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ras NEMOURIA

Occasional Papers of the Delaware Museum of Natural History
NuMBER 21 JuNE 3, 1979

A REVIEW OF GEOGRAPHIC VARIATION IN
CONTINENTAL POPULATIONS OF THE
RUBY-CROWNED KINGLET (Regulus calendula)

M. Ralph Browning *

INTRODUCTION

Four races of Regulus calendula Linnaeus are recognized in the A.O.U.
check-list (1957), including the nominate form (type locality: Philadelphia,
Pennsylvania), which breeds from northwestern Alaska south to central British
Columbia and eastward to eastern North America. The other three races that are
recognized occur in western North America, the first described by Ridgway
(1876) as obscurus (type locality: Guadalupe Island, Baja California), a small,
dark form found only at the type locality. This insular race is distinct from all
other populations of R. calendula and is not considered further in this paper.
The two other races are grinnelli Palmer, 1897 (type locality: Sitka, Alaska) and
cineraceus Grinnell, 1904 (type locality: Strain’s Camp, Mt. Wilson, Los Angeles
County, California). According to the check-list, the former occurs from
southern coastal Alaska to southwestern British Columbia, whereas the latter
occurs from south-central British Columbia eastward to Montana and the Rocky
Mountains and southward to Arizona and California.

The taxonomic status of the continental races has recently aroused discussion
by Phillips et al. (1964), Phillips (1964), and Hubbard and Crossin (1974), with
the status of the race cineraceus the subject of most of the controversy. Ridgway
(1904) was the first to examine the distinction of this form, and as the result he
synonymized cineraceus with nominate calendula. However, he commented that
the race was possibly valid, as he did not have critical specimens available for
comparison. Nonetheless, he listed specimens from California in his table of
measurements and states that the California series was similar to the eastern
specimens in size and color. Bishop (1926) later argued that cineraceus was
distinct, and in the subsequent A.O.U. check-list (1931) the name was accepted
as applicable to those western populations that occur elsewhere than the range
of grinnelli. Hellmayr (1934) also recognized cineraceus, remarking that the
name applied to the large, gray populations of the West.

*National Fish and Wildlife Laboratory Library of Congress
National Museum of Natural History Card No. 77-94139
Washington, D.C. 20560

Z NEMOURIA No. 21

Recently, Phillips (1964) synonymized cineraceus with calendula, while also
describing a new western race, arizonensis (type locality: vicinity of Phelps
Ranger Station, White Mountains, Arizona). Phillips gave the breeding range of
arizonensis as the mountains of Arizona and probably most of the boreal regions
of the western United States. Hubbard and Crossin (1974) followed Phillips’
treatment synonymizing cineraceus, but they questioned the validity of
arizonensis as being any more separable from the nominate form than cineraceus.
Pending a revision of the species, they suggested that all continental populations
except grinnelli be included in the nominate calendula. Because of this lack of
unanimity, I compared specimens in order to better assess the geographic variation
of the continental populations of Regulus calendula.

METHODS

Adult specimens of R. calendula were compared in coloration (N = 500),
wing chord (N = 270), and length of tail (N = 237). Measurements, all in
millimeters, were made of specimens collected on their breeding grounds, which
are here defined as areas of occurrence during the months of May and June,
depending on latitude. Specimens were segregated by age so as to exclude all
first-year birds. This was achieved by examination of crown color (in males),
shape of the rectrices (see Svenson, 1970), and data on the specimen labels. All
specimens that were doubtfully collected on their breeding grounds or were of
uncertain age were excluded.

Statistical comparisons include Student’s t-test and coefficient of difference
(C.D.), the latter a method to depict the percent of joint nonoverlap among
populations (cf. Mayr, 1969).

RESULTS AND DISCUSSION

The races of R. calendula have been characterized on the basis of size and
coloration. Palmer (1897) described grinnelli as sooty dark olive above, with the
area surrounding the crown patch blackish, the throat dusky gray, and the
abdomen yellowish. He described grinnelli as being darker and smaller than
calendula. Grinnell (1904) characterized cineraceus as being larger and grayer
than grinnelli and calendula. Phillips (Phillips et al. , 1964) stated that populations
of R. calendula are subject to a great amount of fading, and he found that in
fresh fall plumage, specimens of cineraceus and calendula are identical in colora-
tion. He concluded that only measurements showed populational differences in
the ranges attributed to these two forms.

The following analysis is my attempt to clarify conflicting statements on the
geographic variation of this species:

Wing chord. Comparisons of wing chord in my samples of R. calendula (Tables
1 and 2) demonstrate only minor differences among populations. In addition,

June 3, 1979 Browning 3

Table 1: Measurements of Wing Chord and Length of Tail of Adult Male
Specimens of Regulus calendula Collected During the Breeding Months

Wing chord Tail
Sample N Mean §S.D. Range N Mean S.D. Range

Alaska 10 59.8 1.4 56.8-61.1 5 47.5 1.3 45.3-48.3
NW Territory 4 58.2 56.1-60.6 4 46.7 43.0—50.5
N British Columbia 7 57.2 1.6 54.3-58.8 6 45.8 2.1 43.0—48.5
S British Columbia 10 59.8 1.3 57.6-61.3 10 45.1 1.5 42.3-47.4
Washington 13 59.6 1.3 57.5-62.6 12 46.1 0.8 44.9-—47.8
Oregon 21 60.0 0.9 57.8-61.6 14 466 1.1 44.4-—48.3
N California! 8 60.3 1.3 58.2-62.9 8 45.8 1.6 43.2—48.2
Central California” 7 60.3 1.0 58.4-61.1 7 46.8 0.8 45.5—47.5
S California? 7 60.2 0.8 59.3~61.5 7 46.0 1.1 44.1-47.9
California total 31 60.5 1.0 58.2-62.9 29 46.4 1.2 43.2-48.8
Idaho 15 60.2 1.0 58.1-61.8 15 46.8 1.3 44.2-—49.2
N Nevada‘ 13 60.5 1.3 58.8-63.2 13 47.6 1.7 45.1-—51.2
S Nevada® 10 60.2 1.6 57.4-62.8 10 47.5 2.4 44.3-51.4
Nevada total 28 60.4 1.3 57.4-63.2 28 47.6 1.9 44.8-51.4
“arizonensis”’ 7 61.6 0.6 60.4-—62.4 7 47.4 11.5 45.5—50.5
Arizona total® 13 60.9 1.1 58.5-62.4 11 46.4 2.5 40.8-48.5
N Alberta 4 57.3 55.6-—58.9 3 45.8 44,9-47.3
S Alberta 6 58.9 1.4 57.3-61.1 4 45.3 1.6 43.6—47.7
N U.S. Rocky Mtns. 4 59.8 57.3-62.1 2 43.6 43.0-44.1
Eastern N America 12 58.3 1.5 55.8-61.4 10 45.6 1.4 44.0-—47.8
grinnelli 14 56.2 1.2 54.1-58.3 14 44.3 1.1 42.0-46.4
1Modoc Co.

2Mono Co.

3San Bernardino and Riverside counties
*Elko and Humboldt counties

5Clark and Lincoln counties

®Includes “arizonensis”’

some of the samples of males from western North America demonstrate clinal
variation; e.g., specimens from Idaho average smaller than those from Arizona
(Fig. 1).

The means of several of the samples were compared using Student’s f-test.
The mean of the sample from eastern North America is significantly (P < 0.05)
smaller than those from Arizona (t = 4.74, P < 0.0001), Alaska (¢ = 2.32,
P < 0.02), and California (t = 5.26, P< 0.0001). However, the eastern sample is
not significantly smaller than those of birds from such areas as Oregon (t = 0.79,
P < 0.40). Student’s t-test of other populations revealed significant differences
between the means of the samples from northern British Columbia and those of
Alaska, between those of northern and southern British Columbia, and between
southern British Columbia and California. Results of t-tests between the means
of other samples are not significant.

4 NEMOURIA No. 21

Table 2: Measurements of Wing Chord and Length of Tail of Aduit Female
Specimens of Regulus calendula Collected During the Breeding Months

Wing chord Tail
Sample N Mean S.D. Range N Mean S.D. Range
Alaska 5 56.2 1.5 53.8-57.9 3 43.5 42.4-45.2
N British Columbia 3 55.3 54.5-55.8 3 43.7 42.1-45.3
Washington 3 56.3 55.5-56.9 3 43.6 43.0-—44.5
Oregon 5 55.6 2.0 52.5-56.8 2 43.8 43.5—44.1
California total 17 $7.1 1.3 $4.6-59.5 16 446 1.5 41.6-—46.7
Idaho 4 56.2 55.7-57.0 4 42.7 42.2-43.0
Nevada total 13 57.0 1.7 54.0-59.2 13 446 1.8 41.0-46.6
Arizona total 1 29681) 235.) 5 225-00.2 4 43.6 40.9-44.9
Alberta total 3 56.1 54.2-57.3 3 44.2 43.2-45.6
N U.S. Rocky Mtns. 3 55.7 54.0-58.2 2 41.4 41.0-41.9
Eastern N America 9 55.8 1.5 53.7-58.0 7 45.4 1.9 42.8-49.0
grinnelli 6 52.9 1.0 51.6-54.3 6 41.3 1.1 39.6—42.7

Although the differences between the mean wing chord in some samples of
the males are statistically significant, there is considerable overlap in measure-
ments between most samples (Table 1). The amount of joint nonoverlap between
several samples was determined with results that are below the level (90%)
suggested by Mayr (1969) for recognizing subspecific differences. This level is
equivalent to saying that less than 75% of the specimens of one sample can be
separated from 97% of another sample of specimens. To compute the C.D., I
purposely compared samples having extremely high or extremely low means in
order to gauge as rapidly as possible the prospects of separation. In all instances
the C.D. values fell below the 90% joint nonoverlap level, except in the case of
the samples compared with grinnelli.

The degree of variation in wing chord between samples of females (Table 2)
is even less than that between males (Table 1). Although the sample from
Arizona has a greater mean in wing chord than in all other samples, the range of
variation is so great as to include even the relatively short-winged eastern sample
(Table 2).

Because variation in wing chord could be related to altitude of breeding area,
I tested a sample of 54 males from Nevada and California for correlation of wing
chord with elevation. There is no correlation (r = 0).

My averages for wing chord of birds from eastern North America are greater
than those from New Hampshire reported by Sawyer (1961), especially for
males, for which our values differ significantly (t = 2.09, P <0.05). However,
comparable measurements reported by Phillips (1964) and Godfrey (1966) are

June 3, 1979 Browning 3)

ee ;
British Columbia
Northern Alberta — Seg er i i gags Naat anata omg =) sora Ss

Y )
en /
sonert S oe | |
British Columbia : .
Oregon a

ee ee Se
America

Figure 1: Comparison of wing chord measurements of adult male Regulus
calendula. Horizontal lines show the ranges. On each side of the mean, shown as
a vertical line, are one standard error, indicated by a solid rectangle, and one
standard deviation, indicated by an open rectangle. The number of specimens in
each sample is listed in Table 1.

similar to mine; e.g., the latter’s mean of 57.8 mm (range 55.0 to 61.1) for adult
male calendula (sensu A.O.U., 1957). My figures also generally agree with those
of Ridgway (1904) of eastern birds although his maxima for eastern specimens
exceed values for specimens measured by me. From the preceding, I conclude
that the sample from New Hampshire measured by Sawyer does not adequately
represent the range of variation of eastern populations.

6 NEMOURIA No. 21

Phillips (1964) gave the range in wing chord of arizonensis as 60.4 to 63.5
(x = 61.5) for summer males and “56.3+” to 58.8 for females. For eastern popula-
tions, Phillips reported maxima of 60.0 and 58.0 for males and females, respec-
tively. Phillips’ measurements of the wing chord of the holotype of cineraceus is
60.0, 0.4 mm less than the minimum in wing chord of his series of arizonensis.

In summary, the geographic variation in R. calendula in wing chord is slight.
The northwestern race, grinnelli, has a shorter wing than all other samples of the
species from the continental populations. Southwestern birds have slightly longer
wings than those in the other samples, but the range of individual variation
includes that of most other samples, including birds from eastern North America.
Based on wing chord, eastern and western populations, other than grinnelli,
cannot be separated at any meaningful subspecific level.

Tail. Comparisons of measurements of length of tail for males and females
(see Tables 1 and 2) do not reveal any significant geographic variation, and
measurements overlap considerably. Means in males demonstrate that birds
having the longest tails are from Alaska and the southwestern United States.
Specimens having the shortest tails are from the northern U.S. Rocky Mountains
and from eastern North America. Comparisons of my measurements with those
of Sawyer (1961) and Phillips (1964) reveal that the means for my samples of
males and females are, on the average, larger. These differences (possibly due to
technique) do not affect my conclusions because I measured all populations and
am not relying on the measurements of others.

Weight. A small number of weights taken from the specimen labels of birds
from western North America is presented in Table 3. Although these are too few
and too geographically scattered to be useful to compare for taxonomic discus-
sion, it is interesting that adult males taken during the breeding months have
weights similar to those of birds collected during the fall. Murray and Jehl
(1964) reported a mean weight of 5.8 grams (N = 66; sexes apparently pooled) for
birds from eastern North America; but birds from New Hampshire, according to
Sawyer (1961), averaged 7.19 and 6.74 grams for males and females, respectively.

Coloration. Plumage of breeding birds becomes extremely worn during
summer and, therefore, this character is useless for making comparisons of
coloration. Except for grinnelli, specimens in fresh plumage are identical in
dorsal and ventral coloration among continental populations. Having examined
over 500 specimens, many in fresh fall plumage, I agree with Phillips that the
populations of R. calendula cannot be separated on the basis of coloration.

CONCLUSIONS

Morphological variation in continental populations of R. calendula is slight,
with differences found in wing chord, length of tail, and plumage color among
some populations. Birds along the coastal regions of southern Alaska south to
southwestern British Columbia having significantly shorter wings are validly

June 3, 1979 Browning 7

Table 3: Weights (in grams) of Adult Specimens of Regulus calendula*

N Mean S.D. Range
S British Columbia 8 6.21 0.33 5.8-6.3
Oregon 13 6.07 0.51 5.2—7.2
California 6 6.23 0.72 5.7-7.5
Nevada 26 6.25 0.77 5.5—9.0
Nonbreeding adults 11 6.19 0.53 5.5—7.3

Females

Oregon 3 6.27 5.3-7.3
California 4 6.03 5.7-6.4
Nevada 16 6.23 0.68 5.2-—8.0

*All samples are from the breeding months unless otherwise indicated.
tFrom Nevada

separated as R. c. grinnelli Palmer. Specimens from the western United States
have longer wings than those of western British Columbia, but are similar to
birds from most of the remainder of Alaska and Canada and from the eastern
United States. The amount of individual variation among these populations
demonstrates extensive overlap; both large and small birds occur east and west of
the Rocky Mountains. Tail length generally parallels wing chord among popula-
tions but demonstrates even fewer differences than the latter. Plumage color is
uniform among continental populations, except that grinnelli is noticeably
darker than other birds.

Based on these data, exclusive of grinnelli, western populations, whether
ascribed to cineraceus (A.O.U., 1957) or to arizonensis (Phillips, 1964), are not
separable from populations of eastern North America. Thus, the names cineraceus
Grinnell, 1904 and arizonensis Phillips, 1964 are hereby considered synonyms of
the name calendula Linnaeus, 1766. The distinctive race grinnelli, the only other
separable continental race, breeds from southern Alaska to Vancouver Island.
Contrary to earlier literature, grinnelli is probably a resident, as my examination
of a large series of migrants collected as far south as southern California revealed
only nominate calendula (sensu lato) southward.

SPECIMENS EXAMINED

Specimens from the following localities and collections (see Acknowledg-
ments for abbreviations of museums) include birds collected on their breeding
grounds and some specimens collected during spring and fall months. Entire
series of specimens of nonbreeding birds from several collections (AMNH,
DMNH, MCZ, MVZ, and USNM) were examined for variation in color but are
not listed.

& NEMOURIA Now]

Alaska (USNM, 20; AMNH, 1; UAM, 1), Alberta (USNM, 3; MCZ, 2; UAMZ,
12), Arizona (USNM, 10; AMNH, 3; DMNH, 28), British Columbia (USNM, 5;
BCPM, 9; MCZ, 21), California (USNM, 13; MVZ, 48; AMNH, 3; MCZ, 1),
Colorado (AMNH 1;MCZ, 1), Idaho (USNM, 80), Labrador (USNM, 4), Mackenzie
(USNM, 5; MCZ, 1), Manitoba (USNM, 1), Montana (USNM, 2; MCZ, 1), Nevada
(USNM, 13; MVZ, 60), Newfoundland (USNM, 9), New Mexico (USNM, 6;
AMNH, 2; DMNH, 1; UMMZ, 2), Nova Scotia (USNM, 4), Oregon (USNM, 11;
MVZ, 16; CMNH, 2; DMNH, 1), Quebec (USNM, 4), Washington (USNM, 34;
BMWSM, 3; UMMZ, 1), Wyoming (USNM, 5; AMNH, 1).

Acknowledgments

For permitting me to examine specimens under their care, I am indebted to
the authorities of the following collections: American Museum of Natural
History (AMNH), British Columbia Provincial Museum (BCPM), Burke Memorial
Washington State Museum (BMWSM) of the University of Washington, Cleveland
Museum of Natural History (CMNH), Delaware Museum of Natural History
(DMNH), Museum of Comparative Zoology (MCZ), Museum of Vertebrate
Zoology (MVZ), United States National Museum of Natural History (USNM),
University of Alaska Museum (UAM), University of Alberta Museum of Zoology
(UAMZ), and University of Michigan Museum of Zoology (UMMZ). I wish to
thank J. P. Hubbard, A. R. Phillips, and G. E. Watson for discussion and sugges-
tions concerning this study. I thank R. C. Banks and J. P. Hubbard for reading
an earlier draft of this paper. Jennifer H. Maglidt typed the manuscript.

BIBLIOGRAPHY
American Ornithologists’ Union
1931 Check-list of North American birds, ed. 4. Amer. Ornith. Union,

Lancaster.
1957 Check-list of North American birds, ed. 5. Amer. Ornith. Union,
Baltimore.
Bishop, L. B.
1926 The distribution of the races of the Ruby-crowned Kinglet. Condor,
28:183.
Godfrey, W. E.

1966 The birds of Canada. Nat. Mus. Canada Bull. No. 203, pp. 1-428.
Grinnell, J.
1904 The Ashy Kinglet. Condor, 6:25.

June 3, 1979 Browning 9

Hellmayr, C. E.
1934 Catalogue of birds of the Americas. Field Mus. Nat. Hist., Zool. Ser.,
13, part 7.
Hubbard, J. P., and R. S. Crossin
1974 Notes on northern Mexican birds. Nemouria, 14:1-41.
Mayr, E.
1969 Principles of Systematic Zoology. McGraw-Hill Book Co., New
York.
Murray, B. G., and J. R. Jehl, Jr.
1964 Weights of autumn migrants from coastal New Jersey. Bird-Banding,
35:253-263.
Palmer, W.
1897 The Sitkan Kinglet. Auk, 14:399-401.
Phillips, A. R.
1964 #Notas sistematicas sobre aves Mexicanas, III. Rev. Soc. Mexico Hist.
Nat., 25:217-242.
Phillips, A. R., J. T. Marshall, Jr., and G. Monson.
1964 The Birds of Arizona. Univ. Arizona Press, Tucson.
Ridgway, R.
1876 Ornithology of Guadeloupe Island, based on notes and collections
made by Dr. Edward Palmer. Bull. Geol. Geogr. Surv. Terr., 2(2):

183-195.
1904 The birds of North and Middle America, part 3. U. S. Nat. Mus.
Bull., 50.
Sawyer, P. J.

1961 Report on the cause of mortality and the morphometry of seventy
Ruby-crowned Kinglets killed at the WENH-TV tower in Deerfield,
New Hampshire. Bird-Banding, 32:162-168.
Svenson, L.
1970 Identification Guide to European Passerines. Naturhistoriska
Riksmuseet, Stockholm.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
Kennett Pike, Greenville,
Delaware 19807

a=
|

th NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 22 OcTOBER 30, 1979

REVIEW OF THE SUBGENUS
PTERYNOTUS (GASTROPODA: MURICIDAE)
IN THE WESTERN ATLANTIC

M. G. Harasewych* and R. H. Jensen**

INTRODUCTION

The genus Pterynotus represents one of the oldest of muricine lineages,
dating back to at least the Paleocene (Vokes, 1964, p. 15; 1970, p. 2). By the
end of the Eocene, the genus had achieved a Tethyan distribution, with species
ranging to Peru and New Zealand. However, it was in the warm, shallow Eocene
seas of western Europe (Paris Basin) that a major radiation, which produced
most of the lineages recognized today, occurred. Some of these lineages have
been given generic or subgeneric status (Pterochelus, Purpurellus, Marchia), while
others, although just as distinct, have been retained in Pterynotus s.s. In this
paper, the supraspecific taxonomy of Vokes (1971, p. 53) is followed.

In a review of the Cenozoic Pterynotus of the western Atlantic region, Vokes
(1970, pp. 1-17) reported that the subgenus Pterochelus first appears in the
lower Oligocene of the area and is represented by a single Recent species, which
is known only from the holotype; that the subgenus Purpurellus is presently
known only from a single early Miocene species; and that the subgenus Pteryno-
tus has been continuously represented since the Paleocene, with three deep-water
species in the Recent fauna. These Recent species have since been synonymized
by some workers (Abbott, 1974, p. 176; Radwin and D’Attilio, 1976, p. 100)
but retained as distinct by others (Fair, 1976, pp. 28, 47, 67). Additionally, a
“lost” species of shallow-water Pterynotus, erroneously believed for many years
to come from the Indian Ocean, has been rediscovered in the Caribbean (Emer-
son and Old, 1972, pp. 350-354).

*Biochemical Systematics Laboratory Library of Congress
Delaware Museum of Natural History Card No. 79-90857
**Department of Mollusks
Delaware Museum of Natural History

p. NEMOURIA No. 22

In addition to the types, we were able to examine more than a hundred spec-
imens of western Atlantic Prerynotus, some with soft parts partially preserved.
Based on this material, a revision of the existing species is proposed; three new
species are described; and the anatomy, ecology, and evolution of these species
are discussed.

Abbreviations used within the text are as follows: AMNH, American Museum
of Natural History; DMNH, Delaware Museum of Natural History; MCZ, Mu-
seum of Comparative Zoology; MHNG, Museum d’Histoire Naturelle, Geneva;
USNM, United States National Museum.

Acknowledgments

We are grateful to the following people, who have assisted us with informa-
tion, loans of types, and other material; K. J. Boss, Museum of Comparative
Zoology, Harvard; W. K. Emerson, American Museum of Natural History, New
York; R. S. Houbrick, United States National Museum, Washington, D.C.; G. L.
Voss, Rosenstiel School of Marine and Atmospheric Sciences, University of
Miami; C. J. Finlay; A. T. Guest; R. and D. Janowsky; J. R. H. Lightbourn;
B. McCormack; E. J. Petuch; A. and T. Van Landingham.

For critical reading of the manuscript we thank A. D’Attilio, W. K. Emerson,
E.H. Vokes, E. J. Petuch, and D. J. Bereza.

We also thank G. Kris Jensen for illustrating the protoconchs.

SYSTEMATICS
Family MURICIDAE
Genus Pterynotus
Subgenus Pterynotus s.s.

TYPE SPECIES: Murex pinnatus Swainson, 1822 (is Purpura alata Roding, 1798)
by subsequent designation, Swainson, 1833.

The Recent species presently assigned to the subgenus Pterynotus span at
least four distinct lineages, three of which are represented in the western Atlan-
tic. Although we are not presently advocating recognition of these lineages as
subgenera, we feel that they do provide valuable insights into the evolution of
the genus. These lineages will therefore be treated separately.

The first is the primitive Pterynotus line, which has remained virtually un-
changed since the Paleocene. It is characterized by a tabled shoulder that leads
to a channelled posterior spine on the varix. The aperture does not open into
this channelled spine. During the middle Eocene, this group gave rise to the sub-
genus Pterochelus, in which the aperture does open into the channelled spine
(Vokes, 1971, p. 42). Ponder and Wilson (1973, p. 395) have raised some ques-
tions as to the validity of Pterochelus as a subgenus.

The ancient Pterynotus line is represented in the Recent fauna by two relict,
deep-water species, P. vespertilio (Kira, 1959) from off southeastern Japan and
P. guestin. sp. from the Straits of Florida.

October 30, 1979 Harasewych and Jensen 3

Pterynotus (Pterynotus) guesti new species

(Figures 6, 13, 14)
DESCRIPTION: Shell of moderate size (to 28 mm), fusiform, thin; spire angle
40-42°; protoconch with one and a half whorls, smooth, conical, light tan color;
juncture with teleoconch poorly delineated, chiefly by change in surface sculp-
ture; six post-nuclear whorls; first post-nuclear whorl with two nodes; first varix
below protoconch-teleoconch juncture, adherent to protoconch; subsequently,
three varices per whorl with single intervarical nodes on the first three or four
post-nuclear whorls; thereafter, no axial sculpture between varices; spiral sculp-
ture consisting of major spiral cord at shoulder, giving sheli a tabulate appear-
ance, and six minor spiral cords below shoulder, producing spines on winglike
varices; major cord producing large, channelled shoulder spine; ventral surface of
varices lamellate; aperture oval; inner lip smooth, attached posteriorly; outer lip
with five to seven denticles; anal sulcus not opening into channelled shoulder
spine; siphonal canal moderate in length, open, slightly recurved dorsally; shell
color light tan with reddish brown blotches, especially on and around varices;
aperture reddish brown; operculum, periostracum, and soft parts unknown.
TYPE SPECIMENS: Holotype—DMNH 122258, length 28.6 mm; paratype—USNM
783318, length 23.4 mm (broken canal).

TYPE LOCALITY: ESE of Key West, Florida, in 275 meters.
RANGE: Known only from the type locality.
MATERIAL EXAMINED: The two specimens in type lot.

DISCUSSION: This new species very closely resembles the Paleocene Pterynotus
mathewsensis (Aldrich, 1886) (Vokes, 1970, pl. 1, fig. 1a,b) and the Eocene P.
stenzeli E. H. Vokes, 1970 (Vokes, 1970, pl. 1, fig. 5a,b). It may be distinguished
from all other Recent Atlantic species by the presence of a tabled shoulder and
a channelled posterior spine. That P. guesti is very similar to the Japanese P.
vespertilio (Kira) (Radwin and D’Attilio, 1976, pl. 9, fig. 2) is not surprising
since there has been so little morphological change in this line since the Paleocene.
Pterynotus guesti differs from P. vespertilio in having a more elongate shell with
narrower varical webbing and by the presence of reddish brown color inside the
aperture. This new species honors Arthur T. Guest of Bailey’s Bay, Bermuda, in
recognition of his assistance in our work on the mollusks of Bermuda.

A second, slightly modified lineage can be traced to P. tripteroides (Lamarck,
1822) of the Paris Basin Eocene. This species group, which contains P. phaneus
(Dall, 1889) and P. leucas (Locard, 1897) as well as the two new species described
in the text that follows, may be recognized as having thin, delicate shells that
lack scabrous surface ornament and are not buttressed on the apertural side of

4 NEMOURIA No. 22

the varices. The subgeneric name Jimbellus (Type: M. latifolius Bellardi) was
proposed for this group by de Gregorio (1885, p. 275). At present, we feel it is
advisable to retain this group in the nominate subgenus of Pterynotus. All Recent
species of this lineage, which first evolved in warm, shallow seas, are members
of upper continental slope communities.

Pterynotus (Pterynotus) phaneus (Dall, 1889)
(Figures 1-5, 13, 16, 17)

SYNONYMY

1889 Murex (Pteronotus) phaneus Dall, Bull. Mus. Comp. Zool. Harvard,
18:201; 1889, Dall, Bull. U. S. Natl. Mus., 37:120, pl. 42, fig. 1;
1890, Dall, Proc. U. S. Natl. Mus., 12:330, pl. 11, fig. 1; 1934, John-
son, Proc. Bost. Soc. Nat. Hist., 40(1):115.

1889 Murex (Pteronotus) tristichus Dall, Bull. Mus. Comp. Zool. Harvard,
18:202, pl. 15, fig. 3; 1889, Dall, Bull. U. S. Natl. Mus., 37:120,
pl kd, fis 3:

1893 Murex (Pteronotus) pygmaeus Bush, Bull. Mus. Comp. Zool. Harvard,
23:213, pl. 1, figs. 3, 4; 1934, Johnson, Proc. Bost. Soc. Nat. Hist.,
40(1):115.

1927 Pteropurpura tristica (Dall), Dall, Proc. U. S. Natl. Mus., 70(2667):58.

1934 Murex (Pteropurpura) tristichus Dall, Johnson, Proc. Bost. Soc. Nat.
Hist., 40(1):115.

1945 Murex (Pterynotus) tristichus Dall, Clench and Perez Farfante, John-
sonia, 1(17):36, pl. 20, figs. 1—4.

1945 Murex (Pterynotus) pygmaeus Bush, Clench and Pérez Farfante, John-
sonia, 1(17):36, pl. 20, figs. 7, 8.

1945 Murex (Pterynotus) phaneus Dall, Clench and Perez Farfante, John-
sonia, 1(17):37.

1964 Pterynotus (Pterynotus) tristichus (Dall), E. H. Vokes, Malacologia,
21): Se

1970 Pterynotus (Pterynotus) havanensis E. H. Vokes, Tulane Stud. Geol.
Paleont., 8(1):13, pl. 3, figs. la,b, new name for Murex tristichus
Dall non Beyrich; 1976, Fair, The Murex Book, p. 47, pl. 13, fig. 164.

Figures 1-5: Pterynotus (Pterynotus) phaneus (Dall, 1889).

Figure 1: Holotype — Murex pygmaeus Bush, MCZ 6918, Blake Station 319,
off Charleston, South Carolina, in 479 m (X2).

Figure 2: Holotype — Murex phaneus Dall, USNM 93256, Albatross Station
2662, off St. Augustine, Florida, in 794 m (X2).

Figure 3: Holotype — Murex tristichus Dall, MCZ 7308, Blake Station 51,
off Havana, Cuba, in 732 m (X2).

Figure 4: USNM 416663, off Key West, Florida, in 174 m (X2).

Figure 5: DMNH 122423, R/V Pillsbury Station P-587, Arrowsmith Banks,
off Isla Mujeres, Quintana Roo, Mexico, in 245-250 m (X2).

Figure 6: Pterynotus (Pterynotus) guesti new species. Holotype, DMNH
122258, ESE of Key West, Florida, in 275 m (X2).

October 30, 1979 Harasewych and Jensen

6 NEMOURIA No. 22

1970 Pterynotus (Pterynotus) bushae E. H. Vokes, Tulane Stud. Geol.
Paleont., 8(1):13-14, pl. 3, figs. 2a,b, new name for Murex pygmaeus
Brush non Muricites pygmaeus Schlotheim;, 1976, Fair, The Murex
Book, p. 28, pl. 13, fig. 163.

1970 Pterynotus (Pterynotus) phaneus (Dall), E. H. Vokes, Tulane Stud.
Geol. Paleont., 8(1):14-15, pl. 3, figs. 3a,b; 1974, Abbott, American
Seashells, second edition, p. 175, fig. 1856; 1976, Fair, The Murex
Book, p. 67, pl. 13, fig. 165; 1976, Radwin and D’Attilio, Murex
Shells of the World, p. 100, pl. 9, fig. 3.

DESCRIPTION: Shell small (to 20 mm), fusiform, thin; spire angle 33-44° ; proto-
conch with one and a half whorls, smooth, translucent, slightly bulbous, ending
in thin varix; teleoconch with five slightly convex whorls; three thin, flaring
winglike varices per whorl; axial sculpture consisting of none, one, two, or three
intervarical nodes (number may vary on single specimen); spiral sculpture of
fine threads, barely discernible, except on dorsal surfaces of varices where four
to nine raised ribs may be produced; laminae on ventral surfaces of varices pro-
nounced to barely noticeable; aperture oval; inner lip smooth, attached poste-
riorly; outer lip may be smooth or may bear four to six denticles; siphonal canal
moderate in length, open, slightly recurved dorsally; shell and aperture color
uniformly white; periostracum very thin, light yellowish brown, generally
abraded away; operculum corneous, light amber, elongate, with terminal nucleus.

SOFT PARTS: A single, partially preserved female specimen (see Figure 5) was
examined; animal uniformly cream-yellow; mantle edge smooth; osphradium un-
equal, right side 1.5 times as wide as left; about 35 leaflets per side of osphradium;
ctenidum of about 85 leaflets tapers anteriorly and posteriorly; large, purplish
hypobranchial gland; capsule gland enlarged, suggesting specimen was ready to
spawn; radular ribbon long and narrow, consisting of 142 rows; rachidian five
cusped, laterals scythe shaped (see Figure 17); ratio of radular length to shell
length 0.136; large valve of Leiblein; paired salivary glands; large, amber-colored
gland of Leiblein.

TYPE SPECIMENS: P. phaneus: holotype—USNM 93256, length 16.9 mm. P.
tristichus: holotype—MCZ 7308, length 15.3 mm; paratype 1—-MCZ 7309, length
7.0 mm; paratype 2—USNM 87084, length 18.5 mm. P. bushae: holotype—MCZ
6918, length 16.0 mm.

Figures 7-9: Pterynotus (Pterynotus) lightbourni new species.

Figure 7: Holotype, DMNH 122259, off St. David’s, Bermuda, in 275 m (X2).

Figure 8: Paratype 2, A. T. Guest collection, off St. David’s, Bermuda, in
275 m (X2).

Figure 9: Paratype 1, USNM 652787, off St. David’s, Bermuda, in 600 m
(X2).

October 30, 1979 Harasewych and Jensen oi

8 NEMOURIA No. 22

TYPE LOCALITY: Albatross Station 2662, off St. Augustine, Florida, in 434
fathoms (794 meters).

RANGE: Outer continental shelf and upper continental slope from off Charleston,
South Carolina, down the east coast of Florida, along both sides of the Straits
of Florida to off the Arrowsmith Banks, Yucatan, Mexico, in 165 to 800 meters.

MATERIAL EXAMINED: Off Charleston, South Carolina, 479 m depth (type
locality of P. bushae), MCZ; off Brunswick, Georgia, 485-530 m depth, MCZ;
off Fernandina, Florida, 794 m depth, USNM; off Key West, Florida (13 speci-
mens, 174-274 m depth), AMNH, DMNH, USNM, Finlay coll., Janowsky coll.;
off Sand Key, Florida (4 specimens, 165-232 m depth), USNM, Janowsky coll.;
off Sambo Reef, Florida (22 specimens, 216-247 m depth), USNM; off West’n
Dry Rocks, Florida, 174 m depth, USNM; off Dry Tortugas, Florida, 192 m
depth, Janowsky coll., Lightbourn coll.; off Havana, Cuba (type locality of P.
havanensis), (5 specimens, 320-823 m depth), MCZ, USNM; off Arrowsmith
Banks, Yucatan, Mexico, 245-250 m depth, DMNH.

DiscussION: P. phaneus is the most widely distributed and variable species in
this lineage. Our examination of 50 specimens supports the conclusion that P.
havanensis and P. bushae are forms of P. phaneus. The chief characters used to
distinguish these forms had been the presence or absence of three intervarical
nodes and the relative degree of varical webbing. We have found varical webbing
to be variable. The number of intervarical nodes can range from none to three
on a single specimen! Specimens from both the northern and southern ends of
the range have heavier shells, greater spire angles, and a higher incidence of inter-
varical nodes.

In addition to the type specimens of Murex tristichus discussed by Vokes
(1970, p. 13), a second paratype from Blake Station 5, which was referred to by
Dall (1889, p. 202), has been located at USNM.

Pterynotus (Pterynotus) lightbourni new species

(Figures 7-9, 13, 15)
DESCRIPTION: Shell large (to 39 mm), fusiform, delicate; spire angle 35-37";
protoconch with one and a half whorls, conical, glassy, translucent, ending in a
thin varix; teleoconch with seven slightly convex whorls; first 11/3 whorls bear

Figure 10: Pterynotus (Pterynotus) radwini new species. Holotype, DMNH
122424, R/V Pillsbury Station P-610, east of Turneffe Islands, Belize, in 296-329
m (X2).

Figure 11: Pterynotus (Pterynotus) phyllopterus Lamarck, 1822, Finlay col-
lection, off Malendure Beach, Guadeloupe, in 6 m (X1).

Figure 12: Pterynotus (Perynotus) phyllopterus Lamarck, 1822, Van Land-
ingham collection, off Guadeloupe in 10-12 m (X1).

October 30, 1979 Harasewych and Jensen

10 NEMOURIA No. 22

BERMUDA
a

lightbourni

GULF OF MEXICO
BAHAMAS

vo
Qxy
Xy

o 6
BARBADOS
q

Figure 13: Geographical distribution of species assigned to Pterynotus s:.s.
in the western Atlantic.

six varices per whorl; subsequently, three varices per whorl with single, prominent
intervarical node between varices; spiral sculpture of 10 to 12 faint cords, pro-
ducing thin ribs on dorsal surfaces of wide, thin varical webs; ventral surfaces of
webs faintly lamellate; aperture oval; inner lip smooth, appressed posteriorly;
outer lip may be smooth or may bear prominent anal sulcus and six denticles;
siphonal canal long, open, curved to animal’s right, but not recurved dorsally;
shell and aperture color white, except for three faint golden tan patches on out-
side of aperture, one at edge of siphon, one medial, and one at posterior margin;
operculum, periostracum, and soft parts unknown.

TYPE SPECIMENS: Holotype—DMNH 122259, length 33.8 mm; paratype 1—USNM
652787, length 38.6 mm; paratype 2—Arthur Guest collection, length 37.0 mm.

TYPE LOCALITY: Off St. David’s, Bermuda, in 275-600 meters.
RANGE: Known only from the type locality.
MATERIAL EXAMINED: The three specimens in type lot.

DISCUSSION: The three apertural color patches, as well as shell size and shape,
serve to distinguish this species from other western Atlantic taxa. It resembles
P. leucas from the eastern Atlantic but differs in having a proportionally smaller
aperture, wider varices, and a single intervarical node. It is also similar to Recent

October 30, 1979 Harasewych and Jensen 11

specimens of P. flemingi Bue, 1967 (Beu, 1970, pl. 2, figs. 23-26) from off New
Zealand. Pterynotus flemingi also has the color patches outside the aperture, as
well as a single intervarical node, but differs from P. lightbourni by having a
much larger aperture and a greater spire angle.

We take pleasure in naming this species in honor of John R. H. Lightbourn,
of Bailey’s Bay, Bermuda, in recognition of his contributions to the studies of the
deep-water mollusks of Bermuda.

Pterynotus (Pterynotus) radwini new species
(Figures 10, 13, 18)

DESCRIPTION: Shell large (31 mm), fusiform, thin; spire angle 33°; protoconch
unknown; adult shell of six slightly convex post-nuclear whorls; three varices
per whorl; no axial sculpture between varices; spiral sculpture of eight equally
prominent cords, producing raised ribs on dorsal surfaces of webbed varices,
these raised ribs projecting as spines beyond edges of varices; ventral surfaces of
varices complexly lamellate; aperture oval; inner lip smooth, attached posteriorly;
outer lip with pronounced anal sulcus and six denticles; siphonal canal long,
open, curved to animal’s right, but not recurved dorsally; shell color pale coffee,
cords and ribs slightly darker; aperture white; periostracum thin, yellowish, but
generally abraded away; operculum comeous, light amber, elongate, with ter-
minal nucleus.

SOFT PARTS: Holotype, a partially preserved female specimen; animal uniformly
cream-yellow; mantle edge smooth; osphradium roughly symmetrical, with 68
leaflets per side; ctenidium of 90 branchial leaflets tapers posteriorly but remains
broad anteriorly; large purplish hypobranchial gland; capsule gland not enlarged;
radula of 110 rows with five cusped rachidian, sicklelike laterals (see Figure 18);
ratio of radular length to shell length of 0.051; pronounced valve of Leiblein and
gland of Leiblein, salivary glands paired.

TYPE SPECIMEN: Holotype—DMNH 122424, length 30.9 mm.

TYPE LOCALITY: R/V Pillsbury station P-610, east of Turneffe Islands, Belize,
in 296-329 meters.

RANGE: Known only from the type locality.
MATERIAL EXAMINED: The holotype.

Discussion: At first glance, this species more closely resembles the ocenebrine
Pteropurpura macroptera (Deshayes, 1839) than any Pterynotus, but closer exa-
mination reveals typically muricine radula and operculum. It can readily be dis-
tinguished from P. lightbourni, its nearest congener, by its coffee color, lack of
apertural color patches, lack of axial sculpture, prominent spiral sculpture, and
strongly digitated varices.

This species is named in honor of George E. Radwin, who would have des-
cribed it were it not for his untimely death.

12 NEMOURIA No. 22

A third lineage is represented in the western Atlantic by P. phyllopterus
(Lamarck, 1822). This line has been traced to the middle Eocene of the Paris
Basin by Vokes (1970, p. 13) and is characterized by its large size (60-100
mm), crenulated varices, flanged outer lip with pronounced denticles, and pres-
ence of medial and siphonal color patches on the varices. This shallow-water
group reached its peak in the Miocene faunas of Europe and North America
but is represented in the Recent fauna by two relict species, P. bednalli (Brazier,
1878) of northwestern Australia and P. phyllopterus of the Antilles.

