Nl
HARVARD UNIVERSITY
mm
LIBRARY
OF THE
Museum of Comparative Zoology
TRANSACTIONS
OF THE
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOLUME 15
1968-1969
Printed from the
W. W. Whitney Publication Endowment
COMMITTEE ON PUBLICATION
KURT K. BOHNSACK JOSEPH R. JEHL, JR.
BAYLOR BROOKS RICHARD P. PHILLIPS
JOHN A. COMSTOCK, M.D. ARNOLD ROSS
EDWARD CREUTZ CHARLES YORK
CARL L. HUBBS HERBERT F. YORK
MUS. COMP. ZOCL.
LIBRARY
OEC 1 ! 1969
HARVARD
UNiVERsj-nj
CONTENTS
1. Geographic and seasonal variation in Smith's longspur, Calcarius pictus.
By Joseph R. Jehl, Jr. 8 January 1968 1-6
2. New Panamic nudibranchs (Gastropoda; Opisthobranchia) from the Gulf
of California. By James R. Lance. 8 January 1968 7-20
3. Thermal relations, distribution, and habitat of Cnemidophorus labialis
(Sauria: Teiidae). By Dennis L. Bostic. 5 June 1968 21-30
4. Castela polyandra, a new species in a new section; union of Holacantha with
Castela (Simaroubaceae). By Reid Moran and Richard Felger. 5 June 1968 31-40
5. Recent data on summer birds of the Chiricahua Mountains area, southeast-
ern Arizona. By J. David Ligon and Russell P. Balda. 5 June 1968 41-50
6. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. I.
General account. By J. Laurens Barnard and John R. Grady. 17 June 1968 51-66
7. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. II.
Benthic polychaetous annelids. By Donald J. Reish. 17 June 1968 67-106
8. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. III.
Benthic Mollusca. By Eugene V. Coan. 25 September 1968 107-132
9. Type specimens of birds in the San Diego Natural History Museum. By
Joseph R. Jehl, Jr. 25 September 1968 133-139
10. Geographic variation in the cling fish, Gobiesox eugrammus Briggs. By
David W. Greenfield and James W. Wiley. 25 September 1968 141-147
1 1 . Five new taxa of Haplopappus (Compositae) from Baja California, Mexico.
By Reid Moran. 12 February 1969 149-164
12. Fossil grouse of the genus Dendragapus. By Joseph R. Jehl, Jr. 12 Februarv
1969 165-174
13. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. IV.
Benthic Amphipoda (Crustacea). By J. Laurens Barnard. 27 June 1969 .... 175-228
14. A recent molluscan fauna from the Caribbean coast of southeastern Pan-
ama. By George E. Radwin. 27 June 1969 229-236
15. Studies on the Tetraclitidae (Cirripedia: Thoracica): Revision of Tetraclita.
By Arnold Ross. 22 September 1969 237-251
16. Type specimens of mammals in the San Diego Natural History Museum. By
Suzanne I. Bond. 3 October 1969 252-263
17. Twelve new dicots from Baja California, Mexico. By Reid Moran. 15 Octo-
ber 1969 265-295
18. Observations on a young pygmy killer whale (Feresa attenuata Gray) from
the eastern tropical Pacific Ocean. Bv William F. Perrin and Carl L. Hubbs.
24 November 1969 297-308
LIBRARY
JAN 25
HARVARD
UNIVERSITY
GEOGRAPHIC AND SEASONAL VARIATION
IN SMITHS LONGSPUR, CALCARIUS PICTUS
JOSEPH R. JEHL, JR.
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 1, 8JANUARY 1968
LIBRARY
GEOGRAPHIC AND SEASONAL VARIATION
IN SMITH'S LONGSPUR, CALCARIUS PICTUS
Joseph R. Jehl, Jr.
JAN 25
HARVARD
UNIVERSITY
ABSTRACT
Color differences that have been used to define races of Smith's Longspur result from
seasonal wear and fading of the breeding plumage. There is no demonstrable geographic
variation in this species.
Smith's Longspur (Calcarius pictns) is an endemic species of the Nearctic sub-
arctic avifauna (Johansen, 1963) that breeds from north-central Alaska eastward to
Cape Henrietta Maria, Ontario (A.O.U., 1957; Gabrielson and Lincoln, 1959; Godfrey,
1966; Kemsies, 1961); apparently a small population breeds in northwestern British
Columbia (Rand, 1948; Weeden, 1960). Owing to insufficient data from Alaska and
central Canada, the breeding distribution of this species cannot be mapped accurately.
The map (Fig. 1) is based on the assumption that throughout its range, as at Churchill,
Manitoba (Jehl, Ms.), Smith's Longspur breeds primarily in the narrow belt of tundra
bordering the treeline {i.e., the forest-tundra of Johansen, 1963).
In a recent study, Kemsies (1961) argued that three subspecies of Smith's Long-
spur should be recognized. The approximate ranges of these races are shown in Figure 1.
According to Kemsies, males in breeding plumage from north-central Alaska (C. p.
roiveorum) are browner dorsally and deeper orange ventrally than males of the nomi-
Figure 1. Presumed breeding range of Smith's Longspur. The distribution of
races recognized by Kemsies (1961) is indicated.
the
: San Diego Society of Natural History Vol. 1 5
nate race, which breeds from extreme northeastern Alaska to the northeastern corner of
Manitoba. Males from the Hudson Bay Coast of Ontario (C. p. mersi) are blacker
dorsally and paler ventrally than nominate pictus. Thus, from west to east male Smith's
Longspurs exhibit increasing blackness of the dorsum but increasing paleness of the
venter. The apparently well-marked racial differences are clearly illustrated in Kemsies'
Figure 1 (1961: 144). Females show parallel though less conspicuous clinal variation.
In the summers of 1965, 1966, and 1967, I studied the breeding biology of Smith's
Longspur at Churchill, Manitoba (Jehl, Ms.). Specimens collected for data on repro-
ductive biology and feeding habits were taken without conscious selection throughout
the summer, except that in 1966 and 1967 I selected for territorial or known breeding
birds. I made no attempt to sample plumage variation present in the population at any
one time, nor to collect birds that appeared to be migrating.
Since Churchill is in the eastern half of the range of C. p. pictus, I was surprised
to discover breeding birds fitting the description of each of the alleged races in my
sample. Further study showed that variation in the Churchill population is clearly
attributable to seasonal wear and fading of the breeding plumage. Birds collected in the
first days of June are brightly colored and can be referred to roweorum; throughout
most of June pictus predominates; by late June the first mersi appear and by mid-July
the entire population is dark-backed and pale-bellied. The seasonal variation in males is
shown in Figure 2 (cf. Kemsies' Fig. 1; 1961: 144). The specimens illustrate approxi-
mately average plumage coloration and wear in the population at the time they were
collected. I emphasize that they represent breeding birds. There is no possibility that
birds taken in June were migrants to Alaska, where nesting commences earlier than
at Churchill (Irving, 1960: 124-125), or that the July birds were post-breeding wan-
derers from Ontario. Females do not show such extreme seasonal variation, presumably
because of their reduced activity during incubation.
A re-examination of Kemsies' data suggests that he was misled by biased sampling
and inadequate material, and that seasonal rather than geographic variation accounts
for the differences he recognized. For example, 14 of the 15 Alaskan specimens that he
assigned to roweorum were collected between 27 May and 9 June (type collected
4 June), when birds are still in relatively unworn plumage. The entire series of mersi
from Ontario (45 specimens) was collected between 22 June and 10 August (type
collected 31 July); most of these specimens can be expected to show advanced stages
of wear and fading.
Inadequate samples plus seasonal variation could also explain several of Kemsies'
additional, and otherwise puzzling, findings.
1. I he existence of three well-marked races along a narrow belt of tundra that is
not interrupted by any obvious barriers to gene flow.
2. The small breeding ranges of roueorum and mersi.
3. The opposite trends in clinal variation of dorsal and ventral coloration noted
above.
4. The abundance of roueorum during migration. Kemsies referred 5/ of 66 speci-
mens collected south of the breeding grounds in April and May to roueorum and the
remainder to pictus. If those assigned to roueorum are other than freshly molted in-
s difficult to explain the far greater abundance of the race with the much
needing grounds, especially since Gabrielson and Lincoln (1959: 818) state that
Smith's longspurs are not common in Alaska.
ate numbers of roweorum also occur in fall. Of 24 specimens taken
breeding grounds between 15 August and 30 October, Kemsies assigned
"?, 3 to pictus, even though he gave no criteria by which the races could
bc distingt the nonbreeding plumage! If he assumed that roweorum would be
1968
Jehl: Variation in Smith's Longspur
Figure 2. Seasonal plumage variation in male Smith's Longspurs collected at Churchill,
Manitoba. Top and bottom, left to right: 2 June 1966; 9 June 1967; 11 June 1966;
16 June 1965; 25 June 1966; 1 July 1966; 11 July 1966.
4 San Diego Society or Natural History Vol. 15
brow [km- at .ill seasons, he would have consequently assigned most of the fresh-plumaged
birds to that race.
5. The virtual absence of mersi away from the breeding grounds. Kemsies lists
only one migrant specimen of mersi (Hamilton, Kansas, 21 March 1914). Since the
breeding range of this subspecies is approximately as large as that of roweorum, and
since mersi would appear to be relatively common on the Ontario coast (26 specimens
collected in 1948, alone), the relative disproportion is especially notable.
6. The relative abundance of the races in Ohio. Smith's Longspurs do not nor-
mally occur east of Ohio and one would predict that many, perhaps most, of the spring
migrants in that state would be referrable to the easternmost breeding population. Yet,
/5 of the 20 specimens listed by Kemsies taken between 19 March and 24 April are
assigned to roweorum and the remainder are classified as pictus. However, if the alleged
geographic variation is seasonal, the racial identifications and the absence of mersi are
not surprising, for at this season birds are molting into fresh plumage and are unworn.
Through the courtesy of the curators acknowledged below, I have been able to
examine most of the specimens from the breeding grounds on which Kemsies based his
descriptions, and to compare them with seasonally comparable material from Churchill.
Specimens examined (see Appendix) included 19 from Alaska, including the type of
you corn in, 21 from the Northwest Territories, 3 5 from Churchill, and 3 8 from the
Ontario coast, including the entire series of mersi paratypes. Of these 6 5 per cent had
been examined previously by Kemsies. My study confirmed that seasonal variation was
the basis for Kemsies' conclusions. All specimens from Churchill and those that Kemsies
had assigned to pictus were indistinguishable from those assigned to roweorum or mersi,
provided they acre compared with specimens taken at the same stage of the breed-
ing cycle and acre in a comparable stage of wear. Moreover, even some slightly worn
specimens of roweorum from Alaska (e.g., U. S. National Museum no. 454878, Summit,
Alaska, 3 June 1952) were inseparable from relatively unworn specimens of mersi (e.g.,
Royal Ontario Mus. Zool. no. 93 5 57, 20 mi. S. Cape Henrietta Maria, 14 July 1948).
Furthermore, I found no differences between several juveniles from Churchill and one
from the Ontario coast, or between Churchill birds in fresh non-breeding plumage and
a specimen from Alkavik, NWT, previously assigned to roweorum. Ontario birds in
non-breeding plumage and Alaskan juveniles were not available.
Kemsies' data did not indicate geographic size variation in this species. I also found
no differences between Churchill and Alaskan birds (Table 1); unworn specimens from
Ontario were not available for comparison. The wing and tail measurements of Churchill
males taken in late July average nearly 2 mm shorter than those collected in early June.
To reduce possible error, the dimensions of males collected after 20 June were not in-
cluded in Table 1.
TABLE 1
Ml ASURIMl-NTS OF SMITH'S LONGSPURS1
I uposed Oilmen Wing (Chord) Tail Tarsus
No. Range and Mean No. Range and Mean No. Range and Mean No. Range and Mean
Churchill males IS 9.9-11.7 (10.6) II 88-94 (91.8) n 59-67 (63.7) 18 20-22 (20.9)
Alaskan males \\ 9.9-11.4 (10.6) 13 87-95 (92.2) 12 57-65 (62.5) 13 20-23.5 (20.9)
Churchill females 9 9.9-11.4 (10.4) 9 84-90 (87.5) 9 57-60 (58.8) 9 20-23.5 (20.9)
Alaska females 4 9.7-10.3 (10.0) 6 87-91 (88.4) 6 58-62 (60.0) 6 20-21.5 (20.6)
'Oilmen measured to n iresr 0.1 mm, tarsus to nearest 0.5 mm, wing and tail to nearest 1 mm. Wing
and tail measurements from Churchill males collected after 20 June are omitted (see text for explanation).
All Alaskan specimens were collected on or before 20 June.
1968 Jehl: Variation in Smith's Longspur >
In summary, Smith's Longspur shows no taxonomically recognizable geographic
variation in either plumage coloration or size. Therefore, acceptance of the proposed
races must be withheld.
Acknowledgments
I am grateful to the Frank M. Chapman Fund of the American Museum of Natural History, The
University of Michigan, the National Science Foundation, and the San Diego Society of Natural History
for supporting my field work at Churchill, Manitoba. The following persons kindly made specimens
available for study: Richard C. Banks and Richard L. Zusi, United States National Museum; Jon C.
Barlow, Royal Ontario Museum of Zoology; Tom J. Cade, Cornell University; W. Earl Godfrey, National
Museum of Canada; Kenneth C. Parkes, Carnegie Museum.
W. Earl Godfrey, Harrison B. Tordoff and D. J. T. Hussell offered helpful suggestions on the
manuscript.
Literature Cited
American Ornithologists' Union
1957. Check-list of North American birds. Fifth ed.
Gabrielson, I. N., and F. C. Lincoln
195 9. The birds of Alaska. Stackpole Co., Harrisburg, Penna., and Wildlife Management Inst.,
Washington, D.C.
Godfrey, W. E.
1966. The birds of Canada. Nat. Mus. Canada Bull. 203.
Irving, L.
1960. Birds of Anaktuvuk Pass, Kobuk, and Old Crow. U.S. Nat. Mus. Bull. 217.
Jehl, J. R, Jr.
Ms. The breeding biology of Smith's Longspur, Calcarius pictu\.
Johansen, H.
1963. Zoogeographical aspects of the birds of the subarctic. Proc. XIII Intern. Ornithol. Cong.:
1117-1123.
Kemsies, E.
1961. Subspeciation in the Smith's Longspur, Calcarius pictus. Canad. Field-Nat. 75: 143-149.
Rand, A. L.
1949. Distributional notes on Canadian birds. Canad. Field-Nat. 62: 175-180.
Weeden, R. B.
1960. The birds of Chilkat Pass, British Columbia. Canad. Field-Nat. 74: 119-129.
Manuscript Received 23 October 1967
Appendix
Specimens Examined
ALASKA. Anaktuvuk Pass (8): June 3, 4 (3), 5, 8, 19, 20. Summit (2): June 3, 9. Giant Creek (1):
June 2. Tolugak Creek (7): May 27, 28 (2), 29, June 4, 5, 9. Contact Creek (1): June 20. MAC-
KENZIE DISTRICT, NWT. Fort Good Hope (3): May 31. Fort Simpson (16): May 18, 20 (10), 21
(2), 25 (2), 26. Aklavik (2): May 26, August 19. MANITOBA. Churchill (35): June 2 (2), 4 (3),
9 (4), 11 (3), 13, 16 (3), 18, 2 5, 3 0, July 1, 9, 11 (2), 13 (2), 15, 16 (2), 22, 24, August 12 (2), 25,
26 (2). ONTARIO. Little Cape (9): July 2 9, 31 (5), August 1 (3). Fort Severn (6): June 2 2 (2).
27 (2), July 1, 14. Winisk (1): June 30. 54 mi. below Hawley Lake (1): June 22. W.icliee Creek (1):
June 25. Cape Henrietta Maria (19): July 2 (2), 3 (3), 5 (3), 7, 9 (2), 12 (2), 14 (6). 20 mi. S. Cape
Henrietta Maria (1): July 14.
/ARD
UNIVERSITY
NEW PANAMIC NUDIBRANCHS
(GASTROPODA; OPISTHOBRANCH1A)
FROM THE GULF OF CALIFORNIA
JAMES R. LANCE
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 2, 8 JANUARY 1968
L ":Y
JAN?
HARVARD
UNIVERSITY
Plate 1. Ncmbrotha hubbsi, sp. nov. Kodachrome by Wesley M. Farmer.
Acanthodoris stohleri, sp. nov. Ektachrome by Walter E. Harvey
es K. Lance.
NEW PANAMIC NUDIBRANCHS
(GASTROPODA: OPISTHOBRANCHIA)
FROM THE GULF OF CALIFORNIA
James R. Lance1
ABSTRACT
Three new species of shell-less euthyneuran mollusks from the northern part of the Panamic
faunal province are described. These are: Chromodorh faycac sp. now, Acanthodoris stohlcri sp.
nov. and Nembrotha hubbsi sp. nov. The last taxon represents the first record for this genus in
the eastern Pacific. Other northeastern Pacific species of the treated genera are compared and dis-
cussed. Anatomical details and geographic ranges are provided.
Although some 1600 species of mollusks have been reported from the Panamic
faunal province (Keen, 1958: 7), only 20 species of nudibranchs (Order Nudibranchia)
are known from the region. This small number of species reflects a general disinterest of
malacologists in observing and collecting shell-less mollusks and, in part, the inaccessi-
bility of the region, rather than a paucity of material. Expeditions to the Gulf of Cali-
fornia undertaken between 1962 and 1967, by the San Diego Natural History Museum,
the Santa Barbara Museum of Natural History, and the writer, have collected large
numbers of these animals. These collections apparently include many new species,
of which three are described below. Numerous individuals of each species from different
geographic localities have been compared and color transparencies have been made of
the animals in life. The illustrations portray typical individuals from the obtained
samples. The systematic treatment follows Odhner's (1939) revision.
Species Accounts
Family Dorididae
Subfamily Chromodoridinae
Chromodoris fayeae sp. nov.
Type material. — Holotype: California Academy of Sciences, Department of In-
vertebrate Zoology no. 314. Collected 6 March 1966 in the rocky, lower intertidal
region at Santa Cruz, Nayarit, Mexico (approximately ten miles south of San Bias);
21°30'N, 105°16'W. Paratypes: Santa Barbara Museum of Natural History nos.
19669 (two specimens collected 6 March 1966) and 2 5 000 (15 specimens collected 4
February 1967).
Description. — The largest specimen measured 47 mm long and 1 1 mm broad
while crawling actively. The notum is moderately expanded in front and behind, nar-
rower in the middle and completely covers the foot except posteriorly where the pointed
tail protrudes (Fig. 1). The notal surface is soft and smooth, its margins entire when
the animal is gliding, but it may become crenulate when the animal is irritated. The
vertical sides of the body are completely covered by the notum, and are not partially
exposed as in HypscloJoris californiensis (Bergh, 1879).
The spectacular color pattern of C. fayeae at once distinguishes it from all other
described northeastern Pacific opisthobranchs. The general ground color is pure white,
not the colorless translucency of many other species. Three distinct, uninterrupted
lines of contrasting color occur around the notal brim: an inner band of opaque white,
which contrasts with the pellucid white of the body, a central line of vivid red, and a
!Stazione Zoologica di Napoli. Present address: 746 Agate St., San Diego, California 92109.
San Diego Society of Natural History
Vol. 15
Fig. 1
10mm
Fig. 2
Fig. 3
1968 Lance: New Panamic Nudibranchs 5
marginal edging of brilliant yellow. This same pattern is repeated around the foot mar-
gins. It is most intense posteriorly and gradually fades anteriorly and disappears at about
the level of the rhinophores. The inner and outer surfaces of the distal portions of the
branchial stems are red. The perfoliate regions of the rhinophores are colored a similar
but darker red, especially on their anterior and posterior axial ridges. Some preserved
specimens retained a faint trace of red after one year in 70 per cent isopropyl alcohol,
but this was lost after the second year. The opaque white and yellow bands faded two
months after preservation.
Anteriorly the foot is expanded and produced into sharp corners (Fig. 2). Its
frontal margin is rounded, deeply bilabiate and entire, not notched. The lateral margins
are nearly parallel except posteriorly, where they converge to form a pointed tail with a
low dorsomedian crest. The anterolateral corners of the head are produced into short,
digitform tentacles, between which the mouth appears as a longitudinal slit (Fig. 2a, b).
The rhinophores are completely retractile into chambers which bear sheaths with
low, smooth margins. These sensory organs are relatively short but deeply perfoliate
with 21 lamellae (in a 36 mm specimen) joined by a longitudinal septum on both the
anterior and posterior faces.
The elegantly developed branchiae are simply pinnate, although some of the anterior
primary stems may bifurcate distally. They are completely retractile within a cavitv
bearing a low, smooth margin and are disposed in an arc with enrolled extremities
similar to those described and figured by Eliot (1904) for several species of Indo-
Pacific chromodorids. The 24 to 32 branchiae gradually decrease in size toward the
extremities of the enrolled arc. Except for momentary periods of quiescence, these ap-
pendages are continually in motion. The movement of each branchium is a rapid
twitching from side to side and is entirely unlike the rhythmical circulatory pulsations
observed in the respiratory appendages of many other cryptobranch dorids. Eliot (1904)
has reported similar branchial motion for Chromodorh annulata, C. runcinata, C. tryoni
and Casella atromarginata. Gohar and Aboul-Ela (1957) have observed sideward vibra-
tions in Chromodor'n pitlchclla and C. annulata from the Red Sea. Such a respiratory
adaptation is obviously advantageous to organisms inhabiting waters low in dissolved
oxygen.
The labial armature consists of a horn-colored, circular band of minute rodlets
arranged in an irregular series. Each rodlet is flexed near its midpoint (from nearly
straight to about 90°) and terminates in a bifid cusp (Fig. 3b). From above, only the
bifid hooks and portions of their supporting rods are evident (Fig. 3a). Bergh (1905,
pi. 16) figures similar labial elements for a number of Indo-Pacific chromodorids.
The radula is broad and deeply grooved medially. The teeth and basal membrane
are colorless in freshly killed individuals. The radular formula from a typical specimen
is 129 x 42. 1. 42. An average dental row consists of a dwarf but robust rachidian tooth
with a blunt cusp (Fig. 4a) flanked on each side by a series of unicuspidate lateral teeth
that very gradually increase in size outward (Figs. 4b,c,d). The inner laterals bear a
linear series of 6 to 8 denticles on the outer sides of the cusps and fewer on the inner.
The denticles decrease in number outward and become obsolete at about the level of the
3 0th tooth. The outermost laterals are smooth hamate hooks with elongated bases.
White, stellate spicules of two principal size classes were found imbedded in the
Figures 1-5. Chromodor'n fayeae. 1. Dorsal aspect of animal in life. 2. Anteroventr.il
aspect of animal in life: a. oral tentacle; b. mouth; c. upper foot lip; d. lower foot lip;
e. foot; f. hyponotum. 3. Elements of the labial armature: a. top view of exposed por-
tion of elements in situ; b. lateral view of three typical elements freed from matrix.
4. Representative teeth from left side of 60th row of radula: a. rachidian; b. 1st lateral;
c. 2nd lateral; d. outermost tooth (42nd). 5. Representative spicules from innermost
layers of notal epidermal tissue; the two size classes are illustrated.
6 San Diego Society of Natural History Vol. 15
inner epiderm.il layers of the notum in freshly killed specimens (Fig. 5). They were
concentrated in a loose band circumscribing the margins of the body cavity. The larger
elements attained a maximum diameter of 140/t while the smaller were about half
that size.
No trace of a penial armature was found.
Distribution. — This species has been taken on the open Pacific coast of mainland
Mexico and from several localities within the Gulf of California: (1) Bahia San Luis
Gonzaga, 23 September 1962, 1 specimen; (2) Isla Angel de la Guarda, 1957, 7 speci-
mens; (3) Isla Santa Cruz, 26 June 1964, 9 specimens; (4) Mazatlan (yacht club
cove), 20 December 1961, 2 specimens; (5) Santa Cruz, Nayarit (type locality),
6 March 1966, 4 specimens; 4 February 1967, 16 specimens; (6) Tentacatita, Jalisco,
6 February 1963, 1 specimen. Vertical distribution of the animals ranged from lower
intertidal to 37 feet.
Name. — The specific patronym fayeae was chosen to honor Faye B. Howard, Re-
search Associate in Conchology, Santa Barbara Museum of Natural History, in recog-
nition of her many outstanding contributions to our knowledge of Panamic mollusks.
Discussion. — Following the opinions of Odhner (1957: 250-253), Gohar and
Aboul-Ela (1957: 204), Marcus and Marcus (1960: 901), Burn (1962: 152), Farmer
(1963: 81) and Marcus and Burch (1965: 247-248), I reject the generic name Glosso-
doris Ehrenberg, 1831, in favor of Chromodoris Alder and Hancock, 1855, for species
of Chromodoridinae with unicuspidate lateral teeth, and Hypselodoris Stimpson, 185 5,
for those bearing bicuspidate ones. This choice is necessary because the radula of Ehren-
berg's Glossodoris has never been examined. For a review of this controversy see Odhner
(1957).
According to these views the present species must be assigned to Chromodoris on
the basis of its gross morphology, dwarf rachidian, and unicuspidate radular teeth.
Marcus (1965: 272-273) observed that elements of the labial armature, which gen-
erally consist of rodlets, either simple or terminating in a bifid hook, or are scale-like
platelets, are of taxonomic significance in the Chromodoridinae. Thus, the flexed rodlets
with terminal bifid hooks of the present species further substantiate its generic
assignment.
This is the third endemic chromodorid described from the Panamic region.
Family Gymnodorididae
Nembrotha hubbsi sp. nov.
