) from
“Choco, Colombia” represents all three changes noted above. It is the
most divergent from the “normal” of all of the so-far named varieties.
9 f. werneri Hering and Hopp, (Iris, 39:191, 1925) from Rio Micay,
Choco, Colombia, and 9 f. johnsoni Talbot, {Bull. Hill Mus., 4:196,
1930-32) from Torqueral, N. Colombia, demonstrate the reduction of red
on the hindwings. On werneri the white extends posteriorly to M2; on
johnsoni beyond that nervule. These two names should be considered
synonyms.
f. pacifica Niepelt, {Int. Ent. Zeit., 17:137, 1924) from Pacifico, S.
Colombia, partially demonstrates the change from white to red sub-
marginal spots. Some of the spots are red, and some red, centered with
white.
For the benefit of those with limited material, the following analysis
of minor variations among the three races is presented:
a. On the forewing there is an apical submarginal row of spots,
usually four in number. The spot in M1-M2 is variable:
1. on helvina it is small, black, and surrounded by a narrow
brown iris;
2. on incanescens it is large, black, and surrounded by a narrow
grey iris;
3. on ocreata it is variable in size, black and surrounded by a
narrow brownish grey iris. Among the females there is a
tendency toward the development of a white spot rather than
an ocellus. One in the U. S. National Museum has a normal
spot on the right wing and just a few white scales on the left
wing. Forms hymettia, werneri, and pacifica, each has a white
spot in this position.
1948
Brown: Taxonomy of Pierella
61
b. On the hindwings there always are at least two spots flanking
the ocellus:
1. on helvina the posterior of these two spots is obsolescent;
2. on incanescens there are four or five spots, two anterior to and
two or three posterior to the ocellus;
3. on ocreata there are always two and often three spots: The
variable spot is in M2-M3.
c. Some forms have the posterior part of the red on the hindwing
extended and show a black line in this red:
1. on helvina and ocreata this line, when present, extends from A
to Cui;
2. on incanescens this line is almost always present and extends
from A to Cu2 and thence along that nervule to the margin.
d. The anterior extension of the red on the hindwing is variable:
1. on helvina the red usually extends to the costal margin, where
it is heavily dusted with dark brown scales: there is no white
in the band ;
2. on incanescens the anterior margin of the red varies between
M2 and Rs: there is no white in the band;
3. on ocrm/a there is white anterior to the red : the boundary for
the two colors lies between Mi and Cui: it lies near Mi on
hymettia and pacifica, near M2 on werneri, near M3 on ocreata
and near Cui on johnsoni: the anterior margin of the white is
usually the costa where it is obscured by brown scales.
e. The posterior terminus of the red on the hindwing is,
1. on helvina and ocreata anterior to A, occasionally at Cu2, and
the inner margin of the colored band lies at or close to the end
of the cell ;
2. on incanescens almost always at A, and the inner margin is
well outside the cell.
IL The hyalinus group.
Characteristics: The upperside of the hindwings have a submarginal
series of spots or ocelli and lack a brightly colored transverse band.
Weymer in Seitz's Macrolepidoptera (5:177, 1911) and Gaide in Lepi-
dopterorum Catalogus (29: 421-423, 1931) recognize only three taxonomic
species in this group. There are really five co-equal taxonomic units
when the anatomy of the forms is considered. All of these are confined to
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the tropical rain forests east of the Andes. They are rarely found above
1000m. On the basis of the collections thus far made, the center for these
species is the middle Amazon basin. In the triangle between Manaos,
Brazil; Iquitos, Peru; and Leticia, Colombia; all five species are found.
3. Pierella hyalinus
This species is usually cited under Huebner’s name, dracontis. It forms
a rather weak connecting-link between the werm-group and this group of
species. It is easily recognized by its abberant wing-form. The margin
of the hind-wing anterior to Ms is “excavated.” This “excavation” is an
exaggeration of the angular shape of the hindwing of helvina. There are
two subspecies found on the continent of South America. Typical ex-
amples of these are readily identified but there is a broad zone where the
two meet in which all sorts of intermediates fly. A third subspecies is
1948
Brown: Taxonomy of Pierella
63
confined to the island of Trinidad and is a continuation of the variation
shown by the more northeastern mainland race.
The three races are recognizable by the following characteristics:
1. hyalinus has light, metallic blue spots on the upperside of the
hindwings. Several of these spots are fused to adjacent spots in
the same series on the males. Such fusion occurs much less fre-
quently among the females.
2. dracontis has deep, blue spots on the upperside of the hindwings.
These spots are well defined and separate. The ground color of
the wings tends to be much darker on this race than in either of the
others.
3. fusimaculata has light, almost silvery, blue spots on the upperside
of the hindwings. These spots are frequently fused and the three
series in turn tend to fuse. Fusion reaches a higher degree among
the females than among the males. This is contrary to the con-
dition in hyalinus.
A short series of specimens of h. hyalinus in the Academy of Natural
Sciences at Philadelphia have labels “San Juan Evangelista, Vera Cruz,
Mexico.” I have examined numerous collections made in the state of
Vera Cruz, some from San Juan Evangelista and have not seen this species
among them. Hoffman’s list of Mexican Butterflies {Annales Inst. Biol.,
11:666, 1940) does not mention the species from Mexico. Godman and
Salvin in the Biologia do not record the species from anywhere in Central
America.
3a. Pierella hyalinus hyalinus (Gmelin)
Original description: 1788-91, Syst. Nat., I, 5:2259.
Earliest figure: 1780, Cramer, Pap. Exot., 4:5, pi. 291 A, B (9 ) as lena.
Other figures: (1813) Huebner, Samm. Exot. Schmett., pi. 83 ( 9 ) as lena.
1835, Lucas, Hist. Nat. Lepid. Exot., pi. 79, f. 3 as lena.
Material seen: 32 cfcT, 11 9 9.
Range: The rain forests of the three Guianas, Venezuela primarily in
the Orinoco basin, and adjacent parts of N.E. Brazil, in the region north
of the Amazon.
This form has long been considered among nomina incognita. Masson
and Weymer used it in Lepidopteren Stuhel's Reise, p. 87, for h. dracontis
and Kaye used it in Trans. Ent. Soc. Bond., 1904, p. 179, for the specimens
he had from Trinidad. Kaye’s use of the name for what I csAl fusimaculata
may invalidate my name in the view of some taxonomists. However,
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Annals of the Carnegie Museum
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Gmelin’s specimens in all probability come from the Guianas and it is for
that form of the species that I retain hyalinus.
The Orinocan-Guiana specimens of the species are recognizable by the
lighter color of the blue spots on the upperside of the hindwings and by
the tendency toward fusion of these spots on the males. The tendency
toward fusion is greatly reduced among the females of hyalinus. The few
specimens that I have seen from the coastal area of Brazil, north of the
Amazon, are intermediate to hyalimis and dracontis.
3b. Pierella hyalinus dracontis Huebner
Original description: 1816, Verz. bek. Schmett., p. 53.
Earliest Figure: (1851) Westwood. Gen. Diurn Lepid., 2: p. 62, f. 2 as
lena.
Other figures: 1888, Staudinger. Exot. Tagf. 2, pi. 77.
Material seen: 14 cf cT', 7 9 9.
Range: Amazon basin, especially south of the Amazon River and west
of the Rio Negro. Very rarely found upstream from the “fall line.” The
westernmost specimens that I have seen with wholly reliable data come
from Leticia, Colombia.
The race is darker and has smaller, better-defined, blue spots on the
hindwings than either of the other races. The maximum reduction in size
of the blue spots occurs around Sao Paulo de Olivenga in the south-
western part of the range of the race. A female, with no definite locality,
in the collection of the Carnegie Museum has the limbal and discal row
of spots on the hindwing white instead of blue.
Weymer {Seitz's Macrolepidoptera, 5.T77, 1911) validated the Staud-
inger manuscript name extincta for a minor variation of this race. The
name applies to those specimens on the underside of which the trans-
verse light band of the forewing is broken into two or more fragments.
3c. Pierella hyalinus fusimaculata, subsp. nov.
This race from the island of Trinidad is more closely related to hyalinus
than to dracontis. The upperside of the hindwings of the species exhibits
three rows of pale blue spots, submarginal, limbal, and discal. On typical
dracontis the spots composing each row and the rows themselves are clearly
defined and isolated. On hyalinus the spots are larger and less well-
defined than in dracontis. There is a tendency toward fusion of the spots
in each row, especially in the males. On fusimaculata the spots of each
1948
Brown: Taxonomy of Pierella
65
row, except the terminal spots, are fused and on many specimens, espe-
cially females, the rows of spots are partially fused. Some specimens
demonstrate this to so high a degree that the hindwing appears to be
silvery blue with a dark base. On the most completely fused specimens of
hyalinus that I have seen, there is a large dark area between the limbal and
discal rows between M3 and Cu2. On fusimaculata this dark area is re-
duced to two interneural spots, M3-CU1, and Cui-Cu2.
Holotype: cf , Hololo Rd., St. Ann’s, Trinidad, Aug. 16, 1933 (Pinckus),
Carnegie Museum.
Allotype: 9 , Arima, Trinidad, (E.W. Rorer), U. S. Nat. Mus.
Paratypes: 1. cf , same data as holotype; 2. 9 , same data as holotype;
3. 9 , same locality as holotype, Aug. 12, 1933; 4. 9 , Tacariqua, Trinidad,
XL. 32 (Katwara) Carnegie Museum; 5. 9 , Carenage, Trinidad, Aug. 12,
1909 (Carricker), Carnegie Museum; 6. c^, St. Ann’s, Trinidad, Feb. 28,
1933 (Pinckus), Carnegie Museum; 7. 9 , St. Ann’s, Trinidad, March 7,
1933 (Pinckus), Carnegie Museum; 8. 9 , St. Ann’s, Trinidad, March 4,
1933 (Pinckus), Carnegie Museum; 9. 9 , Lady Charwell’s Rd., St. Ann’s,
Trinidad, April 21, 1933 (Pinckus), Carnegie Museum; 10. 9 , same as 9,
March 4, 1933; 11. cf, Carenage Trinidad, Aug. 14, 1909 (Carricker), Car-
negie Museum; 12. cT, same as 11, Aug. 18. 1909; 13. 9 , same as 12, Aug.
18, 1909; 14. 9, Port of Spain, Trinidad LV. 33 (Pinckus), Carnegie
Museum; 15. 9, Mt. Taber, Trinidad, (Netting) Carnegie Museum; 16.
9 , Tondes Amande Rd., St. Ann’s, Trinidad, March 12, 1933 (Pinckus),
Carnegie Museum; 17. cf, Sta. Cruz Valley, Trinidad, March 29, 1929
(Huntington), Am. Mus. Nat. Hist.; 18. 9, Trinidad, Reading Public
Museum.
4. Pierella lena
This species and the preceding one were confused for many years by the
older students of butterflies. Cramer, Huebner, Lucas, and Hewitson, all
figured hyalinus forms as lena. Beginning with Hewitson’s Exotic Butter-
flies and Butler’s “catalogues” the species has been properly recognized.
The species resembles hyalinus in pattern but the hindwing is rounded and
not excavated as is that of hyalinus.
The species flies throughout the tropical rain forest area of South
America, east of the Andes. There are two well-defined races that,
anatomically at least, are approaching independence as taxonomic species.
One of these, glaucolena, produces a form, brasilensis, that approaches the
other, lena. Unfortunately the name brasilensis antedates glaucolena.
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Annals of the Carnegie Museum
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A fourth named form, amalia, has been considered a member of this
species-complex. I believe this form to have equal claim with lena to
specific standing and treat it as such.
The two races differ from each other in the blue spots on the hindwings.
These are white-centered on lena and solid blue on typical glaucolena.
4a. Pierella lena lena (Linnaeus)
Original description: 1767, Syst. Nat., XII:487.
Earliest figure: 1779, Cramer, Pap. Exot., 3: pi. 198 f, D. E.
Other figures: 1911, Weymer, Seitz's Macrolepid., 5; pi. 42, line e.
Material seen: 53 crcT, 49 9 9-
Range: Tropical rain forests of lower Rio Orinoco, the Guiana and the
eastern half of the Amazon basin, westward to around Santarem.
Specimens from Mengel’s collection in the Reading Public Museum are
labelled from “Muzo, Colombia,” “Bath, Jamaica,” and “Port-of-Spain,
Trinidad.” The Jamaican data is certainly incorrect and the other two
localities are very doubtful.
This race is reasonably constant. About fifty percent of the specimens
seen bear on the forewings a black dot in M1-M2 flanked by white dots in
the adjoining interspaces. This marking is constant among the females
and occurs occasionally on the males. The white spots composing the
inner two rows of spots on the hindwings are very rarely circled with
blue. The sex patch on the inner margin of the hindwings of the males
is almost invariably wholly yellow in lena lena.
4b. Pierella lena brasilensis (Felder)
Original description: 1862, Wein. Ent. Mon., 6:126. There is no pub-
lished figure of this form.
Material seen: 30 cf cJ', 12 9 9 .
f. glaucolena Weymer
Original description: 1911, Seitz's Macrolepid., 5:177.
Earliest figure: 1911, Weymer, l.c., pi. 42, line e.
Material seen: 47 cTcf, 27 9 9.
Range: P. brasilensis is found in the rain forests of the southwestern
part of the Amazon basin extending into Bolivia, usually not found above
600m. The form glaucolena has a greater range northward into Colombia
and ranges higher into the Andean foothills to a little above 1000m.
1948
Brown: Taxonomy of Pierella
67
It is unfortunate that the name hrasilensis antedates and must take
precedence over glaucolena as the name of the race. The latter insect is
the more typical of the two forms and has by far the greater range.
The decoration of the upperside of the hindwings of hrasilensis ap-
proaches that of lena. However the “white spots” on hrasilensis are
better described as blue spots with white centers while those on lena
rarely show the slightest trace of blue at the periphery. The black dot
practically universal in its occurrence in M1-M2 on the forewing of female
lena and less frequently on males, occurs only cocasionally on hrasilensis
and then only on females. On glaucolena this spot is invariably white on
the males and usually so on the females. The flanking dots of white found
on lena are absent or obsolete on glaucolena. The male sex patch on
glaucolena is frequently dark brown with a yellow border. This condition
is more often found on specimens from the eastern part of the range than
from the western. On these the inner and outer zones on the underside
are more nearly concolorous.
The form hrasilensis is further differentiated from lena by often having
a powdering of blue scales at the base of the hindwings. There are oc-
casionally small interneural marginal spots on the forewings of the females.
The underside of hrasilensis is intermediate to lena and glaucolena, less
contrasting than the former and more so than the latter.
The three forms fly together in the vicinity of Aria on the Rio Purus in
the southern drainage of the Amazon.
A striking variant of glaucolena occurs across the central part of the
range of this form.
f. obsoleta forma nova
This is a form in which the tendency of form glaucolena is carried al-
most to completion. The blue spots on the upper-side of the hindwings
are obsolescent. On the holotype the marginal and submarginal rows of
spots are lacking and those composing the limbal and discal rows of spots
are reduced to a few bluish purple scales. The basic color of the wings is
darker than usual. On the underside the markings are partially obscured
by dark scales.
Holotype: cf , Nova Olinda, Rio Purus, Brazil, May 1922 (S. M. Klages),
Carnegie Museum.
Three other specimens before me approach this form. Two are from the
collections of the Carnegie Museum, one purchased by Holland from
Staudinger in 1885 and the other caught by Klages at Sao Paulo de
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Olivenga in January, 1923. The other is in my own collection. Mac-
Intyre caught it on Dec. 12, 1938, at Canelos on the Rio Bobonaza in
eastern Ecuador. On these the marginal series of spots is missing, the
submarginal series faintly present, the limbal spots reduced and the discal
spots obsolescent. The under surfaces are much more normal, being only
a little darker than is usual in glaucolena.
Fig. 7. Ranges of Pierella lena and amalia.
5. Pierella amalia Weymer
Original description: 1885, Stettiner Ent. Zeit., 46:285.
Earliest figure: 1860, Hewitson, Exot. Butt.., 2: Haetera 1, f. 2. as lena
var.
Other figures: 1888, Staudinger, Exot. Tagf., 2: pi. 77 as leucospila.
Also in volume 1 :220.
Material seen: 1 cf, 1 9 .
Range: The tropical rain forest on the upper Amazon. All specimens
have come from the region west of Pebas and north of the Rio Marahon.
It is probable that this rare insect is not a race or form of lena as it
has been considered ever since Hewitson first figured it in 1860. Its
restricted range falls within the range of the western races of lena. It has
1948
Brown: Taxonomy of Pierella
69
been taken in company with glaucolena. I believe that amalia is a ‘Re-
cently” established species formed from a mutant of ancestral lena.
The male genitalia are very much like those of lena. The most notice-
able differences are at the distal end of the valve. This area is truncate
with a strongly toothed margin in amalia. The figures show this and the
other more subtle differences.
P. amalia and the following species, lucia, resemble each other in having
a large white patch on the outer half of the upperside of the hindwing.
This patch is well basad of the outer margin on amalia and touches or al-
most touches the outer margin of the wing on lucia.
6. Pierella lucia Weymer
Original description: 1885, Stettiner Ent. Zeit.^ 46:285.
Earliest figure: 1860, Hewitson Exot. Butt., 2: Haetera 1, f. 5. as astyoche
var.
Other figures: 1888, Staudinger, Exot. Tagf., 2: pi. 77, as albomaculata
(also in volume 1:129). 1911, Weymer, Seitz's Macrolepid., 5: pi. 42,
line e.
Material seen: 36 cf cf, 15 9 9 .
Range: The tropical rain forests of the upper Amazon, westward from
Pebas into S.E. Colombia, E. Ecuador, and N.E. Peru, rarely above 700 m
altitude.
Previous students of this genus have followed Hewitson and considered
lucia to be a race or form of astyoche. It is sufficiently different in pattern
and anatomy to rank as a taxonomic species in this genus.
This species seems to be much more abundant south of the Rio Mar-
anon than north of the river. This is probably an illusion caused by the
wealth of material in the Bassler collections at the American Museum of
Natural History. The zone in which this species flies is in general very
poorly represented in collections. Every large collection that I have ex-
amined from within the range of this species contains specimens of lucia.
7. Pierella astyoche (Erichson)
Original description: 1848, in Schomburgk’s Reisen Br. Guayana, 3:599.
Earliest figure: 1851, Doubleday and Hewitson, Gen. Diurn. Lepid:
pi. 62, f. 1, as larymna (also on p. 365).
Other figures: 1911, Weymer, in Seitz's Macrolepid., 5: pi. 42, line e.
Material seen: 22 cT'cT, 28 9 9 (and material in U. S. Nat. Mus.).
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Range: Wherever there is tropical rain forest, in the whole of the
Amazon basin below 700m., and in the Guianas.
There is very little variation in this species. That which does occur is
of very minor importance. The three apical spots on the upperside of the
forewing all tend to be white on Guiana specimens while the majority of
the Amazonian specimens have a small ocellus in one of the interspaces.
Specimens from around Sao Paulo de Olivenga, Brazil, at Arima on the
Rio Purus, and on the lower Rio Ucayali in Peru, are darker than usual
and the white spots on the hindwings are reduced, occasionally they are
obsolete.
The bluish sheen seen on the hindwings in a certain light is more com-
mon on Guiana and upper Amazon material than on lower Amazon speci-
mens.
Fig. 8. Ranges of Pierella lucia and astyoche.
Division B
Characteristics: The undersides of both wings are marked with three
almost parallel straight dark lines. There is no marked dilution of the
base color between the outer two lines as there is on species of Division A.
1948
Brown: Taxonomy of Pierella
71
III. The rhea group.
Characteristics: the same as for the division.
The species of this group have the greatest range of the various groups.
Schwanwitsch considers the patterns of these species to be the most
primitive. These two items suggest that the species of the rhea group may
represent the stock from which the genus has developed. One species,
luna, is found from Mexico south into Ecuador and eastward to the
Guianas. The others are all restricted to the region east of the Andes.
Except in the cases of rhea, lamia, and chalybaea, I agree with Weymer
in his grouping the forms in species-complexes. I believe that the three
forms named are members of a single species. This taxonomic species,
rhea, is very confusing and probably deserves more thorough study than
I have given it. I did not see enough material from the eastern part of the
range.
8. Pierella rhea
Five names have been applied to forms of this species. Only one of
these names is a strict synonym, dindymene Cramer, 1779, is the same in-
sect as lamia Sulzer, 1776.
The characteristics that I have selected to use for separating the four
named forms are these:
a. rhea has the uppersides of the forewings and hindwings more or
less concolorous and no steely blue reflections on the hindwings.
There are well-developed ocelli on the hindwings.
b. lamia has the hindwings darker than the forewings, especially
toward the outer margin and there is usually a steely blue reflection on
the basal part of the hindwings. There are well developed ocelli on the
hindwings.
c. chalybaea is like lamia but the steely blue reflections cover almost
the entire hindwings and the submarginal ocelli appear as white dots, the
iris being obscured by the dark margin.
d. columbina is like chalybaea but generally lacks the broadly de-
veloped steely blue gloss.
I have found no populations that are really pure. The tendency is for
Guiana material to be lamia, lower Amazonian, especially north of the
river to be rhea. Middle Amazonian, especially south of the river to be
chalybaea, and material from above “fall-line” from Colombia to Bolivia
to be columbina. I have a feeling that rhea and lamia are forms of one
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subspecies or two poorly-differentiated ones and that the same is true of
chalyhaea and columbina. I shall treat these forms as such until proven
wrong.
There is a specimen in the collection of the Carnegie Museum that is
very puzzling. It was taken by S. M. Klages at Arima on the Rio Purus
in west-central Brazil in November, 1922, It looks like a rhea4orm with
a flush of dull reddish brown on the upperside of the hind wings. It appears
as one might expect a natural hybrid between rhea and hyceta. The male
genitalia bears on the dorsal margin of the valves a strongly developed
comb as in rhea. The valves themselves are much more ‘Rectangular”
than in any rhea specimens that I have examined. Some future worker
with more material than I have seen will have to decide what to do
taxonomically with this specimen.
8a. Pierella rhea rhea (Fabricius)
Original description: 1775, Syst. Ent., p. 467.
Earliest figure: 1911, Weymer, Seitz's Macrolepid., 5; pi. 43, line a.
Material seen: 4 , 1 9 .
Range: The tropical rain forests of the Amazon east of Santarem,
Brazil.
f. lamia (Sulzer)
Original description: 1776, Gesch. Ins., pi. 18, f. 1.
Earliest figure: in original description.
Other figures: 1779, Cramer, Pap. Exot., 3: pi. 198. f. F, G as dindymene,
1911, Weymer, Seitz's Macrolepid., 5: pi. 43, line a.
Material seen: 8 cTcf , 7 9 9.
Range: The tropical rain forests of the Orinoco basin, the Guianas, and
the northern part of the Amazon basin.
This race is characterized by having distinct ocelli on the hindwings.
These are white pupilled and have black irises. The hindwings of f. lamia
are darkest on the margin but not so deeply colored as to obscure the iris
of the ocelli. This form differs from rhea in having a steely blue gloss on
the hindwings, at least at the base.
There are specimens of lamia in the Mengel collection labeled “Bath,
Jamaica,” “Cristobal Colon, Panama” and “Coachi, Colombia.” The
Jamaica and Panama data I consider false, the Colombian record very
doubtful.
1948
Brown: Taxonomy of Pierella
73
8b. Pierella rhea chalybaea Godman
Original description: 1905, Trans. Ent. Soc. Land., p. 185. There are no
published figures.
f. columbina Krueger
Original description: 1925, Ent. Rundschau, 42: 17.
Material seen: chalybaea 18 cfcT, 7 9 9; intermediate 31 cfcf , 12 9 9 ;
columbina 6 cf cf , 2 9 9-
Range: The tropical rain forests of the southern and western Amazon
basin.
These two forms blend so completely that the majority of the speci-
mens that I have seen can be placed in either category. Generally speci-
mens from east of the Rio Huallaga are closer to chalbyaea than to col-
umbina. They show more of the blue gloss on the hindwings the farther
east the habitat. Those from west of the Rio Huallaga progressively ap-
proach close to columbina the higher the altitude of the habitat. I have
seen good columbina from Colombia, Ecuador, Peru, and Bolivia, and good
chalybaea only from Brazil and Peru.
The blue gloss on typical chalybaea is almost silvery. The gloss when
noticeable on columbina has purplish lights. Most specimens of columbina
lack some of the ocelli toward the anal angle and the iris are totally
obscured on those present. Most specimens of chalybaea show all of the
ocelli and there is usually a faint trace of the iris.
9. Pierella stollei
This recently discovered species of Pierella seems to differ from rhea
sufficiently to be accorded good standing. Miranda-Ribiero in describing
the species likened it to astyoche. A careful reading of his description
seems to identify a single pair in the Carnegie Museum collections. These
prove to be more closely related to rhea. My remarks about stollei are
based upon these specimens. A long series of an unnamed Bolivian
Pierella is apparently a western race of the species. I have called these
boliviana.
This species resembles luna and hyceta in having a strong green irides-
cence on the cell of the forewing on the upperside. The iridescent area on
stollei is very large and almost covers the entire wing. The females of
stollei are unique in having a rather broad and sharply defined dark margin
on the upperside of the forewing. The race boliviana has the iridescent
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patch on the males restricted to the cell and immediately adjacent areas
and the females lack the sharply defined dark margin on the forewings.
Fig. 9. Ranges of Pierella rhea and stollei.
9a. Pierella stollei stollei Miranda-Ribiero
Original description: 1931, Bol. Mus. Nac. Rio de Janiero, 7:32. No
figure has been published.
Material seen: 1 cT, 1 9 .
Range: Southeastern Brazil, probably in wooded areas of the tropical
savannahs.
Only five specimens of this form are known to me: two co types from
Jamary, Matto Grosso, one cotype from Rio Tapajoz, and the above pair
from Munez Freire, Espiritu Santo. The two specimens that I have seen
differ from the original description in one point. They are dull grey-
brown while the types are described as olive-brown (“castanho olivaceo”).
9b. Pierella stollei boliviana, subsp. nov.
Upperside: dull grey-brown; a greenish iridescent patch on the cell of
the forewings of the males extending marginad: hindwings darker to-
ward the margin, very rarely with the barest hint of bluish gloss. The
1948
Brown: Taxonomy of Pierella
75
apex of the forewing bears up to three minute, white points between
R4+5-M1, M1-M2, M2-M3. The complete series is rare in males and more
common in females. A series up to five submarginal ocelli on the hind-
wings: Rs-Mi small, sometimes only a white point, occasionally absent;
M1-M2 largest of all, white with a broad black iris; M2-M3 usually absent,
when present a white point with or without a narrow black iris; M3-CU1
second largest rarely absent, white with a broad black iris; Cui-Cu2 small,
usually present, white with a broad black iris, rarely lacking the white
pupil. The usual dark lines for the rhea group are present on the forewings
and the hindwings. The innermost line is broken in the cell on the fore-
wing and faint on the hindwing. The middle line emerges from the cell
of the forewing at or near the origin of Cu2 and crosses the hind wing out-
side of the origins of Mi and Cu2. The outermost line is the heaviest and
runs from near the origin of R4+5 to the inner margin close to the inner
angle on the forewing, and on the hindwing from directly below the ter-
mination on the forewing to near the terminus of Ai.
Underside: Pale brown occasionally with a slight purplish tinge, irro-
rate with darker brown. The ocelli of the upperside are repeated as small,
inconspicuous, white points. The lines on the upperside are repeated and
in addition, on the forewings, there is a short line outside the cell that
roughly parallels the discocellular nervules. There is a small, black spot
in the base of the cell on the forewing and two, sometimes three, in the
base of the cell of the hindwing. There is another basal spot on the hind-
wings between Cug and Ai and a spot on the weak Idc.
There is a small dark-centered sex patch on the males between Ai
and A2.
The margin of the hindwing is rounded and undulate.
Holotype: cf , Rio Yapacani, 600m. E. Bolivia (Steinbach), Sept., 1915.
Allotype: 9 , same data.
Paratypes: 1-5. cfcf, same data; 6. 9, same data; 7-10. 99, same
locality, March, 1915; 11. U, same locality, Feb., 1915; 12, 9, same
locality, Feb., 1915; 13-23. cfcf, same locality, Aug. 1913; 24-25.9 9,
same locality, Aug. 1913.
Additional specimens used but not labelled “para type” are: 3 cfcf , 3
9 9 , Rio Surutu, 350m., E. Bolivia (Steinbach), March to April, 1915; 6
cfd^, 399, Buena Vista, Prov. Sara, 400m., E. Bolivia (Steinbach)
March, 1921, May and June, 1915; 8 cfcT, 4 9 9, Prov. Sara, 450m., E.
Bolivia (Steinbach) May, 1910; 3 9 9, Portachuelo, Rio Palmatillas, E.
Bolivia (Steinbach) April, 1915.
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10. Pierella hyceta
The western Amazon basin is the stronghold of this species. It ranges
higher into the Andean foothills than do most species of the genus, oc-
casionally being found as high as 1200m. There are two clearly defined
races of the species. One of these, latona, with deep rust-red on the disc
of the hindwings, is found in the foothills from Colombia to Bolivia. The
other, hyceta, with ocher-yellow on the disc, is found at a lower altitude in
a more restricted area in the central-eastern part of the range of the species.
The foot-hill material may represent two races. I prefer to consider these
specimens as a single race exhibiting a color dine from north to south. The
northern form latona is less heavily marked on the hindwing than the
southern form cercye. The racial name is latona^ and cercye is here con-
sidered a form of it.
10a. Pierella hyceta hyceta (Hewitson)
Original description: 1860, Exot. Butt., 2: (83), pi. Haetera, fig. 1.
Earliest figure: in original description.
Other figures: 1911, Weymer, Seitz's Macrolepid., 5: pi. 43, line b.
Material seen: 6 (TcT, 3 9 9.
Range: Rain forest of upper Amazon basin below the “fall line.” (All
specimens known to me are from below 300m.)
This form is relatively uncommon in the collections that I have seen.
Bassler’s extensive collection from the lower rivers in northern Peru con-
tains a single specimen from Iquitos. The large collections made by Klages
on the Rio Purus in Brazil, just east of the area collected in by Klug
for Bassler, contain the rest of the material noted above.
10b. Pierella hyceta latona (Felder)
Original description: 1869, Reise Novara, Lep., 3:459. No figure known
to me.
^ I am not sure of the date of publication of Hewitson’s rare Bolivian Butter-
Jiies in which cercye was described. It must be later than 1869, the date of Felder’s
name latona, since it is based on Buckley’s collection made early in the 1870’s. The
Ent. Monthly Mag., 6:219, 1870, reports Buckley at a meeting of the Entomological
Society of London on Jan, 3, 1870, after his first South American trip to Ecuador:
Ent. Mo. Mag., 9:65, 1874, states “Mr. Buckley, who has just been out to Bolivia
for me ” W. C. H.
1948
Brown: Taxonomy of Pierella
77
f. cercye (Hewitson)
Original description: (1870) Bol. Butt: 10.
Earliest figure: 1911, Weymer, Seitz's Macrolepid., 5: pi. 43, line b.
Material seen: latona — 7 cTcf, 2 9 9; intermediate 21 cfcf, 3 9 9 ;
cercye — 2 1 9 .
Range: Rain forests of the western Amazon basin above “fall line.”
(All specimens known to me come from between 700m. and 1200m.)
Typical latona is found from Colombia southward to the Rio Marahon.
Typical cercye is found from Bolivia northward to the Sani Beni region
of Peru. Intermediate specimens are found from the Rio Napo, Ecuador,
south at least to the Rio Marahon in Peru.
The mahogany-red on the disc of the hindwings of cercye is darker than
on latona. The dark border on the upperside of the hindwings of latona
does not involve the row of ocelli ; on cercye it engulfs these markings and
Fig. 10. Ranges of Pierella hyceta and luna.
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obliterates some of them. The great majority of the specimens I have seen
are neither typical latona nor cercye but are about half-way between the
two of them. In the regions where latona and the intermediate to cercye
fly, I have found' that the intermediate forms are more common at the
upper limits of the altitudinal range of the race, and latona more common
in the lower altitudes. I suspect that the deepening of the color, etc., is
a response to heavier rainfall.
1 1 . Pierella luna
Pierella luna is the only species of this group that is found west of the
Andes and in Central America. It resembles hyceta in general markings.
The two species differ in the number of ocelli on the upperside of the
hindwing, luna bearing two, rarely three, and hyceta four or five, rarely
three. I am inclined to believe that the two groups of forms represent a
single biologic species of the pre-Andean uplift era. I am not wholly con-
vinced that they represent independent species today. The form luna
luna probably resembles the ancestral form more closely than do any of
the others. It is interesting to note that in Colombia this form tends to
vary in the direction of the more brightly-colored race leshia found in the
equatorial forests and thus toward the “species” hyceta.
There are three easily recognized races of luna. The race luna, occupy-
ing the central part of the species-range has more or less concolorous wings.
The northern race rubecola Qieracles) from Mexico and Guatemala has a
rusty flush on the hindwings. The southern race from western Ecuador
and southwestern Colombia has a bright rust disc on the hindwings.
11a. Pierella luna luna (Fabricius)
Original description: 1793, Ent. Syst., 3(1); 109.
Earliest figure: 1850, Hewitson, Exot. Butt., 2: Haetera 1, f. 3.
Other figures: 1911, Weymer, Seitz's Macrolepid., 5: pi. 43, line a.
Material seen: 44 cfcf , 13 9 9 .
Range: In the tropical rain forests from eastern Honduras to northern
Colombia; rarely found above 1000m.
Godman and Salvin described a form of this race as pallida {Ann. Mag.
Nat. Hist., (4) 2:142, 1868). Later in the Biologia they relegated this
name to the synonymy of luna. In 1896, Oberthur described and figured
an albinic specimen and dubbed it alhina {Etudes d'Ent., 20:32, pi. 2,
f. 10). As has been noted above, some of the specimens of luna from
1948
Brown; Taxonomy of Pierella
79
Colombia bear a dusting of rusty red scales on the upper side of the hind-
wing. This is most highly developed on females but also occurs on males.
These specimens are not intergrades to leshia. The general impression
given by these specimens is of evolution toward a form similar to ruhecola.
I believe that the form is worth taxonomic recognition but name it with
some misgivings.
f. rubra forma nova
A form of luna luna differing from typical specimens in this respect: the
upperside of the hindwings bear a light dusting of rusty red scales.
Holotype: 9 , Quindio, Colombia (Fassl), Reading Public Museum.
Paratypes: 1. 9 , “Bogota,” Colombia, Carnegie Museum; 2-3. cT’cf’,
“Colombia” (Ovalle), American Museum of Natural History; 4. 9 , same
data as 2 and 3.
There is an interesting aberrant specimen in the collection of the Car-
negie Museum. It is a male collected in May at Don Diego, Colombia.
On it the middle dark line on the underside of the hindwing is irregular and
shifted toward the outer line in the vicinity of the cell (see Schwanwitsch
loc. cit. for a detailed discussion of such shifts).
11b. Pierella luna rubecola Godman and Salvin
Original description: 1868, Ann. Mag. Nat. Hist., (4) 2:142.
Earliest figure: 1880, Godman & Salvin, Biologia Cent-Amer., Rhop.,
3: pi. 6, f. 10, 11.
Material seen: 7 cfcf , 4 9 9 .
Range: Tropical rain forests from Mexico southward to Honduras.
This race is often referred to as heracles Bdv. Boisduval’s description
was published in 1870 in his account of Lepidoptera received by him from
M. d’Orza. Thus it is antedated two years by ruhecola.
There is some variation in the row of ocelli on the upper side of the
hindwing. Typical specimens show two white-pupilled black ocelli.
Occasional specimens bear up to five, the anterior two being large, the
others small. Some specimens have one, two, or three, white dots pos-
terior to the normal complement of ocelli.
11c. Pierella luna lesbia Staudinger
Original description: 1888, Exot. Tagf., 1:219.
Earliest figure: 1911, Weymer, Seitz's Macrolepid., 5: pi. 43a.
Material seen: 12 cf cT, 4 9 9.
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Range: The tropical rain forests on the western slope of the Cordillera
Occidental in Ecuador and S. Colombia.
The only Colombian specimens of which I know are Staudinger’s type
from the Rio San Juan and a female from “Colombia” in the Ovalle col-
lection at the American Museum of Natural History. It appears to be
common in Ecuador.
Division C
Characteristics: The species that compose this division bear a pattern
on the upperside that is abberrant to this genus. On the underside they
are sufficiently close to the other species to be included in Pierella. Struc-
turally this division is clearly Pierella. The division is characterized by
bearing large blue or blue and white areas on the upperside of the hind-
wings and smaller areas on the forewings. Beneath, the characters of the
two preceding divisions are combined. The forewing resembles Division
A in that there is a light transverse band margined with dark lines. The
hindwings are like Division B and bear three straight parallel dark lines.
IV. the hortona group.
Characteristics: those of the division.
The group is restricted to the western part of the Amazon basin in the
tropical rain forests. Two species are generally recognized in the group,
hortona and albofaciata. Typical specimens of these are easily recognized
but each produces forms that approximate the other. It is quite possible
that there is only one biologic species in this group. The two taxonomic
species may be separated on the appearance of the underside. The outer
part of the wings of hortona are essentially the same color as the basal part.
These areas are strongly contrasting in albofaciata except among the
males of the race decepta described below.
12. Pierella hortona (Hewitson)
Original description: 1854, Trans. Ent. Soc. London, (2)2:246, pi. 23, f. 1.
Earliest figure: in original description.
Other figures: 1888, Staudinger Exot. Tagf., 2: pi. 77. 1911, Weymer,
Seitz's Macrolepid., 5: pi. 43 b.
f. hortensia (Felder)
Original description: 1862, Wiener Ent. Mon., 6: 125. No figure known
to me.
1948
Brown: Taxonomy of Pierella
81
Material seen: 30 cf cT, 9 9 9 •
Range: The rain forest of the west-central Amazon basin from S.
Colombia to N. Peru below 1100m. and eastward as far as the junction
of the Rio Negro and Rio Amazon.
This species is quite variable, hardly two specimens being exactly alike.
The variation shows two trends — one, toward elimination of the blue
patch at the end of the cell on the upperside of the forewing, the other
toward enlargement of the blue areas and replacement of the blue scales
with white.
Typical horiona has a well-developed blue spot on the forewing and a
larger one on the hindwing.
Typical hortensia has the spot on the forewing greatly reduced, on some
specimens it is almost gone, and the spot on the hindwing contains a large
area of white scales.
A third form with large white areas in the blue patches on both wings
approaches albofaciata in the appearance of the upperside. I call this form
albopunctata.
f. albopunctata forma nov.
This form differs from typical hortona in having a large white area within
the sub-apical patch on the upperside of the forewings and a large white
center to the blue area on the hindwings. It differs from hortensia in
having a large colored area on the upperside of the forewings.
Beneath, this form is typical of hortona both in coloration and disposi-
tion of the dark lines in the apico-cellular area of the forewings.
Holotype: cT, Upper Rio Huallaga, Peru (Klug) ex Bassler, f 6107,
July 20, 1928, American Museum of Natural History.
Allotype: 9 , same data as holotype but caught on August 19, 1928.
Paratypes: 1. cf, Juanguy, Upper Rio Huallaga, Peru, Nov. 1935,
(Klug) ex Mengel, Reading Public Museum. 2. 9 , Rio Tsaya, Amazonas,
Peru, Oct. 1927, (Klug) ex Mengel, Reading Public Museum. 3. cf,
Manacapuru, Brazil, Sept. 1925, (Klages) Carnegie Museum.
These three forms intergrade completely and have no definite geo-
graphic restrictions. The form hortensia tends to be more common in the
northeastern part of the range, albopunctata in the southeastern part.
There are several other minor variations that do not warrant names.
A good series in the Carnegie Museum indicates that a local form flies
around Sao Paulo de Olivenga on which the underside is materially darker
than on specimens from farther west.
82
Annals of the Carnegie Museum
VOL. 31
One of the differences between hortona and albofaciata is the presence,
on the latter, of a bar across the costal end of the characteristic dark
flanked light band on the underside of the forewing. This bar is light in
color and flanked by dark lines. There is a specimen of hortona in the
Mengel collection at Reading, taken by Klug at San Gabriel on the
Upper Amazon in Brazil, which resembles albofaciata in this one respect.
Fig. 11. Ranges of Pierella hortona and albofaciata.
13. Pierella albofaciata
This species from south of the Rio Mara non is much less frequently
found in collections than is hortona. Its characteristics have been pointed
out above. There are two races of the species, albofaciata from Peru and
decepta from Bolivia. These are alike on the upper surface but differ ma-
terially on the underside.
13a. Pierella albofaciata albofaciata Rosenberg
Original description: 1913, Trans. Ent. Soc. Lond., p. 678.
Material seen: 3 cfcf, 2 9 9.
Range: Tropical rain forest of the tributaries to the Amazon in N. Peru.
On the underside of albofaciata the outer half of the wings is very dark
1948
Brown: Taxonomy of Pierella
83
brown and the inner half dun-colored with a slight pearly luster. On the
upperside there is a large rectangular white bar across the discocellular of
the forewing.
13b. Pierella albofaciata decepta subsp. nov.
Upperside: like albofaciata in both sexes.
Underside: differs from albofaciata in these respects:
Male forewing: dilute dark brown marginad of the light transverse
band and as a suffusion along the inner margin of that band; the basal
half of the wing is a light sand color with a pearly luster; there are two
black points in the cell.
Male hindwing: almost entirely of the light sand color with a strong
pearly luster; the three dark lines are not strong, one crosses the wing
through the middle of the cell, the middle one crosses just outside of the
cell and the outer one passes through the sharp bend in M3; costad of M3
the outer line is suffused margined with dark brown; there is a series of
minute black submarginal dots between the nervules from Rs to Cu2;
there are black dots in the base of the cell and between Cus and Ai at
the base.
Females approach the typical coloration of albofaciata on this surface:
They are a little lighter and more pearly.
Holotype: cf, Rio Yapacani, E. Bolivia, March 1915 (Steinbach),
Carnegie Museum.
Allotype: 9 , the same data except caught in February 1915.
Paratypes: 1-2. (Tcf, same data, caught in August 1913. 3-5. cfcf , same
data, caught in September 1914. 6. 9 , same data, caught in February
1915.
Check-List of the genus Pierella.
1. nereis (Drury)
2. helvina
a. helvina (Hewitson)
b. incanescens Godman and Salvin
f. costaricana Niepelt
c. ocreata Godman and Salvin
9 f. hymettia Staudinger
9 f. werneri Hering & Hopp
9 f. johnsoni Talbot
f. pacifica Niepelt
84
Annals of the Carnegie Museum
VOL. 31
3. hyalinus
a. hyalinus (Gmelin)
b. dracontis (Huebner)
f. extincta Weymer
c. fusimaculata Brown
4. lena
a. lena (Linnaeus)
b. hrasilensis (Felder)
f. glaucolena Weymer
f. obsoleta Brown
5. amalia Weymer
6. lucia Weymer
7. astyoche (Erichson)
8. rhea
a. rhea (Fabricius)
f. lamia (Sulzer)
b. chalyhaea Godman
f. columhina Krueger
9. stollei
a. stollei Miranda-Ribiera
b. holiviana Brown
10. hyceta
a. hyceta (Hewitson)
b. latona (Felder)
f. cercye (Hewitson)
11. luna
a. luna (Fabricius)
ab. albina Oberthur
f. rubra Brown
b. rubecola Godman and Salvin
c. lesbia Staudinger
12. hortona (Hewitson)
f. hortensia (Felder)
f. albopunctata Brown
13. albofaciata
a. albofaciata Rosenberg
b. decepta Brown
1948
Brown: Taxonomy of Pierella
85
Notes on Ecuadorian Pierella.
The species marked with an asterisk in the following list are here re-
ported from Ecuador for the first time. The major portion of the Brown
collection is now at the American Museum of Natural History.
*2c. Pierella helvina ocreata G. & S.
1 9 , Hacienda Lorena, near Santa Domingo de los Colorados, 550m.,
Feb. 19, 1941 (D. B. Laddey) coll. F. M. Brown.
An abberant specimen of f. pacifica Niepelt that approaches 9 f. hymettia
Staudinger.
*4b. Pierella lena brasilensis Felder
1 cf , Canelos, Oriente, 700m., Dec. 12, 1938 (W. C. MacIntyre), coll.
F.M.B.
1. cf, Rio Jondachi, near Archidona, Oriente, 800 m., Nov. 1939
(W.C.M.), coll. F.M.B.
1 d^, Rio Toachi, Pichincha 800m., Nov. 1939 (W.C.M.), coll. F.M.B.
1 cf , Puyo, Oriente, 1000m., Dec. 5, 1938 (F.M.B.), coll. F.M.B.
Sd'd^, Sarayacu, Oriente, 500m. (Schaus) U.S.N.M.
1 d', Rio Arajuna, Oriente 1000m., April 27, 1941 (W.C.M.), coll.
F.M.B.
The Rio Toachi record is very doubtful. MacIntyre purchased the
material from a Quito bird collector who had visited both Rio Jondachi
and Rio Toachi in Novembei 1939. 1 firmly believe the specimen is from
east of the Andes.
*6. Pierella lucia Weymer.
1 d, Canelos, Oriente, 700m., Dec. 12, 1938 (W.C.M.), coll. F.M.B.
2 dd, Sarayacu, Oriente, 500m., (Schaus) U.S.N.M.
4rdd, l9Zumbi, Oriente, 700m., Oct. 28, 1911, Nov. 1941 (D.B.L.),
coll. F.M.B.
*8b. Pierrella rhea chalybaea Godman
Id, Rio Jondachi, near Archidona, Oriente, 800m., Nov. 1939 (W.C.M.),
coll. F.M.B.
3 cfcf, Sarayacu, Oriente, 500m.
These are typical f. columbina Krueger in darkness of color and dis-
position of ocelli; they tend to chalybaea in that there is a faint bluish
gloss on the hindwings.
86
Annals of the Carnegie Museum
VOL. 31
10. Pierella hyceta latona (Felder)
10 cfcT, Puyo, Oriente, 1000m., Feb. 6, Dec. 1938 (F.M.B.), coll.
F.M.B.
1 cT, Rio Tutenongoza, Oriente, 800m., Feb. 10, 1939 (F.M.B.), coll.
F.M.B.
1 d'j 2 9 9 , Huagra-yacu, Oriente, 900m., March 7, April 6, 1941
(W.C.M.), coll. F.M.B.
Id',! 9 Rio Arajuna, Oriente, 1000m., April 23-26, 1941 (W.C.M.),
coll. F.M.B.
5 Hacienda La Mascota, Oriente, 1250m., (Coxey) A.N.S.P.
1 d, Rio Pastaza, Oriente, Oct. 1, 1936 (W.C.M.), U.S.N.M.
1 cT, 19 , Macas, Oriente, Jan. 26 (Feyer?), ex Mengel coll., Reading
Public Museum.
1 d, Zumbi, Oriente 700m., Nov. 11, 1941 (D.B.L.), coll. F.M.B.
1 19, Zamora, Oriente, 1000m., Dec. 20, 1941 (D.B.L.), coll.
F.M.B.
3 “Pacific Slope” (E. W. Rorer), U.S.N.M.
Stiibel collected this species at Pintuc (Puyo). These were reported as
cercye by Weymer in his account of Stiibel’s collection. All of the Ecua-
dorian material with locality data except the Zumbi and Zamara speci-
mens are intermediate to cercye. The three excepted specimens are typical
latona as are the three “Pacific Slope” specimens in the U.S.N.M. I
gravely doubt the veracity of that data. I know Mrs. Rorer and know that
while living on the Pacific Slope, she frequently collected in the “Oriente.”
I feel sure that some slipshod prepara tor put a “Pacific Slope” label on the
specimen without reference to Mrs. Rorer’s locality data.
11c. Pierella luna lesbia Staudinger
1 d Balzapamba, 630m., June 1938 (W.C.M.), coll. F.M.B.
1 d, Playas de Montalvo, 30m., April 1938 (W.C.M.), coll. F.M.B.
2 dd, Santo Domingo de los Colorados, 500m., Dec. 12-19, 1940
(D.B.L.), coll. F.M.B.
1 9 ,La Lorena, 550m., Feb. 25, 1941 (D.B.L.), coll. F.M.B.
1 d*, Rio Toachi, 800m., Sept. 1938 (W.C.M.), coll. F.M.B.
2 9 9 , Palmar, 200m., March 31, April 15, 1941 (D.B.L.), coll. F.M.B.
1 d, Rio Maizito, near Palmar, 200m., May 7, 1941 (D.B.L.), coll.
F.M.B.
Huigra, March 11 (S. N. Rhoads), A.N.S.P. and R. P. M.
^ dd, Dos Puentes (Coxey) A.N.S.P.
1948
Brown: Taxonomy of Pierella
87
Zaruma {ex Dognin) U.S.N.M.
Las Guayas (Rorer) U.S.N.M.
In addition to these specimens of the race lesbia, there are three speci-
mens of the race luna in the Mengel collection at the Reading Public Mu-
seum. One of these is labelled “Guayaquil,” the other two are labelled
“Cuenca.” The latter locality is certainly incorrect. I have collected in the
Cuenca area and it is quite impossible that Pierella is to be found there. It
is totally wrong for the genus climatically, being semi-arid and cool. The
climatic conditions near Guayaquil are better suited to the genus. How-
ever the suitable areas there are well occupied by the race lesbia. Six
excellent collectors, Rhoads, Rorer, Coxey, Laddey, MacIntyre and Feyer
have collected in this region and have not turned up luna unless Feyer
caught the Mengel specimen. Since Mengebs collection contains numerous
mislabelled specimens I am inclined to doubt the validity of the Guaya-
quil label.
Dognin in his report of the lepidoptera from the environs of Loja did
not report lesbia. He did report hyceta. There are no hyceta in that part
of the Dognin collection at the U. S. National Museum. Possibly he mis-
identified the Zaruma specimen as hyceta.
*12. Pierella hortona (Hewitson)
1 cT, 19, Canelos, Oriente, 800m., Dec. 12, 1938 (W.C.M.), coll.
F.M.B.
2 cTcr, Rio Jondachi, Oriente, 800m., Nov. 1939 (W.C.M.), coll.
F.M.B.
1 cT, Rio Tutenongoza, Oriente, 800m., Feb. 10, 1939 (F.M.B.), coll.
F.M.B.
Hacienda La Mascota, Oriente, 1250m., (Coxey), A.N.S.P.
3 cfcT, Sarayacu, Oriente, 500m., (Schaus) U.S.N.M.
1 cf, Macas, Oriente, 1050m., (L. M. Higgins) U.S.N.M,
1 cT, Macas, Oriente, (Feyer?) R.P.M.
1 cf , “Ecuador” {ex Johnson), U.S.N.M.
These are typical hortona except the two from Macas and the “Ecuador”
specimen which are hortensia (Felder).
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ART. 7. THE LAND SNAILS OF WEST VIRGINIA
By Gordon K. MacMillan
(Plates 1-15)
PREFACE
The taxonomy of the species in this paper, especially of the family
PolygyridcB, has been based entirely upon the character of the shell. This
is in direct contrast with the system proposed by Dr. H. A. Pilsbry, Dr.
H. B. Baker, and others, who are using anatomical characters of the
animal as a criterion for classification. In view of this, much of the
classification proposed by Dr. Pilsbry in “The Land Mollusca of North
America (North of Mexico)”, vol. 1, pt. 2, 1940, has not been followed en-
tirely by the author.
In most cases the original description of each species and subspecies has
been copied from the publication in which it appeared. Some of the earlier
descriptions had been written in Latin or in Latin with an English de-
scription following. In this paper the English translations have been used
in all such cases or the English description utilized when accompanied
with the Latin one.
Many of the descriptions of the species and varieties of land snails
contained in this article are adaptations or condensations from the
original descriptions by the describers of the species. Much use was made
of the parts of the “Manual of Conchology” and the volumes of the “Land
Mollusca, etc.,” which pertain to the fauna of West Virginia. Although
these adaptations of Dr. Pilsbry and also those of Dr. H. B. Baker have
not been acknowledged with each species, recognition of their use is made
now. Conchologists will undoubtedly recognize in the descriptions of the
species, some of the characteristic phraseology of Dr. Pilsbry.
In the extension and support of this work. Dr. and Mrs. Stanley T.
Brooks have made extensive collections from the West Virginia line to the
Great Smoky Mountains.
I extend my appreciation to Dr. Harald Rehder, of the United States
National Museum, for his kindness in identifying the Succinea and in
verifying those species that I had already named. I wish to thank the
Note: The term “land snails” used in the title of this paper, includes those
terrestrial forms which possess a visible shell, but does not include the “slugs,”
which possess only the remnant of a shell under the skin in the region of the neck,
or which have no shell at all. Perhaps a more technical title would have been the
“terrestrial shell-bearing mollusca,” since this includes all land forms with a shell,
and still excludes the slugs.
89 FEB 1 1950
Issued February 28, 1949.
90
Annals of the Carnegie Museum
VOL. 31
officials of the West Virginia Academy of Science for their contribution to
this work. Without their support much territory could not have been
covered. I also wish to thank M. Graham Netting, Curator of Herpe-
tology at the Carnegie Museum, and many contributors to the collections
for their work and data.
The list of collectors and others who have contributed toward our
knowledge of the mollusca of West Virginia follow. The initials after
the locality at which the shells have been found, are those of the collector
and reference to the following list will indicate his identity. These sym-
bols are used throughout this paper.
AEO A. E. Ortmann
AFA A. F. Archer
CB C. Brown
CBW C. B. Wurtz
cc Crooke Collection (Ameri-
can Museum of Natural
History)
CG Calvin Goodrich
cjG C. J. Goin
C&A W. J. Clench and A. F.
Archer
Ejc E. J. Court
FML F. M. Love
FRW F. R. Wilson
GB Grace Bogges
GEW G. E. Wallace
GHC. . . . . .G. H. Clapp
GKM G. K. MacMillan
GRH G. R. Hunt
GWT G. W. Tryon
HAP H. A. Pilsbry
HHS H. H. Smith
IHM I. H. Marshall
JHA J. H. Alexander
JPEM J. P. E. Morrison
LL Leonard Llewellyn
Mcz Museum of Comparative
Zoology
MGN M. G. Netting
MSB M. S. Briscoe
MRS. GA. .Mrs. G. Andrews
M&R G. K. MacMillan and
N. D. Richmond
NDR N. D. Richmond
PCB P. C. Bibbee
PR Paul Ridgway
p&i H. A. Pilsbry and J. E.
Ives
p&L W. M. Perrygo and C.
Lingebach
RLF R. L. Fricke
UM .University of Michigan
USFC United States Fish Com-
mission
vs Victor Sterki
wjF W. J. Fox
Family Helicinid^
The members of this family of snails are characterized by a shell that is
globose, the base of which has a callus around the columella; while the
columella is somewhat flattened and rather straight. The aperture is
1949
MacMillan: Land Snails of West Virginia
91
triangularly semi-ovate and entire ; the peristome is either simple, straight,
or thickened, and often widely expanded, with or without teeth. The
operculum is not spiraled and somewhat semi-oval, and is also membran-
ous or testaceous.
This family is represented by only one genus and species in West
Virginia, and is one of the two snails in that state which possesses an
operculum.
Genus Hendersonia Wagner
Shell carinated, narrow and slowly increasing whorls, solid, striated,
imperforate. Aperture small, semi-lunar; the peristome slightly reflected
and thickened internally. Umbilicus covered by reflected peristome.
Whorls 5^21 suture scarcely indented.
Hendersonia occulta (Say)
(Plate 1, fig. 1)
Shell small, rather solid, low conical, acute at apex, cretaceous, obviously
striated ; spire of five nearly plane whorls, the last of which is angular at
the periphery, and this angle continuing up the spire adjacent to the suture,
makes it appear double; the aperture is small, semi-lunar; the peristome
is scarcely reflexed, but is thickened internally ; the columella is very short,
and joins the peristome by a slightly waving curve, without forming an
angle. Diameter 6, height 5 mm.
Type Locality: A rugged and abrupt “bluff,” a half-mile below New
Harmony, Indiana (Say, 1831).
Range: Pennsylvania to Minnesota, south to Missouri and Florida.
West Virginia Records
Grant County; Greenland Gap (ndr) . Greenbrier County; Renick (grh) ,
Renick Valley (ndr). Marion County; Fairmont (ndr). Mineral County;
Keyser (ihm). Ohio County; Wheeling (T. Bland). Pendleton County;
Franklin (ndr) (gkm). Wyoming County; Oceana (m&r).
Remarks: This is the only species in the family Helicinidce in the United
States that is carinated and striated. Although it is found in this country
east of the Rocky Mountains, its distribution is rather local and dis-
continuous.
Hendersonia occulta was first described from a fossil shell, being abund-
ant in the loess formations of Iowa, Nebraska, Missouri, Illinois, and
Indiana. In 1832, Jacob Green described a living species of Helicina,
92 Annals of the Carnegie Museum vol. 31
which is called rubella. Some conchologists consider this a subspecies of
occulta, and others a synonym of that species.
Family Cionellid^ Kobelt
The shell is small or minute, imperforate, elongate, varying from ovate
to turrite or subcylindric, thin, glossy, having little or no sculpture, and
generally it is somewhat transparent. The apex is small and obtuse,
embryonic whorl not differentiated from those following. Aperture ovate
or piriform, simple. The columella is usually sinuate or truncate at the
base. Outer lip not expanded, often thickened within.
Genus Cionella Jeffreys
The shell is oblong-conic or oblong-cylindric, imperforate, smooth and
glossy, composed of from 5 to 7 slightly convex whorls. Aperture small,
ovate, nearly vertical, toothless. Outer and basal lip arcuate, obtuse,
thickened within by a callous rim which is continuous to the upper in-
sertion ; columella short, concave or straightened, very slightly sinuate at
the base; parietal callus very thin throughout.
There is only one species represented in the fauna of North America.
Although two forms of Cionella luhrica have been described, each of
these is hardly distinguishable from the typical species.
Cionella lubrica (Mueller)
(Plate 1, fig. 4)
Shell small, thin, transparent, oblong-oval; epidermis smoky horn-
color, smooth, very bright and shining; whorls 5 or 6, somewhat rounded,
the last equalling two-fifths the shell’s length, rounded at base; apex
obtuse ; suture somewhat impressed ; aperture lateral, oval, its plane nearly
parallel with the axis of the shell; peristome simple, thickened, often
slightly rufous; umbilicus imperforate; columella obsoletely truncated at
base. Length 6 mm.; diameter 2.5 mm.
Type Locality: Not designated. (Mueller, 1774).
Range: Newfoundland; Canada; Alaska; United States.
West Virginia Records
Boone County; Peytona (ndr) (m&r), Seth (m&r). Braxton County;
Gassaway (gkm). Cabell County; Barboursville (m&r), Ona (m&r). Clay
County; Clay (gkm). Doddridge County; West Union (frw). Fayette
1949
MacMillan: Land Snails of West Virginia
93
County; Cliff top (gkm). Grant County; Greenland Gap (ndr), Mount
Storm (ndr). Greenbrier County; Alderson (gkm), Renick (grh), Renick
Valley (ndr), Ronceverte (gkm) (m&r), White Sulphur Springs (p&l).
Jefferson County; Charles Town (msb), Leetown (msb). Harper’s Ferry
(msb) (jpem). Kanawha County; Hudnall (cbw). Lewis County; Jackson’s
Mill (gkm), Weston (gkm). Lincoln County; Branchland (m&r), Myra
(m&r). Logan County; Blair (m&r). Sharpies (m&r). McDowell County;
Avondale (m&r). Marion County; Fairmont (ndr). High Point (ndr).
Mercer County; Bluefield (m&r), Princeton (m&r), Speedway (m&r).
Mineral County; Burlington (gkm). Monroe County; Alderson (gkm).
Salt Sulphur Springs (m&r), Sweetsprings (m&r). Nicholas County;
Lockwood (gkm). Pendleton County; Circleville (ndr), Franklin (ndr)
(gkm). Pocahontas County; Dunmore Spring (gkm), Greenbank (gkm),
Marlinton (gkm). Mill Point (gkm). Putnam County; Poca (gkm).
Randolph County; Huttonsville (gkm). Summers County; Riffle (gkm),
Talcott (gkm). Upshur County; French Creek (gkm). Waijne County;
Fort Gay (m&r). Webster County; Bolair (ndr). Wetzel County; Silver
Hill (ndr). Wyoming County; Pineville (m&r).
Family Haplotrematid^
Shell thin, widely umbilicated, depressed, striated or wrinkled, color
pale yellowish or greenish yellow ; whorls 43/2 to 5, the last broad, depressed,
and moderately deflected in front; the aperture is obliquely ovate, the
peristome somewhat thickened or expanded, the margins approximating,
and the basal shortly reflexed ; the columella is entire.
Genus Haplotrema Ancey
The characters are the same as those of the family Haplotrematidce.
This genus is represented by only one species in West Virginia, although
two forms, major and minor, have been recognized, but are distinguished
from the typical species only by differences in measurements.
Haplotrema concavum (Say)
(Plate 1, figs. 2, 3)
Shell much depressed, suborbicular, horn-color or whitish, immaculate;
volutions five, irregularly wrinkled across, more convex beneath; suture
distinctly impressed ; umbilicus very large, exhibiting all the volutions to
the summit distinctly; aperture large, short; labrum towards the base very
Annals of the Carnegie Museum
94
VOL. 31
slightly and inconspicuously reflected. Greatest width seven-tenths of an
inch (21 mm.).
Type Locality: Not designated (Say, 1821).
Range: Quebec and Ontario, west to Minnesota, south to Arkansas
and east to Georgia.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County; Pey-
tona (ndr) (m&r), Racine (m&r), Seth (m&r). Braxton County; Frametown
(gkm), Gassaway (gkm), Shadyside (gkm). Cabell County; Bourboursville
(ndr) (m&r), Lesage (m&r), Ona (m&r). Calhoun County; Grantsville
(ndr). Clay County; (gew) (gkm). Doddridge County; Sherwood (ndr).
West Union (frw). Fayette County; Clifftop (mrs. gkm) (gkm). Grant
County; Greenland Gap (ndr). Mount Storm (ndr), Petersburg (pr).
Greenbrier County; Alderson (ndr) (gkm). Organ Cave (ndr). Renick
Valley (ndr), Ronceverte (m&r). White Sulphur Springs (p&l). Hamp-
shire County; Romney (gkm) . Hancock County; Pugh town (ll) . Harrison
County; Bristol (ndr). Jefferson County; Bardane (msb), Bloomery
(jpem), Bolivar (msb), Charles Town (jpem), Harper’s Ferry (jpem)
(gwt), Key Gap (jpem), Loudoun Heights (jpem), Reedson (msb),
Shenandoah City (msb). Kanawha County; Alum Creek (ndr), Charles-
ton (gkm), Dunbar (gkm), Hudnall (cbw), St. Albans (um). South
Charleston (ndr). Lewis County; Jackson’s Mill (gkm), Weston (gkm).
Lincoln County; Branchland (m&r), Myra (m&r), Sheridan (ndr). Logan
County; Blair (ndr) (m&r), Davin (ndr). Sharpies (m&r), Slagle (cg).
McDowell County; Avondale (m&r), Panther (m&r). Marion County;
Curtisville (ndr), Fairmont (ndr) (grh). High Point (ndr), Jayenne
(grh), Midway (grh), Mt. Harmony (ndr), Rivesville (ndr). Smith ville
(cbw). Marshall County; Bannon (ndr), Kent (ndr), Powhattan Point
(frw). Mason County; Leon (ndr). Point Pleasant (ndr) (cbw). Mercer
County; Bluefield (m&r), Oakvale (m&r), Princeton (m&r), Spanishburg
(m&r), Speedway (m&r). Mineral County; Burlington (gkm), Keyser
(ihm). Mingo County; Taylorsville (m&r), Williamson (m&r). Monongalia
County; Coopers Rock (cbw), Smithtown (ndr), Sturgisson (ndr). Monroe
County; Alderson (gkm). Gap Mills (m&r). Salt Sulphur Springs (ndr)
(m&r), Sweetsprings (m&r), Waitesville (m&r). Morgan County; Berkeley
Springs (ndr), Largent (ndr). Nicholas County; Craigsville (gkm). Lock-
wood (gkm), Richwood (ndr), Summersville (gkm), Swiss (ndr). Ohio
County; Wheeling (mgn) (ndr) (cbw). Pendleton County; Circleville
1949
MacMillan: Land Snails of West Virginia
95
(ndr), Franklin (ndr) (gkm), Judy Gap (gkm), Upper Tract (mgn) (ndr).
Pocahontas County; Cass (mgn), Dunmore Spring (gkm), Greenbank
(gkm), Hillsboro (ndr) (gkm), Marlinton (mgn) (gkm), Mill Point (gkm).
Preston County; Cascade (ndr), Cranesville (mgn), Manheim (jha). Terra
Alta (mgn). Putnam County; Poca (gkm). Raleigh County; Daniels (m&r),
Shady Springs (m&r). Randolph County; Cheat Bridge (gkm), Helvetia
(vs), Huttonsville (mgn) (ndr). Summers County; Riffle (gkm), Talcott
(gkm). Tucker County; Parsons (mrs. gkm) (gkm). Tyler County; Friendly
(cBw). Upshur County; French Creek (mrs. gkm) (gkm). Wayne County;
Fort Gay (m&r). Webster County; Bolair (ndr) (m&r). Wetzel County;
Silver Hill (ndr). Wirt County; (wjf). Wyoming County; Baileysville
(m&r), Oceana (m&r), Pineville (m&r).
Family Pomatiopsid^e Stimpson
Shell elongated, turreted, thin to thick, smooth, iimbilicated ; aperture
expanded; peristome continuous, thin or slightly reflected; operculum
corneous, subspiral, with spiral sculpture.
This, together with the Helicinidoe, are the only families of Gastropoda
in West Virginia which possess an operculum. Only one genus and two
species are found in the state.
Genus Pomatiopsis Tryon
The characters of the genus are the same as those of the family.
Although most of the representatives of this genus are aquatic, P.
lapidaria and P. prcelonga have been found in moist places and also on
hillsides, sometimes as much as a hundred to two hundred feet from any
body of water.
If the species of this genus are land snails, as many conchologists believe,
they are the first of their kind which possess gills, the characters of fresh-
water gastropoda and not of the land snails, which breathe by means of
lungs.
Pomatiopsis lapidaria (Say)
(Plate 1, fig. 5)
Shell elongated, turreted, color brownish-horn or chestnut; surface dull
to shining; lines of growth crowded, slightly wrinkled in some specimens,
rather fine; apex rounded, flattened, depressed; nuclear whorl not emer-
gent; partly embraced by the second whorl; not much constricted by the
suture, sculpture very finely granular, almost smooth; whorls 7, well
96
Annals of the Carnegie Museum
VOL. 31
rounded, slowly and regularly increasing in diameter; suture deeply im-
pressed; spire acute, about three times as long as the aperture; aperture
elongate ovate, somewhat narrowed and angled above, rounded below,
slightly expanded, somewhat purplish within ; peristome simple or slightly
thickened within, upper terminations connected on the parietal wall by a
callus of greater or less thickness; umbilicus well marked, emargined by
the inner lip which is slightly reflected over the umbilical region; base
rounded. Length 7 mm., width 3.1 mm.
Type Locality: Not designated (Say, 1817).
Range: Quebec and Ontario; New York and New Jersey to Minnesota,
south to Texas and Georgia.
West Virginia Records
Boone County; Peytona (m&r). Calhoun County; Grantsville (ndr).
Greenbrier County; Renick (grh), Ronceverte (m&r). Kanawha County;
Alum Creek (ndr), Dunbar (gkm). Lewis County; Weston (gkm). Logan
County; Sharpies (m&r). McDowell County; Avondale (m&r). Panther
(ndr) (m&r). Marion County; Fairmont (ndr). Mercer County; Bluefield
(m&r), Oakvale (m&r). Speedway (m&r). Mingo County; Taylorsville
(m&r), Williamson (m&r). Monroe County; Waitesville (m&r). Wyoming
County; Oceana (m&r).
Remarks: P. lapidaria may be known by its elongated, brownish shell,
convex and deeply sutured whorls, and the reflected rim of the aperture.
Habits: It is rarely found in water, but abundantly under wet leaves and
in places subject to overflow from creeks and streams.
Pomatiopsis praelonga Brooks and MacMillan
(Plate 1, fig. 6)
Shell elongate, turreted, and narrow. Color brownish-horn. Surface
somewhat shining and somewhat wrinkled, lines of growth very fine and
hardly discernible. Apex slightly rounded, flattened; nuclear whorl
emergent; sculpture worn and smooth. Whorls 8, somewhat rounded,
slowly and regularly increasing in diameter. Suture deeply impressed.
Spire acute, about three and a half times as long as aperture. Aperture
ovate, somewhat narrowed and angled above, rounded below. Peristome
thickened and extremities connected. Umbilicus well marked, ermargined
by inner lip which is reflected over umbilical region. Base rounded. Length
7.5, width 2.25 mm. .
Type Locality: Hillside, along Elk River, 1)/^ miles south of Clay, Clay
Co., West Virginia.
Range: West Virginia Record ; Clay County; Clay (gkm).
1949
MacMillan : Land Snails of West Virginia
97
Remarks: This is a much longer and narrower species of Pomatiopsis
than lapidaria. P. prcElonga has a coarser sculpture, more thickened
peristome, and much flatter whorls than lapidaria. In the number of
whorls and the length of the shell, prcBlonga is much like P. scalaris, but it
is much narrower. P. prcelonga is an existing species while P. scalaris is a
fossil shell from the loess of the Wabash River near New Harmony,
Indiana.
Habits: P. prcelonga was found under the dead fronds of ferns near the
bases of plants, where the immediate hillside was rocky and covered with
dead leaves.
Family Valloniid^ Pilsbry
Minute snails with perforate or umbilicate shells of few whorls, dis-
coidal to ovate conic in shape, often with spaced cutical ribs, and without
internal laminae; the peristome either expanded, thickened, or simple,
and toothless.
Genus Vallonia Risso
Shell minute, 1.8-3. 5 mm. greater diameter, openly umbilicated, with
flat or moderately elevated spire, colorless or yellowish to reddish horn-
color, transparent or diaphanous, without bands or markings of any kind.
Surface striated or with membranous ribs; whorls 3-4, rounded; aperture
nearly circular or somewhat transversely elongate; peristome everted or
straight, with or without a whitish lip.
The last whorl, in the majority of species, descends in front to the
aperture, either as a whole or at the suture only, while the back of the
whorl keeps straight to the very margin. Also in a number of species the
last whorl ascends distinctly before it descends.
Vallonia pulchella (Mueller)
(Plate 1, fig. 7)
Shell widely umbilicated, depressed, slightly convex above, thin and
transparent; epidermis colorless; whorls 33^-4, very minutely striated;
the last large and spreading at the aperture like a trumpet; aperture or-
bicular, a little dilated ; peristome much thickened, white, reflected, mak-
ing nearly a continuous circle, ends approaching; umbilicus large, exhibit-
ing all the volutions. Greater diameter 3, lesser diameter 2.5 mm.; height
1.5 mm.
Type Locality: Not designated (Mueller, 1774).
Range: Canada and the United States.
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Annals of the Carnegie Museum
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West Virginia Records
Greenbrier County; Ronceverte (m&r). Hampshire County; Romney
(gkm). Jefferson County; Aurora (Heideman), Bakerton (msb), Bolivar
(msb), Charles Town (msb), Engle (msb), Harper’s Ferry (ndr) (gwt)
(ejc), Lee town (msb), Meyerstown (msb), Rippon (msb), Silver Grove
(msb). Marshall County; Moundsville (vs). Monongalia County; Morgan-
town (hhs). Monroe County; Salt Sulphur Springs (ndr) (m&r). Union
(gkm). Willow Bend (m&r). Pendleton County; Judy Gap (gkm).
Vallonia excentrica Sterki
(Plate 1, fig. 8)
Moderately umbilicated, the umbilicus elongate, rapidly widening for
the last Yj, whorl ; slightly convex above ; pale horn-colored, transparent or
slightly opaque, with a somewhat fatty gloss, smooth or very finely and
irregularly striate, smooth at the nucleus. Whorls 3-33^, rather rapidly
increasing, with a moderately deep suture, the last comparatively large,
well rounded, expanding towards the aperture, not descending in front.
Aperture moderately oblique and inclined, forming five-sixths of a circle,
subangular at the base, the umbilical margin somewhat protracted ; peris-
tome everted only in the inferior part, with a rather strong white lip,
shining throughout the shell. Height 1.1, greater diameter 2.3, lesser 1.8
mm.
Type Locality: Quebec (Sterki, 1892).
Range: Canada, Maritime Provinces, Quebec, Ontario; New England
States, New York and New Jersey to Indiana and Michigan.
West Virginia Records
Jefferson County; Aldridge (msb), Bakerton (msb), Bloomery (msb),
Duffield (msb), Halltown (msb). Harper’s Ferry (ejc) (msb), Kabletown
(msb), Kearneysville (msb), Lee town (msb), Mechanicstown (msb), Mid-
dleway (msb), Millville (msb), Morgan Grove (msb), Mt. Pleasant (msb),
Piperstown (msb), Rippon (msb), Shepherdstown (msb). Silver Grove
(msb), Skeetersville (msb). Summit Point (msb), Uvilla (msb). Marshall
County; Moundsville (vs).
Remarks: This species is distinguishable from pulchella by its size,
averaging a little smaller, by the somewhat elongated outline of the shell
and umbilicus, the last whorl more expanding towards the aperture, the
smaller and less elevated spire; the less deep suture; the peristome little
and not abruptly everted.
1949
MacMillan: Land Snails of West Virginia
99
Vallonia costata (Mueller)
(Plate 2, fig. 1)
Moderately umbilicated, more widely so for the last half whorl, depressed
convex above, or nearly flat, grayish to light or reddish horn-colored, with
rather regularly set membranous ribs and fine striae between them, nu-
cleus with fine revolving lines. Whorls 33^, slightly flattened above and
below the periphery, at the circumference somewhat angular, with a deep
suture, rather rapidly increasing; the last whorl expanding to the aper-
ture, shortly and moderately descending in front, more so at the suture
than on the back. Aperture rather oblique and inclined, almost circular, a
little flattened above, ends of margin much approximating, slightly pro-
tracted and auricled, connected by a thin callus; peristome strongly and
abruptly everted, with a strong white lip. Height 1.1, greater diameter 2.5,
lesser 2.1 mm.
Type Locality: Fridrichsdal, Denmark (Mueller, 1774).
Range: Canada; United States as far south as California, New Mexico,
Kansas, and Virginia.
West Virginia Records
Jefferson County; Aldrich (msb), Bakerton (msb), Bloomery (msb),
Charles Town (msb), Duffield (msb), Engle (msb), Halltown (msb), Har-
per’s Ferry (msb), Jamestown (msb) Kabletown (msb), Kearneysville
(msb). Keystone (msb), Leetown (msb), Loudoun Heights (jpem), Me-
chanicstown (msb), Meyerstown (msb). Middleway (msb), Millville (msb),
Morgan Grove (msb), Rippon (msb), Shepherdstown (msb), Skeetersville
(msb). Summit Point (msb), Uvilla (msb). Monroe County; Salt Sulphur
Springs (ndr). Pendleton County; Franklin (ndr).
Remarks: Until recently V. costata had been considered a variety of
pulchella. One of the main distinguishing features is the presence of very
fine, raised revolving lines on the nuclear, or the embryonal whorls,
while those of pulchella are smooth. The generally much more depressed
spire, the deeper suture besides the rather flattened whorls above, the last
whorl more rapidly expanding towards and descending to the aperture,
especially at the suture, the more approximating margin rendering the
aperture nearly circular, are sufficient to distinguish costata from pul-
chella.
Vallonia perspectiva Sterki
(Plate 2, fig. 2)
Shell small, with very wide perspective umbilicus, more widening for the
last }/2 whorl, flat, or a little elevated above, with rather dense, somewhat
100
Annals of the Carnegie Museum
VOL. 31
regularly set, moderately strong membranous ribs (about 35 on the last
whorl) and with finer striae between them; nucleus without revolving
lines; pale-horn to colorless, thin, translucent; whorls 33^2? gradually in-
creasing, a little flattened above and below the periphery, with a deep
suture, the last rounded, comparatively narrow, little expanding towards
the aperture, rather rapidly descending; aperture very inclined and
oblique, almost tangential, transversely ovoid or oblong; peristome con-
tinuous, shortly but not abruptly everted except near the suture, without
(or with a very thin) lip. Diameter major 2, minimum. 1.7, altitude
0.7 mm.
Type Locality: Knoxville, Tennessee (Sterki, 1893).
Range: West Virginia and Kentucky to Iowa and Montana, south to
Arizona and east to Alabama and Tennessee.
West Virginia Records
Grant County; Greenland Gap (ndr). Greenbrier County; Renick (grh),
Renick Valley (ndr). Hampshire County; Romney (gkm). Jefferson
County; Bloomery (jpem), Harper’s Ferry (jpem). Pendleton County;
Franklin (ndr) (gkm). Judy Gap (gkm). Upper Tract (gkm). Pocahontas
County; Greenbank (gkm).
Remarks: V. perspectiva cannot be mistaken for any other species except
parvula, which it resembles in size, coloration, and the membranous ribs.
But it is sufficiently distinguished by the wider umbilicus, the compara-
tively narrower last whorl, less expanding and descending to the aper-
ture, which is smaller and not circular; the continuous peristome and the
absence of a lip. It holds a singular position also among the species with
strongly descending last whorl and transversely elongated aperture by the
comparatively more distant and regularly standing membranous ribs, and
its small size.
Family Strobilopsid^ Hanna
Shell trochiform, dome-shaped or discoidal, umbilicate, of 4^^ to 6
slowly enlarging whorls. The aperture is small, oblique, with armature of
2 or 3 parietal lamellae and several deeply placed basal folds, all growing
continuously from an early neanic stage. Peristome more or less thickened
and expanded, the ends of the lip remote, joined by a parietal callus.
Genus Strobilops Pilsbry
The shell is small, perforate or umbilicate, trochiform to subdiscoidal,
with rounded, angular, or carinate periphery, of 4^^ to 6 closely coiled
1949
MacMillan: Land Snails of West Virginia
101
whorls. Cavity of the last whorl is obstructed by two or three long parie-
tal lamellae, the upper one emerging to the edge of the parietal callus, the
lower one weaker, emerging or immersed, the intermediate one when
present, smallest and remote from aperture ; a series of two or more short
folds on the basal wall of the cavity deep within the last whorl. Peristome
expanded; usually thickened, the insertion of the lip remote, connected by
a parietal callus.
The members of this genus are characterized by the presence of the
parietal lamellae and basal folds, which separate them from all other shells
in the North American fauna. These lamellae and folds appear very early
in the life of the shell, growing at the forward end of the shell and being
absorbed behind.
Strobilops labyrinthica labyrinthica (Say)
(Plate 2, figs. 4, 5)
Shell conic, dark reddish-brown, body lighter; whorls 5 or 6, with
conspicuous, elevated, equidistant, obtuse lines across, forming grooves
between them; apex obtuse; lip reflected, rounded; pillar lip with a large
lamelliform tooth, which appears to revolve within the shell parallel to
the suture, a smaller raised line revolves nearer to the base within the
shell, but becomes obsolete before it arrives at the pillar lip; umbilicus
rather large. Breadth one-tenth of an inch. (2.5 mm.).
Type Locality: Not designated (Say, 1817).
Range: Quebec, Ontario, and Manitoba: eastern United States from
North Dakota and Louisiana to Alabama and Maine.
West Virginia Records
Boone County; Peytona (m&r), Racine (m&r), Seth (m&r). Braxton
County; Frametown (gkm), Gassaway (gkm). Cabell County; Barboursville
(ndr) (m&r), Lesage (m&r), Ona (m&r). Clay County; Clay (gkm). Grant
County; Greenland Gap (ndr). Greenbrier County; Alderson (gkm). Re-
nick (grh). Renick Valley (ndr), Ronceverte (m&r). Hampshire Comity;
Romney (gkm). Jefferson County; Harper’s Ferry (gwt) (cc), Shenandoah
Junction (msb). Kanawha County; Alum Creek (ndr), Hudnall (cbw).
South Charleston (ndr). Lincoln County; Branchland (m&r), Myra (m&r),
Sheridan (ndr). Logan County; Blair (m&r). Stone Branch (ndr). Mc-
Dowell County; Avondale (m&r), laeger (m&r). Panther (m&r). Marion
County; Curtisville (ndr), Glady Creek (ndr). Mercer County; Oakvale
(m&r), Princeton (m&r), Speedway (m&r). MonroeCounty; h\d^rson{G^M),
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VOL. 31
Salt Sulphur Springs (m&r), Sweetsprings (m&r), Waitesville (m&r),
Willow Bend (m&r). Morgan County; Berkeley Springs (ndr). Nicholas
County; Summersville (gkm). Pendleton County; Franklin (ndr) (gkm).
Pocahontas County; Marlinton (gkm). Putnam Comity; Poca (gkm).
Summers County; Riffle (gkm), Talcott (gkm). Upshur County; French
Creek (mrs. gkm) (gkm). Wayne County; Echo (m&r), Fort Gay (m&r),
Genoa (m&r). Webster County; Bolair (ndr) (gkm) (m&r). Wyoming
County; Baileysville (m&r), Pineville (m&r).
Remarks: S. labyrinthica has longer, more conspicuously unequal basal
folds, the first two much longer than the others, and the series does not
form an even curve as in affinis; the infraparietal lamella generally emerges
more ; the shell is smaller and generally less elevated and the spire is with
a more strongly convex outline. The whole shell, and especially the last
whorl is higher, the outline of the spire more convex, the basal folds are
more numerous, the color is duller, and the shell is less transparent than
in 5. cenea.
Thomas Say mentioned no definite locality for this species in the descrip-
tion of the type specimen, but the four cotypes at the Academy of Natural
Sciences of Philadelphia are labelled “Penna. ; Hyde and Mason,” and came
probably from the immediate vicinity of Philadelphia, which place Dr.
H. A. Pilsbry has selected as the type locality.
Habits: This shell is found under loose bark of logs, in half-decayed
wood, among dead leaves, and in sod at the base of trees.
Strobilops labyrinthica parietalis Pilsbry
(Plate 3, fig. 1)
Similar to 5. labyrinthica in the convexly conic, ribbed shell and weakly
emerging infraparietal lamella, but the lamellae penetrate more deeply,
being two-thirds and three-fourths of a whorl long. There is a blunt
columellar lamella and four basal folds, four within the side wall. The
whole base is ribbed in some examples, or smoothish, nearly finally striate
in others. Height 1.75, diameter 2.35 mm.
Type Locality: Ardsley, Montgomery County, Pennsylvania (Pilsbry,
1927).
Range: Pennsylvania, West Virginia, Tennessee, Florida, Alabama, and
Louisiana.
West Virginia Records
Boone County; Peytona (gb). Braxton County; Gassaway (gkm). Cabell
County; Ona (m&r). Calhoun County; Grantsville (ndr). Grant County;
1949
MacMillan: Land Snails of West Virginia
103
Greenland Gap (ndr). Greenbrier County ; (gkm), Renick Valley
(ndr), Ronceverte (m&r). Hampshire County; Romney (gkm). Kanawha
County; Dunbar (gkm), South Charleston (ndr). Lincoln County; Branch-
land (m&r), Myra (m&r), Sweetland (m&r). Logan County; Blair (m&r),
Sharpies {m&r). McDowell County; hwondsXe (m&r), laeger (m&r), Panther
(m&r). Mercer County; Princeton (m&r). Mingo County ;W\\\\2imson{M&.R) .