Pterynotus (Pterynotus) phyllopterus (Lamarck, 1822)
(Figures 11-13, 19)
SYNONYMY

1822 Murex phyllopterus Lamarck, Anim. sans Vert., 7:164; 1829, Schubert
and Wagner, Martini and Chemnitz, Neues Systematisches Conchylien-
Cabinet, 12:19, pl. 219, figs. 3042, 3043; 1842, Kiener, Coq. Viv.,
7:103~-104, pl. 24, fig. 1; 1843, Deshayes and Milne-Edwards, Anim.
sans Vert., second edition, 9:577 (in part, references to Schubert and
Wagner, 1829, and Kiener, 1842, only); 1856, Kuster and Kobelt,
Systematisches Conchylien-Cabinet von Martini and Chemnitz,
3(2):43—44, pl. 18, figs. 1, 2; 1880, Tryon, Manual of Conchology,
2:114, pl. 34, fig. 373 only; 1883, Poirier, Nouv. Arch. Mus. Hist.
Nat., 10:97; 1971, Cernohorsky, The Veliger, 14:189, fig. 5.

1846 Murex rubridentatus Reeve, Conch. Icon., vol. 3, Murex, pl. 36, figs.
186a,b; 1879, Sowerby, Thes. Conch., 4:22, pl. 11, fig. 105; 1880,
Tryon, Manual of Conchology, 2:85, pl. 40, fig. 507; 1883, Poirier,
Nouv. Arch. Mus. Hist. Nat., 10:96; 1971, Cernohorsky, The Veliger,
14:189, fig. 8.

1970 Pterynotus (Pterynotus) phyllopterus (Lamarck). E. H. Vokes, Tulane
Stud. Geol. Paleont., 8:13; 1971, Emerson and Old, The Veliger,
14:350-354, figs. 1-13; 1974, Abbott, American Seashells, second
edition, p. 176, pl. 8, fig. 1862; 1976, Fair, The Murex Book, pp. 67-
68, pl. 13, fig. 157; 1976, Radwin and D’ Attilio, Murex Shells of the
World, pp. 100-101, pl. 7, fig. 8; 1977, Lozet, Shells of the Carib-
bean, p. 73, fig. 88.

Not Murex phyllopterus ““Lamarck” of authors: 1828, Sowerby, The Genera
of Recent and Fossil Shells, 2(30), pl. 224 [is Ceratostoma nuttalli

Figure 14: Protoconch of Pterynotus (Pterynotus) guesti new species. Holo-
type, DMNH 122258.

Figure 15: Protoconch of Pterynotus (Pterynotus) lightbourni new species.
Paratype 1, USNM 652787.

Figure 16: Protoconch of Pterynotus (Pterynotus) phaneus (Dall, 1889), off
Key West, Florida, in 274 m, DMNH 124417. Scale bar equals 1 millimeter for
all drawings.

13

Harasewych and Jensen

October 30, 1979

14 NEMOURIA No. 22

(Conrad, 1837)]; 1845, Reeve, Conch. Icon., vol. 3, Murex, pl. 16, fig.
63 [is Pteropurpura trialata (Sowerby, 1841)]; 1879, Sowerby, Thes.
Conch., 4(33-34):24, pl. 11, fig. 107 [is Pteropurpura trialata (So-
werby, 1841)].

DESCRIPTION: Shell large (to 100 mm), fusiform, moderately heavy; spire angle
40-42°; protoconch of one and a half whorls, smooth, conical; teleoconch with
eight slightly convex whorls; three crenulated, flaring varices per whorl; two pro-
nounced intervarical nodes between varices; spiral sculpture of nine to 12 cords,
extending onto varices, producing dorsal fluting; ventral surfaces of varices
lamellate; aperture lenticular; inner lip smooth, attached posteriorly; outer lip
flanged, appressed to ventral surface of varix; six to eight denticles on outer lip;
siphonal canal straight, open, slightly recurved dorsally; shell color typically tan,
but may be brick-red, white, yellow, lavender, or chocolate-brown; two brown
patches on outside of aperture, one at edge of siphon, one medial; aperture
white; denticles purplish red; periostracum thin, brownish; operculum corneous,
brown, elongate, with terminal nucleus; soft parts unknown except for radula (see
Figure 19), which has five cusped rachidian and scythelike laterals (D’ Attilio,
personal communication).

TYPE SPECIMEN: Holotype—MHNG 1099/27, length 83.4 mm.

TYPE LOCALITY: Martinique, Windward Islands (subsequent designation, Emer-
son and Old, 1972).

RANGE: Known only from the islands of Martinique and Guadeloupe, in 6 to
30 meters. Sand and coral rubble bottom.

MATERIAL EXAMINED: Off Guadeloupe (8 specimens, 6-20 m depth), Finlay
coll., Janowsky coll., Van Landingham coll.; off Martinique (7 specimens, 15-
30 m depth), AMNH, DMNH, Janowsky coll.

DISCUSSION: Although this is one of the largest species of Pterynotus, it has only
recently been rediscovered in the Caribbean, previously having been believed to
come from the Indian Ocean. This relict species more closely resembles the Mio-
cene species P. hoerlei E. H. Vokes, 1970, of Florida and P. rovasendae (Bellardi,
1872) of Italy than any Recent species.

Western Atlantic taxa erroneously referred to Pterynotus:

Murex (Pteronotus) macropterus “Deshayes”’ Dall, 1889, Bull. Mus. Comp.
Zool. Harvard, 18:201; Dall, 1889, Bull. U. S. Natl. Mus., 37:120 is
Pteropurpura bequaerti Clench and Perez Farfante, 1945 (Clench and
Perez Farfante, 1945, p. 40).

Murex abyssicola Crosse, 1865, had been placed in Pterynotus by Clench and
Perez Farfante, 1945, p. 38 and reassigned to Dermomurex by Vokes,
1970, p. 15; 1975, p. 148, pl. 4, figs. 2, 3.

October 30, 1979 Harasewych and Jensen 15

Figure 17: Scanning electron micrograph of radula of Pterynotus (Pterynotus)
phaneus (Dall, 1889), taken from specimen shown in Figure 5. Scale bar equals
20 pm.

Figure 18: Scanning electron micrograph of radula of Pterynotus (Pterynotus)
radwini new species. Holotype. Scale bar equals 20 um.

Figure 19: Drawing of radula of Pterynotus (Pterynotus) phyllopterus (La-
marck, 1822) (courtesy A. D’Attilio).

16 NEMOURIA No. 22

BIBLIOGRAPHY
Abbott, R. T.
1974 American Seashells, second edition. Van Nostrand Reinhold Com-
pany, N.Y.
Beu, A. G.

1970 New Zealand Gastropod Molluscs of the Genus Pteropurpura Jous-
seaume. Trans. R. Soc. N.Z., Biol. Sci., 12(12):133-143.

Clench, W. J., and I. Perez Farfante
1945 The Genus Murex in the western Atlantic. Johnsonia, 1(17):1-56.

Dall, W. H.
1889 Reports on the Mollusca, Part II — Gastropoda and Scaphopoda: Re-
ports on the results of dredging, under the supervision of Alexander
Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea
(1879-80), by the U.S. Coast Survey Steamer “Blake,” Lieut.-Com-
mander C. D. Sigsbee, U.S.N., and Commander J. R. Bartlett, U.S.N.,
Commanding. Bull. Mus. Comp. Zool. Harvard, 18:1-492.

Emerson, W. K., and W. E. Old
1972 On the identity of Murex phyllopterus Lamarck, 1822, a tropical
western Atlantic species (Gastropoda: Prosobranchia). The Veliger,
14(4):350-354.
Fair, R. H.
1976 The Murex Book: an illustrated catalogue of the recent Muricidae
(Muricinae, Muricopsinae, Ocenebrinae). Ruth H. Fair, Honolulu.
de Gregorio, A.
1885 Studi su talune conchiglie mediterranee viventi e fossili con una revista
del gen. Vulsella. Bull. Soc. Malac. Ital., 10:36-288.
Ponder, W. F., and B. R. Wilson
1973 A new species of Pterynotus (Gastropoda: Muricidae) from western
Australia. J. Malac. Soc. Aust., 2(4):395-400.

Radwin, G. E., and A. D’Attilio
1976 Murex Shells of the World: An illustrated Guide to the Muricidae.
Stanford University Press, Stanford, California.

Vokes, E. H.

1964 Supraspecific groups in the subfamilies Muricinae and Tritonaliinae
(Gastropoda: Muricidae). Malacologia, 2(1):1-41.

1970 Cenozoic Muricidae of the western Atlantic region, Part V — Pterynotus
and Poirieria. Tulane Stud. Geol. Paleont., 8(1):1-50.

1971 The geologic history of the Muricinae and the Ocenebrinae. The Echo,
4:37-54.

1975 Cenozoic Muricidae of the western Atlantic region, Part VI — Aspella
and Dermomurex. Tulane Stud. Geol. Paleont., 11(3):121-162.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
Kennett Pike, Greenville,
Delaware 19807

i
bi rds
NEMOURIA
Occaszonal Papers of the Delaware Museum of Natural History

NUMBER 23 DECEMBER 30, 1979

TWELVE NEW INDO-PACIFIC
GASTROPODS

Edward J. Petuch*

INTRODUCTION

This paper is a compilation of descriptions of new gastropod mollusks from
several subregions of the Indo-Pacific Faunal Province. The new ‘species were
collected in a variety of manners in the various areas: drag-netting by com-
mercial shell collectors in the Central Philippines, dredging by commercial shell
collectors off the Solomon Islands, SCUBA diving by commercial shell collectors
in the Red Sea, trawling by Taiwanese fishing boats off Southeast Africa, and
trawling by exploratory fisheries boats in the China Strait off eastern New
Guinea.

Most of the new species described in the paper are from the upper continen-
tal slope fauna of the Central Philippines; they give mute testament to the fact
that we are still in the “Descriptive Age of Malacology” (Rosewater, 1976, p. 5).
As more exploratory work is done on the lower shelf and upper slope communi-
ties around the world, doubtless many hundreds of new species of mollusks will
be discovered. This is especially apparent when one considers that the slope and
lower shelf areas were virtually unaffected by Plio-Pleistocene sea level fluctua-
tions and temperature changes, and offered stable ecological conditions for
diversification of post-Tethyan faunas. This long-lived stability allowed the
faunas to become partitioned into many more ecological niches than were pos-
sible in the unstable shelf waters. A tiny part of this tremendous unstudied
diversity is seen in the gastropod species described herein.

Ackowledgments

I would like to thank Messrs. Richard M. Kurz, of Wauwatosa, Wisconsin,
and Robert Morrison and John Bernard, of Sarasota, Florida, for supplying a
great part of the type material. Special thanks are given to Messrs. Russell Jensen
and M. G. Harasewych, Delaware Museum of Natural History, and Dr. Gilbert
L. Voss, Rosenstiel School of Marine and Atmospheric Science, University of

*Rosenstiel School of Marine and Atmos- Library of Congress
pheric Science, University of Miami, Card No. 79-92393
Miami, Florida

2 NEMOURIA No. 23

Miami, for critical review and encouragement in my research, and to Sally
Diana Kaicher, St. Petersburg, Florida, for the excellent photographs used in
this paper.

This is a scientific contribution from the Rosenstiel School of Marine and
Atmospheric Science, University of Miami.

NEW TAXA

Turbinidae Olividae

Guildfordia kurzi n.sp. Oliva baileyi n.sp.
Trichotropidae Cancellariidae

Zelippistes eccentricus n.sp. Agatrix (Olssonella) nodosivaricosa n.sp.
Ovulidae Conidae

Pseudocypraea exquisita n.sp. Conus aphrodite n.sp.
Cassidae Conus boholensis n.sp.

Morum (Oniscidia) Kurzi n.sp. Conus fragilissimus n.sp.
Muricidae Conus nereis n.sp.

Axymene philippinensis n.sp. Conus zulu n.sp.

The type material is deposited in the type collection of the Department of
Mollusks, Delaware Museum of Natural History, Greenville, Delaware.

FAMILY TURBINIDAE
SUBFAMILY ASTRAEINAE
Genus Guildfordia Gray, 1850

Guildfordia kurzi new species
(Figures 1 and 2)

MATERIAL EXAMINED: Holotype — Length 49 mm, width 47 mm, spire height
21 mm; 300 m depth off Balicasag, Bohol Is., Philippines; June 1978; DMNH
No. 126389. Paratypes — Lengths 50 mm, 48 mm, 46 mm, 42 mm; same depth,
locality, and date as holotype; DMNH No. 126390.

Figure 1: Guildfordia kurzi n.sp., dorsal aspect of holotype.

Figure 2: Guildfordia kurzi n.sp., ventral aspect of holotype.

Figure 3: Guildfordia triumphans (Philippi, 1841), Shikoku Is., Japan, dorsal
aspect, width 55 mm.

Figure 4: Guildfordia triumphans (Philippi, 1841), ventral aspect of same
specimen shown in Figure 3.

Figure 5: Pseudocypraea exquisita n.sp., dorsal aspect of holotype.

Figure 6: Pseudocypraea exquisita n.sp., ventral aspect of holotype.

Figure 7: Zelippistes eccentricus n.sp., dorsal aspect of holotype.

Figure 8: Zelippistes eccentricus n.sp., ventral aspect of holotype.

Figure 9: Zelippistes eccentricus n.sp., lateral aspect of holotype.

December 30, 1979

4 NEMOURIA No. 23

SHELL DESCRIPTION: Flattened, discoidal; body whorl with 7-9 short, solid
spines; dorsal surface with 9-11 spiral rows of coarse granulations; basal surface
with 5-6 spiral rows of coarse granulations arranged in 2 groups — 2 rows along
periphery and 3-4 rows surrounding umbilical region; wide, deeply incised
sulcus between 2 groups of granulations; umbilicus closed but with sunken area
in center; dorsal color varying from greenish tan to lavender, basal color white;
umbilicus with green or pink tint; operculum shiny, calcareous, flattened,
elliptical in outline.

TYPE LOCALITY: 300 m depth off Balicasag, Bohol Is., Philippines.

DISTRIBUTION: At present known only from deep water in the Central Philip-
pine Trenches.

ECOLOGY: Exploratory trawls have shown that Guildfordia kurzi prefers mud
bottoms at a depth of approximately 200-500 m. The new species is a com-
ponent of the upper slope ecosystem surrounding the steep-sided trenches in
the Central Philippines. Gastropods collected with G. kurzi include: Cypraea
guttata Gmelin, 1791; Cypraea katsuae Kuroda, 1950; Oliva dubia Schepman,
1911; Agatrix (Olssonella) nodosivaricosa new species; Lyria planicostata Sower-
by, 1903; Conus kintoki Habe and Kosuge, 1952; Conus kimioi Habe, 1965;
C. kinoshitai Kuroda, 1956; C. praecellens A. Adams, 1854; and Comitas ensuyen-
sis (Shikama and Hyashi, 1977).

ETYMOLOGY: For Richard M. Kurz, Wauwatosa, Wisconsin, who first recog-
nized the species as new and whose generosity in donation of material for
study has been of great help to workers at many museums and scientific insti-
tutions.

REMARKS: Only four species of Guildfordia were previously known, these
being divided into two groups: Guildfordia s.s. and the subgenus Pseudoastralium
Schepman, 1908 (Wagner and Abbott, 1978, pp. 02-002). The new species
belongs to the nominate group and is most closely related to Guildfordia trium-
phans (Philippi, 1841) (pl. 1, figs. C and D). Guildfordia kurzi differs from this
species by having a higher spire, shorter and solid spines, more numerous and
coarser granular sculpturing, and greener color. The most important character
for separating the two species is seen in the basal sculpturing. Guildfordia
triumphans has a smooth, unornamented base, with only a single row of granu-
lations surrounding the umbilical region; G. kurzi is heavily sculptured, with
coarse granulations and a deeply incised spiral sulcus.

December 30, 1979 Petuch 5

FAMILY TRICHOTROPIDAE
Genus Zelippistes Finlay, 1926

Zelippistes eccentricus new species

(Figures 7—9)
MATERIAL EXAMINED: Holotype — Length 18 mm, width 11 mm, spire height
11 mm; 250 m depth in China Strait, off Samarai Is., Eastern Papua, Papua-New
Guinea; January 1970; DMNH No. 123691.

SHELL DESCRIPTION: Thin, fragile, smooth and glossy; body whorl flattened,
discoidal; early whorls scalariform, producing sharply projecting spire; whorls
rounded, strongly tricarinate; body whorl and early whorls completely uncoiled
and separated from one another; aperture flaring, D-shaped; color uniform pale
pinkish cream; aperture salmon-pink; periostracum thin, smooth, translucent
yellow.

TYPE LOCALITY: 250 m depth in China Strait, off Samarai Is., Eastern Papua,
Papua-New Guinea.

DISTRIBUTION: At present known only from type locality.

ECOLOGY : Zelippistes eccentricus is a component of the organic-rich mud bot-
tom community found at approximately 200 m depth around the Milne Bay
area of Eastern Papua. Exploratory trawling has shown that the bottom is
rich in sponges and scattered beds of brissid-type irregular echinoids. Gastropods
commonly taken with the new species include: Cypraea bregeriana Crosse, 1868;
Distorsio reticulata (Roding, 1798); Murex pecten Lightfoot, 1786; Murex
ternispina Lamarck, 1822; Conus floridulus Adams and Reeve, 1848; C. filicinctus
Schepman, 1913; C. Lynceus Sowerby, 1857; C. poehlianus Sowerby, 1887; and
C. schepmani Fulton, 1936.

ETYMOLOGY: In reference to the rather unusual uncoiled whorls.

REMARKS: Zelippistes eccentricus is the first known tropical member of a
basically antitropical species complex. In shell morphology, the new species is
closest to the temperate-water, South Australian Z. blainvilleanus (Petit, 1851)
(Macpherson and Gabriel, 1962, p. 129, fig. 153) and the Japonic Z. helicoides
(Gmelin, 1791) (Kira, 1962, p. 29, pl. 14, fig. 11). Zelippistes eccentricus
differs from both these species in being larger, having an exerted spire, and
having uncoiled, planulate whorls.

6 NEMOURIA No. 23

FAMILY OVULIDAE
SUBFAMILY EOCYPRAEINAE
Genus Pseudocypraea Schilder, 1927

Pseudocypraea exquisita new species
(Figures 5 and 6)

MATERIAL EXAMINED: Holotype — Length 9 mm, width 5 mm; app. 250 m
depth off Panglao, Bohol Is., Philippines; June 1978 ; DMNH No. 126392.

SHELL DESCRIPTION: Thin, small, ovately pyriform; dorsum humped; body
whorl with numerous fine, evenly spaced spiral threads; outer lip thickened,
smooth, shiny; anterior and posterior terminals well produced, prominent;
aperture narrow; columella and base smooth and polished; outer lip with 20
well developed teeth; columella with 24 fine teeth; posterior 5 labial teeth more
developed than the others, projecting beyond lip margin; color pale tan with
large, irregular patches of wine-red; base white with 2 pale red bands.

TYPE LOCALITY : 250 m depth off Panglao, Bohol Is., Philippines.

DISTRIBUTION: At present known only from deep water in the Central Philip-
pines.

ECOLOGY: Pseudocypraea exquisita is a component of the lower shelf, mud
bottom ecosystem that surrounds the Central Philippine Trenches. Some gastro-
pods taken with this new species include: Chicoreus nobilis Shikama, 1977;
Murexiella martini Shikama, 1977; Conus boholensis new species; C. cancellatus
Hwass, 1792; C. gloriamaris Chemnitz, 1777; C. grangeri Sowerby, 1900; C.
neptunus Reeve, 1843; C. nereis new species; C. praecellens A. Adams, 1854;
Latiaxis diadema A. Adams, 1854; Lyria planicostata Sowerby, 1903; and occa-
sionally such rare species as Cypraea leucodon Broderip, 1828; C. porteri Cate,
1966; C. valentia Perry, 1811.

REMARKS: The genus Pseudocypraea was previously considered to be mono-
typic (Cate, 1973, pp. 4-5; Cernohorsky, 1972, p. 91; Keen, 1971, pp. 497-
498); the new species represents the second known member of this eocypraeid
group. Pseudocypraea exquisita closely resembles P. adamsonii (Sowerby, 1832),
the type of the genus, but differs in being more slender, in having the well
developed posterior and anterior terminals, and in having well developed posterior
labial teeth, which give the outer lip a serrated edge. Pseudocypraea adamsonii
has a heavily sculptured columella and base, whereas the new species contrasts
greatly in having a smooth and highly polished basal area. Pseudocypraea exquisita
is also a much more brightly colored shell; the deep wine-red blotches of the
new species contrast with the pale tan markings of P. adamsonii. The new
species has a finer body sculpture and lacks the prominent longitudinal cross-
hatching seen in P. adamsonii.

December 30, 1979 Petuch )

FAMILY CASSIDAE
Genus Morum Roding, 1798
Subgenus Oniscidia Morch, 1852
Morum (Oniscidia) kurzi new spectes
(Figures 10-13)
MATERIAL EXAMINED: Holotype — Length 23 mm, width 15 mm; 250 m depth
off Panglao, Bohol Is., Philippines; June 1978; DMNH No. 126393.

SHELL DESCRIPTION: Thick, stocky, pyriform with wide shoulder; spire elevated,
exerted; body whorl with 12 raised cords — 8 large, prominent on body whorl
proper and 4 small on anterior canal; 8 varices per whorl; large hooked spines
at juncture of spiral cords and varices; at shoulder, each varix has erect, sharply
pointed spine; parietal shield large, covering whole columellar area, covered with
numerous pustules; outer lip crenulate, with numerous large primary and sec-
ondary teeth; protoconch erect, papillate (see Figure 13); color pale cream with
3 tan bands, 1 on shoulder, 2 on either side of midbody line; base color pattern
overlaid with numerous dark brown fleckings; protoconch and first three whorls
bright pink-purple; parietal shield bright orange-pink with white pustules; outer
lip orange with white teeth; interior of aperture white. In Figure 12, the holo-
type is coated with magnesium oxide to enhance the characteristic sculpture
pattern.

TYPE LOCALITY : 250 m depth off Panglao, Bohol Is., Philippines.
DISTRIBUTION: At present known only from the Central Philippines.
ECOLOGY : Same as that of Pseudocypraea exquisita.

ETYMOLOGY: For Richard M. Kurz, Wauwatosa, Wisconsin, who first recog-
nized the species as new.

REMARKS: The remarkable new species resembles no other known Indo-Pacific
Oniscidia. The only species with which it appears to share any morphological
characters is Morum (Oniscidia) praeclarum Melvill, 1919, from Southeast Africa
and the Seychelles Islands (Emerson, 1977, p. 84; Kilburn, 1975, p. 50). Morum
kurzi differs from M. praeclarum in being less pyriform, by having a less devel-
oped parietal shield, and by having a higher spire. Probably the most striking
difference is seen in the colors of the parietal shields: M. kurzi has a bright
orange shield, whereas that of M. praeclarum is pure white (Emerson, 1977,
p. 83). In this last character, the new species somewhat resembles a miniature
version of the Caribbean M. dennisoni (Reeve, 1842).

8 NEMOURIA No. 23

FAMILY MURICIDAE
SUBFAMILY TROPHONINAE COSSMANN, 1903
Genus Axymene Finlay, 1927
Axymene philippinensis new species
(Figures 14 and 15)
MATERIAL EXAMINED: Holotype — Length 16 mm, width 8 mm; app. 250 m
depth off Panglao, Bohol Is., Philippines; June 1978; DMNH No. 126394.

SHELL DESCRIPTION: Thin, fragile, elongate, fusiform; spire elevated, pro-
tracted; shoulder rounded, suture distinct; body whorl with 15 large, raised
spiral cords, these crossed by axial growth lines, giving cancellate appearance;
spire whorls with 4 spiral cords; apex small, papillate, 15 whorls; outer lip thin,
crenulate, with small tooth at anterior end; columella well defined, glazed;
color pure white; interior of aperture white; operculum unknown. |

TYPE LOCALITY : 250 m depth off Panglao, Bohol Is., Philippines.
DISTRIBUTION: At present known only from deep water in Central Philippines.

ECOLOGY: Same as the previous two species, Pseudocypraea exquisita and
Morum kurzi.

ETYMOLOGY: For the type locality, the Philippine Islands.

REMARKS: The genus Axymene was previously thought to be monotypic (Rad-
win and D’Attilio, 1976, pp. 178-179). This new species represents the second
known species.

Axymene philippinensis closely resembles the New Zealand A. turbator
(Finlay, 1927), type of the genus. The new species differs from A. turbator
by being larger, by having a rounded shoulder instead of an angled one, and by
being pure white instead of chocolate-brown. The presence of an apertural
tooth was not reported in Axymene. By having this character, A. philippinensis
may prove to belong to a separate genus. Until more specimens are collected,
however, it seems best to keep the new species in Axymene.

Figure 10: Morum (Oniscidia) kurzi n.sp., dorsal aspect of holotype.

Figure 11: Morum (Oniscidia) kurzi n.sp., ventral aspect of holotype.

Figure 12: Morum (Oniscidia) kurzi n.sp., detail of protoconch.

Figure 13: Morum (Oniscidia) kurzi n.sp., ventral aspect of holotype, coated
with magnesium oxide to enhance sculpture.

Figure 14: Axymene philippinensis n.sp., dorsal aspect of holotype.

Figure 15: Axymene philippinensis n.sp., ventral aspect of holotype.

December 30, 1979 Petuch 9

10 NEMOURIA No. 23

FAMILY OLIVIDAE
Genus Oliva Bruguiére, 1789

Oliva baileyi new species

(Figures 16-19)
MATERIAL EXAMINED: Holotype — Length 30 mm, width 8 mm; trawled from
180 m depth off Russell Is., Solomon Islands; October 1978; DMNH No. 126395.
Paratypes — Length 28 mm; same depth, locality, and date as holotype; DMNH
No. 126396.

SHELL DESCRIPTION: Highly polished, thickened, oblong, cylindrical; spire
short, flattened, calloused, with incised suture; callus of last whorl produced
at posterior end of aperture; aperture narrow, straight; columella with 12-16
low teeth; posterior end of columella without teeth; color bright yellow-tan
crossed diagonally by widely separated brown bands that extend from shoulder to
anterior tip; diagonal brown bands never coalescing; spire callus bright purple,
contrasting greatly with yellow shell; columella and teeth white; interior of
aperture lavender.

TYPE LOCALITY: 180 m depth off Russell Is., Solomon Islands.

DISTRIBUTION: At present known only from deep water near the Solomon
Islands.

ECOLOGY: Oliva baileyi is a component of the lower shelf, soft bottom eco-
system near the Solomon Islands. Other gastropods commonly taken with the
new species include: Murex heros Fulton, 1936; Pterynotus loebbeckii (Kobelt,
1879); Siratus propinquus Kuroda and Azuma, 1961; Lyria santoensis Ladd,
1975; Conus armadillo Shikama, 1971; C. excelsus Sowerby, 1908; and C.
pretiosus G. and H. Nevill, 1874.

ETYMOLOGY: For Mr. Brian Bailey, Solomon Islands, who trawled the type
specimens and recognized the species as new.

REMARKS: Oliva baileyi most closely resembles O. woolnoughi Ladd, 1934, and
O. lauensis Ladd, 1945, from the Pliocene of the Fiji Islands (Ladd, in Ladd
and Hoffmeister, 1945, p. 368, pl. 53, figs. A and B). Of the two latter species,
O. baileyi most closely resembles O. lauensis, which probably represents the
direct ancestor.

Of the Recent Olividae, the new species appears closest to O. rufula Duclos,
1835, from shallow water in the Indo-Malay Archipelago and the Philippines.
Oliva baileyi differs from O. rufula in being more slender and having a flatter
spire and narrower aperture. The color patterns of the two species differ in an
interesting way: the yellow and brown-banded body color of O. baileyi is the
“photo negative” of the dark brown and white-banded body color of O. rufula.
The strikingly bright purple spire callus and purple aperture of O. baileyi are
unique characters, setting it aside from all other known Olividae.

December 30, 1979 Petuch 11

FAMILY CANCELLARIIDAE
Genus Agatrix Petit, 1967
Subgenus Olssonella Petit, 1970
Agatrix (Olssonella) nodosivaricosa new species
(Figures 26 and 27)
MATERIAL EXAMINED: Holotype — Length 11 mm, width 9 mm; 300 m depth
off Balicasag, Bohol Is., Philippines; June 1978; DMNH No. 126397.

SHELL DESCRIPTION: Thin, fragile, inflated; suture indented, spire protracted,
somewhat scalariform; body whorl with 12 rounded varices, crossed by numerous
raised axial threads; varices ornamented with large, undulating, raised knobs,
giving shell nodulose appearance; columella with 3 large plaits; umbilicus closed;
outer lip with strong lirations; color pale cream with 3 tan bands, one at shoul-
der, one at midbody line, one at anterior end; aperture pale yellow with tan
bands showing through in interior; periostracum thin, tufted at shoulder.

TYPE LOCALITY : 300 m depth off Balicasag, Bohol Is., Philippines.
DISTRIBUTION: At present known only from the Central Philippines.

ECOLOGY: The same as the previous new species, Guildfordia kurzi, with which
the new cancellariid is sympatric.

ETYMOLOGY: In reference to the heavily nodulose varix characteristic of the
new species.

REMARKS: Agatrix nodosivaricosa is closest to A. deroyae (Petit, 1970) from
150 m off the Galapagos Islands. The new species differs from A. deroyae by
having a higher, scalariform spire; a deeply indented suture; 3 tan color bands;
and larger and more pronounced nodules on the varices.

FAMILY CONIDAE
Genus Conus Linnaeus, 1758

Conus aphrodite new species

(Figures 34 and 35)
MATERIAL EXAMINED: Holotype — Length 21 mm, width 11 mm; app. 250
m depth off Panglao, Bohol Is., Philippines; June 1978; DMNH No. 126398.
Paratype — Length 13 mm; same locality, depth, and date as holotype; DMNH
No. 126399.

SHELL DESCRIPTION: Thin, delicate, lightweight, glossy; outline straight sided,
elongate, tapered toward the anterior end; shoulder smooth, sharp, slightly
carinate; anterior one third with 6-8 faint spiral sulci; color lilac-purple with
3 bands of chestnut-brown flammules; base color pattern overlaid with 12
revolving rows of white and brown dots and dashes; one row of dashes just

12 NEMOURIA No. 23

anterior of midbody line always more prominent than others; spire purple with
alternating brown flammules; edge of shoulder with alternating brown and white
dashes; aperture purple; periostracum thin, smooth, translucent yellow.

TYPE LOCALITY: Approximately 250 m depth off Panglao, Bohol Is., Philip-
pines.

DISTRIBUTION: Ranging from the Philippines to at least the Taiwan region
(Walls, 1979, p. 516) at the preferred depth.

ECOLOGY: Conus aphrodite is a component of the lower shelf, mud bottom
ecosystem that surrounds the Central Philippine Trenches and is sympatric with
Pseudocypraea exquisita n.sp., Morum kurzi n.sp., Axymene_ philippinensis
n.sp., Agatrix nodosivaricosa n.sp., Conus boholensis n.sp., C. nereis n.sp., and
other associated gastropods.

ETYMOLOGY: For Aphrodite, Greek goddess of beauty and love, often associated
with the sea.

REMARKS: Walls (1979, p. 516, lower left figure) illustrates in color a specimen
of Conus aphrodite. However, he misidentifies the new species as C. otohimeae
Kuroda and Ito, 1961, a closely related species. Conus aphrodite is the newest
member of an interesting species complex comprising C. nadaensis (Azuma and
Toki, 1970) from Japan and the Ryukyu Islands and C. memiae (Habe and
Kosuge, 1970) and C. otohimeae from deep water off Japan, the Philippines,
and the Solomon Islands (Walls, 1979, p. 707). In general shell morphology, the
new species is closest to C. otohimeae. Conus aphrodite differs from C. otohimeae
in being smaller, having a higher spire, having a consistent purple shell color,
and having a proportionately larger protoconch (a feature that shows up well
on Walls’ plate).

Conus boholensis new species
(Figures 20 and 21)

MATERIAL EXAMINED: Holotype — Length 38 mm, width 18 mm; app. 250 m
depth off Panglao, Bohol Is., Philippines; June 1978; DMNH 126400. Paratypes —
Lengths 26 mm, 17 mm; same depth, locality, and date as holotype; DMNH
No. 126401.

Figure 16: Oliva baileyi n.sp., dorsal aspect of paratype.
Figure 17: Oliva baileyi n.sp., ventral aspect of paratype.
Figure 18: Oliva baileyi n.sp., dorsal aspect of holotype.
Figure 19: Oliva baileyi n.sp., ventral aspect of holotype.
Figure 20: Conus boholensis n.sp., dorsal aspect of holotype.
Figure 21: Conus boholensis n.sp., ventral aspect of holotype.

December 30, 1979 Petuch 13

14 NEMOURIA No. 23

SHELL DESCRIPTION: Thin, delicate, glossy; outline straight sided, very elon-
gate, tapering to anterior end; shoulder sharp, extremely carinate, bladelike;
slight constriction just anterior to shoulder carina; early whorls strongly coronate;
spire whorls excavated because of well developed carina; spire scalariform with
carinae of previous whorls projecting beyond suture; body whorl sculpture
consisting of 25-35 deeply incised spiral sulci; spire smooth with only faint
radiating growth lines extending from suture to carina; color pure white with
scattered red-brown dashes; spire color white with only a few scattered, cres-
cent-shaped, red-brown flammules; aperture white; periostracum unknown.

TYPE LOCALITY: Approximately 250 m depth off Panglao, Bohol Is., Philip-
pines.

DISTRIBUTION: At present known only from around Bohol Island in the Central
Philippines.

ECOLOGY: Conus boholensis, like C. aphrodite, is a component of the lower
shelf, upper slope, mud bottom ecosystem surrounding the Philippine Trenches
and is sympatric with the same gastropods.

ETYMOLOGY: For Bohol Is., the type locality.

REMARKS: The new species most closely resembles Conus cancellatus Hwass,
1792, a widespread offshore Indo-Pacific species (Walls, 1979, pp. 267, 270). In
the Philippines area the two species are sympatric. Conus boholensis differs
from the pyriform C. cancellatus in being consistently straight sided, being more
elongate, having a scalariform spire, and having an up-turned, extremely well
developed carina and accompanying excavated spire — all features not found
in C. cancellatus. The latter species has a spire sculpture composed of revolving
spiral sulci, a character not seen in the smooth-spired C. boholensis.

Conus fragilissimus new species
(Figures 22-25)

MATERIAL EXAMINED: Holotype — Length 30 mm, width 12 mm; 3 m depth
off south coast of Harmil Is., Dahlak Archipelago, Eritrea Province, Ethiopia,
Red Sea; 1974; DMNH No. 126402. Paratypes — Lengths 34 mm, 31 mm,
26 mm; same depth, locality, and date as holotype; DMNH No. 126403.

Figure 22: Conus fragilissimus n.sp., dorsal aspect of holotype.

Figure 23: Conus fragilissimus n.sp., ventral aspect of holotype.

Figure 24: Conus fragilissimus n.sp., dorsal aspect of paratype.

Figure 25: Conus fragilissimus n.sp., ventral aspect of paratype.

Figure 26: Agatrix (Olssonella) nodosivaricosa n.sp., dorsal aspect of holo-
type.

Figure 27: Agatrix (Olssonella) nodosivaricosa n.sp., ventral aspect of holo-
type.

December 30, 1979 Petuch 15

16 NEMOURIA No. 23

SHELL DESCRIPTION: Extremely thin, fragile, translucent, glossy; outline
ovately cylindrical; sides convex, tapering to anterior end; shoulder wide, angled,
with prominent coronations; spire high, stepped, somewhat scalariform; spire
sculpture consisting of 4-6 fine revolving spiral threads; aperture wide, flaring;
color pale tan with longitudinal brown flammules, flammules often coalescing
into large brown patches; base color pattern overlaid with variable amounts of
dots, dashes, and netlike pattern; spire color pale tan with regularly spaced dark
brown flammules; protoconch and early whorls dark brown; shoulder corona-
tions white; aperture white; periostracum smooth, translucent yellow; operculum
unknown.

TYPE LOCALITY: 3 m depth off south coast of Harmil Is., Dahlak Archipelago,
Eritrea Province, Ethiopia.

DISTRIBUTION: At present known only from the central Red Sea area.

ECOLOGY: According to the commercial divers and collectors, Conus fragilis-
simus is found in coral rubble in shallow reefs throughout the Dahlak Archipelago.
Being a member of a piscivorous species group (Walls, 1979, p. 19), the new
species most probably feeds on small benthic fishes, such as blennies and gobies.
Gastropods commonly collected with C. fragilissimus in the Dahlak Archipelago
include: Cypraea erythraeensis Sowerby, 1837; C. nebrites Melvill, 1888; C.
pantherina Lightfoot, 1786; C. turdus Lamarck, 1810; Strombus fasciatus
Born, 1778; S. tricornis Humphrey, 1786; Homalocantha fauroti (Jousseaume,
1888); Naquetia jickelii (Tapparone-Canefri, 1875); Oliva bulbosa Roding,
1798; Conus cuvieri Crosse, 1858; C. erythraeensis Reeve, 1843; C. excavatus
Sowerby, 1866; C. nigropunctatus Sowerby, 1857; and C. taeniatus Hwass,
1792.

ETYMOLOGY: In reference to the almost paper thinness of the shell.

REMARKS: Conus fragilissimus is the newest member of an exclusively Indo-
Pacific species complex comprising C. eldredi Morrison, 1955; C. geographus
Linnaeus, 1758; C. obscurus Sowerby, 1833; and C. tulipa Linnaeus, 1758. This
complex can be further divided into two subgroups: C. eldredi, C. fragilissimus,
and C. geographus, all with strongly coronated shoulders, and C. obscurus and
C. tulipa, both with smooth shoulders. Of the four related species, C. fragilis-

Figure 28: Conus zulu n.sp., dorsal aspect of holotype.
Figure 29: Conus zulu n.sp., ventral aspect of holotype.
Figure 30: Conus zulu n.sp., dorsal aspect of paratype.
Figure 31: Conus zulu n.sp., ventral aspect of paratype.
Figure 32: Conus nereis n.sp., dorsal aspect of holotype.
Figure 33: Conus nereis n.sp., ventral aspect of holotype.
Figure 34: Conus aphrodite n.sp., dorsal aspect of holotype.
Figure 35: Conus aphrodite n.sp., ventral aspect of holotype.

December 30, 1979 Petuch 17

18 NEMOURIA No. 23

simus most closely resembles C. geographus — so much so that the new species
was often called a “dwarf form of C. geographus” by commercial shell dealers
and collectors. Conus fragilissimus differs from C. geographus by having a smal-
ler adult size (average 30 mm as opposed to 100-150 mm for C. geographus),
by lacking a pinkish red base color, by having distinct white shoulder corona-
tions, and by having early whorls that are dark brown instead of pink or red
(Walls, 1979, p. 507).

Conus nereis new species

(Figures 32 and 33)
MATERIAL EXAMINED: Holotype — Length 23 mm, width 12 mm; app. 250 m
depth off Panglao, Bohol Is., Philippines; June 1978; DMNH No. 126404. Para-
type — Length 14 mm; same depth, locality, and date as holotype; DMNH No.
126405.

SHELL DESCRIPTION: Thin, delicate, smooth, glossy; outline straight sided,
elongate; shoulder smooth, sharp, distinctly carinate; small groove just anterior
to shoulder carina; anterior one third with 6-8 wide, deeply incised spiral sulci;
color pale blue-white with 3 light brown bands, one just anterior to shoulder,
one at midbody, and one anterior to midbody line; 3-4 rows of alternating
brown and white dashes between 3 brown bands; dashes rectangular shaped,
giving shell “checkerboard”’ effect; spire color white with alternating dark brown,
crescent-shaped flammules; flammules extend from suture, across shoulder, and
onto body whorl, producing pattern of alternating white and dark brown dashes
along shoulder carina; aperture pale lilac, deeper in interior; periostracum thin,
smooth, translucent yellow-brown.

TYPE LOCALITY: Approximately 250 m depth off Panglao, Bohol Is., Philip-
pines.

DISTRIBUTION: At present known only from the Central Philippines near
Bohol Island.

ECOLOGY: Like Conus aphrodite and C. boholensis, C. nereis is a member of the
mud bottom community that borders the Philippine Trenches. It is sympatric
with the same gastropods.

ETYMOLOGY: For any of the sea nymphs, daughters of Nereus, the constant
attendants of Poseidon (Neptune); since the new species is sympatric with Conus
neptunus Reeve, 1843, the taxon honors the courtiers of the submarine god.

REMARKS: The new species is closest to the widespread Indo-Pacific Conus
eugrammatus Bartsch and Rehder, 1943 (Walls, 1979, pp. 292-293, 450-454)
but differs from that species by having a lower spire; a broader shoulder; and
revolving bands of broad, checker-shaped markings. The most important dif-
ference between these closely related species is seen in the body whorl sculp-

December 30, 1979 Petuch 19

turing: C. eugrammatus is characteristically a heavily sculptured shell, being
completely covered with deeply incised spiral sulci; C. nereis lacks this sculpture
and has a smooth, polished body whorl surface. The two species are sympatric
in the Philippines (Walls, 1979, p. 451).