(Plate 1)
Type material. — Holotype: California Academy of Sciences, Department of In-
vertebrate Zoology no. 315. Collected 21 April 1962 at a depth of 10 m near Isla
la Ventana, Bahia de Los Angeles, Baja California, Mexico; 28°55'N., 113°32'W.
Paratypes: California Academy of Sciences, Department of Invertebrate Zoology no. 316
(one specimen); Santa Barbara Museum of Natural History no. 23705 (one specimen).
Collecting data for paratypes same as holotype.
Description. -- Dimensions of actively crawling individuals observed by Farmer
and Sloan were: length, 38, 32, 21, 25, 19 mm; width, 9, 9, 4, 4, 3 mm; height, 11,
I I, 3.5, 6, 4 mm, respectively. However, one preserved specimen from Isla la Ventana
measured 5 4 mm in length, indicating that live animals may grow to be at least 60-70
mm long.
I he general body shape is limaciform, rounded in front, highest in the cardiac
region and tapering to a pointed tail. The surface is smooth, not furnished with
tubercles, papillae, or extra-branchial processes. The head region is provided with a
1 frontal veil which passes around the anterolateral corners and becomes obsolete
ibout i Ik- level of the rhinophores. Distinct pallia! ridges are not present.
striking coloration of this species at once separates it from all other north-
tuu idibranchs. The background color is a deep yellow ochre. Five longi-
1968
Lance: New Panamic Nuoibranchs
tudinal cerulean blue stripes of varying width, bordered on each side by a narrow line
of intense blue-black or black, occur on each side of the body and extend for most of
the length. A similar stripe arises medially in the head region, passes back between the
rhinophores, is interrupted in the branchial region, and continues posteriorly. Most of
the lines converge on the tip of the tail. The inner surfaces of the branchiae arc dark
blue-black; their outer faces a deep yellow ochre with a blue line running up the pri-
mary stem. The rhinophores are dark blue-black. The general color pattern is retained
after five years in 70 per cent alcohol.
The prominent rhinophores are perfoliate and retractile within chambers bear-
ing low, smooth, upstanding sheaths. The number of rhinophoral lamellae was not
determined.
There is a circlet of 5 branchiae located medially on the posterior slope of the
cardiac hump. As in other gymnodorids, the branchiae are not retractile within a
subepidermal chamber but are highly contractile. Due to maceration, the configuration
of the finer structures of the branchiae could not be determined.
The radular morphology was determined from one of the larger specimens (41 mm
preserved). The formula at the level of the 10th row was 3 5 x 3-4. 1. 1. 1. 3-4. The
rachidian teeth (Fig. 6a) consist of thickened arched bases with the extremities de-
rachidians reported for other species of Nembrotha (Eales, 1938: 96; Marcus, 1956: 47).
veloped into upstanding subequal prongs, and are much more robust than the delicate
A well developed central cusp, as in N. gracilis Bergh, 1877 (pi. 56) and N. rubro-
papulosa Bergh, 190 5 (pi. 18) is lacking in N. hubbsi. The prominent first laterals are
U-shaped with asymetrical extremities terminating in a pair of long sharp points
(Fig. 6b). The outer side of the shoulder bears a grooved process which apparently
hinges with the first marginal plate. The 3 to 4 marginal plates are greatly reduced
irregular triangles and decrease in size outwards (Fig. 6c, d, e, f). The attached bases
of the rachidians and first laterals, and upper portions of the outer marginal plates are
a dark yellowish-brown; the free portions are colorless. A labial cuticle was not found.
Mr. Farmer (personal communication) has observed this species swimming in an
aquarium by undulating its body from side to side in a manner similar to that of
Flabellinopsis iodinia (Cooper, 1862).
Distribution. — To date this species has been collected only in the northern part
of the Gulf of California. Five specimens were obtained by Prof. Hubbs and party
from: (1) Isla la Ventana (type locality), 3 specimens; (2) reef betwen Isla la Ventana
— ■■■! '.■'■ I I I l i
Figure 6. Nembrotha hubbsi. Typical teeth from 10th row of radula: a. rachidian;
b. 1st lateral; c-f. marginals.
8 San Diego Society of Natural History Vol. 1 5
and [sla Cabeza de Caballo, 1 specimen; (3) Punta Que Malo, 1 specimen. All localities
are within Bahia de los Angeles. Additional material was collected by Wesley Farmer
and John Sloan in May, 1962, and March, 1963, at Isla Cerralvo, Bahia de los Angeles,
and Puerto Refugio on Isla Angel de la Guarda. I have also tentatively identified this
species from color motion pictures taken in the La Paz area during a recent California
Academy of Sciences expedition to the Gulf of California.
Name. — The specific name bubbsi was chosen to honor Prof. Carl L. Hubbs of
the Scripps Institution of Oceanography, who first brought this elegent animal to my
attention.
Discussion. — Bergh (1877) erected the genus Nembrotha to accommodate a
number of colorful, limaciform gymnodorids from the western Pacific, principally the
Philippine Sea. This genus differs from the related Gymnodoris Stimpson, 18 5 5
(—Trevelyana Keelart, 1858) by possessing rachidian teeth, reduced number of laterals,
tower gills, and absence of a prostatic portion of the vas deferens. This last character,
however, applies only at the specific level as a prostate has been demonstrated to occur
in N. lineolata (see Eales, 1938: 97) and N. nigerrima (see Macnae, 1957: 361).
Burn (1962: 98) erected Tambja for those species of Ncmbrotba bearing a
rachidian tooth in the form of a simple rectangular plate, bifid first lateral, a highly
reduced number (3 to 4) of marginal teeth, and a pair of short extra-branchial
processes. This last character, however, is neither figured nor described for Tambja
sagamiana (=N. sagamiana Baba, 195 5) contrary to Burn's (loc. cit.) statement.
The radula of N. bubbsi is most similar to that of N. gracilis Bergh, 1877 (pi. 56)
and N. rubro papulosa Bergh, 1905 (pi. 18), but may be readily distinguished by the
different configuration of the rachidian, lateral, and marginal teeth. Also, the color
pattern is entirely different. Until a comprehensive review of the genus is undertaken,
I prefer to assign the present species to Ncmbrotba (sensu lato) . This species is the first
eastern Pacific representative of this colorful tropical genus.
Family Onchidorididae
Acanthodoris stohleri sp. nov.
(Plate 2)
Type material. — Holotype: California Academy of Sciences, Department of In-
vertebrate Zoology no. 307. Paratype: no. 308 at the same institution. Both specimens
collected at Bahia San Luis Gonzaga (inner side of Isla Willard), Baja California,
Mexico (20° 48'N., 114° 25'W.) on 5 February 1966 by Mr. Gale Sphon.
Description. — The largest specimen obtained from all collections measured, in
life, 21 mm long and 11 mm broad. Average individuals were smaller: 14.5 mm long
and 7.5 mm broad.
I he body is typically acanthodoridiform; subovate, broader in front than behind
(Fig. 7). The mid-dorsal region is somewhat arched and bordered all around by a wide,
flat notal brim. A moderate number of upstanding, slender papillae, more or less equidis-
tant from each other, cover the notum except around the margin. Although taller and
shorter ones are intermingled, the latter occur more abundantly toward the margins. In
the color plate, the papillae are highly contracted.
I lie color pattern immediately distinguishes A. stobleri from all other described
thodoricK. In gross dorsal aspect, the notum is black with a wide, cherry-red mar-
not, il papillae, rhinophore sheaths and rhinophores are of a similar but more
use shade of red. In detail the rhinophores are tipped distally with a highly contrast-
^hite pigment that also occurs sparingly as flecks on some of the notal
lae, branchiae, oral veil, at the junction of the foot and hyponotum, and on the
s mostly colorless except for a few longitudinal streaks of black
most specimens the base of each papilla is surrounded by a colorless
variation in intensity and relative abundance of the pigments were
1968
Lance: New Panamic Nudibranchs
Fig. 7
Figure 7. Acanthodoris stohlcri. Dorsal aspect of animal in life.
observed but the over-all pattern and basic colors were constant. Specimens preserved
one year in 70 per cent isopropyl alcohol retained most of the black and red pigments.
In ventral aspects (Fig. 8) the foot is about two-thirds the width of the notum
and protrudes posteriorly as a rounded, flat tail. Its anterior margin is weakly bilabiate
with the upper lip barely discernible (Fig. 8c). The margins are entire, not notched.
The broad head region is produced into a semicircular veil with unilabiate margins and
is very slightly wider than the greatest width of the foot (Fig. 8b). The anterolateral
corners are expanded into blunt triangles.
As in other members of the genus, the retractile rhinophores are elongate and
deeply perfoliate. In life they are directed forward and outward. About two-thirds of
the length of each rhinophore is occupied by the conspicuous club, which bears 18 to 22
transverse lamellae joined anteriorly by a longitudinal septum. The rhinophore stalks
are slightly inflated at their bases and the entire organs are capable of being retracted
within upstanding sheaths which usually bear 1 to 4 elongate papillae similar in shape
and length to those on the notum. Each rhinophore terminates distally in a knob free
of lamellae.
The 6 to 7 bipinnate branchiae are separate at their bases and are contractile, but
not retractile, within a subnotal chamber. They decrease in size posteriorly. Conspicuous
white globular glands occur within the bases of the primary branchial stems. Unlike
other acanthodorids for which data are available, the present species does not bear notal
papillae within the area circumscribed by the branchial circlet. The anal opening occurs
atop a conspicuous excretory papilla within the branchial arc. Under ten power magnifi-
cation, rhythmically beating cillia may be observed on the inner distal face of the
anal tube.
10
San Diego Society of Natural History
Vol. 15
Fig. 8
0.05mm
Fig 10
Fig. 11
1968 Lance: New Panamic Nudibranchs 1]
The labial disc is characteristic in shape, cuticular and mostly transparent (Fig. 9a).
In a 14 mm specimen it measured 1.5 mm long. The margin joining the two halves is
serrate. The upper portion of each component gives rise to a triangular area supported
along its median margin by an elongated chitinous rod (Figs. 9a, b). The triangles are
composed of rows of minute, horn-colored elements with bluntly denticulate distal
margins (Fig. 9c). Each element is attached to the basement membrane by an imbedded,
thickened base.
The radula is narrow, colorless, and 2 mm long in in average live specimen of
14 mm. The formula is 34 x 4. 1.0. 1. 4. The denticulation is similar to that of other
acanthodorids: the first lateral is a large, erect hamate hook arising from a thickened
base and bears 2 to 3 inconspicuous denticles about half way along the cusp (Fig. 10a).
The four marginals (Fig. 10b) are wedge-shaped, greatly reduced, and decrease in
size outwards.
The spicules, although few in number, are relatively large, attaining a length of
1.0 mm in a 14 mm live specimen. They are simple, thick rods with slightly nodulose
surfaces and tapering ends and occur singly, not forming bundles or a reticulum
(Fig. 11a, b). The greatest concentration is radially oriented around the notal margins,
but they do not attain the edge thus leaving the brim free (Fig. 8e). Similar but smaller
spicules are found scattered in the mid-dorsal integument and rhinophore bases.
The penis bears a series of spines, but details of the armature and other aspects of
the reproductive system have not been determined.
Several individuals of the large collection made at Punta Diggs on 12 February
1967, and transported to Scripps Institution of Oceanography, deposited nidosomes (egg
masses) on the aquarium walls within a few days after capture. Apparently the animals
were undergoing a spawning cycle since many individuals were in a coital position
when collected. Nidosomes, presumably belonging to this species, were common under
or near stones harboring the nudibranchs. No evidence of reproductive activity of speci-
mens collected during other seasons has been observed either in the field or in the
aquarium.
The nidosome, as in many doridiform nudibranchs assumes the shape of a white
ribbon, loosely coiled counterclockwise, attached by one edge to the substratum. The
attached and free margins are of equal length resulting in erect, vertical sides. In
general shape it resembles that of AcanthoJoris nanaimoensis as figured by Hurst
(1967: 2 57) except, curiously, her illustration depicts a dextrally-oriented spiral.
Of the nine nidosomes deposited, the simplest consisted of a single convolution, the
largest of approximately two whorls. The pure white egg capsules contained a single
egg each and were deposited in vertical rows within the colorless, jelly-like matrix. A
typical nidosome measured 9 mm at its greatest diameter and 2.8 mm in height. No
attempt was made to observe embryological development.
Distribution. — This species has been collected only within a very restricted geo-
graphical area along the northeastern shores of the Baja peninsula: (1) Bahia Percebu
(about 15 miles south of San Felipe), 12 February 1965, 9 specimens; 26 November
1965, 1 specimen; (2) Punta Diggs, 12 February 1967, 37 specimens (egg masses
present); (3) Puertecitos, 26 November 1964, 3 specimens, 19 March 1965, 7 speci-
Figures 8-11. AcanthoJoris stohlcri. 8. Anteroventral aspect of animal in life:
a. mouth; b. oral veil; c. anterior foot margin; d. pigment streak; e. spicule (imbedded
in hyponotum) ; f. foot; g. hyponotum; h. dorsal papilla. 9. Labial armature and ele-
ments: a. lateral aspect of the paired membranous plates; b. frontal view of primary
supporting ribs and attached triangular basis anchoring labial elements; c. detail of
elements. 10. Three half-rows from middle of radula: a. denticulate first lateral; b. the
four successive marginals. 11. Spiculation: a. spicules from notal margain; b. detail of
spicule surface.
12
San Diego Society of Natural History Vol. 15
mens (egg masses present); (4) Bahia San Luis Gonzaga (type locality), 5 February
1966, 3 specimens. All animals occurred under stones in the lower intertidal region.
Nunn-, — The specific name stohleri was chosen to honor Dr. Rudolf Stohler of
the University of California at Berkeley, in recognition of his indefatigable efforts in
developing the journal "The Yeliger" into one of the major international publications
in malacology.
Discussion. — Acanthodoris Gray, 1850, is distinguished from the related suctorian
phanerobranch genera Akiodoris Bergh, 1879, Adalaria Bergh, 1878, and Onchidoris
Blainville, 18 16, by the presence of numerous soft notal papillae, a body broader in
front than behind, elongate rhinophores, upstanding papillated rhinophore sheaths, and
a characteristic radula. Only the first two genera bear an armed penis.
In addition to its unique color pattern, Acanthodoris stohleri may be distinguished
from other members of the genus by the absence of notal papillae within the branchial
circlet, and the presence, on the rhinophore sheath margins, of papillae similar to those
on the notum.
Acanthodoris stohleri is the first named subtropical representative of this common
temperate and subarctic genus.
Acknowledgments
I am grateful to the San Diego Natural History Museum and the Santa Barbara Museum of Natural
History for the many courtesies extended to me, and to Faye B. Howard, Gale G. Sphon, Fay Henry
Wolfson, Allan J. Sloan, and Helen Dushane for providing me with specimens and distributional data.
I would especially like to thank Wesley M. Farmer for his data on the new species of Nembrotha. The
San Diego Society of Natural History kindly provided me with facilities at the Vermilion Sea Field Station
and funds for the color plate, so expertly executed by Dr. and Mrs. G Dallas Hanna and Maurice Giles
of the California Academy of Sciences.
SUMARIO
Las recientes colecciones de moluscos Opistobranquios obtenidas en las zonas de
mareas del Golfo de California, comprenden varios Nudibranquios nuevos. Hasta la
fecha, se habian observado en las zonas calidas del Pacifico oriental, solamente 29 especies
correspondientes a estos animates sin concha; de modo que estas colecciones indican, que
dichas poblaciones son mas abundantes y diversas, por lo menos en la parte septentrional
de la provincia Panamica. En el presente trabajo se describen tres especies nuevas de
Nudibranquios.
1. Chromodoris fayeae sp. nov. Dorido grande, bianco, con bordes de tonalidades
brillantes en rojo y amarillo. Se han observado estos animales en muchas localidades
del Golfo de California. Se trata del tercer Chromodorido descripto, que es endemico
en la region Panamica.
2. Nembrotha hubbsi sp. nov. Animales de color brillante, ocre amarillento con listas
azules. Su distribucion abarca por lo menos las regiones septentrionales y meriodionales
del Golfo, Esta especie viene a ser la primera que representa este genero tropical de
vivos colores, en el Pacifico oriental.
3. Acanthodoris stohleri sp. nov. Dorido pequeno, negruzco, con el borde del dorso de
color rojo. Se encontro solamente en una zona restringida (entre Bahia Percebu y
Bahia San Luis Gonzaga) en las costas nordeste del Golfo. Se trata del primer repre-
subtropical de un genero que es comun en las zonas templadas y subarticas.
Literature Cited
Bada, k.
thobranchia of Sagami Bay: Supplement. Iwanami Shoten, Tokyo. 194 p; 20 pis.; 5 6
text figs.
R.
he Untersuchungcn; //; C. Semper: Reisen im Archipel der Philippincn 11: 429-
pls. 54-57.
1968 Lance: New Panamic Nudibranchs 13
1878. Ibid., 14: 603-645; pis. 66-68.
1879. Gattung nordischer Doriden. Arch. Naturgcsch., 45(1): 3 54.
1879. On the nudibranchiate gasteropod Mollusca of the North Pacific ocean, with special reference
to those of Alaska. Proc. Acad. Nat. Sci. Philadelphia 31: 71-132; pis. 1-8.
1880. On the nudibranchiate gasteropod Mollusca . . . Part II. Proc. Acad. Nat. Sci. Philadelphia
32: 40-127; pis. 9-16.
1905. Die Opisthobranchiata. Siboga Reports 50: 1-248; pis. 1-20.
Blainville, H. M. D.
18 16. Prodome d'une nouvelle distribution systematique du regne animal. Bull. Soc. Philom., Paris
(unpaginated) .
Burn, R.
1962. Descriptions of Victorian nudibranchiate Mollusca, with a comprehensive review of the
Eolidacea. Mem. Nat. Mus. Melbourne No. 25: 95-128; figs. 1-24.
Cooper, J. G.
1862. Some genera and species of California Mollusca. Proc. Calif. Acad. Nat. Sci., 2: 202-207.
Eales, B.
193 8. A systematic and anatomical account of the Opisthobranchia. John Murray Exped. Sci. Rep.,
5 (4): 77-122; figs. 1-28; pi. 1.
Ehrenberg, C. G.
1831. Symbolae Physicae animalia evertebrata exclusis Insectis. Berlin. (Not seen).
Eliot, C. E.
1904. On some nudibranchs from East Africa and Zanzibar. Pt. 4. Proc. Zool. Soc. London 1(2):
380-406; pis. 23-24.
Farmer, W. M.
1963. Two new opisthobranch mollusks from Baja California. Trans. San Diego Soc. Nat. Hist.,
13(6): 81-84; fig. 1; pi. 1.
Gohar, H. A. F., and I. A. Aboul-Ela
1957. The development of three chromodorids. Publ. Mar. Bio. Sta. Al-Ghardaqa Red Sea. No.
9: 203-228; pis. 1-5.
Gray, J. E.
1850. Figures of molluscan animals, hi Guide to the systematic distribution of Mollusca in the
British Museum. Pt. 1. London, 18 57. (Not seen).
Hurst, A.
1967. The egg masses and veligers of thirty northeast Pacific opisthobranchs. Veliger 9 (3): 255-287;
figs. 1-31; pis. 26-38.
Keelart, F. E.
185 8. Description of a new Ceylonese nudibranch. Ann. Mag. Nat. Hist., 3 (1): 2 57.
Keen, A. M.
1958. Sea shells of tropical west America. Stanford Univ. Press, Stanford, Calif, xi + 624 pp.; illus.
Macnae, W.
1957. The families Polyceridae and Goniodorididae (Mollusca, Nudibranchiata) in southern Africa.
Trans. Roy. Soc. South Africa 35 (4): 341-372; pis. 17-18.
Marcus, Ernst
195 6. Notes on Opisthobranchia. Bol. Inst. Oceanografico 7 (1-2) 31-79; pis. 1-8 (published 1958).
1965. Some Opisthobranchia from Micronesia. Malacologia 3 (2): 263-286; figs. 1-43.
Marcus, Ernst, and J. B. Burch
1965. Marine euthyneuran Gastropoda from Eniwetok atoll, western Pacific. Malacologia 3 (2):
235-262; figs. 1-43.
Marcus, Ernst, and Eveline Marcus
1960. Opisthobranchia aus dem Rotcn Meer and von den Malediven. Akad. Wiss. Lit. Main/,
Math. Natur. Kl., 1959 (12): 871-934; figs. 1-86.
Odhner, N. H. J.
193 9. Opisthobranchiate Mollusca from the western and northern coast of Norway. Kg. Norske
Vidensk. Selsk. Skr., No. 1: 1-93; figs. 1-5 9.
1957. ChromoJoris contra Glossodorh. A systematic controversy. Proc. Malacol. Soc. London 32 (6):
250-253.
Stimpson, W.
185 5. Descriptions of some of the new marine Invertebr.ua from the Chinese and Japanese Seas.
Proc. Acad. Nat. Sci. Philadelphia 7: 378-389.
Manuscript Received: 22 June 1967
JlJn 1 8 me
THERMAL RELATIONS, DISTRIBUTION, AND
HABITAT OF CNEMIDOPHORUS LAB1ALIS
(SAURIA: TEIIDAE)
DENNIS L. BOSTIC
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 3, 5 JUNE 1968
THERMAL RELATIONS, DISTRIBUTION, AND
HABITAT OF CNEMIDOPHOKUS LABIALIS
(SAURIA: TEIIDAE)
Dennis L. Bostic
Abstract. Cnemidophorus labialis, a lizard endemic to Baja California, Mexico, was studied in 1966
and 1967. This species inhabits a narrow coastal strip of Sonoran Desert from Punta San Jose south
to Miller's landing, an area subject to frequent northwest winds, fog, predominantly winter rainfall,
and summer temperatures similar to those of San Diego, California. It is usually encountered in
areas of sparse but varied vegetation and well-drained soil. Its range may be restricted by the oc-
currence of C. hyperythrus. The normal activity range (NAR) of C. labialis is 30- 42C. Its mean
body temperature is 37. 2C, nearly 1C lower than any previously reported for Cnemidophorus.
The small size and low body temperature of this lizard may reflect thermal adaptation to its coastal
habitat. There were no statistically significant differences between the MBTs of males and females,
or of adults and immatures; nor were their important seasonal or daily variations. Body tempera-
tures were more closely correlated with substrate than air temperatures. Diurnal activity began
between 0800 and 1000 and at soil temperatures of 32C; peak activity occurred between 1000
and 1300. Completely overcast skies, strong steady winds, and prolonged precipitation interrupted
activity. Adults enter hibernation from late July through September, immatures in December.
Active individuals were first observed in March.
Resumen. Cnemidophorus labialis, lagarto endemico de Baja California, Mexico, ha sido estudiado
durante 1966 y 1967. Esta especie habita una estrecha banda costera del desierto de Sonora,
desde Punta San Jose en el Sur, hasta el desembarco de Miller (Miller's Landing) ; en una zona
afectada por vientos frecuentes del Noroeste, nieblas, lluvias invernales dominantes, y temperaturas
estivales similares a las de San Diego, California. Estos animales se encuentran usualmente en re-
giones de escasa, pero variada vegetacion y en terrenos con buen drenaje. Su distribucion esta posi-
blemente restringida por la presencia de C. hypcrythrus. La amplitud normal de actividad (NAR)
de C. labialis abarca de los 30°C a los 40°C. La temperatura media del cuerpo es de 37.2 C, casi
un grado menos que las observadas previamente para Cnemidophorus. La talla pequena de estc lagarto
y la temperatura baja del cuerpo podi.in indicar una adaptacion termica a su habitat costero. No-
se presentan diferencias de significado estadistico, en la temperatura media del cuerpo (MBT) de
machos y hembras, o entre individuos adultos y jovenes; ni tampoco variaciones importantes, ya sea
estacionales o diarias. La temperatura del cuerpo podria relacionarse mas directamente con la tem-
peratura del substrato que con la del aire. La actividad diurna se inicia entre las 0800 y 1000 horas,
cuando la temperatura del suelo es de 32CC; y la maxima actividad entre las 1000 a las 13000 horas.
La actividad de estos animales se interrumpe cuando el cielo esta cubierto, y con vientos y precipi-
taciones fuertes y persistentes. Los adultos entran en hibernacion de finales de Julio a Septiembre, y
los jovenes en Diciembre. En Marzo empiezan a observarse individuos en actividad.
Cowles and Bogert (1944) and Bogert (1949) pointed out that diurnal species of
lizards maintain body temperatures at characteristic levels relatively independent of air
and substratum temperatures primarily by behavioral means, and that the ability to
maintain these temperatures is essential for survival. They demonstrated that species in
the same genus have similar body temperatures even though they exist in markedly dif-
ferent environments and, conversely, that lizards of different genera maintain different
temperature levels although they occur within the same habitat. Interpretations of these
San Diego Soc. Nat. Hist., Trans., 15 (3): 21-30, J Jim 9
22 San Diego Society of Natural History Vol. IS
species differences, and temperature responses of reptiles in general, have been reviewed
from historical, behavioral, ecological and physiological standpoints by Schmidt-Nielsen
and Dawson (1964).
The present study was undertaken to determine the thermal relations of Cnemido-
phorus labialis (Baja California Striped Whiptail). C. labialis, endemic to Baja Califor-
nia, Mexico, is the only member of the sexlineatus species group inhabiting the peninsula,
and because it inhabits a relatively inaccessible strip of coastline it is the least known
member of the group. During this study additional data on the distribution and habitat
of C. labialis were gathered.