Monroe County; Salt Sulphur Springs (m&r), Sweetsprings (m&r), Waites-
ville (m&r). Willow Bend (m&r). Morgan County; Berkeley Springs (ndr).
Pendleton County; Franklin (gkm). Pocahontas County; Marlinton (gkm).
Summers County; Talcott (gkm). Upshur County; French Creek (gkm).
Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County; Bolair
(ndr) (gkm) (m&r). Wyoming County; Baileysville (m&r).
Strobilops affinis Pilsbry
(Plate 3, figs. 2, 3)
The shell is convexly conic with obtusely angular periphery; the base
moderately convex, becoming rather strongly so in its last third; glossy
brown, with pale apex; narrowly umbilicate, contained about 7)/^ to 8
times in the diameter. There are 6 moderately convex whorls, increasing
very slowly, the first two smooth, the rest sculptured with narrow, some-
what retractive ribs. The first half of the base is smooth, the ribs barely
passing over the peripheral angle and reappearing within the umbilicus,
but they continue weakly over the last half. The aperture is semi-lunar,
oblique. Peristome is well expanded, thickened within, its face convex
and fleshy-brown in color. Parietal callus moderately strong. The
parietal lamella emerges to the edge of the callus and penetrates inward
about two-thirds of a whorl. The infraparietal lamella is low and weak,
deeply immersed, not visible in front or basal view. Interparietal lamella
short and very weak. A third of a whorl within there is an oblique radial
series of about 8 folds: a short low lamella on the columellar axis, followed
by two folds larger and higher than the rest, and an oblique series running
to the suture, composed of 4 to 7 short subsequent folds. Height 2.5,
diameter 2.75 mm.
Type Locality: Not designated (Pilsbry, 1893).
Range: Canada; Maine to Minnesota, south to Texas, and east to Ala-
bama and New Jersey.
West Virginia Records
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r).
Braxton County; Gassaway (gkm). Cabell County; Lesage (m&r), Ona
(m&r). Calhoun County; Grantsville (ndr). Fayette County; Clifftop (gkm).
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Annals of the Carnegie Museum
VOL. 31
Grant County; Greenland Gap (ndr), Mount Storm (ndr). Greenbrier
County; Ronceverte (gkm) (m&r). Hampshire County; Romney (gkm).
Kanawha County; Alum Creek (ndr), Charleston (gkm), Dunbar (gkm),
Institute (gkm). Lincoln County; Myra (m&r), Sweetland (m&r). Logan
County; Sharpies (m&r). Stone Branch (ndr). McDowell County; laeger
(m&r). Panther (m&r). Mason County; Leon (ndr). Mercer County;
Princeton (m&r). Speedway (m&r). Mingo County; Williamson (m&r).
Monroe County; Salt Sulphur Springs (m&r), Sweetsprings (m&r). Morgan
County; Berkeley Springs (ndr). Nicholas County; Summersville (gkm).
Pendleton County; Franklin (ndr) (gkm). Upper Tract (gkm). Poca-
hontas County; Marlinton (gkm). Summers County ;Tdi\cott (gkm). Upshur
County; French Creek (mrs. gkm). Wayne County; Echo (m&r). Fort Gay
(m&r), Genoa (m&r). Wetzel County; Silver Hill (ndr). Wyoming County;
B alley sville (m&r), Oceana (m&r).
Remarks: This shell is somewhat larger than labyrinthica, thinner, with
the infraparietal lamella more deeply immersed and the baso-palatal folds
less unequal, all being rather short and forming a regular curve across
the base and up the outer wall. The outline of the spire is somewhat less
convex than in labyrinthica but more convex than in cenea.
Strobilops aenea Pilsbry
(Plate 3, fig. 4)
The shell is narrowly umbilicate, the width of umbilicus contained
about 6y2 times in the diameter of the shell, low conic, with obtuse,
rounded summit, the periphery distinctly but bluntly angular. The base
is somewhat flattened below the periphery, elsewhere moderately convex.
Whorls convex, slowly increasing, the first 13^ smooth, corneous, the
rest dark brown with red-gold gleam; sculptured with narrow riblets
which are somewhat oblique, retractive, rather fine and close. The base is
smoothish, marked with growth-strise only, except on its last third, where
the riblets of the upper surface continue over the base. The aperture is
semilunar, low but wide. Outer and basal lips brown, well expanded,
somewhat thickened, the columellar margin dilated. The parietal lamella
emerges to the edge of the parietal callus, penetrating inward a half-
whorl. Infraparietal lamella weakly emerging. Midway between the
lamellae there is a very weak, low, deeply-placed interparietal lamella.
These lamellae are nodose far within, the nodes roughened, shortly prickly.
The internal barrier, situated one-third of a whorl from the aperture, is
radial, but slightly oblique; it consists of a short, weak columellar fold
and four basal folds, visible through the shell; the second and fourth folds
from the axis are long, the first short, the third fold weak or sometimes
1949 MacMillan: Land Snails of West Virginia 105
wanting; there is no fold above the periphery. Height 1.5-2, diameter
2.4-2.75 mm.
Type Locality: Cazenovia, New York (Pilsbry, 1926).
Range: Massachusetts to Minnesota, south to Arkansas and Louisiana,
east to Florida, and north to New Jersey.
West Virginia Records i
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Brax-
ton County; Gassaway (gkm). Cabell County; Lesage (m&r), Ona (m&r).
Calhoun County; Grantsville (ndr). Fayette County; Cliff top (gkm).
Grant County; Greenland Gap (ndr), Mount Storm (ndr). Greenbrier
County; Ronceverte (gkm) (m&r). Hampshire County; Romney (gkm).
Kanawha County; Alum Creek (ndr). Charleston (gkm), Dunbar (gkm).
Institute (gkm). Lincoln County; Myra (m&r), Sweetland (m&r). Logan
County; Sharpies (m&r). Stone Branch (ndr). McDowell County; laeger
(m&r), Panther (m&r). Mason County; Leon (ndr). Mercer County; Prince-
ton (m&r). Speedway (m&r). Mingo County; Williamson (m&r). Monroe
County; Salt Sulphur Springs (m&r), Sweetsprings (m&r). Morgan County;
Berkeley Springs (ndr). Nicholas County; Summersville (gkm). Pendleton
County; Franklin (ndr) (gkm). Upper Tract (gkm). Pocahontas County;
Marlinton (gkm). Summers County; Talcott (gkm). Upshur County;
French Creek (mrs. gkm). Wayne County; Echo (m&r). Fort Gay (m&r),
Genoa (m&r). Wetzel County; Silver Hill (ndr). Wyoming County; Baileys-
ville (m&r), Oceana (m&r).
Family Succineid^
Shell ovate or oblong, whitish to greenish horn-colored, very thin and
fragile, imperforate, smooth or minutely wrinkled or striate; whorls 2}/^
to 4, the last usually very large and much expanded, more or less oblique;
spire short, acute, or elevated; suture moderately to deeply impressed;
aperture oblique, elongate-oval to roundly-oval or broadly rounded;
peristome simple, acute; columella deep, folded, or arcuate.
Genus Succinea Draparnaud
The characters of the genus are the same as those of the family.
The habits of the animal do not vary much from those of Helix. They
have been described as being amphibious, but are not in any sense this
way, as they live upon land exclusively and are air-breathers. Some of
them occupy situations very distant from any body of water, and others
106
Annals of the Carnegie Museum
VOL. 31
inhabit wet localities at the borders of swamps and ponds, attaching them-
selves to the leaves of plants growing out of the water.
The animals retire into their shells on the approach of winter and during
seasons of drought; every part of the body is retracted within in the plane
of the aperture, and over it is extended a membranous epiphragm. They
cannot, however, retract the body much beyond the plane of the mouth,
and the foot is never wholly drawn into the aperture of the mantle and
concealed by it.
Succinea ovalis Say
(Plate 3, fig. 5)
Shell ovate, pale green, yellowish-green, or amber colored to cinereous,
very thin and fragile, pellucid, sometimes roseate at apex; periostraca
shining, minutely wrinkled or striate ; whorls rather more than 3, the last
very large and much expanded and more or less oblique; spire very small,
not prominent nor pointed; suture distinct, impressed; aperture oval,
large, and expanded, more or less oblique ; columellar margin with a slight
testaceous glazing; columella thin, sharp, narrowed; peristome thin, its
edge blunted by the reflection of the periostraca. Greatest length 25 mm. ;
ordinary length 18 mm.
Type Locality: Not designated (Say, 1817).
Range: Newfoundland; Canada; United States, Maine to Montana,
south to Oklahoma, east to Florida, and north to New Jersey.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r). Braxton County; Shady Side (gkm). Cabell County;
Lesage (m&r). Clay County; Clay (gew) (gkm). Doddridge County; West
Union (frw). Fayette County; Clifftop (gkm). Jefferson County; Charles
Town (msb), Leetown (msb), Meyerstown (msb), Kanawha County; Nitro
(gkm). Lewis County; Jackson’s Mill (gkm). Logan County; Blair (ndr)
(m&r). Sharpies (m&r). Stone Branch (ndr). McDowell County; Avondale
(m&r). Panther (ndr). Mason County; Leon (ndr). Monongalia County;
Smithtown (ndr). Morgan County; (p&i). Nicholas County; Richwood
(ndr) (cjg). Pendleton County; Franklin (ndr). Pocahontas County; Mill
Point (ndr) (gkm). Preston County; Cranesville (mgn). Randolph County;
Cheat Bridge (ndr), Huttonsville (ndr) (gkm). Tucker County; Parsons
(mrs. gkm). Webster County; Bolair (m&r). Wyoming County; Baileysville
(m&r).
Remarks: This species lives under leaves and other forest debris that
1949 MacMillan: Land Snails of West Virginia 107
affords concealment during the day. In wet weather it may often be
seen on tree trunks at considerable heights above the ground, and some-
times in dry weather it may be found clinging to these high places, await-
ing more favorable conditions.
Succinea avara Say
(Plate 3, fig. 6)
Shell rather small, very thin and fragile, straw-colored, rosy, amber-
colored, or greenish; periostraca shining or presenting minute hairy pro-
cesses in the young; whorls 3, very convex, separated by a deep suture;
last whorl rather large, not much expanded ; spire very prominent, acute ;
aperture ovate, rounded at both extremities, about half as long as the shell.
Extreme length about 6 mm.
Type Locality: Northwest Territory (Say, 1822).
Range: Newfoundland; Canada; and the United States.
West Virginia Records
Boone County; Seth (m&r). Cabell County; Barboursville (gkm) (m&r).
Greenbrier County; Organ Cave (ndr). Renick Valley (ndr), Ronceverte
(m&r). Hampshire County; Romney (gkm). Jefferson County; Leetown
(msb), Meyerstown (msb). Kanawha County; Alum Creek (ndr), Dunbar
(gkm), Nitro (gkm), Tornado (gkm). Lewis County; Jackson’s Mill (gkm).
Lincoln County; Myra (m&r), Sweetland (m&r). Logan County; Sharpies
(m&r). McDowell County; Panther (m&r). Marion County; Curtisville
(ndr). High Point (ndr), Mt. Harmony (ndr), Rivesville (ndr). Mercer
County; Oakvale (m&r), Princeton (m&r), Spanishburg (m&r). Mingo
County; Taylorsville (m&r). Monongalia County; Coopers Rock (cbw),
Morgantown (hhs). Monroe County; Salt Sulphur Springs (ndr) (m&r),
Sweetsprings (m&r), Waitesville (m&r). Willow Bend (m&r). Pendleton
County; Circleville (ndr), Franklin (ndr), Judy Gap (gkm), Ruddle (gkm).
Upper Tract (ndr). Raleigh County; Daniels (m&r). Randolph County;
Huttonsville (gkm). Summers County; Riffle (gkm). Tucker County; Par-
sons (gkm). Wayne County; Fort Gay (m&r). Webster County; Bolair
(gkm) (m&r). Wyoming County; Oceana (m&r), Pineville (m&r).
Remarks: In the young shells the periostraca is covered with numerous
fine, hairy processes, which accumulate particles of dirt, and in this way
the shell is coated over its entire surface.
It is found under stones and fragments of wood in moist places, and
often on hillsides and other positions far removed from water.
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VOL. 31
Family Polygyrid^ Pilsbry
Shell discoidal or lens-shaped to globose-conic, wrinkled to striate,
usually with 53^ whorls, which are rounded or keeled; aperture broadly
open to narrowly transverse; lip reflected and expanded, toothless to
tridentate; umbilicus perforate or imperforate.
The members of this family are snails of humid country ; most eastern
Polygyridce being woodland snails living under dead wood or leaves or
under stones, chiefly in limestone areas, but also in sandstone and shaly
areas, and feeding mainly upon the mycelix of fungi. At night or after a
rain they may be found wandering about, but this habit is taken up more
freely with the young than the adults, often being found on plants where
the adults are under cover.
Genus Mesodon Rafinesque
Shell umbilicated or with the umbilicus closed, subglobose or orbicu-
larly depressed, thin, delicately striate, with sometimes a decussated
sculpture; whorls 5-6, regular; aperture rotundly lunar, sometimes nar-
rowed by a small denticle on the parietal wall and a lamellar-like denticle
or tooth on the inner edge of the peristome; peristome thickened, white,
expansively reflexed.
Mesodon albolabris albolabris (Say)
(Plate 4, fig. 1)
Shell thin, fragile; convex, imperforate; with six volutions, whorls
obtusely wrinkled across, and spirally striated with very fine impressed
lines, a little waved by passing over the wrinkles, both becoming extinct
towards the apex, which is perfectly smooth; aperture lunate, not angu-
lated at the base of the column, but obtusely curved, lip contracting the
mouth abruptly, widely reflected, flat and white. Breadth one inch
(25 mm.).
Type Locality: Harrigate, Country estate of Jacob Gilliams, outside of
Philadelphia (Say, 1816).
Range: Quebec and Ontario; Minnesota, Kansas and Louisiana east-
ward to Atlantic Ocean.
West Virginia Records
Barbour County: Nestorville (gkm), Philippi (gew) (gkm). Boone
County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Braxton County;
1949
MacMillan; Land Snails of West Virginia
109
Frametown (gkm), Gassaway (gkm), Shady side (gkm). Cabell County;
Barboursville (m&r), Ona (m&r). Calhoun County; Grantsville (ndr).
Clay County; Clay (gew) (gkm). Doddridge County; West Union (frw).
Fayette County; Cliff top (gkm). Grant County; Greenland Gap (ndr),
Petersburg (pr), Mount Storm (ndr). Greenbrier County; Alderson (gkm),
White Sulphur Springs (p&l). Hampshire County; Lehew (j. A. G. rehn),
Romney (gkm). Hancock County; Pugh town (ll). Hardy County; Moore-
held (ll), Wardensville (ghc). Harrison County; Bristol (ndr). Jefferson
County; Aldridge (msb), Bardane (msb), Bolivar (msb), Charles Town
(jpem), Engle (msb), Halltown (msb). Harper’s Ferry (ejc) (msb) (jpem)
(gwt), Jamestown (msb), Kearneysville (msb), Millville (msb), Mt.
Mission (jpem), Mt. Pleasant (msb), Morgan Grove (msb), Reedson
(msb), Shenandoah Junction (msb). Summit Point (msb), Uvilla (msb)
Kanawha County; Alum Creek (ndr). Charleston (gkm), Dunbar (gkm),
Hudnall (cbw). Institute (gkm). Lincoln County; Branchland (m&r),
Myra (m&r), Sheridan (ndr), Sweetland (m&r). Logan County; Blair
(m&r), Davin (ndr), Sharpies (m&r), Slagle (cg). McDowell County;
Avondale (m&r), laeger (m&r). Panther (m&r). Marion County; Curtis-
ville (ndr), Fairmont (ndr), Glady Creek (ndr), Mt. Harmony (ndr),
Rivesville (ndr). Mason County; Point Pleasant (ndr) (cbw). Mercer
County; Athens (mgn), Blueheld (m&r), Oakvale (m&r), Princeton
(m&r), Speedway (m&r). Mineral County; Keyser (ihm). New Creek (ndr).
Mingo Taylorsville (m&r), Williamson (m&r). Monongalia County;
Cheat Mt. (hhs). Coopers Rock (mgn) (ndr) (cbw), Sturgisson (ndr).
Monroe County; Sweetsprings (m&r). Morgan County; Berkeley Springs
(ndr). Nicholas County; Belva (ndr), Craigsville (gkm), Richwood (cjg)
(ndr), Swiss (mcz). Ohio County; Wheeling (mgn). Pendleton County;
Franklin (mgn) (ndr) (gkm), Judy Gap (gkm). Upper Tract (mgn) (gkm).
Pocahontas County; Dunmore Spring (gkm), Hillsboro (mgn), Marlinton
(mgn) (gkm). Mill Point (mgn) (ndr). Preston County; Cascade (ndr),
Corinth (mgn). Terra Alta (rlf) (mgn). Putnam County; Poca (gkm).
Raleigh County; Eccles (m&r). Flat Top Mt. (p&l). Shady Spring (m&r).
Randolph County; Helvetia (vs), Huttonsville (gkm). Summers County;
Hinton (usFc), Talcott (gkm). Tucker County; Parsons (gkm). Tyler
County; Sistersville (ghc). Upshur County; French Creek (mrs. gkm).
Wayne County; Echo (m&r). Fort Gay (m&r), Genoa (m&r). Webster
County; Bolair (ndr). Wetzel County; Silver Hill (ndr). Wirt County;
(wjf). Wood County; Boaz (cbw). Wyoming County; Baileysville (m&r),
Oceana (m&r), Pineville (m&r).
no
Annals of the Carnegie Museum
VOL. 31
Mesodon albolabris dentatus Tryon
(Plate 4, fig. 2)
Mesodon albolabris with a tooth on the parietal wall of the aperture,
closely resembling M. zaletus, but distinguished by its larger size, less
convex body whorl, broader lip, more transverse aperture, and generally
lighter color.
Type Locality: Not designated (Tryon, 1867).
Range: Much the same as M. albolabris.
West Virginia Records
Barbour County; Philippi (gkm). Greenbrier County; Alderson (gkm).
Logan County; Slagle (cg). Nicholas County; Richwood (ndr). Pocahontas
County; (ghc). Upshur County; French Creek (gkm).
Mesodon profundus (Say)
(Plate 4, fig. 3)
Shell pale horn-color; spire convex, very little elevated; whorls 5,
regularly rounded and wrinkled transversely, body whorl with a single
revolving line, which is almost concealed upon the spire by the suture,
but which passes for a short distance above the aperture; aperture dilated;
labrum reflected, white, and excepting near the superior angle flat, a
slightly projecting callus near the base on the inner edge; umbilicus large,
profound, exhibiting all the volutions to the apex. Transverse diameter
19/20 of an inch (23 ^ mm.).
Type Locality: Not designated (Say, 1821).
Range: New York to Minnesota, south to Louisiana, east to Alabama,
and north to Maryland.
West Virginia Records
Barbour County; Nestorville (gkm). Boone County; Racine (m&r).
Braxton County; Frametown (gkm). Brooke County; Bethany (mgn).
Cabell County; Kelleys (usfc). Calhoun County; Grantsville (ndr). Clay
County; Clay (gkm). Grant County; Bayard (ndr). Greenbrier County;
Renick Valley (ndr). Jefferson County; Harper’s Ferry (B. Chambers)
(gwt). Kanawha County; Hudnall (cbw). Logan County; Blair (ndr)
(m&r), Davin (ndr). McDowell County; Avondale (m&r). Panther (m&r).
Marion County; Fairmont (ndr), Smithville (cbw). Marshall County;
Bannon (ndr), Cameron (gb), Powhattan Point (frw). Mercer County;
Speedway (m&r). Mineral County; Keyser (ihm). Monongalia County;
1949
MacMillan: Land Snails of West Virginia
111
Cheat Mt. (hhs). Monroe County; Sweetsprings (m&r). Nicholas County;
Swiss (mcz). Ohio County; Wheeling (mgn) (cb) (ndr) (cbw). Pocahontas
County; Durbin (mgn). Wirt County; (wjf). Wyoming County; Oceana
(m&r), Pineville (m&r).
Mesodon mitchellianus (Lea)
(Plate 4, fig. 4)
Shell imperforate, depressed, conoid-globose, thin, without crowded
striae and very crowded, decussating, microscopic lines, pellucid, horn-
color, polished, spire briefly conoid ; whorls 5, moderately convex, gradually
increasing, the last ventricose, subconstricted and briefly deflected an-
teriorly ; aperture diagonal, lunate, subperlaceous within ; peristome white,
thickened, its terminations slightly converging, subequally reflected, that
of the columella narrow, adherent, or subdilated and spreading. Greater
diameter 163^, lesser 143^ mm., height 10 mm.
Type Locality: Ohio (Lea, 1839).
Range: New York, Pennsylvania, Ohio, Michigan, Indiana, Illinois,
Kentucky, West Virginia, and Virginia.
West Virginia Records
Brooke County; Bethany (cbw). Mercer County; Spanishburg (mcz).
Ohio County; Wheeling (mgn) (ndr) (cbw) .
Remarks: This species resembles M. clausus closely in size, texture, and
color, but may be known by the somewhat different shape and the closed
umbilicus, the columellar terminations of the lip being appressed over it,
while in M. clausus a cleft always remains open.
Mesodon thyroidus (Say)
(Plate 4, fig. 6)
Shell thin, fragile, convex, umbilicate; whorls wrinkled or rather with
equidistant, gradually elevated, obtuse lines, and spirally striate with
numerous, impressed lines; lip widely reflected, white and flat before,
partially concealing the umbilicus; pillar lip furnished with a very oblique
tooth. Breadth four-fifths to nine-tenths of an inch. (19-22 mm.).
Type Locality: Not designated (Say, 1817).
Range: Quebec and Ontario; Maine to Minnesota, south to Texas, east
to Alabama, and north to New Jersey.
West Virginia Records
Barbour County; Philippi (mrs. gkm) (gkm). Berkeley County; (mgn).
Boone County; Peytona (ndr) (m&r), Seth (m&r). Braxton County; Frame-
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VOL. 31
town (gkm), Gassaway (gkm), Shadyside (gkm). Brooke County; Bethany
(CBw). Cabell County; Barboursville (gkm) (m&r), Lesage (m&r), Milton
(aeo) (cbw). Calhoun County; Grantsville (ndr). Clay County; Clay
(gew) (mrs. gkm) (gkm). Doddridge County; Smithburg (ndr), West
Union (frw). Grant County; Greenland Gap (ndr), Maysville (ll),
Mount Storm (ndr), Petersburg (pr). Greenbrier County; Alderson (ndr)
(gkm), Organ Cave (ndr), Renick Valley (ndr), Ronceverte (gkm) (m&r).
Hampshire County; Romney (gkm). Hardy County; Moorefield (ll).
Jefferson County; Aldridge (msb), Bardane (msb), Bloomery (msb),
Bolivar (msb), Charles Town (msb), Duffield (msb), Engle (msb), Gibson-
town (msb). Hall town (msb), Harper’s Ferry (ejc) (msb) (gwt), Kable-
town (msb), Kearneysville (msb). Keystone (msb), Leetown (msb),
Loudoun Heights (jpem). Middleway (msb), Millville (msb), Morgan
Grove (msb), Ranson (msb), Shenandoah City (msb), Shenandoah Junc-
tion (msb), Shepherdstown (msb) (jpem), Skeetersville (msb), Uvilla
(msb). Kanawha County; Alum Creek (ndr), Dunbar (gkm), Hudnall
(cbw), Nitro (gkm). South Charleston (ndr), Tornado (gkm). Lewis
County; Weston (gkm). Lincoln County; Branchland (m&r), Myra (m&r),
Sheridan (ndr), Sweetland (m&r). Logan County; Davin (ndr). Stone
Branch (ndr). McDowell County; Panther (ndr). Marion County; Fair-
mont (ndr), High Point (ndr), Rivesville (ndr), Smithville (cbw).
Marshall County; Cameron (gb), Kent (ndr). Mason County; Leon (ndr),
Maysville (cbw). Mercer County; Princeton (m&r), Spanishburg (m&r).
Speedway (m&r). Mineral County; Burlington (gkm), Ridgeley (aeo).
Mingo County; Williamson (m&r). Monongalia County; Cheat Mt. (aeo),
Morgantown (hhs). Monroe County; Alderson (gkm). Salt Sulphur Springs
(m&r), Sweetsprings (m&r). Willow Bend (m&r). Nicholas County; Belva
(ndr), Richwood (cjg). Ohio County; Wheeling (mgn) (ndr) (cbw).
Pendleton County; Circleville (ndr), Franklin (ndr) (gkm). Ruddle (gkm).
Upper Tract (mgn) (ndr) (gkm). Pocahontas County; Dunmore Spring
(gkm), Marlinton (gkm). Preston County; Cascade (ndr). Putnam County;
Poca (gkm). Red House (cbw). Randolph County; Elkins (ghc), Huttons-
ville (gkm). Ritchie County; Pennsboro (frw). Summers County; Riffle
(gkm), Talcott (gkm). Tucker County; Parsons (gkm). Tyler County;
Friendly (cbw). Wayne County; Echo (m&r). Fort Gay (m&r), Sandy
River (Sanderson Smith). Webster County; Bolair (ndr) (gkm). Wetzel
County; New Martinsville (cbw). Wirt County; (wjf). Wood County; Boaz
(cbw), Williamstown (frw). Wyoming County; Baileysville (m&r).
Remarks: Many specimens of M. thyroidus possess a small tooth on the
1949
MacMillan: Land Snails of West Virginia
113
parietal wall of the aperture, which, apparently, was not present on the
specimen upon which Say based his description of this species. Dr. Pilsbry
believes that thyroidus was described from a small form of this species in
and around Philadelphia and has selected one from near the mouth of
Wissahickon Creek as the neotype.
Mesodon pennsylvanicus (Green)
(Plate 4, fig. 7)
Shell subglobose ; spire elevated ; whorls six or seven, with numerous
oblique wrinkles or striae, suture deeply impressed; epidermis smooth and
of an olive brown color; umbilicus closed or masked; aperture slightly
contracted at base, a small callosity on the inner margin of the lip, near
its lower angle; shell rather more than half an inch in diameter (17 mm.).
Type Locality: Near Chartiers Creek, Washington County, Pennsyl-
vania (Green, 1827).
Range: Pennsylvania, Ohio, Indiana, Michigan, Illinois, Missouri,
Kentucky, and West Virginia.
West Virginia Records
Doddridge County; West Union (frw). Marion County; Fairmont (ndr),
Smithville (cbw). Marshall County; Cameron (gb). Monongalia County;
Morgantown (hhs). Monroe County; Sweetsprings (m&r). Ohio County;
Wheeling (cb) (mgn) (ndr) (cbw).
Remarks: M. pennsylvanicus is somewhat like M. clausus in size and
color, but it is perforate, the whorls are more closely coiled, the aperture
of more irregular shape and the basal lip is straight or obscurely toothed.
The spire is usually high, much as in M. elevatus, but this varies.
The type locality in Washington County, Pennsylvania, is near Canons-
burg, as M. pennsylvanicus was collected, along with Amby stoma jeffersoni-
ana, in the moist ground near Chartiers Creek in the vicinity of Jefferson
College, which is now a part of Washington and Jefferson College at
Washington, Pa.
Mesodon zaletus (A. Binney)
(Plate 4, fig. 9)
Shell imperforate, convex, somewhat ventricose ; epidermis of a uniform
yellow horn- or russet color; whorls between 5 and 6, with fine parallel
striae crossing them obliquely; body whorl large and ventricose; suture
114
Annals of the Carnegie Museum
VOL. 31
well marked and distinct; aperture rounded, contracted by peristome, the
plane of the aperture making a considerable angle with the plane of the
base; peristome thickened, white, reflected, the basal portion subdentate,
parietal wall with a prominent, white, oblique tooth; umbilicus covered.
Greater diameter 28, lesser 23 mm,; height 17 mm.
Type Locality: Cincinnati, Ohio (A. Binney, 1837).
Range: Lake Erie Islands, Ontario; New York to Wisconsin, south to
Arkansas, east to Alabama, and north to Maryland.
West Virginia Records
Barbour County; Nestorville (gkm). Boone County; Racine (m&r).
Braxton County; Shadyside (gkm). Calhoun County; Grantsville (ndr).
Grant County; Greenland Gap (ndr), Mount Storm (ndr). Greenbrier
County; Renick Valley . Hampshire County; Romney {gkm) . Kanawha
County; Dunbar (gkm), Hudnall (cbw). Logan County; Blair (m&r),
Davin (ndr). Sharpies (m&r), Stone Branch (ndr). McDowell County;
Avondale (m&r). Panther (m&r). Marion County; Fairmont (ndr), Ham-
mond (ndr), Kingmont (grh), Mt. Harmony (ndr). Marshall County;
Bannon (ndr), Cameron (gb). Mercer County; Speedway (pcb) (m&r).
Monongalia County; Cheat Mt. (hhs). Coopers Rock (cbw), Smithtown
(ndr), Sturgisson (ndr). Nicholas County; Richwood (ndr). Ohio County;
Wheeling (mgn) (ndr) (cbw) . Pendleton County; Onega (mgn) . Pocahontas
County; Cass (mgn), Durbin (mgn), Marlinton (gkm). Mill Point (mgn)
(ndr) (gkm). Preston County; Cold Spring Park (grh). Randolph County;
Huttonsville (mgn) (gkm). Summers County; Talcott (gkm). Taylor
County; Valley Falls (ndr). Tucker County; Parsons (gkm). Wayne County;
Fort Gay (m&r). Webster County; Bolair (ndr) (m&r), Webster Springs
(J. A. Moore), Wyoming County; Pineville (m&r).
Remarks: Though resembling M. albolabris in many respects, M.
zaletus differs in being smaller, more convex, and the body whorl is more
ventricose. The peristome is less flat and broad and is sometimes a little
grooved. The aperture is more round, and the plane of the mouth, instead
of being flattened in the direction of the plane of the base, is much more
upright, making a considerable angle with the base of the shell. However,
many -forms of albolabris have the plane of the aperture at a considerable
angle with the base of the shell, nearly as much as in zaletus; the only
difference being the absence of a tooth in albolabris and the presence of it
in zaletus.
1949
MacMillan: Land Snails of West Virginia
115
Mesodon sayanus (Pilsbry)
(Plate 4, fig. 10)
Shell somewhat depressed; spire convex, very little elevated; whorls 5,
rounded, regularly but not prominently wrinkled, and grooved trans-
versely; aperture moderate; labrum reflected, with a slightly projecting
dentiform callus near the base on the inner edge ; labrum with an oblique
tooth on the middle; umbilicus rather large, profound, exhibiting all the
volutions. Breadth rather more than seven-tenths of an inch (27 mm.).
Type Locality: New York (Say, 1824).
Range: Quebec and Ontario; Maine to Michigan, south to Tennessee
and North Carolina, and north to Maryland.
West Virginia Records
Barbour County; Nestorville (gkm). Boone County; Peytona (m&r),
Racine (m&r). Cabell County; Barboursville (ndr), Ona (m&r). Clay
County; Clay (gkm). Fayette County; Cliff top (gkm). Greenbrier County;
Alderson (gkm). Kanawha County; Hudnall (cbw). Lincoln County;
Myra (m&r). Logan County; Blair (ndr) (m&r), Davin (ndr). Sharpies
(m&r), Slagle (cg). McDowell County; Avondale (m&r). Panther (m&r).
Marion County; High Point (ndr). Marshall County; Cameron (gb).
Mercer County; Athens (mgn). Mingo County; Taylorsville (m&r). Monon-
galia County ; (hhs). Coopers Rock (ndr) (cbw). Maiden Spring
Run (grh), Sturgisson (ndr). Nicholas County; Lockwood (gkm), Rich-
wood (ndr) (cjg). Snake Den Mt. (ndr), Swiss (mcz). Pendleton County;
Franklin (ndr) (gkm), Judy Gap (gkm). Upper Tract (mgn). Pocahontas
County; Arbovale (mgn), Dunmore Spring (gkm), Durbin (mgn). Marlin-
ton (mgn) (gkm). Mill Point (ndr) (gkm), Minnehaha Springs (mgn).
Travellers Repose (ghc). Preston County; Cranesville (mgn). Randolph
County; Helvetia (vs), Huttonsville (gkm). Webster County; Bolair (ndr)
(m&r). Wyoming County; Oceana (m&r), Pineville (m&r).
Remarks: The large umbilicus, light color, thin shell, and small but
constant tooth at the base of the columella, are characteristics of this
species. M. sayanus was first described by Thomas Say (1824) as Helix
diodonta, but, as this name had been used previously, H. A. Pilsbry (1906)
changed the terminology to its present status, sayanus.
Mesodon dentiferus (A. Binney)
(Plate 5, fig. 1)
Shell imperforate, flattened, convex on the upper surface, convex below;
epidermis yellowish horn-color, immaculate; spire depressed; whorls 5,
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Annals of the Carnegie Museum
VOL. 31
with delicate, parallel, oblique striae ; sutures distinct, not deeply impressed ;
aperture contracted by the peristome, flattened towards the plane of the
base; peristome thickened, white, broadly and abruptly reflected; parietal
wall with a prominent, white, toothlike process, nearly parallel with the
lower margin of the aperture, not projecting towards the umbilicus, base
convex. Greater diameter 23, lesser 18 mm.; height 10 mm.
Type Locality: Eastern slopes of Green Mountains of Vermont. (A.
Binney, 1840).
Range: Quebec and Ontario; New England States; New York and
Pennsylvania, south to Tennessee and North Carolina.
West Virginia Records
Barbour County; Philippi (gkm). Braxton County; Shadyside (gkm).
Clay County; Clay (gkm). Fayette County; Clifftop (gkm). Grant County;
Greenland Gap (ndr). Mount Storm (ndr). Marion County; Fairmont
(ndr). High Point (ndr), Mt. Harmony (ndr). Monongalia County; Cheat
Mt. (hhs), Sturgisson (ndr). Monroe County; Alderson (gkm). Nicholas
County; Richwood (ndr) (cjg), Snake Den Mt. (ndr), Summersville
(gkm). Pendleton County; Onega (mgn). Pocahontas County; Cass (mgn),
Durbin (mgn), Hillsboro (mgn) (gkm), Mill Point (mgn) (ndr) (gkm),
Seebert (cjg). Preston County; Cranesville (mgn), Terra Alta (mgn).
Randolph County; Cheat Bridge (gkm), Helvetia (vs), Huttonsville (mgn)
(gkm). Summers County; Talcott (gkm). Tucker County; Parsons (gkm).
Webster County; Bolair (gkm) (m&r). Wirt County; (wjf).
Remarks: M. dentiferus can be distinguished from M. albolabris dentatus
by the more depressed and thinner shell, narrower aperture, broader lip,
and more conspicuous parietal tooth.
Mesodon clausus (Say)
(Plate 4, fig. 5)
Shell subimperforate, conoidly semi-globose, rather solid, with crowded,
rib-like striae, yellow horn-color; spire sub-regularly conoid; whorls S]/^,
rather convex, gradually increasing, penultimate subangular, the last
rounded, anteriorly subconstricted, and briefly deflected; umbilicus nar-
row, almost covered by the reflected peristome; aperture diagonal, sub-
regularly lunate; peristome with a heavy white thickening, uniformly
subangularly reflected, its columellar portion subdilated. Greater di-
ameter 183^^, lesser 16 mm.; height 113^ mm.
Type Locality: Illinois (Say, 1821).
Range: Pennsylvania to Minnesota, south to Oklahoma, east to Georgia,
and north to Maryland,
1949
MacMillan: Land Snails of West Virginia
117
West Virginia Records
Berkeley County; Martinsburg (ghc). Fayette County; Oak Hill (um).
Remarks: This species might be confused with the small form of M.
thyroidus known as M. thyroidus hucculentus, but M. clausus has no
parietal tooth in contrast with M. thyroidus. Dr. Pilsbry believes that in
some forms of thyroidus the umbilicus may become closed. In clausus the
ere vis always remains open.
Mesodon appressus appressus (Say)
(Plate 4, fig. 8)
Shell depressed, brownish horn-color; whorls five, depressed, forming an
angle on the external one, more acute near the superior angle of the labrum,
with numerous transverse, elevated, equidistant lines, with interstitial
grooves; umbilicus covered over with calcareous matter, but concave
within; aperture moderate; labrum dilated, reflected, white, margined
with brownish ; near the base, appressed to the body-whorl, and covering
umbilicus; a slight projecting dentiform angle on the inner middle; labrum
with a strong, prominent, oblique, compressed, white tooth, which gradu-
ally slopes and becomes obsolete towards the umbilicus. Breadth three-
fifths of an inch. (18 mm.).
Type Locality: Not designated. (Say, 1821).
Range: Pennsylvania to Indiana, south to Alabama, and north to
West Virginia.
West Virginia Records
Braxton County; Frametown (gkm). Cabell County: Barboursville (gkm)
(m&r), Lesage (m&r), Milton (cbw). Kanawha County; Hudnall (cbw),
Nino (gkm). South Charleston (ndr). Tornado (gkm). Lincoln County;
Branchland (m&r), Sheridan (ndr). Logan County; Stone Branch (ndr).
McDowell County; Avondale (m&r), laeger (m&r). Panther (ndr) (m&r).
Mason County; Leon (ndr). Mercer County; Bluefield (m&r). Spanishburg
(m&r). Speedway (mgn) (pcb) (m&r). Mingo County; Taylorsville (m&r),
Williamson (m&r). Monongalia County; Morgantown (H. Kahl). Summers
County; Hinton (usfc), Talcott (gkm). Wayne County; Fort Gay (m&r),
Genoa (m&r).
Remarks: Thomas Say, in his description of M. appressus, did not
notice the minute sculpture of the shell consisting of papillae. The micro-
scopic papillae are found between the rib striations and are strung into
spiral lines on the last whorl; on the base they are closely scattered.
118
Annals of the Carnegie Museum
VOL. 31
Dr. Pilsbry has selected specimens from Ohio as the neotypes, designat-
ing Gallipolis as the type locality. Say stated in his description, “Inhabits
banks of the Ohio and Missouri. This species is very common on the banks
of the Ohio below Gallipolis; I also found it near Council Bluff.” His
description was drawn evidently from the Ohio forms in which there is no
upper tooth.
Mesodon appressus perigraptus (Pilsbry)
(Plate 4, fig. 11)
The shell is depressed, glossy, chamois-colored. Striations very fine
and close, weaker on the base in front, the later whorls cut by engraved
spiral lines throughout. The broadly reflected lip has no tooth in the outer
arc, basal margin with a blade-like thickening within, truncate at its outer
end. Parietal tooth short and high, widely separated from the axial callus,
but generally a very low callus ridge runs nearly to the latter (but this is
often weak or wanting). Height 8.5-12.6 mm., diameter 15.8-23.4 mm.;
whorls 5^.
Type Locality: Woodville, Jackson County, Alabama. (Pilsbry, 1894).
Range: West Virginia, Kentucky, Tennessee, North Carolina, South
Carolina, Georgia, Alabama, Mississippi, Louisiana, and Arkansas.
West Virginia Records
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r).
Cabell County; Barboursville (ndr), Ona (m&r). Calhoun County; Grants-
ville (ndr). Clay County; Clay (gew) (gkm). Fayette County; Cliff top
(gkm). Grant County; Stony River Dam (mgn). Greenbrier County; Alder-
son (gkm). Kanawha County; St. Albans (um), South Charleston (ndr),
Tornado (gkm). Lincoln County; Branchland (m&r), Myra (m&r). Logan
County; Blair (ndr) (m&r), Davin (ndr), Sharpies (m&r), Slagle (cg).
McDowell County; laeger (m&r). Panther (m&r.) Mingo County; Ta^ylors-
ville (m&r). Nicholas County; Belva (ndr), Lockwood (gkm). Rich wood
(ndr), Summersville (gkm). Raleigh County; Daniels (m&r). Shady Spring
(m&r). Wayne County; Fort Gay (m&r). Wyoming County; Baileysville
(m&r), Pineville (m&r).
Remarks: The smooth engraved lines, usually developed strongly down
to the umbilical callus, distinguishes this subspecies from M. appressus.
Also, there is no trace of a superior tooth on the outer arc of the lip.
1949 MacMillan: Land Snails or West Virginia 119
Genus Triodopsis Rafinesque
Shell moderate to large, umbilicate or imperforate, depressed and
carinate to subglobose-conic. Whorls 5-7, the last wider, more or less
deflexed in front. Surface may be striate, with or without spiral lines or
hirsute. Aperture lunate, typically obstructed by three teeth, two on the
lip and one on the parietal wall but any or all are often absent; peristome
reflected and thickened within.
Most of the species live around decaying logs and under and upon de-
caying leaves in forests. Some prefer shady, leaf-carpeted and rocky
bark of other trees. Most species come from their hiding places in the
warm days of early spring, and during rainy weather in summer. They
may be found crawling upon the dead leaves, or ascending nettles, the
leaves of which they eat. They may also be found adhering to the lower
surfaces of nettle leaves in the summer after a rain, but never have been
known to ascend trees.
Triodopsis platysayoides (Brooks)
(Plate 4, figs. 13, 14)
The five whorls are flattened and only very slightly convex; base flat-
tened and slightly inflated at the aperture. The shell is thin and translu-
cent but not fragile. The color is light-horn with a yellowish area on the
exterior surface of the peristome which exhibits a punctate appearance.
This area of punctation extends back past the constriction of the peris-
tome and over the first four or five oblique striae. The whorls striated
obliquely, terminating in the wide, inverted cone-shaped umbilicus which
exhibits all of the volutions to the apex. The first whorl is smooth. The
umbilicus seems slightly excentric due to the deflection of the body whorl
at the aperture. The aperture is oblong-lunate. The lip is reflected, flat,
white, and quite heavy; the basal edge forming quite a straight line,
thickened with a deposit of callus. The basal angles thickened and ex-
tending into the body whorl, visible in the umbilicus as a whitish band for
the full length of the body whorl. On the parietal wall is a thick, obtusely
pointed, tongue-shaped tooth, its apex directed tangentially out and away
from the aperture ; it is quite similar to the parietal tooth in M. dentiferus.
Greater diameter 22 mm., lesser 18 mm.; height 8 mm.
Type Locality: Coopers Rock, Monongalia County, West Virginia
(Brooks, 1932).
Range: West Virginia Record; Monongalia County; Coopers Rock
(mgn) (ndr).
Remarks: It is a Triodopsis in which the lip teeth have become de-
generate as in some forms of T. tridentata.
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Annals of the Carnegie Museum
VOL. 31
Triodopsis fraudulenta fraudulenta (Pilsbry)
(Plate 5, fig. 6)
“The shell is similar to T. tridentata but slightly less depressed, cin-
namon-buff. The whorls are rather closely coiled, the last descending
steeply in front, strongly contracted behind the spreading lip, with a dis-
tinct external impression at the position of the outer lip-tooth. The em-
bryonic whorl has short radial lines below the suture (when not worn).
Last whorl is regularly striate, with traces of spiral lines below the suture
(or none), and densely, minutely papillose at umbilicus and below suture;
aperture is distinctly dished, nearly closed by the large teeth. The lip is
rather widely reflected in its outer and basal arcs, the basal margin
straightened. Within the outer lip is a wide, receding tooth; a narrow
tooth in the basal margin, and a high, nearly straight parietal tooth.
Height 7.7-9 mm., diameter 14.7-16.7 mm., 6 whorls.”
Type Locality: Morgan County, West Virginia (Pilsbry, 1894).
Range: Pennsylvania, West Virginia, and Virginia.
West Virginia Records
Braxton County; Gassaway (gkm), Shadyside (gkm). Grant County;
Greenland Gap (ndr). Mount Storm (ndr). Greenbrier County; Organ
Cave (ndr). Renick Valley (ndr), Ronceverte (gkm) (m&r). Hampshire
County; Capon Bridge (jpem). Hancock County; Pugh town (ll). Jefferson
County; Mt. Pleasant (msb). Lewis County; Weston (gkm). Mercer County;
Princeton (m&r). Mineral County; Keyser (ihm). New Creek (ndr),
Ridgeley (aeo). Monroe County; Alderson (gkm). Gap Mills (ndr). Salt
Sulphur Springs (ndr) (m&r), Waitesville (m&r). Morgan County; Ber-
keley Springs (ndr), Largent (ndr). Nicholas County; Belva (ndr),
Summersville (c&a) (mcz). Ohio County; Wheeling (mgn). Pendleton
County; Franklin (ndr) (gkm), Judy Gap (gkm). Upper Tract (mgn).
Pocahontas County; Cass (mgn), Hillsboro (mgn) (gkm), Marlinton (mgn)
(gkm). Mill Point (ndr) (gkm), Minnehaha Springs (mgn). Randolph
County; Huttonsville (gkm). Summers County; Riffle (gkm), Talcott
(gkm). Tucker County; Canaan Valley (mgn). Webster County; Bolair
(ndr).
Remarks: T. fraudulenta was based upon a mountain form characterized
by the development of a heavy, nearly straight ledge within the basal lip,
upon which the basal tooth stands. In an oblique view this tooth appears
to pass down the inner side of the straight callus ledge. The basal lip struc-
ture is sometimes similar to T. fallax, but the decidedly wider umbilicus
and the straight parietal tooth of fraudulenta, as well as the greater size,
1949
MacMillan: Land Snails of West Virginia
121
differentiates them at once. The outer lip tooth is very broad and deeply
receding, while the parietal tooth is long and straight.
Triodopsis fraudulenta vulgata Pilsbry
(Plate 4, fig. 12)
“The shell is cream buff to sea-foam yellow. It differs from T. tridentata
and T. tridentata juxtidens by having the upper lip-tooth wider than its
fellows, and distinctly bent inward, and the peristome is more or less
dished. Compared with T. fraudulenta fraudulenta the teeth of T. f.
vulgata are not so large and the aperture is more open. The parietal tooth
is straight or nearly so, and not so long as in fraudulenta, leaving much
more space between it and the two lip teeth. The basal lip, while well
thickened within, does not have the prominent straight callus of frau-
dulenta. The bay between the lip teeth is more symmetrical. The um-
bilicus is somewhat well-like beyond the enlargement at the last whorl,
and wider than in T. t. juxtidens, showing the first whorl plainly at the
bottom. Height 7.3-10.4 mm.; diameter 13.5-19.5 mm.”
Type Locality: Columbus, Ohio (Pilsbry, 1940).
Range: Ontario; New York to Michigan and Illinois, south to Missouri
and Alabama.
West Virginia Records
Barbour County; Nestorville (gkm). Grant County; Greenland Gap
(ndr). Greenbrier County; Alderson (ndr), Renick Valley (ndr). White
Sulphur Springs (p&l). Jefferson County; Middleway (msb). Kanawha
County; Hudnall (cbw). Marshall County; Kent (ndr). Mineral County;
Burlington (gkm). Monroe County; Alderson (gkm), Nicholas County;
Craigsville (gkm), Richwood (ndr). Ohio County; Wheeling (cbw).
Pendleton County; Franklin (ndr), Judy Gap (gkm), Onega (mgn). Upper
Tract (ndr). Pocahontas County; Durbin (mgn). Mill Point (gkm).
Randolph County ; Cheat Bridge (ndr) (gkm). Mill Creek {gril). Summers
County; Talcott (gkm). Tucker County; Canaan Valley (mgn). Webster
County; Bolair (gkm). Wood County; Boaz (cbw).
Remarks: The aperture of this variety is more dished than in T. t.
juxtidens, and there is an arcuate impression behind the peristome in the
region back of the outer lip-tooth, which is lacking in juxtidens.
Triodopsis rugeli (Shuttleworth)
(Plate 5, fig. 2)
Shell imperforate, orbicularly’ convex, with granulate striations and few
hairs, waxen horn-color; spire short, obtuse; whorls S}/2y rather convex,
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Annals of the Carnegie Museum
VOL. 31
the last suddenly falling in front and strongly contracted at the aperture ;
aperture depressed, narrowed by a tongue-shaped, flexuose, strong parietal
denticle; peristome reflected, within thickened, its right termination with a
large, obtuse, very deeply seated tooth (whose position is marked on the
exterior of the shell by a groove or pit), the basal terminus furnished with
a smaller, transverse, submarginal denticle. Greater diameter 13, lesser
113^ mm.; height 634 rnni.
Type Locality: Tennessee (Shuttleworth, 1852).
Range: West Virginia; Virginia; Kentucky; Tennessee; North Carolina;
South Carolina; Georgia, and Alabama.
West Virginia Records
McDowell County; Avondale (m&r), Panther (ndr) (m&r). Mercer
County; Bluefield (m&r), Spanishburg (mcz).
Remarks: T. rugeli is distinct from T. inflecta by the decidedly more
deeply immersed and larger tooth on the outer lip and more strongly
curved parietal tooth.
Triodopsis denotata (Ferussac)
(Plate 5, fig. 3)
Shell depressed, with elevated lines, forming grooves between them;
epidermis fuscous rugose with very numerous minute tuberculous acute
prominences; volutions five, depressed above, beneath rounded, forming an
obtuse angle exteriorly, which is more acute near the termination of the
labrum, umbilicus covered with a white callous; aperture contracted by
the labrum ; labrum reflected widely, white, two profound, obtuse sinuses
on the inner side above the middle, forming a prominent distinct tooth
between them and a projecting angle near the middle of the lip; labium
with a large, prominent, white tooth, placed perpendicularly to the whorl,
and obliquely to the axis of the shell, and nearly attaining the umbilical
callus. Greatest breadth four-fifths of an inch (20 mm.).
Type Locality: Illinois (Say, 1821).
Range: Ontario; Maine to Wisconsin, south to Arkansas and Missi-
ssippi, east to Alabama, and north to New Jersey.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Braxton County;
Frametown (gkm), Gassaway (gkm), Shadyside (gkm). Brooke County ^
Williamsburg (cbw). Cabell County; Barboursville (m&r). Calhoun County;
Grantsville (ndr). Doddridge County; West Union (grh). Grant County;
Greenland Gap (ndr). Mount Storm (ndr). Hampshire County; Romney
(gkm). Kanawha County; South Charleston (ndr). Logan County; Blair
1949
MacMillan: Land Snails of West Virginia
123
(m&r), Davin (ndr), Sharpies (m&r). McDowell County; Avondale (m&r)^
Panther (m&r). Marion County; Curtisville (ndr), Fairmont (grh), High
Point (ndr). Marshall County; Bannon (ndr), Kent (ndr). Mercer
County; Speedway (m&r). Mineral County; Keyser (ihm), New Creek
(ndr). Mingo County; Williamson (m&r). Monongalia County; Cheat Mt.
(hhs), Coopers Rock (cbw), Sturgisson (ndr). Nicholas County; Rich-
wood (ndr) (cjg) . Ohio County; Wheeling (mgn) (ndr) (cbw) . Pendleton
County; Circleville (ndr). Pocahontas County; Marlinton (gkm). Preston
County; Cascade (ndr). Randolph County; Helvetia (vs), Huttonsville
(gkm). Tucker County; Parsons (gkm). Upshur County; French Creek
(MRS. gkm) (gkm). Webster County; Bolair (ndr). Wood County; Boaz
(cbw). Wyoming County; Pineville (m&r).
Remarks: T. denotata is very distinct from all others except T. ohstricta,
which is much more keeled at the periphery. The surface is roughened by
stiff periostracal hairs, more strongly developed than in any other Helix
of the region.
This species was first described by Thomas Say in 1821 as Helix pal-
liata. However, J. D. W. Hartmann had given this same specific name to a
Helix in 1807. By this means, H. palliata of Say became a synonym of
H. denotata of Ferussac, mentioned by him in 1821. Since that name was
not validated by a description or figure, H. notata was substituted, as this
appeared to be the first published name to be defined, and that by De-
shayes in 1830. It is possible that notata was an error for denotata, or an
emendation of it, as Deshayes attributed it to Ferussac, and referred to his
"Tableau Systematique.”
Triodopsis tridentata tridentata (Say)
(Plate 5, fig. 4)
Shell depressed, spire very little raised, brownish or horn-color; whorls
5, crossed by numerous raised, equidistant, acute lines, separated by regu-
lar grooves; aperture lunate, three toothed; teeth placed triangularly, one
on the pillar lip situated diagonally; outer lip abruptly contracting the
aperture, widely reflected and white, furnished with two of the teeth re-
sembling projecting angles; umbilicus moderate. Half an inch wide
(14-16 mm.).
Type Locality: Middle States (Say, 1816).
. Range: Quebec and Ontario; Maine to Minnesota, south to Arkansas,
east to Georgia, and north to New Jersey.
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Annals of the Carnegie Museum
VOL. 31
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gew) (mrs. gkm) (gkm).
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Braxton
County; Frametown (gkm), Gassaway (gkm). Brooke County; Bethany
(CBw), Williamsburg (cbw). Cabell County; Barboursville (ndr) (gkm)
(m&r), Lesage (m&r), Milton (aeo) (cbw), Ona (m&r). Calhoun County;
Arnoldsburg (grh), Grantsville (ndr). Clay County; Clay (gew) (mrs.
gkm) (gkm). Doddridge County; Sherwood (ndr), Smithburg (ndr), West
Union (frw). Fayette County; Cliff top (gkm). Oak Hill (afa) (mcz).
Grant County; Greenland Gap (ndr), Maysville (ll). Mount Storm (ndr).
Greenbrier County; Alderson (gkm). Organ Cave (ndr), Renick Valley
(ndr), White Sulphur Springs (p&l). Hampshire County; Capon Bridge
(jpem). Hardy County; Moorefield (ll). Harrison County; Bristol (ndr).
Jackson County; Odaville (cbw). Jefferson County; Aldridge (msb),
Bakerton (msb), Bloomery (msb), Charles Town (jpem), Halltown (msb).
Harper's Ferry (ndr) (gwt) (jpem), Jamestown (msb), Kabletown (msb),
Loudoun Heights (msb), Mechanicstown (msb), Middleway (msb), Morgan
Grove (msb), Mt. Mission (jpem), Shepherdstown (msb), Uvilla (msb).
Kanawha County; Alum Creek (ndr). Charleston (gkm), Dunbar (gkm),
Hudnall (cbw). Institute (gkm), South Charleston (ndr). Tornado (gkm).
Lewis County; Jackson’s Mill (mrs. gkm) (gkm), Weston (gkm). Lincoln
County; Branchland (m&r), Myra (m&r), Sheridan (ndr), Sweetland
(m&r). Logan County; Blair (ndr) (m&r), Davin (ndr), Sharpies (m&r)
Stone Branch (ndr). McDowell County; Avondale (m&r), laeger (m&r).
Panther (ndr) (m&r). Marion County; Curtisville (ndr), Fairmont (ndr)
(grh), Glady Creek (ndr), Hammond (ndr). High Point (ndr), King-
mont (grh), Smithville (cbw). Marshall County; Bannon (ndr), Cameron
(gb), Kent (ndr), Moundsville (cbw), Powhattan Point (frw). Mason
County; Leon (ndr). Point Pleasant (ndr) (cbw). Mercer County; Blue-
field (mgn) (m&r), Oakvale (m&r), Princeton (m&r), Spanishburg (m&r)
(afa) (c&a) (mcz). Speedway (pcb) (m&r). Mineral County; Keyser (ihm).
New Creek (ndr). Mingo County; Taylorsville (m&r), Williamson (m&r).
Monongalia County; Cheat Mt. (hhs). Coopers Rock (mgn) (ndr) (cbw),
Morgantown (hhs). Smith town (ndr), Sturgisson (ndr). Monroe County;
Alderson (gkm). Gap Mills (m&r). Salt Sulphur Springs (ndr) (m&r),
Sweetsprings (grh) (m&r), Waitesville (m&r). Willow Bend (m&r).
Morgan County; Berkeley Springs (ndr), Largent (ndr). Nicholas County;
Lockwood (gkm), Powell Mt. (mrs. ga). Rich wood (cjg), Summersville
1949
MacMillan: Land Snails of West Virginia
125
(gkm). Ohio County; Wheeling (mgn) (ndr) (cbw). Pendleton County;
Franklin (ndr) (gkm), Judy Gap (gkm), Onega (mgn). Pocahontas
County; Cass (mgn), Greenbank (gkm), Hillsboro (gkm), Marlinton (gkm).
Mill Point (ndr). Travellers Repose (ghc). Preston County; Cascade
(ndr). Cheat Mt. (grh), Cranesville (mgn). Terra Alta (mgn). Putnam
County; Poca (gkm). Raleigh County; Daniels (gkm) (m&r), Eccles (m&r).
Randolph County; Cheat Bridge (ndr) (gkm), Helvetia (vs), Huttonsville
(gkm), Nettie (ndr). Rich wood (ndr). Ritchie County; Pennsboro (frw).
Summers County; Riffle (gkm), Talcott (gkm). Taylor County; Valley
Falls (ndr). Tucker County; Canaan Valley (ndr), Parsons (mrs. gkm)
(gkm). Tyler County; Friendly (cbw). Upshur County; French Creek
(mrs. gkm) (gkm). Wayne County; Echo (m&r). Fort Gay (m&r). Webster
County; Bolair (ndr) (gkm) (m&r), Hodon Creek (grh). Wetzel County;
Silver Hill (ndr). Wirt County; (wjf). Wood County; Boaz (cbw), Parkers-
burg (frw). Wyoming County; Baileysville (m&r), Oceana (m&r). Pine-
ville (m&r).
Remarks: Dr. Pilsbry believes that Thomas Say described the small
form of this species common on the gneiss around Philadelphia and has
selected a specimen from Montgomery County, near the northern edge of
Philadelphia, as the neotype. On calcareous soils it is often larger than the
typical specimens, attaining diameters up to 18.5 mm.
T. tridentata is more abundant on limestone soils, but it lives everywhere
wherever there is some shade, with moderate moisture, herbage, dead
leaves or woodshelter. Though hilly country is preferred, it is not found
in the higher Appalachians.
Triodopsis tridentata juxtidens (Pilsbry)
(Plate 5, fig. 5)
This form is distinguished from typical tridentata by the lower position
of the upper lip-tooth, the latter part of the parietal lamella being directed
towards this tooth or to a point above it, whilst in the type the lip-teeth
are more separated and the latter portion of the parietal lamella is di-
rected towards a point on the peristome between them. The variety is
more coarsely striated also. The number of whorls (5) is the same in the
variety and type. Altitude 6-7, diameter 11-14 mm.
Type Locality: Cave Town, Maryland (Pilsbry, 1894).
Range: New Jersey to Ohio, south to Tennessee and North Carolina.
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Annals of the Carnegie Museum
VOL. 31
West Virginia Records
Boone County; Peytona (m&r), Seth (m&r). Braxton County; Frame-
town (gkm), Gassaway (gkm), Shadyside (gkm). Brooke County; Bethany
(mgn). Cabell County; Barboursville (m&r). Calhoun County; Grantsville
(ndr). Clay County; Clay (gkm). Fayette County; Cliff top (gkm). Grant
County; Petersburg (pr). Greenbrier County; Alderson (ndr), Organ Cave
(ndr). Hampshire County; Romney (gkm). Jefferson County; Aldridge
(msb), Bolivar (msb), Duffield (msb), Gibsontown (msb), Halltown (msb),
Harper’s Ferry (ejc), Kabletown (msb), Keystone (msb), Mechanicstown
(msb), Millville (msb), Morgan Grove (msb), Shenandoah Junction (hap)
(msb), Shepherdstown (msb), Skeetersville (msb), Uvilla (msb). Kanawha
County; Dunbar (gkm). Institute (gkm). Lewis County; Jackson’s Mill
(gkm), Weston (gkm)). Lincoln County; Branchland (m&r). Logan County;
Davin (ndr), Slagle (cg). Marshall County; Cameron (gb). Mercer
County; Princeton (m&r). Mineral County; Burlington (gkm). Monon-
galia County; Cheat Mt. (hhs). Monroe County; Sweetsprings (grh).
Morgan County; Berkeley Springs (ndr), Largent (ndr). Nicholas County;
Craigsville (gkm). Rich wood (ndr), Summersville (gkm). Pendleton
County; Circleville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm),
Macksville (mgn). Upper Tract (mgn) (ndr) (gkm). Pocahontas County;
Dunmore Spring (gkm), Greenbank (gkm), Marlinton (gkm). Mill Point
(gkm), Travellers Repose {gy^c) . Randolph County; Helvetia (vs), Huttons-
ville (gkm). Summers County; Talcott (gkm). Wayne County; Fort Gay
(m&r), Genoa (m&r).
Triodopsis rugosa Brooks and MacMillan
(Plate 5, fig. 8)
Shell somewhat depressed, costate, reddish horn-color, with a narrow
umbilicus. Whorls 534> flattened above and rounded below, body-whorl
rounded. Suture impressed. Rib-striations prominent and continue
undiminished into the umbilicus; interstitial striae few and faint; spiral
striae weak, hardly discernible ; a few granulations are visible on the upper
whorls, becoming more numerous on the basal part of the shell. Stria-
tions on embryonic whorl and a quarter very weak, on next whorl and a
half they become broken into short bars, and on remaining whorls they
become gradually larger until they are rib-like on the last two whorls.
Aperture lunate, tridentate; peristome white, broad, thickened within;
outer margin bearing a small, narrow, squarish tubercle which is bent
slightly inward; basal lip bearing a marginal tubercle; parietal denticle
1949 MacMillan: Land Snails of West Virginia 127
tongue-shaped, entering slightly into the aperture, and separated from the
umbilical margin of the peristome by a narrow channel. Umbilicus nar-
row, deep, the inner whorls hardly perceptible. Greater diameter 10.9,
lesser diameter 9.2, height 5.6 mm.
Type Locality: Damp Ravine, Blair Mountain, one mile southwest of
Blair, Logan County, West Virginia (Brooks and MacMillan, 1940).
Range: West Virginia records; Logan County; Blair (m&r), Slagle (cg).
Nicholas County; Richwood (ndr), Summersville (gkm). Putnam County;
Buffalo (cBw).
Remarks: The essential features of this species are the thickening of
the outer lip, which produces a sloping buttress below the outer lip and the
rough sculpture of the shell produced by the prominent and well-separated
rib-striations. The parietal tooth is strongly developed as in T. fallax
and T. vannostrandi.
Triodopsis fallax (Say)
“The shell is narrowly umbilicate, the umbilicus about 9 times in the
diameter, depressed, with dome shaped or conoidal spire; deep olive-buff
(or dilute snuff-brown). Whorls closely coiled, the last most convex
above the middle, deeply constricted behind the peristome. First whorl
smooth with a band of short radial striae below the suture. Later whorls
with little gloss, closely thread-striate, with minute papillae around the
umbilicus. The aperture is irregularly trilobed. The reflected peristome
is white, thickened within, with a blunt, rather wide inwardly bent tooth
in the outer margin, and a callous ledge surmounted by a transverse tu-
bercle in the middle of the basal margin. Parietal tooth rather long and
strongly arcuate or angularly curved. On the columellar axis there is a
white ridge (sometimes rising to form a tubercle) formed by a continua-
tion inward of the ledge upon the basal lip. Height 7. 5-8. 2 mm. ; diameter
11.8-12.4 mm.; S}/2 whorls.” (Pilsbry).
Type Locality: Vicinity of Philadelphia, Pennsylvania (Say, 1825).
Range: New Jersey and Pennsylvania, south to Tennessee and Georgia.
West Virginia Records
Hampshire County; Junction (ndr). Jefferson County; Jamestown
(msb). Summit Point (msb). Mineral County; Burlington (gkm).
Remarks: T. fallax differs from T. fraudulenta by having the upper
tooth smaller and not as deeply seated in the aperture, possessing a nar-
rower umbilicus, and the presence of a callus tubercle within the axis of
the lip. There is also a callous ridge from the basal tooth to the base of the
columella within. The parietal tooth is somewhat tongue-shaped.
128
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VOL. 31
Genus Stenotrema Rafinesque
Shell small, compact, imperforate or umbilicate; subglobose, globose-
depressed, or lens-shaped, the periphery varying from rounded to acutely
keeled ; surface dull, smooth, generally hairy. Whorls 5-6, closely revolving,
the last suddenly deflected in front. Aperture basal, narrow, obstructed
by an oblique blade-like parietal tooth parallel to the reflexed basal lip,
the latter often notched in the middle. Last whorl generally having in its
last fourth a short transverse partition on its axis. Embryonic whorls
generally with a dense pattern of radially lengthened granules, or some-
times radially striate.
Stenotrema stenotrema (Ferussac)
(Plate 5, fig. 7)
The imperforate shell is depressed-globose, with low conoid spire,
strongly convex base and rounded periphery; from buckthorn brown to
cinnamon brown in color. Embryonic whorls with the usual sculpture,
the later whorls unevenly striate, set with papillae bearing rather short
hairs in forwardly descending trends ; these continue over the base, where
they are fine near the axis. The narrow aperture has buff to brown borders.
Parietal tooth high, but not rising to the level of the basal lip, leaning
towards the latter, gently curved, its outer end turning into the inter-
denticular sinus (and often further curved into a very short hook at the
end), a low and inconspicuous buttress between parietal tooth and termi-
nations of the outer lip. The basal lip has a thin, wholly adnate outer
margin ; inner margin nearly straight in basal view, with a small but well-
marked median notch, with slightly raised, callous border. The inter-
denticular sinus is moderate, deep and narrow. The outer lip bears a low
tooth or none. The fulcrum is well developed with convex edge. Height
5.3-8.6 mm., diameter 7.8-12.8 mm.; 5-5^ whorls.
Type Locality: Indiana (Ferussac, 1842).
Range: Virginia to Missouri, south to Louisiana and Georgia.
West Virginia Records
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r).
Braxton County; Frametown (gkm). Cabell County; Barboursville (ndr)
(m&r), Milton (aeo) (cbw), Ona (m&r). Calhoun County; Grantsville
(ndr). Fayette County; Clifftop (gkm), Gauley Bridge (grh). Jefferson
County; Harper’s Ferry (jpem). Kanawha County; Charleston (gkm),
Hudnall (cbw), St. Albans (um), South Charleston (ndr). Lincoln County;
Branchland (m&r), Myra (m&r), Sheridan (ndr), Sweetland (m&r).
1949
MacMillan: Land Snails of West Virginia
129
Logan County; Blair (ndr) (m&r), Davin (ndr), Sharpies (m&r), Slagle
(cg). McDowell County; Avondale (m&r), laeger (m&r), Panther (ndr)
(m&r). Mercer County; Princeton (m&r), Spanishburg (mcz) (c&a) (m&r).
Speedway (pcb) (m&r). Mingo County; Taylorsville (m&r), Williamson
(m&r). Monroe County; Gap Mills (m&r). Salt Sulphur Springs (m&r).
Nicholas County; Rich wood (ndr), Summersville (mrs. ga). Putnam
County; Poca (gkm). Raleigh County; Eccles (m&r). Summers County;
Talcott (gkm). Wayne County; Fort Gay (m&r). Wyoming County; Baileys-
ville (m&r), Pineville (m&r).
Remarks: S. stenotrema is larger than S. hirsutum, with a decidedly
narrower aperture, a higher parietal tooth, a wider basal lip, and the in-
terdenticular sinus is narrower and deeper.
Stenotrema hirsutum (Say)
(Plate 5, fig. 12)
The shell is depressed globose with rather low, convexly conoid spire,
rounded periphery and strongly convex base, cinnamon-buff to clay color.
After the initial smooth stage the embryonic shell has close, radially
lengthened granules. The later whorls have short, moderately stiff hairs
with rounded bases, arranged in oblique series, over the usual microscopic
lineolation. The parietal tooth is slightly bowed, rather high but lower
than the level of the basal lip, slightly sinuous in the outer third, the
end not turning towards the interdenticular sinus. The basal lip is rather
broad, its outer edge closely appressed, the calloused inner edge having a
large and deep, slightly oblique notch with slightly raised edges. The
interdenticular sinus is rather broadly rounded. Tooth in the outer lip
is rather well developed, bluntly conic. Height 4.3-5. 7 mm., diameter
6.2-9. 6 mm.; 5 whorls.
Type Locality: Not designated (Say, 1817).
Range: Ontario; Maine to Minnesota, south to Louisiana and Georgia.
West Virginia Records..
Barhour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (gb) (m&r), Seth (m&r). Braxton County; Gassaway (gkm).
Cabell County; Barboursville (gkm) (m&r), Lesage (m&r), Milton (cbw),
Ona (m&r), Calhoun County; Grantsville (ndr). Clay County; Clay (mrs.
gkm) (gkm). Doddridge County; West Union (frw). Grant County; Green-
land Gap (ndr). Mount Storm (ndr). Greenbrier County; Alderson (gkm),
Renick Valley (ndr). Hampshire County; Junction (ndr), Romney (gkm).
Harrison County; Bristol (ndr). Jefferson County; Harper’s Ferry (gwt)
130
Annals of the Carnegie Museum
VOL. 31
(msb), Mt. Mission (jpem). Kanawha County; Dunbar (gkm), Hudnall
(cBw), Institute (gkm), South Charleston (ndr), Tornado (gkm). Lewis
County: Jackson’s Mill (gkm). Lincoln County; Branchland (m&r), Myra
(m&r), Sweetland (m&r). Logan County; Blair (m&r), Stone Branch (ndr).
McDowell County; laeger (m&r). Panther (m&r). Marion County; Fair-
mont (ndr), Glady Creek (ndr). High Point (ndr), Kingmont (grh).
Mineral County; Burlington (gkm). New Creek (ndr), Ridgeley (aeo).
Mingo County; Williamson (m&r). Monongalia County; Coopers Rock
(cBw). Monroe County; Gap Mills (m&r). Salt Sulphur Springs (m&r),
Sweetsprings (m&r) . Morgan County; (afa) . Ohio County; Wheeling (ndr)
(CBw). Pendleton County; Circleville (ndr), Franklin (ndr) (gkm), Judy
Gap (gkm). Upper Tract (ndr) (gkm). Pocahontas County; Cass (mgn),
Marlinton (gkm). Mill Point (ndr) (gkm). Putnam County; Buffalo (cbw),
Red House (cbw). Raleigh County; Flat Top Mt. (p&l). Randolph County;
Helvetia (vs), Huttonsville (gkm). Ritchie County; Pennsboro (frw).
Summers County; Talcott (gkm). Taylor County; Grafton (frw). Tucker
County; Parsons (mrs. gkm) (gkm). Tyler County; Friendly (cbw).
Upshur County; French Creek (mrs. gkm) (gkm). Wayne County; Echo
(m&r). Fort Gay (m&r), Genoa (m&r). Webster County; Bolair (ndr)
(gkm) (m&r). Wetzel County; Proctor (vs). Wirt County; (afa) (wjf).
Wyoming County; Baileysville (m&r), Oceana (m&r).
Remarks: This species is smaller than S. stenotrema; the parietal tooth
is not so high and its outer end not at all turned into the interdenticular
sinus. The notch is generally wider.
Stenotrema edvardsi (Bland)
(Plate 5, figs. 10, 11)
Shell imperforate, lenticular, carinate, the carina obsolete near the
aperture, rather thin, beneath the epidermis pale brown, the epidermis
dark-chestnut color, with numerous minute curved hair-like processes
lying flat upon, and attached to the epidermal surface of the upper whorls
in the direction of the incremental striae, the epidermis at the base covered
with acute, raised, transverse tubercles, most numerous, and having erect
bristles near the aperture; spire convex-conoid; whorls five, flattened,
gradually increasing, the last gibbous above, suddenly but slightly de-
flected ; apex minutely granulate; base convex, little indented in the umbili-
cal region, and with impressed spiral lines beneath the epidermis; suture
deeply impressed; aperture oblique, transverse, auriform, narrowed by a
slender slightly arcuate lamelliform parietal tooth extending across from
the umbilical axis, and terminating with a short angular deflection within
1949 MacMillan: Land Snails of West Virginia 131
the aperture; upper margin of the peristome acute, scarcely reflected, lower
margin slightly arcuate, depressed, slightly reflected, and partially ap-
pressed to the body whorl, with a tooth-like callus within, having an al-
most obsolete notch in the center. Diameter major 9, minor 8, alti-
tude 5, mm.
Type Locality: Mountains, Fayette or Greenbrier Counties, West Vir-
ginia (Bland, 1858).
Range: West Virginia, Virginia, Kentucky, and Tennessee.
West Virginia Records
Boone County; Peytona (m&r), Racine (m&r). Fayette County; (T.
Bland), Cliff top (mrs. gkm) (gkm)., Greenbrier County; (T. Bland).
Kanawha County; Alum Creek (ndr), Hudnall (cbw), Kanawha City
(afa) (T. Bland). Lincoln County; Myra (m&r), Sheridan (ndr). Logan
County; Blair (m&r), Davin (ndr). McDowell County; Panther (m&r).
Mercer County ; (afa) (m&r). Speedway (pcb) (m&r). Mingo
County; Taylorsville (m&r), Williamson (m&r). Nicholas County; Lock-
wood (gkm), Richwood (ndr). Raleigh County; Eccles (m&r). Wayne
County; Fort Gay (m&r). Wyoming County; Baileysville (m&r), Oceana
(m&r), Pineville (m&r).
Remarks: The peripheral carination, the rather low parietal tooth, not
projecting as far as the level of the basal lip, the very small basal notch,
shallow interdenticular sinus and absence of a buttress between parietal
tooth and upper end of lip, are its main characters. It is very near S.
barbigerumj but higher and with less developed fringe.
Stenotrema leai (Ward)
(Plate 6, fig. 1)
The umbilicate shell is depressed with low, convexly conoid spire of
narrow, very closely coiled whorls; very bluntly subangular or rounded at
the periphery, which is above the middle, the base convex. Dilute Isa-
bella color, slightly translucent and somewhat glossy. Embryonic whorls
with the usual fine sculpture of radially lengthened granules, often running
into striae, and generally almost or quite effaced in adult shells. Later
whorls with faint lines of growth, the last having very short, delicate
hairs rising from little acute bases, which alone remain in most adult
shells. The oval-lunate, oblique aperture has a brownish or white peris-
tome thickened within, well reflected in its outer and basal margins.
Parietal tooth short, white, straight, standing obliquely on the thin parie-
tal callus, and typically not prolonged towards and aperture. The fulcrum
is quite short with convex edge. Height 3.9-5. 7 mm., diameter 6.7-9.4
mm.; 5^-6Fi whorls.
132
Annals of the Carnegie Museum
VOL. 31
Type Locality: Above Thunder Bay, Lake Huron, Michigan (Rackett,
1822).
Range: New Brunswick, Ontario; Maine to North Dakota, south to
Texas and Alabama.
West Virginia Records
Barbour County; Philippi (gew) (gkm). Clay County; Clay (gkm).
Doddridge County; West Union (frw). Greenbrier County; Ronceverte
(m&r). Kanawha County; Hudnall (cbw). Lewis County; Weston (gkm).
Marion County; High Point (ndr). Mineral County; New Creek (ndr).
Monroe County; Salt Sulphur Springs (m&r). Willow Bend (m&r.)
Nicholas County; Rich wood (ndr), Summersville (gkm). Ohio County;
Wheeling (mgn) (cbw). Pendleton County; Circleville (ndr), Franklin
(ndr) (gkm). Pocahontas County; Dunmore Spring (gkm), Marlinton
(gkm), Mill Point (gkm). Randolph County; Huttonsville (gkm). Upshur
County; French Creek (mrs. gkm) (gkm).
Remarks: S. leai differs from S. fraternum by the closer coils of the
spire, usually having one more turn in specimens of about equal size ; the
umbilicus is typically larger; the surface is smoother, more or less glossy,
hairs or their scars being delicate and generally almost or quite absent in
adults; and the parietal tooth is short and does not extend to and into the
umbilicus.
Stenotrema fraternum fraternum (Say)
(Plate 6, fig. 2)
The shell is imperforate or nearly covered perforate, with convexly
conoid spire of closely coiled whorls, which are noticeably wider than in
S. leai the rather strongly convex base is impressed around the axis;
the rounded periphery is above the middle. Color of cartridge buff to
tawny olive. Embryonic whorl closely covered with radially lengthened
granules, often somewhat indistinct. Later whorls densely covered with
short hairs or their bases. Aperture much as in S. leai: parietal tooth
short, rather long, nearly straight but with the ends often a trifle turned
towards the basal lip. Basal lip well thickened within. The fulcrum is
rather short. Height 5. 2-6.9 mm., diameter 7.8-10.5 mm.; 53^-5 2/3
whorls.
Type Locality: Pennsylvania (Say, 1824).
Range: Ontario; Maine to Minnesota, south to Missouri and Alabama.
1949
MacMillan: Land Snails of West Virginia
133
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gew) (mrs. gkm) (gkm).
Boone County; Peytona (m&r), Racine (m&r). Braxton County; Gassaway
(gkm), Shadyside (gkm). Cabell County; Lesage (m&r). Calhoun County;
Grantsville (ndr). Doddridge County; Smithburg (ndr). Grant County;
Greenland Gap (ndr), Mount Storm (ndr). Greenbrier County; Organ
Cave (ndr), Renick Valley (ndr), Ronceverte (m&r). Hampshire County;
Junction (ndr). Jefferson County; Harper's Ferry (jpem). Kanawha
County; Dunbar (gkm), Hudnall (cbw). Lewis County; Weston (gkm).
Lincoln County; Sweetland (m&r). Logan County; Blair (m&r). Marion
County; High Point (ndr). Mason County; Leon (ndr). Mercer County;
Oakvale (m&r), Princeton (m&r). Mineral County; New Creek (ndr).
Monongalia County; Coopers Rock (ndr) (cbw), Morgantown (hhs).
Monroe County; Alderson (gkm), Gap Mills (m&r), Salt Sulphur Springs
(ndr) (m&r). Willow Bend (m&r). Morgan County; Largent (ndr).
Nicholas County; Craigsville (gkm), Summersville (gkm). Pendleton
County; Circleville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm). Upper
Tract (ndr) (gkm). Pocahontas County; Dunmore Spring (gkm), Hills-
boro (gkm), Marlinton (gkm), Mill Point (ndr). Raleigh County; Shady
Spring (m&r). Randolph County; Helvetia (vs), Huttonsville (gkm).
Summers County; Talcott (gkm). Taylor County; Valley Falls (ndr).
Tucker County; Parsons (gkm). Tyler County; Friendly (cbw). Upshur
County; French Creek (mrs. gkm) (gkm). Wayne County; Genoa (m&r).
Webster County; Bolair (ndr) (m&r). Wetzel County; Silver Hill (ndr).
Wyoming County; Baileysville (m&r).
Remarks: Differs chiefly from S. leai by the noticeably wider whorls in
an apical view and the short, but long parietal tooth that extends to
and into the umbilicus.
Stenotrema fratemum cavum (Pilsbry and Vanatta)
(Plate 6, fig. 3)
Shell larger than S.fraternum; more openly umbilicate, deeply impressed
or excavated around the umbilicus. Parietal tooth short, its ends about
equally remote from the termination of the lip. Fulcrum well developed,
notched above and below. Whorls 53^. Pilosity fine and short, about as in
S. fraternum. Altitude 6, diameter 10.5 mm.
Type Locality: Cazenovia, Madison County, New York (Pilsbry and
Vanatta, 1911).
134 Annals of the Carnegie Museum vol. 31
Range: Ontario, Quebec, and New Brunswick; Maine to Minnesota,
south to Iowa and Maryland.
West Virginia record: Jefferson County; Harper’s Ferry (gwt).
Family ENDODONTiDiE Pilsbry
Shell globose to depressed, rugose to costulately striate, or radially
striate, and widely umbilicated. Whorls Z]/2 to 63/^, gradually increasing,
rounded to carinate, or bicarinate. White, straw-colored to dark reddish-
horn, or greenish ; with or without brownish revolving bands, interrupted
and obliquely arranged reddish patches and spots, or yellowish lines.
Aperture semilunate or rhomboidal, or stirrup-shaped, oblique, furnished
within with or without a single, basal tooth, or 1 to 3 pairs of minute,
conical, white teeth on external wall ; peristome simple, acute, extremities
joined by a thin to heavy callus. Umbilicus perspective to funnel-shaped,
showing all inner volutions.
Genus Punctum Morse
Shell minute, thin, subdiscoidal but with convex spire, openly umbili-
cate; unicolored; whorls about 4, convex, the apical 13^ smooth, rather
distinctly demarked from the following whorls, which have oblique striae
or irregular riblets and excessively fine spiral striae ; the last whorl cylindri-
cal, not descending in front. Aperture lunate, rounded; lip simple, thin.
Punctum minutissimum (I, Lea)
(Plate 6, fig. 4)
Shell subglobose, above obtusely conical, below convex; Brown horn-
colour, very minutely striate, umbilicate; spire short; sutures impressed;
whorls four, round ; lip acute. Diameter 0.06., length 0.04 of an inch. (1.5
X 1 mm.).
Type Locality: Cincinnati, Ohio (I. Lea, 1841).
Range: Newfoundland; Canada; United States.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r), Racine (m&r), Seth (m&r). Braxton County; Frametown
(gkm), Gassaway (gkm). Cabell County; Barboursville (m&r), Lesage
(m&r), Ona (m&r). Fayette County; Clifftop (gkm). Grant County; Green-
1949
MacMillan: Land Snails of West Virginia
135
land Gap (ndr), Mount Storm (ndr). Greenbrier County; Alderson (ndr)
(gkm), Renick (grh), Renick Valley (ndr). Hampshire County; Romney
(gkm). Jefferson County; Bloomery (jpem), Shepherdstown (usb) . Kanawha
County; Tornado (gkm). Lewis County; Jackson’s Mill (gkm). Lincoln
County; Branchland (m&r), Myra (m&r), Sweetland (m&r). Logan County;
Blair (m&r), Sharpies (m&r). McDowell County; Avondale (m&r), laeger
(m&r). Panther (m&r). Marion County; Curtisville (ndr), Glady Creek
(ndr). High Point (ndr). Mason County; Leon (ndr). Mercer County;
Bluefield (m&r), Oakvale (m&r), Princeton (m&r), Speedway (m&r).
Mineral County; Burlington (gkm). New Creek (ndr). Mingo County;
Williamson (m&r). Monroe County; Alderson (gkm), Sweetsprings (grh)
(m&r). Willow Bend (m&r). Nicholas County; Lockwood (gkm), Sum-
mersville (gkm). Pendleton County; Circleville (ndr), Franklin (ndr)
(gkm), Judy Gap (gkm), Upper Tract (ndr). Pocahontas County; Dun-
more Spring (gkm), Greenbank (gkm), Marlinton (gkm). Mill Point
(gkm). Putnam County; Poca (gkm). Randolph County; Cheat Bridge
(gkm), Helvetia (vs). Summers County; Riffle (gkm), Talcott (gkm).
Tucker County; Parsons (gkm). Upshur County; French Creek (mrs.
gkm). Wayne County; Eccho (m&r). Fort Gay (m&r). Webster County;
Bolair (ndr) (m&r) . Wetzel County; Silver Hill (ndr) . Wyoming County;
Pineville (m&r).
Remarks: Due to the small size of this species it can easily be overlooked
by collectors. However, it may be found beneath the bark of fallen trees,
on small limbs and twigs of fallen timber, and in forest debris.
Punctum vitreum H. B. Baker
(Plate 6, figs. 5, 6)
Shell: minute, depressed turbinate ; light corneous to vitreus and al-
most transparent. Whorls: 43^2 (maximum), later ones rounded but de-
pressed {i.e., elliptical in outline); last whorl slightly descending; suture
strongly impressed (although more broadly so than in P. minutissimum) .
Embryonic whorls: 1^, quite high; first whorl irregularly wrinkled but
almost smooth; last ^ whorl with fine spiral striae becoming more and
more distinct. Later whorls: major growth riblets high and thin, weakly
arcuate just above periphery; interspaces with 5-10, very low but sharp,
minor growth ridges, which are crossed by fine, spiral lines so that minute
squares are formed. Umbilicus: open, about 3.3 times in major diameter
of shell. Aperture: depressed; altitude 0.44, diameter 0.52 mm., slightly
oblique (about 25° to axis of shell). Peristome: simple and sharp; weakly
arcuate above the periphery. Pallial complex as in P. conspectumjaliscoense.