Conus zulu new species
(Figures 28-31)

MATERIAL EXAMINED: Holotype — Length 62 mm, width 35 mm; app. 40 m
depth off the mouth of the Umfolozi River, Zululand Coast, Natal, South
Africa; February 1973; DMNH No. 126406. Paratypes — Lengths 58 mm, 62
mm, 65 mm, 68 mm; same depth, locality, and date as holotype; DMNH No.
126407. Length 74 mm; 35 m depth off Inhaca Is., Mozambique; January 1975;
DMNH No. 126408.

SHELL DESCRIPTION: Heavy, glossy, pyriform, tapering to anterior end; an-
terior third of body whorl heavily sculptured with deeply incised spiral sulci,
posterior two thirds smooth; shoulder rounded, widest part of body whorl just
anterior to shoulder; spire rounded, smooth, with early whorls exerted; color
varying from yellow to gray-brown to deep gray, anterior half of shell darker
than posterior half; shoulder with clear orange or red band; base color overlaid
with numerous spiral rows of dark brown dots and dashes; anterior third with
6-10 prominent raised cords marked with white and brown dashes; spire color
cream-gray with intermittent, regularly spaced, dark brown, crescent-shaped
flammules; columella gray-white; aperture white, turning pale brown or violet
in interior; periostracum thin, brown, with fine longitudinal striae.

TYPE LOCALITY: Approximately 40 m depth off the mouth of the Umfolozi
River, Zululand Coast, Natal, South Africa.

DISTRIBUTION: From Natal northward to central Mozambique.

ECOLOGY: The new species apparently prefers sand and mud bottoms at depths
ranging from 30 to 50 m‘depth along the Southeast African coast. Gastropods
collected with Conus zulu include: Cypraea lisetae Kilburn, 1975; Murex purdyae
Radwin and D’Attilio, 1976; Turbinella laffertyi Kilburn, 1975; Festilyria
duponti Weaver, 1968; Conus eumitis Tomlin, 1931; C. typhon Kilburn, 1975;
C. visagenus Kilburn, 1975; and Turris tanyspira Kilburn, 1975.

ETYMOLOGY: The taxon honors the renowned Zulu nation of the Republic
of South Africa and southern Mozambique.

REMARKS: Conus zulu is the newest member of an exclusively Indo-Pacific
complex comprising the wide-ranging C. betulinus Linnaeus, 1758; C. figulinus
Linnaeus, 1758; and C. loroisii Kiener, 1845, from India and the Bay of Bengal
region, as well as C. glaucus Linnaeus, 1758, and C. suratensis Hwass, 1792,
from the Southwest Pacific. Of this species group, C. zulu is closest to C. figuli-

20 NEMOURIA No. 23

nus — so much so that Walls (1979, p. 474) considered the African species to be
merely a color form. Conus figulinus is basically flat spired, often with sunken
early spire whorls. Conus zulu, on the other hand, is characteristically high
spired, with the early whorls exerted and well defined. The color patterns of the
two species differ considerably: C. figulinus is dark gray-brown, with a lighter-
colored midbody band, this color base in turn being overlaid with numerous,
fine, uninterrupted bands of black or dark brown. C. zulu is light gray, darker
on the anterior one third, this color base being overlaid with dots and dashes,
much like C. suratensis or C. glaucus. The spire color patterns of the two species
differ greatly, C. figulinus having a completely dark brown or black spire, almost
always devoid of any pale markings, but C. zulu having a characteristic pale spire
with regularly spaced: dark crescents, producing a sun burst effect. Conus zulu
has a thinner and more elongate shell than does C. figulinus.

Conus zulu is a member of the poorly known and highly endemic fauna
found in the Mozambique Channel area. More extensive trawling in this region
will no doubt bring to light many new discoveries.

BIBLIOGRAPHY
Cate,C.N.
1973 A systematic revision of the Recent Cypraeid Family Ovulidae (Mol-
lusca: Gastropoda). The Veliger, 15 (supplement).
Cernohorsky, W. O.
1972 Marine Shells of the Pacific, vol. 2. Pacific Publications, Sydney,
Australia.
Emerson, W. K.
1977 Notes on some Indo-Pacific species of Morum (Gastropoda: Ton-
nacea). The Nautilus, 91(3):81-86.
Keen, A. M.
1971 Sea Shells of the Tropical West America, ed. 2. Stanford University
Press, Stanford, California.
Kilburn, R.N.
1975 The rediscovery of Morum praeclarum Melvill (Cassidae). The Nau-
tilus, 89(2):49-SO0.
Kira, T.
1962 Shells of the Western Pacific in Color. Hoikusha Publishing Co.,
Ltd., Osaka, Japan.
Ladd, H.S., and J. E. Hoffmeister
1945 Geology of Lau, Fiji. Bulletin 181, Bernice P. Bishop Museum,
Honolulu, Hawaii.

December 30, 1979 Petuch pl

Macpherson, J. H., and C. J. Gabriel
1962 Marine Molluscs of Victoria. Melbourne University Press, Melbourne,

Australia.

Radwin, G. E., and A. D’ Attilio
1976 Murex Shells of the World: An Illustrated Guide to the Muricidae.
Stanford University Press, Stanford, California.
Rosewater, J.
1976 William Healey Dall — The Legacy He Left for Malacology. Bulletin
of the American Malacological Union for 1975: 4-6.
Wagner, R.J.L., and R. T. Abbott
1978 Standard Catalog of Shells, ed. 3 (with supplements). American
Malacologists, Inc., Greenville, Delaware.
Walls, J. G.
1979 Cone Shells: A Synopsis of the Living Conidae. T. F. H. Publications,
Inc., Neptune City, New Jersey.

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Published by the
DELAWARE MUSEUM OF NATURAL HISTORY

Kennett Pike, Greenville,
Delaware 19807

NEMOURIA
Occasional Papers of the Delaware Museum of Natural History

NUMBER 24 JANUARY 15,1980

NOTES ON PHILIPPINE BIRDS (NO. 5)
BIRDS OF BURIAS

John E. duPont

INTRODUCTION

The island of Burias lies in the mouth of Ragay Gulf, west of the Luzon
Province of Albay and east of the Sibuyan Group. The island is approximately
70 km by 15 km, with a maximum altitude of 458 m, and is at present 50 per-
cent forested. There is no pronounced maximum rain season and no dry season.
Basically, the island is made up of tertiary sediments.

Dr. Kenneth C. Parkes brought to my attention the fact that Burias was the
largest ornithologically unexplored island in the Philippine Group. The Delaware
Museum of Natural History sponsored field work at Pinamasigan, San Pascual,
Burias, in January 1977, resulting in the collection of 29 species of birds. Three
species (Phylloscopus borealis, Muscicapa griseisticta, and Lanius cristatus) are
well known migratory visitors to the Philippines. The 26 resident species may
be broken down as follows: 13 belong to species that do not vary geographically
within the Philippines, seven belong to widely distributed subspecies occupying
most of the archipelago, one (Sarcops calvus) most closely resembles the sub-
species of adjacent Luzon, four (7Jreron pompadora, Tanygnathus lucionensis,
Copsychus saularis, Cyornis rufigaster) rather surprisingly belong to subspecies
of the central and southern islands rather than to that of nearby Luzon, and one
(Hypsipetes philippinus) is represented on Burias by a distinctive new endemic
race.

Library of Congress
Card No. 79-92780

2 NEMOURIA No. 24

FAMILY ACCIPITRIDAE HAWKS AND EAGLES
Elanus caeruleus hypoleucus Gould, 1859

DMNH-1 6
FAMILY RALLIDAE RAILS
Amaurornis olivaceus olivaceus (Meyen,.1834)
DMNH-~1 ?

FAMILY COLUMBIDAE PIGEONS AND DOVES

Treron pompadora canescens Parkes, 1965

DMNH-1 6,22
Although Burias lies close to Luzon, these specimens show the grayer color
typical of canescens of the southern Philippines and are not axillaris of the
adjacent parts of Luzon.

Ptilinopus leclancheri leclancheri (Bonaparte, 1855)
DMNH-—1 @

Ducula aenea aenea (Linne, 1766)
DMNH-1 6

Geopelia striata striata (Linne, 1766)
DMNH-1 @

Chalcophaps indica indica (Linne, 1758)
DMNH-1 6,1 2

FAMILY PSITTACIDAE PARROTS

Tanygnathus lucionensis salvadorii (Ogilvie-Grant, 1896)
DMNH-1 2
As in the case of Treron pompadora, this specimen is referable to the form
inhabiting the southern islands, lacking the blue on head and back typical of
lucionensis of Luzon.

FAMILY CUCULIDAE CUCKOOS

Eudynamys scolopacea mindanensis (Linne, 1766)
DMNH-—1 6

January 15, 1980 duPont 3

FAMILY ALCEDINIDAE KINGFISHERS

Halcyon chloris collaris (Scopoli, 1786)
DMNH™~"*4 4, 3 in alcohol

Halcyon smyrnensis gularis (Kuhl, 1820)
DMNH-1 6

FAMILY CORACIIDAE ROLLERS

Eurystomus orientalis cyanocollis Vieillot, 1819
DMNH-1 2?

FAMILY CAMPEPHAGIDAE CUCKOO-SHRIKES

Lalage nigra chilensis (Meyen, 1834)
DMNH-~1 4, 3 9, 1 in alcohol

FAMILY ORIOLIDAE ORIOLES

Oriolus chinensis chinensis Linnaeus, 1766
DMNH-1 4, 2

FAMILY CORVIDAE CROWS

Corvus macrorhynchos philippinus (Bonaparte, 1853)
DMNH-1 6

FAMILY PYCNONOTIDAE BULBULS
Hypsipetes philippinus
This species presently has three recognized races. A fourth heretofore un-
known and characterized by its very brightly streaked throat may be called:

Hypsipetes philippinus parkesi new subspecies

Type DMNH 60646, 6 Barrio Pinamasigan, San Pascual, Burias, P.I., 20-
100, January 1977. Wing 110; tail 103; bill 25; tarsus 20.

DIAGNOSIS: Differs from H. p. philippinus (J. R. Forster, 1795) by having a
duller rufous throat heavily streaked with white, also much larger in overall
size but not as large as guimarasensis (Steere, 1890). Differs from guimarasensis
of islands to the south and west by averaging slightly smaller and by having
the throat heavily streaked with white. Differs from mindorensis in having a
rufous rather than deep olive throat. The sexes do not differ and this population
is generally closer to guimarasensis in color than to philippinus.

4. NEMOURIA No. 24

RANGE: Burias

SPECIMENS EXAMINED: philippinus — Cebu 2 d, 2 ¢; Leyte 7 6, 1 9; Lubang
5 6, 4 9; Marinduque 6 6, 5 9; Mindanao (eastern) 25 6, 29 9; Polillo 5 6,2 9;
Samar 4 6, 2 ?. guimarasensis — Masbate 6 6, 1 9; Negros 7 6, 8 2; Panay 5 6,
4 2. mindorensis — Mindoro 15 6, 6 9. parkesi — Burias 13 6,4 2,1 0,9 in
alcohol.

ETYMOLOGY: This new subspecies is named for Kenneth C. Parkes of the
Carnegie Museum of Natural History.

FAMILY TURDIDAE THRUSHES

Copsychus saularis mindanensis (Boddaert , 1783)

DMNH~1 ¢, 1 &, 1 in alcohol
The female lacks the finely barred buff flanks that distinguish C. s. deuw-
teronymus of Luzon.

FAMILY SYLVIIDAE OLD WORLD WARBLERS
Cisticola exilis semirufa Cabanis, 1872
DMNH-2 6

Phylloscopus borealis ssp.
DMNH-1 4, 1 9, 1 in alcohol

FAMILY MUSCICAPIDAE OLD WORLD FLYCATCHERS

Rhipidura javanica nigritorquis Vigors, 1831

DMNH~—1 4g, 1 9, 3 in alcohol
Cyornis rufigaster philippinensis Sharpe, 1877

DMNH-3 4, 2 9, 2 in alcohol

These specimens are typical of the dark southern race philippinensis rather

than the paler simplex of adjacent Luzon.
Muscicapa griseisticta (Swinhoe, 1861)

DMNH-1 6,12
Hypothymis azurea azurea (Boddaert, 1783)

DMNH-~1 @, 1 in alcohol

FAMILY ARTAMIDAE WOOD-SWALLOWS

Artamus leucorhynchus leucorhynchus (Linnaeus, 1771)
DMNH-2 6

January 15, 1980 duPont 5

Figure 1: Hypsipetes philippinus philippinus
Figure 2: Hypsipetes philippinus parkesi
Figure 3: Hypsipetes philippinus guimarasensis
Figure 4: Hypsipetes philippinus mindorensis

’

6 NEMOURIA No. 24

FAMILY LANITDAE SHRIKES

Lanius cristatus lucionensis Linnaeus, 1766
DMNH~—4 4g, 3 2, 5 in alcohol

FAMILY STURNIDAE STARLINGS
Aplonis panayensis panayensis (Scopoli, 1783)
DMNH-3 ¢, 2 9, 1 0, 1 in alcohol

Sarcops calvus (Linnaeus, 1766)

DMNH-5 6
The birds match those from southern Luzon; however, they show some
darkness in the middle of the back, which shows a slight tendency towards
melanonotus.

FAMILY NECTARINIIDAE SUNBIRDS
Nectarinia jugularis jugularis (Linnaeus, 1766)
DMNH-—12 6, 8 2, 11 in alcohol

FAMILY DICAEIDAE FLOWERPECKERS

Dicaeum australe australe (Hermann, 1783)
DMNH-1 6

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Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
Kennett Pike, Greenville,
Delaware 19807

NEMOURIA
Occasional Papers of the Delaware Museum of Natural History

NuMBER 25 OcToBER 30, 1980

THE EXTENT AND SEQUENCE OF THE
MOLTS OF THE YELLOW-RUMPED WARBLER

John P. Hubbard*

INTRODUCTION

The Yellow-rumped Warbler (Dendroica coronata) is a breeding species of
northern and western North America, ranging as such from Alaska to Guatemala.
This study was undertaken as part of an investigation of the overall relationships
and evolution of this complex of variously differentiated populations (see
Hubbard 1967, 1969, 1970), and it focuses on the descriptive aspects of the
molts. Aside from Foster’s study (1967) of the Orange-crowned Warbler (Vermi-
vora celata), 1 know of no detailed work that has been published on the molts of
a member of the family Parulidae occurring over such an extended range. Treated
here are all the races of D. coronata, that is, D. c. coronata (includes D. c. hooveri),
breeding in northern North America; D. c. auduboni (includes D. c. memorabilis),
breeding in western United States and adjacent Canada; D. c. nigrifrons, apparently
resident in northwestern Mexico; and D. c. goldmani, resident in Guatemala and
adjacent Mexico.

The molts of D. c. coronata have been treated by Dwight (1900), whereas
those of D. c. auduboni are covered in a general way in Bent (1953), Swarth
(1926), and Phillips et al. (1964). The molts of D. c. nigrifrons and D. c. goldmani
are not treated in the literature, and they appear not to have been previously
studied. In general, the extent and sequence of molt among the various races are
closely similar, with the exceptions being pointed out in the discussions that
follow.

MATERIALS AND METHODS

I examined almost 2000 specimens for molt, including 68 of the rare D. c.
nigrifrons and 26 D. c. goldmani. Of these, 246 in active molt were selected to
document the extent and sequence of molts in this complex. These included 110

2016 Valle Rio Library of Congress
Santa I’e, New Mexico 87501 Catalog No. 80-68398

2 NEMOURIA Now 25

coronata, 120 auduboni, and 16 nigrifrons, with 43 in postjuvenal (or first pre-
basic) molt, 95 in prenuptial (or prealternate) molt, and 108 in postnuptial (or
prebasic) molt. No specimens were available of birds in the postnatal molt;
hence, it is not discussed in this paper.

The selected specimens were examined under a high-intensity Tensor lamp
with a 60 watt bulb, and most were viewed under magnification. Feathers were
gently probed with a dissecting needle, and detailed comparisons and notes were
made on each of the specimens. Many other molting specimens were examined
in a more superficial manner, mainly to check the documented data and to pro-
vide other insights into molt. Examined in this manner were several hundred
additional specimens, especially those in the postjuvenal and postnuptial molts.

In order to facilitate comparisons with the molts of Vermivora celata, the
nomenclature used for feather tracts in this paper follows Foster (1967). Primaries
are numbered from the inside out, with the outermost (vestigial) primary being
number 10. Secondaries are numbered from the outside in, with the innermost
(tertials) being 7, 8, and 9.

Postjuvenal Molt

The postjuvenal body molt generally begins between the time the tail is one-
quarter and one-half grown. As a rule it initially involves the upper and lower
marginal wing coverts and often the crural tract (Table 1). Molt then starts in
the anterior humeral tract and, either simultaneously or slightly later, in the
adjacent sternal region of the ventral tract. As the tail nears full length, additional
loci appear on the forehead, chin, and/or throat, and in the pelvic portion of the
dorsal tract, the latter giving rise to the yellow rump. Also at this time or slightly
later, molt begins in the femoral tract and in the cervical region of the ventral
tract. The spread and anastomosing of the ventral molt loci progressively give
rise to the inverted-Y pattern of new feather growth characteristic of the ventral
area in the juveniles of many passerines. Additional molt then appears on the
posterior capital tract and dorsal region of the spinal tract; and the replacement
of the greater and, later, the middle secondary coverts begins. Molt finally spreads
from established loci to the sides of the head, the spinal cervical region, abdominal
region, and tail coverts; and these are among the last areas to acquire the winter
plumage. Retained at this molt are the remiges, rectrices, primary coverts, and
alular feathers. The statement in Phillips et al. (1964:151) that this molt (“‘pre-
basic’) is complete in young birds refers to the body plumage only (A. R.
Phillips, pers. comm.).

Postnuptial Molt

Generally, the replacement of body plumage in this molt begins at the time
of the loss of the first or second primary (Table 2). Simultaneously or closely
following this loss is initial molt in the anterior marginal coverts, although in a

October 30, 1979 Hubbard 3

TABLE 1: The Sequence of Inception of the Postjuvenal Molt

Alar tract (marginal coverts)

Crural tract

Humeral tract (anterior)

Ventral tract (sternal region)

Capital tract (frontal, malar regions)

Ventral tract (cervical, interramal, submalar, axillary regions)
Spinal tract (pelvic region)

Femoral tract

Capital tract (coronal, loral, occipital regions)
Spinal tract (dorsal, interscapular regions)
Alar tract (greater secondary coverts)

Capital tract (auricular, superciliary regions)
Ventral tract (abdominal region)

Spinal tract (cervical region)

Alar tract (middle secondary coverts)

Caudal tract (tail coverts)

NO =

8.
2.

TABLE 2: The Sequence of Inception of the Postnuptial Molt

Alar tract (primaries 1, 2; marginal coverts)

Alar tract (primary 3; secondary 8)

Caudal tract (rectrices 1, ye 3° tail coverts)

Spinal tract (pelvic region)

Femorla tract

Alar tract (primary 4)

Spinal tract (pelvic region)

Femoral tract

Humeral tract (anterior)

Alar tract (primary 5; secondary 9; greater secondary coverts)
Caudal tract (rectrices 4, 5*, 6*)

Ventral tract (cervical region)

Crural tract

Alar tract (primary 6; secondaries 1*, 7*)

Spinal tract (interscapular region)

Ventral tract (interramal, submalar regions)

Capital tract (occipital, malar regions)

Alar tract (primary 7; middle secondary coverts)
Ventral tract (abdominal region)

Alar tract (primary 8; secondaries 2, 3*, 4*)

Spinal tract (cervical region)

Capital tract (coronal, frontal, loral, superciliary, auricular regions)
Alar tract (primaries 9, 10; secondaries 3, 4*, 6", 5*)
Alar tract (alular feathers)

‘ May be affected somewhat later.

4 NEMOURIA No. 25

few cases I have not been able to detect molt there as late as the loss of the third
or (very rarely) fourth primary. The first sign of body molt is usually in the pelvic
region (rump) of the spinal tract; this is followed by the femoral tract, generally
coinciding with the loss of the third or fourth primaries. Also with the loss of
the third primary is that of the eighth secondary (middle tertial) and first to third
pairs of rectrices; molt of the tail coverts usually also begins at this time. Generally
with the loss of the fourth primary, additional loci appear in the sternal region
of the ventral tract and in the humeral tract; molt then moves anteriorly from
the pelvic to the dorsal region of the spinal tract. As the fifth primary is dropped,
usually the ninth secondary is also lost, and molt begins in the crural region. At
the same time, molt in the ventral tract moves from the sternal region to the
cervical region. Often by this time many birds have lost all, or all but the outer
pair or two, of the old rectrices. With the loss of the sixth primary, molt generally
begins on the head—especially on the chin, throat, and the lateral and posterior
occipital areas. At the same time in the spinal tract, molt spreads anteriorly
into the interscapular region. With the loss of this primary or the seventh, the
first and seventh secondaries are also dropped, as well as the last of any remaining
old rectrices. Rectrix growth is rapid, and the tail is usually nearly full grown
before the seventh primary reaches its full length. Starting with the loss of the
eighth primary, the remaining secondaries are gradually dropped in the order
2-3-4-6-5. With the molting of the eighth, ninth, and vestigial (tenth) primaries,
molt begins on the forehead, auriculars, middle of the crown, and cervical region
of the spinal tract. These areas, along with the chin and the abdomen, are generally
the last in which body molt is completed.

The greater secondary coverts are generally lost with the dropping of the fifth
or sixth primaries, and the middle ones are lost after the greater coverts are
partially erupted—usually by the time the eighth primary has been dropped. The
primary coverts fall in sequence with their attendant primaries, the first new
ones usually not appearing before the fourth primary is dropped. The ailular
feathers are the last of the wing to be replaced. In the final stages of remix
growth, the outer primaries and the inner secondaries reach full growth nearly
simultaneously.

Although the power of flight is retained during the course of this molt, there
is a reduction in the wing area that probably inhibits flight to a degree. Some
estimate of this reduction can be made by calculating the percent that each of
the major remiges (excluding the tiny tenth primary and ninth secondary) con-
tributes to the wing area, and then summing the areal reduction in specimens at
different stages of molt. Thus, one finds that primaries 5, 6, and 7 each comprise
about 7 percent of the wing area; primaries 1, 2, and 3 and secondaries 4, 5, 6,
7, and 8 are each 5 percent, and the remaining remiges 6 percent. In 47 heavily
molting specimens, the maximum reduction in wing area ranged from 10 to 30
percent, with the median 20 percent. The reduction of wing area with the loss
of given remiges varies considerably because of slightly different rates of replace-

October 30, 1979 Hubbard 5

ment of other remiges; however, the greatest reduction is generally at the time of
loss of primaries 7, 7+8, and 8+9. Large reductions of wing area (25 to 30 percent)
also occurred with the loss of primaries 3+4, 4+5, 5+6, and 6; but generally the
loss of single or pairs of primaries in this range caused lesser reductions (10 to
25 percent). Assuming that the loss of wing area reduces flight capacity and
makes molting individuals more vulnerable to predation, one can appreciate the
increased secretiveness reported in many passerines during their postnuptial molt.

Prenuptial Molt

Generally, the prenuptial molt begins on the upper midback and the middle
and upper breast (Table 3). This is usually followed by molt beginning first on
the sides of the anterior part of the crown and then on the lores and forehead.
At this time, replacement of the greater and middle secondary coverts begins,
and this aspect of molt produces the characteristic wing covert patterns of the
various subspecies. Usually the inner five or six pairs of coverts are renewed,
whereas the distal three or four pairs are retained. With the beginning of molt
on the throat, the loci elsewhere show progression anteriorly and posteriorly.
In the final stages, molt includes the feathers of the chin, central crown, auriculars,
nape, and anterior scapulars. Generally retained at this molt, in addition to the
flight feathers, are most of the wing coverts, the tail coverts, the lower scapulars,
and the feathers of the rump and abdomen. In females the molt is often less
extensive than in males, with some specimens showing very few, if any, signs of
a prenuptial molt. However, in most females, there is at least a partial replace-
ment of the dorsal and ventral body plumage, particularly on the head, back,
and breast.

TABLE 3. The Sequence of Inception of the Prenuptial Molt

1. Spinal tract (interscapular region)
Ventral tract (cervical, sternal regions)
2. Spinal tract (dorsal, cervical regions)
Capital tract (frontal region)
3. Ventral tract (submalar region)
Capital tract (loral, malar regions)
Humeral tract (anterior)
Alar tract (greater and middle secondary coverts)
4. Ventral tract (interramal region)
Capital tract (coronal, auricular, superciliary, occipital regions)
Humeral tract (anterior scapulars)

6 NEMOURIA Noi25

DISCUSSION
Comparison of Molts within Dendroica coronata

The postnuptial, postjuvenal, and prenuptial molts differ from one another
both in the extent of plumage replacement and in the sequence with which
feather tracts are affected. In the relative extent of plumage replacement, the
molts may be ranked as follows: postnuptial, postjuvenal, and prenuptial.

The postnuptial molt replaces all of the plumage, whereas the postjuvenal
replaces all except the remiges, rectrices, primary coverts, and alular feathers.
The prenuptial molt replaces a more variable amount of plumage, depending on
sex and possibly age and subspecies. In males of coronata and auduboni, most of
the body plumage is replaced (but usually not on the rump, tail coverts, and
abdomen), whereas on the wings only the inner five or six pairs of greater and
middle secondary coverts are renewed.

The extent of the molt in males of the races nigrifrons and goldmani is less
certain, mainly because of the absence of sufficient specimen material. It is clear
that first-year males of both races undergo an extensive prenuptial molt, because
their femalelike winter plumage differs from their breeding plumage; thus, the
latter can hardly be assumed by wear alone. On the other hand, adult males of
goldmani appear to be identical in winter and breeding seasons, and it is con-
ceivable that they do not have a prenuptial molt. The only “‘summer”’ adult male
of this race I have seen is from early May. The plumage wear in this specimen is
not pronounced, which may indicate that a prenuptial molt does occur. The
winter plumage of adult males of nigrifrons wears to resemble that of the breeding
season, but the lack of pronounced abrasion in June specimens suggests that
they, too, have an extensive prenuptial molt. In addition, I have examined three
male specimens of nigrifrons that do show active molt in the spring (one adult and
two first-year birds), which proves the existence of this molt but not its frequency.

Females of coronata and audubonishow various gradations from no prenuptial
molt to one as complete as in males. Most show at least a partial replacement of
body plumage, usually on the head, back, and breast but averaging less than in
males. I have not seen females of nigrifrons and goldmani in active molt, but the
presence of apparently new feathers (in the same area as affected in coronata
and auduboni) suggest that they, too, undergo a partial prenuptial molt.

In the sequence of body plumage and covert replacement there are both
differences and similarities among the three molts in the D. coronata complex.
In the postnuptial and the postjuvenal molts, the sequence of replacement is
similar in the marginal and secondary coverts and in the humeral, ventral, and
capital tracts. On the other hand, in the postnuptial molt, the spinal, femoral,
and caudal (tail coverts) tracts are affected earlier and the crural tract later than
in the postjuvenal molt. Most of these differences are slight, but crural and tail
covert molt are markedly different, with these areas being affected at reversed
and opposite extremes of their cycles. In Vermivora celata, Foster (1967) found
the sequences of these two molts to be alike. The prenuptial molt in the Den-

October 30, 1979 Hubbard a

droica coronata complex is more similar in sequence to the postjuvenal molt,
although in the former the spinal and ventral tracts precede head molt. In the
postjuvenal molt, molt in the humeral and ventral tracts precedes head molt.

In the sequence of molt within individual feather tracts of Dendroica coronata,
there is an overall similarity among the three molts. In the spinal tract, the post-
nuptial and postjuvenal molts begin in the anterior pelvic region and move
anteriorly and posteriorly. In the prenuptial molt, the pelvic region usually is not
affected; instead, replacement begins in the dorsal and interscapular region and
moves anteriorly. In the ventral tract, all three molts begin in the sternal region,
closely followed (simultaneously in the prenuptial) by the cervical region and
then the submalar and/or interramal regions. Molt then moves outward from
these loci, and the abdominal area is molted last (this area is usually not affected
in the prenuptial molt). In the postjuvenal and prenuptial molts, the capital tract
usually begins molt on the foreparts of the head and progresses posteriorly toward
a secondary locus on the crown. In the postnuptial molt, the secondary locus is
on the forehead, and the primary one is on the crown.

Comparison of Molts with Vermivora celata

As indicated earlier, only the Orange-crowned Warbler has been studied in
sufficient detail (Foster, 1967) to offer good comparison in its molts to those of
the Dendroica coronata complex. In V. celata the postjuvenal molt may begin in
any or all of three loci, that is, the spinal, ventral, or alar tracts (marginal coverts).
D. coronata begins molting in the marginal coverts, whereas molt in the ventral
tract occurs in the second stage and in the spinal tract in the third. The second
stage of molt in V. celata involves the caudal tract, in addition to the three tracts
already mentioned; in D. coronata, caudal molt (tail coverts) occurs in the last
stage. The third stage of molt in V. celata involves all tracts, with the addition of
molt in the crural and capital tracts. In D. coronata, the capital tract is affected
earlier, usually half-way through the molt cycle and before caudal molt begins;
molt usually begins in the crural tract in the initial stage. Within tracts, the
sequence of molts is generally similar between the two species, although there
are a few differences. For example, the spinal and ventral molts begin more
posteriorly in D. coronata than in V. celata, usually in the pelvic and sternal
regions, respectively, in D. coronata. In the capital tract, the molt of D. coronata
also appears to differ, mainly in that the malar and loral regions are involved
earlier than in V. celata.

The postjuvenal molt of V. celata appears to be more telescoped than that of
the D. coronata complex, with a less clearcut distinction between the inception
of feather replacement in different tracts and regions. This accounts for many of
the differences in the molts between the two species, although there remain
those involving early or late extremes in the molt cycle. Thus, in D. coronata
there is early molt in the crural tract and late molt in the caudal tract, compared
to the reverse in V. celata. The difference in caudal molt may be related to the

8 NEMOURIA No. 25

fact that some V. celata replace one or more pairs of rectrices, but the significance
of delayed crural molt is not apparent.

The postnuptial molt of V. celata follows the same sequence in the nonflight
feathers as its postjuvenal molt, and it begins after the partial replacement of
primary 1+2 (Foster, 1967). This molt differs from the postnuptial molt of D.
coronata less than it does from the latter’s postjuvenal molt, and again the differ-
ences stem mainly from the telescoped sequence of V. celata. Basically, the
differences are that molt in the ventral tract and the greater coverts is com-
paratively retarded in D. coronata, whereas humeral molt is slightly advanced. In
the timing of crural and tail covert molt, the postnuptial molt of D. coronata
agrees with V. celata, that is, the caudal molt coming earlier and the crural molt
later in the sequence; in the postjuvenal molt of D. coronata this sequence is
reversed. In molt of the flight feathers, the sequence is identical in the two species
in the remiges and similar in the more variably molted rectrices. Loss of the
earliest secondary (the ninth) usually coincides with the loss of the fifth primary
in D. coronata and usually occurs between the loss of the fourth and fifth in
V. celata.

ACKNOWLEDGMENTS

First, I am grateful to the many people involved in my use of specimens
from the following institutional and private collections: American Museum of
Natural History; University of Alberta; University of Arizona; D. Boag; Provincial
Museum of British Columbia; British Museum (Natural History); University of
California at Los Angeles (Dickey Collection); California Academy of Sciences;
National Museum of Canada; Carnegie Museum of Natural History; Field Museum
of Natural History; Denver Natural History Museum; University of Kansas
Museum of Natural History; Los Angeles County Museum; Louisiana State
University Museum of Natural History; University of Michigan Biological Station;
University of Michigan Museum of Zoology; Museum of Comparative Zoology;
Museum of Vertebrate Zoology; University of New Mexico; New Mexico State
University; Occidental College (Moore Collection); Ohio State University; Royal
Ontario Museum; University of Oregon; O. S. Pettingill, Jr.; R. Sale; San Diego
Natural History Museum; G. M. Sutton; United States National Museum of
Natural History; University of Utah; Virginia Polytechnic Institute and State
University (Bailey-Law Collection); and University of Washington. Individual
contributions of many are also appreciated, including those of J. Barlow, N. L.
Ford, S. D. MacDonald, G. G. Musser, and R. W. Storer. Mercedes Foster, David
M. Niles, Kenneth C. Parkes, and Ralph J. Raitt read and commented on earlier
drafts of this paper. I am grateful for financial aid from the Chapman Fund and
the National Science Foundation and to officials in Canada, Mexico, and the
United States for collecting permits. Final thanks go to my wife Claudia, who
helped bring this study to its conclusion.

October 30, 1979 Hubbard 0

BIBLIOGRAPHY
Bent, A. C.
1953 Life histories of North American wood warblers. Bull. U. S. Nat.
Mus., 203.
Dwight, J., Jr.

1900 Sequence of plumages and molts in the passerine birds of New York.
Ann. N. Y. Acad. Sci., 13:73-360.
Foster, M. S.
1967 Molt cycles of the Orange-crowned Warbler. Condor, 69:169-200.
Hubbard, J. P.
1967 A systematic study of the Dendroica coronata complex (Aves.). Un-
published Ph.D. thesis, University of Michigan.
1969 The relationship and evolution of the Dendroica coronata complex.
Auk, 86:393-432.
1970 Geographic variation in the Dendroica coronata complex. Wilson Bull.,
82:355-369.
Phillips, A. R., J. Marshall, and G. Monson
1964 The Birds of Arizona. Univ. Arizona Press, Tucson.
Swarth, H. S. |
1926 The Audubon’s Warbler. Bird-lore, 28:82-85.

4 3

i

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY

Kennett Pike, Greenville,
Delaware 19807

OCT 2 61981

LIBRARIES

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 26 OCTOBER 16, 1981

A TAXONOMIC REVIEW OF THE
SPOTTED-BREASTED ORIOLE

Robert W. Dickerman*

INTRODUCTION

The Spotted-breasted Oriole (Icterus pectoralis) is a resident of semi-
arid Pacific lowlands from Guerrero, Mexico south to northwestern Costa
Rica and of xeric habitats in interior areas of Guatemala and Honduras. Four
forms have been described: nominate pectoralis, Wagler 1829 (from Mexico),
guttulatus, Lafresnaye 1844 (from Mexico, but see beyond), anthonyi,
Griscom 1930 (from northwestern Guatemala), and espinachi, Ridgway
1882 (from northwestern Costa Rica). Monroe (1968, p350), the latest author
to discuss geographic variation in the species, wrote, “In an extensive series
from all parts of Central America, size variation ranges clinally from larger
northern to smaller examples from Costa Rica. All color characters seem to be
bridged by individual variation; these characters include intensity of orange,
...and amount of pectoral spotting’. He recognized only nominate pectoralis.
Hellmayr (1937) earlier recognized pectoralis (with guttulatus as asynonym),
anthonyi and espinachi. Blake (1968) added anthonyi to the synonomy of
pectoralis and recognized espinachi.

Unfortunately, Monroe and others apparently did not realize the extent
to which variation in size and plumage is due to age or wear. For example, the
holotype and most of the type series of espinachi are immature birds, thus
accounting in part for their smaller size and their paler color. Not realizing this

*Department of Microbiology Library of Congress
Cornell University Medical College Catalog No. 81-69250
New York, New York 10021

and

Department of Ornithology

American Museum of Natural History
New York, New York 10024

ps NEMOURIA No. 26

led Griscom (1930) to believe that the Guatemala birds (his series consisted
mostly of adults) were slightly larger in size than espinachi. They may be, but
the difference between my series is small and statistically insignificant. Failure
to consider size variation with age also accounts for Monroe’s supposed cline
in size. No geographically ordered variation in size can be found if specimens
are properly segregated by age and sex (and degree of feather wear) and if only
definitive-plumaged adults are measured (Table 1, Fig. 1 and 2). Taxonomi-
cally useful characters in this species are limited to the general intensity of
color of head, rump and underparts, and to the extent of breast spotting.

In the accounts of subspecies specimens of adults that were measured are
presented as totals by country (or state) for I._p. guttulatus and by locality for
the other subspecies. Specimens examined in predefinitive or in worn (and
unmeasurable) definitive plumage are not cited.

Museum acronyms and abbreviations used beyond are identified in the
section of acknowledgements.

SUBSPECIES ACCOUNTS

Icterus pectoralis espinachi Ridgway. Proc. U.S. Nat. Mus. 5:392, 1882.
Type locality: La Palma de Nicoya, Guanacaste, Costa Rica. Holotype:
immature male, USNM 87361 (examined).

DIAGNOSIS: Head and rump orange-yellow, similar to nominate pectoralis
but slightly brighter; much yellower in those areas than J. p. guttulatus.
Slightly paler orange on belly than guttulatus, but this is seen only in series.
Pectoral spots generally isolated from one another, i.e., not coalesced (Fig. 3).

RANGE: Northwestern Costa Rica (Guanacaste and Puntarenas Depart-
ments) and probably adjacent Nicaragua in the arid zone of Rivas Department
from whence no specimens have been seen.

DISCUSSION: It is interesting that the pale populations at the northern and
southern ends of the species range are most similar. One of six male espinachi
(UMMZ 160665, Fig. 3) is as deeply colored as guttulatus, and two (CM
26877 and AMNH 521932) have heavily marked breast bands similar to
those of the Guerrero population named beyond. In size espinachi may be
slightly smaller than the “anthonyi” compoment of guttulatus but is consider-
ably smaller than the nearer Honduras subpopulation of guttulatus. In any
case there is no clinal increase in size from south to north as proposed by
Monroe (1968).

SPECIMENS EXAMINED: 16. Costa Rica: Guanacaste Department,
Bebedero 4 males, 4 females; Bolson 3 males, 1 female; Philadelphia 1 male, 1
female; Santa Cruz Mujal 1 male. Puntarenas Department, Boca de Barranca
Bonilla 1 male.

October 16, 1981 Dickerman 3
MALES

|. P. CAROLYNAE ——_

|. R PECTORALIS
|. PR, GUTTULATUS

“ANTHONY! —
MOTAGUA AND ———
NEGRO VALLEYS

HONDURAS

NICARAGUA a
|. P, ESPINACHI a

90 95 100 105 ite) NS

FEMALES

P. CAROLYNAE _
I. P, PECTORALIS —|—___
P.

. GUTTULATUS
TYPE

x
"ANTHONY' maga —
MOTAGUA AND a

NEGRO VALLEYS

HONDURAS imearDgaaetes —
NICARAGUA XX

I. P, ESPINACHI —

Fig. 1. Statistical analysis of wing chord measurements (rounded to nearest
mm) for subspecies and populations of Icterus pectoralis. For each
sample the range, mean (vertical line), and one standard deviation on
either side of the mean (solid bar) are presented. See Table 1 for
sample sizes.

4 NEMOURIA No. 26
MALES

|. P, CAROLYNAE TT

|. PB, PECTORALIS

|. P. GUTTULATUS

"ANTHONYI" a
MOTAGUA AND ae

NEGRO VALLEYS

HONDURAS

NICARAGUA See RENE
|. P. ESPINACHI ———

85 90 95 100 105 Ke) 15
FEMALES

|. P. CAROLYNAE a
|. P. PECTORALIS —+
|. P, GUTTULATUS

TYPE x

"ANTHONY" a

MOTAGUA AND me -

NEGRO VALLEYS

ERRORS =
NICARAGUA xX

|. P. ESPINACHI a5

85 90 95 100 105 Te) NS

Fig. 2. Statistical analysis of tail measurements (rounded to nearest mm) for
subspecies and populations of Icterus pectoralis. The format and
symbols are as with Figure 1.

October 16, 1981

Table 1. Measurements (mm) of subspecies and populations of Icterus
pectoralis, with mean and standard deviation.

Ip. carolyneae

Lp. pectoralis

Lp. guttulatus

type specimen

“anthonyi”

Motagua and

Negro Valleys

Honduras

Nicaragua

Lp. espinachi

Dickerman

Males

Wing Tail
100-111 100-110
106.3 (3.0) 103.6 (2.5)
n=17 n=14
108-113 103-109
109.6 (1.8) 105.5 (2.6)
n=7 n=6
100-112 96-106
105.0 (2.9) 101.3 (3.4)
n=20 n=20
105-114 104-112
109.6 (3.2) 108.2 (3.8)
n=5 n=5
108-114 103-110
109.4 (1.9) 106.2 (2.8)
n=11 n=11
103-113 96-112
108.8 (4.2) 105.0 (6.8)
n=5 n=5
98-106 94-102
103.5 (2.9) 97.2 (3.3)
n=10 n=10

>

Females
Wing Tail

97-100 95-97
97.8 (1.30) 95.6 (0.8)
n=6 n=5
98-106 98-101
101.5 99.6
n=4 n=3
101 90
95-105 89-101
98.6 (3.6) 95.2 (3.1)
n=20 n=19
99-106 95-106
103.6 (2.7) 101.4 (4.4)
n=5 n=5
102-113 103-112
106.4 (4.8) 106.0 (4.6)
n=17 n=12
102 & 103 100 & 100
n=2 n=2
90-100 89-93
95.7 (3.3) 91.2 (1.3)
n=6 n=

6 NEMOURIA No. 26

Icterus pectoralis guttulatus Lafresnaye. Mag. Zool. no. 2, vol. 6, pl.
52, 1844. Type locality: ““Mexico” = error, I designate San Jose, Dept.
Escuintla, Guatemala. Holotype: adult sex unknown, MCZ 76120
(examined).