Methods
During the spring, summer, and fall of 1966 and 1967, the body temperatures
(BT's) of HO C. labialis were recorded in the field, and under a variety of ambient
conditions, throughout much of the species' known range in Baja California Del Norte,
Mexico. Individuals were collected by shooting them with six-inch rubber bands. A
Schultheiss rapid equilibrium thermometer was used to record BT's. The thermometer
was inserted at least one centimeter into the lizard's cloaca within five seconds after
capture, and was read (to the nearest 0.2C) when the mercury column reached equi-
librium. When recording BT's, I wore a cloth glove to minimize heat conduction to or
from my hand. BT's were not taken for lizards which had been chased, showed evidence
of internal bleeding, were mutilated, or showed abnormal behavioral patterns due to the
observer's presence.
Other temperatures usually recorded at the site of capture were air temperature
(4- 5 mm above substratum) and substral temperature (3mm beneath the surface). The
bulb of the thermometer was shaded for air temperature readings. Soil temperature
above SOC was recorded as SOC +. Additional data frequently recorded for each capture
included time (Standard), per cent cloud cover, wind velocity (Beaufort wind scale),
sex, snout-vent length (SVL), and behavior of lizard prior to capture (e.g. basking,
foraging).
The more important terms as summarized by Brattstrom (1965) and used
throughout this paper are: voluntary minimum (theoretically the lowest body tempera-
ture voluntarily tolerated; in practice, the lowest body temperature recorded of an
active individual); voluntary maximum (theoretically the highest body temperature
voluntarily tolerated; in practice, the highest recorded body temperature of an active
individual) ; normal activity range (the range of body temperatures from the voluntary
minimum to the voluntary maximum) ; bashing range (the range of body temperatures
within the normal activity range for basking individuals); mean body temperature
(the mean of body temperatures within the normal activity range).
The term "significant" is used throughout to denote a statistical difference at the
95 'y confidence level. All mean body temperatures (MBT's) have been rounded off to
the nearest 0.1C.
Distribution
Based on data furnished by Belding, collector of the type series of C. labialis,
Stcjcgner (1890) recorded the type locality as Cerros (=Cedros) Island. However,
C. labialis has not subsequently been collected on Cedros, despite several visits by her-
petologists; and Savage (1954) suggests that the type specimens were collected at
Bahia San Quintin on the peninsular mainland, where Belding also collected. If Savage
Ls correct, the only published localities for C. labialis are Arroyo Seco (Bostic, 1966a),
Bahia San Quintin (Burt, 1931; Tevis, 1944; Walker, 1966; Lowe, Wright and Norris,
1968
Bostic: Cnemidophorus labialis
23
"
,J
tNSENADA
ARROYO SANTO TOMAS
SANTO TOMAS
PUNTA DE SAN JOSE
SAN VICENTE
ERENDIRA
ARROYO SECO
8 PUNTA COLNETT
9 CAMALU
SAN QUINTAN
EL SOCORRO
EL CONSUELO
EL ROSARIO
MILLER'S LANDING
VIZCAINO DESERT
BAJA CALIFORNIA
\— I t-H 1=3
100
IhhM -l
l - ■ . I . . ,
Figure 1. The recorded distribution of Cnemidophorus labialis in Baja California,
Mexico. Collecting sites and general localities are indicated by X and O, respectively.
24 San Diego Society of Natural History Vol. 15
1966), Socorro (Murray, 1955; Lowe et al., 1966; El Consuelo (Walker, 1966) and
Miller's Landing (Tevis, 1944; Duellman and Zweifel, 1962; Lowe et al., 1966) (Fig. 1).
I have also collected specimens at Punta San Jose, Erendira, Punta Colnett, Camulu,
and Socorro (Fig. 1).
All collecting localities, excluding Miller's Landing and El Consuelo, fall within
the vegetative parameter Shreve (1936) termed the Chaparral — Sonoran ecotone (the
region between Arroyo Santo Tomas and 20 miles north of El Rosario). If the ecotone
is not delimited, this area (based on the dominant forms and composition of the flora
and fauna) is Sonoran Desert (Short and Crossin, 1967). Miller's Landing and El Con-
suelo occur within the Central Desert phytogeographic area (Wiggins, 1960), and
Shreve's and Wiggins' (1964) rrAgave-Franseria" Region. The dominant coastal vege-
tation, several species of Agave, Franseria chenopodifolia, Yucca valida, Opuntia cholla
and Dudleya spp., is sparse and widely spaced (Wiggins, 1960).
Punta San Jose, approximately seven miles south of the Arroyo Santo Tomas, is
the northernmost recorded locality for C. labialis. This general area probably represents
the true northern limit for the species. Farther north, the coastal terrain is rugged and
mountainous, in direct contrast to the flat coastal terraces and plains to the south
that are inhabited by whiptails.
Arroyo Seco, (approximately ten miles from the coast) is the most inland capture
locality for C. labialis. This area is on the fringe of the coastal cloud cover and fog.
I suspect that the eastern and southern distributional limits of C. labialis probably con-
form to coastal areas subject to much fog, frequent northwest winds, predominately
winter (November to April) rainfall distribution, and a summer temperature similar
to that of San Diego, California. These environmental conditions, excluding precipita-
tion distribution, prevail along the west coast of Baja California as far south as the
Magdalena Plains (Aschmann, 19 59).
I consider the environmental conditions that occur throughout the recorded range
of C. labialis suboptimal for C. hypcrythrus, the only other small Cnemidophorus with
which it is sympatric. C. hyperythrus is most frequently observed in the coastal foothill
region, where the duration and intensity of solar radiation is greater, and the frequency
and duration of coastal cloud cover and fog less. In the narrow belt of sympatry these
species are regularly observed, but in disproportionate numbers. For example, at Camalu
only five of the 150 Cnemidophorus collected were C. hypcrythrus. At Arroyo Seco, the
habitat preferences of the two species differed. C. labialis occupied only the sandy fringe
of the coastal sage-scrub community preferred by C. hypcrythrus, and was most com-
monly collected in the bordering, sandy-bottomed arroyo where C. hypcrythrus was
absent. C. hyperythrus was present in the surrounding areas of Agave and gravelly soil,
but C. labialis was absent.
The southernmost capture locality, Miller's Landing, is 18 5 mies south of El Con-
suelo. Duellman and Zweifel (1962) stated that this distribution gap is undoubtedly
related to the course of the main road which turns inland at Socorro ( 1 1 miles N. of
El Consuelo) and does not closely approach the coast to the south until Miller's Land-
ing. The southernmost known locality of C. labialis coincides with the northern limit
of the Vizcaino Desert (Fig. 1). Here, through the Magdalena Plains region, coastal
precipitation is not predictable, and is primarily a summer (May to October) phe-
nomenon (Aschmann, 1959). The presence of C. hypcrythrus in this area may also affect
the southward spread of C. labialis, but this is still unstudied.
Habitat
Sand dunes (Tevis, 1944), a sandy plain with scattered Agave (Murray, 1955),
and a humid coastal prairie (Walker, 1966) have been recorded as habitats for
1968 Bostic: Cnemidophorus labialis 2 5
C. labialis. During 1963-64, I found C. labialis abundant at Arroyo Seco (a wide sandy-
bottomed river bed) and in the bordering coastal sage-scrub community. Following the
torrential rains of 196 5 and the concomitant flooding of the arroyo, the population was
considerably reduced. The dominant plants of the river bed were Pluchea sericea, Juncus
sp., and Franseria chenopodifolia. Less conspicuous plants included Eriogonum spp.,
Isomeris arborca and Mesembrianthemum chrystalinum. C. labialis was frequently ob-
served in or near Mesembrianthemum at most collecting localities. In sandy soils of the
coastal areas Mesembrianthemum appeared to be perennial.
C. labialis was present, though scarce, on gravelly and sandy substrata at the
coastal terrace sites of Punta de San Jose, Erendira, Punta Colnett, and El Socorro
(Fig. 1). The dominant vegetation at these localities included Agave shauii and
Euphorbia misera. Also present but less conspicuous were Rosa minutiflora, Atriplex sp.,
Opuntia sp., Machaerocercus gummosus, Myrtillocactus cochal, Dudleya spp., Mesem-
brianthemum, and numerous annuals.
At Camulu, an agricultural community on the Llano de Hamulu, whiptails arc
unusually abundant. This nearly flat plain borders the Pacific for approximately 30
miles from near San Telmo to Santo Domingo, and is delimited a few miles inland by
the foothills of the interior mountains. The soil is fine and compacted, but well drained.
Franseria chenopodifolia and Agave shauii are the dominant plants. Less conspicuous
were Euphorbia misera, Opuntia serpentina, O. echinocarpa, Mammilaria dioca, Ber-
gerocactus emoryi, Myrtillocactus cochal, Mesembrianthemum, and Atriplex sp.
C. labialis exhibited a clumped pattern of local distribution at all collecting sites.
This was particularly evident at Camalu and Arroyo Seco, where the majority of col-
lecting occurred. A similar pattern of distribution was reported by Tevis (1944) and
Walker (1966). It appeared that this pattern was correlated with the availability of
suitable egg laying sites (areas covered with a variety of annuals, with light sandy soil
exposed to direct solar radiation in the morning and afternoon), and adjacent shrub
cover. The vegetational cover was not dense nor unvaried (i.e., predominately Agave).
Dense cover does not satisfy their thermal requirements and homogeneous vegetation,
probably owing to the sparse leaf litter, does not offer a varied food supply. Such sites
are relatively sparse, and in the Camulu, Arroyo Seco, and Bahia San Quintin areas are
becoming scarcer because of agricultural activity.
Thermal Relations
Temperature Preferences. — The normal activity range (NAR) for C. labialis is
30C to 42C (Table 1). This NAR includes the low, often isolated BT's of lizards con-
sidered by Cowles and Bogert (1944) to be "basking" in order to resume "normal
activity." I consider the "basking range," unless quantitatively measured (basking
duration), to be arbitrary (Bostic, 1966b). C. labialis was frequently observed foraging
at temperatures throughout the NAR. Similarly, lizards which basked for short periods
had BTs between 30C - 39C.
The voluntary minimum temperature tolerated is 1-4C lower than those reported by
Fitch (1958), Carpenter (1961), McCoy (1965), and Bostic (1966b) for other
species of Cnemidophorus. These investigators included the low BT's of active in-
dividuals in their calculation of the NAR. The voluntary maximum of 42C is similar
to those values previously recorded for Cnemidophorus.
The previously recorded high and low mean body temperatures of Cnemidophorus
are 41. 3C for C. tigris (Bogert, 1949) and 38C for C. sexlineatus (Carpenter, 1961).
respectively. This variability, in part, reflects the omission of "basking range" BT's
from the NAR.
26
San Diego Society of Natural History
Vol. 15
TABLE 1
A Summary of Published Data on Temperature Relations in Cnemidophorus
Species
Group
Species Normal
Activity
Range (NAR)
X ±
NAR
SE
N
Critical
Thermal
Maximum
(CTM)
X N
CTM
Authority
Lemniscatus
C.
lemniscatus
34.6-42.3
39.0
31
45.2-46.6
46.2
Brattstrom, 1965
C.
lemniscatus
34.5-41.0
38.5
13
Brattstrom, 1965
Hyperythrus
c.
ceralbensis
37.4-42.5
40.35
23
Soule, 1963
c.
hyperythrus
beldingi
3 3.4-42.2
38. 5±
.14
138
Bostic, 1966b
c.
hyperythrus
hyperythrus
38.3-46.6
3 9.9±
.25
14
Soule, 1963
Tessellatus
c.
tigris
37.4-43.5
41. 3±
.24
33
Bogert, 1949
c.
tigris
3 5.0-44.6
40.4
47
Brattstrom, 1965
c.
tigris
46.0 1
Cole, 1943
c.
tigris
septentrional
34.0-43.0
tit
3 9.8
88
McCoy, 1965
Sexlineatm
c.
gularis
37.8-41.0
3 8.9
3
Brattstrom, 1965
c.
lahialis
30.0-42.0
37. 2±
.26
136
This paper
c.
sexlineatus
32.0-43.0
3 8.0
Carpenter, 1961
c.
sexlineatus
34.6-41.5
39.5
45
Fitch, 195 6
c.
sexlineatus
38.5-43.0
41. 0±
.47
Bogert, 1949
c.
sexlineatus
31.0-44.0
40.0-41
.0
100
Fitch, 195 8
The MBT of C. lahialis is 37. 2C (SE = 0.201; SD = 2.35), which is 0.8-1.3C
lower than the MBT's of other Cnemidophorus calculated in a similar manner (see
Carpenter, 1961; Bostic, 1966b). This low MBT may reflect thermal adaptation to the
fog and cloud shrouded coastal habitat of C. lahialis. The favorable volume to surface
ratio of this small lizard may also aid in thermal adaptation. Walker (1966) reported
"that only C. hyperythrus espiritensis (maximum SVL 59.5mm) of Isla Espiritu Santo,
Gulf of California, is of smaller size than C. labialh" However, I examined a male
C. hyperythrus espiritensis (SDSNH no. 528 52) which has a SVL of 64mm.
There was no significant difference between the MBT's of male and female
C. lahialis. The MBT for 77 males and 59 females was respectively 37. 2C (SE = 0.25 5;
SD^2.24) and 37.1C (SE = 0.268; SD = 2.06). Bogert (1949) found no intersexual
differences in the MBT's of C. sexlineatus or C. tigris and Bostic (1966b) found no
significant differences in the MBT's of male and female C. hyperythrus beldingi. Body
temperature variability among species of Cnemidophorus is apparently not due to dif-
ferences in sex.
The MBT of 14 juveniles (SVL < 40mm) is 36. 3C (SE = 0.363; SD = 1.36),
which is 0.9C lower than that of 37. 2C (SE = 0.201; SD = 2.35) recorded for adults.
This difference may be biologically important, although it is not statistically significant.
The ability of juveniles to function optimally at a lower MBT than adults would be
advantageous. Juveniles hatch in August and are active through December, when soil
and air temperatures are low; adults begin hibernating in July. The high correlation
~ 0.85) between juvenile BT's and soil temperature reported by Hardy (1962) for
C. sexlineatus suggests such an advantage. Concomitantly, competition between adults
1968
Bostic: Cnemidophorus labialis
27
a.
Q
O
42
41
40
39
38
37
36
35
34
33
32
31
30
29
0 0 o »
U 0 0 8
0 0 n 0 w 0
0 0 0 0 8o °0
0 0 00 0
S°o° °
w 0 0 u
g 0 0 0 0
0 „° 0 0
0 °^ 0
o 0U 2
0 8
000
0 °
0 0° o °/o
0
/
/
/
8"° 6V "°8 q 8 ° °° °
21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41
AIR TEMPERATURE C
Figure 2. Scatter diagram comparing body temperatures of 13 5 Cnemidophorus
labialis with air temperatures 4- 5 mm. above the substrate. The dashed lines connect the
isothermal points. Single individuals are represented by an open circle, and two lizards
by half-shaded circles.
and juveniles would be reduced throughout much of their activity seasons since all
adults have hibernated by late September. The low MBT would also promote fuller
utilization of shaded areas for foraging and protection.
There was no significant difference between morning and afternoon MBT's. The
morning series (N = 86) had a MBT of 37.4C (SE = 0.248; SD = 2.30), and the
afternoon series (n = 61) a MBT of 37. 3C (SE = 0.232; SD=1.81). I also found no
significant monthly differences in body temperatures. Bostic (1966b) reported no sea-
sonal acclimation in C. h. beldingi. Bostic (1966b) reported similar findings for C. h.
beldingi.
Response to Environmental Tern peratu res. — There is no significant correlation
between BT and air temperature (Fig. 2). Only nine lizards possessed BT's within IC
of the air temperature. Seven individuals were recorded with BT's 1-9C less than air
temperature, and two lizards maintained BT's 14. 8C above air temperature. The majoritv
of whiptails were active at air temperatures ranging from 26C to 32C. No lizards were
active at air temperatures above 41C nor below 2lC.
There is a closer correlation between BT and substrate temperature (Fig. 3). Below
a substrate temperature of 42C the majority of lizards possessed BT's 1 -4C above or
below the substrate temperature. One lizard maintained a BT 5.4C above that of the
substrate. The limit of SOC imposed by using the Schultheiss thermometer to record soil
temperatures makes any delineation of the greatest deviation between BT and substrate
temperature impossible.
28
San Diego Society of Natural History
Vol. 15
42
4 1
40
39
u
38
UJ
a:
3
»—
37
<
OL
UJ
a.
36
i
LU
t—
>■
35
o
o
CO
34
33
32
31
30
29
00
8
e o
o o
o
8 0„
0 0
0 °
°0 0 00
0
o 0o
0 0
0 0
n 0 00
0 0 0
0
0
e
o e
0
0
°o o
0
31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50*
SUBSTRATE TEMPERATURE C
Figure 3. Scatter diagram comparing body temperatures of 134 Cnemidophorus
labialis with substrate temperatures. The dashed lines connect the isothermal points.
Single individuals are represented by an open circle, and two lizards by half-shaded
circles.
Daily Activity Cycle. — C. labialis. commenced activity between 0800 and 1000 at
a soil tmperature of about 32C. This initial activity usually coincided with the dissi-
pation of the early morning fog or coastal cloud cover. Milstead (1957), McCoy
(1965), Bostic (1966b) and Echternacht (1967) recorded similar emergence times and
soil temperatures for other species of Cnemidophorus. The major activity peak for
C. labialis was between 1000 and 1300. This usually coincided with the period between
early morning and late afternoon low clouds, wind, and high (5 0C +) substrate tem-
peratures. No late afternoon activity peak recorded for other species of Cnemidophorus
by Milstead (1957), Carpenter (1961), Bostic (1966b), and Echternacht (1967) was
observed for C. labialis; activity virtually ceased at 1500.
As would be expected, the increase in ambient temperatures from spring through
summer that occurred in inland areas of the peninsula was not as pronounced along the
coast. Subsequently, no obvious seasonal fluctuation in the daily cycle was evident.
The usual daily cycle of C. labialis was altered by the varying meteorlogical con-
ditions. Partial cloudiness did not interrupt activity. No lizards, however, were ob-
served on those days that the sun did not shine intermittently. Winds up to seven mph
did not seem to adversely affect whiptail activity, and intermittent winds of twelve mph
had little affect. Steady winds above 12 mph resulted in the disappearance of most
lizards. Whiptails were not active immediately after a prolonged rain, and generally did
not resume activity until most of the moisture content in the subsoil had disappeared
following one to three days of sun.
1968 Bostic: Cnemidophorus lahialh 29
Seasonal Activity Cycle. — C. labialis was recorded emerging from hibernation i:i
mid-March. Adult whiptails enter into dormancy in the latter part of July, and by late
September nearly all adult whiptails have hibernated. All immature whiptails disappeared
in late December. A similar pattern of seasonal activity has been reported for
C. h. beldingi (Bostic, 1966b), and C. sexlineatus (Hardy, 1962).
C. labialis collected in March and early April had very small, paired, dull yellow
coelomic fat bodies. As the season progressed the coelomic fat bodies became greatly
enlarged, and were bright yellow in color. Apparently the differential times of dor-
mancy recorded for whiptails is correlated with the amount of stored fat. A similar
pattern of fat storage and utilization was recorded for C. sexlineatus (Hardy, 1962;
McCoy, 1965) and C. byperythrus (Bostic, 1966b).
Acknowledgments
I express my appreciation to Mr. Allan J. Sloan and Mr. Wade F. Snyder for their editorial com-
ments and criticisms of the manuscript at various stages of its completion. I am particularly indebted
to Mr. William Hite for his frequent assistance in the field and laboratory. In addition, I thank Mr. Sam
Angcve for his field assistance, Mrs. Aileen Scallet for assistance with the statistical analyses, Mr. Gilbert
Voss for assistance in identifying plants, and Mrs. Norrine G. Gearheart for typing the manuscript.
The specimens collected during this study are deposited in the San Diego Natural History Museum.
This study, in part, was made possible by a Sigma Xi Grant-in-Aid of Research to study the
testicular histology of C. labialis.
Literature Cited
ASCHMANN, N. H.
195 9. The central desert of Baja California: Demography and ecology. Univ. Calif. Press, Berkeley
and Los Angeles. 315 p.
Bogert, C. M.
1949. Thermoregulation in reptiles, a factor in evolution. Evolution 3: 195-211.
Bostic, D. L.
1966a. Threat behavior in Cnemidophorus byperythrus and Cnemidophorus labialis. Herpetolcgica
22: 77-79.
1966b. Thermoregulation and hibernation of the lizard, Cnemidophorus byperythrus beldingi (Sauria:
Teiidae). Southwestern Natur. 11: 275-289.
Brattstrom, B. H.
1965. Body temperatures of reptiles. Amer. Mid. Natur. 73: 376-422.
Burt, C. E.
1931. A study of the teiid lizards of the genus Cnemidophorus with special reference to their
phylogenetic relationships. U.S. Nat. Mus. Bull. 154. 286 p.
Carpenter, C. C.
1961. Temperature relationships of two Oklahoma lizards. Okla. Acad. Sci. Proc. 41: 72-77.
Cole, L. C.
1943. Experiment on toleration of high temperature in lizards with reference to adaptive coloration
Ecology 24: 94-108.
Cowles, R. B., and C. M. Bogert
1944. A preliminary study of the thermal requirements of desert reptiles. Amer. Mus. Natur. Hist.
Bull. 83: 261-296.
Duellman, W. E., and R. G. Zweifel
1962. A synopsis of lizards of the sexlineatus group (genus Cnemidophorus). Amer. Mus. Nat.
Hist. Bull. 123: 155-210.
ECHTERNACHT, A. C.
1967. Ecological relationships of two species of the lizard genus Cnemidophorus in the Santa Rita
Mountains of Arizona. Amer. Mid. Natur. 78: 448-45 9.
Fitch, H. S.
195 6. Temperature responses in free-living amphibians and reptiles of northeastern Kansas. L niv.
Kan. Pub!. Mus. Nat. Hist. 8: 417-476.
1958. Natural history of the six-lined racerunner (Cnemidophorus sexlineatus). Univ. Kan. Publ.
Mus. Nat. Hist. 11 (2): 11-62.
30 San Diego Society of Natural History Vol. 15
Hardy, D. F.
1962. Ecology and behavior of the six-lined racerunners, Cnemidopborus sexlineatus. Univ. Kan.
Sci. Bull. 43: 3-73.
Lowe, C. H., J. W. Wright, and K. S. Norris
1966. Analysis of the herpetofauna of Baja California, Mexico. IV. The Baja California striped
whiptail, Cnemidopborus labialis, with key to the striped — unspotted whiptails of the south-
west. Ariz. Acad. Sci. J. 4: 121-127.
McCoy, C. J.
1965. Life history and ecology of Cnemidopborus tigris septentrionalis. Unpub. Ph.D. thesis, Univ.
microfilms Inc., Ann Arbor, Michigan. 66-2814. 167 p.
Milstead, W. W.
1957. Some aspects of competition in natural populations of whiptail lizards (genus Cnemidopborus) .
Texas J. Sci. 9: 410-447.
Murray, K. F.
195 5. Herpetological collections from Baja California. Herpetologica 11: 33-48.
Savage, J. M.
1954. Notulae herpetologicae 1-7. Kans. Acad. Sci., Trans. 57: 326-334.
Schmidt-Nielsen, K., and W. R. Dawson
1964. Terrestrial animals in dry heat: desert reptiles, p. 467-480. /;; D. B. Dill (ed.). Handbook
of Physiology, Vol. 4, Adaptation to the environment. Washington, D.C. Amer. Physiol. Soc.
Short, L. L., Jr., and R. Crossin
1967. Notes on the avifauna of northwestern Baja California. San Diego Soc. Nat. Hist., Trans.
14(20): 281-300.
Shreve, F.
1936. The transition from desert to chaparral in Baja California. Madrono 3: 257-264.
Shrfve, F., and I. L. Wiggins
1964. Vegetation and flora of the Sonoran desert. Stanford University Press, 1: 840 p.
Soule, M.
1963. Aspects of thermoregulation in nine species of lizards from Baja California. Copeia (1):
107-115.
Stejneger, L.
1890. Description of a new lizard from Lower California. U.S. Natl. Mus., Proc, 12: 643-644.
Te\is, L., Jr.
1944. Herpetological notes from Lower California. Copeia (1): 6-18.
Walker, J. M.
1966. Morphology, habitat, and behavior of the teiid lizard, Cnemidopborus labialis. Copeia (4):
644-650.
Wiggins, I. L.
1960. The origin and relationships of the land flora, p. 148-165. /;; symposium: The biogeography
of Baja California and adjacent seas. Syst. Zool. 9 (3-4).
Manuscript Received 10 January 1968
Life Sciences Department, Palomar College, San Marcos, California.
^
MUS. COMP. ZOOL
LIBRARY
JUN 1 8 1968
HARVARD
UNIVERSITY.
CASTELA POLYANDRA, A NEW SPECIES
IN A NEW SECTION; UNION OF HOLACANTHA
WITH CASTELA (SIMAROUBACEAE)
REID MORAN AND RICHARD FELGER
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 4, 5 JUNE 1968
113°
^de-Jos
Vtngeles
<P
28°
n , . . , Santa Rosalia+v
Castela polyandra \ Jsla San
» Castela emoryi
(closest known locality)
fc
27°
Marcos
Mulege-p
^X
Figure 1. Distribution of Castela polyandra; the closest (and southernmost) known
locality for C. emoryi is also shown.
CASTELA POLYANDRA, A NEW SPECIES
IN A NEW SECTION; UNION OF HOLACANTHA
WITH CASTELA (SIMAROUBACEAE)
Reid Moran and Richard Felger
Abstract. Castela polyandra is a new species of shrub of Isla Tiburon, and of east central
Baja California, Mexico. It resembles Holacantha emoryi A. Gray in its nearly leafless determinate
spinose branches without spur shoots and in its petals, larger in staminate than in pistillate flowers;
it differs from Holacantha and resembles Castela sensu stricto in its 4 (-5)-merous flowers and
its deciduous fruits; and it differs from both in having 2-3 times as many stamens as sepals. Since
Holacantha now seems less distinct than before, it is made a section of Castela. The section
Eremacantha is proposed for C. polyandra.