136 Annals of the Carnegie Museum vol. 31
Type Locality: Pleasure Gardens, New Braunfels, Comal County,
Texas (H. B. Baker, 1930).
Range: New Jersey to Iowa, south to Texas and Alabama.
West Virginia Records
Boone County; Peytona (m&r). Braxton County; Gassaway (gkm).
Hampshire County; Romney (gkm). Jefferson County; Harper’s Ferry
(jpem). Kanawha County; Tornado (gkm). McDowell County; Avondale
(m&r), Panther (m&r). Marion County; High Point (ndr). Mason County;
Leon (ndr). Monroe County; Sweetsprings (m&r). Pendleton County;
Franklin (ndr) (gkm), Judy Gap (gkm). Webster County; Bolair (m&r).
Remarks: This species is about the size of minutissimum but has more
depressed whorls and the sculpture of the Toltecia group {i.e., the contrast
between the major and minor growth riblets is much greater and the for-
mer are more widely spaced). P. vitreum also has thinner and more lightly
colored epidermis, and its sculpture is more delicately and sharply cut.
Under the binocular microscope, P. minutissimum looks as if it were
moulded out of bronze, but P. vitreum appears as if it were cut from yel-
lowish crystal.
Genus Helicodiscus Morse
Shell small, disk or coin-shaped, with flat spire and broad, shallow
umbilicus. Whorls numerous (3-5), convex and closely coiled, smooth,
slightly wrinkled, or spirally striated or lirate, the last lacking or possess-
ing one or several pairs of tubercular teeth within, situated upon the basal
or outer wall. Aperture lunate, lip thin, simple.
The species of this genus are found under bark or in the interstices of
wet and decaying wood, and under layers of wet leaves and stones in
damp places in forests.
Helicodiscus parallelus (Say)
(Plate 7, fig. 1)
Shell widely umbilicated, discoidal; epidermis greenish; whorls 4,
visible on the base of the shell as well as above, with numerous equidistant,
parallel, raised lines revolving upon them; suture much impressed; aper-
ture remote from axis, semi-lunate, narrow, not expanding; peristome
acute, thin ; umbilicus wide, forming a concave depression of the base, each
volution visible to the apex; within the aperture, on the external circum-
ference, are placed from 1 to 3 pairs of minute, conical, white teeth, the
1949 MacMillan: Land Snails of West Virginia 137
first pair in sight when looking into the aperture, the others more remote.
Greater diameter mm., lesser 3 mm.; height \]/2 n^m.
Type Locality: Near Philadelphia, Pennsylvania (Say, 1817).
Range: Newfoundland; Quebec to Manitoba; United States east of
Rocky Mountains.
West Virginia Records
Barbour County: Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Braxton County; Frame-
town (gkm), Gassaway (gkm). Cabell County; Barboursville (gkm) (m&r),
Lesage (m&r), Ona (m&r). Calhoun County; Grantsville (ndr). Clay
County; Clay (gkm). Grant County; Greenland Gap (ndr). Greenbrier
County; Alderson (ndr) (gkm), McClungs Cave (A. M. Reese), Organ
Cave (ndr). Renick (grh). Renick Valley (ndr), Ronceverte (gkm)
(m&r). Hampshire County; Junction (ndr), Romney (gkm). Jefferson
Cowwii/; Aldridge (msb), Bakerton (msb), Bardane (msb), Bloomery (msb)
(jpem), Bolivar (msb), Charles Town (jpem), Duffield (msb), Engle
(msb), Halltown (msb). Harper’s Ferry (jpem) (ndr) (msb) (gwt) (cc),
Jamestown (msb), Kabletown (msb), Kearneysville (msb). Keys Gap
(jpem), Leetown (msb), Loudoun Heights (jpem), Mechanicstown (msb).
Middleway (msb), Morgan Grove (msb), Mt. Mission (jpem), Reedson
(msb), Rippon (msb), Shenandoah City (msb), Shenandoah Junction
(msb), Skeetersville (msb), Uvilla (msb). Kanawha County; Alum Creek
(ndr), Dunbar (gkm), Hudnall (cbw), Nitro (gkm). Tornado (gkm).
Lewis County; Jackson’s Mill (gkm). Lincoln County; Branchland (m&r),
Myra (m&r), Sheridan (ndr), Sweetland (m&r). Logan County; Blair
(m&r), Davin (ndr). Sharpies (mrs. gkm) (m&r). Stone Branch (ndr).
McDowell County; laeger (m&r). Panther (m&r). Marion County; Curtis-
ville (ndr), Fairmont (ndr), Glady Creek (ndr). High Point (ndr).
Marshall County; Bannon (ndr). Mason County; Leon (ndr). Point
Pleasant (ndr). Mercer County; Bluefield (m&r), Princeton (m&r),
Spanishburg (m&r). Speedway (m&r). Mineral County; Burlington (gkm).
New Creek (ndr). Mingo County; Taylorsville (m&r), Williamson (m&r).
Monongalia County; Coopers Rock (ndr), Smithtown (ndr), Uffington
(ndr). Monroe County; Alderson (gkm). Salt Sulphur Springs (ndr)
(m&r), Sweetsprings (grh) (m&r), Union (gkm), Waitesville (m&r).
Morgan County; Berkeley Springs (ndr), Largent (m&r). Nicholas County;
Craigsville (gkm), Lockwood (gkm), Richwood (ndr), Summersville
(gkm). Ohio County; Wheeling (ndr). Pendleton County; Circleville
(ndr), Franklin (ndr) (gkm), Judy Gap (gkm). Upper Tract (ndr) (gkm).
138
Annals of the Carnegie Museum
VOL. 31
Pocahontas County; Black Mt. (p&l), Dunmore Spring (gkm), Greenbank
(gkm), Hillsboro (gkm), Marlinton (gkm), Mill Point (gkm). Putnam
County; Poca (gkm). Raleigh County; Eccles (m&r). Flat Top Mt. (p&l).
Randolph County; Cheat Bridge (ndr) (gkm). Cheat Mt. (p&l), Huttons-
ville (gkm). Summers County; Talcott (gkm). Tucker County; Parsons
(mrs. gkm) (gkm). Tyler County; Friendly (cbw). Upshur County;
French Creek (gkm). Wayne County; Echo (m&r). Fort Gay (m&r), Genoa
(m&r). Webster County; Bolair (ndr) (gkm) (m&r). Point Mt. (grh).
Wetzel County; Silver Hill (ndr). Wyoming County; Oceana (m&r), Pine-
ville (m&r).
Remarks: The most salient characters of this species are the parallel,
spiral ridges encircling the shell and the large, excavated umbilicus.
Genus Discus Fitzinger
Shell widely umbilicated, depressed, discoid, turbinate, or globosely-
depressed, rugose or costulately striate ; whorls 33^-63^^, equal or gradually
increasing, rounded to carinate, or bicarinate. White to reddish-horn color,
with or without yellowish lines. Aperture lunately rounded, rhomboidal,
or stirrup-shaped, oblique, furnished within with or without a single basal
tooth. Peristome simple, acute, extremities joined by a thin callus.
Umbilicus wide, showing all inner volutions.
Discus patulus patulus (Deshayes)
(Plate 6, fig. 7)
Shell broadly and perspectively umbilicated, orbicular, scarcely convex
above, excavated below, thin, reddish horn-color, regularly ribbed ; whorls
gradually increasing; aperture small, lunately subcircular, within
furnished with a single, subprominent tooth on the base of the shell;
peristome simple, acute, its extremities separated widely. Greater di-
ameter 8, lesser 73^ mm.; height 3 mm.
Type Locality: Environs of New York. (Deshayes, 1830).
Range: Ontario; United States east of the Rocky Mountains.
West Virginia Records
Boone County; Peytona (m&r), Racine (m&r), Seth (m&r). Brooke
County; Bethany (cbw), Williamsburg (cbw). Cabell County; Ona (m&r).
Calhoun County; Grantsville (ndr). Doddridge County; West Union (grh).
Fayette County; Clifftop (gkm). Grant County; Greenland Gap (ndr).
Mount Storm (ndr). Greenbrier County; Alderson (gkm). Renick Valley
1949
MacMillan: Land Snails of West Virginia
139
(ndr). Jackson County; (W. M. Gabb). Kanawha County; Alum Creek
(ndr), Hudnall (cbw), South Charleston (ndr), Tornado (gkm). Lewis
County; Weston (gkm). Lincoln County; Branchland (m&r), Myra (m&r),
Sheridan (ndr), Sweetland (m&r). Logan County; Blair (m&r), Davin
(ndr), Sharpies (m&r). McDowell County; Avondale (m&r), laeger (m&r),
Panther (m&r). Marion County; Curtisville (ndr), East Fairmont (ndr),
Fairmont (ndr) (grh), Glady Creek (ndr). High Point (ndr), Kingmont
(grh). Mill Creek (grh), Smithville (cbw). Marshall County; Bannon
(ndr), Cameron (gb). Mason County; Point Pleasant (ndr) (cbw). Mer-
cer County; Bluefield (m&r), Oakvale (m&r), Princeton (m&r). Speedway
(m&r). Mineral County; New Creek (ndr). Mingo County; Williamson
(m&r). Monongalia County; Cheat Mt. (hhs). Coopers Rock (cbw).
Monroe County; Alderson (gkm), Gap Mills (m&r). Salt Sulphur Springs
(m&r), Sweetsprings (m&r). Willow Bend (m&r). Nicholas County; Lock-
wood (gkm). Snake Den Mt. (ndr), Summersville (gkm). Ohio County;
Wheeling (mgn) (ndr) (cbw). Pocahontas County; Hillsboro (gkm), Mar-
linton (gkm). Preston County; Cascade (ndr). Putnam County; Poca
(gkm). Raleigh County; Daniels (gkm), Eccles (m&r). Randolph County;
Huttonsville (gkm). Summers County; Riffle (gkm), Talcott (gkm).
Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County; Bolair
(ndr) (m&r). Wetzel County; Burton (grh). Wood County; Boaz (cbw).
Wyoming County; Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Remarks: In 1817 Thomas Say described a new American snail as
Helix perspectiva. This same specific name had been given to a Helix
described by Megerle von Muhlfeld in 1816. As Megerle’s species had been
described first, it had the priority over Say's species. Consequently Say's
name became a synonym of Helix patula of Deshayes.
This snail selects drier stations than those of related species, and is
commonly found under dead trees, under bark close to the ground, or
between the bark and wood; it is occasionally found under flat stones. It
is particularly abundant under the bark of decaying trees, sometimes
found there in vast numbers.
Discus patulus carinatus, MacMillan
(Plate 6, fig. 8)
Shell slightly elevated, reddish-horn color, dull; spire elevated; whorls
S]/2, angulated; embryonic whorl smooth; surface heavily ribbed above,
the rib-stria tions forming a carina as they pass over the periphery; rib-
striations smooth on top; intermediate striae faint; rib-stria tions below
140
Annals of the Carnegie Museum
VOL. 31
continue into umbilicus; umbilicus moderately wide, deep, showing all
inner whorls; aperture somewhat rounded; internal columellar tubercle
medium in size. Diameter 8 mm.; height 3.5 mm.
Type Locality: Paint Rock, Madison County, North Carolina (Kutchka,
1938).
Range: Michigan and Illinois to western Pennsylvania; western Mary-
land; West Virginia, Kentucky, Tennessee, North Carolina, South Caro-
lina, Georgia, and Alabama.
West Virginia Records
Barbour County; Philippi (gkm). Boone County; Peytona (m&r), Racine
(m&r), Seth (m&r). Braxton County; Frametown (gkm). Cabell County;
Barboursville (m&r), Ona (m&r). Calhoun County; Grantsville (ndr).
Fayette County; Cliff top (mrs. gkm) (gkm). Grant County; Greenland
Gap (ndr). Mount Storm (ndr). Greenbrier County; Renick Valley (ndr).
'Kanawha County; Alum Creek (ndr), Hudnall (cbw). South Charleston
(ndr). Lewis County; Weston (gkm). Lincoln County; Branchland (m&r),
Myra (m&r), Sheridan (ndr), Sweetland (m&r). Logan County; Blair
(m&r), Davin (ndr). Sharpies (m&r). McDowell County; Avondale (m&r),
laeger (m&r). Panther (m&r). Marion County; Curtisville (ndr), Fair-
mont (ndr), Glady Creek (ndr). High Point (ndr). Mason County; Point
Pleasant (ndr). Mercer County; Bluefield (m&r), Oakvale (m&r), Prince-
ton (m&r), Spanishburg (m&r). Speedway (m&r). Mineral County; New
Creek (ndr). Mingo County; Williamson (m&r). Monroe County; Gap
Mills (m&r). Salt Sulphur Springs (m&r), Sweetsprings (m&r), Waitesville
(m&r). Willow Bend (m&r). Nicholas County; Lockwood (gkm). Ohio
County: Wheeling (ndr). Pocahontas County; Marlinton (gkm). Raleigh
County; Eccles (m&r). Randolph County; Huttonsville (gkm). Summers
County; Riffle (gkm), Talcott (gkm). Taylor County; Valley Falls (ndr).
Tucker County; Parsons (mrs. gkm). Upshur County; French Creek (gkm).
Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County; Bolair
(ndr) (gkm) (m&r). Wyoming County; Baileysville (m&r), Pineville
(m&r).
Remarks: This subspecies was first described as Discus patulus angulatus
by G. M. Kutchka (the present author) in 1938. Ten years earlier F. C.
Baker described Gonyodiscus macclintocki angulatus. Since the Interna-
tional Zoological Congress has passed the ruling that “specific and sub-
specific names .... are coordinate,” angulatus of Discus patulus has
been superceded by angulatus of Discus macclintocki, and therefore be-
1949
MacMillan; Land Snails of West Virginia
141
came untenable. Subsequently the author {Nautilus, 1940, vol. 53, p.
143) re-named this form as Discus patulus carinaius.
Discus patulus carinatus can readily be distinguished from the typical
Discus patulus by the angulation of the periphery; the rib-striations, in
passing over the periphery, form a carina, which is more or less promi-
nent.
Discus cronkhitei (Newcomb)
(Plate 7, fig 2)
Shell remarkably thin, somewhat depressed; of a very delicate horn-
color, transparent ; whorls 4, very finely striated transversely ; spire scarcely
elevated; suture moderate; aperture nearly rounded ; labrum not reflected
nor thickened. Umbilicus not remarkably large, in diameter not equal to
the body whorl ; transverse diameter one-fifth of an inch. (Greater diameter
6 mm., lesser 5^^ mm.; height 5 mm.).
Type Locality: Near Strawberry Mansion, Fairmont Park, Philadelphia,
Pennsylvania (Pilsbry, 1906).
Range: Newfoundland; Canada; United States.
West Virginia Records
Jefferson County; Bakerton (msb), Charles Town (msb). Harper’s Ferry
(jpem), Shenandoah Junction (msb). Monongalia County; Morgantown
(hhs). Pendleton County; Franklin (ndr). Tucker County; Parsons (gkm).
Wayne County; Ceredo (cbw).
Remarks: Dr. H. A. Pilsbry changed the terminology of this species
from Helix striatella to Pyramidula cronkhitei, as striatella (Anthony, 1840)
had been used by Rang in 1831 for a species of Helix.
D. cronkhitei differs from patulus as being altogether a more delicate
shell in structure and markings, the number of whorls is two and one-half
less, the color is lighter, and the shell smaller; the sharp external edge is
also more conspicuous, and looking into the throat, there is no tooth-like
thickening which exists within the lower margins of patulus.
Genus Anguispira Morse
Shell rather large and solid, globose to depressed, with convex spire and
open umbilicus; whorls 5-63^, rounded or carinated at the periphery.
Surface wrinkled, striate, or rib-striate, obliquely flamed, or spirally
banded. Aperture oval to rhomboidal; peristome thin, simple, acute, its
142
Annals of the Carnegie Museum
. VOL. 31
extremities joined by a thin to heavy callus. Umbilicus open, deep, show-
ing all inner volutions.
Anguispira alternata alternata (Say)
(Plate 7, fig. 3)
Shell broadly umbilicated, orbicularly depressed; thin, smoky horn-
color varied with red, interrupted, obliquely arranged patches and spots,
roughened by crowded, elevated, rib-like striae, smoother below; whorls
53^, flattened, .... “rounder,” pervious; aperture very oblique. Innately
rounded, banded within; peristome simple, acute, its terminations joined
by a very thin, transparent callus, that of columella subreflected. Greater
diameter 21, lesser 19 mm.; height 10 mm.
Type Locality: Middle States (Say, 1816).
Range: Canada; Maritime Provinces, Quebec, and Ontario; United
States; east of the Rocky Mountains.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Braxton County;
Gassaway (gkm). Brooke County; Bethany (cbw), Williamsburg (cBw).
Cabell County; Lesage (m&r). Grant County; Greenland Gap (ndr). Mount
Storm (ndr). Greenbrier County; Alderson (gkm), Ronceverte (m&r).
Hampshire County; Romney (gkm). Jefferson County; Bakerton (msb),
Bloomery (msb), Bolivar (msb). Hall town (msb). Harper’s Ferry (hap)
(msb), Leetown (msb), Loudoun Heights (jpem). Middleway (msb),
Shenandoah City (msb). Kanawha County; Charleston (um), Hudnall
(cbw). Logan County; Davin (ndr). McDowell County; Panther (m&r).
Marion County; Curtisville (ndr), Fairmont (ndr). High Point (ndr),
Mt. Harmony (ndr), Rivesville (ndr), Smithville (cbw). Marshall
County; Powhattan Point (frw). Mercer County; Bluefield (mgn) (m&r),
Oakvale (m&r), Princeton (m&r). Mineral County; Burlington (gkm),
Ridgeley (aeo). Monongalia County; Cheat Mt. (hhs). Coopers Rock
(cbw), Sturgisson (ndr), Uffington (ndr). Monroe County; Alderson
(gkm). Gap Mills (m&r). Salt Sulphur Springs (m&r). Morgan County;
Cherry Run (ndr). Nicholas County; Summersville (gkm). Ohio County;
Wheeling (cbw). Pendleton County; Circleville (ndr), Franklin (ndr)
(gkm), Judy Gap (gkm), Onega (mgn). Upper Tract (mgn) (ndr) (gkm).
Pocahontas County; Mill Point (gkm). Travellers Repose (ghc). Preston
County; Cheat River (hhs), Cranesville (mgn). Raleigh County; Flat Top
Mt. (p&l). Randolph County; Helvetia (vs). Summers County; Talcott
1949
MacMillan: Land Snails of West Virginia
143
(gkm). Upshur County; French Creek (gkm). Wood County; Boaz (cbw).
Wyoming County; Pineville (m&r).
Remarks: The best collecting places for this species are on hillsides
covered with timber, sticks, and forest debris. Among the hardwoods and
aspens it is found under leaf mould, the underside of rotten logs, fallen
bark, rotten stumps, ferns, and poison ivy. In pine groves it lives under
rotten wood that is frequently covered with lichens. In bog woods it is
often collected at the base of Arbor vitce and spruce. It is not uncommon
in open country in moist situations, where shelter is found under logs and
stumps. After rains it has been observed to crawl over reindeer moss in
considerable numbers. In late autumn it often congregates in great num-
bers under flat rocks, hibernating in such situations, and sometimes bur-
rowing in the loose soil for a distance of one or more inches.
Anguispira alternata angulata Pilsbry
(Plate 7, fig. 4)
Shell broadly umbilicated, orbicularly depressed, thin, smoky horn-
colored with red, interrupted, obliquely arranged patches and spots,
roughened by crowded, elevated, rib-like striae; smooth below. Whorls
53^, flattened, the last strongly carinated at its periphery. Aperture very
oblique, lunately rounded except where the periphery is carinated, whitish
within. Peristome simple, acute, its terminations joined by a very thin,
transparent callus. Umbilicus large, pervious. Greater diameter 21,
lesser 19 mm.; height 10 mm.
Type Locality: Emory River, Harriman, Tennessee (Pilsbry and
Rhoads, 1896).
Range: New York to Indiana, south to Alabama, and north to Mary-
land.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (mrs. gkm) (gkm). Boon^
County; Peytona (ndr) (m&r), Racine (m&r). Braxton County; Gassaway
(gkm). Brooke County; Bethany (mgn). Cabell County; Barboursville (ndr)
(m&r), Lesage (m&r). Calhoun County; Grantsville (ndr). Clay County;
Clay (gkm). Doddridge County; Smithburg (ndr). Grant County; Green-
land Gap (ndr). Mount Storm (ndr). Greenbrier County; Alderson (ndr)
(gkm). Organ Cave (ndr). Renick Valley (ndr), Ronceverte (gkm) (m&r).
White Sulphur Springs (p&l). Jefferson County; Bolivar (msb), Halltown
(msb). Middleway (msb), Millville (jpem), Shenandoah City (msb). Silver
144 Annals of the Carnegie Museum vol. 31
Grove (msb). Kanawha County; Dunbar (gkm), South Charleston (ndr).
Lewis County; Jackson’s Mill (gkm). Lincoln County; Myra (m&r). Logan
County; Davin (ndr), Sharpies (mrs. gkm) (m&r), Slagle (cg). McDowell
County; Avondale (m&r). Panther (m&r). Marion County; Curtisville
(ndr), Fairmont (ndr), Glady Creek (ndr). High Point (ndr), Mt.
Harmony (ndr), Rivesville (ndr). Mercer County; Oakvale (m&r),
Princeton (m&r), Spanishburg (m&r). Mineral County; Burlington (gkm),
Keyser (ihm). Mingo County; Williamson (m&r). Monongalia County;
Morgantown (hhs), Sturgisson (ndr), Uffington (ndr). Monroe County;
Alderson (gkm). Gap Mills (m&r). Salt Sulphur Springs (ndr) (m&r),
Sweetsprings (m&r), Waitesville (m&r). Nicholas County; Richwood (ndr),
Summersville (gkm). Ohio County ;'SN\\eC\mg (mgn) (ndr) (cbw). Pendleton
County; Circleville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm). Upper
Tract (mgn) (ndr) (gkm). Pocahontas County; Mill Point (ndr) (gkm).
Preston County; Cascade (ndr). Randolph County; Helvetia (vs). Sum-
mers County; Talcott (gkm). Upshur County; French Creek (mrs. gkm)
(gkm). Wayne County; Fort Gay (m&r). Webster County; Bolair (ndr).
Wetzel County; Silver Hill (ndr). Wyoming County; Pineville (m&r).
Remarks: In this form the periphery has a distinct, bluntly angulated,
but not acute, carina; the earlier whorls are quite strongly ribbed, but the
ribs diminish in strength on the penultimate whorl and the last whorl
becomes finely and closely striate. The striae pass over the carina and
continue into the umbilicus, but become much weaker below the carina,
and in fully mature specimens they become sub-obsolete towards the aper-
ture. The younger shells are quite strongly carinated and somewhat
excavated below the carina. Between the ribs there is developed a fine
vertical striation.
Anguispira alternata mordax (Shuttleworth)
(Plate 7, figs. 5, 6)
Shell widely and perspectively umbilicated, depressed, sublenticular,
carinate, thin, yellow-horn color, ornamented with interrupted red streaks
in bands, beautifully sculptured with strong, flexuous ribs above and be-
low; whorls flat; aperture very oblique, angularly crescentric, oval;
peristome simple, acute. Great diameter 18, lesser 16, height 6 mm.
Type Locality: Mountains of North Carolina (Shuttleworth, 1852).
Range: West Virginia; Virginia; Kentucky; Tennessee; North Carolina;
South Carolina; Georgia; and Alabama.
1949
MacMillan: Land Snails of West Virginia
145
West Virginia Records
McDowell County; Avondale (m&r), laeger (m&r), Panther (m&r).
Mercer County; Speedway (m&r).
Remarks: This subspecies is very similar to alternata, but is distinguished
from it by the stronger ribs, which are 1 mm. apart at the periphery and
which extend on the under side into the umbilicus. The carina is well de-
veloped, though rather more rounded than in angulata, but the under side,
just below the carina, is distinctly concave, as in Discus bryanti. This
disappears in full-grown specimens towards the aperture, but is always
evident on the first two-thirds of the last >vhorl, and is particularly marked
in young shells. The micro-sculpture is well developed, consisting of very
fine irregular lines of growth between the ribs, which are cut transversely
by numerous fine revolving lines. It differs from A. cumberlandiana in
having the shell strongly striate and costate.
Anguispira alternata mordax has been reported occuring in Greenbrier
and Jefferson Counties, but I am dubious regarding these localities and the
correct identification of the shells found in those two counties. It is my
contention that they are specimens of A. alternata angulata and I have
added them to the localities of that species. The true mordax has been
collected only in McDowell and Mercer Counties, the two southernmost
counties of West Virginia.
Anguispira kochi (Pfeiffer)
(Plate 7, fig. 7)
Shell broadly umbilicated, globosely depressed, coarse, solid, diaphan-
ous, obliquely and crowded wrinkled, from white to dark-reddish horn-
color, with from two to three brownish revolving bands; whorls 6, convex;
suture deep; aperture roundly lunate, pearly white and banded within;
peristome simple, acute, the ends joined by a thin transparent callus, that
of the columella dilated, subreflected. Greater diameter 25, lesser 22
mm.; height 15 mm.
Type Locality: Unknown (Pfeiffer, 1845).
Range: Western Pennsylvania and western West Virginia, central
Mississippi River Valley west to Pacific Ocean; British Columbia; Islands
in western Lake Erie, Ontario.
West Virginia Records
Brooke County; Bethany (cbw). Marshall County; Cameron (gb), Kent
(ndr). Ohio County ; (mgn) (ndr) (cbw).
146
Annals of the Carnegie Museum
VOL. 31
Remarks: The specific name of kochi is now used in place of solitaria
Say (1821), as Helix solitaria was pre-empted by Poiret in 1805. The type
locality of kochi is unknown, the shell coming from the collection of H.
Cuming, Esq. ; that of Say’s solitaria is Lower Missouri.
The animal buries itself from five to six inches or more in the soft soil
of moist hillsides to begin its period of hibernation. It issues forth when
the first warm rays of the vernal sun penetrates its hiding place. During
the summer it is usually found in great numbers under leaves and partially
buried, an inch or two, in the soft soil.
Family Zonitid^
Shell small to large, conic to depressed, thin, shining. Whorls 2-8,
regularly to rapidly increasing, rounded to keeled; smooth, polished,
covered with strongly marked, curved wrinkles or equidistant, elevated,
oblique ribs, or indented with delicate striae of growth or deep, equidistant,
radiating parallel lines; also sculptured with microscopic revolving lines;
furnished with or without rows of upright denticles on floor of whorls.
Color, whitish to greenish-horn or chestnut. Sutures slightly to deeply
impressed. Aperture large to narrow, oblique, nearly circular to transversely
rounded, with or without rows of very minute, white teeth or lamelliform,
elongated, nearly parallel teeth. Peristome simple, thin, acute, slightly or
not thickened within, terminations joined by a slight to heavy callus de-
posit. Umbilicus large to narrow, slightly covered by reflected peristome
or entirely covered, impressed.
Subfamily Kaliellin^
Genus Guppya Moerch
Shell perforate, depressed-conic, rather solid or thin, glossy, pale rusty-
brown or yellowish, with or without numerous faint lines of growth;
spire slightly elevated or depressed, having 4 or 5 closely revolving, well-
rounded whorls, separated by a very deep suture; periphery rounded;
base flattened or convexly rounded and excavated around a small, deep,
open or closed umbilicus; aperture wide, nearly circular or semi-lunate;
peristome simple, slightly expanded, and at the columellar region de-
cidedly reflexed.
1949 MacMillan: Land Snails of West Virginia 147
Guppya sterkii (Dali)
(Plate 7, figs. 9, 10, 11)
Shell minute, thin, yellowish translucent, brilliant, lines of growth
hardly noticeable, spire depressed, four-whorled ; whorls rounded, base
flattened, somewhat excavated about the center, which is imperforate;
aperture wide, hardly oblique, not very high, semi-lunate, sharp edged,
the upper part of the columella slightly reflected; upper surface of the
whorls roundish, though the spire as a whole is depressed. Maximum
diameter. 044 inch.; altitude 0.026 inch. (1 x .6 mm.).
Type Locality: New Philadelphia, Ohio (Dali, 1888).
Range: New Jersey to Ohio, south to Louisiana and Florida.
West Virginia Records
Barbour County; Nestorville (gkm). Boone County; Peytona (m&r),
Racine (m&r). Braxton County; Gassaway (gkm). Cabell County; Ona
(m&r). Greenbrier County; Alderson (ndr) (gkm). Renick (grh). Renick
Valley (ndr). White Sulphur Springs (p&l). Hampshire County; Romney
(gkm). Lincoln County; Branchland (m&r), Myra (m&r), Sweetland
(m&r). Logan County; Blair (m&r). Sharpies (m&r). McDowell County;
Avondale (m&r), laeger (m&r). Panther (m&r). Marion County; High
Point (ndr). Mercer County; Bluefield (m&r), Oakvale (m&r), Princeton
(m&r). Mineral County; Burlington (gkm). Mingo County; Taylorsville
(m&r), Williamson (m&r). Monongalia County; Sturgisson (ndr). Monroe
County; Alderson (gkm). Salt Sulphur Springs (ndr) (m&r), Sweetsprings
(grh), Waitesville (m&r). Nicholas County; Craigsville (gkm), Lockwood
(gkm), Summersville (gkm). Pendleton County; Franklin (gkm), Judy Gap
(gkm). Pocahontas County; Black Mt. (p&l), Greenbank (gkm), Marlinton
(gkm). Mill Point (gkm). Preston County; Cheat Mt. (grh). Putnam
County; Poca (gkm). Raleigh County; Flat Top Mt. (p&l). Summers
County; Riffle (gkm). Webster County; Bolair (ndr), Wyoming County;
Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Remarks: This snail, due to its small size, is easily overlooked by most
collectors. It is most readily found in the sifted samples of forest loam
collected under fallen logs or slabs of sandstone and limestone.
Genus Euconulus Reinhardt
Shell imperforate or very narrowly perforate, turbinate, arctispiral;
whorls 43^ to 7, rounded or bluntly angular, minutely striated or in-
148
Annals of the Carnegie Museum
VOL. 31
distinctly wrinkled ; suture moderate to distinct ; yellowish-white to brown-
ish-amber in color; aperture narrow, nearly transverse or depressed-
lunar; periphery simple, acute; base with or without 1 to 3 low, radial
teeth.
Euconulus fulvus (Mueller)
(Plate 8, figs. 1, 2)
Shell imperforate, subconical, thin, pellucid ; epidermis smooth, shining,
minutely striated, amber colored ; whorls 5 or 6, bluntly angular, very nar-
row, suture distinct and deep; aperture transverse, narrow; peristome
simple, acute; base convex; umbilical region indented, umbilicus closed.
Greater diameter 4, lesser 33^2 i^im. ; height 3 mm.
Type Locality: Fridrichsdal, Denmark (Mueller, 1774).
Range: Greenland; Newfoundland; Canada; Alaska; United States.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm), Boone County;
Peytona (gb), Racine (m&r). Braxton County; Gassaway (gkm). Cabell
County; Barboursville (m&r), Ona (m&r). Fayette County; Clifftop (gkm).
Grant County; Bismark (ndr). Mount Storm (ndr). Greenbrier County;
Renick (grh). Renick Valley (ndr), Ronceverte (gkm). Hampshire
County; Romney (gkm). Jefferson County; Harper’s Ferry (gwt). Kanawha
County; Hudnall (cbw). Lewis County; Jackson’s Mill (gkm), Weston
(gkm). Lincoln County; Branchland (m&r), Myra (m&r), Sweetland (m&r).
Logan County; Blair (m&r). Sharpies (m&r). McDowell County; Avondale
(m&r). Panther (m&r). Marion County; Glady Creek (ndr). High Point
(ndr). Monroe County; Alderson (gkm), Sweetsprings (m&r), Waitesville
(m&r). Nicholas County; Summersville (gkm). Pendleton County; Circle-
ville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm). Pocahontas County;
Dunmore Spring (gkm), Marlinton (gkm). Summers County; Riffle (gkm).
Tucker County; Parsons (gkm). Upshur County; French Creek (gkm).
Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County; Bolair
(ndr). Wetzel County; Silver Hill (ndr). Wyoming County; Pineville
(m&r).
Remarks: The shell is much less elevated than in chersinus, with the 5
whorls not so closely coiled, and the last one distinctly but bluntly angular
at the periphery. It is also distinguished from chersinus and its varieties
by the fewer, wider whorls, and generally less elevated countour.
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MacMillan: Land Snails of West Virginia
149
Euconulus chersinus chersinus (Say)
(Plate 8, figs. 4, 5)
Shell subglose-conic, pale yellowish-white, pellucid; convex beneath;
volutions about six, wrinkles not distinct; spire convex-elevated; suture
moderate; body whorl rounded; mouth nearly transverse, unarmed, the
two extremities nearly equal; labrum simple; umbilicus none. Breadth
1/10 of an inch (23^ mm.).
Type Locality: Sea Islands of Georgia (Say, 1821).
Range: Maine to Wisconsin, south to Mississippi and Florida.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r), Racine (m&r). Braxton County; Frame town (gkm).
Cabell County; Ona (m&r). Clay County; Clay (gkm). Greenbrier County;
Renick Valley (ndr), Ronceverte (m&r). Hampshire County; Romney
(gkm). Jefferson County; Bolivar (msb). Harper’s Ferry (gwt), Reedson
(msb). Kanawha County; Alum Creek (ndr). Lincoln County; Sweetland
(m&r). McDowell County; Avondale (m&r), laeger (m&r). Panther (m&r)
Mason County; Leon (ndr). Point Pleasant (ndr). Mineral County; New
Creek (ndr). Mingo County; Taylorsville (m&r), Williamson (m&r).
Monongalia County; Morgantown (hhs). Monroe County; Alderson (gkm).
Salt Sulphur Springs (m&r), Sweetsprings (grh) (m&r), Waitesville (m&r),
Willow Bend (m&r). Nicholas County; Lockwood (gkm), Summersville
(gkm). Pendleton County; Franklin (ndr) (gkm), Judy Gap (gkm).
Pocahontas County; Hillsboro (gkm). Putnam County; Poca (gkm). Up-
shur County; French Creek (mrs. gkm). Wayne County; Echo (m&r),
Fort Gay (m&r). Webster County; Bolair (ndr) (gkm). Wyoming County;
Baileysville (m&r), Pineville (m&r).
Remarks: Euconulus chersinus is a smoother shell then Euconulus
fulvus, and the whorls are round in comparison to the bluntly angulated
whorls of fulvus.
Euconulus chersinus dentatus (Sterki)
(Plate 8, figs. 8, 9)
Rather small, with narrow whorls of the species, the last whorl con-
taining 1 to 3 low, radial teeth, forming transverse barriers on the basal
wall, and appearing when the shell is viewed from the base as white radial
stripes. The radiating “teeth” are exactly the same type found in Ven-
tridens lamellidens.
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Type Locality: Jackson County, Alabama (Sterki, 1893).
Range: Indiana; Ohio; Maryland; Virginia; West Virginia; North
Carolina; South Carolina; Alabama; Mississippi; Louisiana; and Arkansas.
West Virginia Records
Boone County; Seth (m&r). Cabell County; Ona (m&r). Hampshire
County; Romney (gkm). Lincoln County; Branchland (m&r), Myra (m&r),
Sweetland (m&r). Mingo Williamson (m&r), Monongalia County;
Morgantown (hhs). Nicholas County; Summersville (gkm). Pendleton
County; Franklin (ndr) (gkm). Wayne County; Fort Gay (m&r).
Remarks: The “teeth” found on the base of the shell of this variety are
placed at somewhat irregular intervals, are rounded or elongate in a
radial direction. They are not high, but tooth-like and quite distinct and
in the form of radial bars. When two or three are present, they are always
of the same character, either round or transverse.
Euconulus chersinus polygyratus (Pilsbry)
(Plate 7, fig. 8)
Similar to Euconulus chersinus trochulus, but less elevated, with nar-
row aperture; whorls over 6, very narrow, the last one rounded; upper
surface with a luster of silk; base glossy, with a silky band around the outer
margin. Color generally deep, brownish amber. Altitude 2.2, diameter
3 mm.
Type Locality: Hamilton, Ontario, Canada (Pilsbry, 1899).
Range: Ontario to British Columbia; Maine to Minnesota, south to
Arkansas and Alabama.
West Virginia Records
Boone County; Peytona (m&r), Racine (m&r), Seth (m&r). Braxton
County; Gassaway (gkm). Calhoun County; Grantsville (ndr). Clay
County; Clay (gkm). Grant County; Greenland Gap (ndr). Jefferson
County; Bolivar (msb). Lincoln County; Branchland (m&r). Logan County;
Blair (m&r). McDowell County; laeger (m&r). Marion County; High Point
(ndr). Mason County; Leon (ndr). Mercer County; Oakvale (m&r),
Princeton (m&r). Mineral County; New Creek (ndr). Monroe County;
Alderson (gkm), Sweetsprings (m&r). Morgan County; Berkeley Springs
(ndr). Nicholas County; Summersville (gkm). Pendleton County; Franklin
(gkm). Pocahontas County; Marlinton (gkm). Raleigh County; Eccles
(m&r).
1949
MacMillan: Land Snails of West Virginia
151
Remarks: Euconulus chersinus polygyratus is a smaller shell, with nar-
rower and more numerous whorls than Euconulus chersinus.
Subfamily Zonitin^
Genus Omphalina Rafinesque
Shell globose to depressed -globose, thin, shining, wrinkled to striated,
reddish to reddish-chestnut in color, and umbilicated. Whorls 4-53/^,
rounded or flattened, regularly and rapidly increasing, the last very large
and ventricose. Sutures slightly to moderately impressed. Sculpture
consisting of fine and close striae or wrinkles to coarse and regularly
spaced rib-striations above; spiral sculpture consisting of microscopic
granules, prominent above and obsolete below. Spire short, smooth, conic
or depressed. Aperture large, circular and oblique; parietal wall covered
with or without a thin callus deposit. Peristome simple, thin, acute, and
slightly reflected over the umbilicus. Umbilicus deep and narrow, and
sometimes slightly expanded.
Omphalina cuprea Rafinesque
(Plate 8, fig. 6)
Shell thin, depressed on upper surface, epidermis dark, approaching to
chestnut-color, sometimes almost black, shining and wrinkled ; whorls 43/^,
rapidly increasing, with irregular, oblique wrinkles, the last whorl very
voluminous and expanding transversely towards the aperture; suture very
little impressed; aperture very oblique, ample, lunate-oval, within pearly
or iridescent; peristome simple, thin, brittle, with a light, testaceous de-
posit within, the two terminations approaching each other very nearly,
that of the columella somewhat reflected; umbilicus deep, not much ex-
panded. Greater diameter 26, lesser 22 mm.; height 13 mm.
Type Locality: Kentucky (Rafinesque, 1831).
Range: Ontario; Maine to Minnesota, south to Louisiana and Florida.
West Virginia Records
Barbour County; Nestorville (gkm). Boone County; Peytona (ndr)
(m&r), Racine (m&r), Seth (m&r). Braxton County; Frametown (gkm),
Gassaway (gkm), Shadyside (gkm). Cabell County; Barboursville (ndr)
(m&r), Lesage (m&r). Calhoun County; Grantsville (ndr). Clay County;
Clay (gkm). Fayette County; Cliff top (gkm). Grant County; Greenland Gap
(ndr). Mount Storm (ndr). Greenbrier County; Alderson (gkm), Charmco
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VOL. 31
(ndr). Hampshire County; Romney (gkm). Kanawha County; Alum Creek
(ndr), Charleston (gkm), Hudnall (cbw), Institute (gkm). Lewis County;
Weston (gkm). Lincoln County; Branchland (m&r), Myra (m&r), Sheridan
(ndr), Sweetland (m&r). Logan County; Blair (m&r), Davin (ndr),
Sharpies (m&r), Slagle (cg). Stone Branch (ndr). McDowell County;
Avondale (m&r), laeger (m&r). Panther (m&r). Marion County; Curtis-
ville (ndr), Fairmont (ndr), Glady Creek (ndr). High Point (ndr),
Jayenne (grh), Midway Park (grh). Marshall County; Cameron (gb),
Kent (ndr), Powhattan Point (frw). Mercer County; Bluefield (m&r),
Princeton (m&r), Spanishburg (mcz) (c&a) (m&r). Speedway (mgn) (m&r).
Mineral County; Burlington (gkm), Keyser (ihm). New Creek (ndr).
Mingo County ;Ta,y\oYsw\\\% (m&r), Williamson (m&r). Monongalia County;
Coopers Rock (ndr) (cbw), Smithtown (ndr), Sturgisson (ndr). Monroe
County; Gap Mills (m&r), Waitesville (m&r). Morgan County; Largent
(ndr). Nicholas County; Belva (ndr), Lockwood (gkm), Richwood (ndr)
(cjg), Snake Den Mt. (ndr), Summersville (gkm). Ohio County; Wheeling
(mgn) (ndr) (cbw). Pendleton County; Circleville (ndr), Franklin (ndr)
(gkm), Judy Gap (gkm), Onega (mgn). Upper Tract (mgn) (ndr). Poca-
hontas County; Marlinton (gkm). Mill Point (gkm). Preston County;
Cascade (ndr), Laurel Mt. (grh), Manheim (jha). Terra Alta (mgn).
Putnam County; Poca (gkm). Raleigh County; Daniels (gkm), Eccles
(m&r). Flat Top Mt. (p&l). Randolph County; Helvetia (vs), Huttonsville
(gkm). Roane County; Walton (ndr). Tucker County; Parsons (gkm).
Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County; Bolair
(ndr). Wetzel County; Burton (grh). Wirt County; .Wyoming County ;
Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Remarks: This species was known as Helix fuliginosus for a long time,
but it is now called Omphalina cuprea. The name fuliginosus had been
applied to a species of Helix by Griffith in a letter to A. Binney, which
was published in the Journal of the Boston Society of Natural History,
1841, vol. 3, p. 417, pi. 24. Rafinesque applied the name cuprea to a
species of Omphalina in 1831. As this species was similar to Helix fuligi-
nosus, it had priority over Griffith's species.
Genus Oxychilus Fitzinger
Shell very depressed and almost discoid, more convex beneath, thin and
brittle, very glossy, smooth, and semitransparent, of an amber or yellow-
ish-horn color above, and whitish, with often a greenish or bluish tinge
around the umbilicus, which is comparatively narrow, but deep and
1949
MacMillan: Land Snails of West Virginia
153
slightly overspread by the apertural margin ; lines of growth shallow and
indistinct, slightly puckered at the sutures and microscopically striate in
the spiral direction; suture with a channeled aspect and showing often
as a darker line; epidermis comparatively thick, whorls 5-6, regularly but
slowly increasing; spire slightly raised; aperture broadly lunate, and
slightly oblique, with simple and direct margins, the basal margin slightly
reflected over and encroaching on the umbilicus, the upper margin pro-
jecting beyond the lower.
Oxychilus drapernaldi (Beck)
(Plate 8, fig. 11)
Umbilicated, thin, irregularly impressed striate, pellucid, shining,
smooth, yellowish corneous or greenish, base becoming whitish; whorls
5}/2, sutures impressed, the last whorl much wider, subdepressed. Dia-
ameter 14 mm.
Type Locality: France (Draparnaud, 1801).
Range: California; Pennsylvania; West Virginia; and Virginia.
West Virginia Records
Jefferson County; Bolivar (msb), Charles Town (msb), Jamestown
(msb), Ranson (msb).
Oxychilus cellarius (Mueller)
(Plate 8, fig. 12)
Shell very much depressed, thin, fragile, pellucid; epidermis light-
greenish horn-color; smooth, lightly polished; whorls 5, slightly rounded,
with minute and almost imperceptible oblique striae; aperture not dilated,
the transverse diameter the greatest; umbilicus moderate, regularly
rounded, deep; base rounded, thickened within by a testaceous deposit;
bluish-white; peristome simple, acute. Greater diameter 13, lesser 11)^
mm., height 5 mm.
Type Locality: Havnia (Norway?) (Mueller, 1774).
Range: Ontario and Quebec; Maine; New York; Pennsylvania; West
Virginia; South Carolina; and California.
West Virginia Records
Jefferson County; (msb). Monroe County; Alderson (gkm). Summers
County; Riffle (gkm).
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Genus Mesomphix Beck
Shell depressed-globose to flattened. Epidermis yellowish-horn, green-
ish-horn, or chestnut colored, shining, thin. Whorls 43^ to 6, rather
flattened, rounded, regularly increasing, last whorl moderately to widely
expanding. Suture moderately impressed. Sculpture consisting of minute,
delicate wrinkles, or regular, oblique striae ; sometimes minute microscopi-
cal granulations are present on the upper surface, but absent below; spiral
sculpture of minute and microscopical lines. Aperture oblique to slightly
oblique, transverse to lunar in shape, with or without a white testaceous
deposit within. Peristome simple, thin, acute, its extremities slightly
approaching, lower extremities inserted into center of the base and slightly
reflected. Umbilicus small, round, deep.
Mesomphix inornatus (Say)
(Plate 8, fig. 3)
Shell depressed ; epidermis yellowish horn-color, smooth, shining, with
very minute lines, not breaking the smoothness of the surface; whorls 5;
suture not much impressed ; aperture transverse, scarcely oblique, oblique
lunar, with a thick, white testaceous deposit around its whole inner sur-
face, a little distant from the margin; peristome thin, acute, fragile, its
ends somewhat converging, the columellar margin reaching to the center
of the base, subdilated above; umbilicus small; base rather flattened,
indented in the center. Greater diameter 16, lesser, mm.; height 6
mm.
Type Locality: Pennsylvania (Say, 1822).
Range: Quebec, Ontario; United States, east of the Mississippi River.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (mrs. gkm) (gkm). Boone
County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Braxton County;
Frametown (gkm), Gassaway (gkm), Shadyside (gkm). Cabell County;
Barboursville (ndr) (m&r), Lesage (m&r), Milton (aeo) (cbw), Ona (m&r).
Calhoun County; Arnoldsburg (grh), Grantsville (ndr). Clay County;
Clay (gew) (gkm). Doddridge County; Sherwood (ndr), Smithburg (ndr).
West Union (grh) (frw). Fayette County; Clifftop (mrs. gkm) (gkm).
Grant County; Greenland Gap (ndr). Mount Storm (ndr). Greenbrier
County; Alderson (gkm). Organ Cave (ndr). Renick Valley (ndr). White
Sulphur Springs (p&l). Hampshire County; Romney (gkm). Harrison
County; Bristol (ndr), Salem (grh). West Milford (grh). Jackson County;
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MacMillan: Land Snails of West Virginia
155
(W. M. Gabb). Kanawha County; Alum Creek (ndr), Charleston (gkm),
Hudnall (cbw), St. Albans (um), South Charleston (ndr). Lewis County;
Jackson’s Mill (mrs. gkm) (gkm), Weston (gkm). Lincoln County; Branch -
land (m&r), Myra (m&r), Sheridan (ndr), Sweetland (m&r). Logan County;
Blair (ndr) (m&r), Davin (m&r). Sharpie (mrs. gkm) (m&r), Slagle (cg).
McDowell County ; h-Yonddle (m&r), laeger (m&r). Panther (m&r . Marion
County; Chesapeake (ndr), Curtisville (ndr), Fairmont (ndr) (grh),
Glady Creek (ndr), Hammond (ndr). High Point (ndr), Kingmont
(grh), Mt. Harmony (ndr), Norway (grh), Smithville (cbw). Marshall
County; Cameron (gb), Powhattan Point (frw). Mason County; Point
Pleasant (ndr) (cbw). Mercer County; Bluefield (mgn) (m&r), Oakvale
(m&r), Princeton (m&r), Spanishburg (m&r). Speedway (m&r). Mingo
Taylorsville (m&r), Williamson (m&r). Monongalia County; Cheat
Mt. (hhs), Coopers Rock (ndr) (mgn) (cbw), Morgantown (Dr. H. Kahl),
Smithtown (ndr), Sturgisson (ndr). Monroe County; Alderson (gkm). Gap
Mills (m&r). Salt Sulphur Springs (ndr) (m&r), Sweetsprings (m&r),
Waitesville (m&r). Willow Bend (m&r). Nicholas County; Craigsville
(gkm), Lockwood (gkm), Nettie (ndr). Rich wood (cjg) (ndr), Snake
Den Mt. (ndr), Summersville (gkm). Ohio Wheeling (mgn) (ndr)
(cbw). Pendleton County; Franklin (ndr) (gkm), Judy Gap (gkm). Poca-
hontas County; Cass (mgn), Hillsboro (mgn) (gkm), Marlinton (gkm),
Mill Point (mgn) (ndr) (gkm). Preston County; Caddell (grh). Cascade
(ndr). Cheat Mt. (grh), Manheim (jha). Terra Alta (mgn). Raleigh
County; Daniels (gkm) (m&r). Flat Top Mt. (p&l). Shady Spring (m&r).
Randolph County; Cheat Bridge (gkm), Helvetia (vs), Huttonsville (mgn)
(ndr) (gkm). Roane County ;'^2Xton (ndr). Summers County; Riffle (gkm),
Talcott (gkm). Taylor County; Grafton (frw). Tucker County; Canaan
Valley (mgn). Parsons (mrs. gkm) (gkm). Upshur County; French Creek
(mrs. gkm) (gkm). Wayne County; Fort Gay (m&r), Genoa (m&r). Webster
County; Bolair (ndr) (gkm) (m&r). Wetzel County; Burton (grh). Silver
Hill (ndr). Wirt County; (wjf). Wood County; Boaz (cbw). Wyoming
County; Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Mesomphix perlaevis vulgatus H. B. Baker
(Plate 8, fig. 10)
Shell somewhat convex, often depressed, epidermis greenish horn-color,
shining, thin; whorls 5, rather flattened, rapidly enlarging, with beautiful
and regular oblique striae and revolving microscopic lines, the last whorl
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VOL. 31
expanding towards the aperture, not descending; aperture transverse,
broadly lunar, ample, with a testaceous deposit within; peristome thin,
acute, straight, extremities approaching, its lower extremity inserted into
center of base and somewhat reflected; base smooth perforate. Greater
diameter 18, lesser 15, height 9 mm.
Type Locality: Kentucky (Rafinesque, 1821).
Range: New York to Ohio, Kentucky to Arkansas, south to Florida,
and north to Maryland.
West Virginia Records
Barbour County; Nestorville (gkm). Doddridge County; West Union
(grh). Grant County; Greenland Gap (ndr). Harrison County; Bristol
(ndr). Kanawha County; Hudnall (cbw). McDowell County; Panther
(m&r). Marion County; Curtisville (ndr), Fairmont (ndr), Glady Creek
(ndr), High Point (ndr), Jayenne (grh), Mt. Harmony (ndr). Monon-
galia County; Cheat Mt. (hhs). Coopers Rock (ndr) (cbw), Smithtown
(ndr), Sturgisson (ndr). Ohio County ; Wheeling (mgn) (ndr) (cbw) (grh).
Preston County; Cascade (ndr). Summers County; Talcott (gkm). Tucker
County; Parsons (gkm). Wetzel County; Burton (grh).
Remarks: M. p. vulgatus is a more globose shell than M. inornatus; it
also differs from inornatus by its coarser sculpture.
This subspecies was described in 1821 by Rafinesque as Helix Icevigata.
However, Linne used the same specific name for a Helix in 1766, thereby
making Rafinesque’s name a nude name. For this reason it is now known
as Mesomphix perlcevis vulgatus, a name given to it by H. B. Baker in 1933.
Mesomphix rugeli oxycoccus (Vanatta)
(Plate 8, fig 7)
Shell depressed-globose, perforate, thin, delicately wrinkled, greenish
horn-color. Spire elevated, somewhat rounded. Whorls 6, rounded, the
last globose. Sculpture consists of first 3}/2 whorls slightly striate; the
striae gradually becoming weaker to form wrinkles, comparatively smooth
below; the spiral sculpture consists of densely microscopical granules
above but much weaker below. Aperture large, rounded, somewhat
oblique; peristome simple, thin, acute, its ends approaching slightly.
Umbilicus narrow, and slightly covered by reflected peristome. Greater
diameter 17, lesser 16^ mm.; height 13 mm.
Type Locality: Cranberry, Avery County, North Carolina (Vanatta,
1903).
Range: North Carolina and West Virginia.
1949
MacMillan: Land Snails of West Virginia
157
West Virginia Records
Logan County; Davin (ndr). Mercer County; Bluefield (m&r). Mingo
County; Taylorsville (m&r). Wyoming County; Pineville (m&r).
Remarks: The shell is microscopically granulate above and nearly
smooth on the base, which distinguishes it from M. rugeli.
Genus Paravitrea Pilsbry
Shell polished, corneous to hyaline; usually with major and minor
series of growth-striae, which are less prominent on umbilical side; whorls
overlapping (sutural spiral usually much contracted) ; peristome simple
and sharp; internal armature commonly reduced in adults.
Paravitrea multidentata (A. Binney)
(Plate 8, figs. 13, 14, 15)
Shell depressed, sub-planulate above, very thin, pellucid; epidermis
smooth, shining; whorls 6, narrow, slightly convex, increasing but slowly
in diameter, lines of growth hardly visible; suture impressed; aperture
semi-lunate, narrow; lip acute; umbilicus very small, round, not exhibiting
any of the volutions; base convex, indented around umbilicus. Two or
more rows of very minute, white teeth, radiating from the umbilicus, are
seen through the shell, within the base of the last whorl. Greatest trans-
verse diameter one-eighth of an inch (1334 mm.).
Type Locality: Not designated (A Binney, 1840).
Range: Quebec and Ontario; Maine to Wisconsin; south to Arkansas
and Georgia.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r), Racine (m&r), Seth (m&r). Braxton County; Gassaway
(gkm). Cabell County; Lesage (m&r). Clay County; Clay (gkm). Doddridge
County; West Union (frw). Fayette County; Clifftop (gkm). Grant County;
Greenland Gap (ndr). Mount Storm (ndr). Greenbrier County; Alderson
(grh) (gkm). Organ Cave (ndr). Renick (grh). Renick Valley (ndr),
Ronceverte (m&r). White Sulphur Springs (p&l). Jefferson County;
Harper’s Ferry (jpem). Lewis County; Jackson’s Mill (gkm), Weston
(gkm). Lincoln County; Branchland (m&r), Myra (m&r), Sweetland (m&r).
Logan County; Blair (m&r), Davin (ndr). Sharpies (m&r). Stone Branch
(ndr). McDowell County; Avondale (m&r). Marion County; Curtisville
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(ndr), Fairmont (ndr), High Point (ndr). Marshall County; Bannon
(ndr). Mason County; Leon (ndr). Mercer County; Oakvale (m&r), Prince-
ton (m&r), Spanishburg (m&r). Mingo County; Taylorsville (m&r), Wil-
liamson (m&r). Monongalia County; Morgantown (hhs). Monroe County;
Alderson (gkm), Salt Sulphur Springs (ndr) (m&r), Waitesville (m&r).
Willow Bend (m&r). Nicholas County; Lockwood (gkm), Richwood (ndr),
Snake Den Mt. (ndr), Summersville (gkm). Pendleton County; Franklin
(ndr) (gkm), Judy Gap (gkm). Pocahontas County; Black Mt. (p&l). Dun-
more Spring (gkm), Marlinton (gkm), Mill Point (gkm). Putnam County;
Poca (gkm). Raleigh County; Eccles (m&r). Randolph County; Helvetia
(vs), Huttonsville (gkm). Ritchie County; Pennsboro (frw). Summers
County; Riffle (gkm), Talcott (gkm). Upshur County; French Creek (mrs.
gkm) (gkm). Wayne County; Fort Gay (m&r). Webster County; Bolair
(ndr) (gkm). Wetzel County; Silver Hill (ndr). Wyoming County; Baileys-
ville (m&r), Pineville (m&r).
Remarks: The numerous narrow whorls on the upper surface, together
with the minute, rounded umbilicus, and narrow aperture, are sufficient
to distinguish this species. There are from two to four rows of very min-
ute, delicate teeth on the lower side of the interior of the last whorl,
radiating from the center. One row is usually so near the aperture that it
can be seen with the aid of a microscope ; the other rows are more or less
remote; each row contains from five to six distinct teeth, which are visible
through the shell.
Paravitrea lamellidens (Pilsbry)
(Plate 8, figs. 16, 17; Plate 9, fig. 1)
Shell small, thin, depressed, glossy, cinnamon-buff to darker reddish
brown in color, umbilicate; spire low-conoid ; suture moderately impressed ;
whorls 6)^, closely coiled, rounded; sculpture: first whorl smooth, re-
maining closely, regularly, and strongly striate, the striations much
weaker on peripheral and basal surfaces ; periphery rounded ; aperture nar-
rowly lunate, cavity obstructed by one to three white, curved, obliquely
protractive radial barriers within the outer and adjacent basal walls;
peristome thin, acute, simple, terminations widely separated, joined by a
very thin callus deposit, dilated close to columellar insertion; base flat-
tened, indented ; umbilicus narrow, deep, funnel shaped, not exhibiting all
inner volutions to apex. Diameter 3.5-3. 7 mm.; height 1. 6-1.9 mm.
Type Locality: Thunderhead, Great Smoky Mountains, Blount County,
Tennessee (Pilsbry, 1898).
Range: Tennessee; North Carolina; and West Virginia.
1949
MacMillan: Land Snails of West Virginia
159
West Virginia Records
Braxton County; Gassaway (gkm). Doddridge County; West Union
(frw).
Paravitrea capsella (Gould)
(Plate 9, figs. 2, 3)
Shell quite small, planorboid, pellucid, glistening, amber, colored.
Spire nearly plane, composed of about six and one-half, closely revolving,
flattened whorls. Surface with distant, impressed, radiating striae. Suture
margined. Aperture narrow, semilunar; lip simple, not thickened by a
callus within. Base perforated by a deep, rather small, funnel-shaped
umbilicus. Diameter one-fifth of an inch; axis one-tenth of an inch (5 x
2.5 mm.).
Type Locality: Tennessee (Gould, 1848).
Range: Alabama; Tennessee; Kentucky; West Virginia.
West Virginia Records
Boone County; Peytona (m&r), Racine (m&r), Seth (m&r). Clay County;
Clay (gkm). Grant County; Mount Storm (ndr). Kanawha County; Alum
Creek (ndr), Dunbar (gkm), Tornado (gkm). Logan County; Blair (m&r).
Sharpies (m&r). Stone Branch (ndr). McDowell County; Avondale (m&r).
Panther (m&r). Mercer County; Bluefield (m&r), Spanishburg (m&r).
Speedway (m&r). Mingo County; Taylorsville (m&r), Williamson (m&r).
Nicholas County; Lockwood (gkm). Rich wood (ndr). Pendleton County;
Franklin (gkm), Judy Gap (gkm). Preston County; Cheat Mt. (grh). Cold
Spring Park (grh). Summers County; Riffle (gkm), Talcott (gkm). Taylor
County; Grafton (frw). Wyoming County; Baileysville (m&r), Oceana
(m&r), Pineville (m&r).
Remarks: This species has the size and color of Retinella indentata, and
is similarly striated above. The whorls are numerous and closely con-
voluted, as in Ventridens suppressus, which it most nearly resembles; but
it has a larger umbilicus, like Ventridens lasmodon, and has no thickening,
or plate, within the aperture.
Paravitrea placentula placentula (Shuttleworth)
(Plate 9, figs. 4, 5)
Shell widely umbilicated, very much depressed, arctispiral, very shin-
ing, marked by irregular, distant, impressed striae, horn-color, diaphanous.
160
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VOL. 31
below of uniform color; whorls 7, most gradually increasing, scarcely con-
vex, the last convex below, subexcavated around the umbilicus; aperture
oblique, lunate; peristome simple, acute. Greater diameter 7^, lesser
634 ; height 3 mm.
Type Locality: Tennessee (Shuttleworth, 1852).
Range: Arkansas; Tennessee; North Carolina; Virginia; Kentucky; and
West Virginia.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r), Racine (m&r). Kanawha County; Hudnall (cbw). Logan
County; Blair (m&r), Davin (ndr). Sharpies (m&r). McDowell County;
Avondale (m&r), laeger (m&r). Panther (m&r). Mercer County; Bluefield
(m&r), Spanishburg (m&r). Mingo County; Taylorsville (m&r), William-
son (m&r). Monroe County; Sweetsprings (m&r). Pendleton County; Frank-
lin (gkm). Summers County; Talcott (gkm). Tucker County; Parsons
(mrs. gkm). Webster County; Bolair (m&r). Wyoming County; Pineville
(m&r).
Paravitrea placentula lacteodens (Pilsbry)
(Plate 9, figs. 6, 7)
The shell is similar to P. capsella, except that most specimens have one
to three pairs of tubercular teeth within the last whorl. The sutures are a
trifle less impressed, and the striations perceptibly closer. It differs from
P. significans in the usual persistence of the teeth in the adult stage, and
the median position of the periphery. In fully adult significance the peri-
phery is sub-basal, and there are no teeth. Altitude 2.6, diameter 5 mm.;
whorls 634-
Type Locality Cove,” Tuskeegee Mts., North Carolina (Pilsbry,
1903).
Range: Florida; North Carolina; and West Virginia.
West Virginia Records
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Clay
County; Clay {gkm) . Kanawha County; Alum Creek (ndr), Hudnall (cbw).
Lincoln County; Sweetland (m&r). Logan County; Blair (m&r), Davin
(ndr). Sharpies (m&r). Stone Branch (ndr). McDowell County; Avondale
(m&r), laeger (m&r), Panther (m&r). Marion County; Glady Creek (ndr),
High Point (ndr). Mercer County; Bluefield (m&r), Oakvale (m&r),
Spanishburg (m&r). Mingo County; Taylorsville (m&r), Williamson (m&r).
1949
MacMillan: Land Snails of West Virginia
161
Monroe County; Alderson (gkm), Sweetsprings (grh) (m&r), Willow Bend
(m&r). Nicholas County; Lockwood (gkm). Summers County; Riffle (gkm),
Talcott (gkm). Wayne County; Fort Gay (m&r). Wyoming County; Baileys-
ville (m&r), Oceana (m&r), Pineville (m&r).
Remarks: This variety was first considered a subspecies of Paravitrea
capsella, but many conchologists have now made it a subspecies of P.
placentula. It is distinguished from the former by the persistence of the
pairs of teeth in the adult shells.
Paravitrea petrophila (Bland)
(Plate 9, fig. 8)
Shell broadly umbilicate, depressed; subglobose, thin, shining, trans-
lucent, whitish, irregularly striated ; suture moderately impressed ; whorls
53^-6, rather convex, the last more convex, not descending; umbilicus
widely excavated externally; pervious; aperture roundly lunate; peristome
simple, somewhat thickened, often rose-colored, the columellar margin
slightly reflected. Diameter major 6, minimum 5-534; altitude 3 mill.
Type Locality: The Cliffs, Knoxville, Tennessee (Bland, 1883).
Range: Arkansas; Georgia; South Carolina; North Carolina; Tennessee;
West Virginia.
West Virginia Records
Marion County; Fairmont (ndr). Mason County; Leon (ndr). Monon-
galia County; Sturgisson (ndr). Pendleton County; Franklin (ndr) (gkm).
Summers County; Talcott (gkm). Webster County; Bolair (m&r).
Remarks: This species, in general form, is closely allied to Zonitoides
arboreus, but the color is different, the sutures are more developed, and the
umbilicus is much wider.
Paravitrea reesei J. P. E. Morrison
(Plate 9, figs. 9, 10, 11)
Shell small, subdiscoidal, polished, the last 5^ whorls (of type) closely
wound. Spire low, with shallow sutures. Periphery well rounded above
and below, is an almost even curve from suture to umbilicus. Sculpture
consisting of irregular spaced growth wrinkles or radial grooves; spiral
sculpture indistinct above and below. Umbilicus deep, well-like, exhibit-
ing all the whorls to the apex, contained about 5 times in major diameter
of the shell. Aperture transverse-lunate; lip thin, simple. Internal arma-
ture consisting in the smallest shell seen (of 2 to 234 whorls and 1 to 1.3
mm. major diameter) of two conical teeth in a radial row, dividing the
162
Annals of the Carnegie Museum
VOL. 31
periphery into three almost equal sectors. In a specimen of 3 whorls and
1.4 mm. major diameter, two other teeth appear, a third prominent coni-
cal tooth basal to the earlier pair, and a fourth which is an indistinct
callous or tubercle just beneath the suture. All teeth are retained in the
largest (adult) specimens; the uppermost prominent tooth is at the
periphery, the two others in each row are evenly spaced on the base of the
whorl. Height 1.6 mm.; major diameter 3.1 mm.; minor diameter 3 mm.;
aperture height 1.2 mm.; aperture width 1.45 mm.; umbilicus diameter
0.6 mm.
Type Locality: West Virginia Record; Monroe County; Peters Mt.
(Morrison, 1937) (grh).
Remarks: This new form may be easily distinguished by its small size ;
three prominent teeth in a radial row retained in large shells; and by an
umbilicus narrower than that of P. pilshryana.
Genus Retinella Fischer
Shell small, subglobose to depressed, thin, pellucid; epidermis pale or
shining, almost colorless to brownish horn-color ; finely wrinkled or striated,
or with regular equidistant impressed transverse lines, with or without
microscopic spiral wrinkles or impressed lines; whorls ?>]/2 to 7, rounded,
convex or planulate, regularly increasing, the last globose ; suture moder-
ately impressed; aperture large, transverse, depressed, lunate to sub-
orbicular in shape; peristome simple, acute; umbilicus covered by re-
flected peristome or broadly umbilicated and showing all inner whorls.
Retinella electrina (Gould)
(Plate 9, fig. 12; Plate 10, fig. 1)
Shell umbilicated, small, depressed, thin, fragile; epidermis pale, or
brownish horn-color, wrinkled, shining; whorls 4, the last rapidly en-
larging towards the aperture; aperture transversely rounded; peristome
simple, its edge rather thickened, not acute; umbilicus small, but well
marked and constant. Greater diameter 5, lesser 4^ mm.; height 2 mm.
Type Locality: Borders of Fresh Pond, Cambridge, Massachusetts
(Gould, 1841).
Range: Newfoundland; Canada; Alaska; United States.
West Virginia Records
Barbour County; Philippi (gkm). Boone County; Peytona (m&r), Racine
(m&r). Cabell County; Lesage (m&r). Grant County; Greenland Gap (ndr).
Greenbrier County; Alderson (gkm). Jefferson County; Bolivar (msb).
1949
MacMillan; Land Snails of West Virginia
163
Meyerstown (msb), Shenandoah Junction (msb). Lincoln County; Branch-
land (m&r). Marion County; Glady Creek (ndr). Mingo County; William-
son (m&r). Monongalia County; Morgantown (hhs). Monroe County;
Waitesville (m&r). Nicholas County; Craigsville (gkm), Summersville
(gkm). Ohio County; Wheeling (ndr). Pendleton County; Franklin (ndr)
(gkm). Wayne County; Echo (m&r). Webster County; Bolair (ndr). Wyom-
ing County; Pineville (m&r).
Remarks: In size, depressed-conical shape of the upper surface, the
number of whorls, and the rapid enlargement of the last whorl, this
species corresponds with R. indentata. It differs in the darker, smoky
horn-color, its constant umbilicus, its rather thick and shining lip, and its
whitish wrinkles, which, instead of being remote, are crowded as in other
species. From Zonitoides arbor eus it differs in having one whorl less, the
last one dilating; its apex not being depressed, its thinner, more shining
structure, and its somewhat smaller umbilicus. In Z. arboreus the outer lip
has a flexuous curve, but is nearly a direct section of the whorl in this.
Retinella electrina, described as Helix electrina (Gould, Invert. Mass.,
1841), has been known also as Helix radiatula (Aid., Trans. Nat. Hist.
Soc. Northumb., 1830), Helix viridula (Menke, Sym., ed. 2, 1830), and
Helix hammonis (Strom., Trondl., 1765). It seems that electrina has been
selected for the American species to distinguish it from the European and
other exotic species.
Retinella binneyana (Morse)
(Plate 10, figs. 2, 3)
Shell umbilicated, subglobose, transparent, almost colorless, shining,
smooth, with microscopic wrinkles of growth and still more delicate
oblique wrinkles; spire not much elevated; whorls about 4, rounded,
gradually enlarging, the last globose, broadly umbilicated below ; aperture
oblique, subcircular, large; peristome simple, acute, extremities not ap-
proaching, that of columella subreflected. Greatest diameter 4 mm.;
height 2 mm.
Type Locality: Maine (Morse, 1864).
Range: Quebec to British Columbia; Alaska; northern United States
from Maine and West Virginia to California.
West Virginia Records
Boone County; Racine (m&r). Clay County; Clay (gkm). Hampshire
County; Romney (gkm). Kanawha County; Dunbar (gkm). McDowell
164
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VOL. 31
County; Panther (m&r). Mercer County; Princeton (m&r). Mingo County;
Taylorsville (m&r). Monroe County; Salt Sulphur Springs (ndr), Waites-
ville (m&r). Pendleton County; Judy Gap (gkm). Pocahontas County;
Hillsboro (gkm). Randolph County; Cheat Bridge (gkm), Huttonsville
(gkm). Tucker County; Parsons (mrs. gkm) (gkm). Webster County; Bolair
(m&r).
Remarks: This species is nearest allied to R. electrina; it differs from that
species in the following particulars; it is nearly one-third smaller, and its
color is quite different, being white with a greenish tinge.
Retinella burringtoni (Pilsbry)
(Plate 10, figs. 4, 5)
The shell is depressed, umbilicate, glossy, somewhat translucent, of a
warm buff tint. It resembles R. rhoadsi but differs by having the retrac-
tive radial grooves less widely spaced, minor grooves and wrinkles be-
tween them more strongly developed ; on the latter part of the last whorl
the grooves become closely though somewhat irregularly spaced. Under
the compound microscope the surface is seen to be covered with fine,
distinct, weakly beaded spiral striae, not seen in R. rhoadsi. The umbilicus
is contained about 4.4 times in the diameter. The spire is slightly convex;
four rapidly widening whorls. The aperture is lunate, shaped much as in
R. rhoadsi. Height 2, diameter 4 mm.
Type Locality: Near and at Natural Bridge, Virginia (Pilsbry, 1928).
Range: Connecticut and New York, south to West Virginia and Vir-
ginia.
West Virginia Records
Barbour Nestorville (gkm), Philippi (gkm). Boone County; Pey-
tona (m&r), Seth (m&r). Braxton County; Frametown (gkm), Gassaway
(gkm). Cabell County; Lesage (m&r). Fayette County; Clifftop (gkm).
Greenbrier County; North Cad well (grh), Renich Valley (ndr), Ronceverte
(gkm) (m&r). Hampshire County; Romney (gkm). Jefferson County;
Charles Town (msb). Harper’s Ferry (jpem), Mt. Mission (jpem), Reedson
(msb). Kanawha County; Dunbar (gkm). Institute (gkm). Lewis County;
Jackson’s Mill (gkm). Lincoln County; Myra (m&r), Sweetland (m&r).
Logan County; Blair (m&r). McDowell County; Avondale (m&r). laeger
(m&r). Panther (m&r). Mason County; Point Pleasant (ndr). Mercer
County; Oakvale (m&r), Princeton (m&r). Speedway (m&r). Mingo County;
Taylorsville (m&r), Williamson (m&r). Monongalia County; Sturgisson
(ndr). Monroe County; Salt Sulphur Springs (m&r), Sweetsprings (m&r),
1949
MacMillan: Land Snails of West Virginia
165
Waitesville (m&r). Morgan County; Berkeley Springs (ndr). Pendleton
County; Circleville (ndr), Franklin (ndr) (gkm), Seneca Rocks (ndr),
Upper Tract (gkm). Pocahontas County; Dunmore Spring (gkm). Marlin-
ton (gkm). Putnam County; Poca (gkm). Summers County; Riffle (gkm).
Tucker County; Parsons (mrs. gkm) (gkm). Wayne County; Echo (m&r),
Fort Gay (m&r), Genoa (m&r). Webster County; Bolair (ndr) (gkm).
Remarks: Fresh shells of R. hurringtoni show distinct spiral striae and
and widely spaced, radial lines. The shell also has a dull sheen, produced
by the more sharply cut sculpture, while that of R. electrina looks like it
has been varnished. In addition hurringtoni has more depressed whorls
and usually develops a lower spire.
Retinella circumstriata (J. W. Taylor)
(Plate 10, figs. 6, 7)
Shell small, depressed, perforate, with impressed lines. Whorls 4^,
rounded, regularly and gradually increasing, the last one large. Sculpture
consisting of regularly spaced, crowded impressed lines above, fainter
below, and fine and crowded spiral impressed lines above and below.
Amber colored. Aperture large, oblique, subcircular; peristome simple,
acute. Umbilicus narrow, deep, exhibiting all inner volutions. Greater
diameter 334, lesser 2 9/10 mm.; height 1}^ mm.
Type Locality: Wetumpka, Alabama (Taylor, 1908).
Range: Arkansas; Alabama; and West Virginia.
West Virginia Records
Barbour County ; Si estorwiW^ (gkm), Philippi {gkm) .Boone County
(m&r), Racine (m&r), Seth (m&r). Braxton County; Gassaway (gkm).
Cabell County; Barboursville (m&r), Lesage (m&r), Ona (m&r). Fayette
County; Clifftop (gkm). Greenbrier County; Alderson (gkm). Renick (grh).
Lewis County; Jackson’s Mill (gkm). Lincoln County; Branchland (m&r),
Myra (m&r), Sweetland (m&r). Logan County; Blair (m&r). Marion
County; High Point (ndr). Marshall County; Bannon (ndr). Mercer
County; Bluefield (m&r), Princeton (m&r). Speedway (m&r). Mingo
County; Williamson (m&r). Monroe County; Gap Mills (m&r). Nicholas
County; Craigsville (gkm), Summersville (gkm). Pendleton County; Judy
Gap (gkm). Pocahontas County; Dunmore Spring (gkm), Marlinton (gkm).
Mill Point (gkm). Putnam County; Poca (gkm). Raleigh County; Eccles
(m&r). Randolph County; Huttonsville (gkm). Summers County; Talcott
(gkm). Tucker County; Parsons (mrs. gkm) (gkm). Upshur County; French
166
Annals of the Carnegie Museum
VOL. 31
Creek (mrs. gkm) (gkm). Wayne County; Echo (m&r), Genoa (m&r).
Wetzel County; Silver Hill (ndr). Wyoming County; Pineville (m&r).
Remarks: J. W. Taylor’s original description {Vitrea radiatula electrina
circumstriata) is as follows: “It is, therefore, proposed to regard as strictly
typical of the race {electrina), only those specimens possessing an amber-
coloured shell and upon which the microscopic striation is apparently
absent or but faintly visible. . . . and to apply the term circumstriata to
the shells clearly exhibiting the deep and distinct revolving striae. . .
According to H. B. Baker, Proc. Acad. Nat. Sci. Phila., vol. 82, 1930, pp.
203, 204, this species should be based on Dr. Bryant Walker’s paper,
Terrestrial Shell-hearing Mollusca of Alabama, 1928, p. 78, which seems to
be the first use of a trinomial that is accompanied by a recognizable de-
scription, since a quadrinomial has no status in nomenclature. I believe,
however, that this species should be dated on J. W. Taylor’s paper, even
if it was there described under a quadrinomial, as Bryant Walker copied
Taylor’s original description in his paper.
Retinella wheatleyi (Bland)
(Plate 10, figs. 8, 9)
Shell umbilicated, depressed, thin, shining, pelucid, brownish horn-
colored, finely striated; spire subplanulate, suture slightly impressed;
whorls little convex, the last more convex at the base, rapidly increasing
at the aperture, scarcely descending; umbilicus pervious; aperture de-
pressed, obliquely lunate; peristome simple, acute, the margins approxi-
mating, joined by a thin callus. Diameter major 5, minimum S}/^; alti-
tude 2 millimeter.
Type Locality: The Cliffs, Knoxville, Tennessee (Bland, 1883).
Range: Pennsylvania; Illinois; south to Tennessee, North Carolina, and
Albama. West Virginia Record: Mingo County; Williamson (m&r).
Remarks: R. wheatleyi is more nearly allied to R. electrina than any other
North American form, but differs from that species especially in the form
of the aperture, in the descending last whorl, and in having a wider
umbilicus.
This species was collected on ledges of rocks among dead leaves at the
type locality where the cliffs are very steep and rocky, and face north,
and are almost always shady, damp, and covered with mosses and ferns.
It also lives in grass and under stones in fields and other types of open
country, and is quite often found buried in plant trash in shrubby thickets.
1949
MacMillan: Land Snails of West Virginia
167
Retinella virginica J. P. E. Morrison
(Plate 10, figs. 10, 11, 12)
Shell markedly depressed, somewhat flattened above and below, um-
bilicate, vitreous, pinkish-horn colored. The radial grooves (major
growth wrinkles) are rather closely but irregularly spaced ; minor growth
wrinkles less prominent; with minute spiral strise above and below, less
distinct than in R. hurringtoni. The spire is lower than that of any of the
related species, in some examples approaching a plane; whorls 5 to 6 in
adult shells. The earlier whorls seen from above slowly increasing and
closely wound; the last whorl not rapidly expanding as in R. wheatleyi.
Umbilicus funicular, rapidly widening by the centrifugal growth of the
body whorl, in immature shells contained about 5 times in major diameter
of shell; in adult shells about 3^ times in major diameter of shell. Aper-
ture transverse, wider than high; upper end of peristome meeting the
penultimate whorl horizontally well above its periphery. Height 2.1
mm.; major diameter 5.3 mm.; minimum diameter 4.6 mm.; aperture
height 1.7 mm.; aperture diameter 2.1 mm.; umbilicus diameter 1.4 mm.
Type Locality: West slope of Blue Ridge, Clarke County, Virginia (Mor-
rison, 1937).
Range: Virginia and West Virginia. West Virginia Record: Jefferson
County; Loudoun (jpem).
Remarks: This species may be distinguished by its larger size; by the
proportionately wider umbilicus of adults; by the greater number of more
slowly increasing whorls, with a lower spire and proportionately smaller
aperture.
Retinella lewisiana (Clapp)
(Plate 11, figs. 1, 2, 3)
Shell small, depressed widely, perspectively umbilicate, all whorls
showing to the apex, umbilicus contained about five times in the diameter
of the shell ; yellowish-white, translucent, the inner whorls showing through
the body of the shell, highly polished; the delicate growth lines are very
regularly spaced and close together ; smooth below. Spire much flattened ;
suture well impressed; margined; whorls slightly convex, the last
wide. Aperture oblong-lunate, depressed above, lower margin parallel
with the base, lip simple. Greater diameter 33^, lesser 2.8, altitude 13^ mm.
Type Locality: Monte Sano, near Huntsville, Alabama (Clapp, 1908).
Range: Alabama; Tennessee; and West Virginia.
West Virginia Records
Cabell County; Barboursville (m&r). Lewis County; Jackson’s Mill
(gkm). Lincoln County; Branchland (m&r). Logan County; Blair (m&r).
168
Annals of the Carnegie Museum
VOL. 31
McDowell County; Avondale (m&r), laeger (m&r). Pendleton County;
Judy Gap (gkm). Putnam County; Poca (gkm).
Remarks: The color, and particularly the very regular close lines of
growth, at once distinguishes this shell from all other species. It is per-
haps nearest to R. dalliana, but differs in color, shape, and sculpture. In
dalliana there is a very minute spiral sculpture, only visible under a
magnification of about 60 times.
Retinella raderi (Dali)
(Plate 11, figs. 4, 5, 6)
Shell depressed, four-whorled, smooth except for faint, rather regularly
spaced, incremental lines above, of a pale waxen whitish color; spire raised
above the last whorl, which is much the largest; periphery evenly rounded,
suture appressed, base moderately rounded, the umbilical slope of the last
whorl somewhat flattish; umbilicus very wide, exhibiting all the volutions;
aperture wider than high, the upper margin slightly in advance of the
lower lip, the two connected by a thin wash of callus over the body.
Altitude 1.5, maximum diameter 4, minimum diameter 3 mm.
Type Locality: Cumberland, Maryland (Dali, 1898).
Range: Maryland and West Virginia.
West Virginia Records
Cabell County; Barboursville (m&r). Grant County; Greenland Gap
(ndr). Greenbrier County; Alderson (ndr). Logan County; Sharpies (m&r).
Marion County; High Point (ndr). Mercer County; Speedway (m&r).
Mingo County; Williamson (m&r). Monroe County; Salt Sulphur Springs
(m&r), Waitesville (m&r). Pendleton County; Franklin (ndr) (gkm),
Upper Tract (ndr). Pocahontas County; Marlinton (gkm).
Remarks: The nearest relative of this species is R. wheatleyi, which is a
larger shell with higher spire, more rounded whorls, and a much smaller
and more steep-sided umbilicus.
Retinella rhoadsi rhoadsi (Pilsbry)
(Plate 11, figs. 7, 8)
Similar to R. indentata, but differing from that species in the distinct
umbilicus, about one-half millimeter wide, showing the penultimate whorl
within; radial grooves more numerous, and therefore closer. The same
character, and the smaller size, separate rhoadsi from R. carolinensis.
Altitude 2.5, diameter 4.8 mm., or somewhat smaller.
1949 MacMillan: Land Snails of West Virginia 169
Type Locality: White Pond, Warren County, New Jersey (Pilsbry,
1899).
Range: Ontario; Michigan; Maine to Pennsylvania, south to Tennessee
and North Carolina.
West Virginia Records
Barbour County; Philippi (mrs. gkm) (gkm). Boone County; Peytona
(m&r), Racine (m&r). Braxton County; Gassaway (gkm). Cabell County;
Lesage (m&r), Ona (m&r). Clay County; Clay (gkm). Grant County; Green-
land Gap (ndr). Greenbrier County; Alderson (gkm), Ronceverte (gkm).
Hampshire County; Capon Bridge (jpem), Romney (gkm). Jefferson
County; Bloomery (msb) (Bartsch & Morrison), Charles Town (jpem).
Harper’s Ferry (jpem). Kanawha County; Alum Creek (ndr), Hudnall
(cBw). Lewis County; Weston (gkm). Lincoln County; Branchland (m&r),
Myra (m&r), Sweetland (m&r). Logan County; Blair (m&r). Sharpies
(m&r). McDowell County; Panther (m&r). Marion County; East Fairmont
(ndr), Fairmont (ndr). High Point (ndr). Mineral County; Burlington
(gkm). New Creek (ndr). Mingo County; Williamson (m&r). Monongalia
County; Coopers Rock (ndr). Monroe County; Salt Sulphur Springs (ndr)
(m&r), Sweetsprings (grh) (m&r), Waitesville (m&r). Nicholas County;
Summersville (gkm). Ohio County; Wheeling (ndr). Pendleton County;
Franklin (gkm), Judy Gap (gkm). Pocahontas County; Marlinton (gkm).
Raleigh County; Shady Spring (m&r). Randolph County; Huttonsville
(gkm). Summers County; Riffle (gkm), Talcott (gkm). Tucker County;
Parsons (gkm). Upshur County; French Creek (mrs. gkm) (gkm). Wayne
County; Fort Gay (m&r). Webster County; Bolair (m&r). Wirt County;
(wjf). Wyoming County; Pineville (m&r).
Retinella rhoadsi austrina H. B. Baker
(Plate 11, figs. 9, 10)
Shell quite small, hyaline, thin and transparent; spire more depressed
than in rhoadsi; umbilicus smaller (8 to 9 times in major diameter) ; spiral
sculpture quite weak; radiating lines rather closely spaced (41 on last
whorl) .