Icterus pectoralis anthonyi Griscom. AMNH Novit. 438:18, 1930.
Type locality: Finca El Cipres near Ocos, Dept. San Marcos, Guatemala.
Holotype: adult male, AMNH 398802 (examined).

DIAGNOSIS: Resembling I. p. espinachi but more intensely colored on head
and nape, rump and breast (orange rather than orange-yellow). Size variable
(see Fig. 1, Table 1). Spotting as in espinachi (Fig. 3).

RANGE: Central (Matagalpa) and northwestern (Chinindega) Nicaragua
the more mesic foothills to Rancho Sol y Luna, Oaxaca. Also in the arid
interior of Honduras and in Guatemala in the arid portions of the drainages of
the Rios Motagua and Negro.

DISCUSSION: Lafresnaye’s type of I. guttulatus is an unsexed adult in
moderately worn plumage now in the Museum of Comparative Zoology. It was
compared with near topotypes of espinachi (from Bolson and Bebedero, Costa
Rica) and a series of “anthonyi” including paratypes from Guatemala. The
holotype is inseparable in color or size from Guatemalan specimens and
differs from nominate pectoralis, as does the Guatemalan series, and as do
being more orange. Allan R. Phillips pointed out to me that the holotype of
guttulatus undoubtedly came from Guatemala rather than Mexico. The
Pacific lowlands of Chiapas, the only area in Mexico in which the specimen
could have been taken, were essentially inaccessable during the period in
which the holotype must have been collected (1830-1842). There are no
ornithological collections from that region of Chiapas during that time period.
Ixtapa (then spelled Ystapa) was the major Pacific port for Guatemala up to
1853, and the road from Ixtapa to Guatemala City passed close to the present
day port of San Jose (see Dickerman, 1981). At least one major collection, that
reported by Bonaparte (1837), was made on the Pacific lowlands and slope of
Guatemala during the 1830’s.

Within the range herein ascribed to guttulatus, specimens from Guate-
mala and southern Chiapas are the smallest, while birds from interior
Honduras and the Rio Motagua Valley average larger. Nicaragua specimens
are intermediate in size. Were a better series of measurable adults from
Honduras and Nicaragua available, additional overlap with “anthonyi” would
probably be shown. There is no variation in color among those populations.
There is thus no justification for formal subspecific separation of the somewhat
larger Honduras birds.

October 16, 1981 Dickerman 7

SPECIMENS EXAMINED: 104. Nicaragua 5 males, 2 females; Hon-
duras 11 males, 17 females; E] Salvador 1 male, 1 female; Guatemala 35
males, 19 females; Chiapas 5 males, 5 females; Oaxaca 1 male, 2 females.

Icterus pectoralis pectoralis Wagler. Isis, 1829, col. 755. Type
locality: “Mexico” (= Totolapan, Oaxaca).

DIAGNOSIS: Similar to I. p. guttulatus but paler on crown, venter and rump
(yellowish orange rather than orange). Spotting as in guttulatus and espinachi
(Fig. 3).

RANGE: Restricted to the arid Pacific lowlands of the Isthmus of Tehuantepec,
south to Tonala, Chiapas.

Fig. 3. Four subspecies of Icterus pectoralis. Left to right: I.p. espinachi,
male, typical coloration UMMZ 133347, male extreme coloration
UMMZ 150665; Ip. guttulatus, male UMMZ 103951; Lp. pectoralis,
male UMMZ 139230; and Ip. carolynae, male DEL 30833, female
DE! 30832.

8 NEMOURIA No. 26

DISCUSSION: In the original description, the only location given was
*““Mexico”’. Stresemann (1954) provided the provenience of many of Ferdinand
Deppe’s specimens on which names have been based. ““Totolapa (Oaxaca),
October, 1825” were the data Stresemann provided for the type of I.
pectoralis. Binford (pers. comm.) was unable to ascertain the location of
Totolapa; but, during late October and November 1825, Deppe collected at
San Bartolo (50 mi W of Tehuantepec), at Tehuantepec, and at San Mateo on
the coast south of Tehuantepec (Stresemann 1954, p. 87). The Tehuantepec
region may be thus considered to be the type locality of I. pectoralis.

A specimen (WF 16971) from Rancho Sol y Luna, Oaxaca east of the
Isthmus of Tehuantepec at the base of the foothills (elev. 800’) is as brightly
colored as Chiapas specimens of guttulatus while a specimen from Tonala,
Chiapas (AMNH 521929) is nominate pectoralis in color. Specimens from
Oaxaca west of the Isthmus of Tehuantepec (and from extreme southern
Guerrero) are variably intermediate between nominate pectoralis and the
following subspecies. They are cited under the subspecies described below.

SPECIMENS EXAMINED: 11. Oaxaca: Tehuantepec 3 males, 1 female;
Tapanatepec to 9.5 miles west Zanatepec 3 males, 2 females; San Gabriel
Mixtepec Juquila 1 male. Chiapas: Tonala 1 female.

- Icterus pectoralis carolynae new subspecies. Holotype: DEL 30832,
adult female, collected on the east shore of Laguna de Tres Palos, east of
Acapulco, Guerrero, Mexico, on 17 December 1970 by Sostenes
Romero H; from the collection of Allan R. Phillips.

DIAGNOSIS: Coloration of head, rump and venter pale as in pectoralis,
orange-yellow rather than orange as in comparable areas of guttulatus; slightly
paler in those areas than espinachi. Spotting of breast more extensive, both in
total area and in size of individual spots. These coalesce in some individuals to
form a band connecting with the black of the mantle (Fig. 3).

RANGE: Known only from the Pacific slope of Guerrero and adjacent
Oaxaca.

ETYMOLOGY: It is a pleasure to name this beautiful subspecies for my wife
Carolyn Lyell (Campbell) Dickerman.

DISCUSSION: Five specimens in the Moore Laboratory collection from
Cuajinicuilapa in extreme southern Guerrero are nearer the nominate
subspecies, while four of seven specimens from the Pacific slope of Oaxaca in
the LSU collection are inseparable from carolynae. Late nesting season
specimens of carolynae (August) have spotting reduced through wear.

SPECIMENS EXAMINED: 21. Guerrero: Laguna Tres Palos 3 males, 1
female; Papayo 2 males, 1 female; Ejido Nuevo 2 males; Dos Arroyos 1 male,

October 16, 1981 Dickerman 9

1 female; Joluchuea (= SE of Petalan, SW Guerrero) 2 males, 1 female;
Acupulco 1 male; 28 miles west of Pie de la Questa 1 male; Las Posas (7 miles
north of Coyuca ) 2 males, 4 females; Atoyac de Alvarez 1 female.

Intermediate populations I. p. pectoralis x I. p. carolynae 12. Guerrero,
Cuajinicuilapa 1 female, 4 males. Oaxaca, 6 and 16 miles northwest of Puerto
Escondido 2 males; Minitan (22 roadmiles south of Pinotepa) 1 female, 1
male; 11 miles north of Pochutla 1 female, 2 males. Measurements of
specimens from Oaxaca in the LSU collection are included in the series of
carolynae presented in Table 1 and Figures 1 and 2.

Acknowledgements

I wish to acknowledge the curators of the following collections (abbreviations
refer to those used in the text): American Museum of Natural History
(AMNH); Carnegie Museum of Natural History (CM), Cornell University;
Delaware Museum of Natural History (DEL); Denver Museum of Natural
History; Louisiana State University Museum of Zoology (LSU); Museum of
Comparative Zoology, Harvard University (MCZ); U.S. National Museum
of Natural History (USNM); Peabody Museum, Yale University; University
of Michigan Museum of Zoology (UMMZ), Western Foundation of Vertebrate
Zoology (WF), and Moore Laboratory of Zoology, Occidental College.
Travel to examine specimens was supported in part by the Frank M. Chapman
Memorial Fund.

Allan R. Phillips was an invaluable source of expertise. David M. Niles,
Joe T. Marshall, H. Douglas Pratt, and Joel L. Cracraft provided kind
hospitality during the course of this study.

Some material used in this study was collected during the course of
studies on the ecology of arboviruses in Guatemala under permits from the
Ministerio de Agricultura de Guatemala. That research was supported in part
by U.S. Public Health Service Research Grant AI-06248.

I am grateful to the American Museum of Natural History for preparing
the figures.

10 NEMOURIA No. 26
LITERATURE CITED

Blake, E. R.
1968 Icteridae, in Check-list of birds of the World, Vol. XIV. (R.A.
Paynter, Jr., Ed.) Museum Comp. Zool., Cambridge, Mass.

Bonaparte, C.L.J.L.
1837 Report on a collection of birds formed by Colonel Velasquez de
Leon during a fortnight’s scientific tour in Guatemala. Proc. Zool.
Soc. Lond., pp. 114-119.

Dickerman, R. W.

1981 Preliminary review of the Clay-colored Robin (Turdus grayi) with
redesignation of the type locality of the nominate form and
description of a new subspecies. Bull. Brit. Ornith. Club,
101:285-290.

Griscom, L.
1930 Studies from the Dwight Collection of Guatemala birds. ITI. Amer.
Mus. Nat. Hist. Novitates no. 438, pp. 1-18.

Hellmayr, C. E.
1937 Catalogue of birds of the Americas and adjacent islands. Part 10.
Field Mus. Nat. Hist., publ. 381, Zool. Ser. no. 13, pp. i-v, 1-228.

Monroe, B. L.
1968 A distributional survey of the birds of Honduras, Amer. Ornith.
Union Monogr., 7:1-458.

Stresemann, E.
1954 Ferdinand Deppe’s travels in Mexico in 1824-1829. Condor,
56:86-92.

3

ee

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
Kennett Pike, Greenville,
Delaware 19807

OL.
C//
NM Y¥3L
ae Be,

NEMOURIA

Occasional Papers of the Delaware Museum of Natural History

NUMBER 27 APRIL 29, 1983

A REVIEW OF THE COLUMBARIINAE
(GASTROPODA: TURBINELLIDAE)
OF THE WESTERN ATLANTIC WITH NOTES
ON THE ANATOMY AND SYSTEMATIC
RELATIONSHIPS OF THE SUBFAMILY |

M.G. Harasewych*

INTRODUCTION

The subfamily Columbariinae encompasses a small, poorly known,
morphologically compact group of fusiform marine rachiglossan gastropods
that inhabit areas of the outer continental shelf and upper continental slope.
Known from the Upper Cretaceous (Maestrichtian) of the Netherlands, the
group first appears in the western Atlantic during the Paleocene (Midway
Group of Alabama), but is not represented in the fossil record of the region
after the Eocene.

The group’s taxonomic placement within the Neogastropoda has
undergone considerable peregrination. The status of the seven supraspecific
taxa has also been subject to varying interpretations.

Anatomical investigations have, for the most part, been limited to studies
of radular morphology. Ponder (1973) discussed the alimentary system of two
species of Coluzea in a paper dealing with the evolution of the Neogastropoda.

On the basis of new data provided by anatomy, diet, SEM micrographs of
radulae, and shell characters, I suggest that the group does not merit family
status, but rather, is part of the vermivorous turbinellid radiation that took
place in the late Mesozoic. A revision of the western Atlantic species is
proposed, three new species are described, and the evolution and zoogeo-
graphy of the subfamily are discussed.

*College of Marine Studies Library of Congress
University of Delaware Catalog No. 83-71128
Newark, Delaware 19711

2 NEMOURIA No. 27
Acknowledgments

I am grateful to the following people, who have assisted me with
information and the loan of types and other material: W.K. Emerson,
American Museum of Natural History, New York; R.S. Houbrick,
J. Rosewater, National Museum of Natural History, Smithsonian Institution,
Washington, D.C.; R.H. Jensen, Delaware Museum of Natural History,
Greenville; P.M. Mikkelsen, Indian River Coastal Zone Museum, Fort
Pierce, Florida; R. Robertson, Academy of Natural Sciences of Philadelphia;
R.D. Turner, Museum of Comparative Zoology, Harvard; G.L. Voss,
Rosenstiel School of Marine and Atmospheric Sciences, University of Miami;
R. and D. Janowsky; E.J. Petuch; C. Staples.

I thank G.L. Gruber of the College of Marine Studies, University of
Delaware for his help with the histological preparations, and Drs. M.L. Jones
and K. Fauchald, National Museum of Natural History, Smithsonian
Institution, for identification of polychaete setae. S.D. Kaicher kindly
provided the photograph of the holotype of Fulgurofusus sarissophorus.
Special thanks are due G. Kris Jensen for preparation of the drawings and to
E.K. Wilson, Engineering Test Center, E.I. du Pont de Nemours Co., for her
help with some of the scanning electron microscopy.

I am indebted to Drs. M.R. Carriker, W.K. Emerson, R.S. Houbrick,
E.J. Petuch, and anonymous reviewers for critical reading of the manuscript.

TAXONOMIC HISTORY

Although several species referable to Columbariinae had been previously
described in the genus Fusus (Lesson, 1834; Deshayes, 1835), Martens
(1881) was the first to recognize the group as distinct. He proposed
Columbarium as a subgenus of Pleurotoma |type species Pleurotoma
(Columbarium) spinicincta Martens, 1881] based on Schacko’s (in Martens,
1881) identification of the long, tubular structures taken from the proboscis of
the type species as toxoglossan radular teeth. Some authors accepted this
placement (Fischer, 1883; Tryon, 1884; Martens 1901; Dall, 1918), but
others either retained Fusus (Watson, 1882; Tate, 1888) or placed
Columbarium within Fusidae (=Fusininae) (Cossmann, 1896, 1901).

In 1922 Peile reported that the radula of Columbarium pagoda (Lesson,
1834) was rachiglossate and placed Columbarium near the Muricidae,
further suggesting that Schacko’s figures represented crustacean remains.
Thiele (1925) reexamined Schacko’s slide and concluded that the objects on it
were not radular teeth. After examining the radulae of two additional species,
he placed the genus Columbarium in the Muricidae between Typhis and
Murex.

April 29, 1983 Harasewych 3

Tomlin (1928) erected the family Columbariidae on grounds that, while
sharing some characters with Muricidae and Buccinidae, Columbarium was
not referable to either of these families. The following year, Thiele reduced
Columbariidae to a genus within Muricidae. Finlay (1930) recognized the
family, and included in it the genera Fulgurofusus [erected by Grabau (1904)
for a Paleocene species he believed to be ancestral to Busyconinae] and
Coluzea (a fossil and Recent New Zealand genus that Finlay had proposed in
1927).

With the exception of Wenz (1941), who considered the Columbariinae a
subfamily of Muricidae, subsequent workers have accorded the group family
level status (Iredale, 1936; Finlay and Marwick, 1937). Clench (1944)
reviewed the newly discovered western Atlantic species, Dell (1956) was the
first to describe the radula of Coluzea, and Barnard (1959) added to our
knowledge of the South African species.

In a major revision of the family, Darragh (1969) proposed three new
genera: Hispidofusus, Serratifusus and Histricosceptrum, the first two from
the Miocene of Australia, the last from the Recent of the western Atlantic and
Indian Oceans. Reporting on the mollusks collected during the University of
Miami deep-sea expeditions, Bayer (1971) reduced all supraspecific taxa to
subgenera of Columbarium, and added the subgenus Peristarium to include 3
new species that strongly resemble Fusinus, but have distinctly columbariid
radulae and protoconchs. Ponder (1973) showed that the alimentary system of
Coluzea possessed muricid as well as buccinid features, and concluded that
family status was justified.

MATERIALS AND METHODS

I was fortunate in being able to examine most of the known specimens of
western Atlantic Columbariinae, including all holotypes and all but 3
paratypes. Among the material collected by R/V JOHN ELLIOT PILLSBURY
were several preserved specimens of Fulgurofusus brayi. Most had been
partially dissected and the radulae had been removed, but there remained a
single intact specimen still in its shell. Several fragments of this shell were
broken off near the outer lip, in order to examine shell ultrastructure. The
specimen was then immersed in 10% hydrochloric acid until the shell
dissolved. The operculum was removed and the animal was imbedded in
polyester resin, sectioned at 8 pm and stained with Mallory Triple Connective
Tissue Stain.

Specimens of other species that contained dried animals were reconsti-
tuted in a solution of trisodium phosphate, and gross features of their mantle
cavities, and reproductive and alimentary systems were studied before
extracting the radulae.

4 NEMOURIA No. 27

Radulae and shell ultrastructure were examined using a Zeiss Novascan
30 Scanning Electron Microscope.

With the exception of the holotype of Fulgurofusus sarissophorus, all
shells were coated with ammonium chloride before being photographed.

Abbreviations used within the text are as follows: AMNH, American
Museum of Natural History; BM(NH), British Museum (Natural History);
DMNH, Delaware Museum of Natural History; IRCZM, Indian River
Coastal Zone Museum; MCZ, Museum of Comparative Zoology; UMML
University of Miami Marine Laboratory; USNM, former United States
National Museum, collections in National Museum of Natural History,
Smithsonian Institution.

SYSTEMATICS

Order NEOGASTROPODA
Family TURBINELLIDAE Swainson, 1840

This family, best known for its large, shallow-water members, is
characterized by thick, spindle-shaped shells that have large, bulbous
protoconchs, distinct, sculptured peripheries, and open, axially-oriented
siphonal canals. Shell aperture is generally ovate, the outer lip smooth; and the
columella may or may not have several spiral folds. Opercula are corneous,
elongate, and have a terminal nucleus. Animals have along, narrow proboscis,
a small radula with tricusped rachidia, and an open or partially fused sperm
groove; they lack an accessory salivary gland; and they may or may not have
an anal gland and multicusped lateral teeth. Diet consists largely or
exclusively of polychaetes (Moses, 1923:106) and sipunculids (Taylor et al.
1980:377).

The nomenclatoral history of this family has been discussed by Rehder
(1967:184-185). Current workers divide the family into the subfamilies
Turbinellinae, Vasinae and Ptychatractinae, largely on the basis of shell and
radular characters. Evidence is presented here for the inclusion of an
additional subfamily, the Columbariinae.

Subfamily COLUMBARIINAE Tomlin, 1928

This subfamily, restricted to the bathyal zone in the Recent fauna, may be
recognized by its small, thin, fusiform shell, which has a deviated, paucispiral
protoconch, sculptured periphery, long, narrow siphonal canal and lacks
columellar folds. The distinctive operculum is strongly ovate, with terminal
nucleus and round or ovate attachment area. Anatomical features include a
very long, thin proboscis that retracts into a non-evertable sheath, a tiny radula
with tricusped rachidia and long, monocusped lateral teeth, an open sperm
groove that extends to the tip of the penis, and a distinct anal gland. Rectal
contents indicate a diet of tubiculous polychaetes.

April 29, 1983 Harasewych 5

As only limited anatomical material was available for study, diagnoses of
phyletic relationships between supraspecific taxa are tentative. Recognition of
at least two genera, Fulgurofusus and Columbarium, both well established by
the Paleocene, appears to be warranted on the basis of differences in shell and
radular morphology. Although both genera are represented in Tertiary
deposits along the northern Gulf of Mexico, all Recent western Atlantic
columbarines are attributed to Fulgurofusus s.1. The genus Columbarium
appears to be restricted to the western Pacific in the Recent fauna. Coluzea
and Peristarium are provisionally regarded as subgenera of Fulgurofusus,
based on a general resemblance of shells and radulae, but additional work is
required to more fully delineate the true affinities of these groups.

Cursory examination of the radulae of several South African species
originally described in Columbarium suggests that they will prove to belong to
Fulgurofusus.

Genus Fulgurofusus Grabau, 1904
SYNONYMY
1904 Fulgurofusus Grabau, Smithson. Misc. Collins. 44 (1407): 86.

Type species by original designation: Fusus quercollis Harris,
1896.

1971 Columbarium (Fulgurofusus) Grabau, Bayer, Bull. Mar. Sci.
21(1):170.

This genus was originally proposed in Fusidae (= Fusininae) to include two
Tertiary species that Grabau (1902:922; 1904:87; 1910:11-14) believed to
be ancestral to Busyconinae. One of these species was subsequently transfered
to the genus Columbarium, which differs from Fulgurofusus in having a more
bulbous and deviated protoconch, and a thicker shell, with a strong anterior
carina, triangular spines along the periphery, and scales along the spiral cords.
Axial ribbing, an incised or keeled periphery, and a rachidian in which the
edges of the basal plate are greatly expanded and buttressed are all
characteristic of Fulgurofusus.

Subgenus Fulgurofusus s.s.

This subgenus is represented in the fossil record of the western Atlantic by 3
species: F. quercollis, from the Paleocene of Alabama, F. merriami
(Dickerson, 1916), from the Lower Eocene of California, and F. washing-
tonianus (Weaver. 1912), from the Upper Eocene of Washington. Species in
the Recent fauna inhabit bathyal and abyssal zones along the western Atlantic
as well as off Antarctica and New Zealand. Fulgurofusus (Fulgurofusus)
benthocallis (Melville and Standen, 1907), taken in 3210 m off the South
Orkney Islands, holds the depth record for the subfamily. The flange-like
peripheral keel, which may be crenulate or nodulose, is diagnostic of the
subgenus. Axial sculpture, when present, is limited to weak ribs or nodes in the
early whorls.

6 NEMOURIA No. 27

Fulgurofusus (Fulgurofusus) brayi (Clench, 1959)
Figures 1-2, 4-9, 13-20
SYNONYMY

1959 Columbarium brayi Clench, Johnsonia 3(39):330, pl. 173,
figs. 1,2.
1969 Fulgurofusus brayi (Clench), Darragh, Proc. Roy. Soc. Vict.
83(1):102-103, pl.6, figs. 113,115.
1971 Columbarium (Fulgurofusus) brayi Clench, Bayer, Bull. Mar. Sci.
21(1):173-176, figs. 39 B, 40 A-D; 1974 Abbott, American
Seashells, second edition, p.170.
DESCRIPTION — SHELL: Large for genus (to 73 mm), thin to moderately
heavy, fusiform; spire angle 53°-109°; protoconch (fig.20) with 1% whorls,
bulbous, glassy; transition to teleoconch indistinct, marked by gradual
acquisition of peripheral keel and increase in coarseness of growth striae;
teleoconch with 7 strongly convex whorls; suture impressed; keel, rounded in
first post-nuclear whorl, rapidly extends normal to the coiling axis to form a
peripheral flange that may be short or nearly as wide as the aperture; spiral
sculpture of fine threads to strong cords, usually most prominent adjacent to
peripheral keel, and on body whorl and siphonal canal; axial sculpture limited
to fine sinuate growth striae; aperture strongly ovate; outer lip smooth,
unornamented, except for furrow beneath periphery; inner lip smooth,
appressed posteriorly; siphonal canal long, straight, axial; shell and aperture
color uniformly white; periostracum (fig.8) very thin, yellowish, with short
axial blades.

ULTRASTRUCTURE: shell composed of 2 orthogonal layers of crossed-
lamellar aragonite; outer layer (fig.7,XLC) with lamellar planes parallel to
growing edge, fairly uniform in thickness (150-300 pm), includes spiral
sculpture; inner layer (fig.7,XLT) with lamellar planes perpendicular to

FIGS. 1-2. Fulgurofusus (Fulgurofusus) brayi (Clench, 1959).
1. AMNH 187174, Off Venezuela, in 342 m(1 X).
2. Holotype, MCZ 221601, ATLANTIS cruise 240, Off Cabo Codera,
Venezuela, in 273 m(1 X).

FIG. 3. Fulgurofusus (Fulgurofusus) sarissophorus (Watson, 1882).
Holotype, BM(NH) 1887.2.9.687, CHALLENGER station 122, Off
Pernambuco, Brazil, in 640 m (3 X).

FIGS. 4-6. Fulgurofusus (Fulgurofusus) brayi (Clench, 1959).

4. IRCZM 65:1806, OREGON II station 24263, Off Nevis, Lesser
Antilles, in 512-769 m(1 X).

5. DMNH 153522, PILLSBURY station P-445, ESE of Escudo de
Veraguas, Panama, in 342-346 m (1.5 X).

6. USNM 806987, PILLSBURY station P-781, Off Eastern Colombia, in
567-531 m(1.5 X).

April 29, 1983 Harasewych

8 NEMOURIA No. 27

growing edge, variable in thickness (20-150 pm), thinnest at outer edge of lip,
thickest along columella and inner surface of periphery.

ANIMAL — EXTERNAL FEATURES: The body consists of 24% whorls, of which
the mantle cavity occupies % whorl, the kidney (fig.14,k) % whorl, and the
digestive gland (fig.14,dg) about % whorl. The columellar muscle is large, with
a broad attachment area. The foot is long, narrow, rounded anteriorly and
posteriorly, with a yellow, sharply ovate operculum. Tentacles are short and
blunt, with conspicuous black eyes. The mantle edge is thin and slightly
papillose, the siphon long and narrow.

MANTLE CAVITY: The mantle cavity is narrow and deep, with the pallial
organs situated as in other rachiglossans (fig.14). The short (4 mm), broad
(L/W = 2.2) osphradium (fig.14,os) contains 48 leaflets per side. The
ctenidium (fig. 14,ct), twice as long and slightly narrower than the osphradium,
is composed of about 150 triangular leaflets. To the right of the transversely
folded and unpigmented hypobranchial gland (fig.14,hg) lie the rectum
(fig.14,r) and genital ducts. The rear of the mantle cavity is formed by the
pericardium (fig.14,pc) on the left and the kidney on the right.

ALIMENTARY SYSTEM: The extremely long, narrow proboscis (fig.15,p)
retracts into a non-evertable, muscular proboscis sheath (figs.14,15,ps) that
extends from just behind the rhyncostome to nearly the rear of the cephalic
sinus. It is estimated that the length of the everted proboscis would roughly
equal the shell length. The mouth is situated at the tip of the proboscis, with a
small buccal mass positioned just behind the buccal cavity. The radular ribbon
is short (800 pm) and narrow (75 pm), with about 80 rows of teeth. The
rachidian is broad, with 3 conical cusps compressed toward its center. Lateral
teeth have a single, scythe-shaped cusp. As the only specimen with a radula

FIGS. 7-8. Details of shell structure of Fulgurofusus (Fulgurofusus) brayi.
7. SEM micrograph of fracture surface. Plane of fracture perpendicular
to the outer lip. Scale bar = 100 pm.
8. SEM micrograph of periostracum. Scale bar = 100 pm.
FIG. 9. SEM micrograph of radular ribbon of Fulgurofusus (Fulgurofusus)
bermudezi, taken from specimen shown in figure 25. Scale bar =
10 pm.
FIG. 10. SEM micrograph of radular ribbon of Fulgurofusus (Histroco-
sceptrum) atlantis, taken from the holotype (figure 32). Scale bar =
10 pm.
FIG. 11. SEM micrograph of radular ribbon of Columbarium (Columbarium)
spinicinctum, AMNH 180318. Scale bar = 10 pm.
FIG. 12. SEM micrograph of radular ribbon of Fulgurofusus (Peristarium)
aurora, taken from specimen shown in figure 52. Scale bar = 10 pm.

PER, periostracum; XLC, crossed lamellar aragonite — collabral;
XLT, crossed lamellar aragonite — transverse.

April 29, 1983 Harasewych 9

10 NEMOURIA No. 27

~ BERMUDA

GULF OF MEXICO

ie
t's ane
bermudezi Y (li

ee ae

JAMAICA

: 6 BARBADOS

@ ecphoroides

sarissophorus/w

FIG. 13. Geographic distribution of the western Atlantic species of Fulguro-
fusus S.S.

was serially sectioned, a SEM examination of the radula was not possible.
Bayer (1971:172, fig. 39 C) figured the radula of this species. A section
through the proboscis just behind the buccal mass (fig.16) reveals that the
ducts of the salivary glands (fig. 16,dsg) are imbedded beneath the dorsal folds
(fig.16,df). A thin layer of transverse muscle (fig.16,tms) separates the
esophagus (fig.16,es) from the nerves (fig.16,n) and blood vessel (fig. 16,bv)
that run the length of the proboscis. From the rear of the proboscis, the
esophagus runs anteriorly, enlarging to form the valve of Leiblein (fig. 15,vl)
before passing through the nerve ring (fig.15,nr). Behind the nerve ring lie 2
large asymmetrical salivary glands (fig.15,sg), the larger right gland situated
anterior to the smaller left. The gland of Leiblein (fig.15,gl) lies behind the
salivary glands and to the left of and partially under the proboscis sheath. After
leaving the cephalic sinus, the esophagus enters the stomach (fig.15,sto),
which lies along the dorsal surface of the digestive gland, just behind the
kidney. Longitudinal folds run the length of the U-shaped stomach and into the
intestine, which expands into a broad, spacious rectum (figs.14,15,r) after
entering the mantle cavity. Dorsal to the anterior third of the rectum lies the
anal gland (fig.15,ag). The rectum constricts and detaches from the roof of the
mantle cavity about 1 mm before terminating in the anus (fig.14,15a).

April 29, 1983 Harasewych 11

MALE REPRODUCTIVE SYSTEM: The testis (figs. 14,17,te) is situated along the
right side of the digestive gland, just behind the kidney. It consists of a mass of
tubules that converge on a testicular duct (fig.17,td) that runs anteriorly along
the right ventral side of the visceral mass. Upon reaching the posterior portion
of the kidney, this duct doubles in diameter and becomes convoluted, serving
as the seminal vesicle (fig.17,sv). The anterior portion of the seminal vesicle
runs along the pericardium (fig.17,pc), but no gonopericardial connection
could be detected. Anterior to the pericardium, the duct again constricts to
form the vas deferens (fig.17,vd). This non-glandular, closed duct enters the
mantle cavity along its right wall, just below the rectum, descending to its floor
at midlength, where it becomes a muscular, open groove that runs anteriorly
along the mantle cavity and along the inner edge of the broad, flattened penis
(figs.14,17,pen) to its rounded distal end. Prostatic tissue (figs.18,19,pr) lines
this groove along its entire length.

ETYMOLOGY: Named after W. Scott Bray, Captain of the ATLANTIS.

TYPE SPECIMENS: Holotype — MCZ 221601, length 49.1 mm; Paratype —
MCZ 221602, length 39.2 mm.

TYPE LOCALITY: Off Cabo Codera, Venezuela, 11°00’N,66°0O1’W, in 273 m.

GEOGRAPHIC DISTRIBUTION (fig.13): This is the most widely distributed
Fulgurofusus. It has been collected off Nevis in the Lesser Antilles and along
the southern Caribbean Sea, from eastern Venezuela to Panama.

ECOLOGY: Fulgurofusus brayi inhabits mud bottoms at depths ranging from
273 to 769 m, with a mean station depth (n = 13) of 444 m. Rectal contents
include a large number of polychaete setae attributed to the families
Serpulidae and Chaetopteridae by Drs. M.L. Jones and K. Fauchald of the
National Museum of Natural History, Smithsonian Institution.

FIGS. 14-19 (following pages). Anatomical features of Fulgurofusus
(Fulgurofusus) brayi.

14. Male specimen removed from shell and partially uncoiled, with mantle

and mantle cavity floor opened by mid-dorsal incisions.

15. Alimentary system.

16. Transverse section through the proboscis behind the buccal mass.

17. Male reproductive system.

18. Transverse section through the anterior vas deferens.

19. Transverse section through the middle of the penis.

12 NEMOURIA No. 27

)

Hl

AN

So,

a, anus; ag, anal gland; bv, blood vessel; cm, circular muscle; ct, ctenidium; df,
dorsal fold; dg, digestive gland; dsg, duct of salivary gland; es, esophagus; gl,
gland of Leiblein; hg, hypobranchial gland; k, kidney; ko, kidney opening; Im,
longitudinal muscle; mc, mucous cells; n, nerve; nr, nerve ring; op, operculum;
os, osphradium; p, proboscis; pc, pericardium; pen, penis; pr, prostate gland;
ps, proboscis sheath; r, rectum; rm, retractor muscles; s, siphon; sg, salivary
glands; sgr, seminal groove; sto, stomach; sv, seminal vesicle; td, testicular
duct; te, testis; tms, transverse muscular sheet; vd, vas deferens; vl, valve of
Leiblein.

April 29, 1983

Harasewych

13

14 NEMOURIA No. 27

MATERIAL EXAMINED: OREGON II sta. 24263, Off Nevis, Lesser Antilles,
17°09’N,62°44.8°W, in 512-769 m (IRCZM 65:1806); Dredged off
Venezuela, in 342 m(AMNH 187184); Off Venezuela, in 366 m (AMNH
195603); Off Cabo Codera, Venezuela, 11°00’N, 66°01’ W, in 273 m(MCZ
221601 — holotype; MCZ 221602 — paratype); PILLSBURY sta. P-340, Off
Punto Mosquito, Panama, 9°14’N,77°46’W, in 307-366 m (UMML);
PILLSBURY sta. P-374, NW of Golfo de Morrosquillo, Colombia,
9°57'N, 76°11’W, in 439-377 m (UMML); PILLSBURY sta. P-386, Off
Cartagena, Colombia, 10°30’N,75°42’W, in 275-357 m (UMML);
PILLSBURY sta. P-394, WNW of Golfo de Morrosquillo, Colombia,
9°29°N,76°26’W, in 421-641 m(UMML; USNM 806988); PILLSBURY sta.
P-445, ESE of Escudo de Veraguas, Panama, 9°02’N,81°24’W, in 342-346
m (UMML; DMNH 153522); PILLSBURY sta. P-447, Golfo de los
Mosquitos, Panama, 9°02’N, 81°07’ W, in 664-681 m(UMML); PILLSBURY
sta. P-753, Off Venezuela, 11°18.8’N,68°22’W, in 384-607 m (UMML);
PILLSBURY sta. P-776, Off eastern Colombia, 12°13.3’N,72°50’W, in 408-
576 m (UMML); PILLSBURY sta. P-781, Off eastern Colombia, 11°30.1’N,
73°26.5’W, in 567-531 m (UMML; USNM 806987).

MEASUREMENTS OF SHELL CHARACTERS (n = 10):

spire angle

mean 41.8 mm 69.8°
range 28.5 — 73.0 53° — 109°
sd 12.9 18.4°

COMPARATIVE REMARKS: Although F. brayi is the most variable and widely
distributed member of the subgenus, it can readily be distinguished from all
other species by its large size, broad spire, prominent peripheral keel and lack
of axial sculpture, even in early whorls. Specimens taken off Venezuela tend to
be more globose and have thicker shells (figs.1,2), while those from the
western Caribbean often have depressed spires (fig.5).

FIG. 20. Protoconch of Fulgurofusus (Fulgurofusus) brayi (Clench, 1959).
USNM 806988.

FIG. 21. Protoconch of Fulgurofusus (Fulgurofusus) bermudezi (Clench
and Aguayo, 1938). DMNH 153521.

FIG. 22. Protoconch of Fulgurofusus (Fulgurofusus) ecphoroides new
species. Holotype (figure 27).

Scale bar = 1 mm for all drawings.

April 29, 1983

Harasewych

15

16 NEMOURIA No. 27

Fulgurofusus (Fulgurofusus) sarissophorus (Watson, 1882)
Figures 3,13
SYNONYMY

1882 Fusus (Colus) sarissophorus Watson, J. Linn. Soc. Lond.
16:382.

1886 Fusus sarissophorus Watson, Watson, Challenger Report Zoology
15(2):196-197,p1.14, fig.1.

1944 Columbarium sarissophorum (Watson), Clench, Johnsonia
1(15):1-2,p1.1,fig.1; 1975 Rios, Brazilian Marine Mollusks
Iconography, p.77,p1.21,sp.313.

1969 Fulgurofusus sarissophorus (Watson), Darragh, Proc. Roy.
Soc. Vict. 83(1):102,p1.6,figs. 107,112.

DESCRIPTION: Shell small (19 mm), thin, fusiform; spire angle 53°; proto-
conch glassy, bulbous, with 1% whorls; transition to teleoconch indistinct;
teleoconch with 5 convex whorls; suture impressed; peripheral keel rounded in
transition zone, extending into a short flange normal to the coiling axis in the
first post-nuclear whorl; spiral sculpture of fine threads more prominent
anterior to keel; axial sculpture of sinuate growth striae and 16-20 small
tubercles per whorl along edge of peripheral keel; aperture strongly ovate;
outer lip smooth; inner lip smooth, appressed posteriorly; siphonal canal long,
straight, axial; shell and aperture color porcellaneous white; periostracum,
operculum, and soft parts unknown.

ETYMOLOGY: Gr. sarissa — long pike used by Macedonian foot soldiers +
Gr. phoros — bearing.

TYPE SPECIMEN: Holotype — BM(NH) 1887.2.9.687, length 19.5 mm.

TYPE LOCALITY: CHALLENGER station 122, Off Pernambuco, Brazil,
9°05’S,34°50’W, in 350 fathoms (640 m).
GEOGRAPHIC DISTRIBUTION (fig. 13): Known only from the type locality.

ECOLOGY: This species has only been collected at one station, at a depth of
640 meters, on a substrate of red clay. The only known specimen has a naticid
bore hole.

MATERIAL EXAMINED: The holotype.

MEASUREMENTS OF THE HOLOTYPE:

width

length # whorls

spire angle

19.5 mm

COMPARATIVE REMARKS: The unique holotype resembles Venezuelan
specimens of F. brayi in general shape, but its small size and the presence of
tubercles along the peripheral keel serve to distinguish it from that species. It
differs from F. bermudezi in having a broader spire and finer and more
numerous spiral threads.

April 29, 1983 Harasewych 17

Fulgurofusus (Fulgurofusus) bermudezi (Clench and Aguayo, 1938)
Figures 9,13,21,23-26
SYNONYMY
1938 Columbarium bermudezi Clench and Aguayo, Mem. Soc. Cub.
Hist. Nat. 12(5):383-384,p1.28,fig.7; 1944 Clench, Johnsonia
1(15):2,plh. L fig. 3.
1969 Fulgurofusus bermudezi(Clench and Aguayo), Darragh, Proc. Roy.
Soc. Vict. 83(1):101-102,fig.5,p1.6,figs.120-121.

1971 Columbarium (Fulgurofusus) bermudezi Clench and Aguayo,
Bayer, Bull. Mar. Sci. 21(1):173,figs. 38 B-D, 39 A; 1974 Abbott,
American Seashells, second edition, p.170,fig.1803.

DESCRIPTION — SHELL: Medium in size (to 31 mm), elongate, thin; spire
angle 28°-32°; protoconch (fig.21) glassy, bulbous, of 1% whorls that
gradually merge into teleoconch; teleoconch with 7 slightly convex whorls;
suture impressed; keel rounded in transition zone, nodulate in early whorls,
with nodules fusing by the third post-nuclear whorl to form a crenulated flange
that may be normal to the coiling axis (figs.23,24) or directed anteriorly
(figs.25,26); growth striae fine, sinuate; spiral sculpture of fine threads to
strong cords generally limited to posterior portion of the siphonal canal;
aperture ovate; outer lip smooth; inner lip smooth, appressed posteriorly;
siphonal canal long, straight, axial; shell and aperture color white; perio-
stracum thin, yellowish.

ANIMAL: A single, dried specimen (fig.25) was reconstituted and found to
consist of nearly 3 whorls, of which the mantle cavity ocupied 1% whorl, the
kidney % whorl and the digestive gland %4 whorl. Tentacles short and blunt,
with large black eyespots; foot long, narrow, bearing an elongate, corneous,
terminally nucleate operculum; mantle cavity proportionally narrower and
longer than in F. brayi, as were the pallial organs and penis; radular ribbon
(fig.9) small, containing 73 rows of teeth; rachidian with 3 conical cusps
radiating from the middle half of the tooth; basal plate crescent-shaped, with
broadly expanded ends; lateral teeth with a single, long, curved, tapering cusp
rising from an elongate basal plate.

ETYMOLOGY: Named after P.J. Bermudez, who studied the foraminifera
collected by the ATLANTIS.

TYPE SPECIMENS: Holotype —MCZ 135007, length 27.5 mm; paratype —
MCZ 135008, length 25.0 mm; paratype — Museo Poey, Havana, Cuba;
paratypes — MCZ 135009 fragments.

TYPE LOCALITY: ATLANTIS station 2989, Off Sagua la Grande, Santa Clara,
Cuba, 23°10’N,80°04’W, in 360 fathoms (658 m).

GEOGRAPHIC DISTRIBUTION (fig.13): This species has been reported from
the northern coast of Cuba, the Santaren and Northwest Providence
Channels, and appears to be restricted to the upper continental slope.

NEMOURIA

FIGS. 23-26. Fulgurofusus (Fulgurofusus) bermudezi(Clench and Aguayo,

23.

24.

25

26.

1938).

USNM 806989, GERDA station G-1018, ENE of Dog Rocks,
Santaren Channel, in 556 m (2 X).

Holotype, MCZ 135007, ATLANTIS station 2989, Off Sagua la
Grande, Santa Clara, Cuba, in 658 m(2 X).

USNM 806990, GERDA station G-1015, W of Anguilla Cays,
Santaren Channel, in 525-516 m (2 X).

USNM 806989, GERDA station G-1018, ENE of Dog Rocks,
Santaren Channel, in 556 m (2 X).

April 29, 1983 Harasewych 19

EcoLocy: Fulgurofusus bermudezi has a bathymetric range of 384-1006 m,
with a mean station depth (n = 17) of 600 m. It has been taken on bottoms
ranging from fine sand to mud.