Resumen. Castela polyandra es una nueva especie de arbusto de la Isla Tiburon y de la parte
centro-oriental de Baja California, Mexico. Se parece a Holacantha emoryi A. Gray en tener ramas
espinosas casi afilas, sin braquiblastos, y en sus petalos, los cuales son mas grandes en las flores
estaminiferas que en las pistiladas. Se diferencia de Holacantha y se parece a Castela sensu stricto,
en que sus flores son tetra- o pentameras y sus frutos caducos; diferenciandose de ambas por tener
doble o triple numero de estambres que de petalos. Como Holacantha aparece ahora menos diferenciada,
se la incluye en una seccion de Castela; y se propone la seccion Eremacantha par C. polyandra.
For 37 years botanical collectors in Baja California have been puzzled by a hola-
canthoid shrub, first found by Professors Ira L. Wiggins and James McMurphy, on the
Gulf coastal plain south of Santa Rosalia. Early collectors obtained only sterile plants.
In May 1961 Moran got some insect-damaged staminate flowers there; and in August
1964, he found abundant fruiting material at Bahia de los Angeles, some HO miles to
the north-northwest. In late April of 1966 he found both staminate and pistillate flowers
at the northern locality. Meanwhile, between 1963 and 196 5 Felger made four collec-
tions of it, one with flowers, on the east side of Isla Tiburon, Sonora. The accumulated
material permits the description of the following new species.
Castela polyandra Moran & Felger, spec. nov.
Frutex dioictis, spinosus, fere mtdns, ad 1 .7 nt ciltits et 4.8 in latin. Ramiili rigidi
omnesque spinescentes, 3-6mm diametro, hirtelli, glabrati, quisque ramo ramulisque
ejusdem anni snis monade determinata simitl crescentibus. Folia sparsa, anno prbno
decidua, subadpresse hirtella, brevipetiolata, laminis elliptich usque ad cuneatis, rotundatis
subtruncatisve, apiculatis, 5-24mm longis, 4- 15 m in lafis. Flores spisse paniculati fas-
ciculatiic, rubri, 4 (-5) -inert, pedicellis 2-dmin longis; sepala deltoidea, e\tns hirtella,
l-iy2mm longa; petala elliptica, rotundata, glabra, 3-7mm longa, in fl. $ qnain in
fl. 9 insignife inajora; stamina petalis 2-3-plo pluria, filainentis barbatis; pistila 4 (-5),
stylis connata. Dritpae /-5, stipitatae, subcarnosae, II -14mm longae, put amine duro,
lenticulare. Chromosomata: a=13 Typus: Moran 13072 (SD 62160, 62161). Hola-
cantho emoryo plus qitam speciebits ceteris Castelac ramnlis validis spinescentibus sub-
nudis, foliorum axillarinm absentia, et florum staminatorum petallis majoribus accedit
scd ab illo floribus rubris 4 (-5)-meris fructuque mox deciduo recedit. A speciebits
ceteris Castelae et a Holacantho staminibus petalis 2-3-plo plitribns differt.
Dense but irregular stiff spiny nearly leafless dioecious shrub (Fig. 2 ) , to 1.7m
high and 4.8m wide, various parts at first hirtellous, the trichomes simple, stiff, white,
San Diego Soc. Nat. Hist., Trans., 15 (4): 31-40, 5 Jlm 1968
34
San Diego Society of Natural History
Vol. 15
~-?v*x\ *&*
Figure 2. Shrub of Castela polyandra at the type locality, north of the village, Bahia
de los Angeles.
to 0.4mm long, mostly ascending to subappressed. Stems usually several from the base,
crooked, sympodial, mostly to ca. 5cm thick, rarely the main trunk to 18cm thick, the
lower branches sometimes prostrate and 3 '/?m long, not rooting; bark gray, mostly
smooth but on largest stems coarsely flaking; branches at first gray-green, densely
ascending- to spreading-hirtellous, next season yellow-green and glabrous, 3 -6mm thick,
usually slightly flattened in the plane of the nearest branchlet; branch systems of one
season from upper axils, J4-3 (-7) dm long, J/2-2 (-4) dm wide, the axis somewhat
zigzag, ending in a stout spine, with internodes ca. l-4cm long, all but the lower 1-4
and the upper 1-5 nodes with stiff divaricate or upcurved branchlets that rebranch 0-2
times in the same manner, the branchlets of whatever order each slightly flattened at
the base in the plane of the mother axis and each ending in a stout spine. Leaves falling
the first season, subappressed-hirtellous, short-petiolate, the blade elliptic to cuneate,
rounded to subtruncate, apiculate, 5-24mm long, 4-1 5mm wide, thickish, with ca. 4
pairs of obscure ascending lateral veins. Inflorescence axillary, a compact panicle to 1cm
long or reduced to one or a few fascicles sometimes of 10 or more flowers; bracts deltoid,
ca. Vimm long, densely hirtellous; pedicels 2-6mm long, red, hirtellous but sparsely so
by anthesis; buds globose to obovoid, the sepals open except in the smallest buds, the
petals imbricate. Flowers (Fig. 3) in April to July, red, 8-14mm wide; sepals 4 (-5),
red, deltoid-ovate, acute, 1-1 ^mm long, hirtellous but by anthesis nearly glabrate
except on the margins, persistent in fruit; petals 4 (-5), free, spreading, red, ovate to
obovatc, rounded, cupped, 3-7mm long, 2-4mm wide, glabrous; stamens 8-12 (-14-15?),
4 (-5) antesepalous and 4-8 (-10?) epipctalous, in one circle outside the glandular disk,
the filaments bearded ventrally with stiff white trichomes ca. 0.5 -0.7mm long; glandular
disk yellow, 1 ' .-2mm wide, crenulate opposite the filaments. Pistillate flowers 8-1 lmm
wide; petals 3-4'jmm long, 2-2 '/2mm wide; stamens reduced, ca. 1 ''2111111 long; disk
1968
Moran and Felger: Castela polyandra
35
Figure 3. Flowers of Castela polyandra (Moran 13072), X 6l/z: left, staminate; right,
pistillate.
Figure 4. Fruit of Castela polyandra (Moran 11895), X 3. Usually the drupes are four
or fewer.
36 San Diego Society of Natural History Vol. 15
ca. 2mm wide; gynoecium 2 5/2-3mm high, 2-3 '/^mm wide, the pistils 4 (-5), connate
by the styles, the ovaries free, semiovoid, 1 J/2-2mm high, each with one axillary ovule,
the styles ca. 'imm long, the stigmas spreading, 1 y2-2mm long, ca. J/^mm wide,
papillose. Staminate flowers ll-14mm wide; petals 5-7mm long, 3-4mm wide; filaments
erect, 2-3mm long, 0.5 -0.8mm wide at the base and tapering evenly, bearded ventrally
in the lower half, the anthers yellow, oblong, sagittate, 2I/2-3mm long, ca. 1mm wide,
the basal lobes ca. %mm long; disk ca. 1 ^mm wide, concave; gynoecium vestigial or
mostly absent. Fruits (Fig. 4) in August, solitary or sometimes 2-4 in subumbellate
clusters, falling within a few months; drupes 1-4 (-5), stipitate, pale orange or drying
red, 11- 14mm long, the stipe 2-2 '/^mm long and iVz-imm thick, the body 9-12mm
long, 9- 10mm wide, 6-8mm thick, the exocarp fleshy, bitter, the putamen stony, light
tan, lenticular with obtuse to rounded margins, 6-9mm wide, 4- 5 mm thick. Gametic
chromosome number: w=13^
Holotype. — On gravelly east slope at 75 meters elevation, 3 miles north of the
village of Bahia de los Angeles, on the road to La Gringa, Baja California, Mexico
(near 28° 59'N, 113° 351/2,W), 29 April 1966, Moran 1)072 (SD 62160, 62161);
isotypes (to be distributed): ARIZ, BH, CAS, DS, GH, ICF, K, LAM, LL, MEXU,
MICH, NY, RSA, UC, US.
Distribution. — (Fig. 1.) Locally common on bajadas within 10 miles of the Gulf
of California and below 150 meters elevation in central Baja California and on the east
side of Isla Tiburon, Sonora, Mexico. Other collections: Baja California: type locality,
25 August 1964 [fruiting], Moran 11895 (to be distributed: ARIZ, CAS, DS, ICF,
LAM, MEXU, RSA, SD, UC, US) ; mesa 6 miles south of Santa Rosalia, 8 August 1964
[fruiting], Lindsay 1994 (DS) ; arid slopes 7 miles south of Santa Rosalia, 8 April 1963
[sterile], Wiggins & Wiggins 18214 (DS) ; field at Hacienda San Bruno, 10 March 1935
[sterile], Wiggins 7952 (DS) ; rocky outwash plain, San Bruno, 23 November 1938
[sterile], Gentry 4058 (ARIZ, DS, UC), 18 April 1939 [sterile], Gentry 4462 (DS) ;
3 miles south of San Bruno, 13 March 1934 [sterile], Ferris 9036 (DS) ; bajada 5 miles
south of San Bruno, 2 December 1946 [sterile], Wiggins 11510 (DS) ; 17 miles north
of Mulege, 26 October 1964 [sterile], Hastings & Turner 64-400 (ARIZ); 10 miles
south of San Bruno, 25m, 24 May 1961 [staminate], Moran 8465 (BH, CAS, DS, ICF,
MEXU, SD, UC, US); canyon above gypsum village, Isla San Marcos, 22 April 1952
[sterile], Lindsay 2210 (DS). Isla Tiburon, Sonora: coarse gravelly soil of bajada ca. 1
mile inland, near Santa Rosa, 5m, 26 October 1963 [sterile], Felger 9359 (ARIZ,
LAM, SD) ; coarse gravelly soil of bajada ca. 1 mile inland, near Palo Fierro (= Punta
Tortuga), 10m, 2 May 1964 [sterile], Felger 10011 (ARIZ, LAM, SD), 18 July 1964
[staminate], Felger 101)5 A (ARIZ, LAM, SD), [pistillate] 10135B (ARIZ), 15 Feb-
ruary 196 5 [sterile], Felger 125 57 (ARIZ, LAM, SD).
Castela polyandra has been found only within about 10 miles of the Gulf of Cali-
fornia and almost exclusively on east sloping bajadas. Of the dozen localities cited, only
that on Isla San Marcos apparently is not on an east slope. Despite considerable collect-
ing in coastal Sonora, the plant has so far been found there only on the east side of Isla
Tiburon, which has the only extensive east-sloping bajadas close to the east shore of the
Gulf. Also, several Seri Indians have told Felger that they do not know of it on the
mainland. They do distinguish between, and give different but related names to,
C. polyandra and the somewhat similar Holacantha emoryi A. Gray, which occurs
nearby on the mainland (Fig. 1).
Several plants otherwise confined to Baja California and its islands occur on one or
more of the Sonoran islands but apparently not on the mainland. Besides C. polyandra,
those of Isla Tiburon include Camissonia angeloriun (S. Wats.) Raven, Cassia conftnis
Greene, Drymaria bolosteoides Benth., and Pelucba trifida S. Wats.
1968 Moran and Felger: Castela polyandra 37
The two known areas for C. polyandra on the east coast of Baja California are
nearly 150 miles apart, but the plant is not necessarily absent between: the intervening
coast is mainly roadless and little visited and its flora little known.
At the type locality, C. polyandra grows in gravelly granitic soil of the narrow
bajada, one of the dominant shrubs in an open stand with Fouquieria diguetii (van
Tiegh.) I. M. Jtn., F. splendens Engelm., Olneya tesota A. Gray, Larrca diiaricata Cav.,
and Jatropha cuncata Wigg. & Roll., above the zone of Frankenia palmeri S. Wats. On
the east side of Isla Tiburon, it is limited to an open area of the ecotone between the
lower bajada vegetation, where Frankenia palmeri is dominant, and the middle bajada
vegetation, where Larrea diiaricata and other desert shrubs are dominant. Common
shrubs with the Castela include Fhaulothamnns spinescens A. Gray, Olneya tesota,
Krameria grayi Rose & Painter, Condalia globosa I. M. Jtn., Zizyj)hns lycioides A. Gray,
and Lyciinn spp.
At the type locality in May, Moran spent several hours looking for flowers. Al-
though he examined perhaps several hundred shrubs, only a few of these — probably
less than five per cent — bore any buds or flowers; and usually there were few per plant,
mostly on only one or two branches. (Flowering branches commonly were on the south
side of the plant.) To judge from the several sterile collections, flowering apparently is
uncommon and irregular in spring. On Isla Tiburon in July, just after the onset of
the summer rains, Felger found many of the shrubs in full flower; flowers were abun-
dant in the center of the shrub, where rabbits could not reach them.
Dr. Marion S. Cave kindly made chromosome counts from two collections of
C. polyandra (Moran S465, 15072) and for each reports a gametic number of n - 13.
At each of our localities, plants of C. polyandra at flowering time are infested with
small caterpillars, which festoon the flowering branches with spidery webs and spoil the
flowers for botanical specimens. At Bahia de los Angeles in December 1967, Moran
found numerous larvae of assorted sizes, as well as pupae and one adult. Larvae were
eating leaves and bark. Similar infestations occur on C. penmsularis Rose on the southern
islands of the Gulf of California and on Holacantha emoryi A. Gray at least in western
Imperial County, California. Larvae from Bahia de los Angeles and from Imperial
County, raised by Dr. John A. Comstock and Mr. Charles F. Harbison, proved to be
Atteia exquisita Busck. Adults of this moth had previously been taken by Mr. Harbison
on San Francisco Island, where C. peninsularh occurs, and at Bahia de los Angeles.
Powell, Comstock, and Harbison (Ms.) report on the development of this moth.
Similar insect damage was noted in specimens of C. tortuosa Liebm. and C. tweedii
Planch., but the insects are not known.
Relationships
In the treatment of Engler (1931), the new species can go only in the subtribe
Castelinae, comprising the genera Castela Turp. and Holacantha A. Gray, though it
does not quite ht the description of the subtribe. It approaches both these genera but
does not quite fit in either as defined, for example, by Engler or by Cronquist (1944b).
Vegetatively, the new species resembles Holacantha more than it does Castela sensu
stricto. Since every stem apex becomes a spine, new shoots must be lateral, and growth
is thus sympodial. Each new shoot is a determinate system of two to four generations
of axes, evidently all developing at once rather than the main axis first and the branches
later. The branches mostly are rather stout and more or less elongate, with several evi-
dent nodes; and although the ultimate ones are reduced, they are not clearly differen-
tiated from the others as short slender spines. Although leaves in the new species are
as large as in several species of Castela and may persist for several months, there are no
spur shoots with later leaves, and the general aspect of the shrub is almost as bare as
in Holacantha.
38 San Diego Society of Natural History Vol. 1J
In Castela sensu stricto the branching pattern varies widely from one species to an-
other and considerably within certain species. (For some species it is difficult to gen-
eralize from the few terminal branches seen in herbaria; and we have seen no material
of C. depressa Turp.). In C. spinosa Cronq. and in some specimens of C. erect a Turp.
and C. tortuosa Liebm., as in Holacantha and the new species, all branches (not includ-
ing spur shoots) are spine-tipped; and the complex branch system, with three to five
generations of axes, evidently develops as one unit rather than the main axis first and
the branches later. In other specimens of C. erecta and C. tortuosa, the branch system
is reduced to a main axis with short and often slender axillary spines, and the tip of
the main axis is less uniformly spinose. These three species of Castela, in contrast with
Holacantha and the new species, produce additional foliage leaves like the primary ones
but on accessory spur shoots.
In other species of Castela likewise, the tips of the main branches are less regularly
spinose, though they often are subspinose or abortive. Growth often appears to be sym-
podial and in some species probably is uniformly so; but from herbarium specimens it is
not clear that it is so in all species. In C. peninsularis Rose, the main axis regularly bears
spinose primary and secondary and sometimes also tertiary branches. In C. calcicola
(Britt. & Small) Ekman, C. coccinea Griseb., and C. jacquinifolia (Small) Ekman,
spinose primary and secondary branches are sometimes present. In each of the other
species that we have seen, axillary spines are present in at least some specimens; but
there is no indication of unit development of more complex branch systems. In all these
species except C. jacquinifolia, axillary leaves were seen in at least some specimens; it is
not clear whether they would always be present on some older branches.
Thus in its branching pattern and in its stout and nearly leafless branches, the
new species resembles Holacantha more closely than it does any species of Castela sensu
stricto. From those species of Castela which it otherwise most closely resembles in
branching pattern, it differs, as does Holacantha, in the lack of spur shoots. On the
other hand, the flowers are primarily 4-merous and infrequently S-merous; the few
leaves present are relatively large, and some of them persist for several months; and the
fruits are shed promptly: in these respects it resembles Castela. In plants of the type
collection, flowering under perhaps less than ideal conditions, the inflorescence is reduced,
as usual in Castela; but in those from Isla Tiburon, that flowered at the onset of the
summer rains, it is stouter and more like that of Holacantha.
Raven (1967) reported a somatic chromosome number of 2/7=26 for a cultivated
plant of H. emoryi, originally from Maricopa County, Arizona. For a collection
(Moran 13138) from 6lA miles southeast of Coyote Wells, Imperial County, California,
Dr. Cave reports a gametic number of »=13. Thus H. emoryi has the same chromosome
number as reported above for C. polyandra. Apparently there are no reports for Castela
sensu stricto.
With respect to petal size, greater in staminate than in pistillate flowers, the new
species resembles //. emoryi and seems to differ from other members of both genera.
Apparently in H. steuartii C. H. Muller and so far as we can discover in other species
of Castela, there is no such size difference.
With respect to stamen number, varying from twice to thrice the number of
sepals but commonly falling between, the new species differs both from Castela sensu
stricto and from Holacantha and indeed from most other Simaroubaceae. Although
Holacantha and perhaps some other species of Castela have some tendency to anisomery,
apparently in those species the stamens are rarely more than twice as many as the sepals.
Although Holacantha has always been treated as a genus separate from Castela, the
question of where to place the new species has led us to consider its status more closely.
In some respects the new species resembles Castela sensu stricto and in others Holacantha.
Thus, Holacantha does not now appear as distinct as it did. But the new species also
1968 Moran and Felger: Castela polyandra 39
differs from both. It cannot be placed in either group without broadening the definition
of the group while at the same time somewhat reducing the distinction of the two; yet,
it is so obviously similar to both that there is no question of establishing a new genus
for it. For these reasons, we think it best to reduce Holacantha to a section of Castela
and to place the new species in a separate section. We define Castela and its sections as
follows.
Castela Turp., Ann. Mus. Paris 7: 78, 1806. (Nom. conserv.)
Dioecious shrubs or small trees, mostly with some spinose branchlets, some with all
branches spinose. Leaves alternate, scattered on the main branches or crowded on spur
shoots, simple, entire or toothed, thick, mostly less than 5cm long, sometimes scalelike,
caducous or mostly persistent. Inflorescence axillary or supra-axillary, a panicle or
raceme of cymules or individual flowers or reduced commonly to one or a few fascicles
or even to one flower. Flowers 4-8-merous; sepals 4-8, small, deltoid, persistent; petals
as many, several times larger, imbricate in bud, deciduous; stamens mostly twice to
thrice as manyr, inserted around a glandular disk; pistils mostly as many as the sepals.
each with one ovule, separate below, united by the styles, the stigmas separate. Fruit of
separate drupes, deciduous or persisting for several years.
Key to Sections:
Flowers 4 (-S)-merous; fruits soon deciduous; inflorescence less than 2cm long; leaves
with well-developed blades, mostly persistent.
Stamens twice as many as the sepals; petals not larger in the staminate flowers;
branches spinose or not, leafy; spur shoots commonly present. 1. Castela
Stamens twice to thrice as many as the sepals, the number varying in the same
inflorescence; petals larger in the staminate flowers; branches all spinose, sparsely
leafy; spur shoots absent. 2. Eremacantha
Flowers (S-) 6-8-merous; fruits persistent for several years; inflorescence 2-1 0cm long;
adult leaves scalelike, soon deciduous; branches all spinose; spur shoots absent.
3. Holacantha
Castela section Castela
Neocastela Small, N. Am. Fl. 25: 2 30, 1911.
Castelaria Small, loc. cit.
Main axes abortive and subspinose or in some apparently indeterminate, the pri-
mary (to tertiary) branches commonly spinose and often reduced to axillary spines; less
commonly all branches spinose. Leaves with well-developed blades, persistent, the pri-
mary ones scattered but similar ones commonly produced later on spur shoots, the shrub
thus leafy. Inflorescence a small panicle or raceme with the axis slender and rarely 1cm
long, or commonly reduced to a fascicle or even to one flower. Flowers 4 (-J)-merous;
petals apparently the same size in staminate and pistillate flowers; stamens normally
twice as many as the sepals. Drupes sessile by a narrow base or commonly stipitate, soon
deciduous.
Type Species: C. Jepressa Turp.
In the latest revision, Cronquist (1944a) recognized 12 species, extending from
Texas to Argentina and from the Galapagos Islands to the Antilles.
Castela section Eremacantha Moran & Felger, sect. nov.
Ramitli spinescentes, quisque ramo ramulisque ejusdem aim/ snis monade determinata
sinntl crescent/hits. Flores 4 (-5)-meri, staininihiis senilis 2-1-j'lo pluribus. Drupae
stipitatae, mox Jeciduae. Typus: C. polyandra.
Branches all spinose, the branch system determinate, with 2-4 generations of axe>
developing as a unit; spur shoots none. Leaves with well-developed blades, persistent tor
40 San Diego Society of Natural History Vol. 1 5
several months but sparsely distributed. Inflorescence a compact panicle, with the axis
stout but rarely 1cm long, or reduced to one or a few fascicles. Flowers 4 (-S)-merous;
petals larger in staminate than in pistillate flowers; stamens twice to thrice as many as
the sepals. Drupes stipitate, soon deciduous.
Type Species: C. polyandra Moran & Felger.
The name Eremacanfha is from Greek words for desert and shrub.
Casfcla section Holacantha (A. Gray) Moran & Felger, stat. nov.
Holacantha A. Gray, PI. Thurb. 310, 18 54.
Branches all spinose, the branch system determinate, with 2-4 generations of axes
developing as a unit; spur shoots none. Adult leaves reduced, soon deciduous, the shrub
thus essentially leafless. Inflorescence an open to rather crowded panicle, the branches
stout, like the vegetative branches, the axis 2-10cm long. Flowers (5-) 6-8-merous,
tending to be anisomerous; petals of staminate flowers the same size as or larger than
those of pistillate flowers; stamens twice as many as the sepals or fewer than twice as
many. Drupes sessile by a broad base, persistent for several years.
Type Species: Holacantha emoryi A. Gray.
The section consists of the following two species, native in the southwestern
United States and northern Mexico.
Castela emoryi (A. Gray) Moran & Felger, comb. nov.
Holacantha emoryi A. Gray, PI. Thurb. 310, 18 54.
Castela stewartii (C. H. Muller) Moran & Felger, comb. nov.
Holacantha stewartii C. H. Muller, Madrono 6: 131, 1941.
Acknowledgements
We are grateful to the curators of the herbaria at the Arnold Arboretum, the New York Botanical
Garden, and the United States National Museum, for the loan of specimens. We are indebted to Dr.
Marion S. Cave for the chromosome counts, to Miss Gayle Culver for the map and the drawings, and to
Drs. Peter H. Raven and Ira L. Wiggins for critically reviewing the manuscript. Last but not least, we
sincerely thank each other for help and encouragement during the preparation of this paper.
Literature Cited
Cronquist, A.
1944a. Studies in the Simaroubaceae, I. The genus Casfcla. J. Arnold. Arb. 25: 122-128.
1944b. Studies in the Simaroubaceae, IV. Resume of the American species. Brittonia 5: 128-147.
Engler, A.
1931. Simarubaceae, p. 359-405. /;; Engler, A., and H. Harms, Die natiirlichen Pflanzenfamilien.
2 ed. Vol. 19a.
Powell, J., J. A. Comstock, and C. F. Harbison
Ms. Biology, life history, and geographic distribution of Attcia exquisita (Lepidoptera: Ypono-
meutidae) .
Raven, P.
1967. Holacantha emoryi Gray. //; Anon., Documented chromosome numbers ot plants. Madrono
19: 134-136.
Accepted for Publication 11 April 1968.
San Diego Natural History Museum and Los Angeles County Museum of Natural
History.
MUS. COMP. ZOOL.
LIBRARY
JUN 1 8 ^968
HARVARD
UNIVERSITY.
RECENT DATA ON SUMMER BIRDS OF THE
CHIRICAHUA MOUNTAINS AREA,
SOUTHEASTERN ARIZONA
J. DAVID LIGON AND RUSSELL P. BALDA
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 5, 5 JUNE 1968
RECENT DATA ON SUMMER BIRDS OF THE
CHIRICAHUA MOUNTAINS AREA,
SOUTHEASTERN ARIZONA
J. David Ligon and Russell P. Balda
Abstract. Over four breeding seasons 167 species of birds were recorded from the Chiricahua Mountains
and adjacent areas; 19 of these were winter residents or migrants not seen after May. The status
of several species apparently has changed since the study of Tanner and Hardy (1958). The Golden
Eagle (Aquila chrysaetos) and Prairie Falcon (Falco mexicanus) appear to have decreased, whereas
the Harlequin Quail (Cyrtonyx montezumae) and several of the hummingbirds apparently are more
common than previously. Several cavity-nesting species appear to have increased in numbers and tin
Eastern Bluebird (Sialia sialis) has become established as a breeding species since 1960. Black Swifts
(Cypseloides niger), a species for which there is no specimen for Arizona, were seen on one occa-
sion, and a female Lucifer Hummingbird {Calo thorax lucifer) , another unexpected species, was seen
on several occasions.