Type Locality: Sink Hole, about three miles northwest of Johnson City,
Washington County, Tennessee (Baker, 1930).
Range: Tennessee; North Carolina; Virginia; and West Virginia.
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VOL. 31
West Virginia Records
Calhoun County; Grantsville (ndr). Grant County; Mount Storm (ndr).
Mason County; Point Pleasant (ndr). Morgan County; Berkeley Spring
(ndr). Pendleton County; Franklin (ndr).
Retinella indentata indentata (Say)
(Plate 11, figs. 11, 12)
Shell depressed, pellucid, highly polished ; whorls four, with regular,
distant, subequidistant, impressed lines across, of which there are about
twenty -eight to the body whorl, all extending to the base; suture not
deeply indented; aperture rather large; labrum simple, terminating at its
inferior extremity at the center of the base of the shell; umbilicus none,
but the umbilical region is deeply indented. Greater breadth one-fifth of
an inch (5 mm.).
Type Locality: Harrigate, near Philadelphia, Pennsylvania (Say, 1822).
Range: Canada and the United States.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r), Racine (m&r), Seth (m&r). Braxton County; Frametown
(gkm), Gassaway (gkm). Cabell County; Barboursville (gkm) (m&r),
Lesage (m&r), Ona (m&r). Clay County; Clay (gkm). Doddridge County;
West Union (frw). Fayette County; Clifftop (gkm). Grant County;
Greenland Gap (ndr). Greenbrier County; Alderson (ndr). Renick
Valley (ndr), Ronceverte (gkm) (m&r). Hampshire County; Capon Bridge
(jpem), Romney (gkm). Harrison County; Bristol (ndr). Jefferson County;
Aldridge (msb), Bakerton (msb), Bardane (msb), Bloomery (jpem),
Bolivar (msb), Charles Town (msb). Harper’s Ferry (msb) (ndr) (jpem)
(ejc) (gwt), Jamestown (msb), Kabletown (msb). Keys Gap (jpem).
Keystone (msb), Meyerstown (msb). Middleway (msb), Millville (msb),
Morgan Grove (msb), Mt. Mission (jpem), Reedson (msb), Rippon (msb),
Shenandoah City (msb), Shenandoah Junction (msb), Uvilla (msb).
Kanawha County; Alum Creek (ndr), Dunbar (gkm), Hudnall (cbw).
Institute (gkm), Nitro (gkm), St. Albans (um). South Charleston (ndr).
Tornado (gkm). Lewis County; Jackson’s Mill (gkm). Lincoln County;
Branchland (m&r), Myra (m&r), Sweetland (m&r). Logan County; Blair
(m&r), Davin (ndr). Sharpies (m&r). Stone Branch (ndr). McDowell
County; laeger (m&r). Panther (ndr) (m&r). Marion County; Curtisville
(ndr), Fairmont (ndr), Glady Creek (ndr). High Point (ndr), Jayenne
1949
MacMillan: Land Snails of West Virginia
171
(grh), Marshall County; Cameron (gb), Moundsville (vs). Mason County;
Leon (ndr). Mercer County; Oakvale (m&r), Princeton (m&r), Speedway
(m&r). Mineral County; New Creek (ndr). Mingo County; Taylorsville
(m&r), Williamson (m&r). Monongalia County; Morgantown (hhs). Mon-
roe County; Alderson (gkm), Salt Sulphur Springs (ndr) (m&r), Sweet-
springs (m&r). Union (gkm), Waitesville (m&r), Willow Bend (m&r).
Morgan County; Largent (ndr) . Nicholas County; Richwood (ndr) . Pend-
leton County; Circleville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm),
Seneca Rocks (ndr), Upper Tract (ndr). Pocahontas County; Dunmore
Spring (gkm), Marlinton (gkm). Mill Point (ndr). Putnam County; Poca
(gkm). Raleigh County; Daniels (m&r). Randolph County; Huttonsville
(gkm). Summers County; Riffle (gkm), Talcott (gkm). Tucker County;
Parsons (mrs. gkm) (gkm). Upshur County; French Creek (mrs. gkm).
Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County; Bolair
(ndr) (m&r). Wetzel County; Proctor (vs), Silver Hill (ndr). Wirt County;
(wjf). Wyoming County; Baileysville (m&r), Oceana (m&r), Pineville
(m&r).
Remarks: This species may be mistaken for Zonitoides arboreus, but it
is destitute of the umbilicus, in place of which there is an indented center
to the base in which the labrum terminates. The spire is very much de-
pressed, and the surface radiated by distant impressed lines, the inter-
stices being perfectly smooth.
Retinella indentata paucilirata (Morelet)
(Plate 12, fig. 1)
Shell convexly depressed, horn-colored, shining, smooth, indented.
Whorls 43^, flat, rounded, the last subdilated in front, not deflected.
Sculpture consists of radiating indentations remotely and equidistantly
placed above and below. Aperture oblique, lunately oval ; peristome simple,
acute. Umbilicus closed, but the region around it indented. Greatest
diameter 6 mm. ; altitude 2 mm.
Type Locality: Near Salama, Guatemala (Morelet, 1851).
Range: New Jersey to California, south to Arizona and Florida.
West Virginia Records
Barbour County; Nestorville (gkm). Boone County; Racine (m&r).
Cabell County; Barboursville (ndr) (m&r), Ona (m&r). Clay County; Clay
(gkm). Fayette County; Clifftop (gkm). Grant County; Mount Storm (ndr).
Greenbrier County; Renick Valley (ndr), Ronceverte (gkm). Hampshire
172
Annals of the Carnegie Museum
VOL. 31
County; Romney {gkm). Jefferson County; Rippon (msb). Kanawha County;
Alum Creek (ndr), Dunbar (gkm). Lincoln County; Myra (m&r). Logan
County; Davin (ndr), Sharpies (m&r), Stone Branch (ndr). McDowell
County; Avondale (m&r). Panther (m&r). Marion County; Curtisville
(ndr). High Point (ndr). Mingo County; Taylorsville (m&r), Williamson
(m&r). Monroe County; Salt Sulphur Springs (ndr) (m&r), Waitesville
(m&r). Pendleton County; Circleville (ndr), Franklin (ndr), Judy Gap
(gkm). Upper Tract (ndr). Pocahontas County; Marlinton (gkm). Putnam
County; Poca (gkm). Summers County; Talcott (gkm). Taylor County;
Valley Falls (ndr). Tucker County; Parsons (mrs. gkm) (gkm). Wayne
County; Echo (m&r). Fort Gay (m&r). Wetzel County; Silver Hill (ndr).
Wyoming County; Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Remarks: This variety is distinguished from the typical species by the
radiating lines, which are further apart in indentata paucilirata than in
indentata.
Retinella carolinensis (Cockerell)
(Plate 11, figs. 13, 14, 15)
Maximum diameter 10, altitude 5 mm., whorls 5. Pale-horn, shiny,
semi-transparent, umbilical region somewhat whitened. Surface of shell
with strong transverse growth lines and distinct transverse grooved lines.
The grooved lines number about 26 on body whorl. Umbilicus small,
narrow. Aperture large-lunate, the upper angle much smaller than lower.
Peristome not sinuate.
Type Locality: Not designated (Cockerell, 1890).
Range: Alabama and Georgia, north to West Virginia and Maryland.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Berkeley County;
Hodgesville (ndr). Boone County; Peytona (gb) (m&r), Racine (m&r),
Seth (m&r). Braxton County; Gassaway (gkm). Cabell County; Barbours-
ville (ndr) (gkm) (m&r), Lesage (m&r), Ona (m&r). Calhoun County;
Grantsville (ndr). Clay County; Clay (gkm). Fayette County; Cliff top
(gkm). Grant County; Greenland Gap (ndr). Greenbrier County; Alderson
(gkm). Organ Cave (ndr), Ronceverte (gkm) (m&r). Hampshire County;
Romney (gkm). Jefferson County; Bolivar (msb), Charles Town (msb),
Jamestown (msb), Meyerstown (msb). Kanawha County; Alum Creek
(ndr), Hudnall (cbw). Lincoln County; Branchland (m&r), Myra (m&r),
Sheridan (ndr), Sweetland (ndr). Logan County; Blair (m&r), Davin
1949
MacMillan: Land Snails of West Virginia
173
(ndr), Sharpies (m&r), Stone Branch (ndr). McDowell County; Avondale
(m&r), laeger (m&r), Panther (ndr) (m&r). Marion County; Curtisville
(ndr), Fairmont (ndr), Glady Creek (ndr). High Point (ndr), Rives-
ville (ndr). Mercer County; Bluefield (m&r), Oakvale (m&r), Princeton
(m&r), Spanishburg (m&r). Speedway (m&r). Mineral County; New Creek
(ndr). Mingo County; Taylorsville (m&r), Williamson (m&r). Monongalia
County; Smithtown (ndr). Monroe County; Alderson (gkm), Salt Sulphur
Springs (m&r), Sweetsprings (m&r), Waitesville (m&r). Willow Bend
(m&r). Morgan County; Berkeley Springs (ndr). Nicholas County; Lock-
wood (gkm), Summersville (gkm). Ohio County ;W\ie&\mg (ndr). Pendleton
County; Circleville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm), Upper
Tract (ndr). Pocahontas County; Dunmore Spring (gkm), Greenbank
(gkm), Marlinton (gkm). Mill Point (gkm). Raleigh County; Daniels
(gkm), Eccles (m&r). Randolph County; Huttonsville (gkm). Tucker
County; Parsons (mrs. gkm) (gkm). Upshur County; French Creek (mrs.
gkm) (gkm). Wayne County; Echo (m&r). Fort Gay (m&r), Genoa (m&r).
V/ebster County; Bolair (gkm). Wyoming County; Baileysville (m&r),
Oceana (m&r), Pineville (m&r).
Retinella cryptomphala cryptomphala (Clapp)
(Plate 12, figs. 2, 3)
Shell thin, polished, very light horn-color to white, generally white,
translucent; whorls 5 to 5}^, those of the spire regularly increasing, the
last widening very rapidly and doubling the diameter of the shell. Aper-
ture broadly lunate, sutures well impressed, ail whorls showing through
the shell. Surface sculptured with rather evenly spaced, radiating grooves
continued to the base as in i?. indentata and carolinensis, there being from
23 to 34 on the body whorl. Microsculpture of even, close, clear-cut,
spiral, engraved lines like those of carolinensis, best seen with a magnifica-
tion of 25 diameters or over. Spire much depressed, almost flat, the termi-
nation of the last whorl slightly raised at the lip which is straight on the
lower edge and well curved forward above, projecting about 1 mm. beyond
the lower lip; at the columellar end the lip is thickened and joined to a
tongue-like callus which completely covers the umbilicus at all stages of growth.
There is a thin, microscopically granular parietal callus as in R. indentata.
Base of shell well rounded and less impressed in the umbilical region than
indentata or carolinensis. 5. 1-5.9 x 4.2-5 x 2. 1-2.8 mm.
Type Locality: Knox County, Tennessee (Clapp, 1915).
Range: Alabama; Tennessee; North Carolina; and West Virginia.
174
Annals of the Carnegie Museum
VOL. 31
West Virginia Records
Boone County; Peytona (m&r). Braxton County; Shadyside (gkm).
Cabell County; Barboursville (m&r), Lesage (m&r), Ona (m&r). Calhoun
County; Grantsville (ndr). Clay County; Clay (gkm). Fayette County;
Clifftop (gkm). Grant County; Greenland Gap (ndr). Greenbrier County;
Alderson (gkm), Ronceverte (gkm). Hampshire County; Romney (gkm).
Kanawha County; Hudnall (cbw). Lewis County; Weston (gkm). Lincoln
County; Branchland (m&r), Myra (m&r), Sweetland (m&r). Logan County;
Stone Branch (ndr). McDowell County; Avondale (m&r). Panther (m&r).
Marion County; High Point (ndr). Mason County; Point Pleasant (ndr).
Mercer County; Speedway (m&r). Mingo County; Williamson (m&r).
Monroe County; Sait Sulphur Springs (m&r). Pendleton County; Franklin
(ndr) (gkm), Judy Gap (gkm). Pocahontas County; Marlinton (gkm).
Putnam County; Poca (gkm). Raleigh County; Eccles (m&r). Summers
County; Talcott (gkm). Tucker County; Parsons (mrs. gkm) (gkm). Upshur
County; French Creek (mrs. gkm). Wayne County; Fort Gay (m&r), Genoa
(m&r). Webster County; Bolair (m&r). Wyoming County; Pineville (m&r).
Remarks: The flatter spire, the umbilicus covered at all stages of growth^
the less excavated umbilical region and the wider aperture separates this
species from R. carolinensis.
Retinella cryptomphala solida H. B. Baker
(Plate 12, fig. 4)
Shell: color corneous though fulvous to almost chestnut; typically
larger, heavier and with stronger sculpture than cryptomphala. Whorls
534- Height 4.05, greater diameter 7.5, lesser 6.48 mm.
Type Locality: Along calcareous ledges, south side of Prior Cove, near
Jasper, Marion County, Tennessee (Baker, 1930).
Range: Arkansas; Alabama; and Florida, north to Virginia and West
Virginia. West Virginia Record: Pendleton County; Upper Tract (gkm).
Retinella sculptilis (Bland)
(Plate 12, figs. 5, 6; Plate 13, fig. 1)
Shell scarcely perforate, suborbicular, depressed, subpellucid, pale horn-
color above, of lighter shade beneath, shining, with regular, subequidist-
ant, impressed transverse lines, those on the last whorl extending over the
periphery, and converging in the umbilical excavation; spire very little
elevated, scarcely convex; whorls 7, planulate, the last rapidly increasing.
1949 MacMillan: Land Snails of West Virginia 175
equal at the aperture to one-third of the diameter of the shell, beneath
flattened, and little excavated in the umbilical region; suture lightly im-
pressed; aperture scarcely oblique, depressed, transverse, lunate; peris-
tome simple, acute, sinuate, the columellar margin very rapidly and nar-
rowly reflected over, and almost entirely covering the small perforation.
Diameter major 123/^ minor 11, altitude 5 mm.
Type Locality: Anantchely Mountains, North Carolina (Bland, 1858).
Range: Alabama and Georgia, north to West Virginia and Kentucky.
West Virginia Records
Pendleton County; Judy Gap (gkm). Randolph County; Cheat Bridge
(gkm).
Remarks: In sculpture it is closely allied to R. indentata, but the im-
pressed striae are more numerous, and closer together.
Subfamily Vitrein^
Genus Haw aha Gude
Shell umbilicated, minute, depressed-convex to depressed; epidermis
whitish to whitish-corneous; whorls 3}/2 to 4^^, convex, increasing slowly,
with microscopic wrinkles or fine but rather sharp growth striae and
crowded microscopic spiral striae ; suture very distinctly impressed ; aper-
ture round-lunate ; peristome thin, acute, its columellar margin placed at
one-fourth or two-thirds of the penultimate whorl; umbilicus large, deep,
exhibiting the inner volutions.
Hawaiia minuscula minuscula (A. Binney)
(Plate 12, fig. 9)
Shell minute, depressed-convex; epidermis whitish; whorls 4, convex,
not increasing rapidly in diameter; suture very distinctly impressed;
aperture nearly circular; lip thin, acute; umbilicus large, not spread, deep,
and exhibiting the volutions; base rounded, columella with a thin callus.
Greatest transverse diameter less than one-eighth of an inch. Greater
diameter 23^^, lesser 2}^ mm.; height 1 mm.
Type Locality: Not designated (A. Binney, 1840).
Range: Ontario, Manitoba; entire United States.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (cb) (m&r), Racine (m&r), Seth (m&r). Braxton County ; G2,ssQiW2,y
176
Annals of the Carnegie Museum
VOL. 31
(gkm). Cahell County; Barboursville (gkm) (m&r), Lesage (m&r), Ona
(m&r). Calhoun County; Grantsville (ndr). Grant County; Greenland Gap
(ndr). Greenbrier County; Alderson (ndr) (gkm), Renick Valley (ndr),
Ronceverte (gkm) (m&r). White Sulphur Springs (p&l). Hampshire
County; Romney (gkm). Jefferson County; Bakerton (msb), Bardane
(msb), Bloomery (msb), Bolivar (msb), Charles Town (msb), Engle (msb).
Hall town (msb). Harper’s Ferry (ndr) (msb), Jamestown (msb), Kable-
town (msb), Kearneysville (msb), Keystone (msb), Leetown (msb),
Mechanicstown (msb), Meyerstown (msb). Middleway (msb), Millville
(msb), Morgan Grove (msb), Reedson (msb), Rippon (msb), Shenandoah
City (msb), Shepherdstown (msb), Skeetersville (msb). Summit Point
(msb), Uvilla (msb). Kanawha County; Alum Creek (ndr), Hudnall (cbw).
Tornado (gkm). Lewis County; Jackson’s Mill (gkm). Lincoln County;
Branchland (m&r), Myra (m&r). Logan County; Blair (m&r). Sharpies
(m&r). Stone Branch (ndr). McDowell County; Panther (m&r). Marion
County; Curtisville (ndr), Fairmont (ndr), Glady Creek (ndr). High
Point (ndr). Marshall County; Moundsville (vs). Mason County; Leon
(ndr). Point Pleasant (ndr). Mercer County; Princeton (m&r), Spanish-
burg (m&r). Speedway (m&r). Mineral County; Burlington (gkm). New
Creek (ndr). Mingo County; Taylorsville (m&r), Williamson (m&r).
Monongalia County; Coopers Rock (cbw). Monroe County; Alderson
(gkm). Salt Sulphur Springs (ndr) (m&r), Sweetsprings (grh) (m&r),
Waitesville (m&r). Willow Bend (m&r). Nicholas County; Lockwood
(gkm). Pendleton County; Circleville (ndr), Franklin (ndr) (gkm), Judy
Gap (gkm). Upper Tract (ndr). Pocahontas County; Greenbank (gkm),
Marlinton (gkm). Putnam County; Poca (gkm). Randolph County; Cheat
Bridge (gkm). Summers County; Talcott (gkm). Taylor County; Valley
Falls (ndr). Tucker County; Parsons (mrs. gkm) (gkm). Wayne County;
Fort Gay (m&r), Genoa (m&r). Webster County; Bolair (ndr). Wetzel
County; Proctor (vs). Silver Hill (ndr). Wyoming County; Baileysville
(m&r), Pineville (m&r).
Remarks: This species is slightly larger than Vallonia pulchella, which
it resembles in its upper surface. It also bears some resemblance to
Zonitoides limatulus. H. minuscula has four whorls, with a deep suture;
and a circular aperture, and a large umbilicus, which exhibits about two
volutions. The striae are too minute to be visible to the unaided eye.
1949
MacMillan: Land Snails of West Virginia
177
Hawaiia minuscula alachuana (Dali)
(Plate 12, figs. 7, 8)
Shell larger than H. minuscula. The junction of the inner lip of the
body whorl takes place outside of the middle line or even at the outer third
of the penultimate whorl. The aperture is a little dilated. A much larger
portion of the base of the penultimate whorl is shown and the umbilicus
is wider.
Type Locality: Alachuana County, Florida (Dali, 1885).
Range: Arizona; Texas; Arkansas; Kansas; New Jersey; West Vir-
ginia; and Florida.
West Virginia Records
Barbour County; Philippi (gkm). Boone County; Peytona (cb). Cabell
County; Barboursville (gkm) . Grant County; Greenland Gap (ndr) . Green-
brier County; Alderson (ndr), Ronceverte (gkm). Hampshire County;
Romney (gkm). Kanawha County; Nitro (gkm). Lincoln County; Branch-
land (m&r), Myra (m&r). Logan County; Sharpies (m&r). Marion County;
High Point (ndr), Kingmont (grh). Mason County; Leon (ndr). Mercer
County; Princeton (m&r). Monroe County; Salt Sulphur Springs (ndr),
Sweetsprings (grh) (m&r), Waitesville (m&r). Pendleton County; Franklin
(ndr) (gkm), Judy Gap (gkm), Upper Tract (ndr). Pocahontas County;
Greenbank (gkm). Tucker County; Parsons (gkm). Wayne County; Fort
Gay (m&r). Webster County; Bolair (ndr). Wyoming County; Pineville
(m&r).
Remarks: The larger shell, and the position of the inner lip of the body
whorl with the penultimate whorl, distinguishes this variety from the
typical species.
Subfamily Gastrodontin^
Genus Gastrodonta Albers
Shell small, perforate or imperforate, orbicularly depressed, yellowish-
red to greenish in color, polished. Whorls 5 to 8, regularly increasing,
rounded, sculptured with a few faint, irregularly spaced growth lines,
regular rows of oval beads, or regular, equidistant, elevated, obtuse rib-
striations. Aperture transverse, subcrescentic ; peristome simple, acute;
within furnished with from one to four pairs of lamellae.
178
Annals of the Carnegie Museum
VOL. 31
Gastrodonta interna (Say)
(Plate 13, figs. 2, 3, 4)
Shell very narrowly perforated, depressed, slightly convex; epidermis
reddish-brown, shining; whorls 8, with regular, equidistant, elevated,
oblique, rounded ribs, separated by distinct grooves; suture deeply im-
pressed; aperture flattened, transverse, narrow; peristome thin, acute,
thickened internally; within the base of the aperture, somewhat distant
from the margin are two prominent, sub-lamelliform, white teeth, not
reaching the edge of the peristome; base smooth, polished, umbilical re-
gion indented. Greater diameter 53^ mm.; height 3)^ mm.
Type Locality: Lower Missouri (Say, 1821).
Range: Pennsylvania to Missouri, south to Mississippi and Florida.
West Virginia Records
Braxton County; Frametown (gkm), Gassaway (gkm). Calhoun County;
Arnoldsburg (grh), Grantsville (ndr). McDowell County ; hvondoXe (m&r).
Mercer County; Speedway (m&r). Wayne County; Fort Gay (m&r). Wirt
County; (wjf).
Remarks: The sculpture of this species, consisting of regular, equidistant,
elevated, oblique, rounded ribs, distinguishes it from any other species of
snail known from eastern North America. These ribs are separated by
distinct grooves, and are obsolete on the base of the shell.
Gastrodonta fonticula Wurtz
“Shell distinctly umbilicate with the umbilicus contained about eight
times in the greatest diameter of the shell. The umbilicus scarcely nar-
rowing to the apex of the shell. Inner edge of last ^ (usually) of the body
whorl forming a ridge around the opening of the umbilicus. The base of
the body whorl is strongly convex; the greatest convexity about midway
between the umbilicus and the periphery. Between this and the umbilicus
the surface is slightly concave and ribbed. The concave surface has the
appearance of a broad, shallow groove surrounding the umbilicus. The
inner edge forms a narrowly rounded ridge arqund the umbilicus. Shell
of 7.9 whorls (type specimen). In the apical view the shell is not separable
from G. interna (Say). The sculpturing is identical in the two species. The
spire is dome-shaped, but not as high as in G. interna. Index of h/d-59%
for the type specimen. Aperture as in G. interna. Two teeth appearing
within the aperture as very short lamellae about as long as high. The
teeth are nearly the same size although the outer one shows a tendency
towards a greater height and width. (In G. interna the outer tooth is
appreciably bulkier than the inner tooth). Soft parts of the animal un-
known. Height 3-3.7 mm., greatest diameter 5. 7-6.9 mm. ; 7. 1-7.9 whorls.”
1949
MacMillan: Land Snails of West Virginia
179
Type Locality: On steeply sloping, wooded hillside on the east side of
Paint Creek, Hudnall, Kanawha County, West Virginia (Wurtz, 1948).
Range: Has been reported only from West Virginia.
West Virginia Records
Boone County; Peytona (ndr) (m&r), Racine (m&r), Seth (m&r). Clay
County; Clay (gkm). Kanawha County; Hudnal (cbw). Logan County;
Blair (m&r), Davin (ndr), Sharpies (m&r). McDowell County; laeger
(m&r). Panther (m&r). Upshur County; French Creek (gkm). Wyoming
County; Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Remarks: The most significant feature of this shell is the well-like
umbilicus permitting an uninterrupted view to the apex. This character
most readily allows its separation from G. interna (Say). G. fonticula is
decidedly distinct in its distribution and environmental habits, since G.
interna was not found at the same locality together with this species.
Genus Zonitoides Lehmann
Shell orbicular, depressed, slightly to moderately convex above and
concave below, thin, shining, umbilicate. Whorls 4 to 5, convex, slowly and
regularly increasing; last whorl larger, declining or not declining as it
approaches the aperture. Suture deeply impressed. Sculpture consisting
of delicate and oblique striae to irregular and close rib-striations above,
present or obsolete below. Whitish, greenish, or amber colored. Aperture
oblique, lunate-circular to circular, flattened above; peristome simple,
acute, thin, thickened or not within. Umbilicus narrow to large, deep;
all inner volutions showing or not.
Zonitoides elliotti (Redfield)
(Plate 13, fig. 5)
Shell with a rather narrow umbilicus, depressed-orbiculate, with fine
transverse striae, greenish horn-colored, scarcely translucent, shining
beneath; spire convex, but not much raised; whorls 5, rather convex, last
one sometimes slightly depressed at the aperture ; suture deeply impressed ;
aperture very oblique, lunate-circular; peristome a little sinuate, acute
but thickened within. Diameter major 9, diameter minor 8, altitude 4 mm.
Type Locality: Not designated (Redfield, 1856).
Range: Appalachian Mountains from Alabama to West Virginia.
180
Annals of the Carnegie Museum
VOL. 31
West Virginia Records
West Virginia; (Dr. W. D. Hartman Collection). Mingo County
liamson (m&r). Wayne County; (W. G. Binney).
Remarks: This species is a larger, more coarsely striated, and more ro-
bust shell than Z. arboreus, while its umbilicus spreads less, and the aper-
ture is more oblique. These same features distinguish it from Z. nitidus
and Z. limatulus. Moreover, the inward thickening of the peristome, a
marked feature of elliotti, is wanting in all other species of Zonitoides.
Zonitoides arboreus (Say)
(Plate 13, figs. 6, 7)
Shell umbilicated, depressed, very slightly convex, thin, pellucid;
epidermis amber-colored, smooth, shining; whorls 4-5, with very minute,
oblique striae, apparent when viewed with the microscope; aperture
transversely rounded; peristome thin, acute; umbilical region indented;
umbilicus moderate, well developed, round, and deep. Greater diameter 5,
lesser 4^ ; height 2^4 nim.
Type Locality: Not designated (Say, 1816).
Range: Newfoundland; Canada; Alaska; and United States.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (mrs. gkm) (gkm). Ber-
keley County; Hodgesville (ndr). Boone County; Peytona (cb) (m&r),
Racine (m&r), Seth (m&r). Braxton County; Frametown (gkm), Gassaway
(gkm), Shadyside (gkm). Brooke County; Bethany (cbw). Cabell County;
Barboursville (ndr) (gkm) (m&r), Lesage (m&r), Ona (m&r). Calhoun
County; Grantsville (ndr). Clay County; Clay (gkm). Fayette County;
Clifftop (gkm). Grant County; Mount Storm (ndr). Greenbrier County;
Alderson (gkm). Organ Cave (ndr). Renick (grh). Renick Valley (ndr),
Ronceverte (gkm) (m&r). White Sulphur Springs (p&l). Hampshire
County; Capon Bridge (jpem). Junction (ndr), Romney (gkm). Harrison
County; Bristol (ndr). Jefferson County; Bakerton (msb), Bloomery (jpem),
Bolivar (msb), Charles Town (jpem). Harper’s Ferry (msb) (gwt) (ejc),
Kearneysville (msb). Keys Gap (jpem). Keystone (msb), Meyerstown
(msb), Morgan Grove (msb), Reedson (msb), Shenandoah Junction (msb).
Summit Point (msb), Uvilla (msb). Kanawha County; Charleston (gkm),
Dunbar (gkm), Nitro (gkm). South Charleston (ndr). Tornado (gkm).
Lewis County; Jackson’s Mill (mrs. gkm) (gkm), Weston (gkm). Lincoln
1949
MacMillan: Land Snails of West Virginia
181
County; Branchland (m&r), Myra (m&r), Sweetland (m&r). Logan County;
Blair (m&r), Davin (ndr), Sharpies (mrs. gkm) (m&r), Stone Branch
(ndr). McDowell County; Avondale (m&r), laeger (m&r). Panther (m&r).
Marion County; Chesapeake (ndr), Curtisville (ndr), Fairmont (ndr),
Glady Creek (ndr). High Point (ndr), Kingmont (ndr), Midway Park
(grh), Mt. Harmony (ndr). Poplar Island (ndr). Marshall County;
Bannon (ndr), Moundsville (vs). Mason Comity; Point Pleasant (cbw).
Mercer County; Bluefield (m&r), Oakvale (m&r), Princeton (m&r), Spanish-
burg (m&r). Mineral County; Burlinton (gkm). New Creek (ndr). Mingo
Cowwi?/; Williamson (m&r). Monongalia County; Cheat Mt. (hhs). Coopers
Rock (ndr), Morgantown (hhs). Monroe County; Alderson (grh) (gkm),
Salt Sulphur Springs (ndr) (m&r), Sweetsprings (grh) (m&r). Union
(gkm), Waitesville (m&r). Willow Bend (m&r). Morgan County; Berkeley
Springs (ndr), Largent (ndr). Nicholas County; Craigsville (gkm), Sum-
mersville (afa) (gkm). Ohio County; Wheeling (cbw). Pendleton County;
Circleville (ndr), Franklin (ndr) (gkm), Judy Gap (gkm). Upper Tract
(ndr) (gkm). Pocahontas County; Dunmore Spring (gkm), Greenbank
(gkm), Hillsboro (gkm), Marlinton (gkm). Mill Point (gkm). Preston
County; Aurora (Heideman). Putnam County; Poca (gkm). Raleigh County;
Daniels (gkm) (m&r), Eccles (m&r). Flat Top Mt. (p&l). Shady Spring
(m&r). Randolph County; Cheat Bridge (ndr), Huttonsville (gkm). Sum-
mers County; Bellepoint (gkm). Riffle (gkm), Talcott (gkm). Taylor
County; Valley Falls (ndr). Tucker County; Parsons (mrs. gkm) (gkm).
Tyler County; Friendly (cbw). Upshur County; French Creek (mrs. gkm)
(gkm). Wayne County; Ceredo (cbw). Echo (m&r). Fort Gay (m&r),
Genoa (m&r). Webster County; Bolair (ndr) (gkm) (m&r). Wetzel County;
Proctor (vs). Silver Hill (ndr). Wirt County; (wjf). Wood County; Boaz
(cbw), Parkersburg (frw). Wyoming County; Baileysville (m&r), Pine-
ville (m&r).
Zonitoides nitidus (Mueller)
(Plate 13, figs. 8, 9)
Shell orbicularly depressed, moderately convex above and concave
below, thin, shining, uniform brownish horn-color, with delicate striae of
growth ; whorls 5 or more, convex, separated by a deeply impressed suture,
the outer one disproportionately large, somewhat declining as it approaches
the aperture, and obtusely angular at the periphery, beneath excavated
around a broad, crateriform umbilicus, in which the whorls are displayed
182 Annals of the Carnegie Museum vol. 31
to the apex; aperture oblique, lunate; peristome simple, its basal margin
arcuate. Greater diameter 73^, lesser 6 mm.; height 3^ mm.
Type Locality: Friedricksberg, Germany (Mueller, 1774).
Range: Canada and United States.
West Virginia Records
Barbour County; Philippi (mrs. gkm) (gkm). Boone County; Peytona
(ndr) (m&r), Racine (m&r), Seth (m&r). Braxton County; Gassaway
(gkm). Shady side (gkm). Cabell County; Barboursville (m&r), Ona (m&r),
Calhoun County; Grantsville (ndr). Clay County; Clay (gkm). Fayette
County; Clifftop (gkm). Grant County; Greenland Gap (ndr), Mount
Storm (ndr). Greenbrier County; Alderson (ndr). Organ Cave (ndr).
Renick Valley (ndr), Ronceverte (gkm) (m&r). Hampshire County;
Romney (gkm). Kanawha County; Charleston (gkm), Dunbar (gkm),
Nitro (gkm). South Charleston (ndr). Lincoln County; Branchland (m&r),
Sheridan (ndr), Sweetland (m&r). Logan County; Blair (m&r), Sharpies
(m&r). McDowell County; laeger (m&r). Panther (m&r). Marion County;
Curtisville (ndr), Fairmont (ndr), Glady Creek (ndr), High Point (ndr),
Mt. Harmony (ndr). Marshall County; Bannon (ndr). Mason County;
Leon (ndr). Mercer County; Princeton (m&r). Mineral County; Burling-
ton (gkm). Mingo County; Taylorsville (m&r), Williamson (m&r). Monon-
galia County; Coopers Rock (ndr), Smith town (ndr), Sturgisson (ndr),
Uffington (ndr). Monroe County; Alderson (gkm). Salt Sulphur Springs
(m&r), Sweetsprings (m&r). Union (gkm), Waitesville (m&r). Willow
Bend (m&r). Morgan County; Berkeley Springs (ndr) , Largent (ndr) (grh).
Nicholas County; Nettie (ndr), Richwood (cjg) (ndr), Summersville
(gkm). Pendleton County; Circleville (ndr), Franklin (ndr) (gkm), Judy
Gap (gkm), Upper Tract (ndr) (gkm). Pocahontas County; Dunmore
Spring (gkm), Greenbank (gkm), Hillsboro (gkm), Marlinton (gkm). Mill
Point (gkm). Preston County; Cascade (ndr). Putnam County; Poca (gkm).
Raleigh County; Eccles (m&r). Shady Spring (m&r). Randolph County;
Huttonsville (gkm). Summers County; Riffle (gkm), Talcott (gkm).
Tucker County; Parsons (mrs. gkm) (gkm). Upshur County; French Creek
(gkm). Wayne County; Fort Gay (m&r), Genoa (m&r). Webster County;
Bolair (ndr) (gkm). Wetzel County; Silver Hill (ndr). Wyoming County;
Baileysville (m&r), Oceana (m&r), Pineville (m&r).
Genus Striatura Morse
Shell small, translucent, with a steel gray tinge or pale green to greenish
horn-color, not shining; volutions 3 to 33/2j the outer whorl rapidly en-
1949
MacMillan: Land Snails of West Virginia
183
larging, aperture well rounded, very large, spire slightly elevated. Suture
distinct, and deeply channeled near the apex. Umbilicus small or broadly
open, abrupt, and exhibiting all the volutions. Periostraca minutely
marked with fine revolving lines and lines of increase, which reticulate the
whole surface.
Striatura exigua (Stimpson)
(Plate 13, fig. 10; Plate 14, fig. 1)
Shell broadly umbilicated, depressed, pellucid, greenish horn-color,
marked with delicate revolving lines, and distant longitudinal ribs obli-
quely decussating the incremental striae; spire scarcely elevated, apex
free from striae; whorls 3}^, convex, the last rounded, widely umbilicated
below; aperture oblique, transversely rounded, remote from the axis;
peristome simple, acute, its columellar extremity not reflected. Greater
diameter 23^ mm. ; height Y2 i^im.
Type Locality: Vicinity of Boston, Massachusetts (Stimpson, 1850).
Range: Quebec and Ontario; New England States, Michigan; New
York to Indiana, and West Virginia.
West Virginia Records
Pocahontas County; Greenbank (gkm). Tucker County; Parsons (gkm).
Webster County; Bolair (m&r).
Striatura ferrea Morse
(Plate 13, figs. 11, 12 13)
Shell umbilicated, depressed-globose, transparent, of a very light steel-
gray color, not shining, marked with very delicate incremental wrinkles
and microscopic revolving lines; spire slightly elevated; whorls 3, rounded,
the last rapidly enlarging, globose; aperture large, transversely sub-
circular; peristome simple, acute, its extremities not approaching, that of
the columellar scarcely subreflected. Greater diameter lY mm.; height
134 mm.
Type Locality: Maine (Morse, 1864).
Range: Quebec and Ontario; Maine to Michigan and Illinois, south to
Tennessee and North Carolina.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r). Braxton County; Gassaway (gkm). Fayette County; Cliff-
top (gkm). Hampshire County; Romney (gkm). Lewis County; Weston
184
Annals of the Carnegie Museum
VOL. 31
(gkm). McDowell County; Avondale (m&r). Monongalia County; Coopers
Rock (ndr). Nicholas County; Lockwood (gkm), Summersville (gkm).
Pendleton County; Franklin (gkm). Pocahontas County; Black Mt. (p&l),
Greenbank (gkm), Hillsboro (gkm), Marlinton (gkm). Raleigh County;
Flat Top Mt. (p&l). Shady Spring (m&r). Randolph County; Cheat Bridge
(ndr). Cheat Mt. (p&l), Helvetia (vs), Huttonsville (gkm). Tucker
County; Parsons (mrs. gkm) (gkm). Upshur County; French Creek (gkm).
Webster County; Bolair (m&r).
Remarks: This species resembles very much the young of Retinella
indentata, but the latter is perfectly smooth and exhibits only the faintest
revolving lines; Striatura ferrea also lacks the impressed radiating lines so
characteristic of indentata. The apex of ferrea is marked with distinct
revolving lines, the open umbilicus is constant, and is not half as large as
an adult indentata; finally, the dead color alone is sufficient to distinguish
it from the latter shell.
Striatura milium (Morse)
(Plate 14, fig. 2)
Shell widely umbilicated, depressed, transparent, shining, white, with a
greenish tinge, marked with distinct and regular striae of growth and
microscopic revolving lines, the latter more conspicuous below; spire but
slightly elevated ; whorls 3, rounded, rapidly increasing, the last planulate
above, widely umbilicated below; aperture very oblique, subcircular, re-
mote from the axis; peristome simple, acute, its terminations somewhat ap-
proached, that of the columellar not reflected. Greater diameter mm.;
height 3^ mm.
Type Locality: Mt. Independence, Westbrooke, Maine (Morse, 1859).
Range: Ontario to Manitoba; Maine to Minnesota, south to Missi-
ssippi and Florida.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (m&r), Racine (m&r). Braxton County; Frametown (gkm), Gassa-
way (gkm). Cabell County; Barboursville (m&r), Ona (m&r). Fayette County;
Clifftop (gkm). Grant County; Mount Storm (ndr). Greenbrier County;
Alderson (gkm). Organ Cave (ndr). Renick (grh). Renick Valley (ndr),
Ronceverte (gkm). Hampshire County; Romney (gkm). Jefferson County;
Bolivar (msb). Harper’s Ferry (jpem), Reedson (msb). Kanawha County;
Nitro (gkm). Tornado (gkm). Lewis County; Jackson’s Mill (gkm).
Lincoln County; Branchland (m&r), Myra (m&r). Logan County; Blair
1949
MacMillan: Land Snails of West Virginia
185
(m&r), Davin (ndr), Sharpies (m&r), McDowell County; Avondale (m&r),
laeger (m&r), Panther (m&r). Marion County; Glady Creek (ndr), High
Point (ndr). Mercer County; Bluefield (m&r), Oakvale (m&r), Princeton
(m&r). Mineral County; Burlington (gkm). Mingo County; Taylorsville
(m&r), Williamson (m&r). Monroe County; Sweetsprings (grh), Waites-
ville (m&r). Willow Bend (m&r). Morgan County; Berkeley Springs (ndr).
Nicholas County; Craigsville (gkm), Summersville (gkm). Pendleton
County; Franklin (ndr) (gkm), Judy Gap (gkm). Upper Tract (ndr).
Pocahontas County; Dunmore Spring (gkm), Greenbank (gkm). Marlinton
(gkm). Mill Point (gkm). Raleigh County; Shady Spring (m&r). Randolph
County; Helvetia (vs), Huttonsville (gkm). Summers County; Talcott
(gkm). Taylor County; Valley Falls (ndr). Tucker County; Parsons (gkm).
Wayne County; Echo (m&r), Fort Gay (m&r), Genoa (m&r). Webster
County; Bolair (m&r). Wyoming County; Oceana (m&r), Pineville (m&r).
Remarks: The peculiarities of this species are its diminutive size,
rapidly enlarging and well-rounded whorls, deep and regular striations,
which become obscure at the apex, and the microscopic lines running
parallel with the whorls.
Genus Ventridens Binney and Bland
Shell subperforate or umbilicated, orbicularly convex, diaphanous
horn-color; glossy, more or less wrinkled; whorls 5-7; aperture lunar, al-
most always furnished at its base with fold-like denticles not reaching the
margin; peristome simple, acute.
Ventridens snppressus suppressus (Say)
(Plate 14, fig. 3)
Shell globose, depressed, pale horn-color, polished, somewhat pellucid;
body whorl opaque, whitish near the aperture; volutions six, wrinkled,
spire convex; aperture sublunate, narrow beneath; within, a prominent
tooth near the base, distant from the margin; labrum simple; umbilicus
rather small, orbicular, profound; region of umbilicus indented. Greater
breadth more than one-fifth of an inch (5 mm.).
Type Locality: Germantown, Philadelphia, Pennsylvania (Say, 1829).
Range: Ontario; Maine to Wisconsin, south to Mississippi and Florida.
West Virginia Records
Boone County; Racine (m&r). Braxton County; Shadyside (gkm). Grant
County; Greenland Gap (ndr). Greenbrier County; Alderson (gkm), Renick
186
Annals of the Carnegie Museum
VOL. 31
(grh), Renick Valley (ndr), Ronceverte (gkm) (m&r). Hampshire County;
Capon Bridge (jpem), Romney (gkm). Jefferson County; Bardane (msb),
Bloomery (jpem) (msb), Bolivar (msb), Engle (msb), Gibsontown (msb),
Harper’s Ferry (jpem) (hap) (msb) (ejc) (Dr. E. W. Hubbard) (cc),
Jamestown (msb), Mechanicstown (msb), Meyerstown (msb), Millville
(msb), Morgan Grove (msb), Reedson (msb), Shenandoah Junction (msb),
Shepherdstown (msb). Kanawha County; Hudnall (cbw). McDowell
County; Pcvonddl^ (m&r). Panther (m&r). Mercer County; Bluefield (m&r).
Mineral County; Burlington (gkm). New Creek (ndr). Monroe County;
Sweetsprings (m&r), Waitesville (m&r). Morgan County; (p&i). Pendleton
County; Franklin (ndr) (gkm), Judy Gap (gkm), Seneca Rocks (ndr).
Upper Tract (ndr) (gkm). Pocahontas County; Dunmore Spring (gkm),
Greenbank (gkm), Hillsboro (gkm), Marlinton (gkm). Summers County;
Talcott (gkm). Tucker County; Parsons (gkm). Wayne County; Fort Gay
(m&r).
Remarks: With the exception of size and armature of the aperture, it
resembles somewhat V. ligera; but the capacity of the body whorl near
the aperture is much more remarkable. These characters, combined with
that of the teeth, will readily distinguish this species from others.
Ventridens suppressus virginicus (Vanatta)
(Plate 14, fig. 4)
The adult stage has a long nodule within the columellar lip and a rather
short obtuse lamella within the outer lip, more remote from the columellar
nodule than in the case with the outer tooth of suppressus. Other char-
acters as in V. suppressus. Altitude 3.4 mm.; diameter 6 mm.; slightly
over 6 whorls.
Type Locality: Endless Cavern Hills, near New Market, Shenandoah
County, Virginia (Vanatta, 1936).
Range: New Jersey to Ohio, south to Alabama and Georgia.
West Virginia Records
Braxton County; Gassaway (gkm). Grant County; Greenland Gap (ndr).
Greenbrier County; Alderson (gkm). Organ Cave (ndr), Renick Valley
(ndr), Ronceverte (gkm). Hampshire County; Junction (ndr), Romney
(gkm). Jefferson County; Aldridge (msb), Bloomery (jpem). Blue Ridge
(jpem), Gibsontown (msb). Harper’s Ferry (ndr), Jamestown (msb), Mt.
Mission (jpem), Rippon (msb). Lincoln County; Branchland (m&r),
Sheridan (ndr). McDowell County; Avondale (m&r), laeger (m&r), Pan-
1949
MacMillan: Land Snails of West Virginia
187
ther (m&r). Mineral County; Burlington (gkm), New Creek (ndr). Mingo
County; Williamson (m&r). Monroe County; Sweetsprings (m&r), Waites-
ville (m&r). Morgan County; Largent (ndr). Pendleton County; Franklin
(ndr) (gkm), Judy Gap (gkm), Seneca Rocks (ndr), Upper Tract (ndr)
(gkm). Pocahontas County; Dunmore Spring (gkm), Greenbank (gkm),
Hillsboro (gkm), Marlinton (gkm). Summers County; Talcott (gkm).
Remarks: In some individuals this two-toothed stage is seen in some-
what smaller shells, but the strong callous lining of the throat is usually
not well developed. At an earlier stage, diameter 4.3 mm., more or less, the
columellar tooth is conspicuously bifid, or in some shells trifid; there is a
long and high entering lamella within the outer lip with generally one or
two small laminae above it, and the callous lining is heavy. In old indi-
viduals of V. suppressus there is no tooth within the outer lip, only the
columellar nodule remains. The small laminae above the larger outer tooth
of the young stage disappear in suppressus at a much earlier stage than in
virginicus.
Ventridens suppressus divisidens Pilbsry
Shell subglobose, depressed, pale horn-colored, polished, somewhat
pellucid, moderately umbilicated ; body whorl opaque, whitish near aper-
ture, whorls 6-63^, wrinkled, spire convex; aperture sublunate, narrow
beneath; within, two-toothed, columellar one tubercular or somewhat
lengthened a short distance within base, outer one very shortly lamellar or
tubercular, remote from columella, and situated within periphery; lip
simple, acute, somewhat thickened within, extremities remote, joined by
a very thin callus deposit ; base slightly convex, indented ; umbilicus mode-
rate, deep, hardly exhibiting any inner volutions. Diameter 5. 4-6.4 mm. ;
height 3.5-4 mm.
Type Locality: Marlinton, Pocahontas County, West Virginia (Pilsbry,
1946).
Range: West Virginia records; Pendleton County; Franklin (gkm).
Pocahontas County; Marlinton (gkm).
Remarks: The teeth in V. s. divisidens are farther apart than in V.
suppressus and the basal or peripheral one is usually quite small. It dif-
fers from V. s. virginicus by the simple, not bifid, columellar tooth.
Ventridens gularis gularis (Say)
(Plate 14, fig. 5)
Shell subglobose, pale yellowish horn-color; polished; pellucid, beneath
the aperture whitish-yellow opaque; volutions six or seven, with promi-
188
Annals of the Carnegie Museum
VOL. 31
nent, somewhat regular wrinkles; spire convex, a little elevated; suture
moderate; labrum not reflected; throat far within upon the side of the
labrum, bifid ; teeth lamelliform, of which one is oblique and placed near
the middle, and the other less elongate, placed near the base; umbilicus
none. Breadth more than one-fourth of an inch (6 mm.).
Type Locality: Not designated (Say, 1821).
Range: Pennsylvania and Ohio, south to Alabama and Georgia.
West Virginia Records
Boone County; Peytona (m&r), Racine (m&r). Braxton County; Gassa-
way (gkm). Fayette County; Gauley Bridge (grh). Grant County; Mount
Storm (ndr). Hampshire County; Junction (ndr). Jackson County; (W. G.
Binney). Jefferson County; Bardane (msb). Logan County; Davin (ndr).
McDowell County; laeger (m&r). Panther (m&r). Mercer County; Blue-
field (m&r), Spanishburg (m&r). Mingo County; Williamson (m&r).
Nicholas County; Craigsville (gkm), Summersville (gkm). Pendleton
County; Franklin (ndr) (gkm), Judy Gap (gkm). Pocahontas County;
Mill Point (ndr). Randolph County; Cheat Bridge (gkm). Wyoming
County; Baileysville (m&r), Pineville (m&r).
Remarks: In general form it resembles V. ligera, but may be dis-
tinguished by the absence of the umbilicus and by the presence of teeth,
which are situated far within the aperture.
Ventridens gularis theloides (A. B. Brown)
(Plate 14, fig. 6)
Shell glossy, yellow, perforate, with moderately raised, dome-shaped
spire, composed of 73^-8 narrow, closely coiled whorls; the last hardly
regular at the periphery in adult shells; rather strongly striate above,
nearly smooth beneath, with faint traces of spiral striae near the um-
bilicus, where the base is rather conspicuously excavated. Aperture
somewhat triangidar, the sloping basal lip being straight. The peristome is
acute, strengthened within by a rather wide, low callus rim. Adult shells
are without teeth or lamince. Young shells armed with two strong lamellae
within the aperture, the summit of the outer one curving towards the
inner. Altitude 43/^-5, diameter mm.
Type Locality: North Carolina (Brown, 1902).
Range: West Virginia; North Carolina; and Tennessee. West Virginia
Record: Wyoming County; Oceana (m&r).
Remarks: It differs from V. lawae in the much narrower umbilicus;
from V. gularis it differs in being toothless in the adult stage, and with a
more excavated base and straighter basal lip.
1949
MacMillan: Land Snails of West Virginia
189
Ventridens gularis decussatus (Pilsbry and Vanatta)
(Plate 14, fig. 7)
Shell somewhat dull, brownish, narrowly umhilicate, with moderate or
high, dome-shaped spire, composed of 8 flat whorls, the last angular at
the periphery. Closely, sharply and strongly striate above, less so beneath,
where there are usually traces of spiral strice near the periphery. Aper-
ture lunate, peristome thin and acute, armed within with a blunt tooth
on the middle of the columella, and a high, short, erect lamella, within the
outer basal margin. Altitude 5.5 diameter 7.8 mm.
Type Locality: Mt. Mitchell, North Carolina (Pilsbry and Vanatta
1902).
Range: West Virginia and North Carolina. West Virginia Record:
Boone County; Racine (m&r).
Remarks: It differs from V. gularis and the racial forms of that species
in the strong sculpture, comparatively dull surface, the decussated and
acute carination of the young shells, and the short, high, erect lamella of
the adult stage. Compared with V. collisellus, the variety decussatus is
seen to differ in the closer and flatter whorls of the spire, flattened instead
of swollen base, and the carinated periphery. Young shells resemble V.
intertextus in sculpture.
The young shells are biconvex, depressed, acutely angular or carinate,
more strongly striate beneath than in the adults, and with the striae on the
outer half of the base decussated by many impressed spirals. The internal
lamellae are long.
Ventridens collisella (Pilsbry)
(Plate 14, fig. 8)
Shell rather solid, minutely perforate, above elevated and somewhat
dome-shaped, below rather flattened, the periphery rounded. Surface
glossy, especially beneath, the base being radially finely wrinkled, and
with faint traces of spiral striations in the slightly excavated umbilical
region; upper surface sculptured with irregular, arcuate wrinkle-riblets in
the direction of growth-lines, and stronger towards the base. Whorls 73^,
slightly convex, separated by very shallow sutures, which, under the lens,
seemed margined below by the parietal transparence of the shell. Aper-
ture mainly basal, lunate, with a lining of white callus a short distance
within, heavier and bearing a small tooth on the columellar slope, and a
rather short white lamella towards the outer part of the base. Lip edge
thin and acute, suddenly expanded at the columellar insertion, half
covering the umbilical perforation. Altitude 5^-7, diameter 8>^ mm.
Type Locality: Knoxville, Knox County, Tennessee (Pilsbry, 1896).
190 Annals of the Carnegie Museum vol. 31
Range: West Virginia; Virginia; North Carolina; Tennessee; and Ala-
bama. West Virginia Record : Mercer County; Spanishburg (m&r) .
Remarks: It is most nearly allied to V. tiger a and V, cerinoideus,
being smaller than the former, with different apertural armature, and
larger, duller, and more elevated than the former.
Ventridens demissus (A. Binney)
(Plate 14, fig. 9)
Shell depressed-convex; epidermis yellowish horn-color, shining; whorls
six, with minute lines of growth; spire obtuse; suture impressed; body-
whorl expanded very little towards the aperture; aperture transverse, not
large, a white, testaceous deposit within ; lip thin, acute ; base rather flat,
smooth; umbilicus very small; umbilical region a little impressed. Greatest
transverse diameter more than three-eighths of an inch (934 mm.).
Type Locality: Pennsylvania (Binney, 1843).
Range: Pennsylvania to Kansas, south to Texas and Florida.
West Virginia Records
Barbour County; Philippi (gkm). Boone County; Peytona (m&r), Racine
(m&r), Seth (m&r). Cabell County; Barboursville (m&r). Clay County;
Clay (gew). Fayette County; Clifftop (gkm). Grant County; Mount Storm
(ndr). Greenbrier County; Renick Valley (ndr), Ronceverte (gkm) (m&r).
Kanawha County; Alum Creek (ndr), Hudnall (cbw). South Charleston
(ndr). Lewis County; Jackson’s Mill (gkm). Lincoln County; Myra (m&r),
Sheridan (ndr), Sweetland (m&r). Logan County; Blair (ndr) (m&r),
Davin (ndr). Sharpies (m&r), Slagle (cg), Stone Branch (ndr). McDowell
County; Avondale (m&r), laeger (m&r), Panther (m&r). Marion County;
Curtisville (ndr). High Point (ndr). Mercer County; Bluefield (m&r),
Oakvale (m&r), Spanishburg (m&r). Mineral County; Ridgeley (aeo).
Mingo County; Taylorsville (m&r), Williamson (m&r). Monroe County;
Alderson (gkm). Salt Sulphur Springs (m&r), Sweetsprings (m&r). Nicholas
County; Lockwood (gkm), Richwood (ndr). Pendleton County; Franklin
(gkm), Judy Gap (gkm), Upper Tract (ndr). Pocahontas County; Hills-
boro (gkm). Mill Point (ndr). Preston County; Manheim (jha). Raleigh
Shady Spring (m&r). Randolph Huttonsville (mgn) (gkm).
Summers County; Riffle (gkm). Tucker County; Parsons (gkm). Upshur
County; F rench Creek (gkm) . Wayne County; Echo (m&r) , F ort Gay (m&r) ,
Genoa (m&r). Webster County; Bolair (m&r). Wirt County; (wjf). Wyom-
ing County; Baileysville (m&r), Oceana (m&r), Pineville (m&r).
1949
MacMillan: Land Snails of West Virginia
191
Remarks: The character of the surface and striae is much the same as in
V. ligera; and the base is nearly the same. It resembles V. suppressus
externally, in everything except in size and in being less coarsely striate.
Ventridens ligera ligera (Say)
(Plate 14, fig. 10)
Shell subglobose, pale yellowish horn-color, polished ; body whorl
pellucid, yellowish white, opaque beneath near the aperture; volutions
rather more than six, all except the apical one, wrinkled across; spire
convex, a little elevated; umbilicus very small; suture not deeply im-
pressed; labrum not reflected. Greater length, three-tenths, oblique length
less than nine-twentieths, transverse diameter less than eleven-twentieths
{1)4, X 1134 X 13^ mm.).
Type Locality: Missouri (Say, 1821).
Range: Ontario; Maine to Minnesota, south to Texas and Georgia.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gew) (gkm). Boone
County; Peytona (ndr) (m&r), Racine (m&r). Braxton County; Frametown
(gkm), Gassaway (gkm), Shadyside (gkm). Brooke County; Bethany (mgn)
(cBw). Cabell County; Barboursville (gkm) (m&r), Lesage (m&r), Milton
(cBw). Calhoun County; Grantsville (ndr). Grant County; Mount Storm
(ndr). Greenbrier County; Alderson (ndr) (gkm). Renick (grh). Renick
Valley (ndr), Ronceverte (m&r). Hampshire County; Romney (gkm).
Jefferson County; Bloomery (msb) (jpem), Charles Town (msb), Engle
(msb). Hall town (msb). Harper’s Ferry (msb) (jpem), Kabletown (msb).
Keystone (msb), Loudoun Heights (jpem), Meyerstown (msb), Middle-
way (msb), Morgan Grove (msb), Ranson (msb), Reedson (msb), Shenan-
doah City (msb), Shepherdstown (msb). Summit Point (msb). Kanawha
County; Alum Creek (ndr). Charleston (gkm), Hudnall (cbw), St. Albans
(um). Tornado (gkm). Lewis County; Jackson’s Mill (mrs. gkm) (gkm).
Lincoln County; Myra (m&r), Sweetland (m&r). Logan County; Blair
(ndr) (m&r), Davin (ndr). Sharpies (m&r). McDowell County; Avondale
(m&r). Panther (m&r). Marion County; Fairmont (ndr) (grh). High
Point (ndr), Mt. Harmony (ndr). Smith ville (cbw). Marshall County;
Kent (ndr). Mason County; Leon (ndr). Mercer County; Bluefield (m&r),
Princeton (m&r), Spanishburg (m&r). Speedway (m&r). Mineral County;
Keyser (ihm), Ridgeley (aeo). Mingo County; Taylorsville (m&r), William-
son (m&r). Monongalia County; Coopers Rock (mgn) (ndr), Morgan-
192
Annals of the Carnegie Museum
VOL. 31
town (hhs), Smithtown (ndr), Sturgisson (ndr). Monroe County; Alder-
son (grh) (gkm), Salt Sulphur Springs (m&r), Waitesville (m&r). Morgan
County; Largent (ndr). Nicholas County; Belva (ndr), Richwood (ndr),
Snake Den Mt. (ndr). Ohio County; Clinton (mgn). Wheeling (mgn) (ndr)
(grh) (cbw). Pendleton County; Circleville (ndr), Franklin (gkm), Onega
(mgn). Upper Tract (ndr) (gkm). Pleasants County; St. Mary’s (cbw).
Pocahontas County; Dunmore Spring (gkm), Marlinton (mgn) (gkm).
Mill Point (mgn) (grh) (gkm). Preston County; Cheat Mt. (grh), Man-
heim (jha). Putnam County; Poca (gkm). Raleigh County; Daniels (m&r).
Shady Spring (m&r). Randolph County; Helvetia (vs), Huttonsville (gkm).
Summers County; Riffle (gkm), Talcott (gkm). Taylor County; Grafton
(frw), Thornton (grh). Tucker County; Parsons (mrs. gkm) (gkm). Tyler
County; Friendly (cbw). Upshur County; French Creek (gkm). Wayne
County; Falls of Big Sandy River (Sanderson Smith) (American Museum
of Natural History). Webster County; Bolair (gkm) (m&r). Wetzel County;
New Martinsville (cbw). Wyoming County; Baileysville (m&r), Oceana
(m&r), Pineville (m&r).
Ventridens ligera stonei (Pilsbry)
(Plate 14, fig. 12)
Shell perforate, orbicularly convex; epidermis yellowish horn-color,
shining; whorls 7, finely striated transversely, smooth below; suture not
much impressed; aperture semilunate, rounded, peristome thin, acute;
base and sides of outer whorl, within aperture, thickened and white;
perforation moderate, umbilical region concave, broadly excavated.
Greater diameter 16, lesser 14 mm.; height 10 mm.
Type Locality: New Castle County, Delaware (Pilsbry, 1889).
Range: New Jersey; Delaware; Maryland; and West Virginia.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County; Pey-
tona (m&r). Cabell County; Lesage {yi&U) . Grant County ; Greenland Gap
(ndr). Mount Storm (ndr). Greenbrier County; Renick Valley (ndr), Ron-
ceverte (m&r). Hampshire County; Romney (gkm). Jefferson County; Har-
per’s Ferry (ndr). Kanawha County; Tornado (gkm). Logan County ;Stor\^
Branch (ndr). Marion County; Fairmont (ndr). Mercer County; Oakvale
(m&r), Princeton (m&r), Spanishburg (m&r). Mingo County; Williamson
(m&r). Monongalia County; Coopers Rock (mgn) (ndr). Nicholas County;
Richwood (ndr). Pendleton County; Circleville (ndr), Upper Tract (ndr)
1949
MacMillan: Land Snails of West Virginia
193
(mgn). Pocahontas County; Hillsboro (gkm), Marlinton (gkm), Mill Point
(ndr) {gkm). Preston County; Cascade (ndr). Randolph County; Cheat
Bridge (ndr), Helvetia (vs), Huttonsville (mgn) (ndr) (gkm). Summers
County; Riffle (gkm), Talcott (gkm). Tucker County; Parsons (mrs. gkm)
(gkm). Upshur County; French Creek (gkm). Webster County; Bolair
(gkm). Wyoming County; Oceana (m&r).
Remarks: This subspecies has a more concave and broadly excavated
base than V. ligera, and the umbilicus is much wider.
Ventridens intertextus (A. Binney)
(Plate 14, fig. 11)
Shell sub-pyramidal; epidermis yellowish horn-color; whorls six to
seven, with numerous fine, oblique striae, and very minute, spiral striae;
intersecting each other; outer whorl with a narrow, light-colored band,
and an ill-defined, brownish band below it; aperture rounded, a little trans-
verse; lip thin, somewhat thickened within by a deposition of testaceous
matter, slightly reflected at its junction with the base of the shell; umbili-
cus small, sometimes nearly obsolete; base whiter than upper surface.
Greatest transverse diameter about three-fourths of an inch (18^ mm.).
Type Locality: North Carolina (A. Binney, 1840).
Range: Ontario; Maine to Indiana, south to Louisiana and Florida.
West Virginia Records
Barbour County; Nestorville (gkm), Philippi (gkm). Boo?ie County;
Peytona (m&r). Cabell County; Ona (m&r). Doddridge County; West Union
(frw). Grant County; Mount Storm (ndr). Greenbrier County; Organ Cave
(ndr). Kanawha County; Hudnall (cbw). Logan County; Blair (m&r),
Davin (ndr). Sharpies (m&r). Stone Branch (ndr). McDowell County;
laeger (m&r). Marion County; East Fairmont (ndr). High Point (ndr).
Mingo County ;T2cy\orsY\VL& (m&r), Williamson (m&r). Monongalia County;
Cheat Mt. (hhs). Coopers Rock (cbw), Sturgisson (ndr). Nicholas
County; Craigsville (gkm), Lockwood (gkm), Summersville (gkm). Ohio
County; Wheeling (mgn) (cbw). Preston County; Cascade (ndr), Manheim
(jha). Randolph County; Helvetia (vs). Taylor County; Valley Falls (ndr).
Tucker County; Parsons (gkm). Upshur County; French Creek (mrs. gkm)
(gkm). Wayne County; Fort Gay (m&r), Genoa (m&r). Wyoming County;
Oceana (m&r).
Remarks: This shell resembles some varieties of V. ligera. However,
the spire is less high in the shell of the same size, has a smaller number of
194
Annals of the Carnegie Museum
VOL. 31
whorls, and is more pyramidal in shape. The diameter, in full-grown
specimens, is greater, and the base is flatter. The epidermis is darker and
less shining, the shell is thicker and less pellucid, the deposit of testaceous
matter within the aperture is less. The size of the umbilicus and the shape
of the aperture are the same in both. The principal distinction consists
in the spiral lines, which revolve on the whorls, intersecting the striae of
growth, but so minute as hardly to be perceptible to the naked eye. The
whitish, narrow band, shaded below with rufous, apparent on the outer,
and sometimes on the second whorl, generally aids in identification.
Young specimens are much more depressed than those of V. ligera and
are sometimes distinctly carinated or angulated at the periphery.
Additional localities for the Families
Carychiid^ and Pupillid^
Under the species of these families immediately following this heading
are listed additional localities which are supplementary to those locali-
ties previously listed for these species in papers by Dr. S. T. Brooks and
the present author which appeared in the Annals of the Carnegie Museum ,
vol. XXV, 1937, pp. 155-161 {Carychiidce) and vol. XXVII, 1938, pp.
63-85 {Pupillidce) .
Family Carychiid^
Carychium exile H. C. Lea
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (ndr) (m&r), Seth (m&r). Cabell County; Barboursville (m&r),
Lesage (m&r), Ona (m&r). Fayette County; Cliff top (gkm). Grant County;
Greenland Gap (ndr). Greenbrier County; Alderson (grh)) (gkm). Renick
(grh), Renick Valley (ndr). Jefferson County; Bolivar (msb), Reedson
(msb). Kanawha County; Hudnall (cBw), Institute (gkm). Lewis County;
Jackson’s Mill (gkm). Lincoln County; Branchland (m&r), Myra (m&r),
Sweetland (m&r). Logan County; Blair (m&r), Davin (ndr). Sharpies
(m&r). Stone Branch (ndr). McDowell County; Avondale (m&r). Marion
County; Fairmont (ndr), Glady Creek (ndr). High Point (ndr), King-
mont (grh). Marshall County; Bannon (ndr). Mercer County; Bluefield
(m&r), Oakvale (m&r), Princeton (m&r). Speedway (m&r). Mingo County;
Williamson (m&r). Monroe County; Alderson (gkm). Salt Sulphur Springs
(m&r), Sweetsprings (grh) (m&r), Waitesville (m&r). Willow Bend (m&r).
1949
MacMillan: Land Snails of West Virginia
195
Morgan County; Berkeley Springs (ndr). Nicholas County; Summersville
(gkm). Pendleton County; Circleville (ndr), Franklin (ndr). Pocahontas
County; Mill Point (grh) (gkm). Randolph County; Cheat Bridge (gkm).
Tucker County; Parsons (mrs. gkm) (gkm). Upshur County; French Creek
(gkm). Wayne County; Echo (m&r), Fort Gay (m&r), Genoa (m&r).
Webster County; Bolair (ndr) (m&r). Wyoming County; Oceana (m&r),
Pineville (m&r).
Carychium exiguum (Say)
Cabell County; Ona (m&r). Grant County; Greenland Gap (ndr), Mount
Storm (ndr). Greenbrier County; North Caldwell (grh). Renick Valley
(ndr). Harrison County; Bristol (ndr). Jefferson County; Leetown (msb).
Logan County; Stone Branch (ndr). McDowell County; Avondale (m&r),
laeger (m&r). Panther (m&r). Mercer County; Spanishburg (m&r). Mon-
roe County; Salt Sulphur Springs (ndr). Ohio County; Wheeling (ndr).
Pendleton County; Circleville (ndr). Pocahontas County; Marlinton (gkm).
Preston County; Cascade (ndr). Wayne County; Fort Gay (m&r). Webster
County; Bolair (ndr) . Wetzel County; Silver Hill (ndr) . Wyoming County;
Baileysville (m&r).
Carychium nannodes Clapp
Boone County; Peytona (m&r), Seth (m&r). Cabell County; Lesage
(m&r). Lincoln County; Branchland (m&r). Logan County; Davin (ndr).
McDowell County; Avondale (m&r). Mercer County; Bluefield (m&r),
Oakvale (m&r). Speedway (m&r). Monroe County; Waitesville (m&r).
Wayne County; Fort Gay (m&r). Webster County; Bolair (ndr).
Family Pupillid^ Turton
Genus Gastrocopta Wollaston
Gastrocopta armifera (Say)
Berkeley County; Hedgesville (ndr). Cabell County; Barboursville (gkm),
Lesage (m&r). Grant County; Greenland Gap (ndr). Greenbrier County;
Renick Valley (ndr), Ronceverte (gkm) (m&r). Jefferson County; Bakerton
(msb), Charles Town (msb). Harper’s Ferry (ndr) (gwt) (ejc), Pipers-
town (msb), Shenandoah City (msb), Shenandoah Junction (hap). Marion
County; East Fairmont (ndr), Fairmont (ndr). High Point (ndr). Rives-
ville (ndr). Mason County; Leon (ndr). Mercer County; Spanishburg
196
Annals of the Carnegie Museum
VOL. 31
(m&r). Mineral County; New Creek (ndr). Monongalia County; Coopers
Rock (cBw). Monroe County; Gap Mills (m&r), Salt Sulphur Springs
(ndr) (m&r), Sweetsprings (m&r), Waitesville (m&r). Willow Bend (m&r).
Ohio County; Wheeling (cbw). Pendleton County; Circleville (ndr),
Franklin (ndr). Upper Tract (ndr). Wayne County; Echo (m&r).
Gastrocopta contracta contracta (Say)
Barbour County; Nestorville (gkm), Philippi (gkm). Boone County;
Peytona (gb) (m&r), Racine (m&r), Seth (m&r). Cabell County; Barbours-
ville (ndr) (gkm) (m&r), Lesage (m&r), Ona (m&r). Calhoun County;
Grantsville (ndr). Clay County; Clay (gkm). Fayette County; Cliff top
(gkm). Grant County; Geenland Gap (ndr). Mount Storm (ndr). Green-
brier County; Renick Valley (ndr), Ronceverte (gkm) (m&r). Jefferson
County; Bolivar (msb), Harper’s Ferry (ndr) (gwt) (ejc) (msb), Millville
{msb) . Kanawha County; Alum Creek (ndr), Hudnall (cbw). Lewis County;
Weston (gkm). Lincoln County; Branchland (m&r), Myra (m&r), Sweet-
land (m&r). Logan County; Blair (m&r), Davin (ndr). Stone Branch (ndr).
McDowell County; laeger (m&r). Panther (m&r). Marion County; Curtis-
ville (ndr), Fairmont (ndr), Glady Creek (ndr). High Point (ndr).
Mason County; Leon (ndr). Point Pleasant (ndr). Mercer County; Blue-
field (m&r), Oakvale (m&r), Princeton (m&r). Speedway (m&r). Mineral
County; New Creek (ndr). Mingo County; Taylorsville (m&r), Williamson
(m&r). Monongalia County; Sturgisson (ndr), Uffington (ndr). Monroe
County; Gap Mills (m&r), Salt Sulphur Springs (ndr) (m&r). Sweet-
springs (m&r), Waitesville (m&r), Willow Bend (m&r). Morgan County;
Berkeley Springs (ndr), Largent (ndr). Ohio County; Wheeling (cbw).
Pendleton County; Circleville (ndr), Franklin (ndr). Upper Tract (ndr).
Wayne County; Echo (m&r). Webster County; Bolair (ndr). Wyoming
County; Baileysville (m&r), Pineville (m&r).
Gastrocopta contracta climeana (Vanatta)
(Plate 15, fig. 1)
Shell rimate, ovate-conic, tapering from the last whorl to the obtuse
apex, bluish-milky or spermaceti-colored, imperfectly transparent, glossy,
marked with fine growth lines. Whorls very convex, the last half of
the last whorl straightened, pinched at the base, impressed over the lower
palatal plica, and on both sides of a low rounded ridge which stands a
short distance behind the peristome. Aperture rounded, triangular, al-
1949 MacMillan: Land Snails of West Virginia 197
most closed by large teeth. Angulo-parietal tooth joining lip, angularly
bent to the right near the middle, L-shaped. Columellar lamella large,
thin, very deeply placed, subvertical, the upper end curving forward. A
subvertical callous stands in front of it, near the margin. Palatal plicae
two, connected by a low callous, the lower one obtuse, transverse, more
deeply placed and larger than the tuberculiform upper plica. Peristome
thin, well expanded, continuous. Altitude 2.29, diameter 1.43 mm.
Type Locality: Anderson Landing, on Sunflower River, near Confluence
with the Yazoo, Sharkey County, Mississippi (Vanatta, 1911).
Range: West Virginia; Alabama to Texas, north to Arkansas. West
Virginia Record : Greenbrier County; Alderson (ndr) .
Remarks: The shell is similar to contracta, but the parietal tooth lacks
the inner continuation, being L-shaped.
Gastrocopta holzingeri (Sterki)
Greenbrier County; Renick Valley (ndr). Pendleton County; Franklin
(ndr).
Gastrocopta pentodon (Say)
Barbour County; Nestorville (gkm). Boone County; Peytona (gb) (m&r),
Racine (m&r), Seth (m&r). Cabell County; Barboursville (m&r), Lesage
(m&r), Ona (m&r). Calhoun County; Grantsville (ndr). Clay County; Clay
(gkm). Fayette County; Cliff top (gkm). Grant County; Greenland Gap
(ndr), Mount Storm (ndr). Greenbrier County; Renick Valley (ndr),
Ronceverte (gkm) (m&r). Jefferson County; Gibsontown (msb). Harper’s
Ferry (ndr) (msb). Keystone (msb), Meyerstown (msb), Millville (msb).
Kanawha County; Alum Creek (ndr), Hudnall (cbw). Lewis County;
Jackson’s Mill (gkm). Lincoln County; Branchland (m&r), Myra (m&r),
Sweetland (m&r). Logan County; Blair (m&r), Davin (ndr), Sharpies
(m&r), Stone Branch (ndr). Marion County; Fairmont (ndr), Glady
Creek (ndr), High Point (ndr). Mason County; Leon (ndr). Point
Pleasant (ndr). Mercer County; Bluefield (m&r), Oakvale (m&r). Speed-
way (m&r). Mingo Taylorsville (m&r), Williamson (m&r). Monon-
galia County; Smithtown (ndr). Monroe County; Salt Sulphur Springs
(ndr) (m&r), Sweetsprings (m&r), Waitesville (m&r). Nicholas County;
Richwood (ndr). Snake Den Mt. (ndr). Pendleton County; Circleville
(ndr), Franklin (ndr). Upper Tract (ndr). Pocahontas County; Mill
Point (gkm). Raleigh County; Daniels (m&r). Shady Spring (m&r). Ran-
dolph County; Helvetia (vs), Huttonsville (gkm). Summers County; Tal-
cott (gkm). Tucker County; Parsons (gkm). Wayne County; Fort Gay
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Annals of the Carnegie Museum
VOL. 31
(m&r), Genoa (m&r). Webster County; Bolair (ndr) (m&r). Wetzel County;
Proctor (vs). Wyoming County; Pineville (m&r).
Gastrocopta tappaniana (C. B. Adams)
Boone County; Peytona (m&r). Cabell County; Barboursville (gkm).
Lincoln County; Myra (m&r). Logan County; Blair (m&r), Sharpies (m&r).
Mingo County; Taylorsville (m&r), Williamson (m&r). Monroe County;
Salt Sulphur Springs (m&r). Union (gkm). Willow Bend (m&r). Pendleton
County; Franklin (ndr). Raleigh County; Daniels (m&r). Taylor County;
Valley Falls (ndr).
Gastrocopta corticaria (Say)
Boone County; Racine (m&r), Seth (m&r). Grant County; Greenland Gap
(ndr). Greenbrier County; Renick Valley (ndr), Ronceverte (m&r). Logan
County; Sharpies (m&r). McDowell County; Panther (m&r). Marion
County; High Point (ndr). Mason County; Leon (ndr). Monongalia
County; Uffington (ndr). Monroe County; Salt Sulphur Springs (m&r,)
Waitesville (m&r). Willow Bend (m&r). Morgan County; Berkeley Springs
(ndr). Pendleton County; Franklin (ndr). Taylor County; Valley Falls
(ndr).
Gastrocopta procera (Gould)
Greenbrier County; Renick Valley (ndr). Jefferson County; Aldridge
(msb), Bardane (msb), Bloomery (msb), Bolivar (msb), Kabletown (msb),
Kearneysville (msb). Keystone (msb), Leetown (msb), Mechanicstown
(msb), Meyerstown (msb). Middleway (msb), Millville (msb), Morgan
Grove (msb), Rippon (msb). Summit Point (ndr).
Genus Vertigo Mueller
Vertigo clappi Brooks and Hunt
Greenbrier County; Renick Valley (ndr). Marion County; Fairmont
(ndr).
Vertigo ovata Say
Jefferson County; Meyerstown (msb). Logan County; Sharpies (m&r).
Marion County; Mt. Harmony (ndr). Mingo County; Williamson (m&r).
Monroe County; Salt Sulphur Springs (m&r). Willow Bend (m&r). Poca-
hontas County; Marlinton (gkm).
1949
MacMillan: Land Snails of West Virginia
199
Vertigo gouldii (A. Binney)
Barbour County; Nestorville (gkm). Boone County; Racine (m&r). Cabell
County; Barboursville (m&r). Calhoun County; Grantsville (ndr). Clay
County; Clay (gkm). Grant County; Greenland Gap (ndr). Greenbrier
County; Renick Valley (ndr), Ronceverte (gkm). Jefferson County;
Harper’s Ferry (msb), Meyerstown (msb). Kanawha County; Alum Creek
(ndr). Lincoln County; Myra {m&r). Logan County; Blair (m&r). McDowell
County; Avondale (m&r). Marion County; Fairmont (ndr), Glady Creek
(ndr). Mason County; Leon (ndr). Mercer County; Oakvale (m&r),
Princeton (m&r). Mineral County ; BmXmgton (gkm). New Creek (ndr).
Monongalia County; Sturgisson (ndr), Uffington (ndr). Monroe County;
Alderson (gkm). Gap Mills (m&r), Sweetsprings (m&r), Waitesville (m&r),
Willow Bend (m&r). Morgan County; Largent (ndr). Pendleton County;
Franklin (ndr). Randolph County; Cheat Bridge (gkm). Webster County;
Bolair (ndr) (m&r). Wyoming County; Baileysville (m&r).
Vertigo tridentata Wolf
Jefferson County; Harper’s Ferry (ndr), Shepherdstown (msb), Skeeters-
ville {msb) . Kanawha County; Tornado {gkm) . Lincoln County; Branchland
(m&r). Marshall County; Bannon (ndr). Monroe County; Salt Sulphur
Springs (ndr), Sweetsprings (m&r), Waitesville (m&r). Pendleton County;
Judy Gap (gkm).
Vertigo ventricosa (Morse)
(Plate 15, fig. 2)
Shell umbilicate, ovate, conic, smooth, polished; apex obtuse; suture
deep; whorls four, convex. Aperture, semicircular, with five teeth, one
prominent on the parietal margin, two smaller on the columellar margin,
and two prominent within, contracting the aperture at the base; peris-
tome widely reflected, the right margin flexuous, within thickened and
colored. Length .07 inch, breadth .45 inch. (1.75 x 1.1 mm.).
Type Locality: Not designated (Morse, 1865).
Range: Prince Edward Island; Quebec; Maine to Michigan and Illinois;
and West Virginia.
West Virginia Records
Monroe County; Willow Bend (m&r). Raleigh County; Daniels (m&r).
Remarks: V. ventricosa differs from V. ovata by the constantly smaller
200
Annals of the Carnegie Museum
VOL. 31
size, absence of an angular lamella, and smaller number of teeth. It is of
an auburn color, somewhat transparent, glossy, with only a trace of
striation. Under a high power lens it shows microscopic punctation or
granulation. The basal fold is usually quite small, though sometimes it is
well developed.
Vertigo pygmaea (Draparnaud)
(Plate 15, fig. 3)
The shell is cylindric-oval, sunburn or chestnut-brown, glossy, having
only weak traces of striation, but the surface appears densely weakly-
pitted microscopically. The whorls are moderately convex, the last having
a strong rounded crest a short distance behind the peristome, separated
from it by a concavity, and somewhat paler colored than the rest of the
shell. The parietal lamella is strong but rather short, median. Columellar
lamella deeply placed, short, ascending inwardly. Both palatal folds are
strong, the lower one longer, as usual. They stand on a strong callus. The
basal fold is very small, rarely absent. There is often a low suprapalatal
fold. The peristome is well expanded, somewhat reflected, colored like the
shell. The outer lip is slightly incurved. Length 2, diameter 1.1 mm.; 5
whorls.
Type Locality: France (Draparnaud, 1801).
Range: Maine to Virginia, west to Ohio and Michigan.
West Virginia Records
Mineral County; New Creek (ndr). Pendleton County; Franklin (ndr).
Remarks: The strong, continuous crest behind the well expanded lip
and the absence of a distinctly defined upper arc or sinulus of the outer
lip are its more prominent features. V. gouldii is a paler, much more
sharply striate shell with weaker crest and distinct sinulus. V. ventricosa
has a weaker and interrupted crest.
Vertigo bollesiana (Morse)
(Plate 15, figs. 4, 5)
Shell minutely perforate, cylindrical ovate, delicately striated, sub-
translucent; apex obtuse; suture well defined; whorls four, subconvex;
aperture suborbicular, somewhat flattened on its outer edge; with five
teeth, one prominent and rather curved on the parietal margin, two similar
in form, the lower one smaller, on the columellar margin, and two slightly
elevated lamelliform teeth within and at the base, peristome subreflected
and thickened. Length .065 inch.; breadth .035 inch (1.6 x .9 mm.).
Type Locality: Orono, Maine (Morse, 1865).
Range: Maine to Virginia, west to Indiana and Michigan.
1949
MacMillan: Land Snails of West Virginia
201
West Virginia Records
Braxton County; 4 miles south of the Lewis County Line (ndr). Nicholas
County; Summersville (gkm). Pendleton County; Franklin (gkm).
Remarks: There is an extremely small crest close behind the lip, and
a rather large oblique depression over the palatal folds. The basal fold is
subcolumellar in position, and rarely is it absent. It is smaller, lighter
colored and more transparent and delicate than V. gouldii and less dis-
tinctly striated. The teeth are smaller, especially the palatals.
Vertigo parvula Sterki
(Plate 15, fig. 6)
The shell is minute, subcylindric, tapering very little upwards, the
summit obtuse; thin, subtransparent, slightly yellowish, smooth and
glossy, becoming finely striate behind the outer lip. Whorls are moderately
convex, the last whorl well rounded, slightly impressed behind the pro-
jection of the outer lip. The aperture is somewhat triangular, with three
teeth: parietal lamella rather short and high; lower palatal fold rather
high in front, rapidly becoming lower as it recedes, penetrating to the
dorsal side. Peristome very little everted, slightly thickened, and having a
distinct callus ridge within. The outer lip projects forward and is slightly
bent inward above the middle. Length 1.55, diameter 0.85 mm.; barely
5 whorls.
Type Locality: Summit County; Ohio (Sterki, 1890).
Range: Ohio; West Virginia; North Carolina.
West Virginia Records
Boone County; Peytona (m&r). Grant County; Greenland Gap (ndr).
Greenbrier County; Alderson (gkm). Renick Valley (ndr). Logan County;
Davin (ndr). McDowell County; Avondale (m&r). Monroe County; Aider-
son (gkm). Pendleton County; Franklin (ndr) (gkm), Judy Gap (gkm).
Summers County; Talcott (gkm).
Remarks: It is one of the rarest species, known by the small size,
cylindric shape, and three well-developed teeth, the lower palatal being
quite long and not marked externally by an impression. The auricle or
point of the outer lip projects well forward but is not much bent inward.
It is much smaller than V. tridentata, and more cylindric and lighter
colored than V. perryi, with relatively larger teeth.
The species Vertigo oscariana Sterki recorded as occuring in West
Virginia has been misidentified ; it is V. parvula and all records for V.
oscariana should be placed under this species.
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Vertigo milium (Gould)
Grant County; Greenland Gap (ndr). Greenbrier County; Ronceverte
(m&r). Jefferson County; Bolivar (msb), Harper’s Ferry (ndr). Monroe
County; Sweetsprings (m&r), Willow Bend (m&r).
Genus Columella Westerlund
Columella edentula (Draparnaud)
Calhoun County; Grantsville (ndr). Grant County; Greenland Gap (ndr).
Greenbrier County; Renick Valley (ndr). Logan County; Blair (m&r). Mc-
Dowell County; Avondale (m&r). Marion County; High Point (ndr).
Mineral County; New Creek (ndr). Morgan County; Berkeley Spring
(ndr). Pendleton County; Franklin (ndr). Wyoming County; Baileysville
(m&r).
Genus Pupoides Pfeiffer
Pupoides albilabris (Ward)
Grant County; Greenland Gap (ndr). Greenbrier County; Renick Valley
(ndr), Ronceverte (m&r). Jefferson County; Engle (msb). Harper’s Ferry
(ndr) (gwt) (msb), Millville (msb), Rippon (msb). Marion County; Rives-
ville (ndr). Mingo County; Williamson (m&r). Monroe County; Salt
Sulphur Springs (ndr) (m&r). Pendleton County; Upper Tract (ndr).
1949
MacMillan: Land Snails of West Virginia
203
BIBLIOGRAPHY
Baker, H. B.
1930. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 82, p. 208 {Retinella rhoadsi austrina).
Ibid., vol. 82, p. 213 {Retinella cryptomphala solida).