MATERIAL EXAMINED: ATLANTIS sta. 2981 D, Off Punta Alegre, Cuba,
22°48’°N,78°51’W, in 347-421 m (MCZ 135049); ATLANTIS sta. 2982 A,
Off Punta Alegre, Cuba, 22°48’N,78°50’W, in 384 m (MCZ 135050);
ATLANTIS sta. 2988, Off Sagua la Grande, Santa Clara, Cuba, 23°15’N,
79°57 W, in 695 m (MCZ 135008 — paratype); ATLANTIS sta. 2989, Off
Sagua la Grande, Santa Clara, Cuba, 23°10’N,80°04’W, in 658 m (MCZ
135007 — holotype, MCZ 135009 — paratypes); ATLANTIS sta. 3422, Off
Caibarien, Santa Clara, Cuba, 23°05’N,79°29’W, in 457 m(MCZ 135166);
ATLANTIS sta. 3423, Off Caibarien, Santa Clara, Cuba, 23°50’N, 79°08’W,
in 448 m (MCZ 135167); ATLANTIS sta. 3424, Off Caibarien, Santa Clara,
Cuba, 22°53’N,79°08’W, in 439 m (MCZ 135204); ATLANTIS sta. 3427,
Off Caibarien, Santa Clara, Cuba, 22°52’30”N,79°20’W, in 439 m (MCZ
135169); ATLANTIS sta. 3443, Off Sagua la Grande, Santa Clara, Cuba,
23°22’N,79°53’W, in594 m(MCZ 184514); ATLANTIS sta. 3457, Off Sagua
la Grande, Santa Clara, Cuba, 23°23’N, 80°36’W, in 1006 m (MCZ
135168); ATLANTIS sta. 3459, Off Sagua la Grande, Santa Clara, Cuba,
23°21’N,80°36’W, in 914 m (MCZ 135203); ATLANTIS sta. 3475, Off
Cardenas, Cuba, 23°18’N,80°48’W, in 732 m(MCZ 135250); GERDA sta.
G-190,NW of Great Stirrup Cay, Northwest Providence Channel, 25°57’N,
78°07 W, in 733-897 m (DMNH 153521); GERDA sta. G-524, Northwest
Providence Channel, 26°17’N, 78°41’W, in 513-715 m(DMNH 153520);
GERDA sta. G-1015, W of Anguilla Cays, Santaren Channel, 23°34’N,
79°17 W, in 525-516 m (UMML; USNM 806990); GERDA sta. G-1017,
E of Cay Sal Bank, Santaren Channel, 23°58’N,79°17'W, in 555 m
(UMML); GERDA sta. G-1018, ENE of Dog Rocks, Santaren Channel,
24°07’N,79°28’W, in 556 m (UMML; USNM 806989).

MEASUREMENT OF SHELL CHARACTERS (n = 10):
width # whorls

length Spire angle

mean 27.5 mm 29.2°
range 25.4 — 31.0 28° — 32°
sd 1.8 ae

COMPARATIVE REMARKS: A high spire, axial ribbing on early whorls, and
lack of spiral sculpture above the periphery are characters that distinguish this
species from other members of the subgenus. It resembles F. quercollis in
general body shape, but lacks the pronounced spiral sculpture characteristic of
the Paleocene species. Fulgurofusus bermudezi is quite variable in shell form,
the peripheral keel may be narrow or wide, directed laterally or anteriorly,

20 NEMOURIA No. 27

smooth or crenulated. Unlike F. brayi, this cannot be attributed to clinal
variation, as extreme examples (figs. 23,26) have been taken at the same
station.

Fulgurofusus (Fulgurofusus) ecphoroides new species
Figures 13,22,27-31

DESCRIPTION: Shell small (to 18.5 mm), delicate, fusiform; spire angle 47°-
58°; protoconch (fig.22) bulbous, glassy, of 1% whorls; transition to
teleoconch marked by acquisition of a round keel that rapidly develops into a
short peripheral flange; teleoconch with 5 convex whorls; suture impressed;
spiral sculpture of 2-4 strong cords on body whorl anterior to keel, and 2-6
slightly weaker cords on posterior portion of siphonal canal; keel and cords
undulating, furrowed, each consisting of 2 fine threads in close apposition;
axial sculpture limited to fine, sinuate growth striae; aperture ovate; outer lip
furrowed beneath spiral cords; inner lip smooth, may be posteriorly appressed
or form a raised peristomal plate; siphonal canal long, axial, twisted slightly;
color uniform porcellaneous white; periostracum, operculum, and soft parts
unknown.

ETYMOLOGY: Named for its resemblance to shells of the genus Ecphora
Conrad, 1843 (Gastropoda: Thaididae). Ecphora + Gr. oides — like.

TYPE SPECIMENS: Holotype — USNM 810464, length 18.4 mm; Paratype 1
— DMNH 153515, length 16.0 mm; Paratype 2 — MCZ 281476, length
10.3 mm; Paratype 3 — USNM 821507, length 14.1 mm; 15 additional
paratypes — USNM 821507.

TYPE LOCALITY: OREGON station 4226, 180 miles NE of Tijoca, Para,
Brazil, 0°18’N, 44°17’W, in 150 fathoms (274 m).

GEOGRAPHIC DISTRIBUTION (fig. 13): Known only from the type locality.

ECOLOGy: Of the 19 specimens collected at a depth of 274 m, none were taken
alive. Several were inhabited by hermit crabs, and a number had been drilled
by naticids.

MATERIAL EXAMINED: The 19 specimens in the type lot.
MEASUREMENTS OF SHELL CHARACTERS (n = 10):
# whorls

spire angle

mean 14.0 mm 5.9 mm 5.6 52.6°
range 10.3-18.4 3.9-7.7 4.5-7.0 47°-58°
sd oS 1.0 0.3 3.6°

COMPARATIVE REMARKS: This distinctive new species can be distinguished
from all other members of the subgenus by the presence of 2-4 strong cords on
the body whorl and by the incised furrows that run along the peripheral keel
and all spiral cords. The inner lip may form a raised peristomal plate in mature

April 29, 1983 Harasewych 21

FIGS. 27-31. Fulgurofusus (Fulgurofusus) ecphoroides new species.

27. Holotype, USNM 810464, OREGON station 4226, 180 miles NE of
Tijoca, Para, Brazil, in 274 m (3 X).

28. Paratype 1, DMNH 153515, OREGON station 4226, 180 miles NE of
Tijoca, Para, Brazil, in 274 m (3 X).

29. Paratype 2, MCZ 281476, OREGON station 4226, 180 miles NE of
Tijoca, Para, Brazil, in 274 m (3 X).

30. Apical view of paratype 2 (5 X).

31. Paratype 3, USNM 821507, OREGON station 4226, 180 miles NE of
Tijoca, Para, Brazil, in 274 m (3 X).

specimens (figs.27,31). Darragh (1969:69,71) considered such a plate to be
characteristic of the genus Columbarium, but the absence of other
Columbarium characters such as spines on the periphery, scales on spiral
cords and a strong anterior carina, as well as the presence of a Fulgurofusus
protoconch and early whorls support the placement of this species in
Fulgurofusus.

This species most closely resembles F. sarissophorus, with which it agrees
in size and general shape, but from which it differs in having an incised keel and
Spiral cords.

22 NEMOURIA No. 27

Subgenus Histricosceptrum Darragh, 1969
SYNONYMY

1969 Histricosceptrum Darragh, Proc. Roy. Soc. Vict. 83(1):87. Type
species by original designation: Columbarium atlantis Clench
and Aguayo, 1938.

1971 Columbarium (Histricosceptrum) Darragh, Bayer, Bull. Mar. Sci.
21(1):170.

This group, which is not represented in the fossil record, is very similar to,
and clearly derived from, Fulgurofusus. It is characterized by a relatively
thick shell with strong spiral cords along the body whorl and siphonal canal as
well as by the formation of long, open spines along the periphery. In addition to
3 western Atlantic species, one of which is described herein, a species from the
Indian Ocean has been provisionally included by Darragh (1969:89) who
stated that “further work . . . may show that it would be better placed in
Columbarium’.

Fulgurofusus (Histricosceptrum) atlantis (Clench and Auguayo, 1938)
Figures 10,32,38,39

SYNONYMY

1938 Columbarium atlantis Clench and Aguayo, Mem. Soc. Cub. Hist.
Nat. 12(5):382-383,p1.28,fig.1; 1944 Clench, Johnsonia 1(15):
2-3,p1.1,fig.4.

1969 Histricosceptrum atlantis (Clench and Aguayo), Darragh, Proc.
Roy. Soc. Vict. 83(1):88,fig.1,p1.6,figs.119,122.

1974 Columbarium (Histricosceptrum) atlantis Clench and Aguayo
Abbott, American Seashells, second edition, p.170,fig. 1802.

DESCRIPTION — SHELL: Of medium size (to 36 mm), elongate, moderately
heavy; spire angle 30°-32°; protoconch (fig.39) of 1% whorls, glassy, rotund;
transition to teleoconch indistinct, marked by gradual acquisition of a
peripheral keel, rounded at first, but with spines by second postnuclear whorl;
teleoconch with 7 convex whorls; suture impressed; keel with 8-10 long, open,
anteriorly deflected spines per whorl; spiral sculpture of 4-6 threads and 1 cord
between suture and keel, 3-5 cords on body whorl anterior to keel and 20-25
cords on posterior % of siphonal canal; growth striae fine, sinuate; aperture
semicircular; outer lip furrowed beneath keel; inner lip smooth, covered with
thin glaze; siphonal canal long, stout, straight; color uniformly white;
periostracum unknown.

ANIMAL: Foot long, narrow, bearing a thin, yellow corneous, strongly ovate
operculum with a terminal nucleus and circular attachment area; tentacles
short, with large black eyes; mantle cavity narrow, long of 1%4 whorls; penis
with open, glandular groove; proboscis long, coiled in proboscis sheath; radula
(fig.10) tiny, consisting of approximately 85 rows; rachidian with 3 cusps,

April 29, 1983 Harasewych 23
long, conical, recurved; central cusp flanked by slightly shorter cusps; ends of
basal plate expanded; lateral teeth with long, single, scythe-like cusps.

ETYMOLOGY: Named after the research vessel ATLANTIS, which collected
the only known specimens.

TYPE SPECIMENS: Holotype — MCZ 135004, length 35.2 mm; paratype —
Museo Poey, Havana, Cuba.

TYPE LOCALITY: ATLANTIS station 2999, Off Matanzas, Cuba, 23°10’N,
81°29’W, in 421 fathoms (770 m).

GEOGRAPHIC DISTRIBUTION (fig. 38): This species has been reported from 4
stations off the northwest coast of Cuba. All are within 150 miles (250 km) of
the type locality.

ECOLOGY: This species has been taken at depths ranging from 475 to 770
meters with an average station depth (n = 4) of 650 meters. It has been
collected with F. bermudezi at 2 stations.

MATERIAL EXAMINED: ATLANTIS sta. 2999, Off Matanzas, Cuba, 23°10°N,
81°29’W, in 770 m (MCZ 135004 — holotype); ATLANTIS sta. 3434, Off
Sagua la Grande, Santa Clara, Cuba (MCZ 178190).

MEASUREMENTS OF SHELL CHARACTERS (n = 2):
length # whorls Spire angle

34.0-35.2 mm 6.3-7.1 mm . 30°-32°

COMPARATIVE REMARKS: This species may be distinguished from its
congeners by its thin elongate shell, with spiral sculpture limited to fine, simple
threads, and by the presence of a single row of long, open, anteriorly directed
spines along the periphery.

Fulgurofusus (Histricosceptrum) bartletti (Clench and Aguayo, 1940)
Figures 33-34,38

SYNONYMY

1940 Columbarium bartletti Clench and Aguayo, Mem. Soc. Cub. Nat.
Hist. 14(1):86,p1.14,fig.3; 1944 Clench, Johnsonia 1(15):3-4,
pl.1,fig.5.

1969 Histricosceptrum bartletti (Clench and Aguayo), Darragh, Proc.
Roy. Soc. Vict. 83(1):88-89.

1971 Columbarium (Histricosceptrum) bartletti Clench and Aguayo,
Bayer, Bull. Mar. Sci. 21(1):170,fig.38 A; 1974 Abbott, American
Seashells, second edition, p.170,fig.1801.

DESCRIPTION: Shell of medium size (to 39 mm), elongate, heavy; spire angle
31°-34°; protoconch glassy, bulbous, of 14% whorls; transition to teleoconch
indistinct; keel, rounded at first, becomes nodular, then, by second post-

24 NEMOURIA No. 27

nuclear whorl, spinose; spines long, open, normal to coiling axis, numbering 7-
8 per whorl; teleoconch of 8 convex whorls; suture adpressed; spiral cords
between suture and keel becoming more pronounced, fusing by fourth post-
nuclear whorl to form a channel at the suture; below keel there are 5 cords on
body whorl and 19-23 on posterior half of siphonal canal; growth striae fine,
sinuate, forming axial ribs on sutural channel; aperture ovate to subquadrate;
outer lip furrowed beneath keel; inner lip smooth, appressed posteriorly;
siphonal canal long, stout, axial; color white; periostracum very thin, amber in
color; operculum and soft parts unknown.

ETYMOLOGY: Named in honor of Commander J.R. Bartlett, U.S.N., of the
U.S. Coast Survey Steamer BLAKE.

TYPE SPECIMENS: Holotype — MCZ 104729, length 38.7 mm; paratype —
Museo Poey, Havana, Cuba.

TYPE LOCALITY: BLAKE station 9, Off Homers Cove, Westmoreland,
Jamaica, 18°12’N,78°20’W, in 254 fathoms (465 m).

GEOGRAPHIC DISTRIBUTION (fig.38): The two records of this species are
from off the southwest coast of Jamaica.

ECOLOGY: Based on two records, the bathymetric range of this species is 465-
546 m.

MATERIAL EXAMINED: BLAKE sta. 9, Off Homers Cove, Westmoreland,
Jamaica, 18°12’N,78°20’W, in 465 m (MCZ 104729 — holotype);
PILLSBURY sta. P-1225, SW of Jamaica, 17°42.5’N,78°58’ W, in 546-528 m
(USNM 806991).

FIG. 32. Fulgurofusus (Histricosceptrum) atlantis (Clench and Aguayo,
1938). Holotype, MCZ 135004, ATLANTIS station 2999, Off
Matanzas, Cuba, in 770 m (2 X).

FIGS. 33-34. Fulgurofusus (Histricosceptrum) bartletti (Clench and
Aguayo, 1940).

33. Holotype, MCZ 104729, BLAKE station 9, Off Homers Cove,
Westmoreland, Jamaica, in 465 m (2 X).

34. USNM 806991, PILLSBURY station P-1225, SW of Jamaica, in 546-
528 m (2X).

FIGS. 35-37. Fulgurofusus (Histricosceptrum) xenismatis new species.

35. Holotype, USNM 806992, PILLSBURY station P-1354, Off north-
eastern Nicaragua, in 192-263 m (2 X).

36. Paratype 1, DMNH 153523, PILLSBURY station P-1354, Off north-
eastern Nicaragua, in 192-263 m(2 X).

37. Paratype 2, USNM 806993, PILLSBURY station P-1354, Off north-
eastern Nicaragua, in 192-263 m(2 X).

April 29, 1983 Harasewych

26 NEMOURIA No. 27

BERMUDA

GULF OF MEXICO te
"th ate

> 5) BAHAMAS
\ v

o

c=}
bartletti @®@
JAMAICA

@®@ xenismatis

i)

.)

FIG. 38. Geographic distribution of the western Atlantic species of Fulguro-
fusus (Histricosceptrum) and Fulgurofusus (Peristarium).

MEASUREMENTS OF SHELL CHARACTERS (n = 2):
length # whorls Spire angle

38.7 mm 8.6-8.8 mm 31°-34°

COMPARATIVE REMARKS: The high spire, thick shell, single row of long,
heavy, laterally directed spines along the periphery, and strongly channeled
suture serve to distinguish this species from all other members of the
subgenus.

Fulgurofusus (Histricosceptrum) xenismatis new species
Figures 35-38,40

DESCRIPTION: Shell of medium size (to 34 mm), elongate, thin to moderately
heavy; spire angle 32°-40°; protoconch (fig.40) of 14% whorls, bulbous, glassy;
transition to teleoconch indistinct; keel rounded, becoming nodulose and
finally spinose by the second post-nuclear whorl; periphery with 10-12 short,
open spines per whorl that are directed posteriorly on early whorls, but may be
laterally or even anteriorly deflected on final whorl; spiral cord between suture
and periphery becomes strongly nodulose, and by the fourth post-nuclear
whorl, spinose, producing a second row of spines that are directed posteriorly;
teleoconch with 8 convex whorls; suture impressed; spiral sculpture of 4-5

April 29, 1983 Harasewych 24

cords on body whorl and 15-20 on posterior % of siphonal canal; axial growth
striae fine; aperture ovate; outer lip furrowed beneath both rows of spines;
inner lip smooth, appressed posteriorly; siphonal canal long, stout, axial; color
porcellaneous white; periostracum very thin, yellowish; operculum corneous,
yellowish, strongly ovate, with terminal nucleus; soft parts unknown.

ETYMOLOGY: Gr. xenismatis — amazing.

TYPE SPECIMENS: Holotype — USNM 806992, length 25.8 mm; Paratype 1
— DMNH 153523, length 33.0 mm; Paratype 2 — USNM 806993, length
33.5 mm; Paratype 3 — MCZ 281478, length 27.3 mm; Paratypes 4 & 5 —
UMML 30-8301.

TYPE LOCALITY: PILLSBURY station P-1354, Off northeastern Nicaragua,
14°21°N,81°55’W, in 192-263 m.

GEOGRAPHIC DISTRIBUTION (fig.38): This species has only been taken at
two stations, both off the northeastern coast of Nicaragua.

ECOLOGY: The shallowest verifiable record for this species is 263 m, the
deepest 450 m. Several specimens have muricid and naticid bore holes.

MATERIAL EXAMINED: The 5 specimens in the type lot. Holotype and
paratypes 1-3 from the type locality, paratypes 4-5 from PILLSBURY sta. P-
1357, Off northeastern Nicaragua, 14°35N, 81°32’W, in 450-576 m.

MEASUREMENTS OF SHELL CHARACTERS (n = 3):
# whorls spire angle

mean 30.8 mm : 36.8
range 25.8-33.5 32°-40°

COMPARATIVE REMARKS: A second row of short, open, posteriorly directed
spines between the suture and the periphery serves to separate this new species
from the other members of the subgenus.

Subgenus Peristarium Bayer, 1971
SYNONYMY

1971 Columbarium (Peristarium) Bayer, Bull. Mar. Sci. 21(1):176.
Type species by original designation: Columbarium (Peristarium)
electra Bayer, 1971.

Recognizing only the genus Columbarium, Bayer (1971) included in it his
new subgenus Peristarium. Reasons for reelevating Fulgurofusus to generic
level have already been discussed. Peristarium is provisionally regarded as a
subgenus of Fulgurofusus on the basis of general similarity in shell shape and
protoconch, prominence of axial sculpture, and total lack of spines or scales.
Known from 4 Recent species, one of which is described herein, the subgenus
is characterized by an elongate, moderately heavy shell with spiral ornament

28 NEMOURIA No. 27

of fine threads to low, broad cords, an incised furrow along the periphery, and
12-16 low axial ribs. The siphonal canal may be dorsally deflected in some
species. Gross anatomy is similar to that of Fulgurofusus s.s., but the radula
differs in having the basal plate of the rachidian even more expanded and
buttressed. Unlike most Columbarines, species of Peristarium are steno-
bathic, and do not overlap in their bathymetric ranges.

Fulgurofusus (Peristarium) electra (Bayer, 1971)
Figures 38,41 ,43-45

SYNONYMY

1971 Columbarium (Peristarium) electra Bayer, Bull, Mar. Sci. 21(1):
176-178, figs.39 D,41; 1974 Abbott, American Seashells, second
edition, p.170.

DESCRIPTION: Shell small (to 27 mm), elongate, thin; spire angle 26°-28°;
protoconch of 1% whorls glassy, bulbous, not clearly delimited from
teleoconch of 8 convex whorls; suture adpressed, shoulder rounded at first,
becoming more angulate and bearing nodules within first postnuclear whorl;
periphery of shell slightly furrowed, indicating vestiges of 2 spiral cords in
close apposition; spiral sculpture of 6-14 weak, broad cords above periphery
and 4-5 stronger cords below; siphonal canal with 7-10 weak cords on
posterior %; axial sculpture of 14 low ribs per whorl; growth striae fine,
sinuate; aperture strongly ovate; outer lip smooth; inner lip smooth, covered
with thin glaze; siphonal canal long, stout, axial; color porcellaneous white;
periostracum unknown; operculum strongly ovate, corneous, with terminal
nucleus and elliptical attachment area; radula (Bayer, 1971,p.172,fig. 39 D)
has tricusped rachidian with strongly recurved basal plate and long, single-
cusped, recurved lateral teeth.

ETYMOLOGY: Gr. Electra — daughter of Agamemnon.

TYPE SPECIMENS: Holotype — USNM 701151, length 26.8 mm; paratype —
DMNH 153517; paratypes — UMML.

TYPE LOCALITY: GERDA Station G-289, SSE of Key West, Straits of Florida,
24°11°N,81°36’W, in 604-594 m.

FIG. 39. Protoconch of Fulgurofusus (Histricosceptrum) atlantis (Clench
and Aguayo, 1938). Holotype (figure 32).

FIG. 40. Protoconch of Fulgurofusus (Histricosceptrum) xenismatis new
species. Holotype (figure 35).

FIG. 41. Protoconch of Fulgurofusus (Peristarium) electra (Bayer, 1971).
DMNH 153517.

FIG. 42. Protoconch of Fulgurofusus (Peristarium) aurora (Bayer, 1971),
specimen shown in figure 51.

Scale bar = 1 mm for all drawings.

April 29, 1983 Harasewych 29

30 NEMOURIA No. 27

GEOGRAPHIC DISTRIBUTION (fig. 38): Known only from the type locality.

EcOoLocGy: All known specimens were collected at one station at a depth of
604-594 m.

MATERIAL EXAMINED: The 6 specimens in the type lot.
MEASUREMENTS OF SHELL CHARACTERS (n = 3):
# whorls Spire angle

mean 20.0 mm : Die
range 16.6-26.8 20:5 =2 1.5"

COMPARATIVE REMARKS: This species may be distinguished from other
members of the subgenus by its small size, angulate periphery, and numerous
fine spiral threads.

Fulgurofusus (Peristarium) merope (Bayer, 1971)
Figures 38,53-55

SYNONYMY

1971 Columbarium (Peristarium) merope Bayer, Bull. Mar. Sci. 21(1):
178-180, figs. 39 C,42,43 A; 1974 Abbott, American Seashells,
second edition, p.171.

DESCRIPTION: Shell of medium size (to 36 mm), elongate, moderately thin;
spire angle 28°-32°; protoconch of nearly 2 whorls, bulbous, glassy; transition
to teleoconch indistinct; teleoconch with 8 convex, weakly shouldered whorls;
suture adpressed; periphery passes through rounded and angulate stages to
become nodulose by the second post-nuclear whorl; spiral sculpture of 4-5
cords above periphery, 2 prominent cords separated by a narrow furrow along
the periphery, and 8-10 cords below the periphery, with 20-24 weaker cords
along the siphonal canal; axial sculpture of 12 low ribs per whorl; growth striae

FIGS. 43-45. Fulgurofusus (Peristarium) electra (Bayer, 1971).
43. Holotype, USNM 701151, GERDA station G-289, SSE of Key West,
Straits of Florida, in 604-594 m (3 X).
44-45. Paratypes, UMML, GERDA station G-289, SSE of Key West, Straits
of Florida, in 604-594 m (3 X).
FIGS. 46-49. Fulgurofusus (Peristarium) timor new species.
46. Holotype, USNM 87487, ALBATROSS station 2676, Off Cape Fear,
North Carolina, in 744 m (1.5 X).
47. Paratype 1, USNM 821508, ALBATROSS station 2676, Off Cape
Fear, North Carolina, in 744 m(1.5 X).
48. Paratype 2, Staples collection, Off Cape Fear, North Carolina, in 700
m (2.0 X).
49. Paratype 2, DMNH 153516, Off North Carolina, deep water(1.5 X).

April 29, 1983 Harasewych

32 NEMOURIA No. 27

fine, sinuate; aperture ovate; outer lip crenulated in juveniles, smooth in adults;
inner lip smooth, covered with thin glaze; siphonal canal long, stout, straight;
Shell and aperture porcellaneous white in color; periostracum thin, yellowish,
forming short lamellae along growth lines; operculum corneous, elongate,
slightly unguiculate, with terminal nucleus and small elliptical attachment
area; radula (Bayer 1971,p.172,fig.39 C) with a rachidian in which the basal
plate is not nearly as recurved as in other members of the subgenus, three cusps
occupy a wider portion of the tooth, and lateral teeth appear thinner and more
recurved than in other species.

ETYMOLOGY: Gr. Merope — one of the Pleiades.

TYPE SPECIMENS: Holotype — USNM 701152, length 36.0 mm; paratype —
USNM 806994; paratype — MCZ 281477; paratype — DMNH 156518;
paratypes — UMML 30-6970.

TYPE LOCALITY: GERDA station G-476, SW of Marquesas Keys, Straits of
Florida, 24°14’N,82°24’W, in 549-512 m.

GEOGRAPHIC DISTRIBUTION (fig. 38): Fulgurofusus merope lives along the
upper continental slope in the area south of the Marquesas Keys.

ECOLOoGy: This species has been taken in depths between 512 and 584 m,
with a mean station depth (n = 5) of 546 m.

MATERIAL EXAMINED: GERDA sta. G-970, SW of Marquesas Keys, Straits
of Florida, 24°24’N,82°08’W, in 512 m(DMNH 156518); GERDA sta. G-
440, SW of Marquesas Keys, Straits of Florida, 24°14’N,82°21’W, in 549-
567 m(USNM 806994); GERDA sta. G-966, SW of Marquesas Keys, Straits
of Florida, 24°10’N,82°22’W, in 553-558 m(MCZ 281477); GERDA sta. G-
476, SW of Marquesas Keys, Straits of Florida, 24°14’N,82°24’W, in 512-
549 m(USNM 701152 — holotype); GERDA sta. G-439, SW of Marquesas
Keys, Straits of Florida, 24°14’N,82°29’W, in 566-584 m (UMML 30-
6970).

MEASUREMENTS OF SHELL CHARACTERS (n = 4):
# whorls spire angle
mean 28.9 mm ; 293

range 22.1-36.0 28°-30°
sd 6.2 : 0.9°

COMPARATIVE REMARKS: This species may be separated from Fulgurofusus
electra and F. timor by its stronger spiral sculpture and more prominent axial
ribs. It differs from F. aurora in having an angular periphery and a more
constricted juncture between body whorl and siphonal canal.

April 29, 1983 Harasewych 33

Fulgurofusus (Peristarium) aurora (Bayer, 1971)
Figures 12,38,42,50-52

SYNONYMY
1971 Columbarium (Peristarium) aurora Bayer, Bull. Mar. Sci. 21(1):
180-183,figs.39 E, 43 B, 44; 1974 Abbott, American Seashells,
second edition, p.171.

DESCRIPTION — SHELL: Large (to 43 mm), elongate, moderately heavy;
spire angle 33°-37°; protoconch (fig.42) of 1% whorls, glassy, bulbous;
transition to teleoconch marked by appearance of rounded keel, which
becomes angulate and intersected by axial ribs within the first post-nuclear
whorl; teleoconch of 8 convex whorls; suture adpressed; spiral sculpture of
low, broad bands separated by narrow furrows; 2 bands, wider than the rest,
occur along the rounded periphery, with 4-6 narrower bands above and 7-10
below on body whorl; siphonal canal long, stout, tapering gradually, with up to
25 cords of varying width along most of its length; axial sculpture of 12-16 low
ribs; growth striae fine, sinuate; aperture ovate to nearly elliptical; outer lip
smooth; inner lip smooth, covered with thin glaze; shell dingy white in color;
periostracum thin, yellowish, with axial lamellae that tend to be abraded along
spiral cords.

ANIMAL: Reconstituted soft parts, removed from a paratype (fig.52), of about
2% whorls, with the mantle cavity extending over % whorl, the kidney over 4%
whorl, and the digestive gland over %4 whorl; operculum lenticular, amber
colored, terminally nucleated; mantle cavity containing a broadly expanded
rectum, a muscular open sperm groove, and a long, flat, distally rounded penis,
with an open groove running along its inner edge; proboscis, long, folded,
containing small buccal mass; radular ribbon (fig. 12) small, containing nearly
100 rows of teeth; rachidian with 3 parallel, conical cusps occupying middle
half of tooth; ends of basal plate greatly expanded and buttressed in opposition
to the cusps; lateral teeth with a single, long, scythe-like cusp.

ETYMOLOGY: L. Aurora — goddess of the dawn.

TYPE SPECIMENS: Holotype — USNM 701222, length 41.4 mm; paratypes
— USNM 701153; USNM 806995; USNM 806996; DMNH 153519;
MCZ 281479; UMML.

TYPE LOCALITY: PILLSBURY station P-1309, NE of Fowey Light, Straits of
Florida, 25°40’N, 80°02’W, in 247 m.

GEOGRAPHIC DISTRIBUTION (fig.38): This species inhabits the uppermost
continental slope off Miami, Florida.

EcOLOoGy: The bathymetric range of F. aurora is 247-403 m, with a mean
station depth (n = 5) of 339 m. A dorsally deflected siphonal canal indicates
an infaunal habitat.

MATERIAL EXAMINED: GERDA sta. G-62, SE of Fowey Light, Straits of
Florida, 25°30.5’N,80°00’ W, in 384-403 m (USNM 701153); GERDA sta.

34 NEMOURIA No. 27

G-66, SE of Fowey Light, Straits of Florida, 25°25.5’N,79°59’W, in 366 m
(UMML); GERDA sta. G-67, SE of Fowey Light, Straits of Florida,
25°31°N,79°57'W, in 351 m (USNM 806995); GERDA sta. G-828, E of
Fowey Light, Straits of Florida, 25°34’N,79°57’W, in 333-340 m (USNM
806996); PILLSBURY sta. P-1309, NE of Fowey Light, Straits of Florida,
25°40’N,80°02’W, in 247 m(USNM 701222 — holotype; DMNH 153519;
MCZ 281479).

MEASUREMENTS OF SHELL CHARACTERS (n = 7):

# whorls Spire angle
mean 33.4 mm | . 33.2,
range 22.4-43.0 31 =355

sd 7.3 : 1.6°

COMPARATIVE REMARKS: Fulgurofusus aurora may be distinguished from
its congeners by its thick shell and prominent axial ribs and spiral cords, as
well as by its rounded shoulder and broad siphonal canal.

Fulgurofusus (Peristarium) timor new species
Figures 38,46-49

SYNONYMY

1889 Fusus schrammi Crosse, Dall, Bull. Mus. Comp. Zool. Harvard
18:168; 1889 Dall, Bull. U.S. Natl. Mus. 37:112, 1904 Grabau
[in part], Smithson. Misc. Collns. 44:72-73; 1934 Johnson,
Proc.Bost.Soc.Nat.Hist., 40(1):128. [not Fusus schrammi Crosse,
1865].

1974 Fusinus schrammi (Crosse) Abbott, American Seashells, second
edition, p.230 [not Fusus schrammi Crosse, 1865].

FIGS. 50-52. Fulgurofusus (Peristarium) aurora (Bayer, 1971).
50. Holotype, USNM 701222, PILLSBURY station P-1309, NE of Fowey
Light, Straits of Florida, in 247 m (2 X).
51. Paratype, USNM 806996, GERDA station G-828, E of Fowey Light,
Straits of Florida, in 333-340 m (2 X).
52. Paratype, MCZ 281479, PILLSBURY station P-1309, NE of Fowey
Light, Straits of Florida, in 247 m (2 X).
FIGS. 53-55. Fulgurofusus (Peristarium) merope (Bayer, 1971).
53. Holotype, USNM 701152, GERDA station G-476, SW of Marquesas
Keys, Straits of Florida, in 549-512 m (2 X).
54. Paratype, DMNH 153518, GERDA station G-970, S of Marquesas
Keys, Straits of Florida, in 512 m (2.5 X).
55. Paratype, MCZ 281477, GERDA station G-966, SW of Marquesas
Keys, Straits of Florida, in 553-558 m (2.5 X).

April 29, 1983 Harasewych

36 NEMOURIA No. 27

DESCRIPTION: Shell large (to 54 mm), elongate, moderately heavy; spire
angle 35°-38°; protoconch of nearly 2 whorls, glassy, bulbous, with indistinct
transition to teleoconch of 6 convex, strongly shouldered whorls; suture
adpressed; peripheral keel passing through rounded, angulate, and into
nodulose stage within first post-nuclear whorl; shell periphery with incised
furrow that is most prominent between axial nodes; spiral sculpture of
numerous fine threads on body whorl, with stronger cords appearing on
posterior portion of the siphonal canal; axial sculpture of 12-16 low ribs per
whorl; growth striae fine, weakly sinuate; aperture ovate; outer lip smooth;
inner lip smooth, covered by a thin glaze; siphonal canal long, thin, axial, with
distal portion dorsally deflected; shell color dingy white to very light tan;
periostracum, operculum and soft parts unknown.

ETYMOLOGY: Named after the type locality of Cape Fear, North Carolina.
L. timor — fear.

TYPE SPECIMENS: Holotype — USNM 87487, length 57.1 mm; Paratype 1
— USNM 821508, length 36.3 mm; Paratype 2, Staples collection — length
45.2 mm; Paratype 3 — DMNH 153516, length 53.6 mm.

TYPE LOCALITY: ALBATROSS sta. 2676, Off Cape Fear, North Carolina,
32°39’N,77°01’W, in 407 fathoms (744 m).

GEOGRAPHIC DISTRIBUTION (fig. 38): This species is known only from off the
coast of North Carolina in the vicinity of Cape Fear.

ECOLOGY: The four specimens were taken at depths ranging from 700 to 744
meters.

MATERIAL EXAMINED: ALBATROSS sta. 2676, Off Cape Fear, North
Carolina, 32°39°N,77°01’W, in 744 m (USNM 87487 — holotype; USNM
821508 — paratype); Off Cape Fear, North Carolina, in 700 m (Staples
collection — paratype); Off North Carolina, in deep water (DMNH 153516
— paratype).

MEASUREMENTS OF SHELL CHARACTERS (n = 3):

# whorls spire angle

mean 46.2 mm 12.3 mm ‘ 36.7°
range 452-57 .1 10.4-14.8 35°-38°

COMPARATIVE REMARKS: This species, first collected during the ALBATROSS
cruises, has been misidentified for nearly a century as Fusus schrammi
Crosse, a tropical, subtidal Fusinus from Guadeloupe. Although superficially
resembling a small Fusinus, its columbarine protoconch and incised periphery
show that it is referable to Peristarium. Fulgurofusus (Peristarium) timor
most closely resembles F. electra, but the large size, fine spiral sculpture,
rounded body whorl and prominently furrowed periphery of F. timor
distinguish it from other members of the subgenus.

April 29, 1983 Harasewych 37
DISCUSSION

The lack of a sharp distinction between protoconch and teleoconch, even in
Paleocene species of Fulgurofusus, indicates a long history of direct
development. The smooth shell and dorsally deflected siphonal canal of
Peristarium appear to be adaptations for burrowing, while the wide peripheral
keels and long, delicate spines of Fulgurofusus s.s. and Histricosceptrum
strongly suggest a purely epifaunal habitat. Major predators are crabs and
drilling gastropods, as evidenced by frequent examples of shell repair and
naticid as well as muricid boreholes.

Columbarines are morphologically well adapted for feeding on tubiculous
polychaetes, a diet confirmed by the presence of serpulid and chaetopterid
setae in the rectum of F. brayi. Schacko’s (in Martens, 1881) illustrations of
the ‘“‘radular teeth’? of Columbarium spinicinctum are clearly depictions of
polychaete setae. The long siphon and large osphradium are indicative of
acute chemoreceptive ability, while a long, narrow proboscis, small buccal
mass, and strong retractor muscles extending to the columellar muscle appear
to be adaptations for extracting prey from its tube. The rachidian, with a basal
plate that is greatly expanded and buttressed in opposition to the stout, conical
cusps, is well suited to withstand a pulling force perpendicular to the cusps.
Lateral teeth are long, recurved, and raptorial. Secretions of the large salivary
glands may include a narcotizing agent similar to that found in certain
Buccinaceans (Fange, 1960). The narrow diameter of the esophagus and
nerve ring suggest that prey is either limited to small worms swallowed whole,
or is eaten in fragments. A large, expanded rectum contains an accumulation
of setae.

Little is known about reproductive biology, other than that the open sperm
ducts of the male are a primitive feature (Ponder, 1973:316).

Initial inclusion of certain columbarine species in Turridae was due to the
mistaken identification of polychaete setae, taken from the proboscis, as
toxoglossan radular teeth. Transfer of the group to the Muricidae was based on
superficial similarities of shells and radulae, but the lack of accessory salivary
glands, accessory boring organ and purple hypobranchial gland show that the
Columbariinae are not referable to this calciocavitating family. Although the
Columbariinae’s affinities with the Buccinidae are closer, the presence of an
anal gland and an open sperm groove serve to differentiate it from that family.
The family Columbariidae was erected (Tomlin, 1928:330) and family status
supported (Ponder, 1973:329) on grounds that the group could not be included
in Muricidae or Buccinidae.

Comparison of the morphology of Fulgurofusus (Fulgurofusus) brayi with
that of certain turbinellids [ Turbinella pyrum (Linnaeus), Dall, 1885; Moses,
1923; Turbinella laevigata Anton, Bayer, 1971; Harasewych, unpublished
observations; Cyomesus chaunax (Bayer) Bayer, 1971; Harasewych,
unpublished observations] reveals close correspondence in such features as:
presence of long, folded proboscis and non-evertable proboscis sheath;

38 NEMOURIA No. 27

similarity in radular morphology; lack of accessory salivary glands; presence
of anal gland and open or partially fused sperm groove; diet of tubiculous
polychaetes. These and other similarities leave little doubt that the group
merits no more than subfamily status within the Turbinellidae.

Earliest records of the Columbariinae are from the Upper Cretaceous and
early Tertiary of western Europe, the United States, and New Zealand, and
are roughly contemporary with the first appearance of other turbinellid groups
(Wenz, 1941). The genera Fulgurofusus and Columbarium were well
established by the Paleocene, and were both members of subtropical, offshore
communities. The disappearance of Columbariinae from the post-Eocene
fossil record dates the retreat of the group into deeper water. Australian
Miocene records are from deep water facies (Darragh, 1969:65).

After the closing of the Tethys Sea in the early Tertiary, the genus
Columbarium was restricted to the Indo-Pacific, while Fulgurofusus retained
a wider distribution. Reports of species of Fulgurofusus from off New Zealand
(Darragh, 1969:104) and the South Orkney Islands are here interpreted as
part of an abyssal, circum-Antarctic distribution originating from east
Tethyan stock. Based on an examination of the figures, Columbarium
tomicici McLean and Andrade, 1982, recently described from off western
South America, appears to be referable to Fulgurofusus s.s., and is likely an
offshoot of the Eocene F. merriami or F. washingtonianus.

The subgenus Histricosceptrum, known only from the Recent fauna, is very
similar to Fulgurofusus s.s.. With the exception of a single Indian Ocean
species placed tentatively in this subgenus, all other species are restricted to
the Caribbean. The subgenus Peristarium is also known only from the Recent
of the western Atlantic. Its similarity to certain species attributed to Coluzea,
especially from the Eocene of Europe and the Recent of South Africa,
indicates that further work is required to delineate more precisely the
relationships of this group.

April 29, 1983 Harasewych 39
LITERATURE CITED

Barnard, K.H.
1959 Contributions to the knowledge of South African marine mollusca.
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1971 New and unusual mollusks collected by R/V John Elliott Pillsbury
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1944 The genus Columbarium in the western Atlantic. Johnsonia
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40 NEMOURIA No. 27

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1937 The Wangalloan and associated molluscan faunas of Kaitangata —
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1883 (1880-1887). Manuel de conchyliologie et de paléontologie
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April 29, 1983 Harasewych 41

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1907 The marine mollusca of the Scottish national Antarctic expedition.
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1980 Food specialization and the evolution of predatory Prosobranch
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1925 Gastropoda der deutschen tiefsee-expedition. II. Deutsche-Tiefsee
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Theil I. 376 p.

42 NEMOURIA No. 27

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1928 Reports on the marine mollusca in the collections of the South
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1884 Manual of Conchology 3. Author, Philadelphia.
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1882 Mollusca of the H.M.S. Challenger Expedition. Part XIV. J. Linn.
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Wenz, W.

1941 (1938-1944). Handbuch der Palaozoologie (O.H. Schindewolf,
ed.) Borntraeger, Berlin.

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Editor
JOHN E. DU PONT

Managing Editor
DAVID M. NILES

Editorial Board
DEAN AMADON, American Museum of Natural History, New York, NY
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
Kennett Pike, Greenville,
Delaware 19807

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Occasional Papers of the Delaware Museum of Natural History

NUMBER 28 OCTOBER 28, 1983

NOTES ON VOLES
(MICROTUS, FAMILY CRICETIDAE)
IN NEW MEXICO

John P. Hubbard*, Charles S. Thaeler, Jr.**,
and C. Gregory Schmitt*

INTRODUCTION

Five species of voles of the genus Microtus are known from New Mexico,
these being the meadow vole (M. pennsylvanicus), montane vole
(M. montanus), Mexican vole (M. mexicanus), long-tailed vole
(M. longicaudus), and prairie vole (M. ochrogaster). Distributional and
related data on these species in the state have been presented by various
workers, most extensively by Findley et al. (1975). However, new data have
come to light in recent years and are presented in this paper. Much of this new
information is the result of surveys by the New Mexico Department of Game
and Fish, and the specimens cited are in the Departmental collection
(NMGEF) unless otherwise noted.