Resumen. Se observaron 167 espccies de aves en las Montanas Chiricahua y zonas adyacentes, durante
las cuatro estaciones de cria; 19 de estas especies eran residentes invernales o aves migratorias que no
aparecian despues de Mayo. El estado de varias especies ha cambiado aparentemente desde los estudios
realizados por Tanner y Hardy (1958). El Aguila Real (Acjuila chrysaetos) y el Halcon Cafc; (Falco
mexicanus) han disminuido, mientras que la Codorniz dedilarga (Cyrtonyx montezuma) y varios
colibris son probablemente mas abundantes ahora que antes. Varias especies de las que anidan en
cavidades han incrementando en numero, asi, el Azul de tempestad (Sialia siulis) aparece bien estable-
cido, de modo que cria en esta zona desde 1960. EI Vencejo negro (Cypseloides niger) no se ha
registrado en Arizona, y solamente se ha visto en una ocasion en esta region, asi como una hembra
del Colibri Chupamirto morada grande (Calothorax lucifer) otra de las especies raras en esta zona,
en donde solo ha sido observado en pocas ocasiones.
The Chiricahua Mountains of Cochise County, Arizona, are located near the
Arizona-New Mexico boundary about 20 miles north of the Mexican border. These moun-
tains, along with the Huachuca and Santa Rita mountains, are of special ornithological
interest because the northernmost extension of several essentially Mexican species and the
southern limit of several boreal forms occur here. This, together with the wide variety
of habitats found in the Chiricahuas, ranging from desert scrub to spruce-fir forest,
combine to make this region rich in breeding bird species. In recent years the Chiricahuas
have become relatively well known to ornithologists, largely because the Southwestern
Research Station of the American Museum of Natural History is located in Cue
Creek Canyon.
The avifauna of the Chiricahua Mountains has been popularized by a number of
ornithologists (Peterson, 1948; Brandt, 1951), but little detailed work has been pub-
lished. Tanner and Hardy (1958) briefly describe the clevational distribution of some
of the more common birds and review the avifaunal studies of the region, pointing out
the sparseness of data dealing with breeding birds of the area.
No comprehensive study of plant distribution in the Chiricahuas has been pub-
lished, but Balda (1967) describes in detail the habitats he studied, and W'hittaker and
Niering (1965) present an excellent discussion of plant distribution in the nearby and
similar Santa Catalina mountains.
We spent a great deal of time in riparian communities, habitats not described by
Tanner and Hardy (1958). These are affected by lower temperatures (due to nightly
cold air drainage) and higher moisture conditions than the adjacent typical plant com-
munities of a given elevation. Riparian habitat is best developed in canyon bottoms but
San Diico Soc. Nat. Hist., Trans. 15 (5): 41-50, 5 |i m I •>■
42 San Diego Society of Natural History Vol. 1 5
is present to some degree wherever permanent or temporary streams are found. Plant
species diversity is great in these areas and is reflected in high bird species diversity.
Lower elevation riparian woods are characterized by sycamore, willow, cottonwood,
maple, walnut, ash, and several species of oaks. The Station is located in such an area.
In more restricted canyon bottoms, such as the South Fork of Cave Creek Canyon,
many coniferous species typically found at higher elevations are intermingled with the
deciduous trees. In the Chiricahua Mountains, Arizona cypress is limited to this
community.
Because the avifauna of the Chiricahuas has special interest, we feel that updating
the avifaunal list of Tanner and Hardy (195 8) is worthwhile, even though an excel-
lent state bird book (Phillips et al., 1964) has recently been published. Our data indi-
cate that dramatic changes in the status of some birds have occurred since Tanner and
Hardy's study. Perhaps the best example is the arrival and population increase of the
Eastern Bluebird (Sialia sialis) . In addition, some common species, principally nocturnal
ones, were overlooked.
Ligon camped in Cave Creek Canyon from 17 May to 2 1 July 1964; Balda was
at the Southwestern Research Station from 3 June to 2 July of that year. From 17 Feb-
ruary to 18 August 196 5 Balda conducted a study of the breeding habitats of the birds
of the region. Ligon arrived at the Station on 4 May and remained until 5 August. In
1966 Ligon spent three days in Cave Creek Canyon in April and 18 days there in July.
He was at the Station for several days during May 1967.
We follow the terminology of Tanner and Hardy (195 8) in referring to the status
of each species. Species recorded by them for which we have no additional comments
are simply listed with a one-word designation of status. More detailed information on
abundance is currently being prepared. Those species not recorded by the above authors
are indicated by an asterisk; species for which either nests were found or juveniles seen
are indicated by a dagger. All specimens were taken by Ligon and are deposited in the
collections of The University of Michigan Museum of Zoology. Only species recorded
after 1 May are included in this account.
The village of Portal is located below the mouth of Cave Creek Canyon at an
elevation of 4770 feet. Paradise is about five miles northwest of Portal. Apache is 12
miles south of Rodeo, New Mexico on U. S. Highway 80. Stewart Forest Camp and
Sunny Flats Campground are located in Coronado National Forest between Portal and
the Station (elevation 5400 feet) ; Rustler Park, also in Coronado' National Forest, is
higher (8400 feet).
Systematic List
Oxyitra jamaicensis, Ruddy Duck1'. — A male in breeding plumage was seen on 21 July 1965, in
a cattle tank at the foot of the mountains near Portal.
Ca/hartcs aura, Turkey Vulture. — Common. Large flock of about 50 roosted nightly on huge rock
above Stewart Forest Camp.
Accipiter gentilis, Goshawk^. — An adult female was collected 5 July 1964, nine miles southwest of
Apache. A nest with one flying young was located in the same area the following day at an elevation of
about 5 200 feet. This nest was about 200 yards downstream from an occupied nest of A. cooperii. Gos-
hawks were seen on four occasions over ponderosa pine forest below Rustler Park.
Accipiter striatus, Sharp-shinned Hawk*. — One observed at the station for several days in early
May 1965; captured hummingbirds with apparent ease from the station feeders.
Accipiter cooperii, Cooper's Hawk'. — Common in Cave Creek Canyon from oak woodland to spruce-
fir forest in 1964 and 1965; seven nests found, four examined, each with four or five eggs or young.
Bit/eo jamaicensis, Red-tailed Hawk. ■ — Uncommon.
Bu/eo swainsoni, Swainson's HawkT. — One seen building a nest 20 May 1964. Nest with two well-
grown living young, and a third dead, found 2 July 1964, near Potral.
Buteo albontatus, Zone-tailed Hawk"'. — One seen by Ligon perched on rocks in dry South Fork of
Cave Creek on 21 May 1964. Two seen by Balda along the South Fork of Cave Creek Canyon in
May 1965.
1968 Ligon and Balda: Chiricahua Birds 45
Buteo lagopus. Rough-legged Hawk"'. — Winter visitor; last seen on 5 May.
Aquila chrysactos. Golden Eagle. — Pair seen regularly east of Portal in 1964 and 1965. Not seen in
the mountains.
Circus cyaiwus. Marsh Hawk*. — One was seen occasionally over the grasslands below Rodeo from
March through May 1965.
Falco mexicanus, Prairie Falcon. — Seen irregularly over grassland south of Rodeo and also on west
side of the mountains in 1965.
Falco peregrinus, Peregine Falcon::". — Seen regularly in 1964 and 1965 along high cliffs above
Stewart Forest Camp and Sunny Flats Campground. Believed to have nested. Ligon observed one drive a
Red-tailed Hawk to earth near Sunny Flats Campground.
Falco sparivrius. Sparrow Hawk'. — Uncommon. In 1964 a pair nested in a large dead Cottonwood
one and one-half miles east of Portal. A nest was located near Paradise in 1965. Seen occasionally in pon-
derosa pine forest.
Callipepla squamata, Scaled Quail'. — Common in grasslands containing a scattering of shrubs along
Highway 80 east of the Chiricahuas. Young seen on 8 July 1965. Newly-hatched young captured and
photographed on 11 July 1965. A very late nest located on 17 August contained 12 eggs.
Lophortyx gambclii, Gambel's Quail". — Common in desert scrub surrounding the mountains. Newly-
hatched young captured and photographed 9 June 1964.
Cyrtonyx montezumae, Harlequin Quail""'''. — Not uncommon but not regularly seen in oak woodland
and pine-oak-juniper woodland communities. Four young seen by Balda on 20 July 1965.
Meleagris gallopavo, Turkey'. — Uncommon. Nest, eight eggs located near the Station on 17 May
1965. Seven chicks seen 13 June 1964. Flocks of 8 to 10 birds were seen often in the ponderosa pines
below Rustler Park in 1965.
Cbaradrius lociferus, Killdeer. — Seen irregularly about water tanks.
Actitis macularia, Spotted Sandpiper*. — Five seen on small ponds at the foot of the mountains on
21 July 1965.
Columba fascia/a, Band-tailed Pigeon. — Regular in flocks in Cave Creek Canyon in early summer,
becoming common to abundant at high elevations in mid-July, when pairs were apparently preparing to
nest.
Zcnaida asiatica, White-winged Dove'. — Commonly seen from desert scrub up into oak woodland.
Zenaidura macron ra, Mourning Dove^. — This species was most common in desert scrub where Balda
located 23 nests in 1965. At one nest the young fledged on 4 March. One pair was seen on repeated
occasions in ponderosa pine forest.
Columbigallina passerina, Ground Dove*. — One seen by Ligon on 9 June 1964, one and one-half
miles east of Portal near a watering tank. Phillips ft al. (1964: 43) consider the records for the east
side of the Chiricahuas as representing strays rather than a breeding population.
Geococcyx californianus, Roadrunner. — Uncommon below Portal.
Otus asio, Screech Owl*'. — Common in Cave Creek Canyon, where nests were found in 1965
and 1967.
Otus trichopsis. Whiskered Owl*''. — Common in Cave Creek Canyon. One or more nests were
located near the station in 1964, 1965, 1967.
Otus flamnnolus, Flammulated Owl'. — Uncommon. Two nests were located in 1965, each with one
young. One nest also contained an addled egg.
Bubo virginianus, Great-horned Owl*. — Uncommon from desert scrub up to the lower edges of
ponderosa pine forest.
Glum itlium gnonia, Pygmy Owl*^. — Uncommon. Nest found near the Station on 17 May 1967, in
cavity used by Elf Owls the previous two years. Seen and heard occasionally above Rustler Park.
Micrathene whitneyi, Elf Owl*'''. — Abundant in Cave Creek Canyon where 20 nests were located by
Ligon in 1 965.
Speotyto cuniculnria. Burrowing Owl*'1'. — Rare on foothill grasslands surrounding the mountains.
One pair nested three miles south of Apache in 1965.
Strix occidentalis, Spotted Owl. — Uncommon. Heard calling in Cave Creek Canyon on 11 July 1964
by Ligon, J. P. Hubbard, and B. K. Harris. One seen at Rustler Park by Balda, 14 August 1 96?.
Caprimulgus lociferus, Whip-poor-will1'. — Common along Cave Creek Canyon floor up to the high-
est elevations of the mountains. Female with egg ready for laying collected 31 May. 1964.
Pbalaenoptilus nuttallii, Poor-will. — Common from desert scrub through oak woodland especially ""
the drier slopes. Heard primarily at dusk and dawn. Seen foraging about lights at the Station on several
occasions.
Cbordeiles minor, Common Nighthawk*. — Seen 18 July, 1965, southwest of Apache.
44 San Diego Society of Natural History Vol. 15
Chordeiles acutipciinis. Lesser Night hawk. — Common on desert sciub surrounding mountain and
occasional through oak woodland.
Cypseloides niger, Black Swift*. — Rare. Flock of about a dozen seen above Herb Martyr Dam on
11 July 1964, by Ligon, J. P. Hubbard, and B. K. Harris. They were foraging with Purple Martins,
Violet-green Swallows and White-throated Swifts. There is no satisfactory specimen record for Arizona
(Phillips ct al., 1964: 58).
Aeronantes saxatalis. White-throated Swift. — Common in the mountains. Also foraged over the
desert.
Calothorax lucifer, Lucifer Hummingbird*. — Extremely rare. A female was first seen by Balda on
17 June 1965 southwest of Apache in oak woodland adjacent to xeric slopes covered with manzanita and
agave. This female was seen again on 26 June and 1 July, and was positively identified at a local rancher's
hummingbird feeder. There are only two very old specimen records for the state (Phillips, et al., 1964: 62).
Archilochns alexandri, Black-chinned Hummingbird'. — Uncommon in desert, but very common in
oak woodland, riparian canyons and oak-juniper woodland. Occasionally sighted in lower ponderosa pine
forest. Fourteen nests were located by Balda in 1965, with extreme dates 15 April to 28 June.
Selasphoms platycercus, Broad-tailed Hummingbird^. — Uncommon in oak woodland and oak-juniper
woodland but common in riparian canyon and coniferous forests. An exceptionally early nest found in
South Fork on 4 April contained two eggs. Both young fledged by 1 3 May. The latest nest was located
in ponderosa pine forest and contained two young on 1 August.
Selasphorus nifus, Rufous Hummingbird. — Regular from woodlands through coniferous forest from
mid-July onward. Seen occasionally in desert as well.
Stcllula calliope. Calliope Hummingbird*. — Male shot but net retrieved by Ligon in Rustler Park
on 16 July 1964. One seen on 2 August 1965 in ponderosa pine forest.
Eugenes fulgens, Rivoli's Hummingbird. — Regular at station feeders but uncommon in woodland and
riparian canyons. Seen occasionally at lower edge of ponderosa pine forest.
Lampornis clemenciae, Blue-throated Hummingbird7. — Uncommon in riparian habitats but regular at
the station feeders. Two nests each containing two eggs were located on 28 June 1965, in the South
Fork of Cave Creek Canyon. This species is very aggressive and was seen driving other species from feed-
ing sites and destroying nests of other hummingbirds to obtain nesting materials.
Amazilia verticalis, Violet-crowned Hummingbird*. — First seen at Station in June in 1963, 1964,
and 1965. R. W. Lasiewski captured one in July 1965.
Trogon clegans, Coppery-tailed TrogonT. — Nest found near Sunny Flats Campground on 10 June
1964; the two young fledged on 15 July. This species was seen in 1964, 1965, and 1966. In 1965 it was
common in the South Fork of Cave Creek Canyon, where on 2 3 May Balda observed three males and
two females in close proximity. Males wandered great distances up and down Cave Creek Canyon.
Colaptcs cafer, Red-shafted Flicker7. — Common in woodland and coniferous forest. Eight nests were
found; extremes are 2 May to 24 June.
Melnuerpes formicivorus, Acorn Woodpecker'. — Common to abundant resident of Cave Creek
Canyon. Numerous nests were found.
Dcmlrocopos villosus, Hairy Woodpecker . — Abundant in coniferous forest. Young left nest below
Flys Peak on 24 June. Two nests were located near the Station, one in a walnut, the other in a sycamore,
well below the usual altitudinal breeding range of this species.
Dcmlrocopos scalaris, Ladder-backed Woodpecker7. — Uncommon in desert scrub lowlands surround-
ing mountains. Seen foraging as high as 5 200 feet on xeric slopes. Two nests, each in agave, were located
on 25 May 1964. Both contained three young.
Dcmlrocopos arizonae, Arizona Woodpecker7. — Common in oak and pine-oak woodland, but often
silent and difficult to locate. Nests were found in sycamore, Cottonwood, and walnut trees.
Tyrannus verticalis, Western Kingbird7. — Common in desert but rare in higher woodlands. Three of
four nests in desert scrub were located on the high flower stalk of a yucca. The earliest nests located
were being built on 23 May 1965.
1' \ rtin n us vociferans, Cassin's Kingbird7. — Uncommon in woodlands up to 7000 feet. Two nests were
located in oaks, another in a sycamore.
Myiodynastes luteiventris, Sulfur-bellied Flycatcher7. — Common in the lower portion of Cave Creek
Canyon, less so near the Station. Four nests, all in sycamore cavities, were located between Stewart Forest
Camp and Sunny Flats campgrounds in 1964. In 1966, the first egg was laid in a nest near the Station
on 2 5 July.
Myiarcbus tyrannulus, Wied's Crested Flycatcher*7. — Uncommon in Cave Creek Canyon. Nest con-
taining well-grown young located high in a cavity in a sycamore on 7 July 1966.
Myiarcbus cinerascens, Ash-throated Flycatcher7. — Common in all wooded habitats from desert
through pine-oak woodlands. Highest density found in the latter. Five nests found in cavities or dead
oak stumps. Earliest nest was being constructed on 9 May 1965.
1968 Ligon and Balda: Chiricahua Birds 45
Myiarchus tuberculifer, Olivaceous Flycatcher. — Common in canyons.
Sayortiis nigricans. Black Phoebe'. — Uncommon. In 1964 and 1 96 S occupied nests were located under
the bridge at Stewart Forest Camp. In 196 5 a nest was also located under the bridge near the Station and
on the eave of a station building (four eggs on 8 May).
Sayornis saya, Say's Phoebe'. — Uncommon at lower elevations but common near buildings and other
man-made objects. Nested at Station each year, 1964-67.
Empidonax difficilis, Western Flycatcher'. — Common in ponderosa pine forest; abundant in spruce-
Douglas fir forests on the peaks. Four nests were located in 1965. One nest with three eggs was located
on 1 June in an upturned root system of a large pine. The other nests were located in cavities or attached
to the side of rough-barked trees.
Empidonax fuliifrons. Buff-breasted Flycatcher"'. — One or two pairs spent the summer of 1965 in
the oak-juniper woodland behind the Station. First seen on 8 May.
Contoptts pertinax, Coue's Flycatcher*. — Common in ponderosa pine forest where one nest was
located in 1964 and two in 1965. The earliest nest was being built 20 May. Two stub-tailed young were
taken on 2 8 June 1964.
Contopus sordiduhis, Western Wood Pewee'. — Abundant.
Nu/tallornis borealis, Olive-sided Flycatcher''. — One migrant seen on 30 May 1965, in ponderosa
pine forest.
Pyrocephalus rubinus, Vermilion Flycatcher'. — Rare at lower elevations except near irrigated areas or
human habitation. Male collected at Cave Creek Cabins near Portal on 28 June 1964. Male seen as high
as the Station on three different occasions.
Camptostoma imberbe, Beardless Flycatcher"'. — Rare on the dry slopes and open canyon floors of
Cave Creek. Six sight records by Balda from March through July 1965, from below Stewart Forest Camp
up to South Fork.
Eremophila alpcstris, Horned Lark*. — Common in all grasslands.
Tacbycineta tbalassina, Violet-green Swallow. — Common at all elevations in the mountains.
Hirundo rustica, Barn Swallow* *. — Rarely seen foraging over desert scrub to woodlands. One nest
with three young found in rancher's barn 12 miles south of Apache.
Progne subis, Purple Martin. ■ — Uncommon.
Cyanocitta stelleri, Steller's Jay. — Common in higher riparian and coniferous forest.
Aphelocoma coerulcscens, Scrub Jay*. — Seen rarely in thick foliage at low elevations. One seen one
and one-half miles east of Portal on 20 June 1964; another seen 13 May 1967 near Cave Creek Cabins,
between Portal and the entrance to Coronado National Forest. Three to five seen 10 July 1964, nine miles
southwest of Apache, at the edge of oak woodland.
Aphelocoma ultramarina, Arizona Jay. — Common. Stub-tailed young seen 28 May 1964; one taken
on 3 July 1964. Three nests found in May 1965, all with four eggs.
Con us corax, Raven. — Rare at high altitudes. Seen frequently along the highway 15 miles north of
Douglas, Arizona.
Conns cryptolcucus, White-necked Raven'. — Uncommon on the desert. One nest containing two
young was located in a clump of yucca west of the mountains in 1965.
Pants sclateri, Mexican Chickadee*. — Common but secretive during nesting in the ponderosa pine
forest. Nest located in gambel's oak cavity on 10 June 1964.
Pants inornatus, Plain Titmouse"'". — Uncommon in and around Cave Creek Canyon, but fairly com-
mon in oak woodlands in more xeric areas. One nest was located behind the Station in a cavity of a dead
juniper on 12 June 1965. Young left this nest on 1 9 June.
Pants wollweberi, Bridled Titmouse*. — Common in oak woodland, pine-oak-juniper woodland and
riparian. It overlapped with the Plain Titmouse in the first two communities. Five nests were found in
natural cavities of oaks and sycamore. The earliest nest located had three eggs on 1 May while the latest
nest had seven young on 6 June.
Auriparus flaviceps, Verdin*. — Uncommon on the desert. Four nests were found in 1965 and one
in 1964. A male was observed building a nest, which was never completed, on 3 1 March. A nest with
four eggs was found on 24 May 1964; another, containing one egg was located on 2 June 1965. Im-
matures were observed building roost nests on three occasions.
Psaltiparus minimus, Bushtit*. — Common in oak-juniper and riparian woodlands, uncommon in open
oak woodland. Three nests were found in 1965; the earliest was under construction on 4 April. Young
left this nest about 7 May. The latest nest was found 29 May; the young fledged on 19 June.
Sitta carolinensis, White-breasted Nuthatch*. — Most common in ponderosa pine forest. Breeds sp.n
in riparian habitat, oak-juniper woodland, and spruce-Douglas fir forest. One nest found
17 June in a natural cavity in silver-leaf oak.
46 San Diego Society of Natural History Vol. 1 5
Sitta canadensis. Red-breasted Nuthatch1'. — Common only in spruce-Douglas fir forest where one
nest was observed being built on 24 May 1965.
Sitta pygmaea. Pygmy Nuthatch \ — Abundant in all coniferous forest from 5,000 feet to top of
mountain. Nine nests found in 1965. First young left the nest on 14 June.
Ccrthia familiar is, Brown Creeper. — Abundant in spruce-Douglas fir and common wherever conifers
are present. Immatures move downward in late June. An immature female was collected at Sunny Flats
Campground on 30 June 1964.
Troglodytes aedon, House Wren'. — Common in coniferous forests, where four nests were located in
1965. All contained young by the last week of June.
Tbryomanes beuickii, Bewick's Wren". — Abundant in oak woodland, common in oak-juniper wood-
land. Five nests located in natural cavities of Emory Oak. The earliest nest was under incubation en
19 May, while the latest was being built on 5 June.
Campylorhyncbus brunneicapillum, Cactus Wren7. — Common on desert scrub surrounding the
mountain. Eighteen nests found in 1965; dates ranged from 24 March to 21 July.
Catbcrpcs mexicanus, Canon Wren. — Uncommon.
Sulpirides obsolctus, Rock Wren. — Rare.
Miimts polyglottos, Mockingbird. — Uncommon in desert but common in open oak-woodland.
Dnmetella carolinensis, Catbird". — Migrant, seen by Balda on 9 May 1965.
Toxostoma bendirei, Bendire's Thrasher. — Rare in desert scrub.
Toxostoma currirostre, Curved-billed Thrasher""'. — Uncommon in desert. Nesting range overlaps
with Bendire's Thrasher at the foot of the mountain. Five nests found, four of which were slung between
two yuccas. The earliest nest had three eggs on 4 May, the latest had three eggs on 28 May.
Toxostoma dorsale, Crissal Thrasher^. — Rare from desert up to open oak woodland. Four nests, all
containing eggs, were located; three were found in late May, one on 9 July.
T Urdus migratorius, Robin'. — Common in riparian and ponderosa pine but uncommon in oak-juniper
woodland and spruce-Douglas fir forest. Five nests found in 1965, the earliest with two young on 28 May.
Hylocichla guttata. Hermit Thrush. — Common in coniferous forests where moisture is available,
extending down into moist shaded canyon bottoms. A female with a fresh brood patch was taken along
the South Fork of Cave Creek on 14 May 1967.
Hylocichla ustulata, Swainson's Thrush"'. — Uncommon migrant, seen in all years during the last
two weeks of April and first two weeks of May. One captured and released at the Station on 14 May 1967.
Sialia sialis, Eastern Bluebird*^. — This species was first recorded as a breeding bird in Cave Creek
Canyon in 1960 (Phillips, et al., 1964: 131). It has since become a regular breeding species in the oak-
juniper woodland and riparian habitat near the Station. Young left one nest on 12 June 1965; newly
hatched young were in another on 7 July 1966.
Sialia mexicana, Western Bluebird'. — Abundant in ponderosa pine forest, where four nests were
located. Young fledged in late June and the first two weeks of July.
Myadcsfcs tounsendi, Townsend's Solitaire. — Migrant (but cf. Tanner and Hardy, 1958). Latest
sighting was 20 May 1964, near Herb Martyr Dam, by Ligon.
Polioptila caerulea, Blue-gray Gnatcatcher. — Common in oak woodlands.
Poliop/ila melanura, Black-tailed Gnatcatcher. — Rare in desert scrub.
Regulus satrapa, Golden-crowned Kinglet. — Common in spruce-fir forest but not found elsewhere.
Regains calendula, Ruby-crowned Kinglet. — Abundant in spruce-fir forest but also limited to only
this community.
Pbainopepla nit ens, Phainopepla*. — Rare breeder in desert scrub below Portal and also on the west
side of the mountain. Records from June, July, and August, 1965.
Lanins Indoi icianus, Loggerhead Shrike' . — Uncommon in most desert scrub areas but locally com-
mon. Three nests with four young each were found in 1965. Young left these nests on 12 May, 16 May,
and 20 June.
Vireo huttoni, Hutton's Vireo^. — Uncommon in oak woodlands, more common in riparian situations.
Nest found in sycamore on 15 May 1965.