Occasional Papers of the Museum of Zoology, University of
Michigan, no. 220, p. 9 {Runctum vitreum).
1933. Ibid., no. 269, p. 7 {Mesomphix perloevis vulgatus).
Binney, a.
1837. Journal of the Boston Society of Natural History, vol. 1, p. 492
(Mesodon zaletus).
Ibid., vol. 1, p. 494 {Mesodon dentiferus).
1840. Ibid., vol. 3, p. 413 {Ventridens intertextus) .
Ibid., vol. 3, p. 425 {Paravitrea muUidentata) .
Ibid., vol. 3, p. 435 {Hawaiia minuscula).
1843. Ibid., vol. 4, p. 361 {Ventridens demissus).
Bland, T.
1853. Annals of the Lyceum of Natural History of New York, vol. 6, p.
277 {Stenotrema edvardsi).
Ibid., vol. 6, p. 279 {Retinella sculptilis).
1883. Annals of the New York Academy of Sciences, vol. 2, p. 368
{Retinella wheatleyi).
Ibid., vol. 2, p. 369 {Paravitrea petrophila).
Brooks, S. T.
1932. The Nautilis, vol. 46, p. 55 {Triodopsis platysayoides).
Brooks, S. T., and Hunt, G. R.
1936. Annals of the Carnegie Museum, vol. 25, p. 121 {Vertigo clappi).
Brooks, S. T., and Kutchka, G. M.
1937. Annals of the Carnegie Museum, vol. 25, pp. 155-161 {Occurrence
of the Family Carychiidce in West Virginia).
1938. Ibid., vol. 27, pp. 63-85 {Occurrence of the Family Pupillidce in West
Virginia).
Brooks, S. T., and MacMillan, G. K.
1940. The Nautilus, vol. 53, p. 96 {Pomatiopsis prcelonga, and Triodopsis
rugosa).
Brown, A. B.
1902. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 54, p. 434 {Ventridens gularis theloides).
Clapp, G. H.
1908. The Nautilus, vol. 21, p. 121 {Retinella lewisiana).
1915. Ibid., vol. 29, p. 25 {Retinella cryptomphala).
204
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Annals of the Carnegie MusEyn
Cockerell, T. D. A,
1890. Science Gossip, vol. 26, p. 114 {Retinella carolinensis),
Dall, W. H.
1885. Proceedings of the United States National Museum, vol. 8, p. 270
{Hawaiia minuscula alachuana).
1888. Ihid., vol. 11, p. 214 {Guppya sterkii).
1898. The Nautilus, vol. 11, p. 100 {Retinella raderi).
Deshayes, G. P.
1830. Encyclopedie Methodique, vol. 2, p. 217 {Discus patulus).
Draparnaud, J. P. F.
1801. Tableau des Mollusques terrestris et fluviatiles de la France, p. 57
(Vertigo pygmoea).
Ibid., p. 96 {Oxychilus lucidum).
Ferussac, D’aubelard.
1830. Encyclopedie Methodique, p. 224 {Triodopsis denotata).
1842. Symbolae ad historiam Heliceorum, vol. 2, p. 39 {Stenotrema steno-
trema).
Gould, A. A.
1841. Invertebrates of Massachusetts, p. 183 {Retinella electrina).
1848. Proceedings of the Boston Society of Natural History, vol. 3, p.
38 {Paravitrea capsella).
Green, J.
1827. Contributions of the Maclurian Lyceum to the Arts and Sciences,
vol. 1, p. 8 {Mesodon pennsylvanicus) .
Kutchka, G. M.
1938. The Nautilus, vol. 52, p. 13 {Discus patulus angulatus).
Lea, I.
1837. Transactions of the American Philosophical Society, vol. 5, p. 117
{Succinea retusa).
1839. Ibid., vol. 6, p. 87 {Mesodon mitchellianus) .
1841. Ibid., vol. 2, p. 82 {Punctum minutissimum).
MacMillan, G. K.
1940. The Nautilus, vol. 53, p. 143 {Discus patulus carinatus).
Morelet, P. M. a.
1851. Testacea novissima insulae Cubanae et Americae centralia, vol. 2,
p. 8 {Retinella indentata paucilirata) .
Morrison, J. P. E.
1937. Proceedings of the Biological Society of Washington, vol. 50, p. 55
{Retinella virginica).
1937. Ibid., vol. 50, p. 58 {Paravitrea reesei).
1949
MacMillan: Land Snails of West Virginia
205
Morse, E. S,
1859. Proceedings of the Boston Society of Natural History, vol. 7, p.
28 {Striatura milium).
1864. Journal of the Portland Society of Natural History, vol. 1, p. 13
{Retinella hinneyana).
Ibid., vol. 1, p. 17 {Striatura f err ea).
1865. Annals of the Lyceum of Natural History of New York, vol. 8,
p. 207 {Vertigo ventricosa).
Ibid., vol. 8, p. 209 {Vertigo bollesiana).
Mueller, O. F.
1774. Vermium terrestrium et fluviatilium seu animalium etc., vol. 2,
p. 28 {Oxychilus cellarius).
Ibid., vol. 2, p. 30 {Vallonia pulchella).
Ibid., vol. 2, p. 31 {Vallonia costata).
Ibid., vol. 2, p. 32 {Zonitoides nitidus).
Ibid., vol. 2, p. 56 {Euconulus fulvus).
Ibid., vol. 2, p. 104 {Cionella lubrica).
Pfeiffer, L.
1845. Proceedings of the Zoological Society of London, p. 127 {Anguispira
kochi).
PiLSBRY, H. A.
1889. The Nautilus, vol. 3, p. 46 {Ventridens ligera stonei).
1893. Ibid., vol. 7, p. 57 {Strobilops affinis).
1894. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 46, p. 20 {Triodopsis tridentata juxtidens and T. fraudulenta) .
The Nautilis, vol. 7, p. 140 {Mesodon appressus perigraptus).
1896. Ibid., vol. 9, p. 123 {Ventridens collisella).
1898. Ibid., vol. 11, p. 134 {Paravitrea lamellidens).
1899. Ibid., vol. 12, p. 101 {Retinella rhoadsi).
Ibid., vol. 12, p. 116 {Euconulus chersinus polygyratus).
1903. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 55, p. 211 {Paravitrea placentula lacteodens).
1906. Ibid., vol. 58, p. 153 {Discus cronkhitei anthonyi).
1926. The Nautilus, vol. 40, p. 69 {Strobilops oenea).
1927. Manual of Conchology, vol. 28, p. 24 {Strobilops labyrinihica
parietalis).
1928. The Nautilus, vol. 41, p. 83 {Retinella burringtoni) .
1940. Land Mollusca of North America (North of Mexico), vol. 1, pt.
1, p. 805 {Triodopsis fraudulenta vulgata).
1946. Ibid., vol. 2, pt. 1, p. 442 {Ventridens suppressus divisidens).
1948. Ibid., vol. 2, pt. 2, p. 573 {Anguispira alternata angulata).
PiLSBRY, H. A., AND RhOADS, S. N.
1896. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 48, p. 490 {Anguispira alternata carinata).
206
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PiLSBRY, H. A., AND VaNATTA, E. G.
1902. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 54, p. 436 {Ventridens gularis decussatus).
1911. The Nautilus, vol. 25, p. 12 (Stenotrema f rater num cavum).
Rackett, T.
1822. Transactions of the Linnean Society of London, vol. 13, p. 42
{Stenotrema monodon).
Rafinesque, C. S.
1821. In Ferussac, Tableau systematique des animaux mollusque, suivis
d’un prodrome, etc., p. 41 {Mesomphix loevigatus).
1831. Enumeration et Notice sur quelques objects d’histoire naturelle,
p. 3 {Omphalina cuprea).
Redfield, J. H.
1856. Annals of the Lyceum of Natural History of New York, vol. 6 p.
170 {Zonitoides eUiotti).
Say, T.
1816. Nicholson’s Encyclopaedia, American Edition, p. 7 {Mesodon
alholahris).
Ibid., p. 8 {Triodopsis tridentata and Anguispira alternata).
1817. Journal of the Academy of Natural Sciences of Philadelphia, vol.
1, p. 13 {Pomatiopsis lapidaria).
Ibid., vol. 1, p. 15 {Succinea ovalis).
Ibid., vol. 1, p. 17 {Stenotrema hirsutum).
Ibid., vol. 1, p. 123 {Mesodon thyroidus).
Ibid., vol. 1, p. 124 {Strobilops labyrinthica) .
1821. Ibid., vol. 2, p. 116 {Mesodon profundus).
Ibid., vol. 2, p. 151 {Mesodon appressus).
Ibid., vol. 2, p. 154 {Mesodon clausus).
Ibid., vol. 2, p. 155 {Gastrodonta interna).
Ibid., vol. 2, p. 156 {Euconulus cher sinus and Verltridens gularis).
Ibid., vol. 2, p. 159 {Haplotrema concavum).
Ibid., vol. 2, p. 164 {Helicodiscus parallelus).
1822. Ibid., vol. 2, p. 371 {Mesomphix inornatus).
1824. Major S. H. Long’s Account of an Expedition from Pittsburgh to
the Rocky Mountains, vol. 2, p. 257 {Stenotrema f rater num).
Ibid., vol. 2, p. 260 {Succinea avara).
1825. Journal of the Academy of Natural Sciences of Philadelphia, vol.
5, p. 119 {Triodopsis fallax).
1829. The Disseminator of Useful Knowledge, School of Industry, New
Harmony, Indiana, vol. 2, p. 229 {Ventridens suppressus).
1831. Transylvania Journal of Medicine, vol. 4, p. 528 {Hendersonia
occulta.)
1949 MacMillan: Land Snails of West Virginia 207
Shuttleworth, R. J.
1852. Mittheilungen der Naturforschende Gesellschaft in Berne, p. 194
{Paravitrea placentula).
Ibid., p. 195 {Anguispira alternata mordax).
Ibid., p. 198 {Triodopsis rugeli).
Sterki, V.
1890. The Nautilus, vol. 3, p. 136 {Vertigo parvula).
1892. Manual of Conchology, vol. 8, p. 249 {Vallonia excentrica).
1893. The Nautilus, vol. 7, p. 4 {Euconulus chersinus dentatus).
Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 45, p. 271 {Vallonia perspectiva).
Stimpson, W.
1850. Proceedings of the Boston Society of Natural History, vol. 3, p.
175 {Striatura exigua).
Taylor, J. W.
1908. Land and Freshwater Mollusca of the British Isles, vol. 3, p. 98
{Retinella circumstriata).
Tryon, G. W.
1867. American Journal of Conchology, vol. 3, p. 39 {Mesodon albolabris
dentatus).
Vanatta, E. G.
1903. The Nautilus, vol. 16, p. 106 {Mesomphix rugeli oxycoccus).
1911. Proceedings of the Academy of Natural Sciences of Philadelphia,
vol. 63, p. 525 {Gastrocopta contracta climeana).
1936. The Nautilus, vol. 49, p. 99 {Ventridens suppressus virginicus).
Ward.
1841. In C. B. Adams, American Journal of Science, vol. 40, p. 271
{Pupoides albilabris).
WURTZ, C. B.
1948. The Nautilus, vol. 61, p. 86 {Gastrodonta fonticula).
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EXPLANATION OF PLATE 1
Fig. 1. Render sonia occulta (Say) X 7.
Figs. 2,3. Haplotrema concavum (Say) X 1.25.
Fig. 4. Cionella lubrica (Mueller) X 6.3.
Fig. 5. Pomatiopsis lapidaria (Say) X 4.
Fig. 6. Pomatiopsis prcelonga Brooks & MacMillan X 6.
Fig. 7. Vallonia pulchella (Mueller) X 7.3 (after F. C. Baker).
Fig. 8. Vallonia excentrica Sterki X 7.9 (after F. C. Baker).
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EXPLANATION OF PLATE 2
Fig. 1. Vallonia costata (Mueller) X 6.3 (after F. C. Baker).
Fig. 2. Vallonia perspeciiva Sterki X 20 (Brooks, del.).
Fig. 3. Apertural folds and lamellae of Strobilops (after F. C. Baker).
Figs. ^,5. Strobilops lahyrinthica {Sdiy) X 11.7 (after Pi Isbry).
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EXPLANATION OF PLATE 3
Fig. 1. Strobilops lahyrinthica parietalis 'Pilshry X 19.2 (after Pilsbry).
¥igs.2,3. Strohilops affinis Pilsbry X 11.8 (after Pilsbry).
Fig. 4. Strobilops cenea Pilsbry X 14 (after Pilsbry).
Fig. 5. Succinea ovalis Say X 4.7.
Fig. 6. Succinea avara Say X 8.25.
Art. 7
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Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Figs. 13, 14.
EXPLANATION OF PLATE 4
Mesodon albolabris (Say) X 1.8.
Mesodon albolabris dentatus Tryon X 1.7.
Mesodon profundus (Say) X 1.5.
Mesodon mitchellianus (Lea) X 1.7.
Mesodon clausus (Say) X 1.2.
Mesodon thyroidus (Say) X 1.5.
Mesodon pennsylvanicus (Green) X 1.5.
Mesodon appressus (Say) X 1.7.
Mesodon zaletus (A. Binney) X 1.1.
Mesodon say anus (Pilsbry) X 1.3.
Mesodon appressus perigraptus (Pilsbry) X 1.2.
Triodopsis fraudulenta vulgata 'Pilshry X 1.9.
Triodopsis platysayoides (Brooks) X 1.4.
Plate 4
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Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Figs. 10, 11.
Fig. 12.
EXPLANATION OF PLATE 5
Mesodon dentiferus (A. Binney) X 1.8.
Triodopsis rugeli (Shuttleworth) X 2.5.
Triodopsis denotata (Ferussac) X 1.75.
Triodopsis tridentata (Say) X 1.8.
Triodopsis tridentata juxtidens (Pilshry) X 1.9.
Triodopsis fraudulenta (Pilsbry) X 1.8.
Stenotrema stenotrema (Ferussac) X 3.8.
Triodopsis rugosa Brooks & MacMillan X 4.9.
Aperture of Stenotrema.
Stenotrema edvardsi (Bland) X 1.4.
Stenotrema hirsutum (Say) X 3.9.
Plate 5
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EXPLANATION OF PLATE 6
Fig. 1. Stenotrema leai (A. Binney) X 4.6.
Fig. 2. Stenotrema f rater num (Say) X 3.
Fig. 3. Stenotrema fraternum cavum (Pilsbry & Vanatta) X 3.7.
Fig. 4. Punctum minutissimum (Lea) X 13 (after F. C. Baker).
Fig. 5. Punctum vitreum H, B. Baker X 123 (after H. B. Baker).
Fig. 6. Sculptural details of last whorl of Punctum vitreum (after H. B. Baker).
Fig. 7. Discus patulus (Deshayes) X 9.3.
Fig. 8. Discus patulus carinatus MacMillan X 5.6.
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Fig. 1
Fig. 2
Fig. 3
Fig. 4
Figs. 5
Fig. 7
Fig. 8
Figs. 9
EXPLANATION OF PLATE 7
Helicodiscus parallelus (Say) X 9.3 (after F. C. Baker).
Discus cronkhitei (Newcomb) X 5.5.
. Anguispira alternata (Say) X 1.9.
Anguispira alternata angulata Pilsbry X 1.8.
, 6. Anguispira alternata mordax (Shuttleworth) X 1.9.
. Anguispira kochi (Pfeiffer) X 1.9.
Euconulus chersinus polygyratus (Pilsbry) X 8.25 (after F. C.
Baker).
, 10, 11. Guppya sterkii (Dali) X 10.
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EXPLANATION OF PLATE 8
Figs. 1, 2.
Fig, 3,
Figs. 4, 5.
Fig. 6.
Fig. 7.
Figs. 8, 9,
Fig. 10.
Fig. 11.
Fig. 12.
Figs. 13, 14,
Figs. 16, 17.
15.
Euconulus fulvus (Mueller) X 8.5 (after F. C. Baker).
Mesomphix inornatus (Say) X 1.4.
Euconulus cher sinus (Say) X 9.4 (after F. C. Baker).
Omphalina cuprea Rafinesque X 1.1.
Mesomphix rugeli oxy coccus (Vanatta) X 1.2.
Euconulus cher sinus dentatus (Sterki) X 7.2.
Mesomphix perlcevis vulgatus H. B. Baker X 1.9.
Oxychilus draparnaldi (Beck) X 4.
Oxychilus cellarius (Mueller) X 4.7.
Paravitrea muUidentata (A. Binney) X 7.6.
Paravitrea lamellidens (Pilsbry) X 7.8.
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15
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Fig. ]
Figs.
Figs.
Figs.
Fig.
Figs.
Fig.
EXPLANATION OF PLATE 9
1. Paravitrea lamelUdens (Pilsbry) X 7.5.
2, 3. Paravitrea capsella (Gould) X 6.9.
4, 5. Paravitrea placentula (Shuttleworth) X 8.5.
6, 7. Paravitrea placentula lacteode^is (Pilsbry) X 8.7.
8. Paravitrea petrophila (Bland) X 6.8.
9, 10, 11. Paravitrea reesei J. P. E. Morrison X 8.1 (after Morrison).
12. Retinella electrina (Gould) X 6.4.
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Fig. 1.
EXPLANATION OF PLATE 10
Retinella electrina (Gould) X 7.
Figs. 2, 3. Retinella hinneyana (Morse) X 7.9.
Figs. 4, 5.
Figs. 6, 7.
Figs. 8, 9.
Retinella hurringtoni (Pilsbry) X 7.3.
Retinella circumstriata (J. W. Taylor) X 7.9.
Retinella wheatleyi (Bland) X 6.4.
Figs. 10, 11, 12. Retinella virginica J. P. E. Morrison X 9.1 (after Morrison).
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Figs. 1, 2, 3.
Figs. 4, 5, 6.
Figs. 7, 8.
Figs. 9, 10.
Figs. 11, 12.
Figs. 13, 14,
EXPLANATION OF PLATE 11
Retinella lewisiana (Clapp) X 7.
Retinella raderi (Dali) X 9.7.
Retinella rhoadsi (Pilsbry) X 6.9.
Retinella rhoadsi austrina H. B. Baker X 7.6.
Retinella indentata (Say) X 6.5.
15. Retinella carolinensis (Cockerell) X 6.3.
ANNALS, CARNEGIE MUSEUM, Vol. 31 Art. 7 Plate 11
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EXPLANATION OF PLATE 12
Fig. 1. Retinella indentata paucilirata (Morelet) X 7.4.
Figs. 2, 3. Retinella cryptomphala (Clapp) X 6.
Fig. 4. Retinella cryptomphala solida H. B. Baker X 6.6.
Figs. 5, 6. Retinella sculptilis (Bland).
Figs. 7, 8. Hawaiia minuscula alachuana (Dali) X 9.5.
Fig. 9. Hawaiia minuscula (A. Binney) X 9.3 (after F. C. Baker).
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Fig. 1.
Figs. 2, ^
Fig. 5.
Figs. 6, '
Figs. 8, <
Fig. 10.
Figs. 11,
EXPLANATION OF PLATE 13
Retinella sculptilis (Bland) X 6.8.
!, 4. Gastrodonta interna (Say) X 2.7.
Zonitoides elliotti (Redfield) X 8.4.
'. Zonitoides arbor eus (Say) X 5.7.
>. Zonitoides nitidus (Mueller) X 7.5.
Striatura exigua (Stimpson) X 9.8 (after Morse).
12, 13. Striatura ferrea Morse X 13 (after Morse).
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EXPLANATION OF PLATE 14
Fig. 1. Detailed view of sculpture of Striatura exigua.
Fig. 2. Striatura milium (Morse) X 12 (after F. C. Baker).
Fig. 3. Ventridens suppressus (Say) X 7.8.
Fig. 4. Ventridens suppressus virginicus (Vanatta) X 7.
Fig. 5. Ventridens gularis (Say) X 4.
Fig. 6. Ventridens gularis theloides (A. B. Brown) X 4.25.
Fig. 7. Ventridens gularis decussatus (Pilsbry & Vanatta) X 4.3.
Fig. 8. Ventridens collisella (Pilsbry) X 3.75.
Fig. 9. Ventridens demissus (A. Binney) X 2.9.
Fig. 10. V entridens ligera (Say) X 1.5.
Fig. 11. Ventridens intertextus (A. Binney) X 1.8.
Fig. 12. Ventridens ligera stonei (Pilsbry) X 3.
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EXPLANATION OF PLATE 15
Fig. 1. Gastrocopta contractu climeana (Vanatta) X 32.7 (after Vanatta).
Fig. 2. Vertigo ventricosa (Morse) X 32.6 (after Pilsbry).
Fig. 3. Vertigo pygmcea (Draparnaud) X 27.75 (after Pilsbry).
Figs. 4, 5. Vertigo bollesiana (Morse) X 27 (after Pilsbry).
Fig. 6. Vertigo parvula Sterki X 31 (after Pilsbry).
Plate 15
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V
ART. 8. A REVIEW OF THE PILEATE THELEPHORACE.T;
(FUNGI) OF WESTERN PENNSYLVANIA
By LeRoy K. Henry
(Plates 1-2)
The family Thelephoracese belongs to that class of basidiomycetes in
which the spores are borne upon a smooth, or at most a rugose-wrinkled,
hymenial (spore-bearing) surface. These fungi vary in habit from resupi-
nate (expanded with the hymenial surface facing outward) through effuse-
reflexed (expanded with the hymenial surface facing outward and turned
back at the margin to form a cap) to pileate (having a cap bearing the
hymenium on its underside).
Five genera are included here, of which Hymenochcete and Stereiim
resemble in habit some of the small leathery polypores of the genus
Polyporus; some of the Thelephora have the appearance of Clavaria, but
are darker in color; Sparassis looks like a greatly enlarged Tremella, but is
not waxy; and Craterellus has the form of Agaricus^ i.e., with a definite
cap and central stem.
Sparassis, Hymenochcete, and Stereum (with the exception of two species)
grow upon decaying twigs, branches, logs, and roots, while Thelephora
and Craterellus grow upon decaying organic matter in the soil.
This review is based upon the specimens from Western Pennsylvania
now in the Herbarium of the Carnegie Museum. As the eastern boundary
of Western Pennsylvania, I have arbitrarily chosen the eastern borders of
Potter, Clinton, Centre, Huntingdon, and Fulton counties. The initials
following the collection data of the rare or less frequent species, are to
be interpreted as follows: l.k.h. — L. K. Henry; o.e.j. — O. E. Jennings;
D.R.s. — D. R. Sumstine; m.b.k. — Marie B. Knauz; H.s.w. — H. S. Wieand.
Key to the Genera of the Pileate Thelephorace^
I. Fruiting-body resupinate to effused-reflexed to pileate.
A. Hymenial layer brown to chocolate-brown and roughened by the
setae Hiymenochcete
B. Hymenial layer smooth, light brown, cream, white, smoky to
orange Stereum
II. Fruiting-body usually erect to pileate.
A. Fruiting-body leathery, resupinate, lacerate-lobed, dimidiate,
or semi-pileate Thelephora
239
.1
FEB 1 19§0
ued May 4, 1949.
240
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VOL. 31
B. Fruiting-body semi-fleshy, funnel-shaped, top-shaped, agaricoid,
stipitate ; hymenial layer wrinkled Crater ellus
C. Fruiting-body tough and elastic but fleshy, repeatedly branched
into a thick mass of flat contorted-anastomosing branches, and
growing from rotting roots or stump bases Sparassis
Key to Craterellus
I. Sporophore tubular with cavity extending nearly to the base of the stem;
pileus and stem drab when fresh, darker when old; hymenium ashy drab,
even or rugose-wrinkled C. cornucopioides
II. Sporophore somewhat tubular; pileus yellowish brown to fuscus, 1.5 to 3 cm.
broad; hymenium and stem yellow, even or rugose-wrinkled C. lutescens
III. Sporophore not tubular or clavate; pileus convex then depressed to infundi-
buliform, egg-yellow, 2.8 to 8 cm. broad; stem solid, color of pileus; hymenium
even or rugose-wrinkled C. cantharellus
Craterellus cantharellus Schw. ex Fries (Plate 2, fig. 1)
Allegheny County: 1 mi. w. of Ben Avon Heights; Tarentum; Aspinwall;
Sandy Creek; North Park; Imperial; Y2 s. of Smithdale; 2 mi. w. of
Gibsonia; east branch of Big Sewickley Creek, 2 mi. n.e. of Duff City.
Armstrong County: near Ford City; Kittanning. Beaver County: intersec-
tion of Beaver-Conway and Ambridge-Rochester roads; 23^2 irii- s.w. of
New Sheffield; Raccoon Creek Recreational Area. Butler County: 4 mi.
n.e. of Harmony; Little Buffalo Creek at Monroe Station; 2 mi. s. of
Leasuresville ; near Slippery Rock; near Saxonburg; near Culmersville ;
Stone House on Rt. 8, at junction with Rt. 78. Cambria County: near
Cresson. Centre County: near Woodward. Clarion County; Cook Forest;
near Clarion; near keeper. Clearfield County: State Game Lands 34, s. of
Medix Run. Crawford County: French Creek near Cochranton. Erie
County: Presque Isle. Fayette County: Ohiopyle; Killarney Park, s.e. of
Normalville; Claircrest near Normalville; near Geneva. Greene County:
Deep Valley. Indiana County: 2 mi. n.e. of Chambersville. Lawrence
County: Elliot Mills, 5 mi. s.w. of Slippery Rock. Somerset County: near
Trent; near Jennerstown; 3 mi. w. of Berlin; 2 mi. e. of Berlin. Venango
County: 3 mi. n.e. of Emlenton; near Emlenton; 1 mi. n. of Lisbon. West-
moreland County: near Rector; Jones Mills; Rachelwood, 6 mi. s.e. of New
Florence; near Laughlintown ; Idlewild Park, Ligonier; Derry; Kiski
Campus opposite Saltsburg; 2 mi. e. of Trafford; Lynn Run, 3 mi. s.e. of
Rector; 13^2 e. of New Alexandria, off Rt. 22; 4 mi. e. of Waterford;
Pike Run, n.e. of Hopewell; South Greensburg; Loyalhanna Creek dam,
2 mi. s. of Saltsburg.
1949 Henry: Thelephorace^ of Western Pennsylvania 241
Craterellus cornucopioides L. ex Pers. (Plate 2, fig. 2)
Allegheny County: 1 mi. n.e. of Mt. Nebo; Powers Run, opposite Verona;
Warden Mine region, opposite Sutersville; North Park. Armstrong County:
near Kittanning. Beaver County: Temple Hollow, 1 mi. n.w. of Aliquippa;
intersection of Beaver-Conway and Ambridge-Rochester roads; 2]/2
s.w. of New Sheffield. Bedford County: Sulphur Springs. Blair County:
Yellow Spring, off Rt. 22 east of Altoona. Butler County: Dutilh Church
near Criders Corners; Little Buffalo Creek at Monroe Station; 4 mi. n.e.
of Harmony. Centre County: Pine Hall; Scotia, near Port Matilda; Wood-
ward. Clearfield County: State Game Lands 34, s. of Medix Run. Crawford
County: French Creek near Cochranton. Erie County: Weiss Library
Woods, 8 mi. s.w. of Erie. Fayette County: Ohiopyle. Forest County:
western part of Allegheny National Forest. Somerset County: near Trent.
Venango County: 1 mi. n. of Lisbon; Yi n. of Emlenton. Westmoreland
County: 6 mi. s.e. of New Florence; Jones Mills; Derry; Idlewild Park near
Ligonier; Forbes Forest, 3 mi. s.e. of Rector.
Craterellus lutescens Pers. ex Fries
Armstrong County: Kittanning, d.r.s. Centre County: Shafers Creek,
M.B.K. Forest County: near Brookston, J. Kosinski. Rare.
Key To Hymenochaete
I. Resupinate or reflexed.
A. Fruiting-body effused, confluent, becoming reflexed, leathery, thin, light
brown, concentrically ridged, silky fibrillose at first, becoming glabrous;
hymenium not cracking, velvety, light brown H. Curlisii
II. Usually reflexed, sometimes resupinate.
A. Fruiting-body thin, leathery, tobacco colored with orange-yellow margin;
hymenium snuff-brown to sepia, deeply cracked where resupinate, with
one system of radiating and branching cracks to each centimeter of area.
H. tabacina
B. Fruiting-body leathery, rigid, thick, concentrically sulcate, velvety,
finally glabrous, brown, becoming dusky brown with an ochraceous-
tawny margin; hymenium covered with little round elevations, light
brownish black, with setae conspicuous under a lens H. ruhiginosa
Hymenochaete Curtisii (Berk.) Morgan
Butler County: Saxonburg, d.r.s. Somerset County: 2 mi. w. of Berlin,
M.B.K. Rare.
Hymenochaete ruhiginosa Dicks, ex Lev.
Allegheny County: Coraopolis; 2 mi. e. of Upper Talley-Cavey ; Frick
Park; 1 mi. w. of Mt. Nebo; near Verona. Armstrong County: Kittanning.
242
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Beaver County: along Rt. 18, 1 mi. n. of Mechanicsburg. Bedford County:
Sulphur Springs. Butler County: near Saxonburg; Watson’s Run, 2 mi. s.
of Leasuresville. Cambria County: Ebensburg. Centre County: Woodward.
Clarion County: Cook Forest. Clearfield County: State Game Lands 34, s.
of Medix Run. Indiana County: near Homer City. Somerset County: Laurel
Hill near Trent; near Somerset; Kooser State Park, near Bakersville.
Venango County: 3 mi. n.e. of Emlenton; 1 mi. n.e. of Lisbon. Westmoreland
County: near J. R. Mellon estate, 6 mi. s.e. of New Florence; near Latrobe;
along Rt. 271, 4 mi. e. of Waterford; near Derry.
Hymenochaete tabacina Sowerby ex Lev.
Allegheny County: Warden Mine opposite Sutersville, H. Roslund. Bed-
ford County: Sulphur Springs, d.r.s. Cambria County: Cresson, d.r.s.
Elk County: s. of Kane, d.r.s. Fayette County: Ohoipyle, d.r.s. McKean
County: Tionesta Tract near Brookston, l.k.h. Venango County: Little
Scrubgrass Creek, s.e. of Suttons Mills, m.b.k. Westmoreland County: near
Laughlintown, d.r.s. Infrequent.
Sparassis crispa (Wulf.) Fr. (Plate 2, fig. 3)
There is only one species of Sparassis in our region.
Clarion County: near Clarion, d.r.s. Fayette County: Killarney Park,
near Mill Run, d.r.s.; foot of Laurel Hill east of Hopwood, Edna Higbee;
New Geneva, d.r.s. Rare.
Key to Stereum
I. Fruiting-body stipitate; growing on the ground.
A. Pileus 1 to 2 cm. broad, brown or hazel on drying; stem 2 to 4 mm. thick.
5. diaphanum
B. Pileus less than 1 cm. broad, drying pallid; stem up to 1 mm. thick.
5'. Burtianum
II. Fruiting-body small, convex, or tuberculate, thick, cushion-shaped, not
pileate, rarely up to 0.5 cm. broad.
A. Fruiting-body red or reddish, waxy when fresh; growing on Populus only.
5'. rufum
B. Fruiting-body white or yellowish, hard and rigid when fresh; growing on
Quercus and Castanea only 5. frustulosum
III. Fruiting-body sessile, effused-reflexed to overlapping, or if resupinate more
than 0.5 cm. broad.
A. Sporophore hard and rigid when fresh.
1. Fruiting-body brown, tomentose, concentrically sulcate and zoned
with narrow zones that finally become black; hymenial surface even,
bright cinnamon to buff; growing on dead logs of Quercus. This fungus
reaches its northern limit of distribution in southern Pennsylvania.
5 subpileatum
1949
Henry: Thelephorace^ of Western Pennsylvania 243
B. Sporophore thin and flexible when fresh.
1. Fruiting-body usually overlapping, tobacco brown, more or less zonate,
matted tomentose; hymenial surface white or dirty white.. .S. fuscum
2. Fruiting-body 2 to 6 cm. broad, less than 1 mm. thick, often laterally
coalesced, gray, zonate, covered with a tomentum which wears away,
exposing zones of hazel; hymenial surface gray to smoky or pale
yellowish tan. Important in slash decay 5. fasciatum
IV. Sporophore coriaceous when fresh.
A. Fruiting-body 1 to 2 cm. long and broad, gray, zonate in large specimens,
hirsute or strigose; hymenial surface pallid, in age gray-smoky.
5. hirsutum
B. Fruiting-body overlapping, 1 to 2 cm. broad and up to 1.5 mm. thick, pale
yellowish or brown, hirsute, zonate or subzonate, usually crisped or
folded; hymenial surface smooth, pale brown, becoming darker on dry-
ing, surface bleeding where wounded when fresh 5. gansapatum
C. Fruiting-body 2 to 7 mm. broad, white or gray, soft-hirsute or hirsute-
tomentose; hymenial surface yellow to buff, in age more gray-smoky,
5'. ochraceoflavum
D. Fruiting-body 0.5 to 1 cm. broad, gray-cinnamon-buff, strigose hairy at
base, with radiating fibers, margin often crisped and folded, zonate, espe-
cially in age; hymenial surface cinnamon-buff to ochraceous-orange. Im-
portant in slash decay 5. rameale
E. Fruiting-body gray, villose, zonate; hymenial surface pallid to gray or
smoky; growing on conifers, usually Tsuga 5'. sanguinolentum
F. Fruiting-body 1 to 1.5 cm. broad, whitish or gray, silky with radiating
fibrils; hymenial surface pallid to wood colored; chiefly on dead wood of
Carpinus 5. sericeum
V. Fruiting-body usually resupinate.
A. Fruiting-body occasionally with a narrow tomentose, brown, reflexed
margin; hymenial surface dark brown with a nearly white margin when
young. Anally darkening. Not common in Pennsylvania.. . . 5. versiforme
B. Fruiting-body at times with a narrow reflexed margin, pubescent and
brown in color; hymenial surface velvety, dark brown with a conspicuous
white margin. Rare in Pennsylvania 5. albobadium
C. Fruiting-body sometimes with a narrow reflexed, violaceous-gray to pale
buff, tomentose margin; hymenial surface pale gray-lavender to lilac-
lavender but fading 5. roseocarneum
D. Fruiting-body corky when fresh, rarely reflexed; hymenial surface white
to dirty white, typically much cracked S. Murrayi
Stereum albobadium (Schw.) Fries
Centre County: State Game Lands 23, 3 mi. s.e. of Philipsburg, l.k.h.,
5032. Westmoreland County: near Laughlintown. d.r.s. Rare.
Stereum Burtianum Peck
Fayette County: Ohiopyle, d.r.s.; 3 mi. n.e. of Seaton’s (Deer) Lake,
244
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VOL. 31
L.K.H. Westmoreland County: Lynn Run, 3 mi. s.e. of Rector, o.E.j.; J. R.
Mellon estate, 6 mi. s.e. of New Florence, o.E.j. Rare.
Stereum diaphanum (Schw.) Cooke
Allegheny County: near Shades Ravine, 2 mi. e. of Trafford, o.E.j. West-
moreland County: J. R. Mellon estate, 6 mi. s.e. of New Florence, o.E.j.
Rare.
Stereum fasciatum Schw. (Plate 1, fig. 1)
Allegheny County: near Aspinwall; Frick Park, Pittsburgh; Y2 mi- s.e.
of Wildwood; Falls Run, Glenshaw; 1 mi. n.e. of Ben Avon Heights;
Warden Mine region, opposite Sutersville; near Large; w. of Bradford
Woods; Narrows Run, Moon Twp.; ravine n.w. of Saunders; Carnegie;
Emsworth; 1 mi. n.w. of Mt. Nebo; Laschell’s Hollow; North Park;
Bellevue Reservoir; Beaver Grade Road near Montour; Black’s Run, n.
of Oakmont; mi. s. of Smithdale; near Imperial; 4 mi. e. of Monon-
gahela; Fleming Park; Tom’s Run, Dixmont; Flaugherty Run at crossing
of Rt. 51, \Y n.w. of Carnot. Armstrong County: Kittanning. Beaver
County: 1 mi. s. of Fombell; Legionville Hollow, Economy; Mudlick, near
Beaver; 2Y mi. s.w. of New Sheffield; along Rt. 151, 3-4 mi. from junc-
tion with Rt. 18; along Rt. 30, 3 mi. w. of Clinton. Bedford County:
Sulphur Springs; Felton’s Mill along Raystown Branch of Juniata River,
s.e. of Everett. Butler County: near Saxonburg; near Butler; 4 mi. n.e. of
Harmony; Watson’s Run, 2 mi. s. of Leasuresville ; Nixon Station on
Butler Short Line; 1 mi. s.e. of Whitestown; along Rt. 528, Y mi. s.e. of
junction with Rt. 8; near Slippery Rock; n. edge of Zelienople; Marwood;
Kroll’s Mills, 1 mi. n.e. of West Liberty. Cambria County: near Cresson;
Ebensburg. Centre County: Pine Hall; State College; Woodward; State
Game Lands 33, 3 mi. s.e. of Philipsburg. Clarion County: Cook Forest.
Clearfield County: State Game Lands 34,s.of Medix Run; 1 mi. s. of West-
over. Crawford County: Conneaut Lake; Linesville; Pymatuning Swamp,
Hartstown. Erie County: Erie; Presque Isle; Corry; Mercyhurst College,
s. of Erie; near Wellsburg. Fayette County: New Geneva; Ohiopyle; Kill-
arny Park, s. of Normalville. Forest County: 1 mi. n.w. of Brookston. Ful-
ton County: Sideling Hill Park. Lawrence County: Muddy Creek Falls at
mouth of Muddy Creek; Rock Point near Elwood City. McKean County:
Tionesta Tract near Brookston; valley below Mt. Jewett. Mercer County:
2 mi. n.e. of Grove City. Somerset County: Kooser State Park, near Bakers-
ville; near Jennerstown; 3 mi. w. of Berlin; St. Clair. Venango County:
1 mi. n.e. of Lisbon; 3 mi. n.e. of Emlenton. Warren County: Tionesta
1949 Henry: Thelephorace^ of Western Pennsylvania 245
Tract, near Brookston. Washington County: along Buffalo Creek near
junction with Buck Run; vicinity of Hanlin Station; vicinity of Houston;
along Raccoon Creek, 13^ mi. s. of Murdocksville; New Eagle; near
Amity. Westmoreland County: Laurel Hill, 1 mi. e. of Kregar; along Cone-
maugh River across from Saltsburg; 1}^ mi. e. of New Alexandria, off
Rt. 22 ; 1 mi. e. of Mt. Pleasant ; Derry ; Latrobe ; Seward ; 4 mi. e. of Water-
ford, on Rt. 271; near Congruity; Loyalhanna Creek Dam, 2 mi. s. of
Saltsburg; near Laughlintown; South Greensburg; Jones Mills; 6 mi. s. e.
of New Florence; Idlewild Park, near Ligonier; Lyons Run, e. of Trafford;
gorge of Jacobs Creek.
Stereum frustulosum (Pers.) Fries
Allegheny County: Tom’s Run, Dixmont; Frick Park, Pittsburgh;
Flaugherty Run at crossing of Rt. 51, 13^2 n.w. of Carnot; Carnegie;
along Rt. 910, 3 mi. s.e. of Bakerstown; near Sandy Creek; 1 mi. n.e. of
Ben Avon Heights. Armstrong County: Kittanning; Watson’s Run across
from Johnetta. Beaver County: woods at intersection of Beaver-Conway
and Ambridge-Rochester roads; Legionville Hollow at Economy; along
Rt. 51, 4 mi. w. of Beaver. Butler County: near Saxonburg; near Zelienople; '■
4 mi. n.e. of Harmony; Nixon Station on Butler Short Line. Centre County:
Woodward. Crawford County: Conneaut Lake. Erie County: near Erie.
Fayette County: Ohiopyle. Indiana County: along Crooked Creek, 2 mi.
n.e. of Chambersville. Lawrence County: Westminster College near New
Wilmington; Kennedy’s Mills on Slippery Rock Creek, s. of Harlansburg.
Somerset County: near Jennerstown. Venango County: 3 mi. n.e. of Emlen-
ton; 1 mi. n.e. of Lisbon; along Allegheny River, 1 mi. n. of Perry Run.
Washington County: along Raccoon Creek, 2 mi. s. of Murdocksville; near
Burgettstown ; 1 mi. s.e. of Houston. Westmoreland County: Laurel Hill,
6 mi. s.e. of New Florence; Forbes Forest, 3 mi. s.e. of Rector; along Cone-
maugh River across from Saltsburg; Shades Ravine, 2 mi. e. of Trafford.
Stereum fuscum (Schrad.) Quel.
Allegheny County: Schenley Park, Pittsburgh, d.r.s. ; Frick Park, Pitts-
burgh, D.R.s. Armstrong County: Kittanning, d.r.s. Westmoreland County:
Kingston, d.r.s.; Derry, d.r.s. Infrequent; not collected since 1910.
Stereum gausapatum Fries (Plate 1, fig. 2)
Allegheny County: Frick Park, Pittsburgh; Schenley Park, Pittsburgh;
Guyasuta Hollow, near Aspinwall; Carnegie; 2 mi. n.e. of Mt. Nebo; 1
mi. n.e. of Ben Avon Heights; south branch of Little Sewickley Creek,
246
Annals of the Carnegie Museum
VOL. 31
n.e. of Sewickley; near Bakerstown; North Park; along Rt. 910, s.w. of
Bakerstown ; Flaugherty Run at crossing of Rt. 51, 13^2 irii- n.w. of Carnot;
3/2 s.e. of Wildwood. Armstrong County: Kittanning. Beaver County:
woods at intersection of Beaver-Conway and Ambridge-Rochester roads;
along Rt. 30, 3 mi. w. of Clinton; mouth of Potato Garden Run; Raccoon
Creek Recreational Park. Bedford County: Bedford Springs, along Rush
Creek, 8 mi. s.e. of Everett. Butler County: near Saxonburg; Nixon Station
on Butler Short Line; near Zelienople; near Butler; 4 mi. n.e. of Harmony;
Kroll’s Mills, 1 mi. n.e. of West Liberty. Centre County: Woodward; State
College Campus; 8 mi. n.w. of State College. Clarion County: near Clarion;
Cook Forest. Clearfield County: State Game Lands 34, s. of Medix Run.
Fayette County: Ohiopyle; near Somerfield. Forest County: Cook Forest.
Mercer County: near Transfer. Somerset County: 3 mi. w. of Berlin; Kooser
State Park, near Bakersville. Venango County: 3 mi. n.e. of Emlenton.
Washington County: 2 mi. n.e. of Burgettstown; vicinity of Hanlin Sta-
tion. Westmoreland County: Shades Ravine, 2 mi. e. of Trafford; Jones
Mills; Idlewild Park, near Ligonier; near Laughlintown ; near Waterford.
Stereum hirsutum (Willd.) Fries
Allegheny County: Falls Run, Glenshaw, l.k.h., 3030; 23^ mi. e. of
Ambridge, E. H. Graham; near Wildwood, C. K. Henlen. Clarion County:
near Clarion, d.r.s. Westmoreland County: Laurel Hill along Rt. 271, 4
mi. e. of Waterford, l.k.h., 5213; near Laughlintown, d.r.s. Infrequent.
Stereum Murray! (B. & C.) Burt
Clarion County: near Clarion, d.r.s. McKean County: near Kane, d.r.s.
Somerset County: Kooser State Park, near Bakersville, d.r.s. Rare here,
but supposedly common in the mountainous regions.
Stereum ochraceoflavum Schw.
Beaver County: woods at intersection of Beaver-Conway and Ambridge-
Rochester roads, l.k.h., 5254. Fayette County: near Somerfield, d.r.s.
Rare in our collections, but supposed to be common in Pennsylvania.
Stereum purpureum (Pers.) Fries
Cambria County: near Cresson, Aug. 24, 1907, d.r.s. Rare here, but sup-
posed to be fairly common in Pennsylvania.
Stereum rameale Schw. (Plate 1, fig. 3)
Allegheny County: mi. s.e. of Wildwood; 1 to 2 mi. n.w. of Mt. Nebo;
1 mi. n.e. of Leetsdale; near Carnot; Frick Park, Pittsburgh; Powers Run,
1949 Henry: Thelephorace^ of Western Pennsylvania 247
opposite Verona; out Audubon Road from Magee Road; Falls Run,
Glenshaw; 1 mi. w. of Mt. Nebo; 1 mi. n.e. of Ben Avon Heights; along
Beaver Grade Road, near Montour Run ; south branch of Little Sewickley
Creek, n.e. of Sewickley ; near Bakerstown; Bellevue Reservoir ; Flaugherty
Run at crossing of Rt. 51, 13^ mi. n.w. of Carnot; Union Ave. between
Bellevue and Westview; along Rt. 910, 3 mi. s.e. of Bakerstown; along
Rt. 930, 1 mi. n.e. of Stoops Ferry; along Montour Road, 2 mi. w. of
Gibsonia; 13^ mi. n.e. of Allison Park. Armstrong County: Kittanning.
Beaver County: 1 mi. s. of Fombell; along Raccoon Creek, 2 mi. s.w. of
New Sheffield; woods at intersection of Beaver-Conway and Ambridge-
Rochester roads; along Rt. 30, 3 mi. w. of Clinton. Bedford County: 6 mi.
n.e. of Everett; Sulphur Springs, along Brush Creek, 8 mi. s.e. of Everett.
Butler County: near Culmerville; 1 mi. s.e. of Whitestown; near Butler;
Watsons Run, 2 mi. s. of Leasuresville ; 4 mi. n.e. of Harmony; Nixon
Station on Butler Short Line; Marwood; near Saxonburg; Little Buffalo
Creek at Monroe Station; Krolks Mills, 1 mi. n.e. of West Liberty; along
Rt. 528, 34 nii- s.e. of junction with Rt. 8; Slippery Rock; near Zelienople;
Stone House on Rt. 8, about 2 mi.n. of Muddy Creek crossing. Cambria
County: near Cresson; 134 e. of Patton. Cameron County: along Rt.
872, 10 mi. n.e. of Sinnemahoning. Centre County: Woodward; State Game
Lands 33, 3 mi. s.e. of Philipsburg. Clarion County: Cook Forest; 4 mi.
n.e. of Parkers Landing. Clearfield County: State Game Lands 34, s. of
Medix Run. Crawford County: Conneaut; Hartstown; near Linesville.
Erie County: Mercyhurst College, s. of Erie; Presque Isle. Fayette County:
New Geneva; Ohiopyle; Claircrest. Forest County: Cook Forest; 1 mi. n.w.
of Brookston. Indiana County: along Crooked Creek, 2 mi. n.e. of Cham-
bersville. Lawrence County: near New Wilmington; Muddy Creek Falls
at mouth of Muddy Creek; Wayne, near Elwood. McKean County: near
Kane; Tionesta Tract near Brookston. Mercer County: 2 mi. s.w. of Mercer.
Somerset County: near Jennerstown; near Trent; Laurel Hill, 10 mi. e. of
Indian Creek Reservoir; 34 n.e. of Pleasant Union (3 mi. s.e. of Wit-
tenburg). Venango County: along Allegheny River, 1 mi. n. of Perry Run;
Little Scrubgrass Creek, n.e. of Buttons Mills; 3 mi. n.e. of Emlenton.
Warren County: Tionesta Tract near Brookston. Washington County:
along Buffalo Creek, near junction with Buck Run; vicinity of Houston;
along Raccoon Creek, 134 mi. s. of Murdocksville. Westmoreland County:
along Conemaugh River across from Saltsburg; Kiski Campus near Salts-
burg; Laurel Hill, 1 mi. e. of Kregar; Shades Ravine, 2 mi.e. of Trafford;
along Loyalhanna Creek at dam, 2 mi. s. of Saltsburg; Rock Run, Forbes
248
Annals of the Carnegie Museum
VOL. 31
Forest, 3 mi. s.e. of Rector; Congruity, 4 mi. e. of Delmont; Chestnut
Ridge, Hillside; Jones Mills; South Greensburg; near Laughlintown ;
Loyalhanna Gorge, 2 mi. s.e. of Kingston; along Rt. 271, 4 mi. e. of Water-
ford; J. R. Mellon estate, 6 mi. s.e. of New Florence.
Stereum roseocarneum (Schw.) Fries
Butler County: near Butler, d.r.s. Rare here, but supposed to be rather
common in Pennsylvania.
Stereum rufum Fries
Clearfield County: State Game Lands 34, s. of Medix Run, l.k.h., 3485.
Crawford County: Pymatuning Swamp, Blair Swamp Road, F. H. Beer.
McKean County: 4 mi. n.e. of Clermont, H. Roslund. Rare.
Stereum sanguinolentum Alb. & Schw.
Allegheny County: Falls Run, Glenshaw, l.k.h., 3031. Lawrence County:
Muddy Creek Falls at mouth of Muddy Creek, l.k.h., 263. Rare here,
but supposed to be rather common in Pennsylvania.
Stereum sericeum Schw. (Plate 1, fig. 4)
Allegheny County: mi. s.e. of Wildwood; Powers Run, opposite
Verona; Frick Park, Pittsburgh; 1 to 2 mi. n.w. of Mt. Nebo; along
Beaver Grade Road near Montour Run; Flaugherty Run at crossing of Rt.
51, nii- n-w. of Carnot; along Rt. 910, 3 mi. s.e. of Bakerstown; vicinity
of Warden Mine, opposite Sutersville. Armstrong County: Kittanning.
Beaver County: Mudlick, near Beaver; along Rt. 18, 1 mi. n. of Mechanics-
burg; woods at intersection of Beaver-Conway and Ambridge-Rochester
roads; mouth of Potato Garden Run, near Murdocksville; along Rt. 51,
4 mi. w. of Beaver; Temple Hollow, 1 mi. n.w. of Aliquippa. Bedford
County: Sulphur Springs. Butler County: Marwood; Nixon Station on
Butler Short Line; 4 mi. n.e. of Harmony; along Rt. 528, Y mi. s.e. of
junction with Rt. 8; Watsons Run, 2 mi. s. of Leasuresville ; Little Buffalo
Creek at Monroe Station; 1 mi. s.e. of Whitestown; Kroll’s Mills, 1 mi.
n.e. of West Liberty; near Saxonburg; Stone House on Rt. 8, 2 mi. n. of
Muddy Creek crossing; 3 mi. n.e. of Harmony, along Little Connoqueness-
ing Creek. Cameron County: along Rt. 872, 10 mi. n.e. of Sinnemahoning.
Elk County: along Rt. 555, 2 mi. e. of Medix Run; Middle Fork of Clarion
River, Jones Twp. Erie County: Corry. Fayette County: Killarney Park,
s. of Normalville; Ohiopyle. Forest County: near Brookston. Indiana
County: near Glen Campbell; along Crooked Creek, 2 mi. n.e. of Chambers-
ville. Lawrence County: near New Wilmington. Somerset County: near
1949 Henry: Thelephorace^ of Western Pennsylvania 249
Trent. Venango County: 1 mi. n.e. of Lisbon. Washington County: vicinity
of Hanlin Station. Westmoreland County: Forbes Forest, 3 mi. s.e. of
Rector; Chestnut Ridge, Hillside; Derry; Shades Ravine, 2 mi. e. of Traf-
ford; near Waterford; 6 mi. s.e. of New Florence.
Stereum subpileatum B. & C.
Westmoreland County: Lynn Run, Forbes Forest, 3 mi. s.e. of Rector,
1924, o.E.j. Rare. Reaches its northern limit of distribution in southern
Pennsylvania.
Stereum versiforme B. & C.
Allegheny County: Keown, along Babcock Blvd., Sept. 8, 1940, d.r.s.
Rare.
Key to Thelephora
I. Sporophore soft-leathery, usually with central stem; pileus composed of nar-
row branching, flattened or cylindrical divisions.
A. Fruiting-body 2 to 6 cm. high, branched, glabrous, with fetid odor.
T. palmata
B. Fruiting-body 3 to 5 cm. high, branched, minutely pubescent, no fetid
odor; stem villose T. anthocephala
II. Sporophore leathery, usually with a central stem; pileus more or less funnel-
shaped, cup-shaped, or fan-shaped, often splitting radially into divisions;
hymenium brown to smoky-brown.
A. Fruiting-body 1.5 to 3 cm. high, smoky drab to ashy; pileus funnel-shaped,
deeply cleft into narrow divisions; stem villose T. multipartita
B. Fruiting-body 6 mm. to 2.5 cm. high, pallid to tawny-olive; pileus funnel-
shaped, or divided into triangular divisions, or fan-shaped. . . T. regularis
C. Fruiting-body 1.5 to 5 cm, high, smoky-purple; pileus funnel-shaped,
often double by growth of smaller pilei from disk of the principal pileus
or by wedge-shaped lobes rising from its upper surface; stem central,
villose T. caryophyllea
D. Fruiting-body 2.5 to 6 cm. high, dirty-whitish with a brown center;
pileus cup-shaped, composed of many small pilei which rise from a com-
mon base, or with interior of cup filled with small pilei and lobes; upper
surface roughened with masses of fibers; stem central, when present.
T. vialis
III. Sporophore effuse-reflexed, semicircular or horizontally expanded,
A. Hymenium pale cinnamon-buff.
1. Fruiting-body 2 to 4 cm. in diameter, leathery, spongy when dry,
usually semicircular and somewhat imbricated, sometimes effused-
reflexed, fibrous-tomentose with thick margin, sessile or nearly so,
cinnamon-buff T. albido-brunnea
250
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B. Hymenium rusty-brown to smoky.
1. Fruiting-body in clusters, 5 to 8 cm. in diameter, soft, whitish, then
red-rusty, drying chestnut-brown; pilei imbricated, fibrous with matted
fibers and adnate squammules, margin whitish, fimbriate at first, be-
coming entire and concolorous with pileus; stems lateral and growing
together T. intyhacea
2. Fruiting-body in clusters, 5 to 8 cm. in diameter, leathery-soft, dark
smoky to fawn colored, semi-circular or encrusting and effused-
reflexed; pilei more or less imbricated, fibrous-squamulose and usually
strigose, margin fibrous-fimbriate; sessile or with a short stem-like
base T. terrestris
Thelephora albido-brunnea Schw.
Butler County: 4 mi. n.e. of Harmony, l.k.h. 556. Westmoreland County:
J. R. Mellon estate, 6 mi. s.e. of New Florence, d.r.s. Rare.
Thelephora anthocephala Bull, ex Fries
Allegheny County: Frick Park, Pittsburgh; Guyasuta Hollow, near
Aspinwall. Armstrong County: Kittanning. Bedford County: Sulphur
Springs. Fayette County: Ohiopyle. Indiana County: Juneau. Westmoreland
County: Jones Mills.
Thelephora caryophyllea Schaeff. ex Fries (Plate 2, fig. 4)
Allegheny County: Frick Park, Pittsburgh; ravine at Glenshaw; North
Park. Armstrong County: Kittanning. Butler County: 4 mi. n.e. of Harmony.
Elk County: near Kane. Fayette County: Ohiopyle. Mercer County: near
West Middlesex. Westmoreland County: Rock Run, 3 mi. s.e. of Rector;
13^ mi. e. of New Alexandria, off Rt. 22.
Thelephora intybacea Pers. ex Fries
Armstrong County: Kittanning, d.r.s. Beaver County: Temple Hollow,
1 mi. n.w. of Aliquippa, S. Ristich. Clarion County: near Leeper, d.r.s.
Erie County: Presque Isle, o.e.j.
Thelephora multipartita Schw.
Btdler County: Harmony, July 21, 1904, R. J. Plaum. Centre County:
Milheim, July 5, 1935, d.r.s. Rare.
Thelephora palmata Scop. (Plate 2, fig. 5)
Allegheny County: near Wildwood; Frick Park, Pittsburgh; 1 mi. n.e.
of Ben Avon Heights; Sandy Creek; Powers Run, opposite Verona;
Douthett, near Warrendale; Coraopolis. Armstrong County: Kittanning.
Beaver County: Temple Hollow, 1 mi. n.w. of Aliquippa; intersection of
1949 Henry: Thelephorace^ of Western Pennsylvania 251
Beaver-Conway and Ambridge-Rochester roads. Butler County: 4 mi. n.e.
of Harmony; Little Buffalo Creek at Monroe Station. Cambria County:
Ebensburg. Centre County: Woodward; Alan Seegar region. Elk County:
along Rt. 555, 2 mi. e. of Medix Run. Erie County: mi. e. of Watts-
burg; Weiss Library Woods, 8 mi. s.w. of Erie. Fayette County: Ohiopyle;
3 mi. n.e. of Seaton’s (Deer) Lake. Somerset County: 3^ mi. n.e. of Pleasant
Union (3 mi. s.e. of Wittenburg). Venango County: 3 mi. n.e. of Emlenton.
Warren County: Benson’s Swamp, 5 mi. e. of Columbus. Westmoreland
County: Ligonier; Nawakwa Lodge, Forbes Forest, 3 mi. s.e. of Rector;
Jones Mills; South Greensburg; Latrobe; .Laurel Hill, along Rt. 271, 4
mi. e. of Waterford.
Thelephora regularis Schw.
Allegheny County: 1 mi. n. of Ben Avon Heights, l.k.h., 1213; McKinley
Park, West End, Pittsburgh, G. E. Wheeler; vicinity of Warden Mine,
opposite Sutersville, l.k.h., 3329. Armstrong County: Kittanning, d.r.s.
Butler County: near Slippery Rock, d.r.s. ; 4 mi. n.e. of Harmony, l.k.h.,
4165. Venango County: 1 mi. n.e. of Lisbon, l.k.h., 1079.
Thelephora terrestris Ehrh. ex Fries
Allegheny County: Camp Meeting Extension Road, n.w. Sewickley
Twp., E. M. McClelland. Erie County: Presque Isle, o.E.j. Rare.
Thelephora vialis Schw.
Bedford County: Pa. Turnpike, 3 mi. e. of Kegg, o.e.j. Fayette County:
Gibbon’s Glade, l.k.h.; 2 mi. n.e. of Seaton’s (Deer) Lake, l.k.h. McKean
County: near Kane, d.r.s. Westmoreland County: Derry, d.r.s.; Laurel
Hill, east of Rector, o.e.j.; Idlewild Park, near Ligonier, d.r.s.; J. R.
Mellon estate, 6 mi. s.e. of New Florence, o.e.j.
Summary
A generic key and descriptive keys to thirty-two species of the pileate
Thelephoraceae have been provided, and distributional records in Western
Pennsylvania are given for each species.
The records of distribution show that many counties in the north-
western, eastern, and southwestern sections of Western Pennsylvania are
either not represented at all or are represented by collections made at
only one or two localities.
In order to obtain a more complete record of the distribution of the
family, additional collecting in these areas will be necessary.
252
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VOL. 31
BIBLIOGRAPHY
Burt, Edward Angus
1914. The Thelephoraceae of North America. I. Thelephora. Ann.
Missouri Bot. Garden, 1: 185-228, pi. 4-5.
1914. Ibid. II. Craterellus. Vol. 1: 327-350, pi. 15-17.
1918. Ibid. X. Hymenochaete. Vol. 5: 301-372, pi. 16-17.
1920. Ibid. XII. Stereum. Vol. 7: 81-248, pi. 2-6.
Coker, W. C.
1919. Craterellus, Cantharellus, and related genera in North Carolina.
Jour. Elisha Mitchell Scientific Soc., 35: 24-48, pi. 2-17.
1921. Notes on the Thelephoraceae of North Carolina. Jour. Elisha
Mitchell Scientific Soc., 36: 146-196, pi. 14-35.
Overholts, L. O.
1939. The Genus Stereum in Pennsylvania. Bull. Torrey Bot. Club, 66:
515-537, pi. 14-18.
Rea, Carleton
1922. British Basidiomycetes. The British Mycological Society, Cam-
bridge University Press, London.
A '
7
Si.'
254
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VOL. 31
EXPLANATION OF PLATE 1
Fig. 1. Stereum fasciahim X 7/8.
Fig. 2. Stereum gausapatufu Fries. X 4/5.
Fig. 3. Stereum rameale Schw X 4/5.
Fig. 4. Steremn sericeum Schw. X 5/7.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 8
Plate 1
4
256
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 2
Fig. 1. Craterellus Cantharellus Schw. eyiYviQs. X 2/3.
Fig. 2. Craterellus cornucopioides L. ex Pers. X 5/6.
Fig. 3. Spar as sis crispa (Wulf.) Fr. X 2/3.
Fig. 4. Thelephora caryophyllea Schaeff. ex Fries. X 2/3.
Fig. 5. Thelephora palmat a Scoi). X 5/6.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 8
Plate 2
4
5
ART. 9. LARVAL DEVELOPMENT OF NECTOPIIRY NOIDES
TORNIERI (ROUX), WITH COMMENTS ON DIRECT
DEVELOPMENT IN FROGS
By Grace L. Orton
(Plates 1-3)
The African bufonid genus, Nectophrynoides, to which three species are
now referred, comprises the only known ovoviviparous frogs. The two
East African (Tanganyika Territory) species have long been known;
vivipara was described by Tornier (1905), who referred it to the genus
Pseudophryne, and tornieri was described and referred to Nectophryne by
Roux (1906), Noble (1926) proposed a new genus, Nectophry^ioides, for
these two species, associating them principally because of their unusual
mode of development. The third form, occidentalis, was described by
Angel (1943), from Mt. Nimba in French Guinea, near the Liberian
border.
The remarkable type of development of these frogs was discovered
early, but until recently little detailed information on embryonic stages
was available. Brief notes on embryos of N. vivipara were published by
Tornier (1905) and of N. tornieri by Krefft (1911) and Noble (1927).
Counts of uterine embryos were listed by Barbour and Loveridge (1928).
The study of occidentalis by Angel and Lamotte (1944) is, however, the
only detailed account of the development of these frogs that has come to
my attention.
In the present paper, several stages in the early development of tornieri
are described and compared with occidentalis, and some comments on di-
rect development in relation to larval characters are presented.
Acknowledgments. — Through the courtesy of Mr. Arthur Loveridge and
Mr. Benjamin Shreve of the Museum of Comparative Zoology I have had
the opportunity to examine several series of embryos of Nectophrynoides
tornieri in the MCZ collection. Preliminary study of this material was
made in connection with my doctoral studies under the direction of Dr.
Norman Hartweg, during tenure of a University Fellowship at the Uni-
versity of Michigan. Larvae in the University of Michigan Museum of
Zoology (UMMZ) collection cited in this paper were also studied at that
time. The Nectophrynoides material was re-examined in 1948. I am also
indebted to Mr. Karl P. Schmidt for the recent opportunity to study
257
sued June 28, 1949.
fEB 1 iggff
258
Annals of the Carnegie Museum
VOL. 31
larvae of Rhinoderma danvinii in the collection of Chicago Natural His-
tory Museum (CNHM).
I. Larval Development of Nectophrynoides tornieri
The material studied is divisible into five rather well-differentiated
‘^stages” of development. Some are anatomically younger than the de-
velopmental stages of N. occide^italis described by Angel and Lamotte
(1944), and provide more definite information on certain larval characters.
My studies are not as detailed as those of Angel and Lamotte for I made
no serial sections, determining internal characters only by simple micro-
dissection.
The actual ages of the embryos are not known. In the numerical stage
designations of the series, the term “stage” is used only to facilitate refer-
ence to particular specimens, and is not intended to have an exact morpho-
logical significance.
First Stage. — (PI. 1, fig. 1) In this, the earliest stage available, the
embryo is already well-formed ; head structures are becoming differentiated,
distinct limb bud anlagen are present, and there is a long thin tail. There
is a large and rather ovoid (probably round in life) yolk mass. Unpig-
mented eye and ear capsules are visible, and the nasal pits are present.
Jaw structures are still in a very rudimentary condition, and the mouth is
not yet open. There is no external indication of adhesive organs. Two
pairs of short but very distinct external gills are present. The front leg-
buds are still external. The long, thin, tail appears to be composed chiefly
of notochordal tissue and very low, ridge-like, dorsal and ventral fins.
Some of the specimens have a few faint melanophores on the body, but
they are otherwise unpigmented. Total length ± 5.5 mm.
Second Stage. — (PL 1, fig. 2) The larvae of the second group are con-
siderably advanced over the first series, particularly in the development of
the head structures. The eyeballs are well differentiated and quite heavily
pigmented. The mouth is a small, simple, transverse, slightly crescentic,
opening. The opercular folds are of varying extent in different specimens
and well illustrate the process of enclosure of the forelimb buds and forma-
tion of the spiracle. The morphological relations appear to be entirely as
in a normal aquatic tadpole. In the least well-developed specimens the
operculum already encloses the gill arch region and the base of the foreleg-
bud on each side, and forms a transverse fold entirely across the throat.
In more advanced specimens fusion of the edge of the opercular fold with
the body wall ventro-caudad to the foreleg buds is underway, forming- an
1949
Orton: Larvae of Nectophrynoides
259
opercular cavity and progressively reducing the size of its external open-
ing (PL 2, figs. 1-4). The final step in this sequence of events is the re-
finement of the opercular opening remaining on the left side to form the
spiracle. This is completed in specimens in the third stage (PI. 2, fig. 5).
In the gill region of specimens of the second stage, three small gill arches
and two open gill slits can be seen. The first two arches each bear a single
tiny gill rudiment. The leg buds are much better developed than in the
first stage, and the tail is longer. There is a fine network of melanophores
over the head and back. Total length ± 6 mm.
Third Stage. — (PI. 1, fig. 3; MCZ 12513) The specimens in the third
stage show further advances. Developmentally, they appear to agree most
closely with Angel and Lamotte’s Stage l-j- (“1 bis”) of N. occidentalis.
The eyes are very large and the facial region is small, embryonic in pro-
portions. The small mouth shows definite though rather simple indica-
tions of larval structure. There are no beaks, labial teeth, or papillae,
but there is a short fleshy ridge representing the larval lower lip. The
sides of the upper jaw overhang the lateral part of the lower jaw (PI. 3,
fig. 1). The jaw region of one specimen was dissected, and it was found
that the general pattern of the cartilages and muscles is essentially larval,
although the proportions are, of course, much modified. The quadrate is
shorter than in aquatic tadpoles, but a distinct processus muscularis is
present. A small flat orbitohyoideus muscle connects it with the cera-
tohyal, which is of larval form and position. The other jaw muscles are
more clearly shown in the fourth stage, and will be discussed in detail
under that heading. Two pairs of open gill slits were found in the speci-
men dissected. There is a small but distinct spiracle on the left side. The
dorsal surfaces of the head, body, and base of the tail, are finely speckled
with melanophores. The leg buds are somewhat elongated, the posterior
pair showing an indication of foot and leg segments. The tail is long and
thin, with low rudiments of dorsal and ventral fins. Total length dh 6 mm.
Fourth Stage. — (PI. 1, fig. 4; MCZ 12515) These larvae are approxi-
mately equivalent to Angel and Lamotte’s Stage 2 of N. occidentalis. The
general proportions are much the same as in the third stage, the principal
changes visible externally being the larger and better developed legs and
greater transparency of the skin. A very small sinistral spiracle is present;
the nostrils are quite large; the mouth is still small. The legs and digits are
well developed. The long rat-like tail has about 17 somites; the first 10-12
of them are well developed, the rest are rather irregular. The low, thin
caudal fins are transparent and appear to contain an extensive vascular
260
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VOL. 31
network. The jaw region of one specimen was dissected (PI. 3, figs. 2-4).
The arrangement of the jaw cartilages and muscles was found to be es-
sentially larval, as in the third stage, but the details were more easily
worked out in the present specimen because of its somewhat larger size.
The various structures are small and modified in proportions, in correla-
tion with the embryonic contours of the entire head. The posterior part of
the quadrate is quite short, the processus muscularis being not far an-
terior to the otic capsule. The processus muscularis is quite well dif-
ferentiated although, as would be expected, its size relative to other cranial
structures is much smaller than in aquatic tadpoles. The orbitohyoideus
muscle is thin and transparent, and the underlying origins of the sus-
pensorioangularis and ceratohyoangularis can be seen through it. The
pterygoideus and the adductor mandibulae posterior longus have sepa-
rate, laterally-adjacent origins, and the latter muscle appears to be dif-
ferentiated into two slips distally, as in most aquatic tadpoles. Small
adductor m. subexternus and articularis muscles are present, and have
normal origins. All of the adductor muscles seem to insert on the lower jaw.
In most aquatic tadpoles certain of these muscles insert on the upper
labial cartilage, their insertions shifting to MeckePs cartilage during
metamorphosis. A narrow strap-like subhyoideus muscle and a thin
crescent-shaped intermandibularis are present in the normal larval posi-
tions. The lower jaw is differentiated into the typical Meckelian and
lower labial segments, but histological studies will be needed to deter-
mine whether these cartilages are separate or fused, and whether there
is a distinct upper labial cartilage. The specimen dissected appeared to
have three open gill slits on each side. On the left side, the first gill arch
was larger than the others and bore two rows of minute gill rudiments;
on the right side the first two gill arches were of approximately equal size
and bore no distinct traces of gills. A typical larva in this series has a total
length of 10 mm., head and body length 4 mm.
Fifth Stage. — (PI. 1, fig. 5; MCZ 12512) In this series, metamorphosis
is well underway and the specimens are approximately in Angel and
Lamotte^s Stage 3 of occidentalis. The forelegs have emerged ; anterior to
each a small slit remains, connecting the gill arch region with the outside.
The head proportions are still embryonic, with relatively huge eyes, but the
mouth has begun to enlarge. The rudimentary lower lip has disappeared.
The long, thin, rat-like tail is still present. The hind legs show traces of
dark banding, but the body is still rather weakly pigmented. The total
length at this stage is rb 11 mm., snout-vent length ± 4.5 mm.
1949
Orton: Larvae of Nectophrynoides
261
Discussion
It is instructive to re-assemble the notes on external characters in the
form of a routine diagnosis so the larval stage of N. toniieri can be more
easily compared with conventional tadpoles: Head and body narrow;
head small; facial region of embryonic proportions, with very small jaws;
eyes very large. Spiracle very small and inconspicuous, sinistral. Tail long,
thin, rat-like in proportions, consists principally of a well-developed
notochord partly concealed by about 17 small somites basally and with
very low ridge-like dorsal and ventral fins. Mouthparts greatly simplified;
mouth very small, jaws devoid of horny beaks, with or without a short
flap-like rudiment of lower lip; no tooth ridges, labial teeth, or papillae.
Dorsum weakly pigmented with fine network of melanophores. Tail color-
less or with a few scattered melanophores basally. Maximum total length
before metamorphosis ±11 mm.
Although tornieri is ovoviviparous and undergoes direct development,
it is evident that few of the important characters of the tadpole stage have
actually been lost. There is no external indication of adhesive organs,
labial teeth, or horny beaks in the material studied. Anatomically, these
structures are relatively superficial, and they are commonly lost in species
that undergo direct development or other modifications of the life history.
On the other hand, tornieri has a distinct trace of a larval lower lip (at
least in some specimens), the internal structure of the jaw region is es-
sentially larval in pattern, a spiracle is present and its development is
essentially typical. Tiny but distinct gill arches, gill slits, and external
gills develop. Although the general proportions of the head, body, and
tail have become greatly modified, it is clear that this species still retains
the more fundamental characters of the tadpole stage in recognizable
condition and still undergoes an extensive metamorphosis. Anatomically,
there is a well-defined tadpole stage even though it is no longer free-living.
Angel and Lamotte emphasized the absence of a spiracle in N. occi-
dentalis. From the figures and description of their Stages 1 and 2, however,
it is obvious that opercular folds develop in that species, for the foreleg
buds are internal. During the growth of the operculum in the period
preceding their Stage 1, a spiracle would therefore have been present, al-
though it may subsequently have closed completely. Developmentally,
the spiracle of the tadpole is merely the excurrent opening remaining after
most of the free margin of the opercular fold has fused with the body wall
below and behind the gill region. The size of the spiracle depends on how
complete this fusion is; the more complete the fusion, the smaller the
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VOL. 31
spiracle. It is possible that vestiges of gill arches and associated structures
will also be found in younger embryos of occidentalis when such specimens
become available. The internal structure of the jaw region of embryonic
occidentalis is not yet known. It will be of considerable interest to learn
how much (if any) of the larval arrangement of cartilages and jaw muscles
is still retained in that species. Angel and Lamotte reported no indication
of any external buccal structures.
Angel and Lamotte noted that in the stages of occidentalis available to
them the ovarian eggs are very minute (diameter 0.2 mm.), and there is no
trace of a large yolk mass in the embryo. Although there is a wide develop-
mental gap between the ovarian egg with a diameter of 0.2 mm. and an
advanced embryo with a total length of 4.5 mm., the next stage available
to them, they concluded that the egg remains poorly equipped with yolk
and that virtually all nourishment must therefore be provided through
maternal tissues. Description of the intervening stages (the mature ovum
and pregastrular phases of embryonic differentiation) in this species will
be awaited with great interest.
II. Comments on Direct Development in Frogs
In recent years there has been an increasing interest in problems re-
lating to direct development in frogs. Lutz (1947, 1948) has reviewed the
general field, with particular reference to the complex frog fauna of south-
eastern Brazil. She emphasized, among other points, the extensive paral-
lelism shown in various unrelated groups of frogs that have independently
achieved specialized developmental patterns, and the tendency for non-
functional larval structures to be reduced or entirely lost in non-aquatic
types of larvae that have an increased yolk supply. These trends result in
a more or less direct transition from generalized embryo to frog without
the intervention of a specialized free-swimming larval stage.
The fate of larval structures in non-aquatic development differs widely
among different species, and various degrees of retrogression of these
structures can be demonstrated. In a typical aquatic life history, critical
early phases in the formation of tadpole structures are completed so
rapidly and at such a small larval size that they are difficult to study.
In the comparatively larger embryos of non-aquatic forms, these early
phases (especially in the jaw and branchial regions) are prolonged and the
relative growth rates are altered, with the result that the structural re-
lations are somewhat clarified. Some of the problems in tadpole anatomy
on which this situation affords evidence are discussed below.
1949 Orton: Larvae of Nectophrynoides 263
1. The Tadpole Operculum and Spiracle
The anatomical relations of the operculum and spiracle to the under-
lying structures in the tadpole are complex, but can be better understood
by comparing the developmental sequence in typical aquatic tadpoles
with that in various specialized non-aquatic frog embryos. The so-called
operculum of the typical tadpole has a dual origin. It is derived princi-
pally from the transverse dermal fold growing backward from the hyoid
region, but a low ridge-like fold across the anterior part of the belly is
incorporated into the completed structure. The latter fold develops pos-
terior to the level of the foreleg region and extends vertically to a point
above it, so that the fusion of the two components of the operculum en-
closes not only the gill chamber, but with it the area in which the foreleg-
will develop. Ordinarily, the process is carried to completion, leaving only
a small excurrent opening, the spiracle. In frogs with an abbreviated
larval history, the development of the operculum and spiracle may remain
fairly typical or it may be arrested at some point short of completion.
The process is completed in Nectophrynoides tornieri (PI. 2, figs. 1-5),
Rhinoderma darwinii, Pipa aspersa, and Pipa pipa; in the latter two
forms, symmetrically paired spiracles develop, as in aquatic larvae of other
Pipidae and of Rhinophrynus.
In certain forms in which the operculum remains incomplete, the
homology of the dermal vestiges in the branchial region has been ques-
tioned. Lynn (1942) suggested that the small dermal fold over the base
of the foreleg bud in Eleutherodactylus embryos is not homologous with the
operculum of the typical aquatic tadpole. In various forms of Gastrotheca
and related genera of “marsupial” hylids, a graded series of departures
from the complete pattern of opercular formation can be demonstrated,
and these help to interpret conditions in Eleutherodactylus. As is now
well known from the work of Noble (1927) and others, some marsupial
hylids have an aquatic larval stage and others have direct development,
but in all known forms the young are carried on the back of the female dur-
ing at least the early phases of development. All, so far as known, have the
same highly diagnostic peltate (c/., nasturtium leaf) type of external gills,
which form a thin vascular sheet enveloping the embryo. Because of the
comparatively large size attained by these embryos and the relatively slow
(apparently) completion of branchial development, they are excellent
material for the study of operculum formation. Several series of embryos
of Gastrotheca sp. examined (c.g., UMMZ 68165) were found to have
larval body and tail proportions and buccal apparatus and to have dis-
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VOL. 31
tinct leg buds before operculum formation was completed. The long-
delayed closure of the operculum in these larvae is evidently associated
with prolonged functioning of the relatively huge peltate external gills.
In these large pre-hatching larvae, the composite origin of the operculum
and its relation to the gill arches and to the already well-formed foreleg
buds are very clearly shown.
In the exceptionally large embryos of Ilemiphractus divaricatiis (UMMZ
92106), the various structures are much modified in proportions and some-
what displaced by the relatively huge yolk mass (its diameter rt 10 mm.).
In the stage studied, there is a short, rather thick, flap-like fold across the
throat and overlapping the bases of the long gill stalks; laterally the fold
thins out somewhat and forms a deep pocket surrounding the base of each
foreleg bud. The structural relations here, although much altered in pro-
portions, are not basically different from conditions in the young pre-
swimming larvae of Gastrotheca described above. The developmental
sequence in the series of Gastrotheca is in turn only proportionately dif-
ferent from that of an ordinary tadpole with a typical larval history. The
differences are in degree only; the structures involved are the same in each
of these forms.
In embryos of Elentherodactylus cooki (UMMZ 73550), the presence of a
short dermal fold over the base of the foreleg bud and the absence of a
transverse opercular fold across the throat were observed, as described by
Lynn (1942) and other authors for other species in this genus. Morpho-
logically, this condition appears to be but a further simplification of the
opercular pattern noted above for Hemiphractus divaricatus. The trans-
verse part of the composite operculum has become entirely suppressed
and all that remains is the pocket-like fold over the base of the foreleg
anlage. If the composite origin of the tadpole operculum is granted, it is
probable that the dermal fold over the foreleg bud in EleiitherodactyUis is
homologous with the postero-lateral part that normally encloses the fore-
leg within the gill chamber, and thus the fold may be correctly interpreted
as an opercular vestige.
As noted above, the spiracle is not always formed in non-aquatic frog
embryos. Theoretically, it could be eliminated from the phenotype either
by complete closure of the opercular folds, or by their failure to close at all,
leaving a wide gap through which the branchial region and foreleg buds
remain exposed. Examples of the latter type include Elentherodactylus
and Ilemiphractus. In such forms the forelegs are more or less fully
visible externally throughout their development. Absence of a spiracle
1949
Orton: Larvae of Nectophrynoides
265
as the result of complete opercular closure has been reported in Sooglossus
(Brauer, 1898), Breviceps (de Villiers, 1929 a), Anhydrophryne (Hewitt,
1929), and Arthroleptella (Power and Rose, 1929; de Villiers, 1929 b), and
has been tentatively suggested for Zachaenus parvulus by Lutz (1944). In
Nectophrynoides tornieri and Rhinoderma danvinii the spiracle is so small
and inconspicuous that high magnification and careful handling are re-
quired in order to locate it. Consequently, it is suggested that a spiracle
may prove to be present in some of the forms listed above.
2. Modification of the Mouthparts
The mouthparts of frog tadpoles are a highly complex mechanism that
has undergone many remarkable evolutionary changes. In species having
non-aquatic development, there is a trend toward loss of the typical larval
buccal structures {vide Lutz, 1948), but the degree to which this has been
accomplished differs widely between different species. Study of these
modified types of mouthparts suggests not only some features relating to
sequence of loss of structures, but also some possible clues to the origin of
certain parts of the buccal apparatus.
In the simplification of the mouthparts in non-aquatic larvae, the end
result may be structurally similar in forms having no close taxonomic
affinity. That similarity should, however, be interpreted with reference
to the type of larval mouthparts occurring in related free-swimming tad-
poles. For example, the absence of beaks and labial teeth in embryos of
Pipa pipa does not necessarily mean that they have been secondarily lost
in the course of modifying the life history of this species, for these struc-
tures are likewise absent from all known free-swimming pipid tadpoles.
Similarly, beaks and labial teeth are not known to occur in aquatic larvae
of the Microhylidae, and would therefore not be expected in terrestrial
embryos of microhylids.
Loss of the labial teeth in relation to direct development involves both
the disappearance of the rows of pigmented labial teeth themselves and
the tooth ridges upon which they are borne. Histologically, the teeth, and
probably also the beaks, may remain after there is no longer any readily
observable external indication of their presence. Noble (1927) figured and
discussed the small non-horny labial tooth remnants revealed in sections
through the lip of larval Rhinoderma darwinii. The jaws of Rhinoderma
are typically larval in form and proportions. They tend to be sheathed
with an opaque white layer which, although not hardened, is visually dif-
ferent from adjacent surfaces. The internal jaw structures of Rhinoderma
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have not yet been described. The head contours are typically larval,
however, especially the region of the processus muscularis of the quadrate
and the orbitohyoideus muscle, as can be seen through the partly trans-
parent skin (CNHM 3684).
In non-aquatic embryos in which the beaks and labial teeth are no
longer externally apparent, more or less well-defined traces of the larval
’ips may still remain. Rhinoderma darwinii (CNHM 3684) has nearly
typical lips with normal marginal papillae, and across the lower lip are
either two or three papillose, fringe-like, remnants of labial tooth ridges.
In Hemiphractus divaricatus (UMMZ 92106), the original larval lips are
reduced to a narrow papilla-edged labial fold along the sides of the upper
jaw and continuing along the base of the lower jaw for a varying distance,
reaching the symphysis in some specimens. In this form the upper jaw is
greatly simplified. The small lateral papillae are borne directly on the
edge, the separate dermal margin (upper lip) having been lost. In a series
of embryos of Cryptobatrachus fuhrmanni (UMMZ 47409), a still further
reduction is found, and all that remains of the external mouthparts is a
very short labial fold bearing a few papillae laterally and none at all on
its very low median continuation along the lower jaw. Morphologically,
this is a further step in the trend toward reduction of the mouthparts
shown by Hemiphractus divaricatus.
In Nectophrynoides tornieri the maximum development of external
mouthparts in the material examined is shown in specimens (MCZ 12513)
having a short fleshy fold along the base of the lower jaw, clearly a vestige
of the lower lip of typical aquatic tadpoles (PI. 3, fig. 1). It is not equally
well developed in all specimens in this series, and is probably both transi-
ent in the individual and of variable occurrence in the species. In Eleu-
therodactijlus this morphological trend has apparently reached its logical
conclusion, for no larval lip structure has been demonstrated in this genus
(see especially Lynn, 1942). It is probable, however, that the peculiar
horny egg tooth in Eleutherodactylus embryos is a specialized derivative
of the upper beak of a typical aquatic tadpole, judging from its position
and from Lynn’s (1942) note on its histological structure in E. nubicola.
The greatly reduced lip rudiments in embryos such as Hemiphractus
are structurally much like the simple labial folds of pipid tadpoles. In
these tadpoles, in turn, there is a close resemblance to the type of labial
folds characterizing salamander larvae and many of the fishes. At this
point, the factual basis for comparison becomes rather tenuous, but the
temptation cannot be resisted to suggest that the origin of the complex
1949
Orton: Larvae of Nectophrynoides
267
lips of the specialized aquatic tadpole can be traced to the simple lateral
labial folds that are of such widespread occurrence among the lower
vertebrates. The convergent development of complex lips in certain
South American catfishes and stream-dwelling Asiatic cyprinids provides
supporting evidence of a possible homology of these structures in
amphibians and fishes. Young larvae of the Australian lungfish possess
labial folds very similar in form and position to those of salamander
larvae.