SPECIES ACCOUNTS

Microtus pennsylvanicus (Ord.)

Findley et al. (1975) map this species mainly in northern New Mexico
(San Juan, Taos, Mora, Colfax, Santa Fe, and San Miguel counties), with
outlying populations in Valencia (=Cibola) and Catron counties. It is
particularly the latter two populations that have been of interest to us, as their
existence has not been confirmed since early in this century. Studies of older

*New Mexico Department of Game and Fish
Santa Fe, New Mexico 87503
**New Mexico State University Library of Congress
Las Cruces, New Mexico 88003 Catalog Card No. 83-72994

Z NEMOURIA No. 28

material (Anderson 1961; Anderson and Hubbard 1971) indicate that neither
population is subspecifically distinct, but some differentiation has occurred in
them and their relictual nature makes them of interest and concern.

The Cibola County population was reported by Bailey (1932) as M. p.
modestus (Baird), from the San Rafael area. Three specimens verifying the
existence of this population are at the National Musuem of Natural History
(Biological Survey Collection), according to Findley et al. (1975).

Our attempts to relocate this population of M. pennsylvanicus have been
thwarted by the loss of habitat in the San Rafael vicinity, namely the
eradication of the marshes in which the species was originally found. Marshes
do remain somewhat to the northeast, along Interstate Highway 40 east of
Grants. We trapped in that area in July 1977, but we failed to find
M. pennsylvanicus there — nor did Bruce Hayward find it in 1979. We
suspect that this population is extinct.

The Catron County population of M. pennsylvanicus is known from a
series of specimens taken in 1915 by J. Stokley Ligon in the Tularosa Valley,
seven to eight miles (11-13 km) southwest of Aragon. Five specimens from
this series have previously been reported by Anderson (1961), Anderson and
Hubbard (1971), and Findley et al. (1975), these being housed at the
American Museum of Natural History and the San Diego Museum of Natural
History. To these may be added two additional Ligon specimens, both at Yale
Peabody Museum: adult male (YPM 2034), 15 February 1915, and adult
female (YPM 2035), 16 February 1915; both were taken eight miles
southwest of Aragon.

Our attempts to relocate the Catron County population of M. pennsylvanicus
have been confined to the upper San Francisco Basin, including the drainages
of Tularosa Creek (especially near Aragon and at Tularosa Marsh), Apache
Creek (Fitzgerald Cienaga), and Centerfire Creek (Centerfire Bog). To date,
we have not found M. pennsylvanicus in the area, and we are tempted to agree
with others (e.g. Anderson and Hubbard 1971, Findley et al. 1975) that the
population there may be extinct.

In addition to our efforts in Catron County, others have sought
M. pennsylvanicus there without success. For example, both Sydney
Anderson and Bruce Hayward (pers. comm.) have trapped in the Tularosa
Creek area, taking only M. mexicanus there. J. S. Ligon also took the latter
species there in 1915, along with the aforementioned specimens of
M. pennsylvanicus. More recently, Krausman and Bahti (1980) reported
having taken M. pennsylvanicus at nearby Centerfire Bog, some 24 km
northwest of the Ligon locality. However, their specimen (NMGF 647), an
adult female taken on 13 August 1978, is clearly M. mexicanus in skull
characters, color, and tail length (standard measurements 1 40-28-20-15 mm).

In addition to our searches for M. pennsylvanicus in Cibola and Catron
counties, we have also sought the species in other parts of New Mexico,
particularly in the northeastern third of the state. There we found it in several
areas not recorded by Findley et al. (1975), the most notable being in Union

October 28, 1983 Hubbard, Thaeler, & Schmitt 3

County, where six specimens were taken along Pinabete Creek, 1.5 km north
and 8 km west of Des Moines on 7 March 1978. We have also taken numerous
specimens at new localities between Union County and the foothills of the
Rocky Mountains, in Colfax County (Stubblefield Lake; Cimarron River, 0.8
km south of Springer; UU Bar Ranch, 1.5 km east of Miami Lake), in Mora
County (Watrous area; Cherry Lake), and in San Miguel County (McAllister
Lake) — all in the period 6 May 1977 to 23 March 1978.

Findley et al. (1975) and earlier workers have pointed out the association
of this vole with mesic habitats in New Mexico, and our data generally support
this contention. This is particularly evident in the San Juan Valley, in
northwestern New Mexico, where Schmitt collected numerous specimens
from areas dominated by grasses, sedges, rushes (Juncus), and similar growth
— typically in wet or very wet sites.

In northeastern New Mexico, we found M. pennsylvanicus in a somewhat
broader array of habitats. These ranged from dense stands of graminoids and
forbs along Interstate Highway 25 (Watrous area) to very wet areas of sedges
(e.g. Stubblefield Lake) and periodically inundated areas of Juncus (e.g.
McAllister Lake). In general, we found that broader habitat usage occurred in
M. pennsylvanicus where it was the only Microtus found. Where other
species occurred, we found M. pennsylvanicus at the most mesic end of the
habitat spectrum. For example, M. pennsylvanicus and M. ochrogaster were
found in close proximity near Miami Lake. There the former was taken in
runways at the edge of a live stream in stands of Juncus, while M. ochrogaster
was taken in adjacent dense grass. However, at Springer and Wagon Mound,
where the two species also occur together, we found both in mesic habitats.
M. pennsylvanicus also occurs in close proximity to M. mexicanus in
northeastern New Mexico, although we have not taken them together (also see
M. ochrogaster account).

In general, we find that our specimens of M. pennsylvanicus from north-
eastern New Mexico closely resemble those of the San Juan Valley in color
and size. Hence, we follow Anderson and Hubbard (1971) in including both
populations in the race M. p. modestus (Baird). Interestingly, six (16.2%) of
the 38 specimens in our series from the San Juan Valley have the posterior
loop on M2 obsolete or reduced. By contrast, all of our specimens from
northeastern New Mexico showed this loop to be discrete, if not well-
developed.

Microtus montanus (Peale)

Our surveys in southwestern New Mexico revealed the presence of this
species in Catron County at Centerfire Bog. There, Marshall Conway took a
total of 10 specimens on 27 June and 21 October 1978 and 12 April 1979. We
have assigned these specimens to M. m. arizonensis Bailey, and this
represents the first unequivocal record of this form to be published for New
Mexico.

4 NEMOURIA No. 28

Bailey (1932) records this race of M. montanus from the Jemez and Zuni
mountains, based on single specimens; however, these were subsequently
reallocated by Anderson (1959) to M. longicaudus. The reallocation of the
Zuni Mountains record is regarded as questionable by Don Wilson (in litt. ),
and the specimen — a juvenile taken in 1874 by H. W. Henshaw and now
preserved at the U. S. National Museum of Natural History — may never be
properly allocated to species. Certainly, even if it were M. montanus, it could
not be confidently identified subspecifically. With regard to the Jemez
Mountains, M. montanus does occur there, Bailey's misidentification
notwithstanding. That population, and others in northern New Mexico, have
been assigned by Anderson (1959) to M. m. fusus Hall.

The generally accepted range of M. m. arizonensis is in the White and Blue
mountains of Arizona and their vicinity (Cockrum 1960). Centerfire Bog is
less than 35 km east of the nearest Arizona locality, and therefore the
occurrence of this vole in New Mexico is not unexpected. The New Mexican
population occurs in a wet sedge and grass meadow (elevation ca. 2100 m),
bordering marshes and open water. Because of the restricted range and
vulnerability of its habitat, M. m. arizonensis was listed as endangered in New
Mexico in 1979 (Game Commission Regulation No. 599) and again in 1983
(Regulation No. 624).

We have not found M. montanus otherwise outside the range outlined by
Findley et al. (1975), but Dalquest (1975) reported the species from Colfax
and Union counties in northeastern New Mexico. However, our examination
of his material shows all of his specimens to be M. mexicanus (see below).

Microtus mexicanus (Saussure)

Findley et al. (1975) map this species in New Mexico primarily in the
southern highlands, north to McKinley and Valencia (=Cibola) counties
(Zuni Mountains and the Mt. Taylor range) and Bernalillo County (Sandia
Mountains). However, the species is known to range north to southern Utah
and southwestern Colorado (Hall 1981), and we here report it farther
northeast as well, i.e. in Colfax and Union counties.

As noted in our discussion of M. montanus, Dalquest (1975) erroneously
reported as that species a series of M. mexicanus taken in northeasternmost
New Mexico. These specimens, housed at Midwestern University, Wichita
Falls, Texas, and kindly lent to us by Dalquest, were taken at Raton Pass,
Colfax County, and near Capulin Mountain National Monument, Union
County, in August 1966. They clearly represent M. mexicanus, as Dalquest
(in litt.) now agrees. Dalquest (1975) found numerous vole runways on the
Colorado side of Raton Pass, and he surmised that M. “montanus”
(= mexicanus) would also be found in that area of the state. Armstrong (1972)
does not record M. mexicanus from southeastern Colorado.

M. mexicanus was also taken at Capulin Mountain National Monument by
A. L. Gennaro and his associates in 1975, 1976, and 1977 (Eastern New

October 28, 1983 Hubbard, Thaeler, & Schmitt 5

Mexico Museum of Natural History collection). In addition, we have taken
specimens on nearby Sierra Grande in the period from 1976 to 1982. Finally,
Donald Hoffmeister (in litt.) reports M. mexicanus as having been taken by
Woodrow Goodpaster south of Cimarron, Colfax County, in 1975. These
records extend the range of this species some 180 to 280 km northeast of the
Sandia Mountains, the previous northeasternmost known area of occurrence
in the state. Whether these occurrences in northeastern New Mexico represent
a recent range extension or are the result of past oversight, we cannot say.
M. mexicanus is now also frequent in the Sandia and Manzano mountains,
where previously rare or not found at all (Findley et al. 1975).

Another somewhat unexpected record of M. mexicanus is from the
marshes east of Grants, Cibola County, where we took two specimens on
7 July 1977. Hayward also took specimens of M. mexicanus there in 1979,
and these are housed at Western New Mexico University. Findley et al.
(1975) recorded the species in that region of the state only in montane areas,
except possibly for a series in the American Museum of Natural History
labelled as Thoreau, McKinley County. However, this series is probably
actually from Cottonwood Gulch, a montane area, to the south in the Zuni
Mountains (D.A. McCallum pers. comm.).

In our experience, M. mexicanus occurs in a wide variety of habitats in
New Mexico, from fairly dry montane meadows to rather mesic sites at middle
elevations. In the latter regard, we have found the species in the edges of cattail
(Typha sp.) marshes east of Grants and at Apache Creek, where this was the
only vole taken. At Centerfire Bog, M. mexicanus was not found in such mesic
sites; instead these were occupied by M. montanus arizonensis, while
M. mexicanus occurred in drier graminoid-forb areas, fringing the wettest
sites.

We did not undertake a close examination of possible geographic variation
in M. mexicanus from New Mexico, as this has been addressed by Findley
and Jones (1962) and is under study by Hoffmeister (in litt.). However, we did
note a peculiarity in the dental patterns of some M. mexicanus specimens from
the upper San Francisco River drainage. In nine of our 48 specimens (18.8%),
we found a well to moderately developed posterior loop on M2. This initially
caused some confusion in specimen identification. However, in their reddish
coloration and short tails, the specimens showing this loop are otherwise
typical of M. mexicanus.

Microtus longicaudus (Merriam)

This species occurs in the higher uplands almost statewide in New Mexico.
However, it appears to be absent from several outlying areas, including the
Animas, Guadalupe, Pinos Altos, San Mateo, Magdalena, and Datil
mountains (Findley et al. 1975). It was, therefore, a matter of considerable
interest to find the species on Sierra Grande, Union County, during the course
of our surveys there in 1976. We collected three specimens on 1-2 July and 22
September, in open woodlands of blue spruce (Picea pungens) and aspen

6 NEMOURIA No. 28

(Populus tremuloides). There this species was outnumbered in traps by
M. mexicanus. This locality is about 100 km north-northeast of the Cimarron
area, the nearest previously known site for M. longicaudus.

Microtus ochrogaster (Wagner)

It was a matter of some surprise when Rowlett (1972) reported taking this
prairie species at Philmont Scout Camp, Colfax County, in northeastern New
Mexico in 1968. Until now, that occurrence represented the only published
record from the state, although we later learned that Woodrow Goodpaster
took specimens in the same general area in 1975 and 1977 (Donald
Hoffmeister in litt.). As a consequence of its apparently restricted range and
probably vulnerability, M. ochrogaster was listed as endangered in New
Mexico by the Game and Fish Department in 1975 (Regulation No. 563).
Subsequently, the Department began studies to determine more about the
status of this species in the state, an investigation that led to its being delisted in
1979.

Surveys conducted by the Department in the period 1976-1978 revealed
that M. ochrogaster persisted at Philmont Scout Camp and that it occurred
elsewhere in Colfax County. It was also found in Mora County, where two
Specimens were taken near Wagon Mound on 25-26 October 1976 by
Conway. In Colfax County, collections included two specimens from 8 km
east and 3 km south of Cimarron on 6 May 1977; two from the U U Bar Ranch,
1.5 km east of Miami Lake on 6 May 1977; and a specimen from Springer on
the Cimarron River on 10 March 1978. Surveys elsewhere failed to yield the
species, which appears to be restricted in New Mexico to the uppermost
Canadian River Basin.

Habitat use by M. ochrogaster populations in New Mexico is variable. For
example, we have taken the species in fairly tall and rather rank stands of
grasses and forbs, in dense meadow-like areas, in wet graminoid marshland,
and in saline Juncus flats. Where the species occurs with M. pennsylvanicus,
we have seen evidence both for and against habitat-partitioning (see
M. pennsylvanicus account). We have not taken M. ochrogaster with any
species other than M. pennsylvanicus, but Hoffmeister (in litt.) reports that
Goodpaster took the former and M. mexicanus together in roadside vegetation
at a locality 4 miles (6.5 km) south and 2 miles (3 km) west of Cimarron.

Rowlett (1972) and Findley et al. (1975) did not address the subspecific
status of M. ochrogaster in New Mexico. However, Severinghaus (1976)
assigned this population to M. o. haydenii (Baird), the central and southern
Great Plains race of the species (type locality: Ft. Pierre, Stanley County,
South Dakota). Severinghaus’ concept of this race includes M. o. taylori(type
locality: 0.5 mile [or 0.8 km] north of Fowler, Meade County, Kansas), which
was characterized by its describers as being larger and darker than M. o.
haydenii (Hibbard and Rinker 1943). In a preliminary study of geographic
variation in M. ochrogaster in the central and southern Great Plains, Conway

October 28, 1983 Hubbard, Thaeler, & Schmitt i

(ms) found that topotypical specimens of M. o. taylori average darker than
most populations assigned to M. o. haydenii and that New Mexico specimens
also tend to be dark. However, these southern populations intergrade in this
character with more northern ones through Colorado. In addition, most
mensural characters, including both standard and cranial, show either clinal or
mosaic patterns of intergradation. Under the circumstances, we follow the
subspecific treatment of Severinghaus (1976) — pending more detailed
analysis of the situation.

Acknowledgements

We thank the following for their contributions to this study: S. Anderson,
M. C. Conway, W. W. Dalquest, J. S. Findley, A. L. Gennaro, M. D. Hatch,
B. J. Hayward, S. Judd, P. R. Krausman, T. L. Yates, and owners on whose
lands specimens were taken. We thank Midwestern University and the
universities of Illinois, Kansas, and New Mexico for the loan of specimens.
Sydney Anderson confirmed the identification of the Krausman and Bahti
specimen of M. mexicanus and of the New Mexico series of M. montanus
arizonensis for us. Donald Hoffmeister also confirmed the Dalquest
specimens as being M. mexicanus instead of M. montanus. Finally, we thank
S. Anderson, M. C. Conway, D. A. McCallum, D. E. Wilson, and T. L. Yates
for reading and commenting on earlier versions of the manuscript.

LITERATURE CITED

Anderson, S.
1959 Distribution, variation, and relationships of the montane vole,
Microtus montanus. Univ. Kansas, Mus. Nat. Hist. Publ.
9:415-511.

1961 A relict population of Microtus pennsylvanicus in southwestern
New Mexico. Amer. Mus. Novitates no. 2034: pp. 1-3.

Anderson, S. and J. P. Hubbard
1971 Notes on the geographic variation of Microtus pennsylvanicus
(Mammalia, Rodentia) in New Mexico and Chihuahua. Amer.
Mus. Novitates no. 2460: pp. 1-8.

Armstrong, D.
1972 Distribution of mammals in Colorado. Univ. Kansas Mus. Nat.
Hist., Monograph no. 3: pp. 1-415.

8 NEMOURIA No. 28

Bailey, V.
1932 Mammals of New Mexico. N. Amer. Fauna no. 53: pp. 1-412.

Cockrum, E. L.
1960 The recent mammals of Arizona. Univ. Arizona Press, Tucson,
276 pp.

Dalquest, W. W.
1975 The montane vole in northeastern New Mexico and adjacent
Colorado. Southwestern Nat. 20:138-139.

Findley, J. S. and C. J. Jones
1962 Distribution and variation of voles in the genus Microtus in New
Mexico and adjacent areas. J. Mammal. 43:154-166.

Findley, J. S., A. H. Harris, D. E. Wilson, and C. Jones
1975 Mammals of New Mexico. Univ. New Mexico Press, Albuquerque,
360 pp.

Hall, E. R.
1981 The mammals of North America. John Wiley and Sons, New York.
Vol. II, pp 601-1181 + vii and 90.

Hibbard, C. and G. Rinker
1943 Anew meadow mouse (Microtus ochrogaster taylori) from Meade
County, Kansas. Univ. Kansas Sci. Bull. 29:255-268.

Krausman, P.R. and K. Bahti
1980 Microtus pennsylvanicus in southwestern New Mexico. South-
western Nat. 25:260-261.

Rowlett, R.
1972 First records of Eumops perotis and Microtus ochrogaster in New
Mexico. J. Mammal. 53:640.

Severinghaus, W.
1976 Systematics and ecology of the subgenus Pedomys. Unpublished
Ph.D. thesis, Univ. Illinois, 174 pp.

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DAVID M. NILES

Editorial Board
DEAN AMADON, American Museum of Natural History, New York, NY
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburg, PA
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
P.O. Box 3937, Greenville,
Delaware 19807

AL ae I oo
671
436 heyy aa 0/86

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Occasional Papers of the Delaware Museum of Natural History

NUMBER 29 DECEMBER 31, 1985

BIBLIOGRAPHY OF PHILIPPINE ORNITHOLOGY

Robert S. Kennedy!
Edward C. Dickinson?
Murray D. Bruce?

This bibliography was originally intended to be a section of the Checklist of
Philippine Birds that Dickinson and Kennedy, along with Kenneth C. Parkes,
plan to have completed in the near future. However, as we located more and
more references on Philippine birds, some that could be cited in the Checklist
but many that could not be, it became clear that our contribution to the
knowledge and development of ornithology and conservation in the Philippines
would be greatest if we published citations of all the references we found in a
separate bibliography.

With recruitment of the bibliographic skills of Murray Bruce, who has one
of the most extensive collections of references on South-east Asian birds in the
world, we have attempted to make this as close to a “complete” bibliography of
Philippine ornithology as we could make it. In offering this statement, however,
we realize other researchers will locate references that we have missed. We hope
this will be the case and, in turn, we hope they will send one of us a copy of the
article or will let one of us know about it so we may include it in a future ad-
dendum.

For this bibliography, we define references that deal with Philippine birds as
those specifically written about Philippine birds or those that contribute new
information on the biology, distribution, or taxonomy of Philippine birds,
particularly the endemics. References that briefly mention a Philippine form or
that do not contribute new information, as well as avicultural notes prior to
1970, have usually been excluded.

1Department of Zoology, Library of Congress
Washington State University, Catalog No. 85-073726
Pullman, Washington 99164-4220 USA.
2Chemin du Chano 8, 1802 Corseaux,
SWITZERLAND.
38 Spurwood Road, Turramurra N.S.W.
2074 AUSTRALIA

<qnnTHSONIAyy™

ran 2 8 1886
_LIBRARIES

2 NEMOURIA No. 29

We have not set limits to the publication dates of the references thus they
span nearly three centuries. The earliest reference listed is Camel (1703) and the
most recent ones are those accepted for publication but not in print when this
bibliography was prepared.

PLAN OF THE BIBLIOGRAPHY

The bibliography will be published in two parts. Part 1 (presented here)
consists of three sections: Section I, List of Books and Checklists on Philippine
Birds; Section II, Master List of References; and Section III, List of Recent
General Works. The contents of each section will be outlined below. Part 2,
which will appear as a future number in Nemourza, will contain an Addendum to
Part 1, an Index to Species, and an Index to Islands. The contents of Part 2 will
be treated in that publication.

Reference Numbers. — Entries within each section of the bibliography are
arranged alphabetically by the last name of the authors and by publication date
when two or more references appear under one author. All entries in the
bibliography are sequentially numbered beginning with the first citation in
Section I and ending with the last one in Section III. These numbers will be used
to reference citations in the index sections of Part 2.

Journal Abbreviations.—With few exceptions, we have used the Serial
Sources for the BIOSIS Data Base for journal abbreviations if they were listed in
this publication. If they were not listed, we patterned the abbreviations after
those used in the BJOSIS Data Base. We deviated from the abbreviations in the
BIOSIS Data Base in a few places, namely by shortening Philippi. to Phil. and
Ornithol. to Orn.

Citations.—We generally follow the style used in The Auk for citations
used throughout the bibliography. We deviate from this style by including page
numbers for books or monographs when we were able to obtain this information.
For articles published without a title, we have provided one and have indicated
that we have done so by enclosing it in brackets. We have included issue
numbers usually only when pagination reverts to “1” with each issue within a
volume.

Annotations. — Because many of the articles and books that contain original
descriptions are difficult to locate, we provide a brief annotation for each that
includes the scientific name of the species or subspecies that we expect to be in
the Checklist and the page(s) where the description originally appeared. For book
entries where only specific chapters deal with birds of the Philippines, we
provide an annotation that includes the island name or Philippines, and the first
and last pages of the chapter. For example in no. 57, Barnes and McClure
(1966), we add “[Philippines, pp. 4-5.]” after the regular citation.

December 31, 1985 Kennedy, et al. 3

Section I, Books on Philippine Birds

This section of the bibliography contains titles of books or checklists that
deal partly or entirely with Philippine birds. We wish to point out that
Hachisuka’s The Birds of the Philippine Islands with Notes on the Mammal Fauna
was never completed. In his preface to Vol. I], Part 4, Hachisuka refers to
“Volume III” that would contain the remaining families in the order
Passeriformes not included in Part 4. Regrettably, this volume was never
published and we do not know if the manuscript (if completed) still exists. Also,
we wish to note that Rand’s (1970) unpublished manuscript is on microfilm in
the Division of Birds at the U. S. National Museum.

Section II, Master List of References

The Master List of References contains articles published in more than 100

~scientific or quasi-scientific periodicals around the world. It also contains titles of

obscure books in which species or subspecies of Philippine birds were first
described.

Section III, Recent General Works

In this section we include recent field guides or checklists covering
neighboring regions in Asia; papers, monographs or books on families or orders
of birds with species found in the Philippines; and general references that treat
Philippine birds. Although this list is by no means complete, we provide these
titles to familiarize beginning students of Philippine ornithology with other
references available to them.

ACKNOWLEDGEMENTS

We are particularly grateful to the curators and librarians of the American
Museum of Natural History, Australian Museum Sydney, British Museum
(Natural History), Delaware Museum of Natural History, Field Museum of
Natural History, National Library of Australia at Canberra, U. S. National
Museum, and University of Sydney (Badham and Fisher Libraries) for allowing
access to, and for providing help in the use of, their library facilities.

Specific help has been given at various points during the completion of this
project by Angel C. Alcala, Jesus B. Alvarez, Jr., John E. duPont, Luz C.
Gonzales, Pedro C. Gonzales, Jurgen Heucke, Ben F. King, H. Elliott McClure,
Joe T. Marshall, Jr., Gerlof F. Mees, Kenneth C. Parkes, Ricardo G. Paloma,
Dioscoro S. Rabor, Charles A. Ross, and George E. Watson. All are gratefully
thanked as are those authors who have, over the years, sent copies of their papers
to one or more of us.

Two unpublished papers helped us in our task: Rand (1970), “An annotated

4 NEMOURIA No. 29

list of Philippine birds,” which was kindly made available by the U. S. National
Museum, and an outline bibliography prepared by Kenneth C. Parkes in 1963.

Kennedy was kindly provided support for this bibliography and for his
Philippine research by the Ferdinand E. Marcos Foundation, Philippine Bureau
of Forest Development, World Wildlife Fund, National Geographic Society,
Philippine Airlines, San Miguel Corporation, Filipinas Foundation, The
Haribon Society, and the Frank M. Chapman Memorial Fund of the American
Museum. He was also provided clerical assistance by the Department of Zoology
at Washington State University and the Raptor Information Center of the
National Wildlife Federation.

Finally, we would like to thank the Editorial Board of the Delaware
Museum of Natural History for their encouragement and for their agreement to
publish this bibliography in Nemouria.

December 31, 1985 Kennedy, et al. 5

SECTION I

BOOKS AND CHECKLISTS ON PHILIPPINE BIRDS

1. ALCALA, A. C. 1976. Philippine land vertebrates: field biology.
Quezon City, Philippines, New Day Publ. 176 pp.

2. BRUCE, M. D. 1980. A field list of the birds of the Philippines.
Sydney, Australia, Traditional Explorations. 26 pp.

3. CASTANEDA, P. 1977. A portfolio of Philippine birds. Manila,
Filipinas Foundation, Inc. 5 pp., 30 pls.

4. DANCEL-CORNISTA, P., & L. P. CASTRO. 1975. Philippine animal
wildlife. Manila, McCullough Printing Co. 70 pp.

5. DELACOUR, J., & E. MAYR. 1946. Birds of the Philippines. New
York, Macmillan. 309 pp.

6. DICKERSON, R. E. 1928. Distribution of life in the Philippines.
Monogr. 21. Manila, Bur. Sci. 322 pp. [Birds of the Philippines, pp.
168-213, written by R. C. McGregor.]

7. DUPONT, J. E. 1971. Philippine birds. Greenville, Delaware,
Delaware Mus. Nat. Hist. 480 pp.

8. HACHISUKA, M. 1931-1935. The birds of the Philippine Islands with
notes on the mammal fauna. London, Witherby. Vol. 1, Part 1,
1931, pp. 1-168; Vol. 1, Part 2, 1932, pp. 169-439; Vol. 2, Part 3,
1934, pp. 1-256; Vol. 2, Part 4, 1935, pp. 257-468.

9. MCGREGOR, R. C. 1909 [-1910]. A manual of Philippine birds. Bur.
Sci., Manila. No. 2. Part 1, pp. 1-412; Part 2, pp. 413-769.

10. ——. 1940. An introduction to Philippine birds. Manila, Bur. Printing.
Popular Bull. 16. 180 pp.

11. ——, & E. J. MARSHALL. 1922. Philippine birds for boys and girls.
Manila, Bur. Printing.

12. ——, & -—. 1932. Philippine birds their haunts and _ habits.
Philadelphia, John C. Winston Co.

13. ——, & D. C. WORCESTER. 1906. A hand-list of the birds of the

Philippine Islands. Bur. Govt. Lab., Manila 36:1-123.

14,

bey

16.

NE

18.

NEMOURIA No. 29

MASON, C. R. 1980. Annotated check list of the birds of the Philip-
pines [Based on John E. duPont’s Philippine Birds.]. Orlando,
Florida, Russ Mason’s Natural History Tours, Inc.

RABOR, D. S. 1977. Philippine birds and mammals. Diliman, Quezon
City, Philippines, Univ. Phil. Press. 284 pp. [Birds, pp. 1-200.]

RAND, A. L. 1970. An annotated list of Philippine birds. Ms., xxii, 511
Pp.

WARDLAW RAMSAY, R. G. 1881. Revised list of the birds known to
occur in the Philippines showing their geographical distribution. Pp.
653-660 im The ornithological works of Arthur, Ninth Marquis of
Tweeddale (R. G. Wardlaw Ramsay, Ed.). London, Taylor &
Francis. 760 pp.

WORCESTER, D. C., & F. S. BOURNS. 1898. Contributions to
Philippine ornithology. Part I.—A list of the birds known to inhabit
the Philippine and Palawan Islands, showing their distribution
within the limits of the two groups. Proc. U. S. Nat. Mus.
20(1134):549-566.

December 31, 1985 Kennedy, et al. ql

SECTION II

MASTER LIST OF REFERENCES

19. ADAMS, C. C. 1905. Types of birds in the Beal-Steere Collection of the
University Museum, University of Michigan. Univ. Bull., Univ.
Michigan N.S. 6(2):23-28.

20. AGALOOS, L. C., & P. M. NEPOMUCENO. 1977. Ecology of
Calauit Island. Sylvatrop Phil. For. Res. J. 2:163-178.

21. ALCALA, A. C., & P. ALVIOLA, III. 1970. Notes on the birds and
mammals of Boracay, Caluya, Carabao, Semirara and Sibay Islands,
Philippines. Silliman J. 17:444-454.

22. ——,&E.E. CARUMBANA. 1975. An ecological study of certain game
birds in southern Negros Oriental, Philippines. NRCP Res. Bull.
30:91-119.

23. ——, & ——. 1980. Ecological observations on birds of southern
Negros, Philippines. Silliman J. 27:197-222.

24. ——, & W. M. SANGUILA. 1969. The birds of small islands off the

eastern coast of Panay. Silliman J. 16:375-383.

25. ALCASID, G. L. 1965. Terpsitphone atrocaudata on Mindoro, Philip-
pines. Auk 82:644.

26. ——, & P. GONZALES. 1968. A new race of Naked-faced Spider-
hunter (Arachnothera clarae) from Luzon. Bull. Br. Orn. Club
88:129-130.

27. ——, & ——. 1970. Kalikasan mga ibon: A guide to the National

Museum exhibition of the same title and a preliminary manuscript
about some Philippine birds. Manila, Graphic Publishing Inc. 46

Pp.

28. ALVAREZ, J. B., JR. 1970. A report on the 1969 status of the Monkey-
eating Eagle of the Philippines. Pap. Proc. Int. Union. Conserv.
Nat. Nat. Resour. llth Technical Meeting, New Delhi, India, 25-28
November 1969, Vol. II:68-73.

29. ——. 1978. Change of name of the Monkey-eating Eagle. President’s
letter. Int. Counc. Bird Preserv. No. 44, p. 11.

30.

alt

a2.

33.

34.
35.

36.

Sie

38.

ahek
40.

41.

42.

43.

44.

45.

NEMOURIA No. 29

ALVIOLA, P. L. II, F. F. SANCHEZ, & E. A. BENIGNO. 1973.
Notes on the feeding habits of three species of Philippine weavers of
the genus Lonchura. Kalikasan Phil. J. Biol. 2:149-153.

AMADON, D. 1942. Birds collected during the Whitney South Sea
Expedition, XLIX. Notes on some non-passerine genera, 1. Am.
Mus. Novit. 1175:1-1l.

——. 1943. The genera of starlings and their relationships. Am. Mus.
Novit. 1247:1-16.

——. 1951. Notes on Chinese Egret, Egretta eulophotes (Swinhoe). Phil.
J. Sci. 80:53-54.

—-. 1952. A bird new to Palawan. Phil. J. Sci. 81:139.

——. 1953. Remarks on the asiatic hawk-eagles of the genus Spzzaetus.
Ibis 95:492-500.

—. 1955. In defence of the principle of the “first revisor.” Bull. Br.
Orn. Club 75:21-23.

—. 1956. Remarks on the starlings, family Sturnidae. Am. Mus.
Novit. 1803:1-41.

—. 1959, Remarks on the subspecies of the Grass Owl, Tyto capensis.
J. Bombay Nat. Hist. Soc. 56:344-346.

——. 1962. A new genus and species of Philippine bird. Condor 64:3-5.

—., & J. E. DUPONT. 1970. Notes on Philippine birds. Nemouria 1:1-
14.

—, & S. G. JEWETT, JR. 1946. Notes on Philippine birds. Auk
63:541-559.

—, & G. WOOLFENDEN. 1952. Notes on the Mathews’ collection
of Australian birds. The order Ciconiiformes. Am. Mus. Novit.
1564:1-16.

AMOR, A. J. 1981. The bird man of the Philippines. Filipinas. J. Sci.
Cult. 2:49-53.

ANONYMOUS. 1949. The Monkey-eating Eagle. Zoo Life Suppl.
4(1):1-2.

—. 1965. Philippines Monkey-eating Eagle. Conserv. News S. E.
Asia. No. 6, p. 45.

December 31, 1985 Kennedy, ez al. 9

46. ——.1970. The Philippines: Protection broadened for Monkey- eating
Eagles. Int. Union Conserv. Nat. Nat. Res. Bull. 2:147.

47. ——. 197la. Recovery round-up: Wedge-tailed Shearwater Puffinus
pacificus. Aust. Bird Bander 9:89.

48. —. 1971b. Telling the people in the Philippines [re. Monkey- eating
Eagle]. Oryx 11:97.

49. ——. 1973. Food habits of three species of Philippine weavers of the
genus Lonchura. An. Rep. Rodent Res. Cent., pp. 65-68.

50. ——. 1974. Biological and toxicological studies on the Philippine
weavers. An. Rep. Rodent Res. Cent., pp. 76-83.

51. ——. 1975a. Affenadler (Pithecophaga jeffery1) steht vor der Ausrottung.
Gefiederte Welt 96(7):139.

52. ——. 1975b. Brutversuch mit dem Affenadler (Pithecophaga jefferyi).

Gefiederte Welt 97(9):180.

53. BAKER, E. C. S. 1920. Some notes on the genus Surniculus. Novit.
Zool. 26:291-294.

54. BAKER, R. H. 1948. Report on collections of birds made by U. S.
Naval Medical Research Unit No. 2 in the Pacific war area.
Smithson. Misc. Collect. 107(15):1-74.

55. BANGS, O. 1919. A new striated grass-warbler from the Philippines.
Proc. New England Zool. Club 7:5-6.

56a. ——. 1922. Notes on Philippine birds collected by Governor W.
Cameron Forbes. Bull. Mus. Comp. Zool. 65:77-84.

56b. ——. 1930. Types of birds now in the Museum of Comparative
Zoélogy. Bull. Mus. Comp. Zoél. 70:147-426. [Ceyx cyanopectus,
Serilophus steeret Pericrocotus cinereus, Lalage niger mitifica, Ortho-
tomus ruficeps nuntius, Megalurus palustris forbest, Parus quadri-
vittatus, Zosterops forbes1, Oriolus xanthonotus persuasus].

57. BARNES, C. M., & H. E. MCCLURE. 1966. M.A.P.S. annual
narrative report 1965. [Philippines, pp. 4-5.]
58. BASAN, A. C. 1976. Winged lightning. Int. Wildl. 6(1):18-19.

59. BAUD, F. J. 1976. Oisepux des Philippines de la collection W. Parsons.
I. Cebu, Samar, Romblon, Tablas et Sibuyan. Rev. Suisse Zool.
83:497-513.

10

60.

61.

62.

63.

64.

65.

66.
Cr

68.

69.

70.

Lh,

2,

fey

74.

75.

NEMOURIA No. 29

—. 1978. Oiseaux des Philippines de la collection W. Parsons. II.
Luzon, Mindoro et Palawan. Rev. Suisse Zool. 85:55-97.

BEECHER, W. J. 1978. Feeding adaptations and evolution in the
starlings. Bull. Chicago Acad. Sci. 11:269-298.

BELL, J. 1973. A tale of two eagles [re. Pithecophaga jefferyi]. Anim.
Kingd. 76(4):2-7.

BENIGNO, E. A., P. L. ALVIOLA, III, F. F. SANCHEZ, M. M.
HOGUE, & G. V. LLAGUNO. 1975. Damage potential of three
species of Philippine weavers, (Lonchura). Kalikasan Phil. J. Biol.
4:141-144.

BLASIUS, W. 1888a. Verein ftir Naturwissenschaft. 10. Braunschweig.
Anz. 37:335-336.

—. 1888b. Verein ftir Naturwissenschaft. ll. Braunschweig. Anz.
52:467-468.

— .1888c. Die Végel von Palawan. Ornis 4:301-320.

—. 1888d. [Letter concerning Mr. Sharpe’s nomenclature of Palawan
birds.] Ibis (5) 6:372-375.

—. 1890a. Verein fiir Naturwissenschaft. ll. Braunschweig. Anz.

73:715-716.

—. 1890b. Verein fiir Naturwissenschaft. 12. Braunschweig. Anz.
87:877-878.

—. 1890c. Die wichtigsten Ergebnisse von Doctor Platen’s or-
nithologischen Forschungen auf den Sulu-Inseln. J. Orn. 38:137-144.

—. 1890d. Die von Herrn Doctor Platen und dessen Gemahlin im
Sommer 1889, bei Davao auf Mindanao gesammelten Vogel. J. Orn.
38:144-149.

— . 189la. Mittheilung iiber ein neues Polyplectron. J. Orn. 39:9-10.

—. 1891b. Polyplectron Nehrkornae nov. sp. Die Schwalbe (= Mitt.
Orn. Ver. Wien) 15:1-3.

—. 1897a. Kritische Bemerkungen iiber einige Végel von Mindoro. 10
Jahresbericht des Vereins fiir Naturwissenschaft zu Braunschweig.
pp. 157-162.

—. 1897b. Uebersicht tiber die Vogelfauna von Mindoro. 10
Jahresbericht des Vereins fiir Naturwissenschaft zu Braunschweig.
pp. 181-183.

December 31, 1985 Kennedy, ez al. 1]

76. BLYTH, E. 1842. Report of the Curator of the Museum of the Asiatic
Society of Bengal. J. As. Soc. Bengal 11:444-470.

77. ——. 1846. Notices and descriptions of various new or little known
species of birds [cont.]. J. As. Soc. Bengal 15:280-314.
78. ——. 1849 [=1852]. Catalogue of the birds in the Museum, Asiatic

Society. Calcutta, As. Soc. Bengal. 403 pp. [Prcoides maculatus
validirostris described, p. 64.]

79. ——. 1861. Report of Curator, Zoology Department, May & June. J. As.
Soc. Bengal 30:90-98.
80. ——. 1870. Notes relating chiefly to the brds of India. Ibis (2) 6:157-176.

81. BODDAERT, P. 1783. Table des planches enluminées d’Histoire
naturelle de M. d’Aubenton. Utrecht. 58 + [9] pp. [Descriptions of
Penelopides p. panini, p. 48; Penelopides panini manilloe, p. 54;
Coracina s. striata, p. 38; Copsychus saularis mindanensis, p. 38;
Hypothymis a. agurea, p. 41; Motacilla violacea (syn. for Sturnus p.
philippensis), p. 11; Turdus manilla (syn. for Monticola sohtarius
philippensis). |

82. BONAPARTE, C. L. J. L. 1850a. Conspectus generum avium.
Lugduni Batavorum, E. J. Brill. Vol. 1:1-543. [Descriptions of
Centropus molkenboeri (syn. for Centropus v. viridis), p.108; Halcyon
h. hombrom, p. 157; Pitta sordida muelleri, p. 256; Zosterops m.
meyent (nomen novum for Dicaeum flavum), p. 398.]

83. ——. 1850b. Nouvelles espéces ornithologiques. Premiére partie:
Perroquets. Compt. Rend. Acad. Sci. Paris 30:131-139.

84. —. 1853. Notes sur les collections de M. A. Delattre. Compt. Rend.
Acad. Sci. Paris 37:827-835.

85. ——. 1853-1854. Notes sur les Collections rapportées en 1853, par M.
A. Delattre, de son voyage en Californie et dans le Nicaragua.
Compt. Rend. Acad. Sci. Paris 37:913-925; 38:1-11, 52-66, 258-266,
378-389, 533-542, 570-572, 650-654. [cf. Pachycephala, Artamus,
Dicrurus, Coracina.]|

86. ——. 1854-1855. Coup d’oeil sur ordre des Pigeons. Compt. Rend.
Acad. Sci. Paris 39:869-880, 1072-1078, 1102-1112; 40:15-24, 96-
102, 205-221.

87. ——. 1855a. Notices ornithologiques. Compt. Rend. Acad. Sci. Paris
41:247-249.

12

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59:

90.

ol

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94.

95;

96.

97.

98.

99.

NEMOURIA No. 29

——. 1855b. Principales espéces nouvelles d’oiseaux qu'il vient d’ob-
server dans son récent voyage en Ecosse et en Angleterre. Compt.
Rend. Acad. Sci. Paris 41:651-661.

—. 1855c. Note sur les Salanganes et sur leurs nids. Compt. Rend.
Acad. Sci. Paris 41:976-979.

—. 1857 [1855-1857]. Conspectus generum avium. Lugduni
Batavorum, E. J. Brill. Vol. 2:1-232. [Descriptions of Treron
pompadora axillaris, p. 13; Phapitreron a. amethystina, p. 28.]