Vireo bellii. Bell's Vireo. — Not uncommon in thick mesquite along dry creek beds east of Portal.
Vireo licinior, Gray Vireo. — Of very local distribution. Found by Balda to be uncommon in oak
woodland 20 miles southwest of Apache.
Vireo solitarius. Solitary Vireo^. — Abundant in ponderosa pine forest, and common in oak-juniper
woodland and riparian communities. One nest with young found 11 June 1964; male parent taken. Four
nests were found in 1965.
Vireo gilvus. Warbling Vireo. — Common in aspen stands, which are found at higher elevations.
Seen on a few occasions as low as the Station.
1968 Ligon and Balda: Chiricahua Birds 47
Vermiiora virginiae, Virginia's Warbler. — Uncommon in shrubby areas in botli the coniferous
forests and in riparian communities.
Vermiiora luciae, Lucy's Warbler1'. — Uncommon from desert through the woodlands. Nest found
one and one half miles east of Portal on 9 June 1964.
PcuceJramus taeniatus, Olive Warbler'1". — Surprisingly common in the ponderosa pine forest in 1965;
rare in spruce-Douglas fir forest. Two nests located in 1965 and one in 1967.
DenJroica petechia, Yellow Warbler. — One seen and heard repeatedly at the Station through the
first two weeks of June 1963; no other records.
DenJroica caerulescens, Black-throated Blue Warbler*. — Casual migrant; seen by Baida on 7 Ma)
1965 in riparian habitat.
DenJroica auduboni, Audubon's Warbler"'. — Common in riparian and coniferous forests. One rust
high in a ponderosa pine was located. Young left around 28 June.
DenJroica nigrescens, Black-throated Gray Warbler7. — Abundant in woodland and upper riparian
communities. Six nests were found in 1965. Incubation was under way on 17 May in the earliest nest.
Recently fledged young were common during the second and third weeks of June.
DenJroica townsendi, Townsend's Warbler"". — Uncommon migrant. Seen through the second week
of May.
DenJroica graciae, Grace's Warbler'. — Very common in ponderosa pine forest. One nest with female
incubating was found on 18 May 1965. A stub-tailed juvenile was taken on 21 June 1964.
Oporornis tolmiei, MacGillivray's Warbler"'. — ■ Common migrant. Seen in desert and woodlands into
the third week of May. One captured and released at the Station on 15 May 1967.
Geothlypis tricbas, Yellowthroat"'. — One seen on station grounds on several occasions in May 1965.
CarJellina rubrifrons, Red-faced Warbler". — Uncommon in coniferous forests and higher riparian
communities. One nest found on 7 July 1965 in spruce-Douglas fir forest below Flys Peak contained
four young.
Wiho?iia pusilla, Wilson's Warbler. — Common near the Station in mid-May 1967.
Setopbaga picta, Painted Redstart7. — Abundant in South Fork of Cave Creek Canyon. One nest
located on 15 May contained four eggs. Juvenile just out of nest taken on 3 1 May 1964.
Passer domesticus, House Sparrow. — Common at Portal. One nest found in old woodpecker cavity.
Sturm lla magna. Eastern MeadowlarkT. — Common in grasslands below Apache.
Xantbocepbalus xanthocephalus, Yellow-headed Blackbird"'. — Rare at water tanks in higher desert
scrub in July and August.
Icterus cucullatus, Hooded Oriole'. — Uncommon in desert and oak woodland. One pair, later col-
lected, was observed building a nest in a large Cottonwood one and one-half miles east of Portal on
8 June 1964. Another nest was found in oak woodland in late June 1965.
Icterus parisorum, Scott's Oriole7. — Uncommon in desert but common in oak woodland and on dry
slopes. Five nests found in 1965. All were placed on the side of a yucca under the green leaves. Nest
building was underway by 15 May and young were off the nest by 21 June. An independent juvenile
was taken east of Portal on 28 June 1964.
Icterus bullockii, Bullock's Oriole"'. — Uncommon at Station and lower elevations where water is
present. One nest was found on station grounds in June 1965. Two males were captured and released at
the Station on 14 May 1967.
Euphagus cyanoccpbalus, Brewer's Blackbird'". — Not uncommon in large flocks at cattle feeder lots
until the middle of May.
Molothrus ater. Brown-headed Cowbird'. — Uncommon from desert up through oak-juniper wood-
land. A recently-fledged juvenile that was attended by a female Vermilion Flycatcher was taken 28 June
1964. Three cowbird eggs were found in a Brown Towhee (Pipilo fnscus) nest on 11 June 1965.
Tangai ins aenens, Bronzed Cowbird""7. — Uncommon from desert up to lower edge ot the oak wood-
land. Three taken on 26 May 1964. One male was molting. A Hooded Oriole nest contained two oriole
eggs and one of this species.
Piranga ludoviciana, Western Tanager. — Common in ponderosa pine forest, lev-, common in high
riparian. Abundant at hummingbird feeders at the Station in mid-May 1967, apparently as a result of
severe food shortage.
Piranga flat a, Hepatic Tanager7. — Uncommon in pine-oak woodlands, upper riparian, and ponderosa
pine forest. One nest was located high in a pine on 8 June 1965. The young fledged about 28 June.
Piranga rubra, Summer Tanager"'. — Rare in cottonwoods and high riparian woodlands. One singing
male was present at the Station from late May through June 1965.
RicbmonJena carjinalis. Cardinal. — Common in lower riparian.
Pbeucticus melanocephahis. Black-headed Grosbeak1'. — Common in oak and pine-oak woodlands and
all riparian situations, but uncommon in ponderosa pine forest. Young were seen in mi
48 San Diego Society of Natural History Vol. 15
Guiraca caerulea, Blue Grosbeak""'. — Uncommon in thick mesquite on the desert east of Portal. One
of two adult males collected on 3 June 1964.
Passcrina amoena. Lazuli Bunting. — Pair seen at the Station during the second week of May 1965.
CarpoJacus cassinii, Cassin's Finch"". — Rare late spring migrant.
Carpodacns mexicanus. House Finch. — Common at low elevations.
Spinus pinus, Pine Siskin. — Seen about the Station in May.
Spinas psalfria. Lesser Goldfinch. — Uncommon along willow thickets in open riparian habitats.
Loxia curvirostra. Red Crossbill. — Sporadic in occurrence in 1965, but flocks of up to 50 birds,
including many juveniles, were seen in pine-oak-juniper woodland and ponderosa pine in May and June.
Seen in 1964, 1965, and 1967.
Chlorura chlorura, Green-tailed Towhee. — Seen through May in open riparian and pine-oak-juniper
woodland. One was trapped and released at the Station on 24 May 1967.
Pipilo er \ t hropbt bal m us , Rufous-sided Towhee. — Uncommon but regular from oak woodland through
spruce-Douglas fir forests.
Pipilo fuscus, Brown Towhee1. — Uncommon in all lower elevation shrubby habitats.
Calamospiza melanocorys, Lark Bunting"'. — Large flocks were observed in the lowland shrub and
oak communities until the second week of May. The last individuals left during the last week of May.
Passerculus sandwichensis, Savannah Sparrow"'. — Seen commonly in vicinity of cattle watering tanks
into the first week of May.
Am m od ram us savannarum, Grasshopper Sparrow*t. — Uncommon on desert grassland between Apache
and Douglas. Bob-tailed fledglings seen on 26 July 1965. Male with enlarged testes taken on 30 June 1964.
Ammodramus bairdii, Baird's Sparrow*. — Winter visitor. Uncommon on desert grassland into early
May.
Chondestes grammacus, Lark Sparrow7. — Uncommon in shrub-grassland areas but common in open
oak-woodland where three nests, all on the ground, were located. A nest found on 17 June contained
one egg as did another located on 23 June. On 1 July, a pair was observed building a nest which con-
tained three eggs eight days later. These are exceptionally late breeding records, as Brandt (1951) found
nests in May and early June.
Aimophila ruficeps, Rufous-crowned Sparrow. — Uncommon to common in grassy woodlands.
Aimophila cassinii, Cassin's Sparrow"'. — Rare, possible breeder on the grasslands south of Apache.
Adult male with enlarged testes taken six miles south of Apache on 30 June 1964. Ohmart (1966)
recently discovered A. cassinii breeding in Arizona (cf. statements of Phillips et al., 1964: 200).
Amphispiza bilineata, Black-throated Sparrow^. — Abundant in desert scrub vegetation. Nest with
three eggs found one and one-half miles east of Portal on 29 May 1964. Four nests found in 1965; the
earliest was under construction on 12 May, whereas the latest held two eggs on 14 June.
Jiinco oreganus, Oregon Junco*. — Last seen on 15 May 1965 in pine-oak-juniper woodland and
ponderosa pine forest.
Jiuico caniceps, Gray-headed Junco*. — Migrant. Last seen on 17 May 1965, in pine-oak-juniper
woodland.
Junco phaeonotus, Mexican Junco'''. — Abundant in coniferous forest. Ten nests found there in 1965.
The earliest nest was being built on 17 May; the latest had three eggs on 1 August.
Spizella passcrina, Chipping Sparrow'. — Common in oak woodland and abundant in open pine-oak-
juniper woodland. Seven nests were found in the latter community in 1965. The earliest nest contained
four eggs on 17 May, while the latest contained young on 12 June.
Spizella breueri, Brewer's Sparrow*. — Large flocks were present in low elevation shrubland until
the end of May.
Spizella atrogularis, Black-chinned Sparrow"'. — One seen at lower edge of oak woodland on
5 May 1965.
Zonotrichia leucopbrys. White-crowned Sparrow. — Common migrant and winter resident at low
elevations.
Melospiza lincolnii, Lincoln's Sparrow*. — One seen at the Station during the first two weeks of
May 1965. Two seen at the Station in mid-May 1967.
Discussion
We recorded 167 species, of which 5 7 were not seen by Tanner and Hardy (1958).
(Nineteen species recorded by us were winter residents or migrants not seen after May.)
This is hardly surprising, as we spent much more time in this region, both within a
single year and over a span of several years. Species for which we have breeding records,
1968 Ligon and Balda: Chiricahua Birds 49
or for which there is reason to assume breeding, that were unrecorded by Tanner and
Hardy include: Peregrine Falcon, Harlequin Quail, Screech, Whiskered, Great-Horned,
Pygmy and Elf owls, Wied's Crested Flycatcher, Buff-breasted Flycatcher, Barn Swal-
low, Scrub Jay, Curved-billed Thrasher, Audubon's Warbler, Bullock's Oriole, Bronzed
Cowbird, Blue Grosbeak and Grasshopper Sparrow.
Of special interest are those species whose status apparently has changed during
the recent past. The Golden Eagle appears to have decreased since the studies of Brandt
(195 1) and Tanner and Hardy (1958), whereas the Harlequin Quail apparently has
increased. The Prairie Falcon may have become rarer, judging from the comments of
Phillips ct al. (1964: 26), as a result of movement of the Peregrine Falcon into south-
ern Arizona. Our probable breeding record of the Peregrine in Cave Creek Canyon
agrees with this; Tanner and Hardy recorded only the Prairie Falcon there. The hum-
mingbirds have either increased in numbers or in conspicuousness or both, largely as a
result of the many birders, both inside and outside the station grounds, who put up
hummingbird feeders. The Violet-crowned Hummingbird has extended its range north-
ward, as pointed out by Phillips ct al., (1964: 65) and is now regular at the Station.
Neither Brandt (1951) nor Tanner and Hardy (195 8) recorded the Plain Titmouse
and Phillips ct al. (1964: 111) report it only from the northern Chiricahua foothills.
We found it using natural cavities throughout the woodlands. The Coppery-tailed
Trogon and Buff-breasted Flycatcher apparently occur irregularly in southern Arizona,
and we were fortunate to be in the area in a year (1965) when both were present; the
trogon was actually almost common.
Tanner and Hardy listed many cavity nesters as uncommon or rare. This
was not borne out by the census data of Balda. The density of the Western Bluebird was
20 pairs per 100 acres in ponderosa pine forest. The Red-breasted Nuthatch also was
listed as rare in 1956, but 2 3 pairs per 100 acres were recorded in spruce-fir forest. This
suggests a recent increase in the abundance of cavity nesters. Severe drought conditions
have killed large numbers of trees in the past few years, indirectly providing numerous
new nest sites for these cavity nesters. As mentioned earlier, the Eastern Bluebird
(apparently S. 5. fulva), another cavity nester, has also become established recently in
areas near the Station. Balda found the density of this species to be 3 pairs per 100 acres
in 1964 and 7 pairs per 100 acres in 1965. Ligon (in press) describes nest sites of this
species in Cave Creek Canyon and mentions some factors which possibly serve to limit
its increase.
Acknowledgments
In 1964 Ligon received financial aid, in the form of a summer training grant, from the United States
Public Health Service and from a grant provided by the Josselyn Van Tyne Memorial Fund of tin-
American Ornithologists' Union. Both Ligon and Balda received grants from the Frank M. Chapman
Memorial Fund of the American Museum of Natural History in 1965, supporting independent research in
the Chiricahuas. Ligon also received assistance from a grant to The University of Michigan Museum of
Zoology from the National Science Foundation Training Program in Systematica and Evolutionary Biol) gj
(GB-3 366), and Balda's work was in part supported by the University of Illinois Research Board.
Literature Cited
Balda, R. P.
1967. Ecological relationships of the breeding-birds of the Chiricahua Mountains. Arizona. L'npubl.
Ph.D. thesis, University of Illinois.
Brandt, H.
195 1. Arizona and its bird life. The Bird Research Foundaticn. Cleveland.
Ligon, J. U.
Factors influencing breeding range expansion of the Eastern Bluebird (Sialia sialis). Wilson Bull..
In press.
50 San Diego Society of Natural History Vol. 15
Ohmart, R. D.
1966. Breeding record of the Cassin Sparrow (Aimophila cassinii) in Arizona. Condor, 68: 400.
Peterson, R. T.
1948. Birds over America. Dodd, Mead, and Co. New York.
Phillips, A. R., J. T. Marshall, Jr., and G. Monson
1964. The birds of Arizona. Univ. Arizona Press, Tucson.
Tanner, J. T., and J. W. Hardy
1958. Summer birds of the Chiricahua Mountains, Arizona. Amer. Mus. Novitates. 1866.
Whittaker, R. H., and W. A. Niering
1965. Vegetation of the Santa Catalina Mountains, Arizona: A gradient analysis of the south slope.
Ecology, 46: 429-452.
Accepted for Publication 11 January 1968
Department of Biology, Idaho State University, Pocatello, Idaho, and Department
of Biological Sciences, Northern Arizona University, Flagstaff, Arizona.
^ /V/j'
A BIOLOGICAL SURVEY OF BAHIA DE LOS
ANGELES, GULF OF CALIFORNIA, MEXICO. I
GENERAL ACCOUNT. ^H
J. LAURENS BARNARD AND JOHN R. GRADY
Mue, ce*i* zool.
LffRARY.
JUL 6T968
HARVARD
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 6, 17 JUNE 1968
113° 30
Bahia de Los Angeles
113° 30'
Figure 1. Plan of hydrographic stations in Bahia de Los Angeles; inset, location of bay
in Gulf of California.
A BIOLOGICAL SURVEY OF BAHIA DE LOS ANGELES,
GULF OF CALIFORNIA, MEXICO. I.
GENERAL ACCOUNT
J. Laurens Barnard and John R. Grady
Abstract. — -The benthic invertebrate fauna of the southern end of Bahia de los Angeles, a small bay
on the Gulf of California, Mexico, was sampled during April 1962 and October 1963. 71 stations
were occupied in 1962 and 91 stations in 1963. Invertebrate samples were collected with an orange-
peel grab. The sediments on the bottom of the bay are predominantly brown, micaceous coarse silts
and fine sands; the steep fringes are composed of gray, fine and medium sands on the shoreline facing
the source area, a granite massif; the shore and fringing shelves on the peninsula and barrier islands
fronting the bay are primarily shell fragments and rocks. There is free water circulation in the bay,
although slight hypersaline conditions exist in summer. Dissolved oxygen ranges from 3.0 to 7.0 -f-
ml/L in surface waters and drops to 1.66 ml/L in bottom waters of 75 to 98 m. Low level red-
tides occur in March and April, and these may affect the invertebrate fauna. Lunar tides are of a
mixed semi-diurnal type. Because winter hydrotemperatures drop below 15C the bay has the aspect
of a warm-temperate province rather than a purely tropical region. Large scale reductions in the
invertebrate fauna occurred prior to the start of the survey and between the April 1962 and October
1963 survey. The cause of these reductions is still unknown. Detailed reports on the fauna will
appear elsewhere in this series.
Resumen. — En Abril de 1962 y Octubre de 1963 se obtuvieron muestras de invertebrados bentonicos
en el extremo meridional de Bahia de los Angeles, Baja California, Mexico. En total se ocuparon
71 estaciones en 1962, y 91 en 1963. Los sedimentos del fondo de esta bahia son predominantemente
de color pardo, micaceos, con fango de particulas gruesas y arenas finas. Los bordes pendientes
estan constituidos por una franja de arenas grises de grano medio y fino, en la banda costera
frente a un macizo granitico; de modo que la costa y las plataformas de la ribera de la peninsula
y las islas que forman una barrera a la entrada de la bahia, estan formadas primordialmente de
fragmentos de rocas y conchas. El agua circula libremente en esta bahia, aun cuando en el verano se
presentan ligeramente hipersalinas. El oxigeno disuelto fluctua de 3.0 a 7.0 -\- ml/1 en las aguas
superficiales, descendiendo a 1.66ml/l en las aguas del fondo, a los 75 a 98m de profundidad. Mareas
rojas de poca intensidad se presentan en Marzo y Abril, las cuales posiblemente afectan a esta fauna
de invertebrados. Las mareas lunares son una mezcla del tipo semidiurno. La temperatura del agua
dcsciende en el invierno por debajo de los 15C de modo que esta bahia pertenece mas bien a una
provincia calido-templada, que a una region puramente tropical. Esta fauna de invertebrados sufrio
una reduccion en gran escala antes de iniciar estos estudios y entre los periodos de Abril, 1962, y
Octubre de 1963. Se desconocen las causas que motivaron tales bajas. Un estudio detallado de esta
fauna aparecera en otra parte de estas series.
A study of the marine environment and invertebrate fauna of Bahia de los Angeles,
by the writers and their colleagues, was suggested by the symposium on "The Bio-
geography of Baja California and Adjacent Seas" (Systematic Zoology, vol. 9, pts. 1-3,
1960), and made possible by the establishment of the Vermilion Sea Field Station of the
San Diego Natural History Museum at Bahia de los Angeles. Our desire to study un-
disturbed shallow-water areas have also prompted this study, which followed guidelines
similar to those used in the biological survey at Bahia de San Quintin (Barnard, 1962).
Because quantitative biological studies in a small area in the Gulf of California have
not been made previously, the present survey has concentrated primarily on the occur-
rence, distribution, and density of benthic, soft-bottom, invertebrate populations. A
study of sedimentary structure, bathymetry, and a bi-seasonal assessment of a few hydro-
graphic parameters were made concurrent with the biological survey. This focus on a
small community comes as an extension of the exploratory work of Parker (1964), who
assembled data on the Gulf biotopes based primarily on the molluscan fauna.
San Diego Soc. Nat. Hist., Trans. 15 (6): 51-66, June 1968
J4
San Diego Society of Natural History
Vol. 15
*g*<^».
Figure 2. Aerial photograph of Bahia de Los Angeles.
Figure 3. Portion of village at Bahia de Los Angeles.
1968 Barnard and Grady: Bahia de los Angeles 5 5
Human influence on the bay environment has probably been of little consequence.
The hunting of sea turtles (Chelonia mydas carrincgra) from small boats in the open
Gulf provides the economic base for the small village situated on the central western
shore of the bay. Fishing for the totuava (Cynosdon macdonaldi) appears to be more
important to the north, at Bahia San Luis Gonzaga and at San Felipe, than at Bahia dc
los Angeles. The main tourist attraction is fishing, although there is some deer hunting
in the winter months. Access to the area is largely by small private plane, although
there are scheduled flights from Tijuana.
Physiography
Bahia de los Angeles is situated on the north east shore of Baja California. It is
fronted at some distance off shore by Angel de la Guarda Island and closer, by numerous
small islands (Figs. 1, 2). There are three main, deep channels into the bay.
Protection from easterly winds is rather good, although severe winds can create
large waves passing westward through the channels between islands. A granite massif,
part of the Sierra de San Borjas range, flanks the west shore, (Fig. 3) and apparently
deflects the northerly and westerly winds into winds with primarily westerly and
southerly components as they pass across the shore (data from wind gauge established
in April, 1962). At times, southerly and northwesterly winds meet on the sea in front
of the village and create small water spouts.
The barrenness of the mountains and islands surrounding the bay results in a stark
appearance. Sand or gravel beaches typify the western and southern shores, whereas the
southeastern peninsula and most of the islands are beset by only cliffs and gravel ram-
parts. Despite the ruggedness of the subaerial topography the southern arm of the bay
is remarkably flat (Figs. 4, 5 ) .
The floor of the bay is in the form of a basin. It is tilted slightly west to east, and
on the eastern shore is bounded by a small trench 1-2 meters deep. The basin shoals
gradually to the south, where the initial dip is about 1 30', but is reduced to 0°24'
from there to the geographic center of the southern portion of the bay, where depths
are approximately 40 meters. Slopes on the east side, supported by volcanic extrusives,
run to over 16°, in contrast to those off the sand beaches on the west side which are
generally little over one degree.
Field Methods
Hydrography. — Hydrographic surveys were conducted in April 1962, April 1963,
and October-November 1963. Their timing coincided with the benthic biological sur-
veys of April 1962 and October-November 1963. The April 1963 hydrographic survey
acted as a control on year-to-year variability. The benthic surveys were planned to lag
about two months behind the expected extreme water temperatures of mid-winter and
mid-summer, in order to sample the accumulated results of the fauna's exposure to
winter and summer seasons. Hence, the hydrographic surveys in both seasons represent
approximately intermediate thermal conditions.
Hydrographic stations were established throughout the bay (Fig. 1), and each grid
station was occupied for a two-day period. At each station the following parameters
were studied at water depths of 0, 5, 10, 20, 30, 40, and 50 meters: salinity, temperature,
dissolved oxygen, pH, silicates, phosphates, and water turbidity. Temperature was
recorded on paired protected-unprotected reversing thermometers attached to Nansen
bottles for correlation with other water analyses. Thermoclines were studied with .i
200-foot bathythermograph. A 24-hour sampling program was conducted at anchor to
study relationships of water variables to tidal changes.
A portable tide gauge was erected at a depth of 1.3 meters at the lowest observed
tide, and was maintained from 21 to 28 April, 1962, 13 to 21 April. 1963, and
S6
San Diego Society of Natural History
Vol. 15
113° 30'
ROCKS EXPOSED AT LOW
TIDE OR PINNACLES
SOUNDINGS IN METERS
CONTOUR INTERVAL'
10 METERS
NAUTICAL MILES
KILOMETERS
113° 30'
I igure 4. Bathymetry of Bahia de Los Angeles based on echo-sounding tracks of inset.
1968
Barnard and Grady: Bahia de los Angeles
57
Figure 5. Depth contours and benthic biological stations in south portion of Bahia de
Los Angeles.
2 5 October to 3 November, 1963. The gauge was attached to 5 meters of Orangeburg
pipe suspended from a metal tripod.
A Simrad recording fathometer aboard the "Neptunus Rex" was used to record
38 profiles, totalling about 250 km. (Fig. 4), with radar navigation based on the chart
prepared by Neil F. Marshall and Wendell R. Gayman of Scripps Institution of Ocea-
nography on a scale of 1:2000. Radar navigation was checked continuously by pclorus
readings from the ship's gyroscope system. Tidal observations were not used for adjust-
ing profile records of the bathymetric survey precisely to datum, as other Inaccuracies
were of greater magnitude.
Biology. — A grid of benthic sampling stations was set up in the southern, m
protected portion of the bay, using procedures described by Barnard and Jones ( I960)
and Barnard (1962). Approximately 15 stations per square mile (nautical) were estab-
lished, totaling 161 stations in an area about ten square miles (Fig. 5). Alternate
stations were to be occupied in two seasons, the respective groups of 80 and 81 samples
58
San Diego Society of Natural History
Vol. 15
APRIL 13
NOVEMBER
OCTOBER 25 26 27 28 29 30 31 I 2
12
08
04
0
04
CO
2j 08
1
BAHIA DE LOS ANGELES
UJ
12
08
04
MAZATLAN
04
08
I 0
Figure 6. Marigrams of Bahia de Los Angeles compared with those of Mazatlan
and Guaymas.
1968 Barnard and Grady: Bahia de los Angeles 59
being of sufficient size and replicative variability to represent a matched pair, thus per-
mitting a seasonal survey. If seasonal biotic differences proved to be minor, a better
mapping of bottom areas would result than if 80 stations were duplicated in the two
seasons. Owing to technical difficulties in April 1962 several stations were not occupied,
and several samples were not recovered because of failure of the grab to penetrate the
bottom. However, these stations were reoccupied in October-November 1963, resulting
in groups of 71 (April) and 91 (October) samples. This departure from plan was less
serious than expected. Seasonal differences were found to be so extreme that precise
comparison was not believed to be necessary at this time.
Benthic biological samples were obtained with a Hayward orange-peel grab modified
with canvas skirt and single-cable closing features. The sediments were washed through
a Tyler Standard Screen Scale Sieve of 0.7mm mesh opening. The residues were preserved
in formaldehyde, sorted in the laboratory, and subsequently preserved in alcohol. After
gross examination of dominant organisms the sorted samples were forwarded to special-
ists for study. Detailed faunal reports based on these collections appear elsewhere in
this series.