Very little has been published on the internal anatomy of the jaw region
of non-aquatic larvae, but the scant published data that I have found,
together with the brief exploratory studies made during my doctoral re-
search at Michigan, suggest that in these specialized forms the cartilages
and muscles of the jaw region tend to become simplified. This would, of
course, be expected, in view of the reduction of other larval structures in
direct development. In some forms the jaw pattern is essentially that of a
typical tadpole. The general trend towards simplification seems to include
the following features: basal part of the quadrate shortens, returning the
suspensorium to more nearly the adult position {Cryptohatrachus, Necto-
phrynoides); Meckelian and lower labial cartilages lose their movable
articulation with each other and become a more or less continuous band of
cartilage {Cryptohatrachus, Nectophrynoides) ; the upper labial cartilage
is reduced in size and distinctness, and eventually becomes indistinguish-
able from the general cartilage field of the trabecular region ; the quadrato-
cranial commissure fails to develop in some forms, including Eleuthero-
dactylus mihicola (Lynn, 1942) ; the processus muscularis of the quadrate
becomes greatly reduced in size but, with its orbitohyoideus muscle, it
tends to retain a typically larval appearance {e.g., in Pipa pipa, Cry-
ptohatrachus fuhrmanni, Nectophrynoides tornieri); it is absent in Eleu-
therodactylus nuhicola.
Suppression of larval characters is more nearly complete in Elezi-
therodactylus than in any other frogs with non-aquatic development that
have been reported thus far. Closely related genera that are known or
suspected to have non-aquatic larvae, especially Syrrhophus, Tomodacty-
lus, and Microhatrachylus, should provide valuable comparative data
when their early stages are discovered.
The occurrence of direct development in frogs of very diverse taxonomic
relationships indicates that it is not a single phylogenetic trend, but rather
is a general tendency that has evolved independently in unrelated stocks.
Thus far it is known in a relatively small number of species, but these are
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VOL. 31
distributed through 10 of the 13 families^ that are currently considered
valid.
It is worthy of emphasis that among tadpoles there is evidence of
evolutionary trends of opposite nature. In one direction is the important
general tendency toward elimination of the tadpole stage through direct
development, while in the opposite direction is the widespread tendency
toward ever greater complexity of the tadpole stage, as in the extremely
specialized larvae of Hyla daresignata, Rana rugulosa, Staurois, and many
others. Although this is on a small taxonomic scale (a single order), it
appears to be an excellent example of the random nature of evolutionary
trends.
Summary and Conclusions
Several early stages in the development of the ovoviviparous frog,
Nedophrynoides tornieri, are briefly described and compared with similar
stages of N. occidentalis reported by Angel and Lamotte (1944). Although
the embryos of tornieri are highly modified in many respects, they develop
numerous tadpole characters, and in their structural plan as a whole they
are definitely larval. Small external gills, gill arches, and gill slits develop;
typical opercular folds form and enclose the gill region and foreleg anlagen ;
a spiracle develops; a rudimentary lower lip is present, at least in some
specimens; and there is a typical larval arrangement of internal jaw struc-
tures. Other details of body and tail structure are also mentioned.
Discrepancies between larval characters of tornieri reported here and
those of occidentalis described by Angel and Lamotte (1944) are at least
in part due simply to the availability of earlier stages of the former
species, in which the development of the jaw and branchial regions can
be more clearly traced. The internal position of the forelegs in occidentalis
indicates that in this species, as in tornieri, larval opercular folds and at
least a transitory spiracle develop.
The problem of direct development in frogs is discussed, with a de-
tailed account of modifications of the operculum and spiracle and the
mouthparts in non-aquatic larvae. The process of opercular develop-
ment and spiracle formation is compared in typical aquatic tadpoles and
in embryos of forms with direct development. It is concluded that the
structures concerned and their anatomical relations are the same in both
^ Examples known in Pipidae, Microhylidae, Ascaphidae, Pelobatidae, Lep-
todactylidae, Bufonidae, Atelopodidae {Rhinoderma) , Dendrobatidae (Sminthillus) ,
Hylidae, and Ranidae; probably occurs in some Rhacophoridae ; unknown in
Rhinophrynidae and Discoglossidae.
1949
Orton: Larvae of Nectophrynoides
269
life history types. Successively more simplified departures from the
typical process are traced in several species having direct development, in-
cluding Nectophrynoides tornieri, Gastrotheca sp., Cryptobatrachus fuhr-
manni, Hemiphractus divaricatus, and Eleutherodactylus sp. When these
forms are studied in morphological sequence, it is seen that they conform
to a common developmental pattern. The complex dual origin of the tad-
pole operculum is discussed; the dermal fold above the foreleg bud in
Eleidherodactylus is considered to be a remnant of the postero-lateral, non-
hyoidean, part of the composite operculum.
In the non-aquatic larvae of frogs with direct development, there is a
tendency toward simplification of the mouthparts. The beaks and labial
teeth fail to harden or to form pigment, and eventually fail to develop at
all; the lips and papillae are small and of simplified structure, and in
extreme cases become reduced to narrow labial folds. The internal parts
of the jaw apparatus tend to retain a larval pattern after the external
structures have been lost, but even the jaw cartilages and muscles may
by-pass the larval plan (as in Eleidherodactylus). It is suggested that the
horny egg tooth of Eleidherodactylus was probably derived from the upper
beak of the aquatic tadpole.
Comparison is made between the basic lip structure of tadpoles and
the closely similar labial folds of salamander larvae and certain fishes,
and it is suggested that these structures are probably homologous.
In direct development, the tadpole may be entirely lost ecologically,
but a substantial part of the larval structural pattern usually develops.
Thus, a tadpole stage, with a more or less extensive metamorphosis, is
still present anatomically even though all outward evidence of it dis-
appears before hatching occurs.
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VOL. 31
Literature Cited
Angel, F.
1943. Description d’un nouvel Amphibien anoure, ovo-vivipare, de la
Haute-Guinee Francaise. Bull. Mus. Paris, ser. 2, 15:167-169.
Angel, F., and M. LaMotte
1944. Un Crapaud vivipare d’Afrique occidentale, Nectophrynoides
occidentalis Ange\. Ann. Sci. Nat. Zool., ser. 11, 6:63-89, figs. 1-21.
Barbour, T., and A. Loveridge
1928. A comparative study of the herpetological faunae of the Uluguru
and Usambara Mountains, Tanganyika Territory, with descrip-
tions of new species. Mem. Mus. Comp. Zook, 50:85-265, pis. 1-4,
Brauer, a.
1898. Ein neuer Fall von Brutpflege bei Froschen. Zool. Jahrb., Abt. f.
Syst., 12:89-94.
Hewitt, John
1919. Anhydrophryne rattrayi, a remarkable new frog from Cape Colony.
Rec. Albany Mus., 3:182-189.
Krefft, P.
1911. tiber einen lebendgebardenden Froschlurch Deutsch-Ostrafrikas
{Nectophryne tornieri Roux). Zool. Anz., 37 :45 7-462.
Lutz, Bertha
1944. Biologia e taxonomia de Zachaenus parvulus. Bol. Mus. Nac.,
Zool. ser., no. 17, pp. 1-66, pis. 1-14.
1947. Trends towards non-aquatic and direct development in frogs.
Copeia, 1947 (4): 242-252, pis. 1-2.
1948. Ontogenetic evolution in frogs. Evolution, 2:29-39.
Lynn, W. Gardner
1942. The embryology of Eleutherodactylus nubicola, an anuran which
has no tadpole stage. Cam. Inst. Wash. Publ. No. 541, pp. 27-62,
pis. 1-5, figs. 1-40.
Noble, G. K.
1926, An analysis of the remarkable cases of distribution among the
Amphibia, with descriptions of new genera. Am. Mus. Nov., No.
212, pp. 1-24, figs. 1-9.
1927. The value of life history data in the study of the evolution of the
Amphibia. Ann. N. Y. Acad. Sci., 30:31-128, pi. 9, figs. 1-30.
1949
Orton: Larvae of Nectophrynoides
271
Power, J. H. and W. Rose
1929. Notes on the habits and life histories of some Cape Peninsula
Anura. Trans. Roy. Soc. South Africa, 17:109-115.
Roux, J.
1906.
Synopsis of the toads of the genus Nectophryne B. & P., with
special remarks on some little known species and description of a
new species from German East Africa. Proc. Zool. Soc. London,
(1906): 58-65, pi. 2.
Tornier, G.
1905. P seudopJiryne vivipara n. sp. ein lebendig gebarender Frosch. Sitz.
K. preuss. Akad. Wissensch., Berlin, (1905): 855-857.
De Villters, C. G. S.
1929a. Some features in the early development of Breviceps. Ann. Trans-
vaal Mus., 13:142-151.
1929&. The development of a species o( Arthr ole ptella horn Jonkershoek,
Stellenbosch. South Afr. Journ. Sci., 26:481-510.
272
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VOL. 31
EXPLANATION OF PLATE 1
Several stages in larval development of
Nectophrynoides tornieri (lateral views).
Fig. 1. “Stage 1.” Total length ± 5.5 mm. Small external gills present; operculum
not yet distinct; foreleg bud external.
Fig. 2. “Stage 2.” Total length + 6 mm. Operculum well developed, partly en-
closes foreleg bud.
Fig. 3. “Stage 3.” Total length + 6 mm. Spiracle formation completed, foreleg
bud internal.
Fig. 4. “Stage 4.” Total length 10 mm. Front leg visible through thin skin of
closed operculum.
Fig. 5. “Stage 5.” Total length + 11 mm. Metamorphosis has begun; front leg
has emerged from opercular cavity; mouth enlarging.
EG, external gill; FL, front leg; S, spiracle.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 9
Plate 1
FL
5
i
J
274
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 2
Development of operculum and spiracle in
larval N ectophrynoides tornieri (ventral views).
Fig. 1. Opercular folds distinct; gill arch region concealed, but both front leg
buds still exposed.
Figs, 2-4. Stages in closure of operculum. Fig. 2 shows slight advance beyond
condition in Fig. 1 ; in Figs. 3 and 4 right foreleg bud is entirely covered,
but left foreleg bud remains visible through wide gap in operculum.
Fig. 5. Operculum closure completed, spiracle formed; both foreleg buds con-
cealed.
E, eye; FL, foreleg bud; OA, opercular fold (anterior part); OL, opercular
fold (lateral part); S, spiracle.
ANNALS, CARNEGIE MUSEUM, Vol. 31 Art. 9 Plate 2
5
;•
276
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 3
Anatomy of jaw region in
larval Nectophrynoides tornieri
Fig. 1. Maximum development of external mouthparts. Jaw arrangement con-
forms closely to typical larval pattern, but accessory labial structures are
lacking except for narrow rudiment of lower lip. “Stage 3,” total length
± 6 mm.
Fig. 2. Jaw region dissected (lateral view). Jaw cartilages and muscles have dis-
tinctly larval arrangement. Position of right eye indicated. “Stage 4,”
total length 10 mm.
Fig. 3. Jaw region dissected (antero-lateral view). Muscles of adductor mandibulae
group conform to typical larval pattern. Right eye removed. “Stage 4, ’
total length 10 mm.
Fig. 4. Lower jaw and throat di.ssected (ventral view). Throat muscles very
small, but follow typical larval arrangement. “Stage 4,” total length
10 mm.
AMA, M. adductor mandibulae articularis; AMP, M, adductor mandibulae
posterior longus; AMS, M. adductor mandibulae subexternus; CA, M. cerato-
hyoangularis; CH, ceratohyal; E, eye; EC, ear capsule; FL, front leg bud; GA,
gill arch; GH, M. geniohyoideus; GS, gill slit; IG, internal gill rudiments; IM, M.
intermandibularis; L, rudimentary lower lip; MC, Meckel’s cartilage; N, nostril;
OH, M. orbitohyoideus; PMQ, processus muscularis of quadrate; PT, M. ptery-
goideus; SA, M. suspensorioangularis; SH, M. subhyoideus., UJ, side of upper jaw.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 9
Plate 3
4
i
1
'A'.X
ART. 10. DESCRIPTION OF A NEW RACE OF THE
SALAMANDER PSEUDOBRANCHUS STRIATUS (LE CONTE)
By Coleman J. Coin
University of Florida; Gainesville, Florida
AND
John W. Crenshaw, Jr.^
Emory University Field Station; Newton, Georgia
The population of Pseudobranchus that occurs in northern and western
Florida and southwestern Georgia appears to be a homogeneous one which
differs from both Pseudobranchus striatus striatus and P. s. axanthus and
we propose for it the name,
Pseudobranchus striatus spheniscus, new subspecies
Type: Carnegie Museum, no. 29,015, adult female, collected on July
4, 1948, by George B. Rabb and James E. Mosimann.
Type locality: Seven miles south of Smith ville, Lee County, Georgia.
Paratypes: Twenty-two, as follows:^ Florida; CM 21,440-41, from 30
miles north of Lake City, Columbia County; CM 21,466, from 12.2 miles
west of Wewahitchka, Gulf County; DBUF 1,855 (4 specimens), from
Lake lamonia, Leon County; DBUF 52, from Tallahassee, Leon County;
and CM 20,160 (2 specimens), from 5.4 miles south of Telogia, Liberty
County. Georgia; CAS 15,246 (3 specimens) from Mossy Pond, western
Baker County; USMN 128,140-44 and UMMZ 99,390 (3 specimens) from
Putney Pond, northwestern Baker County; and AMNH 34,626, from
Lakeland, Lanier County.
1 We are indebted to Doris M. Cochran, C. M. Bogert and M. Graham Netting
for the loan of material, to Horton H. Hobbs and George B. Rabb for aid in col-
lecting, and to Melvin H. Goodwin for making possible the junior author’s work
at the Emory Field Station at Newton, Georgia.
2 AMNH, American Museum of Natural History; CAS, Chicago Academy
of Sciences; CM, Carnegie Museum; DBUF, Department of Biology, University
of Florida; UMMZ, University of Michigan Museum of Zoology; USNM, United
States National Museum.
A 62 mm. specimen from Berrien County, Georgia (originally USNM 62,095),
was examined by the senior author several years ago, and although it is in poor
condition and somewhat faded, enough of its characters could be distinguished to
indicate that it is representative of this new race. It has since been sent on ex-
change to the Institute Miguel Lillo, Tucuman, Argentina.
277
Issued November 21, 1949.
1 raw
278
Annals of the Carnegie Museum
VOL. 31
Diagnosis: A slender Pseudobranchus with a narrow, wedge-shaped
head ; the lateral stripes are distinct and tan in color, tending to continue
forward through the eye to the tip of the snout. It differs from striatus
striatus in its more slender body and narrower head. The lateral stripes
are narrower and the coloration more somber. From s. axanthus it differs
in having the lateral stripes tan rather than gray and tending to continue
through the eye to the tip of the snout, and in having the head narrow
and wedge-shaped rather than broad and truncate.
Description of type: Head elongate, sides of head nearly straight but
tapering anteriorly to region of eyes, thence tapering more abruptly to a
sharply rounded snout; no pronounced swelling in parietal region. Head
acute in profile, slightly arched. Mouth small, ventral in position, pos-
terior angle of mouth separated from a line dropped from anterior border
of eye by about the diameter of an eye. Outline of upper jaw convex as
seen from side, due to pendulous upper lip; upper jaw projecting beyond
lower. No can thus rostralis. Loreal region convex. Nostril, an elongate,
longitudinal slit, ventrolateral in position, its anterior corner about on a
line with anterior portion of mouth, not visible from above. Internarial
distance (ventrally) about twice diameter of eye. Eye small, not pro-
truberant, without eyelids but completely covered by a thin membrane,
its diameter about 2/5 its distance from posterior corner of nostril.
Interorbital distance about 4 times diameter of eye. Head width 14.0
times in body length; head length 7.68 times in body length.
Body slender, nearly circular in cross section; no constriction in neck;
a narrow mid -dorsal groove ; a shallow, longitudinal depression along cen-
ter of lateral stripe ; a shallow mid-ventral groove extending posteriorly to
base of tail. Costal grooves 34, distinct, connecting across abdomen and
extending well up on sides. Vent a short longitudinal slit.
Tail 1.5 times in snout-to-vent length, ovoid in vertical cross section
at base; gradually becoming more and more compressed to flat tip. A
narrow fin on distal 2/3 of dorsal surface of tail and a similar fin on distal
34 of ventral surface of tail, fins connecting around tip of tail. Fins never
so broad as tail musculature.
Forelimbs present, minute. Fingers three, short, stout, 2-1-3 in order
of decreasing length, not webbed. Minute horny caps covering tips. No
metacarpal tubercles present.
External gills three. Gills on both sides compacted and covered by
an investing integument so that they do not hang free; extending pos-
teriorly on both sides to base of forelimbs.
1949 Coin and Crenshaw: New Race of Salamander 279
Tongue small, well back in mouth, its acutely-pointed anterior-half free.
A black horny sheath present at anterior end of each jaw, that of upper
jaw very short, oval in shape and arched, that of lower jaw a well-developed
transverse ridge, curving backward at its outer ends. Lower jaw with
rather long, recurved, sharply pointed, well-separated teeth, arranged in
two rather irregular rows on the anterior portion of each mandible. Two
elongate patches of prevomerine teeth, beginning slightly behind horny
sheath in upper jaw. Internal nares long, longitudinal openings slightly
diagonal in position, each deeply buried in a fold of tissue that is ap-
parently capable of tight closure.
Measurements of type (in millimeters): Head length (tip of snout to
base of foremost gill), 9.5; body length, 73.0; tail length, 55.5; total
length, 138.0; forelimb length, 4.0; head width (maximum), 5.2; body
width (maximum), 5.4; body depth (maximum), 5.5.
Coloration of type (preserved) : Top of head and dorsum dark brownish
gray. Head without any distinct markings but with a faint indication of a
light stripe from nostril to base of gill on each side. A yellowish brown
stripe about 1 mm. in width extending along each side from base of gill
posteriorly to base of tail where it becomes broken and continues as an
indistinct broken line to tip of tail ; another more yellowish stripe extends
from base of arm on each side posteriorly to the region of the vent. This
ventrolateral stripe extends on to the ventral surface of the forelimb. The
distance between the lateral and ventrolateral stripes on each side is
about equal to the width of the two stripes. The area between them is
grayish black. The venter is a dark gray with indistinct scattered light
spots of yellowish fuscous. There is no discernible mid-dorsal stripe but
the dorsal margin of the tail fin is edged with fuscous.
Variation: Most of the variation in the typical series seems to be due to
age and state of preservation. Eleven of the specimens are definitely im-
mature. The narrow, wedge-shaped head is the most striking characteristic
and is apparent in all specimens from the smallest to the largest. In the
largest specimen, the type, the head stripes are not so prominent as they
are in the other specimens of the series, but nevertheless more prominent
head stripes seem to be characteristic of this race.
Remarks: This new race is apparently much more difficult to collect
than is axanthus. The senior author has on numerous occasions taken
more axanthus in a single afternoon from some of the hyacinth-filled
ditches and marshes around Gainesville, Florida, than there are specimens
of spheniscus in the type series. This is certainly not due to intensity of
280
Annals of the Carnegie Museum
VOL. 31
collecting alone since he has made several trips to northwestern Florida
for the express purpose of searching for spheniscus but has been able
to collect only one specimen. Intensive collecting in Baker County,
Georgia, by the junior author, in a region where spheniscus is known to
occur, has produced only eleven specimens. We believe this difference may
be due in part to a difference in the habits of the two forms. P. s. axanthus
is abundant among masses of vegetation, especially in the roots of water
hyacinths, in the large marshes of peninsular Florida, but spheniscus
generally occurs in small to medium, shallow, limesink ponds, abounding
in pond cypress {Taxodium ascendens) and black gum (Nyssa biflora).
In so far as we know, the water hyacinth is not found in much of the area.
It certainly does not occur in any of the ponds in Baker and Lee Counties,
Georgia, from which specimens of P. s. spheniscus were collected. Although
much time was spent in seining masses of Cahomha and Myriophyllum
just beneath the surface, no specimens were collected in this manner.
All but one of the southwestern Georgia specimens were collected by
seining and examining bottom detritus. The other was unearthed from the
mud bottom of a dried-up pond. Several of the Florida specimens were
taken by Dr. Horton H. Hobbs while collecting crayfishes with a dip
net. From our limited knowledge of this form we suspect that it has a
tendency to remain burrowed in decaying bottom vegetation.
ART. 11. A REVIEW OF THE ELVEELACE.T (FUNGI)
OF WESTERN PENNSYEVANIA
By LeRoy K. Henry
(Plates 1-2)
The family ElYellacea:,To which the morels and their relatNes belong,
is placed in the order Pezizales of the Ascomycetes. The fungi in this
order are commonly called cup fungi (Discomycetes) l^ecause their fruit-
ing bodies are usually disk- or cup-shaped. However, the fruiting bodies
of the members of this family are fleshy and consist of a distinct stipe
(stem) and a pileus (cap). The pileus is located at the top of the stipe,
with its margin either free or partly attached to the stipe, and it may be
bell-shaped, saddle-shaped, or subglobose. The hymenium (spore-bearing
surface) is e^^en, ridged and pitted, convoluted, or irregularly folded, and
it covers the outer surface of the cap. The stipe is usuall>' hollow and may
be very slender or short and stout, with the surface even, pitted, ridged,
or fluted.
These are chiefly spring fungi, appearing in open woods, orchards, or
along flood-plains, in April or Ma^o
The morels and elvellas are edible, with the exception of one doubtful
species known as the false morel and formerly called Gyromitra escidenia,
but now identified with Elvella injiila. This false morel can readily be
distinguished from the true morels b}^ the somewhat convoluted character
of the cap and its chestnut-red coloring. There has been disagreement in
the literature concerning the edibility of this mushroom; some myco-
phagists claim no ill effects from eating it and others report that they were
poisoned by it. Individual differences may account for these conflicting
statements. Also, the amount of poison ma}-^ vary in plants from different
localities, or there may be two closely related species involved, the one
edible and the other poisonous. The true morels make a very tasty dish,
provided one can find enough of them at any one time.
This famil}^ contains fiv^e genera in eastern North America, three of
which are represented in the Pennsylvania Herbarium of Carnegie Mu-
seum. The majority of the Herbarium collections were contributed by
D. R. Sumstine, O. E. Jennings, and L. K. Henry.
For this paper, as the eastern boundry of WYstern Penns^dvania, I have
arbitrarily chosen the eastern borders of Potter, Clinton, Centre, Hunting-
don, and Fulton counties.
281
FEB 1 ISSO
Issued December 19, 1949.
282
Annals of the Carnegie Museum
VOL. 31
Key to the Genera of the Elvellace.-e
Outer surface of pileus consisting of deep or shallow pits formed by longi-
tudinal or transverse ridges Morcliella
Outer surface of pileus even, ribbed, or convoluted.
Pileus bell-shaped, edge free from stipe Verpa
Pileus lobed, irregularly subglobose, or saddle-shaped; surface even or
convoluted Elvella
Morchella (Morels)
Pileus subglobose or elongated, blunt or acute at tip, the margin
closel}' joined to the stipe at its base in all but one species. Surface of
pileus traversed by irregularly branching ribs, forming rounded or
elongated pits which are lined with the yellow to brown hymenium. Stipe
cylindrical, usually lighter in color than the pileus, often enlarged at the
base. All of our six species are edible.
Key to Morchella
Base of pileus free from the stipe; plant 4-5 inches high; cap bell-shaped,
up to 3 cm. long; ribs whitish; pits 5-10 mm. in diameter, yellow
within; stem 8-10 cm. long, white or yellowish, irregularly pitted
at base and tapering upward AI. hybrida
Base of pileus joined to the stipe.
Ribs of the pileus much darker than the interior of the pits; pileus
narrowly conic and acute at apex, 2-5 cm. long; plant 2-3 inches
high; pits elongated, 4-10 mm. long, yellowish within; ribs black
on edge; stipe of uniform thickness AI. angusticeps
Ribs of pileus similar in color to the interior of the pits (yellowish)
or of a lighter color.
Pileus large, 4-8 cm. or more long at maturity.
Pits large, irregular, up to 1 cm. broad, usually shallow; plant 7-8
inches high ; ribs thin ; stipe stout, distinctly enlarged at base,
10-11 cm. long AI. crassipes
Pits small, deep; ribs thick; stipe slightl}’ enlarged and irregularly
pitted at base.
Plant 2-4 inches high; pits irregular, 5-10 mm. in diameter; ribs
irregularly branching; pileus subglobose, generally a little
longer than broad AI. esculenta
1949
Henry: ELVELLACE.^i: of W'estern Pennsylvania
283
l^lant 3-5 inches high; pileus elongated or strongly attenuated
upwards; pits elongated, 5-10 mm. long; ribs more or less
longitudinally disposed M. conica
Pileus small, not exceeding 2-3 cm. in length at maturity ; pits elon-
gated, ashy to blackish within; ribs longitudinall}’' disposed,
whitish; stipe whitish, irregularl pitted M. deliciosa
Morchella angusticeps Peck (Plate 1, fig. 4)
Allegheny County: Coraopolis. Washington County: 5 mi. s.e. of Houston.
Rare.
Morchella conica Pers. (Plate 1, fig. 5)
Allegheny County: Frick Park, Pittsburgh; 4 mi. e. of Monongahela.
Armstrong County: Kittanning. Beaver County: 1 mi. n. of Alechanicsburg;
Raccoon Creek region; woods at intersection of Beaver-Conway and
Sewickle\^-Rochester roads. Erie County: Presque Isle. Fayette County:
Ohiopyle. Greene County: 10 mi. s.e. of Waynesburg. Lawrence County:
near New Castle. Somerset County: Ursina. Venango County: 3 mi. n. of
Lisbon. IVashington County: Charleroi. Westmoreland County: near Ligo-
nier; Hillside.
Morchella crassipes (Vent.) Pers. (Plate 1, fig. 3)
Allegheny County: near Saunders; flood-plain of Lyons Run; yard in
Pittsburgh. Armstrong County: Kittanning. Cambria County: near Cresson.
Centre County: 1 mi. s. of Boalsburg. Crawford County: near Linesville;
near Hartstown. Washington County: Washington car-line near Center
Church; near New Lagle. Westmoreland County: 3 mi. s.e. of Rector;
Hillside.
Morchella deliciosa Fries (Plate 1, fig. 2)
Allegheny County: Warden Mine region opposite Sutersville; 2.5 mi.
n.e. of Ambridge at Turke}^ Foot. Beaver County: 2 mi. above mouth of
Raccoon Creek. Butler County: under a Peony, 4 mi. n.e. of Harmony.
Crawford County: near Hartstown. Washington County: near Riverview.
Westmoreland County: Kiski Campus near Saltsburg.
Morchella esculenta (L). Pers. (Plate 1, fig. 6)
Allegheny County: Frick Park, Pittsburgh. Armstrong County: Kittann-
ing. Butler County: 4 mi. n.e. of Harmony; near Renfrew; Winfield
284
Annals of the Carnegie Museum
VOL. 31
Junction. Erie County: Presque Isle. Fayette County: Ohiopyle. Somerset
County: near Ursina. Washington County: 5 mi. s.e. of Houston; vicinity
of Hanlin Station. WestmoreJand County: Ligonier; Hillside.
Morchella hybrida (Sow.) Pers. (Plate 1, fig. 1)
Allegheny County: near \Mldwood; 2 mi. n. of Saunders; near Pitts-
burgh; Frick Park, Pittsburgh. Armstrong County: Kittanning. Butler
County: Marwood. Crawford County: Hartstown. Fayette County: Ohiopyle.
Indiana County: 2 mi. n.e. of Clarksburg. Washington County: Charleroi;
Van Voorhis. Westmoreland County: Hillside.
Verpa
Pileus bell-shaped, yellow to brownish, white beneath, often with a
reflexed margin, free from the stipe, 2-3 cm. long by 1-2 cm. in diameter.
Spore-bearing surface (hymenium) folded into longitudinal and branching-
ribs, yellow to brownish. Stipe nearly cylindrical, hollow, even, white or
slightly cottony, 6-8 cm. long.
Verpa bohemica (Krombh.) Schrdt. (Plate 2, fig. 6)
Beaver County: Pine Grove, J. A. M. Stewart; Raccoon Creek State
l^ark. L.K.H. Rare.
El VELA (Helvella)
Pileus mitre-shaped, saddle-shaped or subglol)Ose, e\’en or irregularh'
con\’olute, the margin reflexed and free or more or less joined to the stipe.
Stipe slender or stout, even, pitted or strongly lluted, white, yellow or
smok\L None of the five species in our region can be considered common.
Key to Elvela
Stipe distinctl}’ lluted, stout and usually enlarged at the base; surface
of pileus even or convoluted.
Pileus and stipe remaining cream or yellowish E. crispa
Pileus becoming dark brownish-black; stipe yellowish, becoming
smoky E. mitra
Stipe not distinctly fluted but even or irregularly pitted.
Stipe very slender, usually not over 5 mm. in diameter; surface of
pileus even or more or less convolute.
Pileus yellowish to dark brown, 2-3 lobed, free from the stipe;
stipe yellowish, 5-10 cm. long E. elastica
1949
Henry: Elvellace.e of Western Pennsylvania
285
Stipe stout, usually 1-5 cm. in diameter; surface of cap more or
less convoluted.
Pileus 6-8 cm. broad, chestnut-red, mitre-shaped, saddle-shaped
or variously convoluted; stipe 6-8 cm. long, white to yellowish
even or more or less pitted; spore ellipsoid, smooth.
E. infula {G. escidenUi)
Pileus 5-12 cm. broad, chocolate-brown, irregularly lobed and
folded; stipe usually even, white, 8-13 cm. long; spores large,
rough, lusoid, strongly pointed E. iindenvoodii
Pileus 5-12 cm. broad, brown to brownish black, irregularly
lobed and folded, often reticulate; stipe deeply pitted, 8-10 cm.
long, whitish; spores large, rough and ellipsoid. .E. caroliniaua
Elvela caroliniana (Bose.) Nees. (Plate 2, fig. 4)
Butler County: Ribold; Criders Corners.
Elvela crispa (Scop.) Fries (Plate 2, fig. 1)
Armstrong County: Kittanning; across Buffalo Creek from West Win-
field. Cambria County: near Cresson. Warre}i County: s. of Tidioute.
Westmoreland County: near Rector.
Elvela elastica Bull. (Plate 2, fig. 3)
Bedford County: Sulphur Springs. Centre County: near Woodward.
Westmoreland County: 3 mi. s.e. of Rector.
Elvela infula SchaelT. [Gyromitra esculenta (Pers.) Fr.] (Plate 2, fig. 5)
Allegheny County: Kennywood Park; 3 mi. s.e. of Bakerstown. Butler
County: 4 mi. n.e. of Harmony. Venango County: 1 mi. n. of Lisbon.
Elvela underwoodii Seaver (£. brunnea Underw.) (Plate 2, fig. 2)
Allegheny County: Swissvale; Library; Kennywood Park; Pittsburgh.
Armstrong County: Kittanning. Butler County: near Saxonburg.
286
Annals of the Carnegie Museum
A'OL. 31
EXPLANATION OF PLATE 1
All figures are one-half natural size
Fig. 1. Morchella hyhrida (Sow.) Pers.
Fig. 2. Morchella deliciosa Fries
Fig. 3. Morchella crassipes (Vent.) Pers.
Fig. 4. Morchella angusticeps Peck
Fig. 5. Morchella conica Pers.
Fig. 6. Morchella esctdenta (L.) Pers.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Plate 1
P.E.
Uon<J
Art. 11
5
288
Annals of the Carnegie Museum
VOL, 31
EXPLANATION OF PLATE 2
All figures are one-half natural size
Fig. 1. Elvela crispa (Scop.) Fries
Fig. 2. Elvela underwoodii Seaver
Fig. 3, Elvela elasiica Bull.
Fig. 4. Elvela caroUniana (Bose) Nees
F IG. vS. Elvela infiila Schaeff.
Fig. 6. Verpa bohemica (Krombh.) Schrot.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 11
Plate 2
ART. 12. A NORTHERN RACE OF RED-TAILED HAWTC
By W. E. Clyde Todd
(Plate 1)
Buteo jamaicensis subsp. Red-tailed Hawk.
Buteo borealis (not of Gmelin) Couper, Canadian Sport, and Nat., 1, 1881, 80
(Mingan R., Quebec, nesting, fide Smyth). — Palmer, Proc. U. S. Nat.
Mus., 13, 1890, 261 (Alingan Is., Quebec, summer). — De Puyjalon, Hist.
Nat. a Tusage Chasseurs Canadiens, 1900, 420 ([Canadian] Labrador). —
AIacoun, Cat. Canadian Birds, pt. 2, 1903, 230 (Moose R., near James
Bay, Ontario, June, fide Spreadborough). — Macoun and Macoun, Cat.
Canadian Birds, ed. 2, 1909, 251 (Missanabie River, Ontario, fide Spread-
borough).— Williams, Canadian Field-Nat., 34, 1920, 123 (Hamilton
Rapids [Groundhog R.], Pike Creek [Mattagami R.], and Missanabie R.
[mouth], Ontario, August). — Baillie, Canadian Field-Nat., 43, 1929, 172
(Couper’s record). — Eidmann, Zoogeographica, 2, pt. 2, 1934, 241 (Matamek
region, Quebec, habitat).— Lewis, Canadian Field-Nat., 49, 1935, 114
(Couper’s record). — Ricker and Clarke, Contr. Royal Ontario Mus.
Zook, 16, 1939, 7 (L. Nipissing, summer).
“Red-tailed Hawk” Audubon (M. R.), Audubon Journ., 1, 1897, 394 ([Wapi-
tagun], Quebec, July). — Townsend, Bull. Geogr. Soc. Philadelphia, 11,
1913, 48 (Natashquan R. [40 4- nii. from mouth], July; Audubon and Pal-
mer’s records). — Eidmann, Verh. Orn. Ges. Bayern, 21, 1937, 161 (Trout
Lake and Matamek River region, Quebec, August- September).
Buteo borealis borealis Townsend, Auk, 30, 1913, 6 (Natashquan R.,
Quebec, summer; plum.). — Lewis, Auk, 39, 1922, 512 (Mingan and “Piashte
Bay” [Baie Johan Beetz], Quebec, June). — Snyder, Univ. Toronto Biol.
Ser., 32, 1928, 22 (Ghost R., L. Abitibi, Ontario, June).
Buteo jamaicensis borealis Godfrey, Nat. Mus. Canada, Bull. 114, 1949, 16
(Lakes Mistassini and Albanel, Quebec, June-September).
The Red-tailed Hawk is the New World representative of the European
Buzzard, Buteo buteo. It is one of the commoner Hawks of North America,
where it enjoys an extensive range, from the Isthmus of Panama north
to the limit of trees. Throughout this range it splits up into a number of
geographic races, whose inter-relationships are rendered all the more
puzzling because of an innate tendency in the species to dichromatism — a
tendenc}" which is manifest in varying degrees, and which greatly com-
plicates the discrimination of the several forms. Individual and seasonal
variations are other factors which must inevitably be taken into account.
Until rather recently the species has been known as Buteo borealis,
^The present account is an excerpt from the writer’s manuscript report on '
the birds of the Labrador Peninsula. It has been prepared to show the style of
treatment to be followed. Comment and criticism are invited.
289
ssued April 25, 1950.
I»AY 3
IQcrn
290
Annals of the Carnegie Museum
VOL. 31
from Falco borealis, Gmelin, 1788. The specific name was based on the
^h\merican Buzzard” of Latham (1781). Oddly enough, “Carolina” is the
only definite locality* these authors cite; this has now been restricted to
South Carolina — an incongruous type-locality for a supposedly northern
bird. On the same page, some lines before, Gmelin also named a Falco
jama icen sis, based on the “Cream-colored Buzzard” of Latham (1788),
said to be from Jamaica. His description is by no means clear, and many
authors have ignored this name as unidentifiable. This would be the safe
and sane course, but most recent authorities (including the American
Ornithologists’ Union Committee, following Peters, 1931) have chosen to
resurrect jamaicensis as the specific name for the Red-tailed Hawk.
Latham’s description may be construed to apply to an immature example
of this species in an albinistic condition, and in any event it could scarcely
pertain to ari}’ other species than the small Red-tailed Hawk of Jamaica
and certain other West Indian islands. There is indeed some justification
for ranking this insular form as a species distinct from the continental bird.
At one time I shared this view, but I now think that we shall have to con-
sider them conspecific. The unfortunate outcome of this decision is that,
following the rule of line anteriority, we are compelled to use the utterly
inappropriate name Biiteo jamaicensis borealis for a bird which was first
described from South Carolina.
To the critical and discriminating study made by the late Percy A.
Taverner (Vuctoria Mem. Mus. Bull. No. 48, 1927, 1-20, 3 pis; and Con-
dor, 38, 1936, 66-71), we are indebted for much light on this interesting
group. His object was to clarify the relationships and taxonomy" of the
forms found in Canada, and particularly in western Canada. He discussed
the western race calurus at some length, but it is interesting to note that
the latest authorities (Hellmayr and Conover, Field Mus. Zool. Ser., 13,
pt. 1, no. 4, 1949, 96-97) synonymize this name with borealis. After an
independent study of the series available to me I find myself in disagree-
ment with this conclusion. Aside from the greater tendency to melanism
in calurus, the race is characterized by its richer coloration as compared
with borealis. Taverner treated the eastern Red-tailed Hawk as a single
entit\y he had no suspicion that more than one race might be involved,
nor had other authors — and this in spite of the observed extent of varia-
tion. In view of the scarcity of breeding specimens from critical areas,
this is not surprising. During a recent study of the Red-tailed Hawks in
the collection of the Carnegie Museum I was interested to find that our
two specimens from near the northern limit of the species’ range stood out
from Pennsylvania breeding birds b>' easily discernible characters.
1950
Todd: New Red-tailed Hawk
291
Through the courtesy of Mr. W. Earl Godfrey of the National Museum of
Canada I have been permitted to examine three additional specimens
showing the same peculiarities. These five examples (breeding birds)
appear to represent a hitherto unrecognized race of this species, which may
be called
Buteo jamaicensis abieticola, subsp. nov.
Northern Red-tailed Hawk
Type, No. 57,348, Collection Carnegie Museum, adult female; Sainte
Margaret Falls (above Clarke City), Quebec, May 25, 1917; Olaus J.
Murie. (Taken with a set of two eggs.)
Siihspecific characters. — Similar to Buteo jamaicensis borealis of the
eastern United States and southern Canada, but underparts more heavily
streaked; throat and upper breast darker colored (more brownish, less
rufescent) ; upperparts (including wings externally) darker colored (more
blackish) ; and subterminal black band on tail averaging wider.
Range. — Apparently the spruce-fir belt of Canada, north (at least west
of Hudson Bay) to the limit of trees.
Comparison was first made with seven breeding specimens from western
Pennsylvania, but later these were supplemented, through the courtesy
of the authorities of the U. S. National Museum, by two specimens from
the vicinity of Washington, D. C., and one from Mount Mitchell, North
Carolina. The last comes from as near the type-locality of borealis as
any we can get, and it agrees closely with the Pennsylvania birds. It is
fair to presume, therefore, that the form represented by these specimens
is the one to which the name borealis rightfully belongs. None of the
recognized synonyms of borealis as commonly understood can possibly
apply to the northern race here described, since they all come from
“Carolina’’ or “Virginia,” and the descriptions and figures are not pertinent.
The characters apparent in the present race seemed, on first reading, to
be those given by Grinnell for his Buteo borealis alascensis (Univ. Cali-
fornia Pub. Zook, 5, 1909, 211), described from Glacier Bay, Alaska. Of
this form I have seen no Alaskan specimens, but through the courtesy of
the authorities of the Chicago Natural History Museum I have before
me seven specimens from British Columbia which presumably represent
this form correctly. When Grinnell described alascensis he compared it
with calurus, and very properly so, since in it the characters of cahirus
are merely carried to an extreme. The rufescent shading of the under-
parts is a marked character. In any case, alascensis has nothing to do
with the new race.
292
Annals of the Carnegie Museum
VOL. 31
Besides the type from Quebec, we have a June specimen from Quatre
Fourches Channel, Peace River Delta, Alberta, and I have seen three
more (National Museum of Canada) from Lac la Nonne, Alberta. These
fi\'e breeding specimens of abieiicola, coming as they do from widely
separated points in the north country’, suggest an extensive breeding range
for this form^ — a range probably co-extensive with the spruce and fir forests
of this area. Here the birds must be only summer residents, migrating
southward for the winter. If so, it would be reasonable to expect to find a
certain proportion of fall- and winter-taken specimens from the northern
United States referable to this race. Examination of a series of specimens
shot by gunners some years ago at “Hawk Mountain,” near Drehersville,
Pennsylvania, and preserved as skins by Dr. George M. Sutton, fully
bears out this expectation. These birds were migrants from the north,
and were actually taken in migration. Laid out side by side with season-
abh- comparable specimens from other parts, they show the characters of
abieticola to even better advantage than do the breeding adults. A few
of our fall and winter birds are indeterminate, and might be referred to one
form as easily as to the other, but the general dilTerence between the two
is well marked. In immature plumage, however, the difference is thus far
unsettled, in part because of lack of material. All we can say is, that in
young birds, probably northern-reared, the general coloration appears to
be darker and the markings on the underparts to be heavier, but there is
a great deal of variation in these respects in the specimens examined.
Measurements
No.
Sex
Locality
Date
Wing
Tail
Bill
Tarsus
57,348
9
Ste. Alargaret Falls,
Quebec
May 25, 1917
387
195
37.5
82
101,899
9
Quatre Fourches Chan-
nel, Alberta
June 8, 1927
407
210
37
83
21,263*
9
Lac La Nonne, Alberta
May 26, 1926
415
218
37
86
21,696*
9
7? ;?
July 20, 1926
422
224
37
86
21,250*
cf
?? M ??
May 26, 1926
360
216
37.5
82
23,819
9
Hardin Co., Iowa
Mar. 24, 1905
418
233
37
80
102,681
cT
Towanda, Pennsylvania
Nov. 17, 1925
354
196
34.5
77
102,686
cf
“Hawk Mountain,” Pa.
Oct. 22, 1927
353
190
36
79
102,687
cf'
7? yj
Oct. 22, 1927
344
192
35
75
102,688
9
yy yy ;?
Oct. 22, 1927
373
211
36
78
102,690
9
yy yy yy
Oct. 23, 1927
382
213
38
80
103,560
cf
yy yy yy
Oct. 22, 1927
363
204
34
74
^Collection National Museum of Canada.
1950
Todd; New Red -tailed Hawk
293
\Miile there is thus no significant difference in size between borealis
(as here restricted) and ahieticola, the above table of measurements re-
veals that Alberta specimens run a little larger than eastern birds. It
will be noted that the migrants taken in Pennsylvania agree in smaller size
with the type-specimen from the north shore of the Gulf of St. Lawrence.
A female from Iowa, taken on March 24, is so palpably an example of
ahieticola that I am constrained to believe that it must have been a mi-
grant on its way north.
In his “Labrador Journal,” Audubon records finding a tail-feather of a
young Red-tailed Hawk near Wapitagun, on the north shore of the Gulf
of St. Lawrence, in July, 1838. Had the record remained unconfirmed (as
it did for maii}^ years) it would be open to question, since it is just pos-
sible that this particular tail-feather might have come from an Osprey
instead. In 1881, the Canadian naturalist William Couper published, in
a rare and little-known journal which he edited, a circumstantial account
of the finding of a nest on the Mingan River by Sir Greville Smyth. “It
contained two young, which could feed themselves at the end of July.”
“There is no reason to question this” (H. F. Lewis). In the summer of
1890, William Palmer saw two on the Mingan Islands. The Audubon and
Palmer records were the only ones known to Townsend and Allen when
they wrote their list of Labrador birds in 1907. Later Townsend himself
met with the species on a tributary of the Natashquan River, some forty
miles from its mouth. Just one bird was seen, whose dark coloration he re-
marked at the time. “No nest could be found although the bird’s actions
suggested young.”
This brings us down to 1917, the year when our party from the Car-
negie Museum was camped just above the first falls on the Ste. Alargaret
river, ready to begin the ascent of that stream as the first stage in the cross-
ing of the Labrador Peninsula. A pair of Red-tailed Hawks was in evi-
dence across the river from our camp, and their nest was located on Alay
25. After lunch our whole party (five Indians, three whites) crossed over
to investigate. The nest was built of sticks and twigs, about fifty feet
up on a thirty-inch birch tree. The men built a blind of young spruce
trees below in which Mr. Murie concealed himself, while the rest of us left
the vicinity. Scarcely had we reached camp again before we heard the re-
port of his gun, and upon returning we found that he had brought down
the parent bird as she returned to the nest. Obviously she could not
count. The men felled an adjacent tree against the nest-tree, and Mr.
Murie climbed up and brought down the two incubated eggs the nest
contained. Thus was the type-specimen of the present new race obtained.
294
Annals of the Carnegie Museum
VOL. 31
On May 27 another individual was noted about nine miles upstream, and
on June 6 still another on the Grand Portage, beyond Lac au Poelon.
More recently, Harrison F. Lewis has published June sight records for this
species from Mingan and Baie Johan Beetz, and H. Eidmann one for the
Matamek region. I do not know the source of the English Bay record to
which A. C. Bent (Bull. 167, U. S. Nat. Mus., 1937, 162) refers. Thus far
the species has not been traced farther eastward along the north shore
than the mouth of the Little Mecatina River, where it was noted by Harold
S. Peters (MS.) on June 25, 1938. It is not known to reach Newfoundland.
For Anticosti Island there is only Joseph Schmitt’s notation that it is a
rather rare permanent resident — on its face a questionable statement.
The northern limit of the range of this species in the Peninsula remains
to be worked out. While we failed to find it on the Ste. Margaret River
beyond 50° 44^ N., I have a sight record from Gull Lake on the Hamilton
River at 53° N., July 17, 1939. From this point to Lake Mistassini and
Lake Albanel, where, according to W. Earl Godfre\^ it was the “most
frequently observed hawk” in the season of 1947, from June 20 to Sep-
tember 4, is a long jump. Unfortunately no specimens were taken. To the
westw’ard of Mistassini our party made four records in 1914. One was
noted along the railroad, eighteen miles west of the Bell River, on May
28, by O. J. Murie. One was seen on the Bell River near Kanikwanika
Island on June 2; several on Chensagi Lake on June 9; and one at the
fifth portage (west of Nemiskau) on the Rupert River on June 24. On
July 8, we saw one at East Main; this is the farthest north I have ever
observed it on James Bay. These are presumptive breeding records, as are
also some of those we have from the southern shores of James Bay and
from the rivers of northern Ontario, and which were made on sundry Car-
negie Museum expeditions. Spreadborough observed the species below
Moose Factory on June 6, 1896, and also on the Missanabie River. G. M.
Sutton saw two at Moose Factory on June 4, 1926 and one on June 13.
The species certainly does not nest on Moose Island, although it must do
so at no great distance. It was repeatedly observed in late August and
again in October 1923, by our party at several points along the Abitibi
River, and Williams (1920) saw individuals in August along the Mat-
tagami and Groundhog Rivers.
Spring records are also available for the Missanabie River. These were
made by our party in 1926, as follows: above Skunk Island, May 27, one;
Long Portage to Soweska River, May 31, three pairs and four single birds;
Soweska River (mouth) to about 50° 29' N., two. In 1912, I saw a single
individual about six miles east of Cochrane on May 21, and on September
1950
Todd: New Red-tailed Hawk
295
26, 1935, R. L. Fricke saw one along the road near Ramore. L. L. Snyder
has recorded the species from Lake Abitibi in summer. Although no speci-
mens appear to be available from this general region, I refer all these
records, south to Lake Nipissing, to the race herewith described — this on
geographical considerations.- Probably the Churchill record given by
Taverner and Sutton (1934) belongs here also, since it is this form that
occurs in northern and central Alberta, as actual specimens show.
Dr. Arthur C. Twomey tells me that in the spring of 1942 he discovered
a nesting pair of Red-tailed Hawks about fifteen miles southeast of
AkIa^'ik, in the Mackenzie Delta, but was unable to secure them. Pre-
sumably they also were of the present race, the range of which extends
far to the northward in this part of Canada.
The general habits of this northern race of Red-tailed Hawk are the
same as those of the eastern bird, and its eggs are indistinguishable. Town-
send’s remarks on a melanistic individual he observed on the Natashquan
River (1913) raise the question whether melanism is confined to the race
calurus. Compare also, in this connection, the remarks of Rand (Auk,
65, 1948, 424).
- Since the above was written, the Royal Ontario Museum of Zoology has sent
me for examination six specimens from western Ontario (Port Arthur, Amyot,
Schreiber) and eastern Manitoba (Deer Lodge, Portage la Prairie, Lake St. Martin).
Three were taken in the breeding season, and three in October. Despite some
obvious variation, and a suggestion of krideri influence in the Lake St. Martin
bird, all these are clearly referable to the new race upon comparison. In the
Deer Lodge specimen, dated October 15, the characters reach an extreme.
296
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 1
Left-hand row: BiUeo ja^naicens is abieticola.
Right-hand row: Buteo jamaicensis borealis.
Upper engraving: breeding specimens.
Lower engraving: autumn specimens.
The type of BiUeo jamaicensis abieticola is the lowermost bird
in the left-hand column of the upper engraving.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 12
Plate 1
I
-i
, y *
ART. 13. A NEW RACE OF PAROQUET OF THE SPECIES
ARATINGA ASTEC FROM THE REPUBLIC OF HONDURAS
By Arthur C. Twomey
A comprehensive survey of the bird life of the Republic of Honduras was
begun by the author in 1947 and 1948 and will continue until the country
can be adequately covered.
This extensive field research by the Section of Birds of Carnegie Mu-
seum has been made possible by the generosity and continued interest of
Dr. Matthew T. Mellon, who has encouraged the author in every phase of
this work. During my first preliminary studies of the 1947 and 1948 collec-
tions, a distinct race of Aratinga astec has appeared that warrants sub-
specific recognition. In appreciation of Dr. Mellon’s unfailing support it
is an honor to name this new race for him.
Aratinga astec melloni subsp. nov. Mellon’s Paroquet.
Type: No. 133,003, Collection Carnegie Museum, adult male; La Ceiba,
Honduras, April 30, 1948; Arthur C. Twomey and R. W. Hawkins.
Paratypes: No. 133,007 C.M., from the type locality, April 30, 1948;
No. 132,540 C.M., Trujillo, Honduras, March 30, 1948; No. 132,540 C.M.,
Trujillo, Honduras, March 31, 1948; No. 133,528 C.M., Coyoles, Hon-
duras, June 16, 1948; No. 133,730 C.M., San Esteban, Honduras, June
30, 1948.
Subspecific characters
Differs from Aratinga astec astec from Costa Rica in being decidedly
paler on the breast, abdomen and fianks, the green being duller and
more suffused with a grayish undertone, the olivaceous cast to the breast
and throat duller; the back is paler with an olivaceous cast on the crown,
back and interscapular regions in contrast to the brighter green of Aratinga
astec astec. SimWdiY to Aratinga astec vicinalis but is duller and lacks the
bright greens on abdomen and back. Specimens from British Honduras
are intermediate between Aratinga astec vicinalis and Aratinga astec
melloni.
Average Measurements
Based on six specimens: wing, 132-139 (135); tail, 109-113 (112.5);
culmen, 19-20 (19.7); tarsus, 12-15 (13.5).
297
[ssued April 28, 1950.
298
Annals of the Carnegie Museum
VOL. 31
Specimens examined
All specimens listed below are contained in the collection of the Car-
negie Museum.
Amtinga astec melloni — Honduras: La Ceiba, 2; Trujillo, 1; San Este-
ban, 1 ; Coyoles, 1.
Aratinga astec vicinalis — Mexico: Estado de Tamaulipas, 1.
Aratinga astec vicinalis (intermediate with A. a. melloni) — British
Honduras: Manatee Lagoon, 6; Duck Run (Benque Ceiba), 1.
Aratinga astec astec — Costa Rica: El Hogar, 3; Guapiles, 4.
Present known range: North coast of Honduras inland to Olancho.
) ( V '
'•W ^
ART. 14. A STUDY OF THE SALAMANDER,
AMBYSTOMA CINGULATUM,
WITH THE DESCRIPTION OF A NEW SUBSPECIES
By Coleman J. Coin
University of Florida
(Plate 1)
Amby stoma cingulatum Cope belongs to the group of salamanders of the
genus Amby stoma which have the plicae of the tongue diverging from a
median furrow rather than radiating from the posterior base of the tongue.
Included in this group are cingulatum^ annulatum, texanum {sensu lato),
mabeei, and probably schmidti of Mexico. In addition to the character
of the tongue, annulatum, texanum, and cingulatum are further character-
ized by multiple rows of teeth on the jaws, and in cingulatum and an-
nulatum the individual tooth terminates in a rounded knob rather than
in a point. A. cingulatum differs from annulatum in its smaller size, much
narrower head, and in that the pattern is not composed of just a few
pronounced annuli.
Although it has been more than eighty years since A. cingulatum was
first described by Cope (1867 : 205), remarkably little new information has
been added to the original description, primarily because no one seemed to
know where to look for the species. Within the past several years two
persons have independently found how to collect the form and a fair
amount of recent material has thus become available. Wilfred T. Neill,
formerly of Augusta, Georgia, and now with Ross Allen’s Reptile Insti-
tute, has been successful in collecting the species on the Atlantic Coastal
Plain of Georgia and South Carolina, and Mr. Harvard E. Nygren, a stu-
dent of the University of Florida, has discovered a population of cin-
gulatum just north of Pensacola in Escambia County, Florida. Both of
these gentlemen have kindly collected fresh material for me. More re-
cently, since the habitat of the species has become known, other interested
persons have collected specimens. There has thus been made available to
me more than five times as much material as any previous worker has had.
Examination of this recent material, much of which I have seen alive,
and re-examination of specimens heretofore available in collections, re-
vealed that cingulatum is made up of two well-differentiated subspecies,
one on the Atlantic Coastal Plain and the other on the Gulf Coastal
SEP 2 01950
300
Annals of the Carnegie Museum
VoL. 31
lina, this name must be applied to the Atlantic Coast form. No name ap-
pears available for the Gulf Coast population and I therefore take pleasure
in naming it in honor of my distinguished friend, Dr. Sherman C. Bishop.
Ambystoma cingulatum bishopi, subsp. nov.
Reticulated Salamander
(Plate 1)
1917. Ambystoma cingulatum Stejneger and Barbour, A Check List of North Ameri-
can Amphibians and Reptiles, 1st ed., p. 8 (part). — Dunn, Bull. Mus. Comp.
Zool., Vol. LXII, no. 9, 1918, p. 458. — Loding, Geol. Surv. Ala., Mus. paper no,
5, 1922, p. 11. — Stejneger and Barbour, A Check List of North American Am-
phibians .and Reptiles, 2nd ed., 1923, p. 4 (part). — Stejneger and Barbour, A
Check List of North American Amphibians and Reptiles, 3rd ed., 1933, p. 4 (part).
— Wright, Proc. Nat. Acad. Sci., Vol. XXI, no. 6, 1935, pp. 340, 341 (part). —
Wright, Science, Vol. LXXXI, no. 2106, 1935, p. 463 (part). — Stejneger and
Barbour, A Check List of North American Amphibians and Reptiles, 4th ed.,
1939, p. 7 (part). — Carr, Univ. Fla., Biol. Sci. Ser., Vol. Ill, no. 1, 1940, pp. 30,
46 (part). — Orton, Copeia, 1942, no. 3, pp. 170, 171. — Bishop, Handbook of
Salamanders, 1943, pp. 113, 123-126 (part). — Stejneger and Barbour, Bull.
Mus. Comp. Zool., Vol. XCIII, no. 1, 1943, p. 9 (part). — Allen and Neill, A
checklist of the amphibians and reptiles of Florida, 1949, p. 1 (part).
1940. Ambystoma texanum Carr, Univ. Fla., Biol. Sci. Ser., Vol. Ill, no. 1, pp. 30,
46.
Type: Carnegie Museum, no. 29,137, adult female, collected about five
miles north of Pensacola, Escambia County, Florida, May 7, 1949, by
Harvard E. Nygren, and preserved on December 8, 1949.
Paratypes^: Sixty-two, as follows: Alabama: Mobile ,Co., Mobile;
USNM 42,861, 57,389-90. Florida: Calhoun Co., Scott’s Ferry; ERA-
WTN 14,004 (32); Escambia Co., Pensacola; DBUF 34; about 5 mi. N.
Pensacola, AMNH 54,742, ANSP 26,274-75, SCB, 1,546-47, CM 29,138-
144, CAS, 15,550-51, DBUF, 2,627, CJG 1,869 (alive), MCZ 26,754-55,
Tulane 11,824, UMMZ 100,982-83; Jackson Co., near Cottondale;
UMMZ 77,062 (2); Mariana; UMMZ 73,989. Georgia: Baker Co., 1.5
^ ERA-WTN, E. Ross Allen-Wilfred T. Neill collection; AMNH, American
Museum of Natural History; ANSP, Academy of Natural Sciences of Phila-
delphia; SCB, personal collection of Sherman C. Bishop; CM, Carnegie Mu-
seum; CAS, Chicago Academy of Sciences; JWC, personal collection of John
W. Crenshaw; DBUF, Department of Biology, University of Florida; CJG,
personal collection of Coleman J. Goin; MCZ, Museum of Comparative Zoology;
Tulane, Department of Zoolog3^ Tulane University; USNM, United States
National Museum; UMMZ, University of Michigan, Museum of Zoology.
1950
Coin : Study of Ambystoma cingulatum
301
mi. NNE junction of Early, Miller, and Baker counties; JWC 293; Early
Co., junction of Colquit-Blakely and Hilton-Damascus roads; UMMZ
100,695.
Diagnosis: An Ambystoma with multiple rows of knob-like teeth on
both jaws and on the vomers and with a dorsal pattern of reticulations
formed by narrow, orange-gray lines on a chocolate-black background;
ventrally the pattern may be described as tiny gray flecks on a dark back-
ground giving a salt and pepper appearance. From Ambystoma cingulatum
cingulatum it differs in its more robust build, in having a ventral pattern
salt and pepper in appearance rather than a series of discrete white spots
on a dark background, in having a more reticulate dorsal pattern, and in
showing a tendency toward having the vomerine teeth arranged in two
rows rather than three, and fewer in number (average 13.2 in each patch
in bishopi, 15.6 in each patch in cingulatum).
Description of Type: Skin smooth on dorsum and venter, heavily
wrinkled on sides between axilla and groin. No large glands present. No
discernible rows of mucous pores on head.
Head (as seen from above) somewhat elongate, the sides nearly parallel
behind the eyes, in front of eyes tapering to a rounded snout; as seen from
side, somewhat depressed, only slightly arched above; snout projecting
beyond tip of lower jaw. Outline of mouth nearly straight posteriorly,
curving down anteriorly; angle of mouth below posterior margin of eye.
No canthus rostralis. Nostrils small, semicircular, anterior in position
being located nearly at tip of snout ; distance between nostril and anterior
corner of eye equals length of eye; distance between nostrils equals about
two-thirds length of eye. Eye moderate in size, distance from tip of
snout just slightly exceeding length of eye. Upper eyelid fitting over
lower eyelid both anteriorly and posteriorly. A distinct groove passes
downward and backward from posterior corner of eye to behind base of
jaw where it forks, one branch extending downward and the other branch
curving upward and posteriorly, nearly reaching the gular fold on the
side of the head. The gular fold extends well up on the side of the neck
on each side, ventrally it forms a very obtuse angle directed anteriorly.
Head width 5.42 times in body length; head length 3.35 times in body
length.
Body somewhat slender for an Ambystoma, very slightly flattened dorso-
ventrally. No pronounced constriction at neck. A narrow mid-dorsal
groove; a brqad but shallow mid-ventral groove. Costal grooves 15, ex-
tending dorsally to the mid-dorsal groove and connecting across the
302 Annals of the Carnegie Museum Vol. 31
venter. Costal folds between adpressed toes, two. Vent a short, slightly
elevated slit; internal lips folded.
Tail shorter than head and body; 1.22 times in head and body length;
not constricted at base; ovate in cross section at base becoming gradually
compressed to flattened tip. No tail fins present; posterior fourth of tail
slightly keeled above.
Fore limbs moderately stout; fingers four, moderately slender, un-
webbed; 3-2-4-1 in order of decreasing length; no metacarpal tubercles
present. Hind limbs moderately stout; toes five, moderately slender, un-
webbed, 4-3-2-5-1 on right foot; second toe on left foot amputated; no
metatarsal tubercles present.
Tongue moderate, slightly free on sides, with a longitudinal median
depression from which the plicae originate. Premaxillary-maxillary teeth
rounded, knob-like, arranged in several ill-defined rows, a maximum of
four rows in the middle of the upper jaw, tapering to two rows posteriorly.
Mandibular teeth similar in structure but more uniformly arranged in
rows, essentially three rows present at mandibular symphysis, but at one
or two points a fourth row can be discerned ; tapering to one or two rows
posteriorly. Vomerine teeth rounded, knob-like, arranged in two elongate
patches between posterior margins of internal nares; distance between
patches about one-half the width of an internal naris; neither patch ex-
tending laterally to behind the naris; fourteen teeth in left patch, eleven
teeth in right patch. Internal nares moderate, rounded.
Coloration {preserved): Ground color above, blackish; dorsal pattern
consisting essentially of reticulations which are formed in the following
manner: on each side, from the angle of the jaws to the base of the tail,
there is a creamish-yellow, lichen-like area interspersed with gray ver-
miculations. Between the axilla and groin there arises from this creamish-
yellow area a series of narrow, gray bands which pass upward, generally
one present on each costal fold. As these bands pass upward they fre-
quently anastomose to form reticulations. About one-third of them con-
tinue up and meet their fellows from the opposite side on the mid-line
of the back. The pattern on the top of the head and the dorsal surface
of the tail is essentially the same except that the narrow, gray bands do
not follow costal folds. Ventrally, the pattern of the throat, belly, and
under surfaces of the tail, and limbs, is composed of innumberable tiny
white areas on a ground color of grayish-black, giving a salt and pepper
appearance. The dorsal surfaces of the limbs have narrow, gray bands
similar to those on the back.
1950
Coin : Study of Ambystoma cingulatum
303
Coloration in life: In life the ground color of the dorsum was a chocolate-
black and the narrow dorsal bands were a brownish-gray. The lateral,
lichen-like areas on the sides of the head and between the axilla and groin
were pearl-gray ; hence these lichen-like areas and the dorsal bands arising
from them were appreciably different in coloration. The ventral ground
color was black and the ventral light markings were a diffuse gray.
Measurements {in millimeters): Head width, 8; head length, 13; body
length, 43.5; tail length, 46.5; axilla to groin, 30; fore limb, 12; hind limb,
13; total length, 103.
Variation: Fresh adult material of A. c. hishopi is rather uniform in
coloration, but there is some variation in the dorsal pattern. The ground
color of the dorsum is a chocolate-black and the lateral, lichen-like marks
between the axilla and groin are gray, but the narrow bands which arise
from this lateral, gray area, generally one on each costal fold, are a
definite brownish-gray. Thus in life the dorsal bands seem to arise from,
but not be continuations of, the lateral, gray areas whereas in A. c.
cingulatum the dorsal bands seem to be mere continuations of the lateral,
gray areas. In the majority of the adult specimens of fresh material the
dorsal, light marks tend to divide and anastomose, giving a definitely
reticulated appearance to the dorsal surface. Markings similar to those
of the back are present on the top of the head and sides and top of the
tail. In a specimen which I still have alive in a terrarium (plate 1, fig. 5)
the brownish-gray bands arising from the lateral, gray areas and ascend-
ing on the costal folds show the minimum amount of branching or ana-
stomosing, the great majority of them continuing uninterruptedly to the
mid-line of the back where they meet their fellows from the opposite side,
giving the individual an annulate rather than a reticulate appearance.
Ventrally, the ground color is black with numerous tiny, anastomosing,
gray flecks giving the throat, venter, and under side of the tail, a salt and
pepper appearance. In general, the dark pigment ventrally is more con-
centrated posteriorly so that there is an indefinite gradient in tone with
the animal being lightest under the chin and throat, intermediate on the
venter, and darkest on the ventral side of the tail.
As so much of the heretofore available material was old and in such
poor condition that teeth and costal grooves could not be counted or
accurate measurements made, I list in Table 1 tooth counts, costal groove
counts, and measurements of eleven recently collected and well preserved
specimens of c. hishopi that are 50 mm. or more in head and body
length. From this table it can be seen that the costal grooves (all forks
304
Annals of the Carnegie Museum
VOL. 31
counted) are constant at 15. The axilla to groin length seems to be on the
average greater in relation to the head length and the limb length than
it is in .4. c. cingulatum. In this fresh, adult material of bishopi the num-
ber of costal folds between adpressed toes ranges from 2 to 4 with an
average of 3.23. In axilla to groin length divided by head length, bishopi
ranges from 2.23 to 2.77 with an average of 2.52.
Fig. 1. Left, teeth of Ambystoma c. cingulatum from Emanuel Co., Georgia, CM
29,145. Right, teeth of Ambystoma c, bishopi from Escambia Co., Florida,
CM 29,137 (type).
If all specimens in which I have been able to make accurate counts are
considered, the number of rows of teeth on the premaxillary-maxillary
occur in the following order of frequency: 3, seven times; 4, thirty-five
times; 5, eight times; and 6, once. The numbers of rows of teeth on the
mandibles occur as follows: 3, twenty-two times; 4, twenty-six times; and
5, twice. In all specimens in which accurate counts of vomerine teeth
were possible, the number in each patch ranges from 5 to 20 with an aver-
age of 13.2. In general, the vomerine teeth inA.c, bishopi are arranged in
Costal
Folds Head
between and Axilla
Costal Vomerine adpressed Body Head Head Tail to Total
Museum Number Grooves Teeth toes Length Length Width Length Groin Length
1950
Coin: Study of Ambystoma cingulatum
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fO On On ^
rtH ro fO
OOiOt'»vot^NOr:}<oO
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6 ^
o ^
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" I § g § <
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305
306
Annals of the Carnegie Museum
VOL. 31
two rows whereas in A. c. cingulatum they show a definite tendency to be
arranged in three rows.
In general build, an adult specimen of hishopi usually lacks a definite
constriction at the neck and has a broad, heavy tail. This is in con-
tradistinction to A. c. cingulatum, a form which has a rather definite neck
and generally a more slender tail.
Specimens from Calhoun County, Florida, approach c. cingulatum in
three respects: ventral pattern, vomerine tooth number, and costal
grooves. The thirty-two specimens in this series were unfortunately placed
in a bottle containing preservative too strong, so that they are coiled and
quite stiff, hence impossible to measure accurately. Therefore they are
not included in the table. In ventral pattern, the ventral, light spots are
slightly larger and more scattered than in topotypical hishopi although
there is some variation within the series. In the vomerine tooth-count, the
series ranges from 12-12 in the specimen having the lowest count to 21-21
in the specimen with the highest, and averages 14.5 in each patch, which
is above Pensacola specimens which average 9.7 in each patch, but below
cingulatum which averages 15.6 in each patch. Two of the thirty- two
specimens have 16 costal grooves, a count commonly found in cingulatum.
The specimens JWC 293 from Baker County, Georgia, and UMMZ
100,695 from Early County, Georgia, are like the Calhoun County speci-
mens in having the ventral spots a little larger than they are in the Pensa-
cola specimens. The vomerine tooth-counts are/13-17 and 13-14 respec-
tively. Finally, it should be mentioned that several of the Calhoun
County specimens tend to have the vomerine teeth alined in three rather
than two rows and thus approach c. cingulatum in this respect.
The dorsal pattern seems to be the same as it is in the Pensacola speci-
mens. Although not measurable, in general body form they are compar-
able to topotypical specimens of hishopi and I have therefore included
them in the definition of this race.
This Calhoun County series exhibits a certain degree of abnormality
in feet and hands. Three specimens have some modification of the normal
digital complement. One of these has the middle toe on the left hind foot
bifurcate, and the third toe on right foot amputated. In another speci-
men, the third digit on the right hand has two supernumerary digits grow-
ing out of the side of it, and another specimen has the fifth toe missing
and the fourth very short on the right hind foot. Finally, one specimen
has the left hand amputated.
Sexual dimorphism: In the series of thirty-two specimens from Scott’s
1950
Coin: Study of Ambystoma cingulatum
307
Ferry, Calhoun County, there are fifteen males and seventeen females.
Sexual dimorphism is not pronounced in this species. The males have the
cloacal region slightly swollen but this character is by no means so obvious
in this species as it is in the larger species of Ambystoma. The most con-
venient and accurate method of determining sex is by examination of the
Mullerian duct, which is pigmented in the males and unpigmented in the
females, as is the case in Salamandra (see Francis, The Anatomy of the
Salamander, 1934: 284). The amount of pigmentation is apparently cor-
related to some extent with age, since the ducts in the larger males are
more heavily pigmented than they are in the smaller individuals.
Ambystoma cingulatum cingulatum Cope
Frosted Salamander
(Plate 1)
1867. Amblystoma cingulatum Cope, Proc. Acad. Nat. Sci, Phila., Vol. XVIII,
p. 205. — Strauch, Mem. Imperiale des Sci., St. Petersbourg, Tome XVI, no.
4, 1870, p. 65. — Cope, Bull. U. S. Nat. Mus., no. 1, 1875, p. 26. — Boulenger,
Cat. Bat. Grad., 1882, p. 50 (part). — Davis and Rice, 111. State Lab. of Nat.
Hist., Bull. no. 5, 1883, p. 11. — Yarrow, Bull. U. S. Nat. Mus., no. 24, 1883,
pp. 21, 192. — Garman, Bull. Essex Inst., Vol. XVI, 1884, p. 38.
1886. Amblystoma lepturum Cope, Proc. Amer. Philos. Soc., Vol. XXIII, p. 524,
— Cope, Amer. Nat., Vol. XXI, 1887, p. 88.
1889. Chondrotus cingulatus Cope, Bull. U. S. Nat. Mus., no. 34, p. 100.
1889. Linguaelapsis lepturus Cope, Bull. U. S. Nat. Mus., no. 34, p. 116.
1907. Ambystoma cingulatus Brimley, Jour. Elisha Mitchell Sci. Soc., Vol. XXIII,
no. 4, p. 153.
1917. Ambystoma cingulatum Stejneger and Barbour, A Check List of North Ameri-
can Amphibians and Reptiles, 1st ed., p. 8 (part). — Stejneger and Barbour, A
Check List of North American Amphibians and Reptiles, 2nd ed., 1923, p. 4
(part). — Brimley, Jour. Elisha Mitchell Sci. Soc., Vol. XLII, nos. 1 & 2, 1926,
p. 77. — Wright, Ecology, Vol. VII, no. 1, 1926, p. 82. — Stejneger* and Barbour,
A Check List of North American Amphibians and Reptiles, 3rd ed., 1933, p. 4
(part). — Wright, Proc. Nat. Acad. Sci., Vol. XXI, no. 6, 1935, pp. 340, 341
(part). — Wright, Science, Vol. LXXXI, no. 2106, 1935, p. 463 (part). — Gee,
South Carolina Vert. Fauna, 1936, p. 1. — Brimley, Carolina Tips, Vol. II, no.
4, 1939, pp. 14, 15. — Stejneger and Barbour, A Check List of North American
Amphibians and Reptiles, 4th ed., 1939, p. 7 (part). — Carr, Univ. Fla., Biol.
Sci. Ser., Vol. Ill, no. 1, 1940, pp. 30, 46 (part). — Bishop, Handbook of Sala-
manders, 1943, pp. 113, 123-126 (part). — Stejneger and Barbour, Bull. Mus.
Comp. Zool., Vol. XCIII, no. 1, 1943, p. 9 (part). — Allen and Neill, A checklist
of the amphibians and reptiles of Florida, 1949, p. 1 (part). — Neill, A checklist
of the amphibians and reptiles of Georgia, 1949, p. 1.
1932. Ambystoma angulatum Wright, Frogs of the Okefinokee Swamp, p. 10.
308
Annals of the Carnegie Museum
VOL. 31
Type locality: Grahamville, Jasper County, South Carolina.
Neotype {here selected): USNM 129,396, adult female collected at
Robertsville, Jasper County, South Carolina, Nov. 17, 1947, by Wilfred
T. Neill.
Diagnosis: An Amhy stoma with multiple rows of knob-like teeth on both
jaws and on the vomers, and with a dorsal pattern consisting of grayish-
green, lichen-like marks which tend to form reticulations. From Amhy-
stoma cingulatum hishopi it differs in its more slender, delicate build, in
having the ventral pattern composed of discrete light spots on a dark back-
ground and the dorsal pattern more lichen-like or “frosted” in appearance,
and in a tendency toward having the vomerine teeth arranged in three
rows and more numerous (average 15.6 in each patch in cingulatum, 13.2
in each patch in hishopi).
Variation: The fresh material of A. cingulatum cingulatum which I have
been able to examine is remarkably uniform in general appearance. The
dorsal pattern has a ground color of black wdth extensive, gray, lichen-
like markings concentrated along the side between the axilla and groin.
On each costal fold these gray, lichen-like marks extend dorsally, generally
forking near the mid-line of the back to meet their fellows from the op-
posite side. Similar diffuse lichen-like mai;ks are present on the top of
the head and top and sides of the tail. The general effect of these gray,
somewhat indefinite bands on the black babkground is to give the sala-
mander a lichenose or frosted appearance and Brimley’s use of the com-
mon name “frosted salamander” seems to me to be appropriate for this
subspecies. Ventrally the ground color is black with scattered pearl-gray
light spots.
Since much of the material which has heretofore been available for
study is old or poorly preserved, I give in Table 2 a list of counts and
measurements made on recently collected specimens 50 mm. or more in
head and body length. From this table it can be seen that cingulatum
has a tendency to have sixteen costal grooves (all forks counted) whereas
fifteen is the number in the great majority of hishopi. It likewise tends to
be relatively shorter in axilla to groin length in relation to head and leg
length than does hishopi. In this fresh, adult material of cingulatum the
number of costal folds between adpressed toes ranges from 0 to 3 with an
average of 1.69. In axilla to groin length, divided by head length, cingu-
latum ranges from 2.08 to 2.62 with an average of 2.33.
There is a pronounced tendency in cingulatum for the teeth to be ar-
ranged in rather compact groups. The teeth on both jaws are arranged
Costal
1950
Coin : Study of Ambystoma cingulatum
309
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310
Annals of the Carnegie Museum
VOL. 31
in more or less definite rows and in the specimens in which unquestionable
counts can be made the rows on the premaxillary-maxillary vary from 3 to
6, occurring in the following frequencies: 3, twice; 4, eight times; 5, three
times; and 6, once. On the mandible the number of rows ranges from 2
to 4, occurring in the following frequencies: 2, once; 3, three times; 4,
nine times. In vomerine teeth there is a tendency for three rows to be
formed on each patch and in fresh material there is definitely a greater
number of teeth in each patch than in hishopi. Extreme caution should be
used in counting vomerine teeth since, if a needle is used in the process,
many peripheral teeth may be displaced without leaving a definite gap,
thereby causing future workers to obtain a lower count. Thus, at the
present time, I am able to count only 12 in each patch on USNM 49,432
from Jacksonville, Florida, a specimen which has been examined many
times by many workers, and on USNM 14,583, the type of A. lepturum
Cope, there are now 9 teeth in the left vomerine patch and 10 on the right.
Since in this specimen, however, the vomerine teeth tend to be arranged
in three rows, since a faint trace of a ventral pattern similar to that of c.
cingulatum is still discernible, and since the general build is much more
similar to cingulatum than it is to bishopij I do not hesitate to consider
it a synonym of c. cingulatum.
I have seen two specimens which are recorded in the literature as com-
ing from Pensacola, Florida, which seem to me to be referable to cingu-
latum rather than to hishopi. These are MCZ 204. In both of these
specimens the vomerine teeth are uncountable. The larger specimen has
16 costal grooves on the left and 15 on the right, and the smaller I could
not count accurately. The smaller of the two specimens is certainly re-
cently transformed and the adult pattern is not fully developed. In the
larger, however, although it is somewhat faded dorsally, the ventral pat-
tern is quite distinct and is similar to that seen in fresh material of cingu-
latum and unlike that of any of the fresh material of hishopi I have seen
from Pensacola. When I queried Mr. Loveridge regarding the data of
these two specimens, he told me that the entry in the catalogue of the
Museum of Comparative Zoology reads as ioWo^s:^ Amhly stoma jeffer-
sonianum Bd. var. 2 ex. Pensacola, Fla. Alcoholic. (Collected) 1853.
Col. Deas. Prof. Agassiz. Sent to Prof. Baird Jan. 22nd 1859. (entered
in register) Feb. 20, 1862.” In view of the length of time that elapsed be-
tween collection of the specimens and their entry in the catalogue, the
possibility of error cannot be ruled out.
1950
Coin: Study of Ambystoma cingulatum
311
Life History
Habitat: Both races of A. cingulatum seem to be essentially inhabitants
of slash pine — wire grass flatwoods. Every specimen for which detailed
ecological notes are available was taken in or near one of the small, shal-
low, cypress ponds so characteristic of these flatwoods or in a drainage area
from such a pond.
Habits: Specimens of c. cingulatum can be found under logs around
the margins of the above-mentioned ponds throughout the winter months.
While they appear to be quite capable burrowers, Wilfred T. Neill informs
me that they show a pronounced tendency to occupy crayfish burrows.
The recent specimens of c. bishopi from Escambia County, Florida,
have all been taken near an old, abandoned air field which had been made
in the flatwoods about five miles north of Pensacola. Most of the speci-
mens were found around the margin of a cypress pond, that has become
more or less a dump heap, near the edge of the air field. Specimens have
been taken in April, May, June, September, and November by turning
over the rubbish in contact with the moist ground. In one case, specimens
were found under clods of earth that had been turned up by people dig-
ging fishing worms. The specimen from Baker County, Georgia, was
taken under a cypress log eight to ten inches in diameter in the nearly dry
basin of a similar cypress pond. A specimen of Triturusv. louisianensis
was under the same log.
The individuals that I kept alive in a terrarium in my back yard from
May 7, 1949, to February, 1950, were active burrowers and built exten-
sive tunnels about equal in diameter to that of their bodies through the
four inches or so of dirt in the bottom of the terrarium. Some days they
would remain in the tunnels, while on others they would rest under a
board kept in the terrarium. They generally tended to remain hidden
either in the tunnels or under the board except on rainy nights when they
would come out and wander about on the surface. During February and
March, 1950, the specimen (CJG, 1,869), still alive in the terrarium, had
a burrow which had its entrance at the exposed surface rather than under
the board. While occupying this burrow it would remain hidden during
the day but would often rest at night with just the tip of its head exposed.
They made no attempt to climb the glass walls of the terrarium.
Eggs and Breeding Season: Specimens of both subspecies have been
found with mature, pigmented eggs in the ovaries. A specimen of cingu-
latum from Jasper County, South Carolina, taken on November 7, 1947,
312
Annals of the Carnegie Museum
VOL. 31
had twenty-one pigmented eggs in the left ovary and twelve in the right.
Another specimen, from Emanuel County, Georgia, collected November
24, 1949, had forty pigmented eggs in the left ovary and thirty in the right.
Two specimens oi A. c. bishopi with pigmented eggs seem to indicate a
rather extensive breeding season. A specimen taken in Escambia County,
Florida, in June, 1949, had sixty-three pigmented eggs on the left side of
the body and sixty-four on the right, whereas the specimen taken by John
Crenshaw in Baker County, Georgia, on January 13, 1950, when killed on
January 25, had forty-seven eggs on the left side and fifty-five on the
right.
The series of thirty-two specimens taken at Scott’s Ferry, Calhoun
County, on February 20, 1950, had presumably bred shortly prior to the
time of collection. In none of the females were the ovaries full of pig-
mented eggs and six had obviously spent ovaries in which just a few pig-
mented eggs were retained. In these six specimens the following numbers
of pigmented eggs were found (in each case the first figure is for the left
side of the body, the second for the right): 7-2, 3-0, 0-2, 2-4, 0-1, 1-0.
LarvcB and Immatures: Orton (1942: 170) described and illustrated three
larval specimens of Amby stoma which, on the basis of comparison with
known larvae, she assigned to cingulatum. Upon comparison of these
larvae with recently transformed specimens of both races of cingulatum, I
feel confident that Dr. Orton’s identification of them as belonging to this
species is correct. In addition to the three larvae from Jackson County
mentioned above, I have seen three recently transformed specimens which
retain the larval pattern: AMNH 36,479 (38 mm., head and body length;
65 mm., total length), from Lanier County, Georgia; CM 29,141 (36 mm.,
head and body length; 57 mm., total length) from Escambia County,
Florida; and MCZ 204 B, purportedly from Escambia County, Florida.
I have indicated earlier (p. 310) that the adult specimen numbered MCZ
204 seems to be more like cingulatum than bishopi. Examination of the
recently transformed specimen confirms this opinion.
The specimen of cingulatum from Lanier County, Georgia, is brown in
ground color with a tan stripe extending from axilla to groin and another
tan stripe extending from above the arm posteriorly until it fades out on
the distal third of the tail. There is a black stripe from the nostril which
extends posteriorly through the eye to the vertical extension of the gular
fold on the side of the neck. Above this is a slightly lightened area as
though the uppermost lateral stripe in life continued on to the side of the
head. A narrow mid-dorsal light line extends from the base of the head to
1950
Coin: Study of Ambystoma cingulatum
313
above the insertion of the fore limbs. There is a very narrow black margin
along the sides of the upper lip. The venter is very pale brown. The
same general coloration and pattern can be made out in MCZ 204 B
except that the upper lateral stripe is more indistinct, the adult pattern
is becoming discernible on the tail and the venter is becoming more
heavily pigmented, leaving spots of unpigmented areas. The specimen of
hishopi from Escambia County, Florida, although 2 mm. less in head and
body length and 8 mm. less in total length than the specimen of cingu-
latum, has more of the adult pattern. Both of the lateral stripes are still
present but the stripe between the axilla and groin is breaking up into
patches and the superior lateral stripe is not nearly so sharply defined
as it is in the specimen from Lanier County, Georgia. Furthermore, al-
though the specimen from Lanier County has a definite dorsal fin on the
tail extending from above the vent to the tip, the tail fin is completely
obsolete in the Escambia County specimen of hishopi. The adult dorsal
pattern is beginning to develop between the upper lateral stripes in the
specimen of hishopi; there is no hint of it in the specimen of cingulatum.
There is no trace of a mid-dorsal light line in the former.
If these specimens are at all typical as to the size at which the adult
color pattern is developed, it would seem that hishopi transforms and
attains the adult pattern at a smaller size than does cingulatum. While
the specimen of cingulatum, 38 mm. in head and body length, retains the
larval coloration and tail fin, the specimen of hishopi, 36 mm. in head and
body length, is beginning to lose the larval coloration and attain the
adult pattern and has completely lost the tail fin. Two other specimens
of hishopi, collected at the same time and place as the recently transformed
one described above, and measuring 36 and 38 mm. in head and body
length and 56 and 62 mm. in total length respectively, show the typical
sub-adult pattern and have lost all sign of a tail fin.
The sub-adult pattern of ^4. c. hishopi differs from the adult in that the
dorsal reticulations are less sharply defined and the venter is paler in
coloration. In immature specimens ranging from 36 to 43.4 mm. in head
and body length, there is a definite light area on the ventral surface of the
throat just anterior to the gular fold, and the chin, venter, and under
surface of the tail are paler than in adult specimens. The lichen-like area
between the axilla and the groin seems to be less well developed since in
specimens of this size the light areas are not confluent but consist of ill-
defined but separate light marks.
Growth Rate and Longevity: No information is available regarding A. c.