BONNATERRE, J. P. 1790 [1790-1791]. Tableau encyclopédique et
méthodique des trois régnes de la nature. Paris, Panckoucke. Vol. 1,
320 pp. Vol. 2, atlas of pls. [see Vieillot 1823] [ Tachybaptus ruficollis
philippensis described, p. 58.]

BONNIT, C. B., L. RUNDQUIST, & V. M. RUNDQUIST. 1975.
Status Report: The Monkey-eating Eagle of the Philippines. Mimeo,
Quezon City, Philippines, Dept. Nat. Res., Bur. Forest Dev., 110 pp.

BOURNS, F. S., & D. C. WORCESTER. 1894. Preliminary notes on
the birds and mammals collected by the Menage Scientific Ex-
pedition to the Philippine Islands. Occas. Pap. Minn. Acad. Nat.
Sci. 1:1-64.

BRODKORB, P. 1934. A new Pitta from Palawan, Philippine Islands.
Occas. Pap. Mus. Zool. Mich. 279:1-3.

—,R. B. GONZALES, & D. EMPESO. 1969. White-tailed Tropic-
bird, an addition to the Philippine avifauna. Auk 86:357.

BRONZINI, E. 1978. Bericht iiber einen langlebigen Affenadler
(Pithecophaga jeffery) in CGefangenschaft. Zool. Gart., Jena
48(4):299-300.

BROOKE, R. K. 1971. Geographical variation in the Little Swift Apus
affinis (Aves: Apodidae). Durban Mus. Novit. 9:93-103.

—. 1972. Geographical variation in palm swifts Cypsiurus Spp. (Aves:
Apodidae) Durban Mus. Novit. 9:217-229.

BRUCE, M. D. 1980. Ursula Island bird sanctuary. Pp.20-21 in The
Palawan Expedition, Stage I (M. Bruce, Ed.). Sydney, Australia,
Traditional Explorations. 47 pp.

December 31, 1985 Kennedy, et al. 13

100.

101.

102.

103.

104.

105.

106.

107.

108.

109.

110.

——. 198la. Bird notes from Ursula Island, southern Palawan. Pp. 64-65
in The Palawan Expedition, Stage II (M. D. Bruce, Ed.). Sydney,
Australia, Associated Research, Exploration and Aid Pty Ltd. 139

Pp.

—. 198lb. Final supplement to “A field list of the birds of the
Philippines.” Pp. 97-100 72 The Palawan Expedition, Stage II (M.
D. Bruce, Ed.). Sydney, Australia, Associated Research, Exploration
and Aid Pty Ltd. 139 pp.

BRUGGEMANN, F. 1876. Beitrage zur Ornithologie von Celebes und
Sangir. Abh. Naturwiss. Ver. Bremen 5:35-102. [Pitta kochi
described, p. 65.]

BRUSH, A. H. 1965. The structure and pigmentation of the feather tips
of the Scaled Cuckoo (Lepidogrammus cumingi). Auk 82:155-160.

BUNKER, P. D. 1916. Nesting of the Philippine Glossy Starling. Phil.
JeScrlk267.

BUTTIKOFER, J. 1893. A review of the genus Rhipidura, with an
enumeration of the specimens in the Leyden Museum. Notes
Leyden Mus. 15:65-98.

—. 1895a. A revision of the genus Turdinus and genera allied to it,
with an enumeration of the specimens contained in the Leyden
Museum. Notes Leyden Mus. 17:65-106. [Malacopteron palawanense
(nom. nov. for Turdinus rufifrons), p. 104.]

—. 1895b. On the genus Pycnonotus and some allied genera, with
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December 31, 1985 Kennedy, et al. 17

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NEMOURIA No. 29

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——. 1868. Description of a new species of the genus Ceyx. Proc. Zool.
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—. 1872. On two new species of birds. Ann. Mag. Nat. Hist. (4)
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GRANT, C. H. B. 1954. Ornithological nomenclature and the “first
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—. 1956. The “first reviser” and the name of the Philippine Pelican.
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GRAY, G. R. 1843. Some remarks on the soft-billed Duck of Latham.
[Descriptions of Falco guttatus (syn. for Falco s. severus) and Terp-
stphone cinnamomea untrufa (Tchitrea rufa).| Ann. Mag. Nat. Hist.
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— . 1844. List of the specimens of birds in the collection of the British
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——. 1844-1849. The genera of birds; comprising their generic
characters, a notice of the habits of each genus and an extensive list
of species referred to their several genera. London, Longman etc. 3
vols. [Descriptions of Collocalia troglodytes 1:55; Ptilinopus oc-
cipitalis, 2:467; Ducula p. poliocephala, 2:469.]

—. 1848-1868. List of the specimens of birds in the collection of the
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December 31, 1985 Kennedy, et al. 25

269. ——. 1861. List of species composing the family Megapodiidae, with
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270. GRAY, J. E. 1831. Description of two new species of birds discovered by
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271. GREENWAY, J. C., JR. 1943. Oriental forms of the Pygmy Wood-
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272. ——. 1973. Type specimens of birds in the American Museum of
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273. -——. 1978. Type specimens of birds in the American Museum of
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274. GRIMWOOD, I. R. 1978. Wildlife conservation in the Philippines.
Tigerpaper 5(4):9-11.

275. GRINNELL, J. 1938. In memoriam: Richard C. McGregor or-
nithologist of the Philippines. Auk 55:163-175.

276. GUILLEMARD, F. H. H. 1885a. Report on the collections of birds
made during the voyage of the yacht Marchesa.—I. A provisional list
of the birds inhabiting the Sulu Archipelago. Proc. Zool. Soc. Lond.,
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277. ——.1885b. Report on the collection of birds made during the voyage of
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278. -——.1886. The cruise of the Marchesa to Kamschatka and New Guinea.
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279a. ——. 1889. The cruise of the Marchesa to Kamschatka and New

Guinea, with notices of Formosa, Liu Kiu, and various islands of the
Malay Archipelago. 2nd Ed. London.

279b. GYLDENSTOLPE, N. 1926. Types of birds in the Royal Natural
History Museum in Stockholm. Ark. Zool. 19A:1-116. [Certhia
gularis = Nectarinia jugularis. |}

280. HACHISUKA, M. 1926a. [Remarks on the forms of night herons
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major.| Bull. Br. Orn. Club 46:103.

26 NEMOURIA No. 29

281. ——.1926b. A revision of the genus Gorsachius. Ibis (12) 2:585-592.

282. ——. 1926c. [Description of fifteen new forms from the oriental region. ]
Bull. Br. Orn. Club 47:52-58.

283. ——.1926d. [Description of five new forms.] Bull. Br. Orn. Club 47:68-
69.

284. ——.1928. Notes on some oriental birds. Tori 5:19-24.

285. ——. 1929. Contributions to the birds of the Philippines, No. 1. Orn.
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286. —. 1930. Contributions to the birds of the Philippines, No. 2. Orn.

Soc. Japan Suppl. 14:141-222.

287. ——. 193la. Notes sur les oiseaux des Philippines. I.— Révision des
colombes terrestres. L’Oiseau 1:23-29.

288. ——. 193lb. Notes sur les oiseaux des Philippines. II.— Activités
francaises dans l’histoire ornithologique des Philippines. L’Oiseau
1:208-210.

289. ——. 193lc. Notes sur les oiseaux des Philippines. III.— L’éperonnier de

Palawan (Polyplectron emphanum Temm.). L’Oiseau 1:265-268.

290. ——. 193ld. Notes sur les oiseaux des Philippines. IV.— Remarques sur
quelques grosses espéces de pigeons carpophages (Zonophaps).
LOiseau 1:397-401.

291. ——. 193le. Notes sur les oiseaux des Philippines. V.— Revue du genre
Neoleucotreron. L’Oiseau 1:469-470.

292. ——. 193lf. Description de nouvelles races d’oiseaux de Mindanao (Iles
Philippines). L’Oiseau 1:471-472.

293. ——. 193lg. Notes sur les oiseaux des Philippines. VI.—Le Mont Apo.
L Oiseau 1:589-593.

294. ——. 1932a. Notes sur les oiseaux des Philippines. VII.— L’Aigle des
Singes (Pithecophaga jefferyi). L’Oiseau 2:97-99.

295. ——. 1932b. Notes sur les oiseaux des Philippines. VIII.— Nouvelle
note sur les pigeons. L’Oiseau 2:225-226.

296. ——. 1932c. Notes sur les oiseaux des Philippines. [X.—Les Canards.
L’Oiseau 2:226.

297. ——.1932d. Notes sur les oiseaux des Philippines. X.—Les Perroquets.

L’Oiseau 2:417-418.

December 31, 1985 Kennedy, et al. 2

298. ——. 1933a. Notes sur les oiseaux des Philippines. XI.—Les Martins-
Pécheurs. L’Oiseau 3:69-7]1.

299. ——. 1933b. Notes sur les oiseaux des Philippines. XII.—Les Coucous.
LOiseau 3:201-203.

300. ——. 1933c. Notes sur les oiseaux des Philippines. L’Oiseau et RFO
Extrait. 33 pp. [A separately paginated reprint of Parts I-XII, 1931-
1933.]

301. ——. 1934a. Notes on the birds from the Philippine Islands and Borneo.
Tori 8:220-222.

302. ——. 1936a. On genus Borisia from the Philippine Islands. Mitt. Konig.

Naturwiss. Inst. Sofia, Bulgaria 9:53-56.

303. ——. 1936b. On the Guillemard Zoogeographical Line. Bull. Biogeogr.
Soc. Japan 6(18):183-192.

304. ——. 1936c. Oiseaux rares ou remarquables des iles Philippines.
L’Oiseau 6:185-197, 418-421.

305. ——. 1937a. Zoogéographie du Passage de Sibutu (Borneo). L’Oiseau
7T20E212:

306. ——. 1937b. Lhistoire du Pape 4 téte verte de Manille (Erythrura
viridifacies). L’Oiseau 7:385-388.

307. ——. 1937c. The discovery of Green-faced Parrot Finch. Tori 9:461-463.

308. —. 1938. La Grive Cendrée de Mindoro (Geokichla cinerea Bourns &
Worcester). L’Oiseau 8:1-2.

309. ——. 1939a. New races of a rail and a fruit pigeon from Micronesia and
Palawan. Bull. Br. Orn. Club 59:151-153.

310. ——.1939b. The Red Jungle Fowl from the Pacific Islands. Tori 10:596-
601.

311. ——. 1940. White-breasted Sea Eagle in the Philippine Islands. Tori
10:721-722.

312. ——. 194la. Description of a new species of bird from the Philippine

Islands. Bull. Biogeogr. Soc. Japan I1(1):1-4.

313. ——. 194lb. Description of a new genus and species of sunbird from the
Philippine Islands. Bull. Biogeogr. Soc. Japan I1(2):5-8.

314. ——. 194lc. Description of a new species of bird from the Philippine
Islands. Proc. Biol. Soc. Wash. 54:51-54.

28

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NEMOURIA No. 29

——. 194ld. Further contributions to the ornithology of the Philippine
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—.. 194le. New race of swift from the Philippine Islands. Proc. Biol.
Soc. Wash. 54:169-170.

——. 1942. Birds described by the Japanese authors. Tori 11:270-351.

——. 1944. Description of a new tailor-bird from the Philippine Islands.
Wort 1525-523:

—. 1952. A change of name for a green pigeon from the Philippine
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—, & J. DELACOUR. 1937. [Erythrura viridifacies described.] Bull.
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—. 1889. Notes on woodpeckers. —No. XVI. On some new species of
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HARRISSON, T. 1962. Hoopoe Upupa epops in the Philippines. Ibis
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HARTERT, E. 1890. Vorlaufiges iiber einige neue Végelarten. J. Orn.
38:154-156.

—. 189la. Die bisher bekannten Végel von Mindoro, nebst
Bemerkungen iiber einige V6gel von anderen Inseln der Philip-
pinen- Gruppe. J. Orn. 39:199-206, 292-302.

—. 189lb. Katalog der Végelsammlung im Museum der Sen-
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——. 1893. Bemerkungen tiber einige Capitoniden. J. Orn. 41:133-135.

—. 1895a. A new Prionochilus from the Philippines and note on an
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December 31, 1985 Kennedy, et ai. 29

331. ——. 1895b. List of a second collection of birds from the Natuna
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332. —.1895c. On a small collection of birds from Mindoro. Novit. Zool.
2:486-488.

333. —. 1897a. Aves: Podargidae, Caprimulgidae und Macropterygidae.
Das Tierreich 1:1-89.

334. ——. 1897b. On the birds collected by Mr. Everett in South Flores.
Novit. Zool. 4:513-528,

335. —. 1898. On the birds collected by Mr. Everett in South Flores. Part
II. Novit. Zool. 5:42-50.

336. —. 1902a. Aus den Wanderjahren eines Naturforschers. II. Abschnitt.
Sumatra, Malakka und Indien. Novit. Zool. 9:141-220.

337. —.1902b. Birds of the Kangean Islands. Novit. Zool. 9:419-442.

338. ——. 1903. [Eight new species described from Mindanao.] Bull. Br.
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339. ——. 1904a. [Description of new birds from Angola and Mindanao.]
Bull. Br. Orn. Club 14:72-74.

340. ——. 1904b. [Description of three new forms of Philippine birds.] Bull.
Br. Orn. Club 14:79-80.

341. ——. 1904c. [Dicaeum nigrilore described from Mindanao.] Bull. Br.

Orn. Club 15:8

342. ——. 1905a. [Sztta frontalis palawana described from Palawan.] Bull. Br.
Orn. Club 16:11-13.

343. ——. 1905b. Die Végel der paldarktischen Fauna. Systematische
Uebersicht der in Europa, Nord-Asien, und der Mittelmeerregion
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344, —.1905c. Family Eurylaemidae. In, P. Wytsman, Gen. Avium 1:1-8.

345. ——. 1906a. On the birds of the island of Babber. Novit. Zool. 13:288-
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346. ——. 1906b. Notes on birds from the Philippine Islands. Novit. Zool.
13:755-758.

347. —.1910. Birds of Hainan. Novit. Zool. 17:189-254.

30 NEMOURIA No. 29

348. ——. 19]6a. [lole philippensis saturatior described from Mindanao.] Bull.
Br. Orn. Club 36:58-59.

349, —.1916b. One of the rarest birds. Novit. Zool. 23:335-336..

350. ——. 1918. Further notes on pigeons. Novit. Zool. 25:434-436.

351. ——. 1919. Types of birds in the Tring Museum. B. Types in the
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352. ——. 1920. Types of birds in the Tring Museum. B. Types in the
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353. ——. 1922. Types of birds in the Tring Museum. B. Types in the
general collection. HI. Troglodytidae to Trochili. Novit. Zool.
29:365-412.

354. ——. 1924. Types of birds in the Tring Museum. B. Types in the
general collection. IV. Meropes to Halcyones. Novit. Zool. 31:112-
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355. ——. 1925a. Types of birds in the Tring Museum. B. Types in the
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356. —. 1925b. Review of the genus Cacomantis Mull. Novit. Zool.
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357. ——. 1925c. Types of birds in the Tring Museum. B. Types in the
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358. ——. 1927. Types of birds in the Tring Museum. B. Types in the
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359. ——. 1928. Types of birds in the Tring Museum. C. Additional and
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360. ——. 1929. On various forms of the genus Tyto. Novit. Zool. 35:93-104.

361. ——, & A. T. GOODSON. 1918. Notes on pigeons. Novit. Zool.
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362. ——, & E. STRESEMANN. 1925. Ueber die Indoaustralischen

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—. 198la. Saving the Philippine Eagle. Natl. Geogr. Mag. 159:846-
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—.198lb. The air’s noblest flier. Filipinas J. Sci. Cult. 2:33-48.

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38 NEMOURIA No. 29

459. ——. 1905d. II. Notes on three rare Luzon birds. Bull. Bur. Govt.
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461. ——.1906b. Siphia erithacus Sharpe, preoccupied. Condor 8:29.

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463. ——. 1906d. Notes on birds observed while traveling from Yokohama to
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464. ——. 1906e. Notes on birds collected in Mindoro and in small adjacent
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465. ——. 1906f. Notes on four birds from Luzon and on a species of
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466. ——.1906g. Notes on birds from Apo Island. Phil. J. Sci. 1:767.

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469. ——. 1906j. Notes on a collection of birds from Palawan Island. Phil. J.
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471. ——. 1907b. Notes on a collection of birds from the island of Basilan
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472. ——. 1907c. Descriptions of four new Philippine birds. Phil. J. Sci.
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473. ——.1907d. The occurrence of Blyth’s Wattled Lapwing and the Scaup
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474. ——. 1907e. Note on a bird unrecorded from Mindanao. Phil. J. Sci.
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475. ——. 1907f. Notes on specimens of the Monkey-eating Eagle
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December 31, 1985 Kennedy, et al. 39

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478. ——.1907i. The birds of Bohol. Phil. J. Sci. 2:315-335.

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480. ——. 1908a. Notes on a collection of birds from Siquijor, Philippine
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481. ——.1908b. Some necessary changes in the names of Philippine birds.
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482. ——. 1908c. Philippine ornithological literature, I. Phil. J. Sci. 3:285-
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486. ——. 1910b. Birds from Pauai and Mount Pulog, subprovince of
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487. —. 1910c. Additional notes on birds from northern Mindanao,
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488. ——. 1910d. Note on the migration of the Tic-wee Buzzard in the
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489. ——. 1910e. Philippine ornithological literature, III. Phil. J. Sci. 5:203-
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490. ——.19l0f. Birds from the coast of northern Luzon and from the islands

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493. ——. 1914. Description of a new species of Prionochilus from the
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40 NEMOURIA No. 29

494, ——. 1915. Birds in their economic relation to man. Manila, Bur. Sci.,
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496. —. 1918. New or noteworthy Philippine birds, II. Phil. J. Sci. 13:1-19.

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500. ——. 192lc. New or noteworthy Philippine birds, IV. Phil. J. Sci.
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501. ——. 1924. Birds of Ilocos Norte Province, Luzon. Phil. J. Sci. 25:111-121.

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503. ——. 1926b. Philippine doves and pigeons. Phil. Educ. Mag. 23:20-21,
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504. ——. 1926c. Philippine rails, gallinules, coots and similar birds. Phil.
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505. ——. 1926d. Philippine sea birds. Phil. Educ. Mag. 23:144-145, 180.

506. ——. 1926e. Philippine ploverlike shore birds. Phil. Educ. Mag. 23:210-
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507. ——. 1926f. Philippine snipelike shore birds. Phil. Educ. Mag. 23:276-
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508. ——. 1926g. Philippine herons and similar birds. Phil. Educ. Mag.
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509. —.1926h. Philippine ducks. Phil. Educ. Mag. 23:398-399, 456.

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511. ——.1927b. Philippine hawks and eagles. Phil. Educ. Mag. 23:540-541.

512. ——.1927c. Philippine owls. Phil. Educ. Mag. 23:610-6ll, 636.

December 31, 1985 Kennedy, et al. 4]

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514. ——. 1927e. Philippine kingfishers, hornbills, and bee-eaters. Phil.
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515. ——. 1927f. Philippine goatsuckers and swifts. Phil. Educ. Mag. 24:14-
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516. ——. 1927g. Philippine cuckoos, barbets, woodpeckers, and broadbills.
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517. ——. 1927h. Philippine passerine birds. Phil. Educ. Mag. 24:130-131,
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526. ——, & L. L. GARDNER. 1930. Philippine bird traps. Condor 32:89-
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42 NEMOURIA No. 29

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531. ——. 1933. Notes on the bird predators of Pericyma cruegeri (Butler).
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533. ——.1934b. Food and feeding habits of the barred ground dove. Phil. J.
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534. ——. 1935a. New birds from northern Luzon; Philippine Islands.
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535. ——. 1935b. Life history and economic importance of Cabanis’s
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536. ——. 1936a. Review of Philippine pigeons, I: The genus Phapitreron.
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537. ——. 1936b. New Philippine fruit pigeons. Phil. J. Sci. 59:307-309.

538. ——. 1936c. A review of Philippine pigeons, II: Subfamily
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539. ——. 1936d. A review of Philippine pigeons, III: Subfamily

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540. ——. 1936e. A review of Philippine pigeons, IV: Subfamily
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541. -——. 1937a. Beneficial swiftlet and edible birds’ nest industry in
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542. ——. 1937b. A review of Philippine pigeons, V: Subfamilies
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543. ——. 1937c. The avifauna of Catanduanes. Phil. J. Sci. 63:185-189.

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MEISE, W. 1931. Zur Systematik der Gattung Gerygone. Novit. Zool.
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NEMOURIA No. 29

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December 31, 1985 Kennedy, et al. 49

634. ——. 1909. A revision of the kingfisher genus Ramphalcyon
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635. ——. 1911. A monograph of the flycatcher genera Hypothymis and
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636. ——. 1912. A revision of the forms of the Edible-nest Swiftlet

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637. ——. 19l5a. A review of the subspecies of the Ruddy Kingfisher,
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638. —. 1915b. A synopsis of the races of the Crested Tern Thalasseus
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639. ——. 1917. Diagnosis of a new pycnonotine family of Passeriformes.
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640. ——. 1918. Diagnosis of a new genus of Timaliidae. J. Wash. Acad.
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641. ——. 1919a. A revision of the subspecies of the White-throated

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642. —. 1919b. Diagnosis of a new genus of Bucerotidae. J. Wash. Acad.
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643. ——. 1923. Descriptions of new East Indian Nectariniidae. J. Wash.

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644. ——. 1924a. Descriptions of new Treronidae and other non- passerine
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645. ——. 1924b. Note on the races of Ramphalcyon capensis. Proc. Biol.
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646. ——. 1926. Descriptions of nineteen new East Indian passerine birds.
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647. ——. 1932. The birds of the Natuna Islands. U. S. Nat. Mus. Bull.
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648. OGILVIE GRANT, W. R. 1894a. On the birds of the Philippine
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50

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NEMOURIA No. 29

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—. 1894c. [Descriptions of new birds from northern Luzon.] Bull.
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——. 1894d. [Descriptions of two new species from northern Luzon.]
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—. 1895a. [Descriptions of two new species from the Philippines.]
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—. 1895b. [Zosterops luzonica described from south Luzon.] Bull.
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—. 1895c. On the birds of the Philippine Islands.—Part III. The
mountains of the province of Isabella, in the extreme north-east of
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—. 1895d. [New species of birds discovered by Whitehead in the
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—. 1895e. On the birds of the Philippine Islands.—Part IV. The
province of Albay, south-east Luzon, and the adjacent island of
Catanduanes. Ibis (7) 1:249-267.

——. 1895f. On the birds of the Philippine Islands.—Part V. The
highlands of the province of Lepanto, north Luzon. Ibis (7) 1:433-
472.

—. 1895g. [Descriptions of four new species from the Philippines.]
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—. 1896a. On the birds of the Philippine Islands.—Part VI. The
vicinity of Cape Engafio, N.E. Luzon, Manila Bay, and Fuga
Island, Babuyan Group. Ibis (7) 2:101-128.

——. 1896b. On the birds of the Philippine Islands.—Part VII. The
highlands of Mindoro. Ibis (7) 2:457-477.

—. 1896c. On the birds of the Philippine Islands.—Part VIII. The
highlands of Negros. Ibis (7) 2:525-565.

—. 1897a. On the birds of the Philippine Islands.—Part IX. The
islands of Samar and Leite. Ibis (7) 3:209-250.

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December 31, 1985 Kennedy, et al. 51

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—. 1897c. A hand-book to the game-birds. London, W. H. Allen.
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—. 1902. A review of the species of shrikes of the genus Lamnzus.
Novit. Zool. 9:449-486.

—. 1905. [Six new species described from south-east Mindanao. ]
Bull. Br. Orn. Club 16:16-19.

—. 1906a. [Two new species described from south-east Mindanao. ]
Bull. Br. Orn. Club 16:99-100.

—. 1906b. On the birds collected by Mr. Walter Goodfellow on the
volcano of Apo and in its vicinity, in south-east Mindanao,
Philippine Islands. Ibis (8) 6:465-505.

—, & J. WHITEHEAD. 1898. On the nests and eggs of some rare
Philippine birds. Ibis (7) 4:231-247.

OLIVIER, G. 1947. Note sur Lanius validirostris Grant 1894 des Iles
Philippines. L’Oiseau 17:182-185.

ORENSTEIN, R. I. 1979. The systematic position of the water
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—, & R. S. KENNEDY. 1985. Creeper, Philippine. Pp. 118 i A
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OUSTALET, E. 1876. [Aethopyga f. flagrans described.] J. Institut.,
p. 108.

— . 1880a. [Ptilinopus marchei described.] Le Naturaliste 1:325.

—. 1880b. Description de deux oiseaux nouveaux des Iles Souloo.
Bull. Hebdo. Assoc. Sci. France 2(2):205-206. [Descriptions of
Ninox philippensis rey1 and Anthracoceros montani]|

—. 1885. Description de deux espéces nouvelles faisant partie de la
collection ornithologique du Muséum d’Histoire Naturelle de
Paris. Le Naturaliste 3:108. [Anthracoceros marchei described. ]

— . 1890. Description d’un nouveau martin pécheur des iles
Philippines. Le Naturaliste 12:62-63. [Halcyon winchelhi alfredi
described. |]

PAGE, W. T. 1913a. White-eyes (Zosterops). Bird Notes 4:1-7.

52 NEMOURIA No. 29
679. ——. 1913b. Fairy bluebirds (/rena). Bird Notes 4:95-101.

680. PARKER, S. A. 1981. Prolegomenon to further studies in the
Chrysococcyx ‘malayanus’ group (Aves. Cuculidae). Zool. Verh.
(Leiden) 187:1-56.

681. PARKES, K. C. 1949. Rallus philippensis on Mindoro, Philippine
Islands. Auk 66:200-201.

682. ——. 1952. The races of the Bald Starling of the Philippines. Condor
54:55-57.

683. —. 1957a. Feathers, Filipinos, and fevers. Carnegie Mag. 31:51-54.

684. ——. 1957b. The plumage sequence of the Philippine Coucal Cen-

tropus viridis. Ibis 99:518-520.

685. ——. 1957c. Taxonomic notes on the Lesser Coucal, Centropus
bengalensis. Bull. Br. Orn. Club 77:115-116.

686. ——. 1958a. Nomenclatorial notes on Philippine Pygmy Wood-
peckers. Bull. Br. Orn. Club 78:6-7.
687. ——. 1958b. A new race of the Blue-headed Fantail (Rhzpidura

cyaniceps) from northern Luzon, Philippine Islands. Am. Mus.
Novit. 1891:1-5.

688. ——. 1958c. A revision of the Philippine Elegant Titmouse (Parus
elegans). Proc. Biol. Soc. Wash. 71:95-106.

689. ——. 1958d. Specific relationships in the genus Elanus. Condor
60:139-140.

690. ——. 1958e. Taxonomy and nomenclature of three species of Lon-
chura (Aves: Estrildinae). Proc. U.S. Nat. Mus. 108(3402):279-
293.

691. ——. 1958f. Extra-limital records of the Australian Whiskered Tern.
Emu 58:288.

692. ——. 1959. Subspecific identity of introduced Tree Sparrows Passer
montanus in the Philippine Islands. Ibis 101:243-244.

693. ——. 1960a. Geographic variation in the Lesser Tree-Swift. Condor
62:3-6.

694. ——. 1960b. Notes on Philippine races of Dryocopus javensis. Bull.

Br. Orn. Club 80:59-6l.

December 31, 1985 Kennedy, et al. 53

695. ——. 1960c. Notes on some Philippine tailor-birds. Bull. Br. Orn.
Club 80:76-78.

696. —. 1960d. A drongo new to the Philippine list. Condor 62:296-297.

697. ——. 1960e. New subspecies of Philippine birds. Proc. Biol. Soc.
Wash. 73:57-62.

698. ——. 1960f. Notes on some non-passerine birds from the Philippines.
Ann. Carnegie Mus. 35(15):331-340.

699. ——. 1960g. The races of the Citrine Canary Flycatcher, Culicicapa
helianthea. Proc. Biol. Soc. Wash. 73:215-220.

700. ——. 196la. A substitute name for a Philippine tailor-bird. Bull. Br.
Orn. Club 81:33.

701. ——. 196lb. The Crested Lizard Hawk (Aviceda jerdoni) in the
Philippines. Postilla 51:1-10.

702. ——. 1962a. The Red Junglefowl of the Philippines—native or in-
troduced? Auk 79:479-48].

703. ——. 1962b. New subspecies of birds from Luzon, Philippines.
Postilla 67:1-8.

704. ——. 1963a. The races of the Flaming Sunbird (Aethopyga flagrans).
Bull. Br. Orn. Club 83:7-8.

705. ——. 1963b. A substitute name for the Luzon race of Copsychus
saularis. Bull. Br. Orn. Club 83:50.

706. ——. 1963c. Additional notes on the Philippine Elegant Titmouse,
Parus elegans. Bull. Br. Orn. Club 83:148-150.

707. ——. 1963d. A new subspecies of tree-babbler from the Philippines.
Auk 80:543-544.

708. ——. 1965a. A small collection of birds from the island of Buad,

Philippines. Ann. Carnegie Mus. 38(2):49-67.
709. ——. 1965b. Character displacement in some Philippine cuckoos.
Living Bird 4:89-98.

710. ——. 1965c. The races of the Pompadour Green Pigeon, Treron
pompadora, in the Philippine Islands. Bull. Br. Orn. Club 85:137-
139.

54 NEMOURIA No. 29

711. ——. 1966a. Geographic variation in Winchell’s Kingfisher Halcyon
winchelh, of the Philippines. Bull. Br. Orn. Club 86:82-86.

712. ——. 1966b. A new subspecies of the Yellow-bellied Whistler
Pachycephala philippinensis. Bull. Br. Orn. Club 86:170-171.

713. ——. 1967. A new subspecies of the Wattled Bulbul Pycnonotus
urostictus, of the Philippines. Bull. Br. Orn. Club 87:23-25.

714. ——. 1968. An undescribed subspecies of button-quail from the
Philippines. Bull. Br. Orn. Club 88:24-25.

715. ——. 1969. Subspecific status of the Small Skylark Alauda gulgula in

the Philippines, with notes on age characters and moult. Bull. Br.
Orn. Club 89:117-119.

716. ——. 1970a. A revision of the Philippine Trogon (Harpactes ardens).
Nat. Hist. Bull. Siam Soc. 23:345-352.
717. ——. 1970b. The Philippine races of the Rufous-capped Grass

Warbler, Megalurus timoriensis. Bull. Br. Orn. Club 90:II1-115.

718. ——. 197la. Taxonomic and distributional notes on Philippine Birds.
Nemouria 4:1-67.

719. ——. 197lb. Two new parrots from the Philippines. Bull. Br. Orn.
Club 91:96-98.

720. ——. 197lc. A new subspecies of pitta from the Philippines. Bull. Br.
Orn. Club 91:98-99.

721. ——. 1973. Annotated list of the birds of Leyte Island, Philippines.

Nemozuria 11:1-73.

722. ——. 1974. Geographic variation in the Flame Minivet (Pericrocotus
flammeus) on the island of Mindanao, Philippines (Aves: Cam-
pephagidae). Ann. Carnegie Mus. 45(3):35-41.

723. ——. 1981. A substitute name for a Philippine minivet (Pericrocotus).
Bull. Br. Orn. Club 101:370.
724. ——, & D. AMADON. 1948. The winter range of the Kennicott

Willow Warbler. Condor 50:86-87.

725. ——, & ——. 1959. A new species of rail from the Philippine Islands.
Wilson Bull. 71:303-306.

December 31, 1985 Kennedy, et al. 55

726. PARROT, C. 1907. Beitrége zur Ornithologie Sumatras und der Insel
Banka mit besonderer Zugrundelegung der von Dr. Hagen auf
Banka gesammelten Vogel. Abh. Bayern Akad. Wiss. Mtinchen II
24(1):149-286.

727. PEALE, T. R. 1848. Mammalia and ornithology. Jn, United States
Exploring Expedition during the years 1838, 1839, 1840, 1841,
1842, under the command of Charles Wilkes, U.S.N.
Philadelphia, Sherman. Vol. 8, 338 pp. [see Cassin 1858]
[Descriptions of Centropus nigrifrons (syn. for Centropus melanops),
p. 137; Harpactes radiosternus (syn. for Harpactes ardens), p. 166;
Ducula casta (syn. for Ducula b. bicolor); Cuculus fucatus (syn. for
Cuculus s. saturatus); Buceros obscurus (syn. for Buceros hydrocorax
mindanensis). |

728. PETERS, J. L. 1930. Notes on some night herons. Proc. Boston Soc.
Nat. Hist. 39:263-277.

729. ——. 1937. A new genus for Pseudoptynx solomonensis J. Wash. Acad.
Sci. 27:81-83.

730. ——. 1938. Generic limits of some fruit pigeons. Proc. VIII Int. Orn.
Congr. (1934):371-391.

73la. ——. 1939. Collections from the Philippine Islands: Birds. Bull.

Mus. Comp. Zool. 86:74-128.

731b. ——. 1943. First supplement to the list of types of birds now in the
Museum of Comparative Zoédlogy. Bull. Mus. Comp. Zodl. 92:53-
118. [Hemiprocne comata barbarae, Hypothymis azurea compilator.|

732. POTTER, N. S., III. 1948. Notes on the Yellow-breasted Sunbird.
Wilson Bull. 60:159-163.

733. ——. 1953. The birds of Calicoan, Philippine Islands. Wilson Bull.
65:252-270.

734. PRESCOTT, K. W. 1950. Malay Banded Crake off the island of
Mindanao in the Philippines. Wilson Bull. 62:134.

735. ——. 1973. First report of Pitta e. erythrogaster from Leyte. Bull. Br.
Orn. Club 93:32-33.

736. PUCHERAN, J. 1853. Voyage au Péle Sud et dans l’Oceanie sur les
corvettes L’Astrolabe et La Zélée...1837...1840. Zoologie 3
(Mamm. et Ois.):47-158. [Gallicolumba c. criniger described. ]

56 NEMOURIA No. 29

737. ——. 1854. Etudes sur les types peu connus du Musée de Paris
(Onzieme article.—Passereaux conirostres). Rev. et Mag. Zool. (2)
6:62-74.

738. —. 1855. Mémoire sur les types peu connus de Passereux den-

tirostres de la collection du Musée de Paris. Arch. Mus. d’Hist.
Nat. 7:321-380. [Description of Philedon gularis (syn. for Hyp-
stpetes p. philippinus), p. 344.]

739. PURCHASE, D. 1973. Eighteenth annual report of the Australian
bird-banding scheme, June 1971 to June 1972. An. Rep. Aust.
Bird Band. Scheme 18:1-93.

740. ——. 1974. First overseas recoveries of Wedge-tailed Shearwaters
banded in Australia. Aust. Bird Bander 12:34-35.

741. QUINONES, N. C. 1981. Diurnal rhythm and periodic changes in
the type, quantity and caloric values of food ingested by adult
Tree Sparrow (Passer montanus saturatus Stejneger). Sylvatrop
Phil. For. Res. J. 6:1-18.

742. RABOR, D. S. 1936a. Life histories of some common birds in the
vicinity of Novaliches, Rizal Province, Luzon. I. The Large-
nosed Shrike, Lanius schach nasutus Scopoli. Phil. J. Sci. 59:337-

355:

743. —. 1936b. Life histories of some common birds in the vicinity of
Novaliches, Rizal Province, Luzon, II. Phil. J. Sci. 60:143-156.

744. ——. 1938a. Birds from Leyte. Phil. J. Sci. 66:15-34.

745. ——. 1938b. The avifauna of the Gigante Islands. Phil. J. Sci.
66:267-274.

746. ——. 1952. Distributional notes on some Philippine birds. Auk
69:253-257.

747. ——. 1954a. Notes on the nesting of some Philippine swifts on
Negros and Mindanao. Silliman J. 1:45-58.

748. ——. 1954b. New bird records on Negros Island, Philippines.
Silliman J. 1:77-82.

749. ——. 1955a. Species limits in the Plain-throated Bulbul, Microscels,

of the Philippines. Silliman J. 2:103-104.

December 31, 1985 Kennedy, et al. a

750. —. 1955b. Notes on mammals and birds of the central northern
Luzon highlands, Philippines. Part II: Notes on birds. Silliman J.
2:253-3ll.

751. ——. 1958. Wildlife in the humid tropics picture. Proc. 9th Pac. Sci.
Cong. 20:95-104.

752. ——. 1959. The impact of deforestation on birds of Cebu, Philip-
pines, with new records for that island. Auk 76:37-43.

753. ——. 196l. The status of conservation in the Philippines. Proc. 10th
Pac. Sci. Cong. 1961:236-240.

754. ——. 1963. Zoogeography of Negros Island and the Visayan provinces
of central Philippines. Phil. Geogr. J. 7:179-192.

755. ——. 1964a. New bird records of various islands in the Philippines.
Silliman J. 11:202-216.

756. ——. 1964b. Conservation in the Philippines. 10th Pac. Sci Cong.,
Conserv. News S. E. Asia No. 4:10-12.

757. ——. 1966a. A report on the zoological expeditions in the Philippines
for the period 1961-1966. Silliman J. 13:605-616.

758. ——. 1966b. Conservation in the Philippines. Silliman J. 13:594-604.

759. —. 1968a. Highlights in the discussion on Technical Session IV.—

Threatened species. Int. Union Conserv. Nat. Nat. Resour. Publ.
New Ser. No. 10:260-261.

760. ——. 1968b. The present status of the Monkey-eating Eagle,
Pithecophaga jeffery: Ogilvie-Grant, of the Philippines. Int. Union.
Conserv. Nat. Nat. Resour. Publ. New Ser. No. 10:312-314.

761. ——.1969. The altitudinal distribution of the birds and mammals of the
Philippines. Proc. Nat. Sci. Tech. Week, Nat. Sci. Dev. Board,
Manila, pp. 13-78.

762. ——. 197la. Wildlife conservation in the Philippines. Phil. Geogr. J.
15:46-63.
763. ——. 197lb. The present status of conservation of the Monkey-eating

Eagle of the Philippines. Phil. Geogr. J. 15:90-103

764. ——. 1975. The present status of conservation of the ecosystem of Lake
Lanao and its surrounding region and the degree of exploitation of
its natural resources. Nat. Res. Council Phil. Bull. 57:70-87.

58 NEMOURIA No. 29

765. ——. 1976. Wildlife of Mindanao. Sylvatrop Phil. For. Res. J. 1:249-
254.

766. ——. 1977. No more forests, no more birds. Conserv. Circ. 13(2):4-5.
College, Laguna, Univ. Phil. Los Bafios, College For.

767. ——. 1978a. Philippine wildlife, their conservation and management.
Conserv. Circ. 14(4):5-7. College, Laguna, Univ. Phil., Los Bafios,
College For.

768. ——. 1978b. Wildlife management a factor in hilly land development.

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769. ——. 1978c. Wildlife resources of the Philippines and their con-
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770. ——. 1979. Present status of conservation of Philippine wildlife. Phil.
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771. ——.1980a. For wildlife to come back. Habitat Phil. 1(1):39-42.

772. ——. 1980b. Pests and potential pests among the vertebrates of the

Philippines other than rats. Pp. 259-276 in Development in pest
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773. ——. 1980c. Philippine wildlife, their preservation, conservation and
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774. ——. 1980d. Present status of Philippine ecosystems and the effects of
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775. —. 1980e. Wildlife components of the Philippine ecosystem and their
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776. ——. 198la. The role of wildlife in forest regeneration in southeast Asia.
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777. ——, A. C. ALCALA, & R. B. GONZALES. 1958. A brief list of land
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778. ——, —,& —. 1970. A list of the land vertebrates of Negros Island,

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779. ——, D. GANAPIN, & R. LUCERO. 1978. Declaration on wildlife
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780. ——,&N. C. QUINONES. 1977. Time now to stockfarm and manage

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781. ——,&A.L. RAND. 1958. Jungle and domestic fowl, Gallus gallus, in
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782. RAND, A. L. 1948. Five new birds from the Philippines. Fieldiana
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783. ——. 1950a. A new race of the Philippine creeper, Rhabdornis inornatus
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784. ——.1950b. Three rare Philippine birds. Nat. Hist. Misc. 60:1-5.

785. ——. 1950c. A new race of owl, Otus bakkamoena, from Negros,
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786. ——. 195la. Birds of Negros Island. Fieldiana Zool. 31(48):571-596.

787. ——. 195lb. Review of the subspecies of the sunbird Nectarinia
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790. ——.1954c. Immature females with adult male plumage characters. Auk
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791. ——.1955. Philippine bird names of Blasius. Auk 72:210-212.

792. ——. 1959. Late records of the Cebu Golden-backed Hanging Parrakeet.

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793. —. 1960. A new species of babbling thrush from the Philippines.
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794. ——. 1961. The tongue and nest of certain flowerpeckers (Aves:
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60

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NEMOURIA No. 29

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—., & ——. 1959a. Notes on some Philippine bulbuls. Auk 76:102-104.

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—., & —. 1960. Birds of the Philippine Islands: Siquijor, Mount
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—, & —. 1967. New birds from Luzon, Philippine Islands.

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—, & —. 1969. New birds from Camiguin South, Philippines.
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December 31, 1985 Kennedy, et al. 61

812. RILEY, J. H. 1920. Four new birds from the Philippines and Greater
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813. ——. 192]. Five new genera of birds. Proc. Biol. Soc. Wash. 34:51-54.