Sediment samples were collected at each station for determination of grain size
(median diameter), organic carbon, calcium carbonate contents, and pH at each station.
Water clarity readings were made with a Secchi disc, and surface-water temperatures
with a centigrade thermometer in bucket samples, as well as with a Yellow Springs
model 43TJ thermistor. Hydrographic casts with Nansen bottles and reversing ther-
mometers, and bathythermograph tracings for thermoclines were taken at every second
or third station during the biological survey.
Bottom depths from sea-surface were taken from the fathometer, checked by lead
soundings, and corrected subsequently by construction of a bathymetric chart approxi-
mately to tidal datum.
The various parameters of the stations sampled are summarized in the Appendix.
Physical Observations
Bahia de los Angeles is open sufficiently to Ballenas Channel to permit free water
circulation. Salinities in all three hydrographic surveys were in normal ranges for the
upper Gulf, showing very slight hypersaline conditions in the upper 10 meters owing to
evaporation during summer months. According to Roden and Groves (1959) salinity
values in Ballenas Channel range from 35.2 to 35.3','. In the Bay in April 1963 we
recorded a surface salinity of 35.259^ and at a depth of 30 meters 35.10%; in October
1963 surface salinities were approximately 35.55% and at 30 meters 35.45', .
Water temperatures in the bay are apparently extreme for a warm-temperate lati-
tude. Not many data have been taken, but those records available show minima of less
than 15C in February 1962 and 1963 and maxima of 29. 8C in August 1962, 28. 5C in
August 1961 and 28. 7C in September 1963.
Dissolved oxygen in the bay at depths of 5 0 meters or less always exceeded 3.0 ml 1
in waters from 10 meters above the bottom to the surface. Occasionally, in the bottom
waters from 0 to 10 meters above, in depths exceeding 30 meters, the dissolved oxygen
values were between 2.0 and 3.0ml L. Bottom water in depths of 7 5 to 98 meters
dropped to a low of 1.66 ml L. No attempt was made to sample at greater depths. The
water between the surface and 30 meters was consistently better oxygenated, being is
much as 3.0 ml L higher in spring than in fall. In April 1962 dissolved oxygen at 10
selected stations in 2 5 meters depth ranged from 4.13 to 7.16 ml L, whereas in October-
November 1963 the range in 20 meters was between 4.0 and 5.0 ml L. These differences
presumably result from winds in the spring in addition to other factors.
60
San Diego Society of Natural History
Vol. 15
NAUTICAL MILES
bOO IDOO 1500
Sediment Types
METERS
Figure 7. Sediment distribution in south portion of Bahia de Los Angeles.
Bahia de los Angeles lies in the middle of more than 100 miles of coastline known
for extensive upwelling (Byrne and Emery, 1960; chart compiled in Parker, 1964),
which enriches surface waters and permits plankton blooms. Small but dense patches of
red-tide were observed almost continuously during April of 1962 and 1963, and during
a reconnaissance trip in March 1961.
Marigrams show the tides to be of a mixed semidiurnal type which, during the
periods measured, had a mean range of 1.22 meters and a high water interval of slightly
more than 12 hours (Fig. 6). The maximum range recorded in April 1962 and 1963
was 2.2 meters, and in October-November 1963, 2.0 meters; maximum spring tides did
not occur during these periods. The tides exceed the spring range, which increases
progressively toward the head of the Gulf, of 0.8 meters at the reference tide station at
Guaymas, and 1.3 meters at Mazatlan. In Figure 6, the marigrams for April are con-
trasted with tidal curves from Guaymas, and for the period in October-November with
a plot of the predicted tides at Mazatlan computed from the reference station at San
Diego, California. The mixed tides of Bahia de los Angeles resemble the semidiurnal tides
1968 Barnard and Grady: Bahia de los Angeles 61
at Mazatlan just south of the entrance to the Gulf, more than they do those at Guay-
mas, Bahia Concepcion or La Paz, which are characterized by dominant diurnal
components.
The sediment pattern (Fig. 7) in the bay apparently conforms with water depth.
Gray sands occur on the western and southern shores from the beach seaward to depths
of 20 to 2 5 meters, grading into silty sand and sandy silt at 20-3 5 meters. Brown silt
forms a cover in depths exceeding 3 5 meters. A belt of black silt mixed with sand also
is present along the south margin of the bay (stations 1 to 8). Seaward of the black
sediment is a region of gray, medium and fine sands. The eastern shore is dominated by
sands and silty sand with a high content of shell particles, rocks, and coarse debris. Fine
sediments apparently have their source on the western and southern margins of the bay,
whereas shell fragments remain exposed on the eastern margin where little sediment is
supplied from the land. Barrier islands are also surrounded by shell fragments. The
abundance of mica in the marine sediments probably indicates the chief source of sedi-
ment to be the southern granitic intrusive.
Biological Observations
Benthic samples of April 1962 included large and diverse populations of polychaetes,
small pelecypods, amphipods, mysids and small shrimps on muds in depths exceeding 5m.
Grab samples on sand bottoms of 2-5m depth fringing the bay were poorly populated,
but dredgings accumulated large masses of red algae and numerous crustaceas. Mud
bottom samples taken in October-November 1963 appeared highly impoverished and
the paucity of organisms was confirmed by frequent utilization of a Van Veen grab in
which no losses of mud could occur. (The Van Veen samples are not included in the
results of this survey.) Thus, a severe reduction of the benthic fauna had occurred
between the sampling periods. Seven benthic samples from Bahia Concepcion (150 miles
southeast of Bahia de los Angeles) in early November 1963 yielded less than 10 macro-
scopic organisms (polychaetes, nemertean, crabs, stomatopod). This fauna, therefore,
was even more impoverished than that of Bahia de los Angeles.
The diminutive size of benthic pelecypods in depths of 10-40m in April 1962
suggests that another reduction had occurred in 1961. In late 1963, population densities
of many amphipods and polychaetes were less than half those of early 1962. Samples
from October 1963 showed many shell fragments but few whole shells of pelecypods.
If benthic predators or scavengers (elasmobranchs) had pulverized these shells, a re-
markable efficiency would have to be assumed.
Bottom waters of Bahia de los Angeles and Bahia Concepcion showed no reduction
in dissolved oxygen and no trace of hydrogen sulphide was found in water or muds
during the several surveys. Small patches of red tide were observed in March-April of
1961, 1962, and 1963, and Dr. D. J. Reish has informed us that severe biotic depletion
occurred in Alamitos Bay, California, during and after periods of red-tide. But the red-
tide so far observed at Bahia de los Angeles has been minor. The cause of faunal reduc-
tion is still unknown. However, windrows of large pelecypods and gastropods on the
southwest shore of Bahia de los Angeles suggests that severe depletion of bottom popula-
tions is not an annual phenomenon.
Acknowledgements
This project was supported by matched funds from the National Science Foundation (G-20
and the Beaudette Foundation of California. The laboratory work was completed under auspices of the
Smithsonian Institution. Richard F. Dwyer of Los Angeles generously provided his 6 J -foot convene
shrimp-trawler "Traveler" for the first expedition, and the Beaudette Foundation, through John P.
McNabb of Los Angeles, provided the 97-foot North Sea Trawler "Xeptunus Rex" for the second and
third expeditions. The San Diego Museum of Natural History, through its former director Dr. Geoi
E. Lindsay, provided quarters and work space at the NSF-suported Vermilion Sea Field Station. Dr. and
62 San Diego Society of Natural History Vol. 1 5
Mrs. Carl L. Hubbs generously donated numerous specimens of marine invertebrates collected by their
SCUBA teams and thus provided representatives of species we would not have obtained otherwise. The
U. S. Coast and Geodetic Survey kindly donated a portable tide gauge.
The success of this project is due directly to the generosity of the following persons, each of whom
supported it financially and each of whom untiringly worked in the field: Palmer T. Beaudette, John P.
McNabb, and Richard F. Dwyer.
For various courtesies extended we are grateful to: Allan J. Stover, Jr., James R. Stewart, Donald M.
Dockins, Dr. Richard H. Rosenblatt, Keith W. Radford, Dr. Wayne J. Baldwin, John H. Emmel, David
W. Hearst, W. W. Shannon, Lloyd R. Hales, Jr., and Fillmore T. Condit.
Literature Cited
Barnard, J. L.
1962. Benthic marine exploration of Bahia de San Quintin, Baja California, 1960-61. General.
Pacific Natur. 3: 249-274.
Barnard, J. L. and G. F. Jones
1960. Techniques in a large scale survey of marine benthic biology, P. 413-447. In, E. A. Pearson
[ed.] Waste Disposal in the marine environment. Pergamon Press.
Byrne, J. V. and K. O. Emery
1960. Sediments of the Gulf of California Bull. Geol. Soc. Amer. 71: 983-1010.
Parker, R. H.
1964. Zoogeography and ecology of some macroinvertebrates, particularly mollusks, in the Gulf of
California and the continental slope off Mexico. Vid. Medd. Dansk Nat. Foren. 126: 1-178.
Roden, G. I. and G. W. Groves
195 9. Recent oceanographic investigations in the Gulf of California. Sears Found., Jour. Mar.
Res. 18: 10-35.
Accepted for Publication 26 March 1968
Smithsonian Institution, Washington, D.C., and Bureau of Commercial Fisheries,
Fort Crockett, Texas.
1968 Barnard and Grady: Bahia de los Angeles 63
Appendix
Table 1. Biological Stations in Bahia de los Angeles. L = sample taken from launch.
Sediment types in parentheses indicate changes from field classification owing
to laboratory determinations of sand-silt percentages. Depths represent cor-
rected values from plots of bottom contours adjusted approximately to
tidal datum.
Station Date Depth (m) Temperature (C) Sediment
gray-black sand with silt
gray-black sand with sill
black very fine sand
black very fine sand
black very fine sand
black very fine sand
black very fine sand
gray sand
gray sand
gray-green fine sand
gray-green fine sand
gray-green silty sand
gray-green sand
gray-green silty sand
gray sand
gray silty sand
gray fine sand
hard bottom, no sample
shell sand
gray-green fine sand
gray very fine sand
brown fine sand
gray very fine sand
gray-green silty sand
gray silty sand
brown silty sand
gray silty sand
greenish shelly silty sand
shell sand
greenish fine sand
gray-green very fine sand
gray-green fine sand
gray-green very fine sand
gray-green silty sand
brown silty sand
greenish-brown silty sand
brown silty s.md
hard bottom, no s.imple
brown siltv s.md
brown silty sand
gray medium sand
gray sand
medium mIi \ i s.md)
silty i sand I
silty sand
brc wn silt and tine sand
brown silty very fine sand
brown silty sand
brown sandy silt
brown silty sand
brown silty sand
1 L
10-30-63
2 L
10-30-63
3 L
10-30-63
4 L
10-30-63
5 L
10-30-63
6 L
10-30-63
7 L
10-30-63
8 L
10-30-63
9 L
10-30-63
10
4-24-62
1 1
10-28-63
12
4-24-62
13
10-28-63
14
4-24-62
15
10-28-63
16
4-24-62
17
10-28-63
18 L
4-24-62
10-31-63
19
4-24-63
20
10-28-63
21
4-24-62
22
10-28-63
2}
4-24-62
24
10-28-63
25
4-24-62
26
10-28-63
27
4-24-62
28 L
10-31-63
29
4-24-62
30
10-28-63
3 1
4-24-62
32
10-28-63
3 3
4-24-62
34
10-28-63
3 5
4-24-62
36
10-28-63
37
4-24-62
10-28-63
38
10-28-63
39
4-24-62
40 L
10-31-63
41
4-27-62
42
10-28-63
43
4-27-62
44
10-28-63
45
4-27-62
46
10-28-63
47
4-27-62
48
10-2 8-6 3
49
4-27-62
Surface Water
ith (m)
Temperature (C)
3
10
14
7
13
12
11
4
2
17
19.7
18
25.2
15
19.7
16
25.5
15
19.4
15
25.6
13
19.9
11
25.8
5
20.2
19
19.1
22
25.6
21
18.9
22
25.7
20
19.2
20
2 5.7
17
19.2
18
2 5.7
13
20.0
4
17
19.9
24
25.9
25
18.1
26
26.0
25
18.0
24
25.9
24
18.7
24
25.9
20
20.4
20
26.2
20
26.3
7
21.6
4
21
18.1
27
2 6.4
30
18.1
30
26.3
30
17.6
29
26.5
25
17.9
24
26.2
20
17.9
64
San Diego Society of Natural History
Vol. 15
Table 1. (Continued)
Station
50
51
52 L
53
54
55
56
57
58
59
60
61
62
63
64
65 L
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102 L
103
104
105
106
Date
10-28-63
4-27-62
10-31-63
4-27-62
10-28-63
4-27-62
10-28-63
4-27-62
10-28-63
4-27-62
10-28-63
4-27-62
10-28-63
4-27-62
10-28-63
10-31-63
4-27-62
10-29-63
4-27-62
10-29-63
4-27-62
10-29-63
4-27-62
10-28-63
4-27-62
10-28-63
4-27-62
10-28-63
4-27-62
10-29-63
4-27-62
10-29-63
4-27-62
10-29-63
4-27-62
10-29-63
4-27-62
10-29-63
4-27-62
10-29-63
10-29-63
4-28-62
10-29-63
4-28-62
10-29-63
4-28-62
10-29-63
4-28-62
10-29-63
4-28-62
10-29-63
4-28-62
10-31-63
4-28-62
10-29-63
4-28-62
10-29-63
Surface Water
(m)
Temperature (C)
18
26.2
15
18.7
9
22
18.2
31
26.4
31
18.2
31
26.4
31
18.1
31
26.5
31
17.9
30
26.3
24
18.0
24
26.2
21
18.3
22
26.2
9
25
18.3
27
24.9
33
18.1
33
25.3
33
18.0
31
25.1
30
17.4
31
26.1
30
17.7
3 0
26.0
28
17.3
27
26.0
17
17.5
27
2 5.4
34
17.9
3 5
25.4
3 5
17.6
3 5
24.3
36
17.6
37
2 5.4
34
16.9
3 3
25.2
3 3
17.3
3 3
25.5
9
25.8
34
17.7
3 5
25.8
3 6
17.5
38
25.5
3 8
17.3
3 9
25.5
37
17.8
3 8
25.5
37
18.2
37
25.6
37
18.2
6
30
18.1
38
26.0
3 8
17.1
40
26.0
Sediment
brown silty sand
gray medium sand
gray sand
gray medium sand
brown silty (sand)
silty fine sand
silty (sand)
very fine sand
brown silty (sand)
brown silty sand
brown sandy silt
silty very fine sand
brown sandy silt
silty very fine sand
very fine sand
coarse shell sand
silty medium sand
gray fine silty sand
silty sand
brown silt
brown sandy silt
brown sandy silt
brown sandy silt
brown silt
sandy silt
brown silt
sandy silt
very fine sand
gray medium sand
gray silty fine sand
brown sandy silt
brown silt
brown sandy (silt)
brown silt
greenish-brown sandy silt
brown silt
brown silt
brown silt
brown silt
brown sandy silt
gray sand
brown silty sand
brown sandy silt
brown sandy silt
brown silt
brown silt
brown silt
brown silt
brown silt
brown silt
brown silt
silty sand with shells
gray sand
brown silty (sand)
brown silt
brown silt
brown silt
1968
Barnard and Grady: Bahia de los Angeles
65
Table 1. (Continued)
Station
107
108
109
1 10
1 11
1 12
11)
I 14
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146 L
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
Date
4-28-62
10-29-63
4-28-62
10-29-63
4-28-62
10-29-63
4-28-62
10-29-63
10-29-63
4-24-62
10-29-63
4-22-62
10-29-63
4-22-62
10-29-63
4-24-62
10-29-63
4-24-62
10-29-63
4-24-62
10-29-63
4-24-62
10-29-63
4-24-62
10-29-63
4-22-62
10-29-63
4-22-62
10-29-63
4-22-62
10-30-63
4-22-62
10-30-63
4-24-62
10-30-63
10-30-63
4-23-62
10-30-63
4-2 3-62
10-30-63
10-31-63
10-30-63
4-23-62
10-30-63
4-23-62
10-30-63
10-30-63
4-23-62
10-30-63
4-23-62
10-30-63
4-22-62
10-30-63
4-23-62
4-23-62
10-30-63
4-23-62
10-30-63
Surface Water
(m)
Temperature (C)
40
17.3
40
26.0
40
17.1
40
26.0
40
17.6
40
25.8
40
17.9
39
26.2
37
26.3
6
19.8
30
26.4
34
20.1
40
26.2
40
19.5
42
26.1
42
17.7
42
25.2
42
17.2
42
26.2
42
17.2
42
26.1
42
18.3
18
26.2
7
19.8
24
26.5
25
19.7
35
26.2
38
19.7
42
26.2
41
19.5
42
25.4
46
18.5
42
25.6
44
17.4
2 5.4
46
25.0
46
18.1
46
24.9
44
17.9
38
25.5
6
9
25.6
15
19.5
19
25.6
29
18.9
25.0
41
25.5
42
17.4
44
25.5
44
17.4
44
25.5
46
18.3
46
25.3
46
17.7
17.4
46
25.5
45
17.9
49
25.3
Sediment
brown silt
brown silt
brown silt
brown silt
brown silt
brown silt
green sandy silt
brown sandy silt
very fine sand
gray medium sand
brown silty sand
brown sandy silt
brown silt
brown sandy silt and clay
brown silt
brown silt
brown silt
brown silt
brown silt
brown silt
brown sandy silt
brown sandy silt
reefy material
gray coarse sand, 3 poor grabs
brown silty sand
sandy silt with shells
brown silt
brown silty medium sand
brown silt
brown sandy silt
brown silt
brown silt
brown silt
no sample
brown silt
brown sandy silt
brown sandy silt
brown sandy silt
brown silty sand
fine shelly silty sand
coarse shell sand
gray fine sand
gray fine sand
gray medium sand
hard bottom, no sample
gray fine sand and shell sand
brown sandy silt
brown sandy silt
brown silt
brown silt
brown silt
brown silt
brown silt
brown silt
brown silt
brown sandy silt
brown sandy silt
silt with shell sand
66
San Diego Society of Natural History
Vol. 1J
Table 2. Samples of Scripps Institution of Oceanography, Dr. Carl L. Hubbs and party
Remarks
Piedras Ahogadas, 0.5 mi S Isla Ventana;
SCUBA sample of bottom: shell frags., pebbles
Isla Ventana, S shore of cove on W side of island:
tailings of fish sample
Isla Cerraja, E shore: tailings of fish sample
Isla Ventana, Pta. Ventana: tailings of fish sample
Reef between Isla Ventana and Isla Cabeza de Caballo:
tailings of fish sample
W shore of bay, south of village: tailings of fish sample
on cobble bottom
Cove on E side of S arm of bay: 16-foot fish-trawl
N of village sandspit: beach poisoning
S arm of bay: shrimp trawl on shrimp boat on N-S line.
Station
Date
Depth (m)
SIO-X
4-22-62
24
SIO-62-210
4-18-62
shore
SIO-62-212
4-19-62
shore
SIO-62-215
4-21-62
shore
SIO-62-216
4-21-62
shore
SIO-62-228
4-24-62
2
SIO-62-234
4-25-62
subtidal
SIO-62-2 3 5
4-26-62
subtidal
SIO-62 236
4-26-62
?30
Station
Date
Depth (m)
210
4-27-62
9
211
4-28-62
9-32
212
4-28-62
9-16
213
4-22-62
Intertidal
214
4-22-62
Intertidal
301
3-10-61
6
3 02
3-10-61
4
303
3-10-61
Intertidal
305
3-10-61
Intertidal
306
3-10-61
Intertidal
306A
3-10-61
Intertidal
3 50
1- 1-63
Intertidal
35 1
11- 1-63
Intertidal
3 52
1- 1-63
Intertidal
353
1- 1-63
Intertidal
3 54
1- 1-63
Intertidal
35 5
1- 1-63
Intertidal
910
1- 2-63 2-3
911
1- 2-63 sur
ace Plankton tow
912 1
1- 2-63 sur
ace Plankton tow
Table 3. Miscellaneous samples
Remarks
Dredge: bag 1/3 full of gracilarioid algae, few Polinices
egg-cases, one Pecten, a few bulls-eye shrimp; algae washed
for small crustaceans; a few Astropccteii
75 0m N of village sandspit, dredge; Ulia and gastropod egg
cases, spider-crabs, 1 asteroid
Main W shore of bay, dredge: Ulva, gracilarioids, 2 asteroids
Vermilion Sea Field Station, rocks at low tide
Same locality: tunicates and sponges washed for crustaceans
OPG test sample, 1 mile S of Vermilion Sea Field Station
Same
Vermilion Sea Field Station, rocks: wash of Sargassum sp.
Same locality: wash of Ulia sp.
Same locality: wash of sandy rocks
Same locality: wash of rocks below water level
Same locality: wash of 2 rocks with muddy tubes
Same locality: wash of sponge
Same locality: wash of sponge
Same locality: wash of algal clumps
Same locality: wash of flabellinid alga
Same locality: miscellaneous collections
1.5 mi S of village on W side of bay, diving: Sargassum sp.
A BIOLOGICAL SURVEY OF BAHIA DE LOS
ANGELES, GULF OF CALIFORNIA, MEXICO. II
BENTHIC POLYCHAETOUS ANNELIDS.
DONALD J. REISH
MUS, COMP, ZOOU
LIBRARY
JUL 8 1968
HARVARD
UNIVERSITY
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 7, 17 JUNE 1968
A BIOLOGICAL SURVEY OF BAHIA DE LOS ANGELES,
GULF OF CALIFORNIA, MEXICO. II.
BENTHIC POLYCHAETOUS ANNELIDS.
Donald J. Reish
Abstract. The benthic polychaetous annelids of Bahia de los Angeles were studied quantitatively in
April 1962 and October 1963. Distribution and sediment preferences of the principal species are
summarized. Of the 112 species encountered, ninety were previously unreported from the Gulf ol
California. Six new species are described: Microphthalmus riojai, sp. n., Aricidea {Aricidea) rosea
sp. n., Prionospio longibranchiata sp. n., Pectinaria (Pectinaria) hartmanae sp. n., Euchone barnardi
sp. n., and Euchone cortezi sp. n. In April 1962, 3 9,470 specimens were collected; in October 1963,
7,100. The cause of this reduction is unknown, but high summer water temperatures may be
involved. Previous reports of polychaetes from the Gulf of California are summarized.
Resumen. Los Anelidos Poliquetos bentonicos de la Bahia de los Angeles fueron estudiados cuantita-
tivamente durante Abril de 1962 y Octubre de 1963. Se detalla la distribucion de las especies
principales, y las preferencias que presentan respecto a los distintos sedimentos. De las 112 especies
registradas, solamente 22 han sido observadas previamente en el Golfo de California. Se incluye la
descripcion de seis especies nuevas: Microphthalmus riojai sp. n., Aricidea (Aricidea) rosea sp. n.
Prionospio longibranchiata sp. n., Pectinaria (Pectinaria) hartmanae sp. n., Euchone barnardi sp. n.,
y Euchone cortezi sp. n. En Abril de 1962 se obtuvieron 3 9,470 ejemplares, y solamente 7,100 en
Octubre de 1963. Se desconocen los factores causantes de esta enorme diferencia numerica; sin
embargo, se considera que las temperaturas elevadas registradas durante el verano, pudieron ejercer
una definida influencia en ese fenomeno. Tambien se incluyen las observaciones previas correspondi-
entes a los Poliquetos del Golfo de California.
Early accounts of polychaetes from the Gulf of California have been summarized
by Rioja (1941). Subsequent information has been added by Rioja (1943; 1947a, b),
Hartman (1944a, 1950), Treadwell (1942), Fauvel (1943), Woodwick (1961), and
Parker (1963). However, no quantitative studies have been made. Previous quantitative
studies of eastern Pacific polychaetes by Reish (see references) have dealt with nearly
all the principal bays from Morro Bay, in central California, south to Bahia de San
Quintin, Baja California. This paper, which summarizes the quantitative data on the
distribution of polychaetes in Bahia de los Angeles, a small bay on the west side of the
Gulf of California, and compares the fauna ecologically and zoogeographically with
that of similar areas in southern California, extends the area of quantitative studies into
a geographically disjunct area.
The specimens on which this report is based were collected by the Beaudette Ioun-
dation in April 1962 and October 1963 (see Barnard and Grady, 1968). Quantitative
samples were taken at 79 stations in April 1962 and 82 stations in October 1963
(Fig. 1) using a modified size-one Hayward orange-peel bucket which samples an area
of 0.06m-. Three dredge hauls were made in April 1962. Samples were brought back to
the base camp, washed through a 0.5 mm screen and preserved in formaldehyde for
later sorting. I wish to thank Dr. J. Laurens Barnard for making the collections avail-
able to me. My research was supported in part by a grant from the National Science
Foundation (NSFG- 20909).
Chemical and Physical Data. — Salinity was measured throughout the bay in April
1962, April 1963, and October 1963 at water depths of 0, J, 10. 20, 30, 40, .md JO
meters. All values ranged from about 3 5.0 to 3 5.5 0 00, which coincide with the
salinity values given for the upper Gulf of California by Parker (1963).
San Diego Soc. Nat. Hist., Trans 1$ (7): 67-106, 17 June 1968.