314
Annals of the Carnegie Museum
VOL. 31
cingulatum but a specimen oi A. c. hishopi, measuring 35 mm. in head and
body length and 56 mm. in total length, was put in my terrarium on
May 7, 1949. It fell into a pan of water in the terrarium and drowned,
apparently on the night of June 23, since, when I returned from the meet-
ings of the American Society of Ichthyologists and Herpetologists in
Washington on June 24, I found it in the pan of water, dead but still in
good, fresh condition. At this time, after a lapse of 47 days, it measured 44
mm. in head and body length, an increase of 9 mm., and 73 mm. in total
length, an increase of 17 mm.
Two adult specimens oi A. c. hishopi which were also placed in the ter-
rarium on May 7 thrived until February 7, a period of 9 months, when my
two-year old son released one of them in the leaf mold of my back yard.
The other is still (July 18, 1950) alive.
Food: Apparently both subspecies feed on earthworms. Three speci-
mens o( A. c. hishopi taken in September, and one taken in February, had
the undigested remains of earthworms in the digestive tract, and one
specimen oi A. c. cingulatum from Jasper County, South Carolina, taken
in October, 1947, had no less than seven pieces of earthworm in its
stomach. Since five of these pieces were terminal sections, it is evident
that parts of at least three individual worms were in the stomach at one
time. Many of the series of specimens of hishopi taken in February in
Calhoun County regurgitated earthworms when they were placed in what
was apparently a quite strong preserving fluid, and earthworms were
found in eight of the specimens when their digestive tracts were examined
after preservation. I have twice found tiny fragments of unidentifiable
plant remains in stomachs of hishopi, and the majority of stomachs of
both subspecies which I have examined have contained at least a few
sand grains. This, of course, would be expected if earthworms are a staple
in their diet.
The earthworms removed from the salamanders probably belong to the
genus Diplocardia according to my colleague, Professor E. Ruffin Jones.
I have made no attempt to supply food to the adult specimens which
I have kept alive, but the soil which I put into the terrarium came from
my back yard, which supports a flourishing annelid population, and worms
have frequently been seen in the terrarium. While I have never been able
to observe one of the salamanders in the process of eating, they seemed to
thrive and apparently there was no other food available.
1950 Coin: Study of Ambystoma cingulatum 315
Range
Ambystoma c. cingulatum ranges from Colleton County, South Caro-
lina, and Jefferson County, Georgia, west and south to Lanier County,
Georgia, and Duval County, Florida.
Ambystoma c. bishopi seems to be restricted to a region on the Gulf
Coastal Plain from Baker County, Georgia, and Calhoun County, Florida,
west to Mobile Bay, Alabama. Dr. E. R. Dunn informs me that he has
Fig. 2. Map showing the distribution of the subspecies of Ambystoma cingulatum.
Triangles, Ambystoma c. cingulatum; circles, Ambystoma c. bishopi. Solid
symbols, specimens examined; hollow symbols, literature records.
examined the specimen recorded by Boulenger (1882: 50) from New
Orleans, that it is at present uniform black, and that his notes indicate
that it is most likely a specimen of texanum. Since I have seen a number of
fresh specimens of a small dark form of texanum from the same vicinity,
and since no other specimens of cingulatum have ever been taken from
there, it seems wisest to me to consider Louisiana beyond the range of
bishopi unless and until fresh specimens of this subspecies are collected
from there.
316
Annals of the Carnegie Museum
VOL. 31
The accompanying map indicates the distribution of the two sub-
species. The solid symbols represent localities from which specimens have
been examined by me and the hollow symbols represent literature records.
The records represented by hollow symbols from northern Georgia and
South Carolina are from Wilfred T. Neill’s unpublished manuscript, ‘‘The
Amphibians of Georgia.”
Specimens Examined
Amby stoma cingulatum cingulatum: Nineteen, as follows: South Caro-
lina: Jasper Co., Ridgeland, ERA-WTN 14,007; Robertsville, USNM
129,396 {neotype). Georgia: Burke Co., Midville, ANSP 26,211 Charlton
Co., 1 mi. E. Chesser School, CU 2,936; 2 mi. E. Chesser Island, CU
2,933; Emanuel Co., mi. S. Midville, Burke Co., ANSP 26,276, CM
29,145, DBUF 2,628-29; McKinney’s Pond in northern part of county,
ERA-WTN 14,006; Lanier Co., Lakeland, AMNH 36,479, 37,388;
Screvin Co., Bascom, ERA-WTN 14,005 (2). Florida: Duval Co., Jack-
sonville, USNM 49,431-32; ? MCZ 204 (2) (purportedly from Pensacola,
Escambia Co.). ?? USNM 14,583, no data {type of lepturum).
Amhy stoma cingulatum hishopi: In addition to the type and sixty-two
para types listed, I have seen three additional specimens of hishopi. Two
of these (MCZ 229) were sent several years ago to the University of
Kansas for osteological studies by Mr. Loveridge. The other specimen
(CJG 1,870), I kept alive for nine months before it escaped.
Acknowledgments
I am particularly indebted to Wilfred T. Neill and Harvard E. Nygren
for the gift of many specimens. In addition, Mr. Neill has kindly made his
unpublished manuscript on “The Amphibians of Georgia” available to me.
I wish to thank Dr. E. R. Dunn and Dr. Sherman C. Bishop for their
valuable suggestions and assistance. For the loan of material I wish to
thank E. Ross Allen of Ross Allen’s Reptile Institute, Charles M. Bogert
of the American Museum of Natural History, Doris M. Cochran of the
United States National Museum, Arthur Loveridge of the Museum of
Comparative Zoology, M. Graham Netting and Grace Orton of the Car-
negie Museum, Charles F. Walker of the Museum of Zoology, University
of Michigan, A. H. Wright and Edward C. Raney of Cornell University,
Fred R. Cagle of Tulane University, and John W. Crenshaw of Emory
University Field Station, Newton, Georgia. The drawings are from the
pen of Miss Esther Coogle. To her I owe my thanks.
1950
Coin: Study of Ambystoma cingulatum
317
BIBLIOGRAPHY
Allen, Ensill Ross and Wilfred Trammell Neill.
1949. A checklist of the amphibians and reptiles of Florida. Printed for
Ross Allen’s Reptile Institute by Business Services, Ocala, Fla.,
4 unnumbered pages.
Bishop, Sherman Chauncey.
1943. Handbook of Salamanders. xiv4-555 p., 1 pi., 144 fig. Ithaca:
Comstock Publishing Company.
Boulenger, George Albert.
1882. Catalogue of the Batrachia Gradientia S. Cauda ta and Batrachia
Apoda in the collection of the British Museum. 2nd ed., viii +
127 p., 9 pi. London.
Brimley, Clement Samuel.
1907. The Salamanders of North Carolina. Jour. Elisha Mitchell Sci.
Soc., 23 (4): 150-156.
1926. Revised key and list of the amphibians and reptiles of North
Carolina. Jour. Elisha Mitchell Sci. Soc., 42 (1 and 2): 75-93.
1939. Reptiles and amphibians of North Carolina (continued). Carolina
Tips, 2 (4): 14-15.
Carr, Archie Fairly, Jr.
1940. A contribution to the herpetology of Florida. Univ. Fla. Publ.,
Biol. Sci. Sen, 3 (1): 1-118.
Cope, Edward Drinker.
1867. A review of the species of the Amblystomidae. Proc. Acad. Nat.
Sci. Philadelphia, 18: 166-211.
1875. Check-list of North American Batrachia and Reptilia. Bull. U. S.
Nat. Mus., (1): 1-104.
1886. Synonymic list of the North American species of Bufo and Rana,
with descriptions of some new species of batrachia, from specimens
in the National Museum. Proc. Amen Philos. Soc., 23: 514-526.
1887. The hyoid structure in the Amblystomid salamanders.
Amer. Nat., 21: 87-88.
1889. The Batrachia of North America. Bull. U. S. Nat. Mus., (34):
1-525, 86 pi., 119 fig.
Davis, Nathan Smith, Jr. and Frank L. Rice.
1883. Descriptive catalogue of North American Batrachia and Reptilia
found east of Mississippi River. Illinois State Lab. Nat. Hist.,
Bull. no. 5: 1-67.
Dunn, Emmett Reid.
1918. The collection of Amphibia Caudata of the Museum of Compara-
tive Zoology. Bull. Mus. Comp. Zook, 62 (9): 445-471.
VOL. 31
318 Annals of the Carnegie Museum
Carman, Samuel.
1884. The North American reptiles and batrachians. Bull. Essex Insti-
tute, 16: 3-46.
Gee, Nathaniel Gist.
1936. South Carolina vertebrate fauna. (Salamanders.) [An unpaged
list printed in two columns, from the Department of Biology,
Lander College, Greenwood, S. C., dated October 3, 1936.]
Loding, Henry Peder.
1922. A preliminary catalogue of Alabama amphibians and reptiles.
Alabama Mus. Nat. Hist., paper no. 5: 1-59.
Neill, Wilfred Trammell.
1949. A checklist of the amphibians and reptiles of Georgia. Printed
for Ross Allen’s Reptile Institute by Business Services, Ocala,
Fla. 4 unnumbered pages.
Amphibians of Georgia. (MS)
Orton, Grace Louise.
1942. Notes on the larvae of certain species of Amhystoma. Copeia, 1942
(3): 170-172, 1 fig.
Stejneger, Leonhard and Thomas Barbour.
1917. A Check List of North American Amphibians and Reptiles. 1st
ed., 126 p. Cambridge: Harvard Univ. Press.
1923. Ibid. 2nd ed., x-|-l7l p. Cambridge: Harvard Univ. Press.
1933. Ibid. 3rd ed., xiv + 185 p. Cambridge: Harvard Univ. Press.
1939. Ibid. 4th ed., xvi+207 p. Cambridge: Harvard Univ. Press.
1943. Ibid. 5th ed., Bull. Mus. Comp. Zook, 93 (1): xix+260 p.
Strauch, Alexander.
1870. Revision der salamandriden-Gattungen. Mem. I’Acad. Imp. Sci.
St. Petersbourg, ser. 7, 16 (4): 1-109, 2 pi.
Wright, Albert Hazen.
1926. The vertebrate life of Okefinokee Swamp in relation to the At-
lantic Coastal Plain. Ecology, 7 (1): 77-95, 5 pi.
1932. Life-histories of the Frogs of Okefinokee Swamp, Georgia. New
York, The Macmillan Co., xv-f497, frontispiece, 45 pi.
1935. Some rare amphibians and reptiles of the United States. Proc.
Nat. Acad. Sci., 21 (6): 340-345.
1935. Some rare amphibians and reptiles of the United States. Science,
n. s., 81, (2106): 463. (Abstract of preceding).
Yarrow, Harry Crecy.
1883. Check list of North American Reptilia and Batrachia, with cata-
logue of specimens in the U. S. National Museum. Bull. U. S.
Nat. Mus., (24): 1-249. [This publication bears the date “1883”
on the dust wrapper, but “1882” on the title page.]
1950
Coin: Study of Ambystoma cingulatum
319
Addenda
After the above paper was submitted for publication, four students at
the University of Florida collected a series oi A. c. hishopi which provides
data on abundance. On April 7, 1950, Messrs. David Beecher, Byrum
W. Cooper, Robert Heilman, and Edwin H. McConkey collected forty-
five specimens about three miles south of Mariana, Jackson County,
Florida. These specimens were taken in low slash pine — wire grass flat-
woods in an area approximately by 34 of a mile in extent. The region
had been recently cut over for timber, and logs about six feet in length
had been left lying by the stumps. The four students collected the entire
series between four and six in the evening by looking under these logs.
Some of them had two or three bishopi under them and one had five.
Manculus q. quadridigitatus, Microhyla c. carolinensis, and Storeria d.
wrightorum were found under the same logs as the Ambystoma. Eight
specimens of Pseudotriton m. floridanus were found under logs in damper
places in the same general locality but were not taken under the same
logs with bishopi.
320
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 1
Miss Esther Coogle, del.
Fig. 1. Dorsolateral view of Ambystoma c. cingulatum from Emanuel Co., Georgia,
in life. (CM 29,145)
Fig. 2. Ventral view of same.
Fig. 3. Dorsolateral view of Ambystoma c. bishopi from Escambia Co., Florida,
in life. (CM 29,137, type)
Fig. 4. Ventral view of same.
Fig. 5. Dorsolateral view of Ambystoma c. bishopi from Escambia Co., Florida
showing maximum degree of annulation. (CJG 1,869).
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 14
Plate 1
2
5
/
4 I'' 7. Vi
ART 15. AN ANALYSIS OF COLOR AND PATTERN IN BUTTER-
FLIES OE THE ASIATIC GENUS KARANASA
By Andrey Avinoff
(A posthumous paper. Dr. Avinoff died July 16, 1949.)
Foreword
The major avocation of the late Dr. Andrey Avinoff, Director of the
Carnegie Museum from 1926 to 1945, was the collection of butterflies and
the study of their colors and patterns. To him the ultimate in rarity and
interest was the group of little brown and orange Satyrids of the genus
Karanasa from the high mountains of Turkestan, Afghanistan, and Kash-
mir. Over a period of forty years he studied all of the known specimens and
gathered together everything that had been written concerning them. He
accumulated two imposing collections, the first of which was nationalized
by the Soviets and the second given to the Carnegie Museum. I worked
with him in the study of these butterflies for eight years and the task of
publishing the final report has been placed in my hands. Dr. Avinoff was
constantly devising and revising schemes for recording and analyzing the
color patterns of Karanasa. This one, evolved in 1945, is more or less of
a digression from the main line of study of the genus. I present it sepa-
rately with a minimum of editing.
Walter R. Sweadner,
Curator of Insects, Carnegie Museum
A standardization of properties in coloration and pattern of Karanasa
lends itself to a graphic survey. A chart based on the characteristics of the
front wings has been prepared (plate 1) in order to assist in analyzing the
mutual relationship of coloration and pattern of the upperside. It occurs
to me that the most convenient way of visualizing these characteristics,
as exemplified by all possible variations in Karanasa, is to be obtained by a
table with vertical and horizontal rows: the vertical columns stand for
pattern, the horizontal for pigmentation, thus establishing a system of
co-ordinates. By reducing the variety of patterns to as few types as pos-
sible it was found that the most practical subdivisions would represent the
gradual intensification of the dark parts of the basal portion of the front
wings. Thus, the sequence runs through the following gradations: (1)
a completely light base, (2) a light base bounded by a dark band through
the mesial part of the wing from the middle of the costa to the inner edge,
corresponding in part to the branded portion on the wings of the males in
some species, (3) a more or less uniform dark suffusion of the base, mixed
with the ground color, and without a too contrasting transverse mesial
321
Issued October 31, 1950. A
'I ' ■ s >
322
Annals of the Carnegie Museum
VOL. 31
band, or (4) a completely dark base approximating in intensity the fus-
cous marginal area with usually a still darker outline of the discal, light,
transverse band. These four divisions would practically suffice if there
would not be occasionally some forms displaying a general darkening not
only of the basal part but also the outer portion of the wing; the light band
in such instances is much darkened, especially exteriorly. Examples of this
are few but they have to be accounted for in a diagrammatic scheme and
it is questionable whether such a phenomenon should be classified
under the terms of pattern or coloration. Since this is an extension and
spread of the dark area, it was found more advisable to consider it as a
matter of pattern — in the sense of distribution of the light and dark areas
on the surface of the wings — and treated accordingly as the fifth step in
the horizontal classification of patterns, under (5) general darkening of the
wings including chiefly the exterior portions of the post-discal light band.
The vertical divisions of ground color, besides the dark fuscous pattern
on portions of the wings, would naturally comprise two tints; different
hues of russet, either of a bright brick tint or of an ochreous tone, and a
basic whitish or -ivory color, corresponding to astorica and naryna re-
spectively. A pale ochreous tone, standing between these two types of
coloration, called for a special subdivision which, however, does not
comprise many forms. So the chromatic variation has been broken into
three steps: russet, pale ochreous, and whitish, marked as A, B, and C.
With the five possible variations of pattern, any form of Karanasa can
be thus designated by these co-ordinates in a table covering fifteen
fundamental types. It is to be noted that males and females of the same
form do not fall on the same point of this table of color-pattern co-
ordinates. To make the use of the table clearer, each intersection is illus-
trated by examples of typical males and females having such properties.
That is how it happens that in the majority of cases the two sexes belong
to different divisions in the table. The fifth group of “general darkening”
does not lend itself with certainty to the attribution of russet, ochreous or
whitish and in the case of “russet” could not be illustrated by any female.
So there is no such insect which could be designated by the formula 9 A5.
The advantages of this table lie not only in the simplicity of indicating
the exterior aspect of a form by means of a symbol, expressed in a letter
and a number; it helps also the appreciation of the range of variation, the
component elements of variability, and also the deviation of a given type,
in terms of color and pattern variations. Diagonal variations on the chart
denote naturally the most extreme forms of instability involving a shift in
1950
Avinoff: Color and Pattern in Karanasa
323
both respects. Also the most extreme point on diagonal lines constitutes
the greatest polarity of types as illustrated, for instance, by wilkinsi A1
and kazakstana C4 (excluding the fifth step) and lactea Cl and kafir A4:,
For instance, the amplitude of variations of latefasciata may be made
graphically clear by extending into the three horizontal-chromatic-di-
vision in the fourth step of patterns A4, B4 and C4, with a tendency to
deviate occasionally into B3.
A critical approach to this chart should not minimize a weak side of it,
namely the mixed phylogenetic significance of the horizontal row B. The
ochreous step, introduced merely for practical reasons to show graded
differentiations, contains on the same levels entities which are not wholly
comparable. Some of the ochreous forms are merely insects of the russet
section that have acquired a paler aspect (like leechi and decolorata), or a
regular whitish species, which has gained more color (like ershoffi and
some females of conradti). For purposes of denoting actual kinship, the
level B should be split into a fluctuating line of demarkation segregating
only two basic color groups: russet (sometimes pale ochreous) and whitish
(sometimes with a yellowish tinge). Grum-Grshimailo with his “keen
entomological eye” perceived that it was enough to consider only two
chromatic divisions.
As has been stated above, the chart accompanying this paper is used
both for supplying a precise notation and for stimulating an analytical
appreciation of analogous, but not homologous, properties falling into one
category in accordance with the use of suggested coordinates. A com-
promise in this case is an invitation for a more comprehensive study.
Finally, it is interesting to note the statistical distribution of forms.
Of the males, 45 (3/5ths) belong to A1-A4. The horizonal B and the
vertical 5 both contain the least number of forms. In the A group the
males fall predominantly into the divisions with higher numbers and the
females belong mostly to the lower numbers in successive designations.
For instance, in the division 4 there are males of several forms and very
few females; these females belong to the preceding division 3; males of the
2nd and 3rd divisions often have females of division 1. In B and C the
sexes are more closely paired in the various degrees of matching both as to
coloration and pattern. Whether it could be an indication that the an-
cestral prototype fitted closer into these latter groups, as does the Ameri-
can Neominois, or that one should not attach any phylogenetic impor-
tance to these relationships, is a matter that could not be argued one way
or the other on the ground of such considerations. At any rate the sche-
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Annals of the Carnegie Museum
VOL. 31
matic table of patterns and colors simplifies and clarifies the revision of
forms, and lends itself to interesting comparisons with other properties
of the pattern on the underside of the hind wing.
In order to systematize the variations of the configuration of the
marbled pattern on the underside, I propose the following method, based
on the position and character of dented ante-marginal line ez of Schwan-
witch, and the properties of the dark median band (plate 2). This ante-
marginal toothed line is formed of outward pointed dentations or arrow-
heads between the veins. In some of the butterflies these points are
sharp; in others they are blunted into crescents which tend to produce a
scalloped effect. The position of this dented or scalloped line may vary ; it
occupies either the middle distance between the exterior margin of the
darker median band and the edge of the wing, or else runs closer to the
latter, approximately at one-third of the distance from the edge of the
wing to the median band. These differences in position are respectively
indicated by X and Y, and the acute or obtuse formation of the indenta-
tions are marked as a and o. A further notation of d and 1 for “dark”
and “light” indicates the characteristics of inner portions of these arrow-
heads which might be darkened inwardly or traced on an even, light
background. This element of the pattern is in close co-ordination with the
character of the median band which may be either a shade darker than
the general background, or else suffused with a marbled striation.
A further characteristic recorded in abbreviated notation is the pro-
minence of the white veins: (1) for faintly seen veins (like angrena), (2)
for a medium development (like dissoluta), and (3) for well accentuated
white veins (like aksuensis). The notation of these traits of the underside
is deemed sufficient, although one might also devise special notations for
the variation of the pattern of the base which is either light or tends to
merge in coloration and surface effect with the darker median band. It
may be light as in aksuensis and, blending with the band into an even
striation, spread throughout the whole basal half of the wing.
With these fundamental elements, a simple formula in letters and num-
bers can express the character of the underside and, together with the
notation of the upperside in regard to color and pattern, may give an
abbreviated, compressed description of the insect. For instance the
hiiebneri male is c? A3 y d a v 2.
One could go further into the analysis of the underside of Karanasa
in order to find a language of symbols for the most conspicuous traits-
like the antemarginal portion of the front wings, the relative development
1950
Avinoff: Color and Pattern in Karanasa
325
of the pattern in the discus, and the characteristics of the ocelli. It seems,
however, that the features pointed out so far will be sufficient for a pro-
posed method of ‘‘algebraic” recording. Such a procedure of using a for-
mula of conventional letters and figures could be easily applied and is
recommended for a designation and analysis of forms in any related
group.
For comparative clarity a typical section of the hind wing is taken
into consideration from vein Cu2 to the middle of the next interspace and
inwardly as far as the inner edge of the median band. This irregular sec-
tion is reduced to a rectangular conventionalization so as to show the
characteristics of the band and the position and aspect of the antemarginal
zig-zag line. In such a fashion each form is reduced to an even and com-
parable rectangular band as shown on plate 2. It can enable one to see at
a glance the elements that are recorded in the system of schematic nota-
tions and group similar forms together. Other important characters, the
formation of the band and the aspect of the base, are too variable for a
system of notation, which might tend to become unduly complex.
One of the pioneers of a study of patterns reduced to co-ordinates in a
rectangular conventionalization was Riffart, who analyzed the pattern
of Heliconia and prepared illustrations which look like a species of lepi-
dopterological cubism. The illustrations in plate 2, where patterns are
reduced to tags, is a less extreme adaptation of the same principle where
the actual pattern remains sufficiently recognizable within moderate
conventionalization. The Heliconids in the “pictograms” of Riffart, on
the other hand, require a considerable stretch of the imagination to
visualize the actual insects. It proves, perhaps, that the rule of geometry
over lepidopterology should not be overly dictatorial where the interests
of visual recognition have to be safeguarded.
While it is aside from any proposed system of marking, it is considered
of interest to illustrate a comparison of a few typical bands in Karmiasa
with those of Neominois, Satyrus and even such a distantly related genus
as Neope, as an example of the extreme complexities in the development
of the band. Schwanevitch^ made some extensive studies of significant and
component elements of the pattern of the underside of the hind wings in
Satyrids. His conclusions on the identity of different parts of the pattern
undergoing fluctuations, dislocations, and fusion are highly instructive.
1 Schwanwitsch, B. N. “Evolution of the Wing-Pattern in Palearctic Satyridae”
“1. Genera Satyrus and Oeneis.” Zeit. Morph. Oekei. Tierre; Berlin, Vol. 13, pp,
559-654, 1929.
326
Annals of the Carnegie Museum
VOL. 31
It suffices to state here in this conjunction that the median band travers-
ing this discocellular is based on a phenomenon of certain symmetry in the
pattern of the basal and external part of the wing. Thus for the Satyrids,
as well as for the related group of Nymphalidae, Brassolidae, and Mor-
phidae, one may accept as typical a medium dark band edged symmetri-
cally, inwardly and outwardly, by a lighter line and a darker outline, as
is clearly seen in Neope. These elements may become partly broken up by
veins and the double margin preserved for each open cell, or else they are
undisturbed by the venation. A comparison of Neope muirheadi with three
other species of the genus illustrates those two varieties of reaction of the
pattern to the presence of veins (plate 2). It demonstrates the relative —
not absolute — dependence of the pattern on anatomical properties of the
surface of the wing, namely, the arrangement of veins, and it may show
to a considerable degree what may be termed organic autonomy of the
pattern. In Karanasa the band does not display such regularity of pattern
structure and is of interest mainly by its outline obeying some standard
fluctuations within cellular outlines. A comparison of Karanasa and
Neommois in this respect is instructive. In Neominois the tooth in space
M2-M3 is the most prominent, while in Karanasa it is the one above in
space M2-M1. Furthermore, in space Cu2-Cui in Neominois the band is
distinctly narrowed, but usually of the same width as the rest of the band
in Karanasa. A constriction of the band in this point is typical for some
forms of the conradti-regeli group, namely in turngensis and arpensis and
in korlana to a lesser degree. This trait is also observed in Satyrus geyeri
and it may be taken perhaps as a sign of some primitive character for a
whole group of related Satyrids. It may be another indication that it is the
conradti-regeli type which corresponds to the ancestral form related to the
American Neominois and somewhat similar to another far more removed
branch exemplified by Satyrus geyeri.
The upper part of the band calls for special considerations. The portion
between the first and second subcostal in Karanasa tends to be outlined
by a concave contour on either side with an extension outwardly along
the 2nd subcostal. In Neommois the formation is reversed constituting
a convex outline that produces an oval marking in this interspace. This
divergence is demonstrated on plate 2 and has certain phylogenetic im-
plications. The band formation of Karanasa is in closer correspondence to
the predominantly even or parallel outlines in the pattern of Satyrids in
general. The comparison with Satyrus and Neope is a confirmation of this
effect. So, the formation of the band in Kara7iasa may be considered more
1950
Avinoff: Color and Pattern in Karanasa
327
primitive than the alteration in Neominois, especially in certain extreme
specimens of N. dionysius. The interpretation of the peculiarities in the
relative formation of the upper portion of the band in Karanasa and
Neominois requires a more thorough scrutiny. The extension into outward
points between the veins in the whole exterior outline of the band is
characteristic in both genera and is shared by Satyrus geyeri as well. In
the interspace below the second subcostal, the outline of the band shows
a vestige of this point in Karanasa, just below the extension along the vein.
One may consider that the point situated in the middle of this interspace
in Neominois has relatively moved upward and has produced a partial
disappearance of this protrusion together with an extension alongside the
vein. It corresponds to the general shift of the pattern upward toward the
costa, thus effecting in Karanasa a constriction of the band, while in
Neominois this part of the pattern shows central inflation, producing be-
tween the veins an isolated oval spot. A shift of pattern, independent of
the anatomical properties of the wing, falls into the general category of
the peculiar phenomenon designated as ‘‘pierellisation” by Schwanevitch.
It was described as a particular dislocation of bands and ocelli in the
South American Satyrid genus Pierella. Schwanevitch observed and singled
out analogous phenomena in a number of cases. Something analogous
occurred in the genus Karanasa, and it may be assumed that the con-
servative trends in the formations of the band were complicated and sup-
plemented by an evolutionary process expressed by a slight shift of the
pattern toward the costa.
The character of the surface treatment of the band in Karanasa may
vary substantially from being almost as light and uniform as the rest of
the background of the wing, as in wilkinsi and intermedia, to an even
darkening as in the decolorata group, or to a marbled aspect as in alpherakyi.
Such a reticulation is, in fact, of a dual nature. In part it is the remnants
of the shuffled outlines of the inner portions of the band so clearly ex-
emplified in Neope; and in part it is the ancestral vermiculation of large
areas in the underside of Rhopalocera, of which the family Elymnidae is
the most typical. Such a pattern is evidence of the loss of the numerous
transverse veinlets of the primitive Neuropteroid insects which preceded
the development of Lepidoptera. Primitive forms of Lepidoptera show
well these surviving scars of lost veins, in the pattern, as in the family
Cossidae, for instance. The preservation of such a transverse reticulation
in the costal region of the front wings, not only of Nymphalids and
Satyrids but even in Pierids, is symptomatic of this origin, as has been
328
Annals of the Carnegie Museum
VOL. 31
frequently stated in entomological literature. The costal region is supposed
to have preserved certain atavistic traits in terms of pattern. The strongly
marbled base of the hind wings, including the band, in the pamira group
and in other forms as shown on plate 2, is at least in part an atavistic
character of this order and not merely a splitting and dislocation of
identifiable linear bands. It is practically a double phenomenon of the
same order as the case of Caligo, where the general over-all reticulation
condenses itself to suggest the shading of typical, individual, ante-
marginal bands in zig-zag formation. Analogously, through the maze of
an irregular striation in the pamira group of Karanasa, one can easily
discern the inner outlines of the median band and the accentuation of the
usually obscured basal band.
In one more point the underside of the hind wing of Karanasa requires
some study, namely in regard to vestigial ocellation. The primoridal type
of the Nymphalo-Satyrids possessed presumably a complete row of ocelli
between the median band and the ante-marginal arcuated line. Parenthe-
tically it should be mentioned that these ocelli in their own turn were a
modification and confluence of two bands, transformed into isolated
links. In Neominois there are no traces of such ocelli, while in Karanasa
they appear in rare instances (in abramovi and talastauana groups) on the
upperside between the first and second radial. On the underside, vestiges
of ocelli may be observed as faint white spots just within some of the ante-
marginal arrow points as seen in diagrams in plate 2.
Probably the closest approach to an abortive ocellus is observable in
safeda which shows a dark outline of this white center. So here we see in
Karanasa an ancestral trait which has been lost in the American branch
of the group. The presence of such vestigial pupils of lost ocelli in the
most primitive Karanasa of the regeli group should not be overlooked from
the phylogenetic angle. Altogether one might sum up the characters of the
Karanasa underside as a typical Satyrid pattern somewhat impoverished,
and with a slight upward dislocation of the hind wing pattern. The ut-
most of simplification is reached in intermedia^ which constitutes the ex-
treme manifestation of a gradual loss of patern on the upperside, with a
predominant russet hue always remaining as one of the two most typical
chromatic possibilities in the family of Satyridae. From these various
grounds it would be justifiable to consider intermedia as the most progres-
sive type in the evolution of the Karanasa group.
The fluctuations between the highly pigmented scales with a russet
hue and the tendency toward a whitish color is to be seen in the whole
1950
Avinoff: Color and Pattern in Karanasa
329
realm of Satyrids. It may be individual within certain species like Satyrus
anthe and hriseis where the white and ochreus-orange morphs appear as
aberrations or mutations. It may be a sexual trait, as in Lethe. On the
other hand, the color principle may be a matter of established stability
such as the white background of Melanargia or the russet bands of
Epinephele. In our group the coloration is in itself a trait of uneven phylo-
genetic value. The lighter background— less pigmented with ochreous and
russet — acquires a more ancestral significance only in conjunction with
more archaic pattern traits. It is only in this spirit that one should ad-
vance the arguments in favor of the antiquity of the whitish conradti-
regeli group, recording at the same time the phenomenon of wide chro-
matic variation within the species grumi and the group pamira-alpherakyi,
both of which groups may include either whitish or bright russet forms.
It proves only that the pattern and color traits are not of equivalent value
as factors of heredity; but both should be considered in their interplay of
relative significance.
330
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 1
Table of standards for the classification of the colors and patterns in the genus
Karanasa. Plate prepared by A. Avinoff.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 15
Plate 1
332
Annals of the Carnegie Museum
VOL. 31
EXPLANATION OF PLATE 2
Schema for the analysis of the undersides of the wing in Karanasa and related
genera. Plate prepared by A. Avinoff.
ANNALS, CARNEGIE MUSEUM, Vol. 31
Art. 15
Plate 2
DIAGRAMMATIC NOTATIONS ON THE UNDERSIDE
OF THE HIND VING OF KARANASA
>i»d
1 igfit dark
>\ White veins : i slig-ht
^ I medium
acute obtuse 3 prominent
X
Y
3
X (standard)
a
Y (standard]
Y fn
aKsuensis Yda3
Y
Y
laiefasciala Yda^
regeli
Ydad
SATYRU3
'wyssii
fidia
parfeatis
gejeri
K" - )
t 3
M
mmi
rwr ■
Bi)
--Y/W
.»1
Jeechi
Xla.2
arpensls
■ JKla2
Iskander Xdo2
dissoluta
Ydo2
y'-:Y
wjw
iii:.
li
m.:mm
intermedia
Ylat
erubestens Yda2
cadesia. Ydo2^
astonca
Yda^
M WM
mm
mmm
fc':l
rs
>Y
regulx
Ydoa
angrena
Ydol
b]astai;ana YdaS-
tolorica
Yia^
Yd o3
pami
Ydo3
tmm
m
ornata
mvim.
alpheTaKyt Ydo3
safeda Yda2. rotorovsK}!! Yd&S, hoffinarmi Y]a2
COMPARISON OF THE SHAPE OFDISCAL BANDS
^ m
gilgitica YdaY
voigti
yda2
NEOPE
dissoluta
<1 4.
turugensis
ridingsi
4 i’ -
ridingsi t , Y '
■i
ART. 16. A NEW RACE OF HUDSONTAX CHICKADEE
By W. E. Clyde Todd
Several years ago, with a fine series of 136 specimens of the Hudsonian
Chickadee at my command, by far the larger part of which was collected
expressly for the purpose of this investigation, I prepared an extensive
manuscript on the taxonomy of this species for my projected report on the
birds of the Labrador Peninsula. Presentation of these notes in their
original form is necessarily deferred, but now that we are looking forward
to the publication of a comprehensive stud}' of this species by a competent
Canadian ornithologist, it may be in order to put on record at least that
part of them which deals with an apparently new race from the coast of
Labrador. Since my earlier study, 35 specimens have been added to the
Carnegie Museum series of this species, and I have also had the benefit of
additional material lent for examination by the authorities of the LTnited
States National Museum, the Academy of Natural Sciences of Phila-
delphia, and the Royal Ontario Museum of Zoology, to the authorities of
which institutions I desire to express my thanks. A brief resume of the
taxonomic history of this species is a necessary preliminary.
Pants hudsoniciis was described by Forster (Philosophical Transac-
tions, 62, 1772, pp. 383, 408, 430) from the mouth of the Severn River,
on the west coast of Hudson Bay. Before any specimens from this point
were available, I found that birds from Churchill to the north, and from
James Bay and northern Ontario on the south, were comparable in every
respect — a judgment that was fully confirmed by actual examination of a
series from the type-locality. The range of this, the typical race, extends
eastward into the Labrador Peninsula and northward to the limit of trees.
Specimens from the north shore of the Gulf of St. Lawrence (“wfgHca/zi'”
of Townsend) are not satisfactorily distinguishable from others from Fort
Chimo (“wwgam” Rhoads) and Richmond Gulf. Recent authors (including
Ridgway, 1904), however, recognize liitoralis (Bryant, 1865) for the birds
of Nova Scotia and New Brunswick. With specimens from the type-
locality (Yarmouth, Nova Scotia), collected by myself in 1901, and with
considerable borrowed material from other localities in these provinces,
I must confess that I am very dubious about the validity of liitoralis. In
the specimens examined, there is much variation in exact shade of color,
and the average smaller size seems to be inconsequential. A larger series of
really good specimens, comparable for season, might suffice to establish
its validity, but as matters stand I consider it doubtful.
333
Issued October 27, 1950.
tJCT 3 1 1950
334
With Rhoads’ type-series of columbianus before me, I am quite unable
to distinguish this supposed race, but again my present material is insuf-
ficient for a final conclusion. The northwestern race evura of Coues (1884)
is easily recognizable by its larger size and somewhat different coloration.
Leaving these western birds, and returning to the East, I find that the
recently described mbbittsi of Burleigh (1948) is barely distinguishable in
fresh plumage by the characters assigned in the original description,
namely, the slight grayish cast of the underparts (as compared with
typical hudsoniciis) , the duller and darker brown flanks (nearer snuff
brown than sayal brown), and the slightly duller and darker brown upper-
parts, with less contrast between the pileum and back than in the typical
bird. In breeding dress these distinctions are mostly obscured.
Specimens from the coast of Labrador, however, cannot be referred to
any of the described races. They are certainly as worthy of subspecific
status as are the Newfoundland birds. In naming them herewith I must
explain that, following Ridgway, I continue to use Penthestes for the
generic group instead of Pams.
Penthestes hudsonicus labradorius, subsp. nov.
Ten specimens: Nain, Mokkovik, and Rigolet, Labrador (August 29 —
September 25).
Type, No. 100,125, Collection Carnegie Museum, immature male;
Rigolet, Labrador, September 24, 1926; W. E. Clyde Todd.
Subspecific characters.—SimWea: to Penthestes hudsonicus hudsonicus
(Forster) of Hudson Bay, etc., but upperparts slightly duller brown;
underparts more purely and more extensively white; and brown color of
flanks paler, duller, and more restricted. Wing (type), 67; tail, 61; bill,
10; tarsus, 15.5.
Range. — Coast of Labrador.
Remarks. — As a series, these specimens may readily be separated from
James Bay birds, as their characters stand out sufficiently well, but indi-
vidual birds might be hard to distinguish. This is by far the whitest (below)
of all the forms of P. hudsoniciis. All of the series on which it is based
happen to be birds of the year in fresh plumage, which show the characters
to good advantage. Whether spring and summer specimens can be as
readily discriminated remains to be seen. They are not so satisfactory for
comparison as a rule — an observation which applies with equal force to
all individuals of this species.
ART. 17. A NEW GEOMYID RODENT FR
MIOCENE OF MONTANA
By Albert E. Wood
Biology Laboratory, Amherst College, Amherst, Massachusetts
Recently, Dr. J. LeRoy Kay of the Carnegie Museum sent me a rodent
jaw from the Miocene of Montana for identification. This specimen repre-
sents a member of the Entoptychinae, the dominant Lower Miocene sub-
family of the Geomyidse or pocket gophers. The specimen indicates that
this represents a member of the group which differs from all hitherto des-
cribed specimens sufficiently to warrant its being established as a new
species.
I wish to express my deep appreciation to Dr. Kay for permitting me to
describe this specimen. This study was assisted by a grant from the Marsh
Fund of the National Academy of Sciences.
Gregorymys kayi, sp. nov.
Holoiype: Carnegie Museum No. 8,999, left lower jaw with incisor
and P4-M3.
Horizon and Locality: Miocene, Six Mile Creek, 5 miles west of Toston,
Montana, field no. 43, collected by J. LeRoy Kay, 1948.
Diagnosis: Similar to G. douglassi, but apparently smaller: much greater
development of cement around the roots, extending well up the sides of
the crown, more than in G. douglassi or G. montanensis; enamel greatly
thinned or even absent on anterior faces of molars ; roots present.
This species differs from the other members of the genus* in the very
extensive development of cement. This not only covers the roots, but
extends up the sides of the crown to levels well above the bottom of the
pattern. The enamel is thinned on the anterior side of the teeth much more
than is the case in other species of the genus, though it does not appear
*Wood, Albert E. 1936. Geomyid rodents from the middle Tertiary. American
Museum Novitales, no. 866, 31 pp., 33 figs.
Hibbard, Claude W., and Kendall A. Keenmon. 1950. New evidence of the
Lower Miocene Age of the Blacktail Deer Creek Formation in Montana. Contrib.
Mus. Paleont., Univ. Michigan, vol. 8, no. 7, pp. 193-204, 3 figs., 1 map. According
to a letter from Dr. Hibbard, dated November 21, 1950, there is no cement on the
sides of the crowns of G. montanensis, even well down in the alveoli.
335
Issued December 21, 1950.
5£C % ®
336
Annals of the Carnegie Museum
VoL, 31
to be interrupted at the stage of wear represented by the specimen. How-
ever it seems probable that at a somewhat more advanced stage of wear
the enamel would be interrupted, thus coming within the definition of
Entoptychus as given by Wood (1936). However, the fact that roots are
clearly present, and that the crowns are not exceptionally high, justifies
the inclusion of this form in Gregorymys. The cement extends as a thin
layer along the anterior face of the molars, although interdental wear re-
moves it very largely.
In all the molars, the central valley has been transformed into a cres-
centic lake, with its concave side directed anterad (fig. 1 b). No trace of
separate cusps seems to be visible in the molars. Mi is the largest of the
molars, and M3 is the smallest.
In the premolar, the central valley has the same shape as in the molars,
but opens broadly along the lingual side. There is, however, a lingual
dam some distance down the side of the crown, so that the valley would
eventually be transformed into the same lake as is seen in the molars. In
the talonid, there is no trace of details of the pattern. In the trigonid,
however, there are faint irregularities in the enamel outline, suggesting
the last traces of cusps. At the antero-lingual corner there is a marked
valley, extending only a short distance below the wear surface, which
appears to represent the last remnants of a valley behind the anteroconid.
That is, as in other species of Gregorymys, the trigonid of P4 must have had
a well developed group of accessory cusps. The premolar is the largest of
the teeth.
The incisor has a broad, flat anterior face, with the enamel only just
reaching the median and lateral surfaces (fig. 1 a). The tooth is a broad
and very efficient cutting tool. A peculiarity, presumably of this indi-
vidual, is the narrowing of the incisor from both ends, so that it is dis-
tinctly more slender at a point just inside the alveolus than it is either
nearer the tip of the tooth or nearer the root (see measurements). The
extra-alveolar enamel is orange-brown in color.
The jaw is not exceptionally heavy for a geomyid. The ventral border of
the masseteric crest extends nearly straight laterad, below the cheek teeth,
and runs nearly horizontally (fig. 1 c). The ascending ramus rises steeply
by the middle of M2. The diastema is short and deeply notched. The
mental foramen lies beneath the deepest point of this notch, just ventrad
of the anterior end of the masseteric fossa. The symphysis is at a sharp
angle with the horizontal ramus, so that, viewed from above, the two
jaws must have diverged markedly. It is also heavily pitted and cor-
1950
Wood: New Rodent from Miocene of Montana
337
ruga ted, showing that there was no motion between the mandibles, a
fact in accord with the structure of the incisor. Just below the main part of
the symphysis is a deep pit. The angle is reduced but is markedly inverted,
being of the general entoptychine type, in contradistinction to Thomomys
and Geomys where the angle is non-existent.
This form is obviously similar to Gregormys curtiis, G. montanensis, and
G. doHglassi, the last two also coming from Montana. Although G.
b
Fig. 1. Gregorymys kayi, Carnegie Museum No. 8,999.
All figures, X 4.
a. Incisori left, view of anterior face.
b. P4— Ma left.
c. Lateral view of lower jaw.
douglassi is known only from a skull, the present form is clearly too large
to belong with that skull. A more important difference lies in the great
expanse of the cement in G. kayi, which would seem to be a progressive
character. The sequence G. ciirtiis — G. douglassi — G. kayi seems to repre-
sent a structural line of rather uniform morphology but with gradually
increasing amounts of cement and with a gradual approach toward the
condition found in Ejitoptychus. G. mo7itanensis would represent a re-
lated side-branch in which the amount of cement had been secondarily
reduced, but which otherwise was very close to G. kayi.
G. kayi seems to represent a culmination of the Gregorymys evolutionary
338
Annals of the Carnegie Museum
VoL. 31
line, being somewhat the most specialized member of the genus. In its
specializations, it shows a number of features characteristic of the related
genus Entoptychus. These, however, certainly represent parallelisms within
the Gregorymys line to the trends within the contemporary Entoptychus,
where Entoptychus, at any given period, is structurally more advanced
than the members of Gregorymys, ]ust as the entoptychines as a whole paral-
lel but are more advanced than the contemporary heteromyids.
Measurements of Gregorymys kayi
(Carnegie Museum No. 8,999)
P4 — Mg, alveolar distance
7 . 80 mm.
crown surface
P4 antero-posterior
6.90
2.37
1.80
2.24
1.53
2.26
width trigonid
width talonid
Ml antero-posterior
width trigonid
width talonid
ca. 2 . 30
M2 antero-posterior
1.60
width trigonid
width talonid
over 2 . 05
over 2 . 00
Mg antero-posterior
1.58 (crown)
width trigonid
width talonid
1.83
1.67
2.06
2.04
1.96
2.05
Ii antero-posterior
transverse, at tip
transverse, just inside alveolus
transverse, beneath cheek teeth
y'r-
ART. 18. THE MAMMALS OF THE MAZINAW LAKE REGION
OF ONTARIO; THEIR REPRODUCTION AND
POPULATION DYNAMICS*
By John J. Christian
The Wyeth Institute of Applied Biochemistry
Philadelphia, 30, Pennsylvania
1. Introduction
The mammals of the Mazinaw Lake region of southern Ontario were
investigated during the first two weeks of July, 1949, and July, 1950. The
following is a report on these investigations.
The region lies in the Canadian Precambrian shield section of Lennox
and Addington Counties, Ontario. The particular area covered in this
account is a north and south strip about twenty-five miles long following
highway 41. The west shore of Mazinaw Lake forms the middle of this
strip. The lake itself is seven miles long and from one-half to three miles
wide, and is one of the larger lakes in the region. All trap locations were
within a mile and a quarter from the highway.
The general elevation varies from 850 to 1,250 feet, rising continuously
to the northwest to the Algonquin Park area about sixty miles away.
The region falls away to the south to meet the great lakes plains, and to
the north to meet the Ottawa River valley. The whole region is rolling
and rocky with numerous extensive outcrops and bare spots resulting
from extensive glaciation. Lakes are numerous, and the general altitude of
the low areas is so nearly the same that the drainage from lake to lake
flows down only a slight gradient, resulting in slow-flowing, sluggish
streams. These traverse numerous boggy areas, and recent lake or pond
fills are numerous. The poorly drained, low areas possess a rich acid soil,
while the hillsides and uplands are relatively dry and rocky with a thin,
sandy, subacid soil. The hillsides are drained by numerous, small, rocky
streams which rapidly go dry under drought conditions. Springs are
scarce.
Along the eastern shore of Mazinaw Lake, and extending considerably
north and south, is a relatively recent fault reaching a height of 1,450
feet from the lake bottom to its highest point. This is responsible for a
series of north and south lakes and bogs, and for forming the main drain-
age of the Mazinaw region.
Halliday (1937) puts this area in the Algonquin-Laurentides section of
Issued February 19, 1951.
2 6 fact
340
Annals of the Carnegie Museum
VOL. 31
the Great Lakes-St. Lawrence forest region, which he characterizes as
follows: . the bed-rock is part of the great Precambrian Shield of
Canada, and consists largely of crystalline limestones (Grenville series),
schists, and gneisses of the altered sedimentaries and granite intrusives.
The topography is rough and irregular, and glacial deposits of varied
character, chiefly of somewhat light texture, cover the greater part. In
addition there are some lacustrine deposits from the Nipissing-Great
Lakes and Algonquin periods. A podsol type of soil is to be expected, but
areas of gray-brown and brown forest soils may be present.
“In this section, white pine probably reached its maximum development
in Canada, but extensive lumbering and fire have removed the greater
part. Red pine has also been a prominent species, especially on the Al-
gonquin Highlands. In spite of the previous dominance of these species
and the presence of intrusive conifers from the boreal forest region, the
general character is that of a mixed forest, and the dominant or competi-
tive'association is one of sugar maple, yellow birch, hemlock, and white
pine. In addition there are varying amounts of basswood, white spruce,
balsam fir, beech, (Northern) red oak, elm, white ash, red maple, iron-
wood, white birch and large-toothed aspen. The composition of this asso-
ciation changes somewhat to the north, as hemlock, (Northern) red oak,
and beech decrease numerically and finally drop out before the limits of
the Section are reached and the proportion of yellow birch, white spruce,
balsam fir, and white birch increases. . . . Throughout the Section, areas
of hardwood occur on the ridge tops and on heavier soil deposits, and
black spruce, tamarack, and some cedar are found in swampy depressions.”
The Mazinaw region fits into Halliday’s description for the more north-
ern limits of the Algonquin-Laurentides Section, probably as a result of
its altitude. Beech and hemlock are scarce, and the northern red oak is
confined to the drier areas. The uplands support a hardwood forest with
sugar maple dominant. Cut-over areas have grown up to aspen (both
large-toothed and trembling), white birch, red maple, red oak, and some
white spruce and fir. Pure conifer stands of white spruce and fir occupy
the lower dry areas, while extensive black spruce and tamarack bogs
abound. These latter, when cut over, come up in alder and willow. Black
ash-white cedar bogs are abundant. About seven miles to the west of
Mazinaw Lake there is still a large stand of virgin white pine surrounding
Weslemkoon Lake. This stand, in which pines four feet d.b.h. are re-
portedly common, is being lumbered around its edges at the present time,
but only in winter when the logs can be skidded out. Besides these forest
1951 Christian: Mammals of Mazinaw Region, Ontario 341
types, there are numerous high “balds” of bare rock alternating with
extensive areas of blueberries.
Extensive timbering occurred sixty years ago and the area has been
burned over two or three times since, the last fire having taken place
about twenty years ago. It is evident at present that white pine will again
be the climax forest type, as even now they are beginning to crowd out the
aspens and birches on the hillside areas. Pine seedlings of all sizes abound.
A small area of each general forest type was selected for trapping, and
as far as possible these areas were chosen so that they were in a continuous
habitat of the same type. The following habitats were trapped in, and
further discussions of the vegetation will appear under the heading de-
scribing the specific area.
I. Dry, rocky, mixed deciduous-conifer woods: deciduous vegetation
dominant.
II. Low, wet, deciduous woods with a few patches of conifers, and
boggy areas.
III. Black ash-northern white cedar bog.
IV. Dry old field. Danthonia dominant.
V. Open Vaccinium-sphagnum bog surrounded by spruce and tama-
rack.
VI. Dominantly deciduous woods with alder bogs and small open
sedge bogs.
VII. Small sphagnum-spruce-fir bog in a more extensive mixed woods.
VIII. Open, sedge marsh.
IX. High blueberry bog.
X. Pure, dense, black spruce bog; white spruce and fir on surrounding
drier areas.
The climate of the area is classified by Halliday (1937) as temperate
and humid plus, with moisture abundant at all seasons. Thirty inches of
snow is seldom exceeded at any time. The lowest temperature so far re-
corded at Mazinaw Lake is 52° below zero F., but ordinarily winter tem-
peratures do not go below — 20°F. In 1949, however, drought conditions
prevailed for two months from the end of April until the end of the first
week in July, with the final forty days without any rain whatsoever. Bogs
were for the most part without standing water, and most streams were
either totally dry or with only a slight trickle. More normal conditions of
precipitation prevailed in 1950, and during the first two weeks of July
there was from a foot to eighteen inches of water standing in the bogs
which had been dry the previous year.
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Annals of the Carnegie Museum
VOL. 31
In 1950 a much colder spring and a later summer prevailed than in
1949. This difference was reflected in the plants. In 1949 the hot, dry-
spring was from two to three weeks ahead of the colder, wet spring of
1950. Blueberries and Aralia berries were ripe by July 1, 1949, but were
just beginning to ripen by July 14, 1950, and this difference was true of
many other blooms and fruits.
The small mammals were trapped with regular snap-back traps. The
bait used was either pure ham-fat rubbed on the trap, or a mixture of ham-
fat, peanut butter, and almond extract. Smears of the testis and epididymis
were made of each male and were stained with haematoxylin-eosin
(Christian, 1950a). The reproductive tracts of the females were preserved
entire.
The fur-bearers and large animals were not trapped, although a few in
good condition were picked up from the highway. Information regarding
these mammals was, for the most part, obtained by questioning local
trappers, and residents, and others who were familiar with the local mam-
mals. Most of this information was obtained from Albert Spencer and
Irving Brown, both reliable and experienced trappers in the area, and from
Fred Garbutt, an interested and observant local resident.
The following account has been divided into four sections: the habitats
trapped in, accounts by species, discussion of populations, and informa-
tion on reproduction and its relation to population dynamics.
II. Discussion of Specific Habitats
Ten habitats were chosen for trapping as being representative of the
region, and these are discussed in detail below. All references to habitats
following the discussion of a mammal refer to the designating numeral of
the specific habitat area.
I. The dry, rocky, steep hillside of the west shore of Mazinaw Lake, seven
miles north of Cloyne, Ontario. Elevation goo feet. Trapped in iQ4g and ig^o.
This area is the dry hillside comprising the west shore of Mazinaw Lake,
which is completely wooded except for a few old cleared areas. The forest
is primarily deciduous and is composed of sugar maple, aspen, white birch,
and some white and red pines. Numerous bare boulders and outcrops of
quartzite are scattered through the area providing many cracks and
crevices in which small mammals may take refuge. The hillside is well
drained. The sandy, thin, and subacid soil has been formed by the break-
down of metamorphosed sandstones. Wild sarsaparilla, wintergreen, bush
1951 Christian: Mammals of Mazinaw Region, Ontario 343
honeysuckle, and low sweet blueberry form the principal ground cover.
The area was burned and lumbered in the past and now represents a
maturing second growth.
The difference in seasons between similar dates in 1949 and 1950 is
plainly shown by the various flowering plants. Bunchberry was in berry
in 1949, while at the same time in 1950 it was only in full bloom, even in
warm, open spots. Aralia berries were ripe in 1949, but the plant was just
past blooming in 1950. The bush honeysuckle was well past blooming in
in 1949, but was in full bloom in 1950. Blueberries were ripe the first of
July in 1949, but were only beginning to ripen at the end of the second
week of July in 1950.
The following is a list of the principal plants of the area. The more
dominant forms are preceded by an asterisk in this and all following lists
of flora. More than one asterisk indicates that the form is dominant al-
most to the exclusion of others.
White pine {Pinus strohus)
Red pine {Pinus resinosa)
Balsam fir {Abies balsamea)
Hemlock {Tsuga canadensis)
*Trembling aspen {Populus tremuloides)
*Large-toothed aspen {Populus grandidentata)
*White birch {Betula papyrifera)
Red oak {Quercus borealis)
*Sugar maple {Acer saccharum)
Red maple {Acer rubrum)
*Smooth-leaved shadbush {Amelanchier laevis)
Moose wood {Acer pensylvanicum)
*Bunchberry {Cornus canadensis) — berry, 1949; bloom, 1950.
*Low sweet blueberries {Vaccinium pennsylvanicum) — berry, 1949.
*Bush honeysuckle {Diervilla lonicera) — Bloom, 1950.
*Wintergreen {Gaultheria procumbens)
Running ground pine {Lycopodium complanatum)
Running clubmoss {Lycopodium clavatum)
Rock polypody {Polypodium virginianum)
Bracken {Pteridium latiusculum)
*Heartleaf lily {Maianihemum canadense) — just past bloom, 1950.
*Fireweed {Epilobium augustifolium) — in bloom, 1949 and 1950.
**Wild sarsaparilla {Aralia nudicaulis) — berries ripe, 1949.
Green pipsissewa {Chimaphila umbellata)
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Annals of the Carnegie Museum
VOL. 31
A list of the birds found in the area follows. Both the yellow-bellied
sapsucker and the least flycatcher were known to be nesting in the im-
mediate area, while the others, if not nesting in the immediate area, were
nesting in the vicinity.
Spotted sandpiper {Actitis macularia)
Ring-billed gull {Larus delawarensis)
Whip-poor-will {Antrostomus vociferus)
Kingfisher {Megaceryle alcyon)
Ruby-throated hummingbird {Archilochus colubris)
Flicker {Colaptes auratus)
*Yellow-bellied sapsucker {Sphyrapicus varius)
Crested flycatcher {Myiarchus crinitus) — abundant 1949, scarce 1950.
*Least flycatcher {Empidonax minimus)
Robin {Turdus migratorius) — abundant 1949, less so 1950.
Veery {Hylocichla fuscescens)
*Cedar waxwing {Bombycilla cedrorum)
**Red-eyed vireo {Vireo olivaceous)
Magnolia warbler {Dendroica magnolia) — 1950 only; immature.
Myrtle warbler {Dendroica coronata)
Pine warbler {Dendroica pinus) — 1949 only.
Oven-bird {Seiurus aurocapillus)
Purple finch {Carpodacus purpureus)
Goldfinch {S pinus tristis)
*Chipping sparrow {Spizella passerina)
Song sparrow {Melospiza melodia)
One blue-tailed skink {Eumeces fas ciatus) was collected from among the
rocks at the lake-edge in this habitat.
Mammals: Since the area is relatively dry, there is a large chipmunk
population in and around the rocks. Skunks are common, probably at-
tracted by the garbage from the few cottages in the area. One half-grown
and one very emaciated adult female were collected. Another young
Mephitis, a litter mate of the one collected, was also known to be in the
area, and possibly others were present. Red squirrels were at one time
abundant, but have been completely ‘^shot out.” At least one mink is
known to have been in the area. Porcupines have wandered in and out,
but are shot on sight. One young racoon was found dead, apparently a
highway casualty, as the animal was entire, including its pelt.
In 1949, eighteen traps were set in the area in places looking suitable
for small mammals, such as rock crevices, along fallen logs, and at the
1951
Christian; Mammals of Mazinaw Region, Ontario
345
bases of rocks. These remained for three nights, during which time one
Peromyscus maniculatus gracilis, three Peromyscus leucopus novehoracensis,
and two Blarina brevicauda talpoides were captured, making nine trap-
nights, per catch.
In 1950, no mouse traps were set, but five rat traps were placed in likely
spots for chipmunks. None was caught, but one Blarina fell victim. Later
the traps were moved and baited with bacon rind for flying squirrels.
These traps caught one juvenile Peromyscus maniculatus gracilis the first
night and two nights later took a half-grown Mephitis.
Trapping Summary — 1949
Date: July 3 4
Peromyscus m. gracilis 0 0
Peromyscus 1. novehoracensis 1 1
Blarina b. talpoides 1 1
Total 2 2
6 catches in 54 trap-nights, or l/9 trap-nights.
Summary 1950: 50 trap-nights (rat traps) — 3 catches or l/l7 trap-nights.
II. .4 damp, second- growth, deciduous woods along Bon Echo Creek, five
and one-half miles north of Cloyne, Ontario. Elevation goo feet. Trapped in
only in 1949.
Bon Echo Creek flows from Bon Echo Lake to Mazinaw Lake, where it
empties on the west shore. The gradient from Bon Echo to Mazinaw is
slight, resulting in the creek being a slow-flowing stream traversing a
poorly drained area with many bogs. The soil is rich, black, and acid, and
the creek is stained dark from this acid bog soil through which it flows.
The surrounding forest is composed primarily of second-growth deciduous
trees with some firs, and patches of white and black spruces in the drier
and wetter areas respectively. An occasional white pine still stands. Most
of the growth has developed since the area was burned over about twenty
years ago. A road passes through the area, and ends at an abandoned
lumber camp at the east end of Bon Echo Lake. The undercover is very
dense with a large percentage of alder, aspen, and white birch saplings.
A list of the dominant vegetation follows.
White pine {Pinus strobus)
White spruce {Picea glauca)
*Black spruce {Picea mariana)
*Balsam fir {Abies balsamea)
*Large-toothed aspen {Populus grandidentata)
*Trembling aspen {Populus tremuloides)
Total
1
3
2
6
346
Annals of the Carnegie Museum
VOL. 31
Yellow birch {Betula lutea)
*White birch {Betula papyrifera)
*Speckled alder {Alnus incana)
White elm ( Ulmus americana)
Moosewood {Acer pennsylvanicum)
*Sugar maple {Acer saccharum)
*Red maple {Acer ruhrum)
Sensitive fern {Onoclea sensihilis)
Thin-leafed pyrola {Pyrola elUptica)
This habitat is of a more southern type than any of the others trapped,
and approaches the transition (upper austral) zone in character.
The birds noted in the area are:
Ruffed grouse {Bonasa umhellus) — covey of 6 young.
Woodcock {Philohela minor)
Hummingbird {Archilochus colubris)
Crested flycatcher {Myiarchus crinitus)
Phoebe {Sayornis phoehe)
Robins {Turdus migratorius)
Mammals: Deer tracks were seen along the old dirt road. Every time
the area was visited a Lepus americanus, apparently the same one, was
seen alongside the road in approximately the same spot. On every oc-
casion I was able to approach within a few feet of this animal without
unduly alarming it.
Forty-nine traps were set in the area with thirty- three through the
moist woodland and along the stream in likely looking spots. An especially
promising site for voles was a rocky portion of a grass-grown, abandoned
road. Seven traps were set in a small, bog area with black spruce saplings
and alders growing in a wet black soil, and nine in a dry patch of white
spruces. These traps were left set for only two nights.
Two Peromyscus maniculatus gracilis, one Peromyscus leucopus nove-
horacensis, and four Blarina were caught in this area. One Microtus penn-
sylvanicus was caught in the grassy spot mentioned above, but was eaten,
apparently by a shrew, and could not be saved as a specimen. A total of
eight mammals caught meant 1 catch per 12.25 trap-nights.
Trapping Summary — 1949
Date: July
3
2
4
0
0
2
0
2
Total
Peromyscus m. gracilis
Peromyscus 1. novehoracensis
Blarina b. talpoides
Microtus p. pennsylvanicus
1
2
1
6
2
1
4
1
8
Total
1951 Christian: Mammals of Mazinaw Region, Ontario 347
III. ^ dense, cool, black ash-northern white cedar hog, five and one-half
miles south of Denbigh, Ontario. Elevation Q^o feet. Trapped in IQ4Q and
1950.
A typical, dense, cool, northern white cedar-black ash bog, which has
apparently been by-passed by recent-past fires or lumbering operations.
The actual bog area is approximately fifty yards wide and extends north
and south about a half-mile, draining into a small mill pond at the south.
The bog is criss-crossed with fallen white cedar logs, subdividing the area
into a series of small, disconnected “boglets.” These vary from a few feet
in each dimension up to about twenty by fifty feet. The black, acid,
mucky soil supports a dense growth of sphagnum which continues on,
with other mosses, over fallen logs, stumps, and hummocks. Liverworts
and Seliginella were also included with the mosses. In 1949, there was no
standing water in the bog except in the center of one very large boglet,
where there were a few inches. The area remained wet, however, in spite
of the lack of standing water. In 1950, there were between twelve and
eighteen inches of water throughout the bog, and this was added to in the
first week of July by the almost daily rains. The hills forming the east and
west boundaries of the bog were covered chiefly with white and black
spruces and firs. Black spruces and firs also occurred in the bog, but were
not among the dominant forms. The full grown, standing, white cedars,
and the great majority of the cedars were of this type, averaged a foot
d.b.h., while the fallen trunks of these trees were of this size or larger.
The relatively uniform size of the standing cedars probably indicates the
growth since some past lumbering operations or fires. The surrounding
spruces were mature trees in the neighborhood of fifty or sixty feet tall.
The bog abutted abruptly against a low, fault scarp on the east. This
scarp varied from a few to over thirty feet in height. Under the sphagnum-
covered talus at its base was a continuous “tunnel-run” which produced
the usual high catches of an “edge” habitat. This scarp appears to be a
northward continuation of that forming the eastern shore of Mazinaw
Lake, and is at the northern end of a continuous valley containing a series
of bogs and small and large lakes. It is also the center of drainage for the
immediately surrounding area.
A list of the major plant forms follows:
*White spruce {Picea glauca) — adjoining dry areas.
* Black spruce {Picea mariana)
*Balsam fir {Abies balsamea)
**Northern white cedar {Thuja occidentalis)
348
Annals of the Carnegie Museum
VOL. 31
Trembling aspen {Populus tremuloides)
White birch {Betula papyrifera)
Speckled alder {Alnus incana)
White elm ( Ulmus americana)
Moosewood {Acer pennsylvanicum)
Black alder {Ilex hronxensis)
Alder-leaved buckthorn {Rhamnus alnifolia)
Red baneberry {Actaea rubra) — in berry 1949.
White baneberry {Actaea alba) — in berry 1949.
Squashberry {Viburnum pauciflorum) — in berry 1949.
Labrador tea {Ledum groenlandicum)
**Various mosses and sphagnum.
Thallus liverworts
Bunchberry {Cornus canadensis) — in berry 1949; in bloom 1950.
*Small cx2inherry {Vaccinium oxycoccos)
*Wood sorrel {Oxalis montana) — in bloom 1950.
*Barren strawberry {Waldsteinia fragar aides) — in bloom 1950.
Arctic bramble {Rubus borealis) — in bloom and berry 1950.
Yellow bead lily {Clintonia borealis) — in bloom 1950.
Wild sarsaparilla {Aralia nudicaulis) — in berry 1949; just past bloom
1950.
Cinnamon fern {Osmunda cinnamomea)
Shining club moss {Lycopodium lucidulum v. occidentale)
Running club moss {Lycopodium clavatum) — adjoining dry areas.
Round -branch ground pine {Lycopodium obscurum v. dendroideum) —
dry areas.
From the comments in this list it again becomes evident how much
later the 1950 season was than the same period in 1949.
The birds noted in the area follow:
Horned grebe {Colymbus auritus)
Broad-winged hawk {Buteo platypterus)
Flicker {Colaptes auratus)
Pileated woodpecker {Ceophloeus pileatus) — only in 1949; one seen.
Yellow-bellied sapsucker {Sphyrapicus varius)
Hairy woodpecker {Dry abates villa sus)
Downy woodpecker {Dry abates pubescens)
Least flycatcher {Empidonax minimus)
Eastern wood pewee {Myiochanes virens)
Bluejay {Cyanocitta cristata)
1951 Christian: Mammals OF Mazinaw Region, Ontario 349
Black-capped chickadee {Penthestes atricapillus)
Tufted titmouse {Bacolophus bicolor) — 1949 only.
White-breasted nuthatch {Sitta carolinensis) — 1949 only.
Red-breasted nuthatch {Sitta canadensis)
Brown creeper {Certhia familiaris)
Blue-headed vireo {Vireo solitarius)
Black and white warbler {Mniotilta varia)
Ovenbird {Seiurus aurocapillus)
Canada warblers {Wilsonia canadensis)
Bronzed grackles {Quiscalus quiscula)
Rose-breasted grosbeak {Hedymeles ludovicianus)
Junco {Junco hyemalis)
The drought conditions prevailing during 1949 may account for the
birds (especially the titmice) observed in this area in that year, which
were not seen in 1950. In 1950, however, the breeding bird population was
noticeably higher than in the previous year; for example, there were at
least two pairs of rose-breasted grosbeaks in 1950 in this relatively small,
trapping area. The warblers were conspicuously abundant in the latter
year.
Mammals: In 1949, Tamiasciuris and Marmota were seen in this
habitat. The former were abundant, although efforts to trap them failed.
The red-squirrels were conspicuous by their absence in 1950, nor were any
Marmota seen. Chipmunks were abundant in the dry areas surrounding
the bog in both years.
In 1949 the area was trapped for six nights. Traps were placed along
fallen logs, at openings of runs in the centers of boglets, under over-
hanging hummocks, and in runs along the scarp base. The traps were
concentrated and the area was trapped in two sub-areas at two separate
times. In the first area, for the first night, there were eighty-seven traps
which were reduced to fifty-three for the following two nights by removing
one or more from each station. Sixty-six traps were used for the last three
nights in a part of the bog north of the first area, making a total for the
area of 391 trap-nights. The trapping summary appears below.
One Condylura and one Synaptomys were trapped at the fault base run.
Four Clethrionomys were taken in and around the western edges of the bog
where it adjoined the white spruce-fir covered hillside. Blarina was found
on the dry hillside next to the bog, while the long-tailed shrews {Soricidae)
were all taken in the boglets throughout the swamp. Peromyscus leucopus
novahoracensis was taken mainly in drier areas, while the one Peromyscus
350
Annals of the Carnegie Museum
VOL. 31
maniculatus gracilis was taken in the wet area. A total of sixteen mam-
mals caught meant 1 catch per 24 trap-nights.
Trapping Summary — 1949
Date; July
5 (87)
6 (53)
7
11 (66)
12
13
Total
Condylura c. cristata
1
0
0
0
0
0
1
Sorex c. cinereus
0
0
1
1
0
0
2
Sorex f. fumeus
0
0
0
0
1
0
1
Sorex p. albiharhus
1
0
0
0
0
0
1
Blarina h. talpoides
0
0
1
0
1
0
2
Peromyscus m. gracilis
0
0
0
0
0
1
1
Peromyscus 1. novehoracensis
1
0
0
0
1
0
3
Synaptomys c. cooperi
1
0
0
0
0
0
1
Clethrionomys g. gapperi
0
1
0
3
0
0
4
Total
4
1
2
5
3
1
16
In 1950 this area was re-trapped intensively for several reasons: (1) to
compare the population with last year, (2) to obtain more shrews, and (3)
because of the large number of species obtained in 1949 in such a limited
area and habitat. Much the same trap concentration was used as in 1949,
but a somewhat wider area was covered, with more traps. Due to the stand-
ing water this year, no traps were placed in the boglet bottoms, being
necessarily placed in dry spots around the edges. Traps, 101 in number,
were placed across the bog, along the cliff base, and back across the bog
again for two nights. For the next three nights the same number of traps
were in the bog, but sixty-one of them were removed from the original
sets and re-set in other areas, with about half of these being along the
scarp base. For the following three nights seventy-two of the scarp and
last-set bog traps (in an area similar to the conditions of last year with no
standing water) were left in place, the remaining twenty-nine in the bog
center having been picked up, since no mammals were caught in that
area. This constituted a total of 721 trap-nights under trap concentrations
and movements similar to those of 1949.
No rodents were caught in this area in 1950, and no mammals were
caught in the center of the bog in areas with or without standing water.
One specimen of Microsorex, two of Sorex cinereus, and one of Sorex
fumeus were caught in the scarp-base run. One Blarina was caught in a
dry spot at the western edge. Four of these five mammals were caught the
first two nights, and the fifth {Sorex cinereus) three nights later. This
strongly suggests that the four represented the total small mammal popu-
lation of the bog, and that the fifth was an outside wanderer that came in.
It seems reasonable that the water level made the difference in the loca-
1951
Christian: Mammals of Mazinaw Region, Ontario
351
tion of the shrews within the habitat in the two years. In 1949, with a wet,
black muck and no standing water, the shrews were scattered over the
bog, but in 1950, with abundant water, no shrews were in the center of
the bog, all having been taken at the bog’s eastern margin in the scarp-
base run. The total catch of five mammals was the equivalent of 1 catch
per 144 trap-nights.
Trapping Summary — 1950
Date: July
Microsorex h. inter-
vectus
Blarina h. talpoides
Sorex c. cinereus
Sorex f. fumeus
Totals
5 (61
3 (101) 4 reset)
1 0 0
1 0 0
0 10
0 1 0
2 2 0
6 78 (72)
0 0 0
0 0 0
0 1 0
0 0 0
0 1 0
9 10 Total
0 0 1
0 0 1
0 0 2
0 0 1
0 0 5
IV. A dry, old field, seven miles south of Denbigh, Ontario. Elevation i,ooo
feet. Trapped in only in 194Q.
A dry, grassy, old field with many rocks and piles of old fence-rails
surrounding a field of un-mown rye. A stone fence topped by a broken-
down rail fence separates the two fields. Polytrichum, dry grasses {Dan-
thonia), sweet fern {Myrica asplenifolia) , and bracken {Pteridium latiuscu-
lum) were the dominant plants. An extensive, pure stand of white spruce
surrounds the area. A pair of ravens were once seen in this area, as well as
purple finches, and song and vesper sparrows.
Mammals: Hares {Lepus americanus) were frequently seen along the
highway in this area, and deer (Odocoileus) were seen just south of the
area. The rye field probably was attractive to many mammals.
Only six traps were placed in the area, for three nights. These were
placed along the old stone and rail fence. Three specimens of Peromyscus
leucopus noveboracensis, one Microtus pennsylvanicus, and one Tamias
striatus lysteri were caught. One of the Peromyscus was captured, by the
front foot only, and it remains alive and well at the present writing. This
was the most productive area trapped relative to trap-nights per catch,
with 1 catch per 3.6 trap-nights.
Trapping Summary—
Date: July 5
-1949
6
7
Totals
Peromyscus 1. noveboracensis
3
0
0
3
Microtus p. pennsylvanicus
0
1
0
1
Tamias s. lysteri
0
1
0
1
Totals
3
2
0
5
352 Annals of the Carnegie Museum , vol. 31
\ . An open, sphagnum-V accinium hog, jive miles south of Cloyne.
Elevation 8^0 feet. Trapped in only in 1949.
This area was once a black spruce-tamarack bog but, having been
cleared for a high-power line, it is now primarily an open bog with a
dense mat of sphagnum supporting a heavy growth of Vaccinium and
Kalmia. There are a few black spruce and tamarack seedlings in the
cleared area. On either side of the clearing are stands of mature black
spruces and tamaracks, while on the drier surrounding land are white
spruces and firs. In 1949 this was thoroughly dried out and few spots of
standing water remained ; one in particular was formed by a stone and log
cradle for a power-line pole. Ordinarily the entire area would be quite wet.
The principal vegetation follows:
**Sphagnum
*Pale laurel {Kalmia polifolia)
*Sheep laurel {Kalmia augustifolia)
*Bog bilberry {Vaccinium uliginosum)
*Small cranberry {Vaccinium oxycoccos)
*Canada blueberry ( Vaccinium canadense)
Cattails {Typha sp.)
Tamarack {Larix laricina)
Black spruce {Picea mariana)
Pitcher plant {Sarracenia purpurea)
The birds noted, principally in the black spruce-tamarack, mature
stand, follow:
Kingbird {Tyranus tyr annus)
Wood pewee {Myiochanes vireus)
Barn swallow {Hirundo erythrogaster)
Bluebird {Sialia sialis)
Bronzed grackles {Quiscalus quiscula)
Myrtle warbler {Dendroica coronata)
Vesper sparrow {Pooecetes gramineus)
This type of habitat is common in low areas south of Cloyne and north
of the lake-plain region.
Mammals: In the dried-out portion of the sphagnum bog, there were
abundant old runs containing old, dried (winter or early spring) Synap-
tomys droppings and cuttungs. The spruce edge sphagnum floor had
similar runs with black droppings. Lepus runs (and forms) were abundant,
criss-crossing the area, and one hare was jumped from its form in the
vicinity of the cradle every time the area was visited. The wet areas con-
1951
Christian: Mammals of Mazinaw Region, Ontario
353
tained numerous runs in the sphagnum containing bright green, vole
feces and fresh cuttings of grasses and mosses {Poly trichum). Apparently
these voles had migrated to the wetter spots as the bog dried.
Fifty-three traps were set across the area so that the spruce edge, a
few of the dry spots, and a large portion of the wetter areas with fresh
signs, were trapped. The traps averaged about ten feet distant from each
other. SynaptomySy and possibly Zapus and Sorex, were considered likely
catches. The traps remained set out for two nights.
Two specimens of Synaptomys cooperi coo peri and one Peromyscus
leucopus novehoracensis were caught the first night, comprising the entire
catch, or 1 catch per 35 trap-nights.
Trapping Summary — 1949
Date: July 6
Peromyscus 1. novehoracensis 1
Synaptomys c. cooperi 2
Totals 3
7 Totals
0 1
0 2
0 3
VI. A small creek and a high, alder bog in the dry hills, seven miles north
of Cloyne, Ontario. Elevation 1,050 feet. Trapped in part in ig4Q, further
in 1950.
This area includes more than the specific area trapped, and consists
really of more than one type of habitat. The hill on either side of the
area trapped is included in the following discussion. The hillside itself is
a dry, deciduous, second growth woods with a few remaining white and
red pines and some white spruces. White pines, however, comprise the
great majority of the seedlings in the area, almost to the exclusion of all
other types. Large-toothed and trembling aspens, white birch, and sugar
maple are the dominant mature trees. The middle of the hill is traversed
by a small rock-lined stream with a black mud bottom. This stream suc-
cessively drains an alder marsh, a cattail-sedge marsh, another alder bog,
and alternating bogs of these two types above this. These extensive bog
areas lie in a flat, shallow valley between the hills. An abandoned road
crosses the stream about half-way up the hill, and below the road the
stream becomes much rockier and flows more rapidly. Sugar maples and
herbaceous plants are much heavier along the stream than on the rest of
the hill.
The plants in the general area include:
White pine {Pinus strobus)
Red pine {Pinus resinosa)
354
Annals of the Carnegie Museum
VOL. 31
White spruce {Picea glauca)
Black spruce {Picea mariana)
Balsam fir {Ahies halsamea)
Hemlock {Tsuga canadensis)
*Trembling aspen {Populus tremuloides)
*Large-toothed aspen {Populus grandidentata)
*Speckled alder {Alnus incana)
Black willow {Salix nigra)
Basswood {Tilia americana)
Red osier dogwood {Cornus stolonifera)
Beaked hazelnut {Corlylus rostrata)
Long-beaked willow {Salix rostrata)
*Sugar maple {Acer saccharum)
Red maple {Acer ruhrum)
Moosewood {Acer pennsylvanicum)
Red oak {Quercus borealis)
Sycamore {Plantanus occidentalis)
Black cherry {Prunus serotina)
Smooth-leaved shadbush {Amelanchier laevis)
Bush honeysuckle {Diervilla lonicera)
White ash {Fraxinus americana)
Toothed woodfern {Dryopteris spinulosa)
Cinnamon fern {Osmunda cinnamomea)
Interrupted fern {Osmunda claytonia)
Sensitive fern {Onoclea sensibilis)
*Sphagnum — in the bogs.
Yellow bead lily {Clintonia borealis)
*Wool grass {Scirpus cyperinus) — bogs.
The birds in this area were observed rather closely, and those on the
following list are all breeding birds, noted in both 1949 and 1950. At the
time of the observations of 1950, fledglings and young of all varieties were
abundant. The second growth, the slashings, the bare grassy areas, the
low brush, and the variety of habitats apparently provided ideal breeding
locations for most varieties, especially the brush-loving warblers. This
area had a higher bird population than any other habitat discussed in
regard to both species and individuals.
Ruffed grouse {Bonasa umbellus)¥emQ\Q and covey of young.
Woodcock {Philohela minor)
Whip-poor-will {Antrostomus vociferus)
1951 Christian: Mammals of Mazinaw Region, Ontario 355
Hummingbird {Archilochus colubris) — feeding on sap of white-birch
from sapsucker holes.
Northern flicker {Colaptes auratus)
Yellow-bellied sapsucker {Sphyrapicus varius)
Least flycatcher {Empidonax minimus)
Wood pewee {Myiochanes virens)
Bluejays {Cyanocitta cristata)
Black-capped chickadee {Penthestes atricapillus)
Winter wren {Nannus hiemalis)
Robins {Turdus migratorius) — abundant 1949, scarcer 1950.
Hermit thrush {Hylocichla guttata)
Veery {Hylocichla fuscescens)
Cedar waxwing {Bomhycilla cedrorum)
Red-eyed vireo {Vireo olivaceous)
Black-throated blue warbler {Dendroica coerulescens)
Black-throated green warbler {Dendroica virens)
Chestnut-sided warbler {Dendroica pensylvanica)
Ovenbird {Seiurus aurocapillus)
Redstart {Setophaga ruticilla)
Magnolia warbler {Dendroica magnolia)
Nashville warbler {Vermivora ruficapilla)
Rose-breasted grosbeak {Hedymeles ludovicianus)
Indigo buntings {Passerina cyanea)
Purple finch {Carpodacus purpureus)
Goldfinch {Spinus tristis)
Chipping sparrow {Spizella passerina)
Mammals: Bears {Euarctos americanus) have been shot frequently in
this area, and a barren field on the hill crest contained numerous over-
turned stones where a bear had been looking for ants. Tracks of minks,
racoons, deer, and foxes were seen in a muddy rut between two parts of
an alder bog. The abandoned road in this area apparently provided a
well-travelled animal highway. A skunk was seen walking up this road in
mid-afternoon.
In 1949, fifty-five traps were set; sixteen along the creek and through
an alder bog, eight through a sedge meadow in underground muddy
runs, twenty-two in a dense, alder and deciduous thicket, and nine in a
cattail-sedge marsh. These traps were set for two nights and yielded one
Sorex fumeus, one Blarina b. talpoides, one Peromyscus maniculatus
gracilis, and six examples of Microtus pennsylvanicus. The latter were
356
Annals of the Carnegie Museum
VOL. 31
caught in both the sedge meadow and the dense, alder bog. The average
catch was 1 per 12.2 trap-nights.
Date: July
Trapping Summary — 1949
9
10
Totals
Sorex f. fumeus
0
1
1
Blarina b. talpoides
1
0
1
Peromyscus m. gracilis
1
0
1
Microtus p. pennsylvanicus
5
1
6
Totals
7
2
9
In 1950 twenty-nine traps were set along the same stream starting just
across the abandoned road from the beginning of the line of 1949 and
working downstream for about two-hundred feet. This portion of the
stream had steep banks and was rockier than above. There were numerous,
large boulders with many runs, crevices, and other likely places for small
mammals. These traps were set for three nights and produced one Sorex
fumeus, one Condylura cristata, and two of Blarina brevicauda talpoides,
for an average of 1 catch per 22 trap-nights.
Date: July
Condylura c. cristata
Sorex f. fumeus
Blarina b. talpoides
Totals
Trapping Summary —
8
1
0
1
2
1950
9
0
0
0
0
10 Totals
0 1
1 1
1 2
2 4
VII. .4 small, sphagnum hog in a patch of spruces, 0.6 miles west of route
41 and 1.2^ miles southeast of Massanoga, Ontario. Elevation g^o feet.
Trapped in in ig4g only.
This is a small sphagnum bog in a patch of black spruce and firs in the
center of a larger and more extensive forest of second-growth hardwoods.
In many respects it is much like trapline III except that it is not as ex-
tensive or dense, and in general is an area not as cool. It was at one time
burned over, as many charred stumps and logs cover the ground. The
bog is divided into boglets by fallen logs, and these are covered with
mosses, primarily sphagnum. Standing water persisted in some of the
boglets in 1949. The outstanding feature of this habitat is its present dis-
continuity with the surrounding drier forest, and it is consequently an
island situation.
The principal vegetation is as follows:
*Black spruce {Picea mariana)
Balsam fir {Abies balsamea)
1951
Christian: Mammals of Mazinaw Region, Ontario
357
*Black ash {Fraxinus nigra)
Red baneberry {Actaea rubra)
*Sphagnum, and other mosses and liverworts.
* Yellow bead lily {Clintonia borealis)
Star violet {Dalibarda repens)
Mammals: Twenty-one traps were set in this area in locations very
similar to those in trapline III: in suitable runs, along fallen logs, around
stump bases, and other such spots. One Condylura cristata and one Micro-
tus pennsylvanicus were taken in two nights of trapping. This seemed like
an ideal spot for Clethrionomys and for Sorex, but since none were caught,
the conclusion was reached that the area was too small to support a
typical bog or bog-edge fauna in an extensive dry habitat. There was one
catch for each 21 trap-nights.
Trapping Summary — 1949
Date: July 10
Condylura c. cristata 0
Microtus p. pennsylvanicus 0
Totals 0
11 Totals
1 1
1 1
2 2
VIII. A sedge meadow and a marsh area in an old lake fill, 1.2 miles
southeast of Massanoga and 1.2 miles west of route 41, on the old road to
Mica. Elevation feet. Trapped in only in iQ4g.
This is a completely open ‘^beaver meadow” of grasses and sedges (wool
grass, locally called “beaver hay”), about three-quarters of a mile long
and varying from one-quarter to one-half mile wide. A small lake still
remains unfilled at the western end of the area. A small, muddy, slow-
flowing creek traverses the center of the area after leaving the lake. The
stream is dark and very acid, as are all of the streams in this area. The
wool grass is about three feet tall and forms a dense cover under which
is a thick blanket of sphagnum, broken dead sedges, and other litter.
Surrounding the meadow is a dense zone of speckled alders which, along
with small willows, are now reaching out into the meadow area, and are
thinly scattered throughout, as well as lining the central creek. These
alder thickets seem to be a favorite breeding place for chestnut-sided
warblers. Surrounding the alders, and on slightly higher ground, are
tamaracks, white spruces, black spruces, white pines, and beyond these
are mixed hardwoods in which the white birch is dominant. The vegeta-
tion of the area is listed below:
358
Annals of the Carnegie Museum
VOL. 31
White spruce {Picea glauca) — edges.
Black spruce {Picea mariana) — edges.
Tamaracks (Larix laricina) — edges.