814. -—. 1927. Note on the genus Jrena, Horsfield. Proc. Biol. Soc. Wash.
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815. ——. 1935. Three new forms of birds from the Philippine Islands and

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816. ——. 1936. A new flower-pecker from Palawan Island, Philippines.
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817. RIPLEY, S. D. 1941. Notes on the genus Coracina. Auk 58:381-395.

818. ——. 1942a. A revision of the kingfishers, Ceyx erithacus and rufidorsus.
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819. ——. 1942b. The species Eurystomus orientalis. Proc. Biol. Soc. Wash.
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820. ——. 1949. A new race of shrike from the Philippines. Bull. Br. Orn.
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821. ——. 1950. Comments on specimens in the Hachisuka collection from
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822. -——. 1951. Remarks on the Philippine Mallard. Wilson Bull. 63:189-191.

823. ——-.1952a. Waterfowl notes. Avic. Mag. 58:21-22.

824. ——. 1952b. A note on the species Muscicapa westermanni. Proc. Biol.

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825. -——.1952c. Additional comments on Philippine birds and a new record
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826. ——.1952d. The thrushes. Postilla 13:1-48.

827. ——. 1954. Names proposed in “the thrushes.” Postilla 13(suppl.):1.

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829. ——,&D.S. RABOR. 1955. A new fruit pigeon from the Philippines.
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830. ——, & ——. 1956. Birds from Canlaon Volcano in the highlands of

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62

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NEMOURIA No. 29

—, & —. 1958a. Columba vitiensis anthracina (Hachisuka), a
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—, & —. 1962. New birds from Palawan and Culion Islands,
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—, & — . 1966. Dicaeum proprium, new species (Aves; family
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—, & ——. 1968. Two new subspecies of birds from the Philippines
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RITTER VON FRAUENFELD, G. 1861. Der Aufenthalt auf Manila
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—. 1904. [A new species of parrot from the Philippines.] Bull. Br.
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—. 1929. Bemerkungen tiber die Gruppe Acrocephalus arundinaceus L.
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December 31, 1985 Kennedy, et al. 63

846. ——. 1937. [Notes on the Philippine paradise flycatchers.] Bull. Br.
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847. ——. 1952a. Systematic notes on some Philippine birds. Vidensk.
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848. ——. 1952b. Mindanao. Oplevelser og haendelser under den Danske

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849. ——. 1953a. Et nyt navn for Rhabdornis longirostris Salomonsen. Dan.
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850. —. 1953b. Miscellaneous notes on Philippine birds. Vidensk. Medd.
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851. ——. 1960a. Notes on flowerpeckers (Aves, Dicaeidae). 1. The genera
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852. ——.1960b. Notes on flowerpeckers (Aves, Dicaeidae). 2. The primitive

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853. ——. 1960c. Notes on flowerpeckers (Aves, Dicaeidae). 3. The species
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854. ——. 196la. Noona Dan Papers. |. A new tit-babbler (Stachyris
hypogrammica, sp. nov.) from Palawan, Philippine Islands. Dan.
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855. ——. 196lb. Notes on flowerpeckers (Aves, Dicaeidae). 4. Dicaeum
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856. —. 1962. Noona Dan Papers. 2. The mountain bird fauna of Palawan,
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857. ——. 1973. [Review of] Philippine Birds [by J. E. duPont]. Auk 90:922-
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858. ——.1977. Noona Dan Papers. 132. The Thicket Flycatcher (Muscicapa

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859. ——. 1983. Noona Dan Papers. 141. Revision of the Melanesian
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64 NEMOURIA No. 29

860. SALVADORI, T. 1865. Descrizione di altre nuove specie di uccelli
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861. ——. 1870. Nuove specie di uccelli. Atti. R. Accad. Sci. Torino 5:507-
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862. ——. 1874. Note ornitologiche. I. Atti. R. Accad. Sci. Torino 9:630-635.

863. ——. 1876. Intorno alla supposta femmina del Dicaeum retrocinctum,
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864. ——. 1878. Intorno alla Trerolaema leclancheri, Bp. Atti. R. Accad. Sci.
Torino 13:425-428.

865. ——.1879a. On Melaniparus semilarvatus. Ibis (4) 3:300-303.

866. ——. 1879b. Di alcune specie del genere Porphyrio. Atti. R. Accad. Sci.

Torino 14:1165-1170.

867. ——. 1880. Osservazioni intorno ad alcune specie del Genere Collocalha,
G.R. Gr. Atti. R. Accad. Sci. Torino 15:343-350.

868. ——. 1882. Descrizione di una nuova specie del genere Collocalia ed
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869. ——. 1886. On some Papuan, Moluccan, and Sulu birds. Ibis (5) 4:151-
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870. ——. 189la. Catalogue of the Psittaci, or parrots, in the collection of the

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871. ——. 189lb. On a rare species of lorikeet in the Rothschild Collection.
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872. ——. 1893. Catalogue of the Columbae, or pigeons, in the collection of

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873. SCHEFFER, T. H., & C. COTTAM. 1935. The Crested Myna, or
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SCHUCHMANN, K.-L., & H. E. WOLTERS. 1982. A new sub-
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——. 1910b. [Further notes on Pithecophaga jefferyi.| Ibis (9) 4:758-759.
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NEMOURIA No. 29

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NEMOURIA No. 29

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927. SOUANCE, C. DE. 1856. Catalogue des Perroquets de la collection du
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935. STEINBACHER, J. 1949. Die Auffindung eines dritten Stiickes des
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938. ——.1947a. Land of the Calao. Audubon Mag. 49:12-17.
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— . 1922b. Zur Ornithologie der Philippinen. Orn. Monatsber. 30:88.

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—. 1923. Zur Nomenklatur indo-australischer Végel. Orn. Monat-
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—.. 1924c. Die Gattung Acrocephalus im indo-australischen Archipel.
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972. TAN, B. C., W. S. M. GRUEZO, & L. S. CUY. 1982. Nesting plant
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973. TEMME, M. 1973a. A White Ibis (Threskiornis aethiopica
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974. ——. 1973b. Notizen tiber die Graseule (Tyto capensis) (= Tyto
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976. ——. 1973d. Einige Beobachtungen am Grau-Mennigvogel (Pericrocotus
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977. ——. 1973e. Zum Vorkommen des Stelzenléufers Himantopus h.
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978. ——. 1974a. Finches—another vertebrate pest in Philippine ricefields.
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979. ——. 1974b. New records of Philippine birds on the island of Mindoro.
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980. ——. 1976a. Beitrag zur Kenntnis der Vorkommens ostpaldarktischer
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981. ——. 1976b. First record of a Dusky Warbler (Phylloscopus fuscatus)

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982. ——. 1976c. Beitrag zur Kenntnis der Philippinenente (Anas luzonica
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1075. WIGLESWORTH, L. W. 1891. On the polynesian members of the
genus Prilopus. Ibis (6) 3:566-584.

1076. WILKES, C. 1844. Narrative of the United States Exploring Expedition
during the years 1838, 1839, 1840, 1841, 1842. Vol. 5, 591 pp.
[Philippines, pp. 293-390. ]

1077. WISCHUSEN, E. W., R. S. KENNEDY, & S. E. GAST. 1984. First
record of the Red Phalarope from the Philippines. Auk 101:185.

1078. WOLF, C. E. 1974. [Review of Philippine Birds by John E. duPont.]
Phil. Studies 22:380.

1079. WOLFE, L. R. 1938. Birds of central Luzon. Auk 55:198-224.

80

1080.

1081.

1082.

1083.

1084.

1085.

1086.
1087.

1088.

1089.

1090.

1091.

1092.

NEMOURIA No. 29

WOLFF, T. 1966. The “Noona Dan” Expedition 1961-1962. General
report and lists of stations. Vidensk. Medd. Dan. Naturhist. Foren.
129:287-336.

WOLTERS, H. E. 1967. Uber einige asiatische Carduelinae. Bonn.
zool. Beitr. 18:169-172.

WORCESTER, D. C. 1898. Contributions to Philippine ornithology.
Part II.—Notes on the distribution of Philippine birds. Proc. U. S.
Nat. Mus. 20(1134):567-625.

—. 1907a. On a nesting specimen of Caprimulgus griseatus Walden.
Phil. J. Sci. 2:271-273.

—. 1907b. On a nesting place of Su/a sula (Linnaeus) and Sterna
anaestheta Scopoli. Phil. J. Sci. 2:277.

——. 19lla. Newly discovered breeding places of Philippine sea birds.
Phil. J. Sci. 6:167-177.

——. 19llb. Hybridism among boobies. Phil. J. Sci. 6:179.

——. 1919. A nesting place of Micropus subfurcatus in Mindoro. Phil. J.
Sci. 15:533-534.

WYLIE, S. R. 1974. Notes on egg-laying in the Monkey-eating Eagle.
Auk 91:191.

ZIMMER, J. T. 1918a. A few rare birds from Luzon, Mindanao and
Mindoro. Phil. J. Sci. 13:223-232.

——. 1918b. Some notes on the birds of southern Palawan and adjacent
islands. Phil. J. Sci. 13:327-357.

—,&E. MAYR. 1943. New species of birds described from 1938 to
1941. Auk 60:249-262.

ZISWILER, V., H. R. GUTTINGER, & H. BREGULLA. 1972.
Monographie der Gattung Erythrura Swainson, 1837 (Aves,
Passeres, Estrildidae). Bonn. zool. Monogr. 2:1-158.

December 31, 1985 Kennedy, et al. 81

SECTION III

LIST OF RECENT GENERAL WORKS

1093a. ALDEN, P., & J. GOODERS. 1981. Finding birds around the world.

Boston, Houghton Mifflin. 683 pp. [Philippines, pp. 483-490. ]

1093b. BEEHLER, B. M., T. K. PRATT, & D. A. ZIMMERMAN. 1985.

1094.

1095.

1096.

1097.

1098.

1099.

1100.

OL

1102.

1103.

1104.

Birds of New Guinea. Princeton, New Jersey, Princeton Univ.
Press. ca 370 pp.

BROWN, L., & D. AMADON. 1968. Eagles, hawks, and falcons of the
world. New York, McGraw-Hill. Vol. 1, pp. 1-434; Vol. 2, pp. 435-
945.

BURTON, J. A. [Ed.] 1973. Owls of the world. New York, E. P.
Dutton. 216 pp.

CAMPBELL, B., & E. LACK. 1985. A dictionary of birds. Calton,
England, T. & A. D. Poyser (for the Br. Orn. Union). 670 pp.

CHANG, J. WAN-FU. 1980. A field guide to the birds of Taiwan.
Taichung. 324 pp.

CLARK, R. J., D. G. SMITH, & L. H. KELSO. 1978. Working
bibliography of owls of the world. Washington, D. C., Nat. Wildl.
Fed. 319 pp.

CLEMENTS, J. F. 1978. Birds of the world: A check list. 2nd Ed. New
York, Two Continents Publ. Group, Ltd. 532 pp.

DEIGNAN, H. G. 1963. Checklist of the birds of Thailand. U.S. Nat.
Mus. Bull. 226:1-263.

DELACOUR, J. 1947. Birds of Malaysia. New York, Macmillan. 382
Pp.

——. 1977. The pheasants of the world. 2nd Ed. Surrey, England, Spur
Publ. 395 pp.

DUPONT, J. E. 1976. South Pacific birds. Greenville, Delaware,
Delaware Mus. Nat. Hist. 218 pp.

ETCHECOPAR, R. D., & F. HUE. 1978. Les oiseaux de Chine, de
Mongolie et de Corée: non passereaux. Papeete, Tahiti, Les Editions
du Pacifique. 585 pp.

82

1105.

1106.

1107.

1108.

1109.

1110.

Hea

PZ:

3:

1114.

LETS:

1116.

LI:

1118.

1119.

1120:

NEMOURIA No. 29

—, & ——. 1983. Les oiseaux de Chine, de Mongolie et de Corée
passereaux. Paris, Boubée. 704 pp.

FORSHAW, J. M., & W. T. COOPER. 1973. Parrots of the world.
Garden City, New York, Doubleday. 584 pp.

—, & —. 1983. Kingfishers and related birds. Melbourne, Lan-
sdowne. Vol. 1, pt. 1, 266 pp. [Alcedinidae: Ceryle to Cittura.]

FRITH, C. B. 1979. Ornithological literature of the Papuan Subregion,
1915 to 1976: An annotated bibliography. Bull. Am. Mus. Nat.
Hist. 164(3):377-465.

FRY, C. H. 1984. The bee-eaters. Calton, Poyser. 320 pp.

GOODWIN, D. 1976. Crows of the world. Ithaca, New York, Cornell
Univ. Press. 354 pp.

—. 1982. Estrildid finches of the world. Ithaca, New York, Cornell
Univ. Press. 328 pp.

——. 1983. Pigeons and doves of the world. 3rd Ed. Ithaca, New York,
Cornell Univ. Press. 363 pp.

GORE, M. E. J., & P. WON. 1971. The birds of Korea. Seoul, Royal
As. Soc. & Taewon Publ. Co. 450 pp.

GREENWAY, J. C., JR. 1958. Extinct and vanishing birds of the world.
Am. Comm. Int. Wild Life Prot. N. Y. Spec. Publ. 13:1-518. [2nd
(revised) ed. 1967, New York, Dover. 520 pp.]

HANCOCK, J., & H. F. 1. ELLIOTT. 1984. The herons handbook.
[2nd ed. of 1978 book.] New York, Harper & Row. 288 pp.

HARRISON, P. 1983. Seabirds, an identification guide. Boston,
Houghton Mifflin. 448 pp. [also London, Croom Helm.]

HOWARD, R., & A. MOORE. 1980. A complete checklist of the birds
of the world. Oxford, Oxford Univ. Press. 701 pp. [2nd (revised) ed.
1984, London, Macmillan. 732 pp.]

JOHNSGARD, P. A. 1978. Ducks, geese, and swans of the world.
Lincoln, Nebraska, Univ. Nebraska Press. 404 pp.

—. 1981. The plovers, sandpipers, and snipes of the world. Lincoln,
Nebraska & London, Univ. Nebraska Press. 493 pp.

KING, B. F., & E. C. DICKINSON. 1975. A field guide to the birds of
South-East Asia. London, Collins. 480 pp.

December 31, 1985 Kennedy, et al. 83

1121. MASSEY, J. A.,S. MATSUI, T. SUZUKI, E. P. SWIFT, A. HIBI, N.
ICHIDA, Y. TSUKAMOTO, & K. SONOBE. 1982. A field guide
to the birds of Japan. Tokyo, Wild Bird Soc. Japan. 336 pp.

1122. MAYR, E. 1945. Birds of the Southwest Pacific: A field guide to the
birds of the area between Samoa, New Caledonia and Micronesia.
New York, Macmillan. 316 pp. [Reprinted 1968, Codicote,
Wheldon & Wesley/Verlag J. Cramer: 1977 (Tuttle).]

1123. ——, & G. W. COTTRELL. [Eds.] 1979. Check-list of birds of the
world. Vol. 1, 2nd Ed. Revision of the work of James L. Peters.
Cambridge, Mass., Mus. Comp. Zool. 547 pp. [Procellariidae by C.
Jouanin & J.-L. Mougin, pp. 58-101; Podicipediformes by R. W.
Storer, pp. 140-155; Pelecaniformes by J. Dorst & J.-L. Mougin, pp.
155-193; Ardeidae by R. B. Payne, pp. 193-244; Ciconiidae by M.
P. Kahl, pp. 245-252; Threskiornithidae by J. Steinbacher, pp. 253-
268; Falconiformes by E. Stresemann & D. Amadon, pp. 271-425;
Anseriformes by P. A. Johnsgard, pp. 435-506. ]

1124. ——,&J. C. GREENWAY, JR. [Eds.] 1960. Check-list of birds of the
world. A continuation of the work of James L. Peters. Vol. 9.
Cambridge, Mass., Mus. Comp. Zool. 506 pp. [Alaudidae by J. L.
Peters, pp. 3-80; Hirundinidae by J. L. Peters, pp. 80-129;
Motacillidae by C. H. Vaurie, C. M. N. White, E. Mayr & J. C.
Greenway, Jr., pp. 129-167; Campephagidae by J. L. Peters, E.
Mayr & H. G. Deignan, pp. 167-221; Pycnonotidae by A. L. Rand
& H. G. Deignan, pp. 221-300; Irenidae by J. Delacour, pp. 300-
308; Lantidae by A. L. Rand, pp. 309-365.]

1125. ——, & ——. [Eds.] 1962. Check-list of birds of the world. A con-
tinuation of the work of James L. Peters. Vol. 15. Cambridge, Mass.,
Mus. Comp. Zool. 315 pp. [Ploceidae by R. E. Moreau & J. C.
Greenway, Jr., pp. 3-75; Sturnidae by D. Amadon, pp. 75-121;
Oriolidae by J. C. Greenway, Jr., pp. 122-137; Dicruridae by C.
Vaurie, pp. 137-157; Artamidae by E. Mayr, pp. 160-165; Corvidae
by E.R. Blake & C. Vaurie, pp. 204-282. ]

1126. ——, & R. A. PAYNTER, JR. [Eds.] 1964. Check-list of birds of the
world. A continuation of the work of James L. Peters. Vol. 10.
Cambridge, Mass., Mus. Comp. Zool. 502 pp. [Turdinae by S. D.
Ripley, pp. 13-227; Timaliinae by H. G. Deignan, pp. 240-427.]

1127. MEDWAY, LORD, & D. R. WELLS. 1976. The birds of the Malay
Peninsula. Volume V: Conclusions, and survey of every species.
London, Witherby. 448 pp.

84

1128.

1129:

1130:

1131.

1132.

1133.

1134.

1135.

1136.

NEMOURIA No. 29

MEYER DE SCHAUENSEE, R. 1984. The birds of China.
Washington, D. C., Smithson. Inst. Press. 602 pp.

MORONY, J. J., W. J. BOCK, & J. FARRAND. 1975. Reference list of
the birds of the world. New York, Dept. Orn., Amer. Mus. Nat.
Hist. 207 pp. [Reprinted 1978 with corrections and additions (2
pp.).]

NELSON, J. B. 1978. The Sulidae; gannets and boobies. Oxford,
Oxford Univ. Press. 1012 pp.

PAYNTER, R. A., JR. [Ed.] 1967. Check-list of birds of the world. A
continuation of the work of James L. Peters. Vol. 12. Cambridge,
Mass., Mus. Comp. Zool. 495 pp. [Pachycephalinae by E. Mayr,
pp. 3-51; Paridae by D. W. Snow, pp. 70-124; Sittidae by J. C.
Greenway, Jr., pp. 125-149; Rhabdornithidae by J. C. Greenway,
Jr., pp. 161-162; Dicaeidae by F. Salomonsen, pp. 166-208; Nec-
tariniidae by A. L. Rand, pp. 208-289; Zosteripidae Indo-Australian
taxa by E. Mayr, pp. 289-325; Genus Hypocryptadius by E. Mayr,
pp. 325-326. ]

—. [Ed.] 1968. Check-list of birds of the world. A continuation of the
work of James L. Peters. Vol. 14. Cambridge, Mass., Mus. Comp.
Zool. 433 pp. [Fringillinae by E. Mayr, pp. 202-206; Carduelinae by
T. R. Howell, R. A. Paynter, Jr. & A. L. Rand, pp. 207-306;
Estrildidae by E. Mayr, R. A. Paynter, Jr. & M. A. Traylor, pp. 306-
390.]

—. [Ed.] 1970. Check-list of birds of the world. A continuation of the
work of James L. Peters. Vol. 13. Cambridge, Mass., Mus. Comp.
Zool. 443 pp. [Emberizinae by R. A. Paynter, Jr., pp. 3-214.]

PETERS, J. L. 1934. Check-list of birds of the world. Vol. 2. Cam-
bridge, Mass., Harvard Univ. Press. 401 pp. [Megapodiidae, pp. 3-
9; Phasianidae, pp. 42-133; Turnicidae, pp. 142-149; Gruidae, pp.
150-154; Rallidae, pp. 157-213; Charadriiformes, pp. 226-359.]

—. 1937. Check-list of birds of the world. Vol. 3. Cambridge, Mass.,
Harvard Univ. Press. 311 pp. [Columbidae, pp. 11-141; Psittacidae,
pp. 141-273.]

——. 1940. Check-list of birds of the world. Vol 4. Cambridge, Mass.,
Harvard Univ. Press. [Cuculidae, pp. 12-76; Strigiformes, pp. 77-
174; Caprimulgiformes, pp. 174-220; Apodidae, pp. 220-256;
Hemiprocnidae, pp. 257-259. ]

December 31, 1985 Kennedy, et al. 85

1137. ——. 1945. Check-list of birds of the world. Vol. 5. Cambridge, Mass.,
Harvard Univ. Press. 306 pp. [Trogonidae, pp. 148-164;
Alcedinidae, pp. 165-219; Meropidae, pp. 229-239; Coraciidae, pp.
240-247; Upupidae, 247-249; Bucerotidae, pp. 254-272. ]

1138. ——. 1948. Check-list of birds of the world. Vol. 6. Cambridge, Mass.
Harvard Univ. Press. 259 pp. [Capitonidae, pp. 24-63; Picidae, pp.
86-232. ]

1139. —. 1951. Check-list of birds of the world. Vol. 7. Cambridge, Mass.,

Mus. Comp. Zool. 318 pp. [Eurylaimidae, pp. 3-13.]

1140. RAND, A. L., & E. T. GILLIARD. 1968. Handbook of New Guinea
birds. Garden City, New York, Nat. Hist. Press. 628 pp. [lst
publ. 1967. London, Weidenfeld & Nicolson.]

1141. RIPLEY, S. D. 1977. Rails of the world. A monograph of the family
Rallidae. Toronto, M. F. Feheley Publ. Ltd. 406 pp.

1142, SCHONWETTER, M. & W. MEISE. 1960-. Handbuch der Oologie.
Berlin, Akad.-Verlag. Vol. 1 (1960-1971) in 13 pts., 929 pp. At
1971, vol. 2, pts. 14-19, 384 pp.

1143. SEVERINGHAUS, S. R., & K. T. BLACKSHAW. 1976. A new
guide to the birds of Taiwan. Taipei, Taiwan, Mei Ya Publ. Inc.
222 pp:

1144. SHORT, L. 1982. Woodpeckers of the world. Greenville, Delaware,
Delaware Mus. Nat. Hist. 676 pp.

1145. SMYTHIES, B. E. 1981. The birds of Borneo. 3rd Ed. Kota
Kinabalu, Sabah, The Sabah Soc. 473 pp. [Third Ed. revised by
the Earl of Cranbrook. ]

1146. TRAYLOR, M. A., JR. [Ed.] 1979. Check-list of birds of the world.
A continuation of the work of James L. Peters. Vol. 8. Cam-
bridge, Mass., Mus. Comp. Zool. 365 pp. [Pittidae by E. Mayr,
pp. 310-329. ]

1147. VAURIE, C. 1959. The birds of the Palearctic fauna. Passeriformes.
London, Witherby. 762 pp.

1148. —. 1965. The birds of the Palearctic fauna. Non-Passeriformes.
London, Witherby. 763 pp.

1149. WEBSTER, M., & K. PHILLIPPS. 1977. A new guide to the birds
of Hong Kong. 2nd Ed. Hong Kong, Sino-American Publ. Co.

111 pp.

86 NEMOURIA No. 29

1150. WHITE, C. M. N., & M. D. BRUCE. 1986. The birds of Wallacea
(Sulawesi, Moluccas & Lesser Sunda Islands, Indonesia). Br. Orn.
Union Checklist Ser. No. 7. 576 pp.[In press. ]

1151. WILDASH, P. 1968. Birds of South Vietnam. Rutland, Vermont &
Tokyo, Charles E. Tuttle Co. Inc. 234 pp.

1152. WOLTERS, H. E. 1975-1982. Die Végelarten der Erde. Hamburg,
Paul Parey. 745 pp. [Ongoing publication in parts.]

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Editor
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Managing Editor
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Editorial Board
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Published by the
_ » DELAWARE MUSEUM OF NATURAL HISTORY
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Occasional Papers of the Delaware Museum of Natural History

NUMBER 30 FEBRUARY 28, 1988

THREE NEW SUBSPECIES OF PHILIPPINE BIRDS

Kenneth C. Parkes!

Subsequent to the publication of Philippine Birds (duPont 1971), the
Delaware Museum of Natural History (DMNH) has received shipments of bird
specimens from several islands of the Philippines that were previously
inadequately represented in collections. Through the courtesy of John E. duPont
I have been able to study these collections. Additional material of one species,
Stachyris capitalis, was examined at the National Museum of Natural History,
Washington, D.C. (USNM). These studies have revealed the existence of three
previously undiscovered subspecies.

Measurements and weights in this paper are given in metric units except
for altitudes quoted from collectors’ original labels, given in feet.

Stachyris capitalis (Tweeddale)

When I described S. c. isabelae from Basilan (Parkes 1963), I assumed (as
had all previous authors) that the birds of this species from the island of Min-
danao were identical to those of the island of Dinagat, the type locality of
capitalis. Dinagat, a relatively small (671 km?) island (see duPont and Rabor
1973 for description) lies only about 7.25 km N of the northernmost point of
Surigao, Mindanao. Its faunal affinities have always been judged to be with
eastern Mindanao (see various maps in Dickerson 1928), and the only endemic
bird subspecies from Dinagat hitherto recognized is Macronous striaticeps alcasidi
duPont and Rabor, 1973.

Tweeddale (1877) described ““Mixornis? capitalis” from a single specimen
from Dinagat, but by 1881 it had already been attributed also to the islands of
Panaon and Leyte. I have already shown (Parkes 1963, 1973) that the Leyte
record was erroneous, having been based on a specimen of Macronous striaticeps.

1Carnegie Museum of Natural History, Library of Congress
4400 Forbes Avenue Catalog No. 87-0735631

Pittsburgh, Pennsylvania 15213

2 NEMOURIA No. 30

Edward C. Dickinson has examined the alleged Panaon specimen (British
Museum [Natural History] 1888.4.20.1128), and finds that it, too, is an example
of M. striaticeps. The attribution of Stachyris capitalis to Basilan by Wardlaw
Ramsay (1881), on the other hand, was correct. The species was first collected on
Mindanao in 1889 by the Platens (Blasius 1890), and was never really considered
a Dinagat endemic.

Comparison of the recently received DMNH series from Dinagat with
specimens from many localities on Mindanao reveals that the populations on
that large island differ from true capitalis of Dinagat. Differentiation from
capitalis and from isabelae of Basilan is best developed in southeastern Min-
danao, so a holotype from that area has been selected. The Mindanao race may
be called:

Stachyris capitalis euroaustralis, new subspecies

HoLotTyPpE: DMNH 13770, adult male, collected at Kablon, Mt. Matutum,
Tupi, Cotabato, Mindanao Island, Philippines, on 3 June 1966, by D. S. Rabor
(collector’s number 46899).

Diacnosis: Differs from capitalis of Dinagat and isabelae of Basilan in having the
chest heavily suffused with gray and the sides and flanks extensively gray,
reducing thereby the cream-colored area of the abdomen. The under tail coverts
are predominantly gray rather than of the same cream color as the abdomen. The
crown color is more orange-brown, less chestnut than in capitalis, approaching
that of zsabelae. The dorsal streaking, on the other hand, is more like that of
capitalis, less heavily developed than in isabelae. Sex for sex, culmen (from base)
measurements of Mindanao birds were smaller than those of Dinagat capitals,
showing no overlap in the relatively small series measured. In flattened wing
measurements there was substantial overlap, but again the Mindanao specimens
averaged smaller. In body weight as recorded on labels (sexes combined),
Mindanao birds averaged slightly lighter (see Table 1 for measurements and
weights).

RANGE: Island of Mindanao, Philippines. Some specimens from the north-
easternmost provinces of Surigao and Agusan (i.e, nearest to Dinagat) approach
capitalis in color and are among the largest Mindanao birds measured. One
specimen from Zamboanga (DMNH 13768, Mt. Sugarloaf) is nearest isabelae of
Basilan, but this is not surprising, as Zamboanga Peninsula birds of several other
species are closer to Basilan races than to those of the rest of Mindanao (see
faunal subprovinces map in Dickerson 1928, p. 295).

ETyMo_ocy: The name is taken from the Latin Euroauster, the Southeast wind,
as the characters of the subspecies are best developed in southeastern Mindanao.
SPECIMENS EXAMINED: Dinagat: 11 (DMNH), 2 (USNM). Mindanao, Davao +

February 28, 1988 Parkes 3

Table 1: Measurements and weights of Stachyris capitalis
Measurements in millimeters to nearest 0.5, weights in grams

N Mean S.D. Range

Dinagat

Wing (flat), male 7 i235 1.8 69-75

Wing (flat) female 5 70.3 eal 68.5-74

Culmen from base, male 8 16.3 0.7 15.5-17.5

Culmen from base, female 5) 16.9 0.4 16.5-17.5

Weight (sexes combined) 6 1651 1.6 14.5-19
SE Mindanao

Wing (flat), male 14 70.5 1.9 67-73

Wing (flat) female 5 68.7 2.0 66-70.5

Culmen from base, male 10 14.7 0.7 13.5-15.5

Culmen from base, female 4 15.0 0.9 14-16

Weight (sexes combined) 24 14.6 0.9 13.5-16
NE Mindanao

Wing (flat), male 71.0 ed 69.5-71.5

Culmen from base, male 15.0 0.4 14.5-15.5
Culmen from base, female 15.5 0.8 14.5-16
Weight (sexes combined) 12 14.5 1.1 12.5-16.5

4

Wing (flat) female 6 68.7 0.9 67-69
4
6

Cotabato (SE): 5 (DMNH), 22 (USNM). Mindanao, Agusan + Surigao (NE): 2
(DMNH), 10(USNM). Mindanao, Zamboanga del Sur: 1 (DMNH). Mindanao,
scattered additional localities: 5(DMNH), 1 (USNM).

Orthotomus derbianus Moore

The holotype of Orthotomus derbianus was a specimen in the Liverpool
Museum, collected by Cuming (whose localities were often inadequately or
erroneously recorded), that Moore attributed to the Philippines with a query.
DuPont (1971) restricted the type locality to Luzon. As a similar species of
Orthotomus also occurs on Luzon, a further restriction would seem desirable,
and I hereby designate the type locality of Orthotomus derbianus Moore, 1855, as
“vicinity of Manila, Luzon, Philippine Islands.”’

A minor correction is appropriate here. Parkes (1971) and duPont (1971)
gave the date of publication of Orthotomus derbianus as “1854.” The name
appeared in the proceedings of the Zoological Society of London’s meeting of 12
December 1854, but these proceedings were not published until 8 May 1855.
The date is correctly given as 1855 by Watson (1986).

4 NEMOURIA No. 30

Until 1971, O. derbianus was considered to be conspecific with the widely
distributed O. atrogularis, of which the representative in northern Luzon is O. a.
chloronotus Ogilvie-Grant. I demonstrated (Parkes 1971) that these two tailor-
birds were sympatric in Laguna Province, at the east side of Laguna de Bay. At
that time I did not discuss altitudinal distribution, and have subsequently been
asked whether the two forms might be altitudinally isolated in the areas of
apparent sympatry. Elevations noted on labels of specimens in the DMNH
demonstrate that there is no clearcut altitudinal separation between these forms.
At Pakil, Laguna, chloronotus was collected at 50 and 100 feet elevation and
derbianus at 1100 feet, whereas in the vicinity of Pangil, Laguna, chloronotus was
collected at 1000 feet and derbianus at between 10 and 200 feet and between 1000
and 1200 feet.

With its removal from the species O. atrogularis, O. derbianus was believed
to be monotypic. My study of the recently received DMNH series of this species
from Catanduanes revealed that the tailorbirds of that island constitute a
recognizable subspecies, which may be known as:

Orthotomus derbianus nilesi, new subspecies

HOoLotyPeE: DMNH 67846, adult male, collected at Bate, Bato, Catanduanes,
Philippines, altitude 200’, 20 July 1979, by Filipino collectors for the DMNH.
DiaGnosis: Similar to O. d. derbianus of Luzon, but males appearing darker
below owing to narrower white streaking, especially on throat; olivaceous wash
on flanks found in most derbianus males lacking in nilest. Yellow edging of
remiges and coverts slightly brighter. Females, which are paler than males, differ
similarly, and in fact resemble males of derbianus. The relatively few specimens
with unworn tails suggest that mz/es: has a larger and more distinct blackish
subterminal spot on the rectrices, which would fit in with the increased
pigmentation of the underparts.

RANGE: Catanduanes Island, Philippines.

EtTmo.ocy: This subspecies is named for Dr. David M. Niles of the DMNH,
who has been consistently and generously helpful to my research on Philippine
birds.

SPECIMENS EXAMINED: derbianus: Luzon 43; niles1: Catanduanes, 21.

Dicaeum pygmaeum (Kittlitz)

The Babuyan Isands of Fuga and Calayan lie north of Luzon, and
demonstrate a moderate amount of avifaunal endemism. Salomonsen (1960)
examined a single male of Dicaeum pygmaeum from Fuga and a single female

February 28, 1988 Parkes 5

from Calayan (misspelled “‘Caleyan” by Salomonsen). On this basis, he
suggested the possibility that the populations of this species in the Babuyanes
“may constitute a slightly different subspecies.’”’ He stated that they were
“slightly larger” than typical pygmaeum, but his Fuga male was 1 mm. shorter in
wing length than his largest Luzon pygmaeum, and the Calayan female only 0.5
mm. longer in wing than a female from Sibuyan. It is clear that separation of a
race from the Babuyanes would have to rest on color rather than size characters.
Salomonsen described the Fuga male as having “‘distinctly more gloss on the
mantle and back, a deeper orange-vinaceus [s7c] tinge on the under parts, slightly
darker olive flanks, and the dark longitudinal patch on the center of the lower
breast and abdomen more well marked and distinct than do males of typical
pygmaeum.” He did not describe the Calayan female. In his account of the
Dicaeidae in the “‘Peters’”? Check-list, Salomonsen (1967) listed the Babuyan
populations under nominate pygmaeum as “‘subspecies?’’, although he admitted
D. p. salomonseni Parkes as the subspecies of northern Luzon, i.e., nearest to the
Babuyan Islands.

Receipt by the DMNH of an excellent series of Dicaeum pygmaeum from
Fuga has permitted the evaluation of Salomonsen’s suggestions as to the
distinctness of this population. Although not exactly conforming to Salomon-
sen’s description of the one male he examined, the Fuga birds do prove to
represent a distinct subspecies, which may be called:

Dicaeum pygmaeum fugaensis, new subspecies

HoLotTyre: DMNH 65668, adult male, collected on Fuga Island, Babuyanes
group, Philippine Islands, on 2 May 1979, by Filipino collectors for the
DMNH.

DIAGNOsIs: adult males are nearest salomonseni and pygmaeum, but have flanks
almost completely gray rather than greenish. The entire dorsum is much blacker
than in these races, and is slightly iridescent (especially the crown), thus ap-
proaching the blue-black color of the geographically distant subspecies davao of
Mindanao. The greenish rump area is more restricted and contrasts more with
the anteriorly adjacent area of the back, again an approach to davao, in which the
greenish rump patch is wholly or virtually absent. Adult females and immature
males are grayer, less green on the flanks than salomonseni and pygmaeum. They
are somewhat darker dorsally (blacker and less greenish) than in salomonseni and
much darker than in pygmaeum, with the greenish rump patch much reduced.
Detailed comparisons with the pale race palawanorum Hachisuka of Palawan are
not necessary.

RANGE: The island of Fuga in the Babuyanes group, north of Luzon, Philip-
pines. Specimens from other islands in this group were not examined, but would
be expected to be referable to this subspecies.

6 NEMOURIA No. 30

ETYMOLOGY: named for the type locality.

SPECIMENS EXAMINED: pygmaeum: Southern Luzon, 15; Catanduanes, 12;
Marinduque, 4; Mindoro, 6; Leyte, 3; Cebu, 1. salomonseni: Northern Luzon,
12. fugaensis: Fuga, 14.

REMARKS: The subspecies salomonseni Parkes of northern Luzon, based on
material other than that examined for the present study, proves to be somewhat
less well differentiated from pygmaeum in males than suggested by the original
description, in particular the dorsal and flank colors. However, the sides of the
head and breast of salomonseni are distinctly grayer, less blackish, with less
contrast with the pale throat, just as originally described.

As for Salomonsen’s description of his Fuga male, I fail to find any deep
orange-vinaceous tinge on the underparts, and there is no “dark longitudinal
patch on the center of the lower breast and abdomen”’ in fugaensis (as there is in
davao), although poorly made specimens of both Fuga and Luzon birds may
suggest the presence of such a patch.

Salomonsen (1960) suggested that because of the differences between
nominate pygmaeum and davao in ecology and color pattern, the latter form
might be treated as a full species. With the description of fugaensis, the color
differences are substantially bridged, and species status for davao on that basis
would not be warranted.

Deignan (1961), citing as authority only an unpublished manuscript of his
own, placed Dicaeum davao Mearns as a subspecies of D. ignipectus rather than
of D. pygmaeum. Salomonsen (1960) called attention to “‘striking”’ similarities
between davao and ignipectus, but maintained the former as a subspecies of
pygmaeum. That Salomonsen considered the resemblances between davao and
ignipectus superficial or convergent may be deduced from his inclusion of
pygmaeum (including davao) in a “‘species group Dicaeum concolor’ that did not
include ignipectus; the latter he later (Salomonsen 1961) placed in a “‘super-
species D. hirundinaceum,”’ which he regarded as “‘the most advanced group
within the genus Dicaeum.”’ I have compared davao directly with D. ignipectus,
and find that the former represents merely the intensification of colors and
patterns of other races of pygmaeus, especially the clearly intermediate fugaensis.
The chief resemblance of davao to ignipectus is its black, iridescent back, which is
approached in fugaensis. There is no indication in davao of the red chest of
ignipectus, and the bills of the two species are quite different in shape, that of
ignipectus being shorter and heavier than that of pygmaeum (including davao).

ACKNOWLEDGEMENTS

I am grateful for the hospitality continually afforded me at the Delaware
Museum of Natural History by John E. duPont and David Niles. The
manuscript was critically read by Edward C. Dickinson and Robert S. Kennedy.

February 28, 1988 Parkes 7
SUMMARY

Study of recently received Philippine bird specimens in the Delaware
Museum of Natural History revealed the existence of three undescribed sub-
species: Stachyris capitalis euroaustralis of Mindanao, Orthotomus derbianus nilesi
of Catanduanes, and Dicaeum pygmaeum fugaensis of Fuga and probably other
islands in the Babuyanes group. The type locality of Orthotomus derbianus
Moore is formally restricted to “‘vicinity of Manila, Luzon, Philippines.’’ The
proposal that Dicaewm pygmaeum davao of Mindanao should be considered a
subspecies of D. ignipectus is rejected.

LITERATURE CITED

Blasius, W.
1890 Die von Herrn Dr. Platen und dessen Gemahlin im Sommer 1889 bei
Davao auf Mindanao gesammelten Voegel. J. f. Orn. 38:144-149.

Deignan, H. G.
1961 Type specimens of birds in the United States National Museum. Bull.
U.S. Nat. Mus. 221:1-x, 1-718.

Dickerson, R. E.
1928 Distribution of life in the Philippines. Monogr. 21, Bur. Sci., Manila,

Philippine Is., 322 pp.
duPont, J. E.
1971 Philippine birds. Delaware Mus. Nat. Hist., Greenville, Delaware,
480 pp.

duPont J. E., & D. S. Rabor
1973 Birds of Dinagat and Siargao, Philippines: an expedition report.
Nemouria 10:1-111.

Parkes, K. C.
1963 Anew subspecies of tree-babbler from the Philippines. Auk 80:543-
544.
1971 ‘Taxonomic and distributional notes on Philippine birds. Nemouria
4:1-67.

1973 Annotated list of the birds of Leyte Island, Philippines. Nemouria
1131-73.

8 NEMOURIA No. 30

Salomonsen, F.

1960 Notes on flowerpeckers (Aves, Dicaeidae). 3. The species group
Dicaeum concolor and the superspecies Dicaeum erythrothorax. Am.
Mus. Novit. 2016:1-36.

1961 Notes on flowerpeckers (Aves, Dicaeidae). 4. Dicaeum igniferum and
its derivatives. Am. Mus. Novit. 2057:1-35.

1967 Family Dicaeidae. pp. 166-208 im Check-list of birds of the world, vol.
12. Mus. Comp. Zool., Cambridge, Mass., 433pp.

Tweeddale, Marquis of

1877 Descriptions of some new species of birds. Ann. Mag. Nat. Hist.

4(20):533-538.
Wardlaw Ramsay, R. G.

1881 Revised list of birds known to occur in the Philippine Islands,
showing their geographical distribution. pp. 654-660 im The or-
nithological works of Arthur, Ninth Marquis of Tweeddale, ed. by R.
G. Wardlaw Ramsay. Taylor & Francis, London, 760 pp.

Watson, G. E.

1986 Family Sylviidae (Holarctic and Oriental), 7m Check-list of birds of the

world, vol. 11. Mus. Comp. Zool., Cambridge, Mass., 638 pp.

Editor
JOHN E. pu PONT

Managing Editor
Davip M. NILEs

Editorial Board
KENNETH C. PARKES, Carnegie Museum of Natural History, Pittsburgh, PA
RICHARD W. THORINGTON, JR., National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Published by the
DELAWARE MUSEUM OF NATURAL HISTORY
Kennett Pike, Greenville,
Delaware 19807

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