White pine {Pinus strohus) — edges.
*Speckled alder {Alnus incana)
*Black willow {Salix nigra)
White birch {Betula papyrifera) — edges.
**Sphagnum.
**Wool grass {Scirpus cyperinus) — “Beaver hay.”
Cattails {Typha sp.)
Chestnut-sided warblers {Dendroica pennsylvanica) were breeding in
large numbers in the low willow and alder bushes in the area, especially
along the northern margin, where there was an extensive area of second-
growth white birch and large alders and willows.
Mammals: Beavers were at one time plentiful in the lake at the west
end of the meadow, but are now reduced by trapping to just a few. Ac-
cording to the report of the trapper for this area (Albert Spencer, Cloyne),
they are building up again (1950) as a result of the present Ontario trap-
ping laws. Tamiasciurus was plentiful in 1949, see the account under the
species discussion.
Twenty-eight traps were put out at twenty-one stations about twelve
feet from each other, extending in a straight line across the meadow with
a few scattered on the opposite side in a patch of spruces and alders. Two
of Blarina h. talpoides, one Peromyscus maniculatus gracilis, one Pero-
myscus 1. novehoracensis, and one Microtus pennsylvanicus were caught in
two nights of trapping, or 1 catch per 11.2 trap-nights.
Trapping Summary — 1949
Date: July 10
Blarina b. talpoides 1
Peromyscus m. gracilis 0
Peromyscus 1. novehoracensis 1
Microtus p. pennsylvanicus 1
Totals 3
11
1
1
0
0
2
Totals
2
1
1
1
5
IX. A high, dry, barren rocky hill, 8.5 miles south of Denbigh, Ontario.
Elevation 1,000 to 1,1^0 feet. Trapped in only in iQ^o.
This area represents one of the habitat types missed in 1949. The hill
is a solid eminence of quartzite, and is one of the typical glaciated grani-
toid hills of the region. It is very steep-sided to the east, north, and
south, but slopes more gradually to the west. At one time this now barren
1951 Christian: Mammals of Mazinaw Region, Ontario 359
hill may have been forested, as a few burned logs bear mute evidence
to a previous fire. At present, however, large areas of bare rock lie ex-
posed, especially where the slope is steep. A few aspens have gained a
foothold along with patches of poverty grass, blueberries, brackens, and
a sweet fern. A few other plants are found in the area, as the following
list shows. The dominant form, however, is the blueberry, and this site
is more or less typical of the many so-called blueberry balds or barrens in
this region. Many of the hilltops and higher elevations are of this type of
habitat. The soil, where it has been gathered in the more level spots, is a
breakdown of the altered sandstones which form the hill itself. It is in
these relatively level areas, in cracks, and in such soil-gathering spots that
the plant life is found. The area is very dry and exposed. Blueberries were
ripe at the time of trapping (July 4, 5, and 6), in contrast to the other
areas in this region.
The following plants were noted.
*Lichens.
*Bracken {Pteridium latiusculum)
*Hair moss {Polytrichum sp.)
White pine {Pinus strobus) — very few.
White spruce {Picea glauca) — very few.
Trembling aspen {Populus tremuloides)
Large-toothed aspen {Populus grandidentata)
White birch {Betula papyrifera)
*Sweet fern {Myrica asplenifolia)
* Poverty grass {Danthonia sp.)
Smooth sumac {Rhus glabra)
**Low sweet blueberry {Vaccinium pennsylvanicum v. augustifolium)
Ground bean {Strophostyles helvola)
Pink fume root {Corydalis sempervirens)
Hairy rock cress {Arabis hirsuta)
Whorled loosestrife {Lysimachia quadrifolia)
Barren strawberry {Waldsteinia fragaroides)
Not many birds were noted in the area. Those seen follow:
Bluejays {Cyanocitta cristata)
Black-throated green warbler {Dendroica virens)
Ovenbird {Seiurus aurocapillus)
White-throated sparrow {Zonotricha albicollis)
Mammals: This area was picked as typical of the high blueberry bar-
rens of the region, in the belief that the faunal inhabitants would be
360
Annals of the Carnegie Museum
VOL. 31
representative of this type of habitat. Some runways, old signs, and grass
cuttings were found around the grasses and blueberries. The cuttings and
runs were old. One Tamias was seen feeding on blueberries, otherwise no
mammals were seen.
Forty-seven traps were put out, starting in a blueberry patch at the
base of a steep, rock face about 150 feet below the summit, and extending
on up to the crest with traps placed in all suitable patches of blueberries,
grasses, or brackens, and along the bases of cliffs, and around the few
logs and stumps. A few were put in relatively barren spots. The average
trap interval was ten feet. The traps were baited with a mixture of peanut
butter, ham-fat, and almond extract, and oatmeal was scattered over
each trap. No small mammals were trapped in the three nights the traps
were left set, although a few were sprung by a heavy rain during the
first night.
It seems probable that this type of habitat is marginal in this region and
that during a year of excessively low population, as in 1950, it remains
practically uninhabited. The evidence of runs, signs, and cuttings shows
the area to have been occupied by small mammals in the past, probably
the preceding fall and early winter.
X. A dense, black spruce hog, 4.5 miles sotUh of Denbigh, Ontario. Ele-
vation approximately 1,000 feet. Trapped in in ig^o only.
In 1949, no traps were placed in the climax black spruce bog, so this
area was trapped in during 1950 to fill this gap. This is a typical, boreal,
black spruce bog. The upper story was almost a pure stand of mature
black spruce with a few tamaracks. The understory was a solid carpet of
sphagnum with Labrador tea growing from it. The spruce habitat was
extensive, and has apparently been totally by-passed by fire and timbering
except for a small area at the south end which now consists of an overstory
of speckled alder. Water was standing beneath the sphagnum. The soil
is acid, black muck. Small cranberries are common. The alder portion had
less sphagnum with more of the black muck being exposed. Also, there
was little standing water in this portion of the bog.
The higher ground surrounding the bog was covered almost entirely
with white spruce and fir. Cinnamon ferns and yellow bead-lillies formed a
dense cover around the periphery of the bog.
Floral list of the area:
**Black spruce {Picea mariana)
*Tamarack {Larix laricina)
1951 Christian: Mammals of Mazinaw Region, Ontario 361
*Speckled alder {Alnus incana)
**Sphagnum.
**Labrador tea {Ledum groenlandicum)
*Small cranberry {Vaccinium oxycoccos)
Heart-leaf lily {Maianthemum canadense)
Bunchberry {Cornus canadensis)
*Deer vine {Linnaea americana) — in bloom.
Barren strawberry {Waldsteinia fragar aides)
Arctic bramble {Rubus arcticus) — in berry.
The following birds were noted in the area, although no concentrated
effort was made to obtain a complete list due to the density of the growth
with its attendant poor visibility. All are probably nesters in or near the
area.
Pair of Broad-winged hawks {Buteo platypterus)
Ruffed grouse {Bonasa umhellus)
Red-breasted nuthatch {Sitta canadensis)
Nashville warbler {Vermivora ruficapilla)
Redstart {Setophaga ruticilla)
Purple finch {Carpodacus purpureus)
Mammals: The area was laced with runs of the snowshoe hare and fresh
scats were abundant, although no hares were seen.
Forty-seven traps were set in the area at intervals of approximately
twenty feet. The first fifteen traps were in the alder portion of the bog,
and the remaining thirty-two were in the stand of black spruce. All traps
were set in likely-looking spots with the spacing a secondary consideration ;
so that some were grouped more closely than others. These traps re-
mained set and unmoved for four nights for a total of 188 trap-nights.
One Sorex cinereus was caught on the second night in the alder portion
of the bog. One Sorex cinereus and one juvenile Peromyscus maniculatus
gracilis were caught in the spruce portion of the bog on the fourth night.
Two very hot, dry days preceded the final night, making one wonder if
the two catches on that night hadn’t wandered into the cool, moist, bog
area from surrounding drier territory. The lack of catches in this portion
of the bog for the three preceding nights tends to support the idea that
these animals were transients from outside of the area. Three catches in
188 trap-nights equals 1 catch per 63 trap-nights.
Trapping Summary — 1950
Date: July 7 8
9
10
Totals
Sorex c. cinereus
0 1
0
1
2
Peromyscus m. gracilis
0 0
0
1
1
Totals
0 1
0
2
3
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In addition to the specific trapping areas described above, highway 41
was a collecting locality. In 1949, three Tamias, one Tamiasciurus, and
one Lepus were picked up and saved as specimens. In 1950 two Tamias
were found in a condition good enough for specimens. Others were picked
up which could not be saved. Among the latter were a weasel {M.frenata),
several porcupines, several red squirrels in 1949 and another red squirrel
in 1950. Many sight records were obtained along the highway for Mar-
motaj Mephitis, Mustela vison, Erethizon, Lepus, Odocoileus, Tamias,
and others, as well as for many of the birds. Most of the animals seen or
picked up from the highway came from the hardwoods and brush along
the roadside. Lepus and deer were seen especially about six miles south of
Denbigh in a small grove of spruces below the farm of trap-site IV. In
1950 minks were twice seen crossing the highway in daytime. Road con-
struction on portions of the highway reduced the number of animals seen
in 1950 from that which otherwise might have been present. Hares were
conspicuous by their absence from the road in 1950, only one record hav-
ing been obtained, but that this was a result of the road-work was shown
by their abundance along other roads. The discontinuance of road treat-
ment with calcium chloride further reduced the numbers seen in 1950,
since this salt attracted many mammals, especially hares, even though it
resulted in their death from calcium poisoning.
III. ACCOUNTS BY SPECIES
Condylura cristata cristata. This was the only mole captured in the
area. Three specimens were captured, two in 1949 and one in 1950, in
wet, muddy habitats. One was taken in the tunnel-run at the base of the
scarp in area III, one along a fallen log in the area VII bog, and one at a
run opening at the base of a rock wall in area VI (1950). It may be noted
that all three of these moles were taken in runs at the bases of natural
barriers.
A female captured on July 5, 1949 contained one extremely small
embryo in each horn of the uterus, corresponding to a six day pregnancy
(early implantation) in the rat. The female trapped on July 8, 1950 was
not lactating, was not pregnant, and no implantation scars were found.
A male trapped July 11, 1949 was immature with small abdominal testes.
No smear of the testis was made. These data suggest that these moles
may have two litters a year in this region.
Sorex cinereus cinereus. Six of these shrews were captured, two in
1949 and four in 1950. The two specimens taken in 1949 and the two in
1951 Christian: Mammals of Mazinaw Region, Ontario 363
1950 were trapped in the black ash-white cedar bog (III). The bog had
no standing water in 1949, and the shrews were caught in boglets in the
center of the bog — one at a hole emerging from the black muck bottom,
and the other at the base of an overhanging hummock. In 1950, when the
boglets were under water, these shrews were caught in a tunnel-run at the
base of a scarp at the bog edge, while none were caught in the central
area. The remaining two specimens of Sorex cinereus, taken in 1950, were
trapped in the black spruce-sphagnum bog (X) : one in the black spruce
portion, and the other in the alder portion of the bog. Both were in clumps
of sphagnum.
Sorex cinereus in this region has the tricolor pattern with a distinct
lateral band, described by Jackson (1928) as occasionally occurring. This
pattern was much more conspicuous in dead specimens than in prepared
skins. Another feature quite striking in specimens “in the flesh” is the
much denser, softer, and silkier pelage found on cinereus than on fumeus
when caught in the same place at the same time. The coarser pelage of
fumeus is less pronounced in the dried skins, but it is nevertheless evident.
This suggests that cinereus may have more fossorial habits than fumeus.
All of the cinereus caught in both years had long tail pencils regardless of
obvious age differences.
Both of the cinereus trapped in 1949 were pregnant females, one with
six 9 mm. embryos caught on July 11, and the other with four minute
embryos captured July 7. The distribution of the embryos in the uterine
horns was equal in both shrews. One immature, non-pregnant female was
caught in 1950. No implantation scars were found, the uterus was very
small and juvenile, and the mammae were obviously small and undeveloped.
The three males of 1950 were all adult with scrotal testes and abundant
sperm in the epididymes. The testicular smears, however, showed ad-
vanced degenerative changes, with few or no spermatozoa, chromolytic
changes in the nuclei of the spermatids, and numerous giant cells con-
taining up to twenty spermatid or secondary spermatocyte nuclei.
Occasionally primary spermatocytes were found involved. The fact that
all three males showed these changes is strong evidence that the adult
males cease their reproductive activity at this time of the year.
If combining two years’ findings is a valid procedure, the capture of a
sexually immature female and two pregnant females at the same time
of the year indicates that two litters may be raised. The fact that pregnant
females were caught so late in the season is suggestive of this without any
further evidence, as the young of the year probably have not reached
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maturity by this time. The possibility still remains until disproven. The
degenerative condition of the testes of the male shrews taken in 1950
indicates that the end of the spring breeding season occurs about the first
of July.
With the adult males undergoing testicular degeneration in early July,
the problem arises of determining the ages of shrews caught later in the
summer. It has been noted in Blarina that the degenerate, late fall testes
of adults have a flaccidity which is quite distinct from the small, firm
testes of juvenile animals, and this may be a useful criterion for Sorex
cinereus. Tooth-wear should also be a distinct aid at this point.
The evidence at hand, then, suggests that Sorex cinereus has a limited
breeding season in the spring and early summer and that either (1), two or
more litters may be raised per adult female, or (2), the young of the year
reach maturity soon enough to become pregnant before the adult males
undergo testicular regression early in J uly.
Sorex fumeus fumeus. Four smoky shrews were captured: two in
1949, and two in 1950. One was caught each year in habitats III and VI.
In 1949 one was caught beneath some alder roots along the streams of
VI, while in 1950 another was caught farther down along the rocky bank
of the same stream. The specimen taken in 1949 in the black ash-white
cedar bog was caught at the base of a boglet hummock in the center of the
area, while in 1950 this shrew was caught in the tunnel-run at the fault
base. None was caught in the boreal, black spruce bog. The implication is
that Sorex fumeus is more closely associated with deciduous growth than
cinereus is in this region.
Of the four specimens of Sorex fumeus^ three were females, one with an
undetermined reproductive status, but not pregnant, and the other two
obviously immature shrews with small, undeveloped uteri with no im-
plantation scars. The one male, captured in 1950, was also immature with
abdominal testes containing mainly spermatogonia with a very few pri-
mary spermatocytes. The interstitial cells were small. In contrast to Sorex
cinereus^ it appears from the collected evidence that Sorex fumeus in this
region may have an early spring breeding season with probably only one
litter raised. This is admittedly limited evidence, and Hamilton (1943)
states that these shrews have two spring litters, and less often a third
litter in the fall. The possibility of the young of the year being responsible
for these later litters, however, is not precluded. It does seem, if there were
adult, reproductively active individuals of fumeus about during the 1949
1951 Christian: Mammals of Mazinaw Region, Ontario 365
and 1950 trapping periods, that at least one would have been captured.
Also, Sorex fumeus may have but one litter in the spring in this region,
whereas it has two farther south and at a lower altitude. That the rela-
tive population levels of these animals may reflect changes in reproduc-
tive potentials must also be taken into consideration.
Sorex palustris albibarbis. One male water shrew was captured in
1949 in the black ash-white cedar bog (III). Judging from its cross-wise
and attenuated position in the trap, the animal apparently had been
running along the mud at the base of an overhanging sphagnum hum-
mock in the center of the bog. The habitats and trap locations were al-
most identical with those in which one of these shrews was caught in
Monroe Co., Pa., in May 1949.
This animal was an immature male with abdominal testes measuring
1.0 X 1.5 mm. The testicular smear showed abundant spermatogonia, very
few primary spermatocytes, and no spermatogenesis beyond this stage.
There were no sperm in the epididymis.
None of these shrews was caught in 1950, when most of the bog was
covered with standing water, even though the area was trapped much
more heavily and for a longer period.
Microsorex hoyi intervectus. One pregnant female was caught in
the run at the base of the fault-face of the black ash-white cedar bog
(III) on July 3, 1950. She had three very small embryos, two in the right
and one in the left uterine horns. The mammae were enlarged.
This shrew was captured in the same tunnel-run in which Sorex cinereus
and Sorex fumeus were taken in 1950, and Condylura and Synaptomys in
1949, as well as the same bog in which these two shrews and Sorex palus-
tris were taken in 1949. Thus four species of long-tailed shrews and
Blarina were inhabiting the same bog. The obvious aquatic adaptations
of Sorex palustris probably serve equally as well on the wet, black muck
of the “1949 bog floor.” That Sorex cinereus may lead a more fossorial
existance than Sorex fumeus with slightly different habitat preferences
has already been suggested, but where Microsorex fits in the ecological
picture with these other shrews is difficult to say. That Sorex palustris
albiborbis and Microsorex hoyi intervectus are much less common than the
other two shrews is evident.
The thymus in all of these shrews was found to be extremely large,
compared with most other mammals. It is a flat, leaf-like organ spreading *
over the entire ventral surface of the thorax, from its apex in the superior
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mediastinum. The interscapular, hibernating gland mass, is likewise large
and extends as a continuous mass over the back in the scapular region,
passing posteriorly around the shoulder and into the axilla. In other
mammals the hibernating gland exists as more or less discreet masses in
these various regions.
Blarina brevicauda talpoides. Twelve short-tailed shrews were
captured in 1949, and five in 1950. As elsewhere in their range, these
shrews are probably the commonest mammals in the region. In 1949 they
were caught in all habitats except the dry, old field (IV), the vaccinium-
sphagnum bog (V), and the small, relatively warm, spruce bog, while in
1950 they were not caught on the high, dry bald (IX), or in the black
spruce bog (X). The inference is that they do not occur in extensive, wet
sphagnum bogs or excessively dry spots, but are otherwise ubiquitous.
However, the low population of 1950 must be kept in mind. In years of
peak populations they may possibly occur in these places.
Testicular smears were made of five males in 1949. Three of these
showed all stages of spermatogenesis in the testes, and spermatozoa in the
epididymes. All three had early degenerative changes with numerous,
large giant cells of spermatids and secondary spermatocytes, chromolytic
changes of the spermatid nuclei, and vacuolar degenerative changes of
the spermatogonial series. The other two were immature with no sperma-
togenesis beyond a few primary spermatocytes and no sperm in the
epididymes. Of the four males in 1950, two were immature with smear
pictures similar to the above juvenile shrews. Two were adult with scrotal
testes, sperm in the epididymes, and active spermatogenesis. One of these,
however, showed marked degenerative changes of the testes, involving
particularly the secondary spermatocytes and spermatids. Giant cells,
composed primarily of spermatids, were abundant with up to ten nuclei.
Most mature Blarina, then, probably undergo testicular regression at the
end of the spring breeding season in this region, while a few may remain
in a functional reproductive state, although they, too, may undergo
degeneration at a later date than is covered in this account.
Four females were examined in 1949. None was pregnant; one had
visible placental scars and was lactating, two others were lactating, and
the fourth was obviously immature. The one ^H950 female” was a nulli-
parous juvenile with remarkably prominent lateral scent glands. Since
no pregnant females were trapped, it is probable that the peak of the
breeding season had been passed by the first part of July.
1951 Christian; Mammals of Mazinaw Region, Ontario 367
Chiroptera. No bats were seen in the 1949 period, and sight records
reported by various people all turned out to be chimney swifts. Definite
knowledge of a few was obtained in the report of some which had been
hanging behind the shutters of a lakeside cottage earlier in the summer.
In 1950, one was seen flying with a cicada in its mouth. It was flying
across a clearing above Marble lake at dusk and the species could not be
determined, but it appeared small — not much larger than the cicada.
No others were seen.
Euarctos americanus. The black bear is common in the region. At
this time of the year they are reported to be back in the bush feeding on
blueberries. In 1950, many large stones turned over by a bear searching
for ants were found in a barren field above area VI. The bear had ap-
parently left shortly before my arrival, as the ants were still running
around carrying their eggs to safety. There were others on the east shore
of Mazinaw Lake during the 1949 period. In the early fall of 1947 three
were trapped above the west shore of Mazinaw in area VI. The unanimous
local opinion is that they are a nuisance.
Procyon lotor. Local trappers and residents report that raccoons
are very abundant, but the present low value of their fur keeps them from
being trapped. One dead juvenile, apparently a highway casualty, was
picked up in area I on the west shore of Mazinaw Lake.
Martes americana. Two trappers (A1 Spencer and Irving Brown)
and the Lands and Forests District warden, all report that martens have
been unknown in the region during their lives — about forty years. Mar-
tens were undoubtedly present at one time, but their trap stupidity has
probably long since exterminated them throughout the district.
Martes pennanti. Reports on the fisher are conflicting, but it seems
likely that some still exist about seven miles northwest of Mazinaw Lake
in the enormous, virgin white pine forest surrounding Weslemkoon Lake.
A few are known to be in Algonquin Park, and some could have wandered
down from there through the continuous intervening forest. However,
the area west of Mazinaw is wild and inaccessible enough to have main-
tained a small permanent population, especially in the white pine forest.
Albert Spencer, who traps an area of about seventy square miles between
Mazinaw and the pines of Weslemkoon, has never taken or seen a fisher
in his area, although he thinks they may be present in the pines. Irving
Brown states definitely that fishers are still present “about seven to eight
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miles west of here in the virgin white pine wilderness around Weslemkoon
Lake.” The forest warden said that Brown was the best trapper here-
abouts and knew more than anyone else about the mammals. The warden
himself recalls having seen the last fisher taken in the Plevna region about
forty years ago.
It is unfortunately true that the edges of the virgin pine forest are
already being timbered during the winter, and it is only a matter of time
before the forest will no longer exist. With its passing, the fisher will un-
doubtedly disappear also.
Mustela erminea cicognanii \
Mustela frenata noveboracensis J Both of these weasels are present,
and are distinguished from each other on the basis of size by all trappers
with whom I talked. A large white frenala from this region commands a
good price on the fur market ($3.75 top in 1949; $2-$3, depending on size,
in 1950), hence they are conscientiously trapped. One example oi frenata,
too badly damaged to keep, was killed on the highway in 1949.
Mustela rixosa. According to Anderson (1946), the least weasel
has not been taken in this part of Ontario. Evidence from local trappers
is contradictory. Brown stated that weasels of all sizes had been acci-
dentally trapped, but brought a good price “except for the very small
one with a tail about an inch long, which was worthless.” The forest
warden stated that weasels were very abundant last year (1949-50) in the
Plevna district with “very large, small, and little short-tailed ones.”
Albert Spencer, however, after hearing a careful description of the least
weasel, said that he was totally unfamiliar with the animal, and that per-
haps it did not occur around Mazinaw. The comments of these three re-
liable people are difficult to reconcile. Although in the face of the above
statements it seems very unlikely, it is possible that Brown and the war-
den were referring to small females of Mustela erminea cicognanii.
Mustela vision vision. Minks, along with beavers, are the staples of
the fur business in this region. They are quite common, and I noted mink
tracks along most of the muddy streams or bogs investigated in 1949 or
1950, especially along the creek and alder bogs of area VI. One mink was
seen in mid-afternoon crossing the highway towards the shore of Mazinaw
Lake in 1950. With both mice and muskrats at very low levels, the mink
must have to work hard and travel far for sufficient sustinence. Irving
Brown (1950) says that the mink is getting scarce in the trapping area to
the east and north of Mazinaw Lake. The forest warden reported them
1951 Christian: Mammals of Mazinaw Region, Ontario 369
plentiful in the Plevna district, and trapped a few in 1949 and 1950. The
fur value of a large mink is now from eighteen to twenty-five dollars.
One mink is known to have been living on the west shore of Mazinaw
in Area I several years ago when it attempted to carry off a kitten from
an old trailer. It was driven off by the mother cat after a vicious fight.
Lutra canadensis canadensis. Otters reportedly have been un-
common in the past, but are now increasing. Albert Spencer trapped two
in this area in the 1949-1950 season, but complained that they brought
only fifteen dollars apiece. Irving Brown reports “loads of otter” and he
recently has been trapping more otters than minks in his area, which
includes the river and marshes north of Mazinaw Lake, as well as the east
side of the lake.
It is evident that otters are quite abundant in the area, as one might
expect from the numerous waterways, lakes, and ponds.
*Mephitis mephitis nigra. Skunks are very common and are a pest
on the highway and around cabins, where they visit every night looking
for tidbits. Skunks were feeding largely on ripe blueberries in 1949.
Several were killed on the highway, and more were seen alongside the
road at night. Two half-grown young ones dwelt beneath our cabin in 1950
(area I), and one of these was trapped. An adult female was collected at
Garbutt’s camp in the same area. This skunk was found alive, but dying,
on the edge of the lake with its head partially in the water. Its left, rear
ankle joint was rigid, forcing it to walk on the tips of its toes. This identi-
fied it as the same animal which had been seen abroad on the hill above
the lake (area VI), at noon two days previously. Autopsy revealed this
animal to be extremely emaciated with atrophic muscles and no fat de-
posits whatsoever. No gross pathological changes were noted in any of
the internal organs. Five porcupine quills, however were removed: three
from beneath the skin, one from in the mandibular joint, and one from
the fibrosed ankle. The ankle fibrosis, however, appeared to be from a
much older lesion than the quill could account for, as old callouses on the
foot testify. The animal had been lactating recently, and had three recent
and three old right, and two recent and three old, left, uterine implanta-
tion scars. External parasites, mostly lice with a few ticks and fleas, were
too numerous to count, and were estimated at at least a hundred for each
square inch of skin over the entire body. The probable history of this
skunk would be that she had raised at least one litter this spring, and some-
*Additional information concerning this species is included in the addendum
after the bibliography, page 385.
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time, probably after the birth of her young, encountered a porcupine.
The quill in the jaw, after it had worked into the muscles and joint,
probably prevented feeding. The hind-foot lesion was an old injury, and
probably did not interfere much with her activities. The parasites most
likely became so abundant as a result of her weakened condition. To the
severe stresses of pregnancy, nursing, and inanition, was added the bur-
den of parasites. Sinus worms were also present. All of these severe stres-
sors resulted in the animal’s death. In spite of all these alarming stimuli,
the adrenal glands were found to have a quite high ascorbic acid content.
This seems most paradoxical and cannot be explained, as exhaustion of
her defense mechanisms, sufficient to cause death, most certainly would
be expected to cause complete depletion of the adrenal ascorbic acid.
Vulpes fulva \
Urocyon cinereoargenteus j Red foxes are present in abundance, and
many sight records are in my possession. I neglected to inquire if gray
foxes were also present while listening to the general complaints about
foxes. The region is outside of the normal range of gray foxes, although
Anderson {loc. cit.) reports that a gray fox had been taken six miles west of
Kaladar, which would be in this general region. The foxes present, how-
ever, are probably red foxes with possibly a few grays.
Canis lupus lycaon. Wolves are relatively common in the Mazinaw
district, although their known presence causes considerable local com-
ment. In the spring of 1949, a pack appeared on the east shore of Mazinaw
Lake, and were heard nightly. The number of animals in the pack is not
known. The forest warden saw a wolf about five and a half miles below
Denbigh on highway 41 during late May, 1950. He stated that these
wolves are much darker than the dog-like wolves around Tweed, and
claims they are ‘^true timber wolves” and distinct from the others. The
so-called wolves of Tweed, that he refers to, may be Canis latrans, or
even wild dogs. This same warden has snared wolves in the Plevna dis-
trict.
Two or three times each winter, wolves put in an appearance along the
shores of Mazinaw Lake. The local opinion, given for what it may be
worth, is that the wolves in the area come down from Algonquin Park.
They are known to be common in the park, but this area probably sup-
ports its own wolves.
Lynx canadensis. The lynx is probably unknown in the territory,
or at least extremely rare, judging from reports.
1951 Christian: Mammals of Mazinaw Region, Ontario 371
Lynx rufus. The bobcat is reportedly common, being frequently
shot or trapped, contrary to Anderson’s (1946) statement that they are
extremely rare. No further information was obtained.
Marmota monax rufescens. Woodchucks are common and were
frequently seen along the west shore of Mazinaw Lake above Cloyne.
None was found dead on the highway in good enough condition for a
specimen, although several had been run over. They all appeared small
and quite red, although one picked up below Kaladar, Ontario, was very
large. The density and length of the guard hairs on this animal were
noticeably greater than on more southern animals.
One was seen along the edge of a pond in the center of a black spruce
bog, and others were noted in the woods, although the roadside areas
seemed to be a choice habitat and supported a large population. This
may be apparent rather than real, since they are so much more easily
seen.
Woodchucks were reportedly very abundant in 1950, more so than in
1949, and created havoc with the few truck gardens in the region.
Tamias striatus lysteri. Chipmunks are abundant throughout the
second-growth, dry, rocky, deciduous woods, and many are killed daily
on the highway. The only mammal recorded on top of the rocky bald
(area IX) was a chipmunk feeding on blueberries. None was seen or cap-
tured in any of the cold habitats visited, but they were abundant around
the shores of the lake. Of the four chipmunks obtained in 1949, two fe-
males and one male were immature. The third female was adult, but,
having been run over, the reproductive status could not be determined
other than that it was lactating.
Two chipmunks were picked up from the highway in good condition
in 1950. Both of these were mature males with scrotal testes. The epi-
didymes of both contained abundant spermatozoa. The testis of one
contained all stages of spermatogenesis and showed no evidence of de-
generative changes. The other had a reduced number of spermatids and
secondary spermatocytes with giant cells composed of four or five secon-
dary spermatocytes. These findings are most likely indicative of very
early degenerative changes, and indicate the beginning of the cessation of
breeding activity in at least the one male.
The population of chipmunks appeared to be about the same in 1950
as in 1949, as they were abundant in both years.
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Tamiasciurus hudsonicus loquax. Anderson (1946) states that
loquax and hudsonicus intergrade in the Algonquin Park region of southern
Ontario, and consequently probably do so in the Mazinaw region also.
The red squirrel is one of the common mammals of the region, and was
abundant in 1949. Not a habitat was visited which did not have red
squirrels chattering from the trees, and many were seen crossing the high-
way or lying dead on it. They were especially abundant in mixed spruce-
deciduous woods, and seemed to stay fairly close to the spruces. They
evidently feed primarily on spruce seeds, as abundant cone cuttings
testify. In one grove of black spruce (VIII) the daytime nest of a red
squirrel was found. It was situated in a crotch, formed by a limb and the
trunk, about twelve feet from the ground. Its outside diameter was about
twelve inches, and in the center was a hollowed out, uncovered cavity
four or five inches in diameter. The whole nest was composed of sticks,
thickly lined with sphagnum and other soft mosses and plant fibers. In
general it appeared like a finely constructed, large, bird nest. A rat trap
was set in this nest after the squirrel was seen in it, but to no avail. A
few feet away, and about twenty feet from the ground in the top of a
small spruce, was its permanent, fully roofed-over nest, composed pri-
marily of deciduous leaves.
In 1949 one lactating female was picked up from the highway.
In 1950, red squirrels in general were conspicuous by their absence. Not
one was seen in any of the habitats visited or trapped in, and none was
heard chattering. Several were seen either crossing the highway or lying
dead on it, but few compared with 1949. Although no actual census was
made, it was very evident that the “1950 population” of red squirrels was
way down from 1949.
Red squirrels are shot on sight by the local residents, since there is no
closed season on squirrels, but their numbers appear to be unaffected by
the shooting, probably due to the enormous reserve range in which they
are completely unmolested.
Sciurus carolinensis leucotis. Both “black” and “gray” squirrels
are found in the district, although I did not see or collect any. Their fur
brings a fair market price, and they are trapped or shot for this reason.
They are reportedly “abundant.”
Glaucomys sabrinus macrotis. Flying squirrels are known to be
abundant even though none was collected. One nest was located in an old
sapsucker hole about eight feet from the ground, in an aspen along the
1951 Christian: Mammals of Mazinaw Region, Ontario 373
shore of Mazinaw Lake. At Northbrook a pair of young ones had been
captured about two weeks prior to our arrival in 1949, and were kept as
pets. Their mother had been shot.
Castor canadensis. Beavers are one of the mainstays of the Mazinaw
fur business, and were rapidly approaching extinction until a few years
ago when the present Ontario trapping laws were put into effect. These
have given the trapper an incentive not to “trap out” a locality by allow-
ing only him to trap in his registered area. Beavers are now increasing
throughout the territory. Mr. I. Brown has been taking many during the
past two years (1949, 1950). Mr. A. Spencer had one beaver house on his
area in 1947, but now (1950) has fifteen, and they are still increasing
rapidly. He limits his catch to one beaver to each house in order to increase
the population. He permits no destruction of dams or houses which bene-
fits both the beavers and the muskrats. One of his colonies is in Essen’s
Lake, slightly west of area VI.
One beaver skull was collected from a trapper’s “carcass dump.”
Peromyscus maniculatus gracilis. Three males and three females
were collected in 1949, and two juvenile females in 1950. One of the males
had completely gray pelage, but was sexually mature with sperm in the
epididymes and all stages of spermatogenesis in the testes. The other
five “ 1949 mice” were adult with respect to both pelage and reproductive
functions. One female had six 23 mm. embryos, and the mammary glands
were post-lactationally enlarged and pigmented. These embryos would
then represent at least her second litter of the year. The same is true for
another which had seven recent and five old implantation scars, and was
lactating. The third had ten uterine scars, and was lactating.
Of the three “1949 males,” the testes of the two with adult pelage
showed marked degenerative changes with very numerous giant cells of
secondary spermatocytes and spermatids. Spermatids with vacuolar and
chromolytic changes were abundant. These two animals were obviously
regressing from full reproductive functioning. The male with juvenile
pelage showed no degenerative changes, but the number of mature sperm
was low.
Peromyscus m. gracilis was captured in such diverse areas as the middle
of a sedge marsh (VHI), an alder bog (VI), a patch of spruces (II), a
black spruce bog (X), a black ash-white cedar bog (HI), and dry mixed
woods (I). The common ecological denominator here is elusive, except
that all habitats are either forested or wet, with a dense cover. The dry
374
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open habitats produced none of these mice, nor did the sphagnum-
Vaccininium bog (V). They were not as abundant as leucopus was in 1949,
but two were caught in 1950 when no leucopus were caught.
Peromyscus leucopus noveboracensis. This is probably the com-
monest mouse in the region. Twelve were taken in 1949, in every type of
habitat trapped, but none was caught in 1950. Of the four females trapped,
only one was adult with a perforate vagina, and this one was not pregnant,
or lactating, or with implantation scars.
Of the eight males, two had abundant sperm in the epididymes. One
of these possibly showed very early degenerative changes of the testes,
while the other had a completely normal smear. One adult had a few sperm
in the epididymes, and the testes showed marked degeneration with chro-
molysis of the spermatids and secondary spermatocytes. Two others, ap-
parently recently matured, had few sperm in the epididymes, but showed
no degenerative changes. The remaining three were juvenile with normal
testicular smears and no spermatozoa in the epididymes. The testis of one
of these contained a few small giant cells.
With only one of the five adult specimens of Peromyscus 1. novenhoracen-
sis showing any marked degenerative changes in the testes, it appears
that these mice either do not follow the same reproductive pattern as
Peromyscus m. gracilis, or that the changes are slower in developing. The
latter seems the likely answer, since at least some of the animals did show
mild degeneration.
One of these mice was captured alive and fed blueberries and other
foods so that it had a choice. At first only blueberries were placed in the
cage and these were eaten. Then blueberries were put in the cage along
with wintergreen berries, and were selected and eaten prior to the winter-
green berries. Berries of Aralia were left untouched, when put in alone or
with other berries. A fly, a beetle, and a bumblebee were quickly seized
and eaten in preference to any kind of berries, or bread, or prepared dog-
food, or a cracker. The berries and an apple were selected in preference to
any kind of prepared food. Black cherries were left untouched! Peanuts
were nibbled, as were various grass seeds. A few stomach examinations
revealed insects primarily, with spruce and pine seeds, and some undeter-
mined vegetable remains. This mouse thrives well on a diet of dried dog-
food and lettuce fourteen months after its capture.
Peromyscus 1. noveboracensis has taken the place of the house mouse,
and invades cottages extensively. Soap seems to be a dietary delicacy.
1951 Christian: Mammals of Mazinaw Region, Ontario 375
Neotoma sp. (?). Although this region is well out of the known
range of the wood rat, it is possible that they are present. Mr. Fred Gar-
butt and his wife described two rats which their cat had brought in several
years ago as being very distinct from Rattus, but about that size. The
exceptional appearance was immediately noticed, and the animal was
described as a rat with large deer-like ears resembling in general a deer-
mouse, only larger. This information was entirely voluntary. Mr. Garbutt
also saw a pair of these rats in a gravel pit about six years ago. Three rat
traps placed in this pit for two nights caught nothing. The greater amount
of fur on the tail of these rats was also mentioned. The fact that this in-
formation was wholly unsolicited lends validity to the description.
Synaptomys cooperi cooperi. Three of these animals were trapped
in 1949; two in a sphagnum-Vaccinium bog (V), and the other in a run at
the base of the rock face in a white cedar-black ash bog (III). They ap-
parently were much more common prior to the extremely dry weather in
the sphagnum-Vaccinium bog, as old droppings were found over the
entire bog, and runs filled with old cuttings literally undermined the
sphagnum of the whole area, most of which was quite dry at the time of
trapping. Fresh feces and cuttings were found only in the remaining moist
areas. One area in particular was littered with fresh green signs and cut-
tings, but no specimens of Synaptomys were captured here. The feces of
the animal in the black ash-white cedar bog were black instead of the
usual bright green, like those found in the open sphagnum bog. Due to
the numerous and extensive spruce and sphagnum bogs in the area,
Synaptomys is probably one of the more abundant mammals in the region.
The period of rainless weather preceding the trapping period in 1949 may
have reduced their numbers to some extent. None were caught in 1950.
The only female trapped had five minute embryos, as well as eight old
implantation scars; so that the young in utero represented at least her
second pregnancy of the year. The mammae were enlarged and post-
lactational.
Both males had abundant epididymal sperm. One of these had a few,
very small degenerative giant cells in an otherwise normal smear. The
testes of the other had numerous giant cells with up to fifteen nuclei of
primary secondary spermatocytes, or spermatids. Late and early sper-
matids were markedly degenerate, and secondary spermatocytes were
undergoing chromolysis. These findings are indicative that Synaptomys
may undergo testicular regression in this region at this time of year.
376
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Clethrionomys gapperi gapped. Four of these voles, of which all
were taken in 1949, came from around the edges of a white cedar-black
ash bog under fallen logs and hummocks. They are probably restricted to
such habitats, and on this basis this vole could not be considered a very
abundant mammal in the region. A higher, less extensive, and less boggy
spruce bog, although containing many fallen moss-grown logs and ap-
parently similar microhabitats, did not yield any of these voles. None
was caught in 1950.
One female was immature; the other was mature, lactating, and con-
tained five right and no left implantation scars or corpora lutea. No
transmigration of ova from one uterine horn to the other had taken place.
Both males were reproductively mature with abundant sperm in the
epididymes. The testes of both of these red-backed mice showed marked
degenerative changes with giant cells of up to thirty secondary sperma-
tocytes or spermatids. Some giant cells involved primary spermatocytes.
All individual spermatids had obvious degenerative changes. The picture
was one of extremely rapid degeneration in early July at the height of
reproductive activity.
Microtus pennsylvanicus pennsylvanicus. Ten specimens were
captured in 1949 ; one in a small spruce bog, one in moist hardwoods, one
in a dry, old field, one in a sedge marsh, and six in an alder bog and sedge
marsh. All but one of these were taken in moist habitats, all but two of
which were wooded. This seems somewhat contrary to the usual type of
habitat associated with Microtus pennsylvanicus. In this region, where
cultivated or fallow fields are scarce, Microtus apparently inhabits sites,
represented by the wet alder and hardwood bogs and sedge marshes,
which are the type they originally occupied prior to the invasion of agri-
culture. Microtus appears to occupy the wetter spots with deciduous
cover, Clethrionomys the coniferous edges of bogs, and Synaptomys the
extensive sphagnum bogs and their edges. No specimens of Microtus were
trapped in 1950.
Three of these voles were very small, apparently fresh from a nest
beneath a snarl of alder and white pine roots, as I surprised them while
out wandering in the afternoon. They were caught while running franti-
cally for cover. Possibly many small mammals are frightened into cross-
ing traps. One of these was sexually mature (Christian, 1950b).
One fully grown male contained abundant sperm in the epididymis,
the other was immature with no epididymal sperm, although the testes
1951 Christian: Mammals of Mazinaw Region, Ontario 377
were scrotal, but small. The adult mammal showed advanced degenera-
tive changes similar to the Clethrionomys above with extremely abundant
giant cells and vacuolar changes. It may be that adult males of Microtus^
functionally active in the spring, undergo testicular regression at this
time.
Ondatra zibethica. Muskrats are one of the principal furbearers in
this region, but not as much so as farther south where there are more
extensive habitats. Extensive cattail bogs are not numerous in this area;
so the population of muskrats is scattered over numerous, small, suitable
areas, such as marshy areas, shallow lakesides, small ponds, and the
larger, slow-flowing streams. There was apparently a high population in
1949, but a severe drop had occurred by 1950, at the time of the spring
trapping period. Albert Spencer had expected a catch of 500 muskrats,
but caught only 150, or a little better than twenty-five percent of the ex-
pected catch. Irving Brown likewise stated that his catch was only twenty-
five percent of the usual number, and further that the catch for the entire
Lake Ontario region was down to a quarter of the usual catch. Apparently
there was a general decrease of three quarters in the expected muskrat
catch throughout the entire area. Muskrats are prime in late March or
early April in this territory.
Mus musculus. The house mouse apparently has not invaded this
area, partly due to the abandoning of most of the houses and cabins for a
good part of the year, especially the winter.
Rattus norvegicus. According to reports, rats are found in Cloyne
and about its dumps. By their destruction of foods stored in the cellars,
they constitute a serious economic pest, and a real threat locally during
the winter, when they move indoors.
Zapus hudsonius 1
Napaeozapus insignisj The “jumping mice” are apparently not
common in the area, as none was caught or seen. The “kangaroo” mouse
was described very accurately by Mr. Fred Garbutt from one his dog had
caught on the rocky, west shore of Mazinaw Lake two years ago. The
animal was most likely Zapus, since Mr. Garbutt was certain there was
no white tip on the tail, and the rest of his description was so accurate that
it does not seem likely that he would have missed it. This is the only one
he has seen in sixteen years of residence in this area.
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Erethizon dorsatum dorsatum. Porcupines are very common in the
area, and they were frequently killed on the highway. Due to their destruc-
tive habits and the danger to dogs, they are shot on sight. They are not,
apparently, destructive to the pine or other timber trees in the area, as
several people evinced surprise when told that porcupines are known to
“bark” pines and other trees.
Lepus americanus americanus. The snowshoe hare is the common
“rabbit” of the region, and was very common both in 1949 and 1950.
Reportedly they were much more abundant in the latter year than in
1949. This abundance is impressive when all reports indicate that there
were none to be seen in 1947, and very few in 1948. They were seen
nightly along the highway in 1949, to which they were drawn by the
calcium chloride. In 1950 the practice of spreading this chemical had been
discontinued and work was progressing on the road, so not so many were
seen. Numerous hare signs, forms, and runs were noted in all of the marshy
areas trapped in both years, and the hares were flushed in habitats II
and V in 1949. One lactating female hare was collected.
The ruffed grouse population in this area has exactly paralleled the hare
population for at least the last four years, and they were abundant and
accompanied by large coveys of young in both 1949 and 1950.
Odocoileus virginianus borealis. Deer are relatively abundant,
though much less common than in irruptive areas of the United States,
such as Pennsylvania. The deer average larger than those further south,
and 250-pound bucks are not at all uncommon. At present there is much
local agitation to have no closed season on deer due to crop damage.
Should this come to pass, I doubt if the deer would suffer very much, as
most of the country is relatively inaccessible and extremely difficult to
hunt. Even now most deer hunting is done with dogs, which would
probably be disastrous elsewhere.
Alces americanus. Moose are uncommon in the summer, but are
frequently seen in the other three seasons. Several were seen along Mazi-
naw Lake in 1949, and the fire rangers see them from time to time back in
the bush. Moose tracks were seen about two miles west of Mazinaw Lake
about two weeks before the trapping period of 1950. They are most fre-
quently seen in the spring.
1951
Christian: Mammals of Mazinaw Region, Ontario
379
IV. Discussion of Populations
From the foregoing account it is evident that the small mammal popu-
lation in the Mazinaw region decreased considerably in 1950 from the
previous year. Examination of the trapping summaries in the section on
habitats reveals the marked decrease in trap line productivity in 1950
as compared to 1949. It is true that trap-night figures will not give a
quantitative picture of an area, but they will give a fairly accurate picture
of relative changes under the conditions of this study. These conditions
are that one person did all of the trapping, reducing the subjective
variables of trap setting to a minimum; the same period of time is involved
in both years; only three of the ten habitats were trapped in both years,
but in both the same general area was involved; an effort was made to
set the traps in as nearly the same concentration and manner in 1950 as
in the preceding year; traps were moved often enough to avoid catching
more than a minimum of stragglers; and the same bait was used each year.
For the sake of uniformity, the figures for 1950 for area I have been
omitted from the succeeding discussion, as the rat traps used there may
have altered the situation. I believe that the following figures represent a
valid picture of the relative population changes from 1949 to 1950.
In 1949, 54 small mammals of all species were trapped in 870 trap-
nights, or one catch per 16 trap-nights. In 1950, only 12 mammals were
caught in 1,127 trap-nights, or 1 catch in 94 trap-nights. This means that
the small mammal population for the Mazinaw region in 1950 was only
17% of the 1949 level. The following figures give the breakdown into
species using trap-night figures. The long-tailed shrews are lumped in one
group, as are the mice and voles. The SoricidcB were 108% of the 1949
level which, considering the number of shrews involved, represents an
unchanged population. Blarina was down 75% from its 1949 figure, and
the mice and voles, represented by one Peromyscus m. gracilis, were down
98%. The population drop, then, was due to Blarina, Peromyscus m.
gracilis, Peromyscus 1. novehoracensis, Synaptomys, Clethrionomys, and
Microtus. A similar, but not so marked, drop in population was ex-
perienced in census lines from the spring of 1949 to the spring of 1950 at
Philadelphia, Pa. In this instance, Blarina dropped 71%, Microtus 86%,
and Peromyscus 1. novehoracensis increased 67% of the 1949 levels, to
make an overall decrease of 48%. The principal difference seems to be
in Peromyscus, which decreased in Ontario and increased in Pennsylvania.
Peromyscus was in phase with the other small mammals (other than
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shrews and moles) in Ontario, while it was out of phase with the others
in Pennsylvania. The principal reason for bringing these figures in at this
point is to show how much more severe the drop was in Ontario than it
was 400 miles further south, and at 1,000 feet lower altitude, although the
drop was experienced simultaneously in both places.
It is interesting that the small mammal decline coincided with the
decline of muskrats throughout southern Ontario, where the spring catch
was 25% of the expected level. However, snowshoe hares, ruffed grouse,
woodchucks, and the predatory fur bearers of the region, were all abundant
and reportedly increased over 1949 levels. Red squirrels were obviously
way down from the 1949 level in this region, but no estimate can be made
of the degree of the decline. This probably is a local change, as they have
reportedly remained abundant in Algonquin Park (Fowle, 1950), al-
though the other declines were also noted there. Based on sight observa-
tions, chipmunks were apparently at the same high level in 1950 as in
1949. Due to present Ontario trapping laws, beavers are increasing
rapidly, and should reach high levels in the near future.
The Ontario laws license each trapper to trap in a specified area. No
one else may trap in this area, and the trapper may keep it as long as he
traps in it. This has added a powerful incentive for the trappers to be
conservationally minded. The number of animals which may be removed
without endangering the next year’s catch is checked by each trapper, so
that complete removal of all muskrats and beavers has ceased. In most
instances, the trappers have been planting wild rice and cattails to in-
crease muskrat forage. Also, wardens are assisted in the apprehension of
poachers by the trappers concerned. All in all, these trapping practices
have produced healthy results in the region, and have won the support
of the trappers, but it remains to be seen what the final results will be in
terms of population dynamics. In spite of this care, the muskrats declined,
and it is conceivable that beavers, if allowed to increase excessively, will
become cyclic and experience severe declines (Christian, 1950c).
V. Discussion of Data on Reproduction in Relation to
Population Dynamics
As a preamble to the discussion of the Canadian data it is necessary to
discuss briefly some findings from the Philadelphia area in April, 1949,
and April, 1950. In the spring of 1949, as already mentioned, the popula-
tion of small mammals in the Philadelphia region was high, while in the
same period of 1950 the population of small mammals, except Peromyscus,
1951 Christian: Mammals of Mazinaw Region, Ontario 381
was at a low ebb. Smears of the testes were made of twelve adult male
mammals {Microtus, Blarina, Peromyscus) trapped in this area in 1949.
The testes of nine of these animals showed advanced degenerative changes
with very numerous giant cells of up to thirty degenerate spermatids,
secondary spermatocytes, or occasionally primary spermatocytes. Chro-
molytic changes were noted in a large portion of the spermatids and
secondary spermatocytes. In some there was a marked reduction of sper-
matozoa and spermatids. The testes of all of these mammals were scrotal,
and some or many spermatozoa were found in the epididymes. Two
animals had mild degenerative changes with fewer and smaller giant
cells, while one had a normal smear. In the spring of 1950, however, only
one testicular smear in eight showed any marked degenerative changes,
although the time of the year and the climatic conditions were about the
same.
In a previous paper (Christian, 1950c), I discussed the possible effect
of high population stresses on the reproductive potential of the species
involved. It seems likely that the severe stresses attending the spring
breeding season of a high population level seen in 1949 account for the
degenerative changes found in the testicular smears. By contrast the nor-
mal smears of 1950 reflect a low population with a minimum of stressors.
That various environmental, psychological, or traumatic stresses may
produce these testicular changes, of which giant cell formation is char-
acteristic, now seems well established (Selye, 1950). This degeneration
is apparently caused by the withdrawal of pituitary gonadotrophin
(hypophystectomy is the most potent producer of these changes) to favor
the production of adrenocorticotrophin. The census taken in the fall in
the same area revealed the population to be at the same level as in the
early spring, instead of being increased. This would be predictable from
the appearance of the spring testes, since the degenerative changes were
indicative of a lower reproductive potential. Whether these changes would
result in a complete cessation of breeding in each animal, or only in a
reduction in the number of sperm produced, would probably depend on
the severity of the stress in the animal. Even though sperm were pro-
duced, a lower count could be too low for successful fertilization. The
effects of these factors on the female in the wild is not known, but ex-
perimental stress causes the ovaries of white rats to atrophy and the
animal to go into permanent anestrus (Selye, 1939).
In contrast with the testes of mammals taken at Philadelphia in early
spring of 1949 and 1950, adult males of Sorex cinereus, Blarina, Tamias,
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VOL. 31
Peromyscus m. gracilis, Synaptomys, Clethrionomys, and Microtus taken
in Ontario, all showed marked degenerative changes of the testes during
the first two weeks of July in both years. Peromyscus 1. novehoracensis
was an exception in that only two of five with sperm in the epididymes,
and one of three juvenile males showed any degenerative changes. De-
generation was especially severe in the microtines and soricids. Five
adult males of Blarina were examined in the two years, and four of these
showed degenerative changes. Adults of the other mammals were examined
in one year or the other; Sorex and Tamias in 1950, and the others in 1949.
All of the adults of Sorex cinereus, Peromyscus m. gracilis, Synaptomys,
Cleihrionomys, and Microtus examined showed degenerative changes.
It would seem, since the testicular changes were found in both years of
peak and low populations in the Ontario mammals, that something is
working on all species other than the stress effects of a high population,
as has been suggested for the “1949” Philadelphia animals. Since all
species are involved, and the degenerative changes range from none (in
Peromyscus 1. novehoracensis) to advanced stages, it would seem that the
causative agent had probably not been operative for very long. Logically,
the passing of the summer solstice with the resultant decreasing day-
lengths would appear to be the causative factor. The relation of light and
other exteroceptive factors to reproduction has been previously referred
to (Christian, 1950c), and the subject has been well summarized by Bur-
rows (1949). It is clear that the daily added light increment is the im-
portant element in bringing an animal to full reproductive activity and to
maintaining it, while decreasing light resulted in testicular regression in
every case investigated. Since stress probably can be ruled out as a result
of similar findings in years of divergent populations, light must the the
factor involved in this early-summer male reproductive regression found
following the summer solstice in all of the species of small mammals con-
sidered. The timing certainly seems more than coincidental.
The possibility of this being an artifact resulting from autolytic changes
while the animal was in the trap has been ruled out by experiments in
which one testis of a normal mouse was “smeared” immediately after
death and the other at various intervals after standing at from 70 to
80°F. No differences between the smears of the two testes were observed
at any time interval up to eight hours. Technical artifacts have been
eliminated by frequent smears of laboratory mice, which were normal in
every instance, as well as by comparison with normal smears in the wild-
trapped mammals.
1951 Christian: Mammals of Mazinaw Region, Ontario 383
It is well known that the spring breeding season reaches a peak, and is
followed by a sharp fall in reproductive activity, best expressed as the
per cent of females which are pregnant. Coventry (1937) reviews this
situation in connection with his own material from Ontario, and explains
the fall in the per cent of pregnant females captured, by postulating a
decrease in activity as a result of almost one hundred per cent of preg-
nancies earlier in the season. The present testicular findings indicate,
however, that the testes in the males which were adult during the spring
breeding season undergo regression following the solstice. This time would
be extended three weeks for pregnant females; also, female reproductive
activity may be similarly affected. The gradually increasing but low level
of pregnancies during the balance of the summer could well be accounted
for by young of the year reaching maturity, or by a few adults which did
not regress markedly. C. D. Fowle (1950) reports that few, if any, preg-
nant individuals of Sorex cinereus or Blarina are taken after the middle
of July in Algonquin Park, Ontario. Coventry’s (1937) peaks of per cent
pregnancies for Peromyscus m. gracilis and Clethrionomys g. gapperi from
Temagami, Ontario, correspond exactly with the present findings, if
conception is dated at an average of two weeks prior to his peak (preg-
nancy first being grossly detectable at six days with a twenty-one day
gestation period).
The present material indicates, then, that the spring breeding season is
terminated by the decreasing day-lengths following the summer solstice
in Sorex cinereus, Blarina brevicauda talpoides, Tamias striatus lysteri,
Peromyscus maniculatus gracilis, Peromyscus leucopus noveboracensis (at
least to some extent), Synaptomys cooperi, Clethrionomys gapperi, and
Microtus pennsylvanicus from the Mazinaw Lake region of Ontario.
In view of the above discussion it is evident that an actual die-off is
not essential for a marked population decline. All that would be required
to seriously curtail the population would be sufficient stress to markedly
reduce reproductive activity of adults from the start of the spring breed-
ing season until the summer solstice. With this occurring, along with a
normal or increased death rate, the number of animals produced would
be markedly reduced, and consequently the number surviving the suc-
ceeding winter to breed in the following spring would be correspondingly
reduced. A population decline of this type would be more likely to occur
in more temperate regions, while the actual marked die-off would be
expected in the more northern regions which have severer stresses. This
would certainly be true for the limited species-food ecology of the sub-
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arctic. A greater reproductive potential farther north (Christian, 1950c)
might still be expected, with a much more intense activity during the
spring breeding season. Some evidence of this has been noted, but it is,
as yet, inconclusive.
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Anderson, R. M.
1946. Catalogue of Canadian Recent mammals. Nat. Mus. Can., Bull.
102, Biol, sen, 31, pp. v + 238.
Burrows, H.
1949. Biological actions of sex hormones. 2nd Ed. Cambridge Univ.
Press, pp, ix + 615.
Christian, J. J.
1950 a. A field method of determining the reproductive status of small
male mammals. Journ. Mammal., 31, pp. 95-96.
1950 b. A precocious male Microtus pennsylvanicus. Journ. Mammal.,
31, pp. 195-196.
1950 c. The adreno-pituitary system and population cycles in mammals.
Journ. Mammal., 31, pp. 247-259.
Coventry, A. F.
1937. Notes on the breeding of some Cricetidae in Ontario. Journ.
Mammal., 18, pp. 489-496.
Fowle, C. D.
Personal communication.
Halliday, W. E. D.
1937. A forest classification for Canada. Canad. Dept. Mines and
Resources, Forest Service Bull., 89, pp. 1-50.
Hamilton, W. J., Jr.
1943. The mammals of eastern United States. Comstock Publ. Co.,
Ithaca. Pp. 1-432.
Jackson, H. H. T.
1928. A taxonomic review of the American long- tailed shrews. U. S.
Dept. Agric., Bur. Biol. Surv., N. A. Fauna No. 51, pp. 1-238.
Selye, H.
1939. The effect of adaptation to various damaging agents on the fe-
male sex organs in the rat, Endocrin., 25, pp. 667-672.
1950. Stress. Acta Inc., Montreal, pp. xx -f- 822.
1951
Christian: Mammals of Mazinaw Region, Ontario
385
Addendum
The normal skunk {Mephitis mephitis nigra) adrenal is a rounded, un-
lobulated, somewhat biconcave disc. The zonation is similar to that found
in other mammals. The glomerulosa, however, is distinctly divided from
the outer fasciculata by a thin connective tissue septum which is clearly
defined in ordinary preparations, and is richly invested with vascular
channels. The glomerulosa of the normal animal is about a fifth the width
of the entire cortex. Dividing the reticularis from the medulla is a thick,
well-defined hyaline connective tissue septum containing numerous veins
and small blood sinuses. This connective tissue boundary is denser and
much more pronounced than has been seen by the writer in any other
species of mammal. The medulla contains the usual granule-filled chro-
maffine cells. The microscopic appearance of the cortical cells is similar to
that seen in other normal, healthy animals.
In the dying female skunk from Mazinaw Lake, Ontario, the cells of
the glomerulosa are shrunken and the nuclei pyknotic. In some areas the
glomerulosa is replaced by small foci of hyaline necrosis. The thin band of
connective tissue between the glomerulosa and fasciculata is not seen.
The changes in the fasciculata are very marked. The outer half of this
zone is necrotic, having formed a coalesced mass of granular cytoplasm
containing pyknotic and karyorhetic nuclei. In a few places this zone of
necrosis is interrupted by a few cords of relatively normal appearing
fasciculata cells. Central to this band of necrosis the cells of the inner half
of the fasciculata and the entire reticularis are shrunken, and the sinusoids
are widely dilated. The cytoplasm of approximately half of these cells
have the characteristic foamy appearance of fine vacuolation. In several
places, involving about one sixth of the entire gland area, wedge-shaped
zones of hyaline necrosis extend from the medulla to the outer fasciculata.
There has been extensive hemorrhage into these zones of necrosis, and all
zones are heavily congested.
The connective tissue separating the medulla from the cortex is enorm-
ously widened, almost the entire width of the cortex in one gland and ten
to thirty-fold in the other. This band and all of the thinner connective
tissue septa are hyalinized, and scattered through them are large pha-
gocytes filled with ingested red blood cells. Arteries, other than a few in
the areas of hyalinosis, appear to be normal.
The medullary chromaffine cells are small, deeply staining, and have
relatively little cytoplasm, which is free of the characteristic baso-
386
Annals of the Carnegie Museum
VOL. 31
philic granules. The nuclei appear normal with distinct nucleoli. Through-
out the medulla, however, there are islands of cortical cells replacing over
half of the chromaffine tissue. A few of these islands are glomerulosa
cells, but the majority are fasciculata cells containing abundant small
cytoplasmic vacuoles. None of these misplaced cortical cells are necrotic,
and appear to be normally functional.
Both of the adrenals of this skunk present the same appearance, both
exhibiting the same marked degree of destruction and cortical metaplasia.
The degree of damage is suggestive of the amount seen in cases of clinical
Addison’s disease; and, although there were limited numbers of normal
appearing cortical cells, the destruction of both the cortex and the me-
dulla must have been severe enough to seriously reduce the functional
capacity of the glands, as well as impair the circulatory supply. I have
already indicated that a semi-quantitative test showed these adrenals
contained a moderate amount of ascorbic acid. This can only be accounted
for by hyperactivity of the remaining normal cells.
The marked degree of hyalinosis, along with hyperplasia and meta-
plasia, which has occurred in these glands supports the previous assump-
tion of long-standing stress with the more acute necrosis resulting from the
final exhausting demands on the gland. The microscopic appearance of the
adrenals of this skunk is a striking demonstration of the effect of excessive
stress in a wild mammal.
INDEX
ji '-l (^\ji '^ 5-
abieticola, Buteo jamaicensis, 291
abieticola, Odontia, 26
abramovi, Karanasa, 328
Adrenal glands, 370
adustum, Steccherinum, 27
aksuensis, Karanasa, 324
alascensis, Buteo borealis, 291
albescentior, Certhiaxis cinnamomea,
38
albido-brunnea, Thelephora, 250
albobadium, Stereum, 243
albofaciata, Pierella, 82
albofaciata, Pierella albofaciata, 82
Alces americanus, 378
alpherakyi, Karanasa, 327, 329
amalia, Pierella, 68
Ambystoma cingulatum, 299-320
annulatum, 299
texanum, 299
mabeei, 299
schmidti, 299
angulatum, 307
americana, Martes, 367
americanus, Alces, 378
americanus, Euarctos, 367
americanus, Lepus americanus, 378
amicus, Calodon, 23
Angola, Two new Passerine birds from,
by James P. Chapin, p. 1-4
angrena, Karanasa, 324
angulatum, Ambystoma, 307
angusticeps, Morchella, 283
Anhydrophryne, 265
annulatum, Ambystoma, 299
anthe, Satyrus, 329
anthocephala, Thelephora, 250
Aratinga astec melloni, 297
arpensis, Karanasa, 326
Arthroleptella, 265
aspersa, Pipa, 263
astec melloni, Aratinga, 297
astorica, Karanasa, 322
astyoche, Pierella, 69
atroviride, Hydnum, 25
Automolus infuscatus purusianus, 40
melanopezus, 41
pallidigularis pallidigularis, 41
Avinoff, Andrey, An analysis of color
and pattern in butterflies of the
Asiatic genus Karanasa, 321-332
axanthus, Pseudobranchus striatus, 277
bannermani, Terpsiphone rufocinerea, 3
Benesh, Bernard, Sphoenognathus cur-
vipes (Coleoptera, Lucanidae),
a new species from Bolivia, 45-48
bishopi, Ambystoma cingulatum, 299-
320
Blarina brevicauda talpoides, 366
bohemica, Verpa, 284
boissonneautii, Pseudocolaptes bois-
sonneautii, 39
boliviana, Pierella stollei, 74
borealis, Buteo jamaicensis, 290, 291
borealis, Odocoileus virginianus, 378
borealis, Falco, 290
boultoni, Bradypterus mariae, 1
Bradypterus mariae boultoni, 1
brasilensis, Pierella lena, 66
Breviceps, 265
Brinkmanni, Grandinia, 24
briseis, Satyrus, 329
Brown, F. Martin, Taxonomy and dis-
tribution of the genus Pierella
(Lepidoptera), 49-88
brunnescens, Sclerurus rufigularis, 42
Burtianum, Stereum, 243
Buteo borealis alascensis, 291
Buteo buteo, 289
buteo, Buteo, 289
Buteo jamaicensis, 289
Buteo jamaicensis abieticola, 291
MATS
388
Annals of the Carnegie Museum
VOL. 31
Buteo jamaicensis borealis, 290, 291
[Biiteo jamaicensis] caliirus, 290
Buzzard, American, 290
Buzzard, Cream-colored, 290
Buzzard, European, 289
Cabomba, 280
Caldesiella ferruginosa, 23
Caligo, 328
Calodon amicus, 23
scrobiculatus, 23
velutimis, 23
zonatus, 23
calurus, [Buteo jamaicensis], 290
calvum, Mucronella, 26
Camp>lorhamphus procurvoides suc-
cessor, 13
canadensis, Castor, 373
canadensis, Lutra canadensis, 369
canadensis, Lynx, 370
Canis lupus hxaon, 370
cantharellus, Craterellus, 240
caroliniana, Elvela, 285
caryopliyllea, Thelephora, 2, 50
Castor canadensis, 373
caurensis, Xiphorhynchus pardalotus, 9
Certhiaxis cinnamomea albescentior, 38
certhiolus, Lcpidocolaptes angustiros-
tris, 13
chalybaea, Pierella rhea, 73
Chapin, James P., Two new Passerine
birds from Angola, 1-4
Chickadee, Hudsonian, 333
Chiroptera, 367
Chondrotus cingulatus, 307
Christian, John J., The mammals of the
Mazinaw Lake region of Ontario;
tlieir reproduction and population
dynamics, 339-386
cicognanii, Mustela erminca, 368
Cinclodes fuscus oreobates, 34
cinercoargenteus, Urocj'on, 370
cine re us cinereus, Sorex, 362
cingulatum, Amb3"stoma bishopi, 299-
320
cingulatus, Chondrotus, 307
cinnamomeus, Irpex, 25
cinnamomea, Synallaxis, 38
claresignata, Plyla, 268
Clethrionomys gapperi gapperi, 376
columbianus, [Penthestes hudsonicus],
334
concolor, Dendrocolaptes, 7
Condjdura cristata cristata, 362
conica, Morchella, 283
connectens, Xiphorhjmchus guttatus, 8
conradti, Karanasa, 323, 329
cooki, Eleutherodactylus, 264
cooperi, Sjmaptomys cooperi, 375
coralloides, Hericium, 24
cornucopioides, Craterellus, 241
crassipes, Morchella, 283
Craterellus cantharellus, 240
cornucopioides, 241
lutescens, 241
Crenshaw', Jr., John W., and Coin,
Coleman J., Description of a new
race of the salamander Pseudo-
branchus striatus (Le Conte),
277-280
crispa, Elvela, 285
Sparassis, 242
cristata cristata, Condylura, 362
cristatum, PIj'dnum, 25
Cryptobatrachus, 267
Crj^ptobatrachus fuhrmanni, 266
Curtisii, Hymenochsete, 241
curvipes, Sphoenognathus, 45
darwinii, Rhinoderma, 263, 265, 266
decepta, Pierella albofaciata, 83
decolorata, Karanasa, 323, 327
deliciosa, Morchella, 283
Dendrocincla merula obidensis, 16
meruloides neglecta, 16
Dendrocolaptes certhia juruanus, 6
Dendrocolaptes certhia ridgw'ayi, 6
concolor, 7
Dendroplex picus dugandi, 12
Dendroplex picus rufescens, 11
Dentium repandum, 24
diaphanum, Stereum, 244
1951
Index
389
dionysius, Neominois, 327
Diplocardia, 314
Direct development (in frogs), 257-276
dissoluta, Karanasa, 324
distinctus, Xiphorh3mchus triangularis,
8
divaricatus, Heiniphractus, 264, 266
dorsatum dorsatum, Erethizon, 378
dracontis, Pierella hyalinus, 64
dugandi, Dendroplex picus, 12
duidae, Lepidocolaptes laj’ardi, 12
egg tooth, 266
elastica, Elvela, 285
Eleutherodactylus, 263, 266
Eleutherodactylus cooki, 264
nubicola, 267
Elvela caroliniana, 285
crispa, 285
elastica, 285
infula, 285
underwoodii, 285
Elvellaceae, 281
Epinephele, 329
Erethizon dorsatum dorsatum, 378
erinaceus, Hericium, 24
ershoffi, Karanasa, 323
esculenta, Morchella, 283
Euarctos americanus, 367
Eumeces fasciatus, 344
Evolution (tadpoles), 267-269
evura, [Penthestes hudsonicus], 334
exilis, Furnarius leucopus, 33
Falco borealis, 290
Falco jamaicensis, 290
fasciatum, Stereum, 244
fasciatus, Eumeces, 344
ferruginosa, Caldesiella, 23
Fishes (labial folds), 266-267
fragillissima, Mycoacia, 26
frustulosum, Stereum, 245
fuhrmanni, Cryptobatrachus, 266
fuligineo-violaceum, Hj^dnum, 25
fulva, Vulpes, 370
fumeus fumeus, Sorex, 364
furfurosus, Sclerurus rufigularis, 41
Furnarius leucopus exilis, 33
torridus, 33
fusco-atra, Odontia, 26
fuscum, Stereum, 245
fusimaculata, Pierella hyalinus, 64
gapperi, Clethrionom3-s gapperi, 376
Gastrotheca, 263, 264
gausapatum, Stereum, 245
Geomyidae, 335
gej’eri, Satyrus, 326, 327
Glaucomys sabrinus macrotis, 372
Gloiodon strigosus, 24
Glyphorynchus spirurus, 13
Glyphorynchus spirurus subrufescens,
14
Goin, Coleman J. and Crenshaw, Jr.,
John W., Description of a new
race of the salamander Pseudo-
branchus striatus (Le Conte),
277-280
Goin, Coleman J., A study of the sala-
mander, Ambystoma cingula-
tum, with the description of a
new subspecies, 299-320
gracilis, Peromyscus maniculatus, 373
Grandinia Brinkmanni, 24
Gregorymys kayi, 335
griseonota, Synallaxis albescens, 37
grumi, Karanasa, 329
habits of spheniscus, 280
Hawk, Northern Red-tailed, 291
Hawk, Red-tailed, 289
Heliconia, 325
helvina, Pierella, 57
helvina, Pierella, helvina, 58
Hemiphractus divaricatus, 264, 266
Henry, LeRoy K., A review of the
Hydnaceae (Fungi) of Western
Pennsylvania, 19-32
Henry, LeRoy K., A review of the
pileate Thelephoraceae (Fungi)
of Western Pennsylvania, 239-
256
390
Annals of the Carnegie Museum
VOL. 31
Henry, LeRoy K., A review of the
Elvellaceae (Fungi) of Western
Pennsylvania, 281-288
Hericium coralloides, 24
erinaceus, 24
laciniatum, 25
Himantia, Mycoacia, 26
hirsutum, Stereum, 246
hortona, Pierella, 80
hudsonicus, Parus, 333
hudsonicus, Penthestes, 334
hudsonius, Zapus, 377
huebneri, Karanasa, 324
H)'aline necrosis, 385
hyalinus, Pierella, 62
hj’alinus, Pierella hyalinus, 63
hybrida, IMorchella, 284
hyceta, Pierella, 76
hyceta, Pierella hyceta, 76
H37dnaceae (Fungi) of western Penn-
S5dvania, A review of, by LeRoy
K. Henry, 2 pis., 19-32
Hydnuni atroviride, 25
cristatum, 25
fuligineo-violaceum, 25
imbricatum, 25
Underwoodii, 25
Hjda claresignata, 268
H^mienochaete Curtisii, 241
rubiginosa, 241
tabacina, 242
hypoleuca, Synallaxis albescens, 37
imbricatum, Hj^dnum, 25
incanescens, Pierella helvina, 58
infula, Flvela, 285
intermedia, Karanasa, 327, 328
intervectus, Microsorex hoyi, 365
int3^bacea, Thelephora, 250
Irpex cinnarnomeus, 25
mollis, 25
jamaicensis, Buteo, 289
jamaicensis, Falco, 290
juruanus, Dendrocolaptes certhia, 6
kafir, Karanasa, 323
Karanasa, 321-330
kayi, Gregorymys, 335
kazakstana, Karanasa, 323
korlana, Karanasa, 326
labradorius, Penthestes hudsonicus, 334
laciniatum, Hericium, 25
lactea, Karanasa, 323
latefasciata, Karanasa, 323
latona, Pierella hyceta, 76
leechi, Karanasa, 323
lena, Pierella, 65
lena, Pierella lena, 66
Lepidocolaptes angustirostris, 13
layardi duidae, 12
Lepidocolaptes layardi madeirae, 12
lepturum, 316
lapturus, Linguaelapsis, 307
Lepus americanus americanus, 378
lesbia, Pierella luna, 79
Lethe, 329
leucotis, Sciurus carolinensis, 372
Linguaelapsis lepturus, 307
littoralis, [Penthestes hudsonicus], 333
Littoralis, Synallaxis albescens, 36
loquax, Tamiasciurus hudsonicus, 372
lotor, Procyon, 367
lucia, Pierella, 69
luna, Pierella, 78
luna, Pierella luna, 78
lutescens, Craterellus, 241
Lutra canadensis canadensis, 369
lycaon, Canis lupus, 370
Lynx canadensis, 370
rufus, 371
lysteri, Tamias striatus, 371
mabeei, Ambystoma, 299
MacMillan, Gordon K., The land snails
of West Virginia, 89-238. Index
to this paper is found immedi-
ately following page 238
macrotis, Glaucomys sabrinus, 372
madeirae, Lepidocolaptes layardi, 12
Marmota monax rufescens, 371
1951
Index
391
Martes americana, 367
pennant i, 367
Mazinaw Lake region, mammals of,
339-386
Melanargia, 329
melanopezus, Automolus, 41
melloni, Aratinga astec. 297
Mephitis mephitis nigra, 369, 385
Microbatrachylus, 267
Microhylidae, 265
Microsorex hoyi intervectus, 365
Microtus pennsylvanicus pennsyivani-
cus, 376
mollis, Irpex, 25
Morchella angusticeps, 283
conica, 283
crassipes, 283
deliciosa, 283
esculenta, 283
hybrida, 284
Mouthparts, tadpole, 265-269
Mucronella calvum, 26
muirheadi, Neope, 326
multiguttatus, Xiphorhynchus obsole-
tus, 10
multipartita, Thelephora, 250
Murrayi, Stereum, 246
Mus musculus, 377
musculus, Mus, 377
Mustela erminea cicognanii, 368
frenata noveboracensis, 368
rixosa, 368
vision vision, 368
Mycoacia fragillissima, 26
Himantia, 26
setosa, 26
Myriophyllum, 280
naryna, Karanasa, 322
necopinus, Xiphorhynchus, 10
Nectophrynoides, 257, 267
Nectophrynoides occidentalis, 257, 268
vivipara, 257
Nectophrj-noides tornieri, 257-262, 263,
265, 266, 268
neglecta, Dendrocincla meruloides, 16
Neominois, 323, 325, 326, 327, 328
Neope, 325, 326, 327
Neotoma sp. (?), 375
nereis, Pierrella, 57
nigra. Mephitis mephitis, 369
nigricans, [Penthestes hudsonicus], 333
noveboracensis, Mustela frenata, 368
noveboracensis, Peromyscus leucopus,
374
norvegicus, Rattus, 377
nubicola, Eleutherodactylus, 267
Nyssa biflora, 280
obidensis, Dendrocincla merula, 16
obscurior, Synallaxis cabanisi, 37
occidentalis, Nectophrynoides, 257, 268
ochraceoflavum, Stereum, 246
ochraceum, Steccherinum, 27
ocreata, Pierella helvina, 59
Odocoileus virginianus borealis, 378
Odontia abieticola, 26
fusco-stra, 26
Ondatra zibethica, 377
orbiculare, Radulum, 27
orenocensis, Xiphocolaptes orenocensis,
6
oreobates, Cinclodes fuscus, 34
ornatus, Xiphorhynchus elegans, 10
Orton, Grace L,, Larval development of
Nectophrynoides tornieri (Roux) ,
with comments on direct de-
velopment in frogs, 257-276.
Oven-birds, Critical remarks on, by
W. E. C. Todd, p. 33-43
Ovoviviparous frogs, 257
pallidigularis, Automolus pallidigularis,
41
palmata, Thelephora, 250
palustris albibarbis, Sorex, 365
pamira, Karanasa, 328, 329
Paroquet, a new race of, 297
Parus hudsonicus, 333
parvulus, Zachaenus, 265
Passerine Birds from Angola, Two new,
by James P. Chapin, p. 1-4
392
Annals of the Carnegie Museum
VOL. 31
pennanti, Martes, 367
penns3Tvanicus, Microtus pennsylvani-
cus, 376
Penthestes hudsonicus, 334
[Penthestes hudsonicus] columbianus,
334
[Penthestes hudsonicus] evura, 334
Penthestes hudsonicus labradorius, 334
[Penthestes hudsonicus] littoralis, 333
[Penthestes hudsonicus] nigricans, 333
[Penthestes hudsonicus] rabbittsi, 334
[Penthestes hudsonicus] ungava, 333
Peromyscus leucopus noveboracensis,
374
Peromyscus maniculatus gracilis, 373
perpallida, Synallaxis albescens, 36
Philydor erythrocercus suboles, 39
Phlebia radiata, 26
strigosa-zonata, 26
Pierella, 327
Pierella albofaciata, 82
Pierella albofaciata albofaciata, 82
amalia, 68
astyoche, 69
stollei boliviana, 74
lena brasilensis, 66
rhea chalybaea, 73
albofaciata decepta, 83
hyalinus, dracontis, 64
hyalintis fusimaculata, 64
helvina, 57
Pierella helvina helvina, 58
hortona, 80
hyalinus, 62
Pierella hyalinus hyalinus, 63
hyceta, 76
Pierella hyceta hyceta, 76
Pierella helvina incanescens, 58
hyceta latona, 76
lena, 65
Pierella lena lena, 66
Pierella (Lepidoptera), Taxonomy and
distribution of the Genus, by
F. Martin Brown, 49-87
Pierella lucia, 69
luna, 78
Pierella luna lesbia, 79
Pierella luna, luna, 78
Pierella luna rubeola, 79
stollei, 73
Pierella nereis, 57
helvina ocreata, 59
rhea, 71
Pierella rhea rhea, 72
Pierella stollei stollei, 74
Pipa aspersa, 263
pipa, 263, 265, 267
pipa, Pipa, 263, 265, 267
Population dynamics of mammals, 339-
386
Population stresses, 381
Procyon lotor, 367
propinqua, Synallaxis, 38
Pseudobranchus striatus striatus, 277
axanthus, 277
spheniscus, 277-288
Pseudocolaptes boissonneautii boisson-
neautii, 39
pulcherrimum, Steccherinum, 28
purpureum, Stereum, 246
purusianus, Automolus infuscatus, 40
quercinum, Radulum, 27
rabbittsi, [Penthestes hudsonicus], 334
radiata, Phlebia, 26
Radulum orbiculare, 27
quercinum, 27
rameale, Stereum, 246
Rana rugulosa, 268
Rattus norvegicus, 377
regeli, Karanasa, 326, 328, 329
regularis, Thelephora, 251
repandum, Dentium, 24
Reproductive potential, 381
rhea, Pierella, 71
rhea, Pierella rhea, 72
Rhinoderma darwinii, 263, 265, 266
Rhinophrynus, 263
ridgwa^d, Dendrocolaptes certhia, 6
rixosa, Mustela, 368
roseocarneum, Stereum, 248
1951
Index
393
rubeola, Pierella lima, 79
rubiginosa, Hyinenochaete, 241
riifesceiis, Dciidroplex picus, 11
rufescens, Marmota monax, 371
rufiim, Stereum, 248
rufus. Lynx, 371
rugiilosa, Rana, 268
safecla, Karanasa, 328
Salamander laruse (labial folds), 266-
267
Salamandra, 307
sanguinolentum, Stereum, 248
Satj'rus, 325, 326, 329
schmidti, Amb3’stoma, 299
Sciurus carolinensis leucotis, 372
Sclerurus rufigularis brunnescens, 42
rufigularis furfurosus, 41
vscrobiculatus, Calodon, 23
sericeum, Stereum, 248
setosa, Mycoacia, 26
Sittasomus amazonus sordidus, 15
griseicapillus viridior, 15
Skink, blue-tailed, 344
Smears of the testes, 381
Snails of West Virginia, The Land. In-
dex to this paper found follow-
ing plate 15, page 238
Sooglossus, 265
sordidus, Sittasomus amazonus, 15
Sorex cinereus cinereus, 362
fumeus fumeus, 364
palustris albibarbis, 365
Sparassis crispa, 242
spetentrionale, Steccherinum, 28
spheniscus, Pseudobranchus striatus,
277
Sphoenognathus curvipes, 45
spirurus, Gh'phor^mchus, 13
Staurois, 268
Steccherinum adustum, 27
ochraceum, 27
pulcherrimum, 28
septentrionale, 28
Stereum albobadium, 243
Burtianum, 243
Stereum diaphanum, 244
fasciaturn, 244
frustulosum, 245
fuscum, 245
gausapatum, 245
hirsutum, 246
M array i, 246
ochraceoflavum, 246
purpureum, 246
rameale, 246
roseocarneum, 248
rufum, 248
sanguinolentum, 248
sericeum, 248
subpileatum, 249
versiforme, 249
Stereum rufum, 248
stollei, Pierella, 73
stollei, Pierella stollei, 74
Stressors, 370
striatus, Pseudobranchus striatus, 277
strigosa-zonata, Phlebia, 26
strigosus, Gloiodon, 24
suboles, Philydor erythrocercus, 39
subpileatum, Stereum, 249
subrufescens, Gljqihorj’nchus spirurus,
14
successor, Campylorhamphus procur-
voides, 13
Synallaxis cinnamomea, 38
albescens griseonota, 37
albescens hypoleuca, 37
albescens littoralis, 36
cabanisi obscurior, 37
albescens perpallida, 36
propinqua, 38
Synaptom^’^s cooperi cooperi, 375
Sj^rrhophus, 267
tabacina, Hyinenochaete, 242
Tadpoles, 257-276
Tadpole operculum and spiracle, 263-
265, 268-269
talastauana, Karanasa, 328
talpoides, Blarina brevicauda, 366
Tamias striatus h^steri, 371
394
Annals of the Carnegie Museum
VOL. 31
Tamiasciurus hudsonicus loquax, 372
Taxodium ascendens, 280
Teeth of Ambystoma c. cingulatum, 304
Ambystoma c. bishopi, 304
Terpsiphone rufocinerea bannermani, 3
terrestris, Thelephora, 251
texanum, Ambystoma, 299
Thelephora albido-brunnea, 250
anthocephala, 250
caryophyllea, 250
intybacea, 250
multipartita, 250
palmata, 250
regularis, 251
terrestris, 251
vialis, 251
Thelephoraceae, 239
Todd, W. E. Clyde, Critical remarks on
the Wood-hewers, 5-18
Todd, W. E. Clyde, Critical remarks on
the Oven-birds, 33-43
Todd, W. E. Clyde, A northern race of
Red-tailed Hawk, 289-296
Todd, W. E. Clyde, A new race of
Hudsonian Chickadee, 333-334
Tomodactylus, 267
tornieri, Nectophrynoides, 257-262, 263,
266, 268
torridus, Furnarius, 33
triangularis, Xiphorhynchus triangu-
laris, 8
turugensis, Karanasa, 326
Twomey, Arthur C., A new race of
Paroquet of the species Aratinga
astec from the Republic of
Honduras, 297-298
underwoodii, Elvela, 285
ungava, [Penthestes hudsonicus], 333
Urocyon cinereoargenteus, 370
velutinus, Calodon, 23
Verpa bohemica, 284
versiforme, Stereum, 249
vialis, Thelephora, 251
vicinalis, Xiphorhjmchus eytoni, 7
viridior, Sittasomus griseicapillus, 15
vision, Mustela vision, 368
vivipara, Nectophrynoides, 257
Vulpes fulva, 370
wilkinsi, Karanasa, 323, 327
Wood, Albert E., A new geomyid ro-
dent from the Miocene of Mon-
tana, 335-338
Wood-hewers, Critical remarks on the,
by W. E. C. Todd, 5-18
Xiphorhynchus elegans ornatus, 10
eytoni vicinalis, 7
guttatus connectens, 8
necopinus, 10
obsoletus multiguttatus, 10
orenocensis orenocensis, 6
Xiphorhynchus orenocensis paraensis, 5
Xiphorhynchus pardalotus caurensis, 9
Xiphorhynchus triangularis distinctus,
8
Xiphorhynchus triangularis triangu-
laris, 8
Zachaenus parvulus, 265
Zapus hudsonius, 377
zibethica, Ondatra, 377
zonatus, Calodon, 23
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