68
San Diego Society of Natural History
Vol. 15
35' 34'
33'
32'
31
II3°30'
28°
58'
1 / 1
152
154
28°
58'
148
\ 147 149 150 151
•
153
•
1^5
156
•
157
158
159
160
161 C\~ ^v
\ 130 132 134
\ • • • • •
\ 131 133
•
135
136
•
r37 •
138
•
•
139
140
141
142
14*3
144 ) ^~
/^T4S~ i46
57'
\ 116 118
\ • • • •
\ 117 119
120
•
•
121
122
123
124
•
•
123
126
•
•
127
128
I29S
57'
\ 103 105
)<» 104
106
107
108
109
110 •
III
•
•
112
113
•
•
114
• \
~"5 \
91
93
95
97
99
101
/ 90 92
94
96
98
100
\ 78 80
82
84
86
88
56'
79
\ 66
V * *
^. 67
ei
68
•
69
83
70
•
•
71
85
72
•
73
87
74
•
75
89
76
•
•\^ — )
77 / /•)
56°
•
34
55
«
•
56
57
•
56
59
•
60
61
•
•
62
63 — ^_^
• • • )
64 63 k
41
•
•
42
V
•
44
45
•
46
47
•
&
49
•
51 /^
• • ft
50 32/
55'
Station Locations \
29
•
30
31
•
3*2
33
•
•
34
33
•
•
36
•
37
39 /
M 1
55'
Number abov* dot ■ April, 1962
Number below dot • October, 1963
19
•
•
20
21
•
•
22
23
•
•
24
25
•
26
27 /
28 /
r
10
•
II
12
•
•
13
14
•
1*5
16
1*7 1*8 )
54'
8 A h 1 A DE LOS ANGELES
1
2
3
4
5
6
7
54'
0 5 i
NAuTiCAL WILCS
0 500 '000 I5O0
,
,
|
|
KH.0ME T£«5
l l
35' 34'
33'
32'
3
II3°30'
Figure 1. — Map of Bahia de los Angeles showing station locations.
Dissolved oxygen was measured at the same stations. Generally the dissolved oxygen
values exceeded 3.0 ml/1 with higher readings in April than in October. Bottom waters
at depths of 75 to 98 meters reached lows of 1.66 ml/1.
Surface water temperatures were taken at the majority of the stations. They varied
from 16.9 to 21.6 with a median of 18.2C. In general, warmer waters were found
inshore in shallow waters and colder waters were encountered near the entrance of the
bay. The warm temperatures occurred after the high summer air temperatures, and the
colder temperatures followed the winter months. The warm water temperatures are
believed to have influenced the marked reduction in the number of specimens collected
in October 1963 as compared to the April 1962 survey.
Four kinds of sediments occur on the bottom of Bahia de los Angeles: sand, silty
sand, sandy silt, and silt. Sand bottoms (40 stations) are present primarily along the
shores of the bay. These are replaced by silty sand, (46 stations) and sandy silt bottoms
(30 stations) in deeper waters. Silts (45 stations) are limited to the outer part of the
bay (Barnard and Grady, 1968).
1968 Reish: Bahia de los Angeles Annelids 69
Discussion
Zoogeography. — A total of 1 12 species were encountered in this study. Specific iden-
tification was not possible for one species of Arenicola that was represented by a single
incomplete specimen. Only 22 of these species, had previously been reported from the
Gulf of California. Of the 90 species previously unknown from the Gulf of California,
72 have been taken from southern California waters, two from northern California, one
from British Columbia, and one from Alaska (also Acapulco). Northern distributional
extensions include four from the Pacific side of southern Mexico (including the one
also known from Alaska), one from Panama, one from Peru, and two from the Central
Pacific Ocean.
Table 1 (see Appendix) summarizes the distributional data on the polychaetcs of
the Gulf of California. Species previously unknown from the area are indicated by an
asterisk; literature citations and locality data are included. Only included in Table 1 are
those reports in which specific identification was made by earlier workers; excluded are
reports to only family or genus. No attempt was made to determine or evaluate the
identification of others, since their material was not at hand. However, evaluations or
opinions of subsequent workers are included. Presently, 3 00 species of polychaetes are
known from the Gulf of California. The quantitative survey at Bahia de los Angeles
therefore increased the number known from 210. Probably this total will be increased
by other quantitative studies.
Relationship of polychaetes to sediments. — As noted, four kinds of sediments were
encountered on the benthos of Bahia de los Angeles. The relationships of the percentage
occurrence and the average number of specimens per station of the principal polychaetes
(based on the presence at 3 5 or more stations) to the general sediment type are given
in Table 2. Since the number of stations characterized by a particular sediment varied,
the occurrence of a species on a particular substrate is given in percentage units rather
than according to the number of times it was found. In addition, the average number
of specimens per station is a further indication of substrate preference or tendency. On
the basis of this analysis, the most abundant species of polychaetes may be grouped as
follows:
Preference for sand: Pareulepis fimbriata, Ophiodroiniis pugettensis, Pilargis hama-
fits, Langerhansia hetcrochaeta, Ceratoncreis mirabilis, Nereis procera, Glycera
tesselata, Lumbrineris minima, Chaetozone corona, Armandia bioculata, Chone mollis.
Preference for silty sand: Eteone dilatae, Ancistosyllis bassi, Tharyx pan ns.
Megalotnma pi gm en turn.
Preference for sand and silty sand: Prionospio pinnata, Magelona californica,
Telcpsavits costaritm.
Preference for sand, silty sand, sandy silt: Haploscoloplos elongatus, Praxillella
affinis pacifica.
Preference for sandy silt: Prionospio pygmaeits.
Preference for silty sand and sandy silt: Capitata ambiseta.
Preference for sandy silt and silt: Onuphis zebra, Prionospio cirri f era.
Preference for silt: Aricidea rosea, Cossnra Candida.
No apparent preference for specific sediments: Aglaophamus dicirrus, Glycindc
armigcra, Polydora socialis plena, Heteroclymene glabra, Amacana occidentals.
Euchone barnardi.
These data indicate that the commonest species of polychaetes occur near sandy
sediments. Furthermore, the finer the sediments the fewer the number of species en-
countered, both with respect to percent occurrence and the average number of specimens
encountered per station.
Substrate domination by polychaetes. — Dominant species were defined as those
represented by the largest number of specimens. These were compared for substrate
70 San Diego Society of Natural History Vol. 1 5
preferences. Five species constituted the dominant bay species during the April 1962
survey and nine species in October 1963. Only two species, Prionospio cirrifera and
Tharyx parvus, were dominant in both surveys.
April 1962. — Sandy substrates: Cap/fata ambheta, Chaetozone corona, Praxillella
affinis pacifica, Haploscoloplos elongatus, Spiophanes bombyx. Silty sands: C. ambheta,
H. elongatus, Aricidea rosea, and Tharyx parvus. Sandy silts and silts: A. rosea, C. am-
bheta. Silts: Prionospio cirrifera, A. rosea, and C. ambheta.
October 7 963. — Sandy substrate: Prionospio pinnata, Armandia bioculata, Glycera
tesselafa. Silty sands: P. pinnata, Magelona californica, Prionospio cirrifera, Tharyx
parvus. Sandy silts: Aglaophamus dicirris, Onnphis zebra, P. cirrifera. Silts: O. zebra,
Cossura Candida.
Seasonal Differences. — Of 46,570 polychaetes taken during this study, 39,470 were
collected in April 1962 and 7,100 in October 1963. The cause of this 89 per cent
reduction in numbers of specimens is unknown, but some explanations can be offered.
Summer water temperatures may be too high for this primarily temperate fauna. Median
surface-water temperatures were 18. 2C in April 1962 and 25. 9C in October 1963.
Greater extremes have been measured (Barnard, personal communication). Minima of
15C (possibly 12C) were measured in February and 29. 8C in August during the years
1962 and 1963. A second possibility is that red tides might lower the dissolved oxygen
below minimal values for survival of benthic animals. Such a phenomenon has occurred
in Alamitos Bay, California (Reish 1963b). Collections of benthic animals in Alamitos
Bay were made before and after an extensive bloom of the dinoflagellate, Gonyaulax
polyhedra Stein. Approximately 5 0 per cent of the animals, the majority of which were
polychaetes, were eliminated; of the remaining species nearly two- thirds decreased in
abundance. Dense red tides were observed in Bahia de los Angeles by Dr. J. L. Barnard
in March 1961, April 1962, and April 1963, but whether they contributed to the
summer decrease in the fauna is unknown. The causative organism of these red tides in
Bahia de los Angeles is unknown.
The number of specimens per station was plotted for both surveys (Figs. 2, 3).
In both surveys the largest populations occurred in the inner middle part of the bay
and along the north shore. A marked decrease in polychaete numbers was evident in the
October survey. In April 1962, samples ranged from 66 to 1722 individuals (median
500), but in October 1963 only 14 to 425 (median 46) were collected. The decrease
in the number of species collected, from 98 to 90, was not as noticeable. Considering
only the 7 5 species in which ten or more individuals were collected, the inter-sample
variation can be classified as follows.
Species presently only in April 1963 (6). — Enmida sanguinca, Syllidia liniafa,
Pionosyllis gigantia, Platynereis bicanaliculata, Orbinia jobnsoni, Polycirris perplcxus.
Species which decreased from April 1962 to October 1963 (41). — Eteone dilatae,
Paranaitis polynoides, Microphthalmns riojai, Ancisfrosyllis bassi, Langerhansia hetero-
chacta, Ccratoncreis mirabilis, Nereis procera, Pilargis hamatus, Aglaophamus dicirris,
Nephtys panamensis, Glycera americana, Lnmbrineris minima, Haploscoloplos elongatus,
Aricidea rosea, Paraonis gracilis oculata, Nerinides acuta, Polydora socialis plena,
Prionospio cirrifera, P. pygmaeus, Spiophanes bombyx, S. missionensis, Telepsavus cos-
tarum, Caulleriella alata, Chaetozone corona, Tharyx parvus, T. tesselafa, Polyophthalmus
pictus, Travisia gigas, Anotomastus gordiodes, Capifafa ambheta, Notomastus
(Clistomastus) tenuis, Heteroclymene glabra, Maldane sarsi, Praxillella affinis pacifica,
Am phisamytha bioculata, Amphicteis scaphobranchiata, Amaeana occidentals, Tere-
bellides stroemi, Chone mollis, Euchone bamardi, Megalomma pigmentum.
Species present only in October 1963 (3). — Nereis riisei, Laonice cirrata, Fabricia
li mni cola.
1968
Reish: Bahia de los Angeles Annelids
71
"^TT
x^x-x-x:x:-x-x:x::;x;::Xx.xx :•:•:■:::•:::
v.x:::x'::x-X\-x-xx+xxoxx :xxxxx;x:;.
Specimens per sample
of alt Polychaete species ^
Apr. I 1962
9
'4% 10-19 8#2e0«99
i:;;^ 50-99 ^g 500-999
Jgg! 100 249 HlOOO-
B * — » or .Ci Mt E l t s
Figure 2. — Map of Bahia de los Angeles showing the number of specimens of poly
chaetes collected per station in April 1962.
Figure 3. — Map of Bahia de los Angeles showing the number of specimens of poly
chaetes collected per station in October 1963.
72 San Diego Society of Natural History Vol. 1 5
Species which increased from April 1962 to October 1963 (16). — Pareulepis
fimbnata, Chloeia viridis, Pareurythoe califomica, Ophiodromus piigettcnsis, Exogone
lonrei, Eunice vittata, Protodorvillea gracilis, Prionospio malmgreni, P. pinnata, Magelona
califomica, Cirriformia sphabrancha, Armandia bioculata, Ouenia fusiformh collaris,
Pista cristata.
Species in approximate equal numbers in both surveys (9). — Glycinde armigera,
Onupbis zebra, Pbylo felix, P. nudits, Scoloplos acmeceps, Aricidea uscbakoui, Poecilo-
cbaetus jobnsoni, Cossura Candida, Pberusa neopapillata.
Therefore, more species were reduced or eliminated from April 1962 to October
1963 than the reverse. If this reduction is an annual phenomenon, then repopulation —
presumably from the deeper waters of the gulf — must be rapid, judged by the large
samples obtained during the April survey. Bahia de los Angeles specimens are smaller
than those of the same species from southern California. Possibly many are immature,
but some with ova developing within the coelom were observed. A study of the causes
of the reduction and subsequent increase in the fauna should be undertaken.
In the following list, the number of stations in at which each species was collected,
and the number of specimens collected (in parentheses) is given for each survey. The
detailed station listing of species and numbers of specimens has been filed with the
American Documentation Institute, Auxiliary Publication Service, Library of Congress,
Washington, D.C. Holotypes and paratypes, and additional specimens of the new species
have been deposited in The United States National Museum; the remaining material
studied is in the author's collection.
SYSTEMATIC LIST
Family Polynoidae
Lepidouotus purpureas Potts
Lepidonotus purpureus. Potts, 1910: 334; Hartman, 1951: 91. Lepidouotus hedleyi Benham. Hartman,
1939a: 40.
Material. — April 1962: 3(4).
Ecology. — Two specimens from sandy substrates, two from silt-sand mixture.
Distribution. — Indian Ocean, Australia, South Pacific, Gulf of California.
Panthalis pacifica Treadwell
Panthalis pacifica. Treadwell, 1914: 184; Hartman, 1939a: 87-88.
Material. — April 1962: 2 (2).
Ecology. — ■ Both specimens from silty substrates.
Distribution. — -Southern California to Baja California, questionably Panama (Hartman, 1939a).
Polyodontes from Hartman
Polyodontes frons. Hartman, 1939a: 84-86.
Material. — April 1962: 1 (1). October 1963: 1 (1).
Distribution. — Previously known only from type collected from Pinas Bay, Panama.
Family Sigalionidae
Stenclanella uniformis Moore
Stelelanella uniformis. Moore, 1910: 391-395; Hartman, 1939a: 69-70.
Material. — October 1963: 3 (9).
Ecology. — Eight specimens from sandy substrates, one from silty sand.
Distribution. — Southern California to Ecuador. This is the first report from the Gulf of California.
Stenolepis fimbriarum (Hartman)
Leanira fimbriarum. Hartman, 1939a: 70-72. Stenolepis fimbriarum. Hartman, 1965b: 15.
Material. — April 1962: 5 (8).
Ecology. — Three stations sandy, two silty.
Distribution. — Southern California to Ecuador; Gulf of California.
Thalenessa lewisii (Berkeley and Berkeley)
Sigalion lewisii. Berkeley and Berkeley, 1939: 226-228. Eusigalion hancocki. Hartman, 1939a: 59-60.
Thalenessa lewisii. Hartman, 195 9: 122.
Material. — April 1962: 2 (8); October 1963: 1 (1).
Ecology. — Eight specimens taken from sandy substrates, and one from silty sand.
Distribution. — Gulf of California south to Ecuador and Galapagos Islands.
1968 Reish: Bahia de los Angeles Annelids 73
Family Pareulepidae
Pareulepis fimbriata (Treadwell)
Eulepis fimbriata. Treadwell, 1901: 190. Pareulepis fimbriate Hartman, 1939a: 79-80; 1961: 54-55.
Material. — April 1963: 13 (29); October 1963: 21 (37).
Ecology. — -Shows a slight preference for sandy sediments (Table 2).
Distribution. — Southern California to Ecuador; West Indies; New Caledonia; Madgascar.
Family Chrysopetalidae
Paleano/us bellis (Johnson)
Heterophil- bellis. Johnson, 1897: 163. Paleanotus belln. Hartman, 1961: 57-58.
Material. — April 1962: 1 (1); October 1963: 1 (1).
Ecology. — Silty sands.
Distribution. — British Columbia to western Mexico. This is the first report from the Gulf of California.
Family Amphinomidae
Chloeia liridis Schmarda
Cloeia liridis. Schmarda, 1861: 144. Hartman, 1940: 205.
Material. — April 1962: 5 (8); October 1963: 5 (18).
Ecology. — Occurs in all types of substrates.
Distribution. — West Indies, Gulf of California, south to Panama, Galapagos Islands, and Cocos Islands.
Pareurythoe californica (Johnson)
Eurythoe californica Johnson, 1897: 159. Pareurythoe californica. Hartman, 1940: 203-204; 1961: 11.
Material. — April 1962: 3(4); October 1963: 4 (13).
Ecology. — Preferred silty sand substrates.
Distribution. — Central and southern California; the distribution is extended into the Gulf of California.
Family Phyllodocidae
Anaitides mucosa (Oersted)
Pbyllodoce mucosa. Oersted, 1843: 31. Anaitides mucosa. Hartman, 1961: 12.
Material. — April 1962: 4 (4).
Ecology. — Taken from various substrates.
Distribution. — Cosmopolitan. This is the first report from the Gulf of California.
Anaitides williamsi Hartman
Anaitides williamsi. Hartman, 1936: 126; Reish, 1963: 408.
Material. — April 1962: 3 (5).
Ecology. — Sandy sediments.
Distribution. — Oregon to Pacific Baja California. This is the first report from the Gulf of California.
Eteone dilatae Hartman
Eteone dilatae. Hartman, 1936: 130-13 1; Reish, 1963a: 408.
Material. — April 1962: 35 (85); October 1963: 7 (14).
Ecology. — Preference for silty sands (Table 2), which agrees with the finding of Hartman (1961) and
Reish (1963a).
Distribution. — ■ California and San Quintin Bay. This is the first report from the Gulf of California.
Eu mid a san guinea (Oersted)
Eulalia sanguinea. Oersted, 1843: 28. Eumida sanguinea. Hartman, 1961: 13.
Material. — April 1962: 17 (30).
Ecology. — ■ This species was taken from a variety of substrates.
Distribution. — -Cosmopolitan. Rioja (1947a) reported this species from La Paz.
Paranoids polynoides (Moore)
Anaitis polynoides. Moore, 1909b: 339. Paranaitis polynoides. Hartman, 1961: 13.
Material. — April 1962: 16 (18); October 1963: 1 (1).
Ecology. — Specimens collected from a variety of substrates.
Distribution. — British Columbia to southern California. The distribution is extended into the Gull 01
California.
Family Typhloscolecidae
Typbloscolex mulleri Busch
Typhloscolex mulleri. Busch, 1851: 115; Tebble, 1960: 231-236.
Material. — October 1963: 1 (1).
Remarks: — Typbloscolex mulleri is a pelagic cosmopolitan species. Presumably, this specimen was collected
while the orange-peel bucket was being brought to the surface.
74 San Diego Society of Natural History Vol. 1 5
Family Hesionidae
Genus Microphthalmias Mecznikow, 1865, emended.
Type: Microphthalmia sczelkouii Mecznikow, 1865.
Remarks. — -All known species of the genus possess three antennae and a pygidial plate (posterior end
unknown in M. aciculata Hartman-Schrbder, (1962a). Since the pygidial plate is distinctive to this genus
of hesionids, the generic diagnosis of Microphthalmus is emended to include species with two antennae
as well as three antennae
Diagnosis. — ■ Prostomium with two or three frontal antennae, a pair of small ventral palpi; eyes present
or absent; segments 1 to 3 each with a pair of dorsal and ventral tentacular cirri, no setae; parapodia
unequally a single simple notoseta; dorsal and ventral cirri short and digitiform; neurosetae compound;
pygidium with an expanded plate and two dorsal anal cirri.
Microphthalmus riojai, new species
Figure 4, a-f
Material. — Stations, April 1962: 5 (12); October 1963: 3 (5).
Description. — Length up to 3 mm; width including setae up to 0.2 mm; setigerous segments up to 30;
prostomium and dorsal surface with reddish brown pigment; prostomium (Fig. 4a) trapezoidal in shape,
widest posteriorly; two eyes at the posterior-lateral margin; a pair of antennae at the anterior margin of
the prostomium; a pair of simple ventral palpi, nature of proboscis unknown; no jaws seen; segments
1 to 3 asetigerous, with 6 pairs of long tentacular cirri; parapodial lobes unequally biramous (Fig. 4c),
neuropodial lobe conical in shape; one simple, curved lyrate notoseta (Fig. 4d) per lobe, each provided
with 8 teeth; 4 to 6 compound neuroseta per lobe (Fig. 4e, f ) , each terminate in a delicate bidentate tip;
pre-anal segment asetigerous, provided with a pair of dorsal and ventral knob-shaped cirri; pygidial plate
(Fig. 4b) broadly concave with no indication of a bilobed condition; anal cirri extend beyond the margin
of the anal plate.
Remarks. — Ten species are previously known to belong to this genus, three of which were recently
described from the Pacific side of South America by Hartmann-Schroder (1962a, b) . Microphthalmus riojai
differs from the other species by the absence of the median unpaired antennae. It belongs to that group
of species characterized by the presence of lyrate setae in the notopodium. Other species known to possess
this type of notosetae include M. sczelkowii Mecznikow, (1865), M. s. marmanica Zachs (in Uschakov,
1955), and M. nrofimbriata Alikunhi (1943.)
Ecology. — Associated with the coarser sediments; eleven specimens were taken from a sandy substrate
and six from silty sand.
Type material. — The holotype and 7 paratypes have been deposited in the U. S. National Museum.
Type locality. — Bahia de los Angeles, Gulf of California, Baja California. This species is named in honor
of Dr. Enrique Rioja, in recognition of his contribution to the knowledge of Mexican polychaetes.
Ophiodromus pugettensis (Johnson)
Podarke pugettensis. Johnson, 1901: 395; Rioja, 1947a: 202. Ophiodromus pugettensis. Hartman, 1961:
67-68; Reish, 1963a: 423.
Material. — April 1962: 29 (46); October 1963: 19 (101).
Ecology. — -Found on sandy or silty substrates (Table 2).
Distribution. — -Japan and from British Columbia to Peru.
Oxytlromns arenicolus glabrns Hartman
Oxydromus arenicolus glabrus. Hartman, 1961: 68-69.
Material. — April 1962: 4 (9).
Ecology. — ■ Specimens from silty sands or sandy silts.
Distribution. — Previously known from California. The distribution is extended into the Gulf of California.
Oxydromus brunnea Hartman
Oxydromus brunnea. Hartman, 1961: 69-70.
Material. — April 1962: 2 (2).
Ecology. — Sand and silty sand.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Syll/dia liniata Hartmann-Schroder
Syllidia liniata. Hartmann-Schroder, 1962a: 115-116.
Material. — April 1962: 8 (55); October 1963: 3 (6).
Ecology. — Silty sands at most stations; however, 30 specimens were taken at a station consisting of sands.
Distribution. — Peru; the distribution is extended into the Gulf of California.
Family Pilargidae
Ancisfrosyllis bassi Hartman
Ancistrosyllis bassi. Hartman, 1947b: 50 1-504.
Material. — April 1962: 4 (7); October 1963: 35 (52).
Ecology. — A. bassi showed a slight preference for silty sand sediments (Table 2).
Distribution. — Known from California and Florida; the distribution is extended into the Gulf of
California.
1968
Reish: Bahia de los Angeles Annelids
75
.07mm
.006 mm
K 1
.008 mm
Figure 4. — Microphthalmus riojai, n. sp.: a, anterior end, dorsal view; b, posterior
end, dorsal view; c, parapodium from mid-region, magnification as figure b; d, lyre seta
from notopodium; e-f, compound neurosetae from neuropodium.
76 San Diego Society of Natural History Vol. 1 5
Loandalia fauveli Berkeley and Berkeley
Loandalia fauveli. Berkeley and Berkeley, 1941: 30-3 1, pi. 5, figs. 4-6.
Material. — April 1962: 4 (4); October 1963: 5 (5).
Ecology. — Taken from all substrate types.
Distribution. — Previously known only from southern California; the distribution is extended into the
Gulf of California.
Pilargis bamatus Hartman
Pilgaris bamatus. Hartman, 1960: 88-89.
Material. — April 1962: 26 (63); October 1963: 9 (16).
Ecology. — Pilargis bamatus showed a preference for either sands or silty sands (Table 2).
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Family Syllidae
Braniit limbata (Claparede)
Grubea limbata. Claparede, 1868: 52. Brania limbata. Reish, 1959a: 79; Hartmann-Schroder, 1962b: 103.
Material. — April 1962: 2 (3).
Ecology. — Sandy sediments.
Distribution. — Known from Europe, southern California, and Chile. This is the first report from the
Gulf of California.
Exogonc lourci Berkeley and Berkeley
Exogone lourci. Berkeley, 1938: 44-47; Reish, 1959a: 80.
Material. — April 1962: 5 (13); October 1963: 11 (231).
Ecology. — This species showed a preference for either sandy or silty sand sediments; 161 specimens were
taken from a station composed of silty sands.
Distribution. — This species is known from British Columbia to southern California and Acapulco,
Mexico. The distribution is extended into the Gulf of California.
Langcrhansia beterocbaeta (Moore)
Syllis (Ehlersia) beterocbaeta. Moore, 1909b: 322-325. Langerhansia beterocbaeta. Hartman, 1959: 210,
1961: 16.
Material. — April 1962: 21 (70); October 1963: 16 (40).
Ecology. — -This species showed preference for either a sandy cr silty sand substrates (Table 2).
Distribution. — Known from British Columbia to southern California. This report extends its distribution
into the Gulf of California
Otfontosyllis phosphor e a Moore
Odontosyllis phosphorea. Moore, 1909b: 327-329; Hartman, 1961: 76-77.
Material. — October 1963: 1 (1).
Ecology. — Sandy sediments.
Distribution. — British Columbia to southern California; this is the first report of the species from the
Gulf of California.
Pionosyllis gigantca Moore
Pionosyllis gigantea. Moore, 1908: 325-328; Hartman, 1961: 16.
Material. — April 1962: 8 (27).
Ecology. — All but 3 specimens from stations possessing sandy sediments.
Distribution. — ■ Known from British Columbia to California; the distribution is extended into the Gulf
of California.
Sphaerosyllis crinaceus Claparede
Spbacrosyllis crinaceus. Claparede, 1863: 45; Rioja, 1943: 211-214; Day, 1954: 13-14; Pettibone, 1963;
135-136.
Material. — April 1962: 1 (1); October 1963: 1 (6).
Ecology. — Substrate sand or silty sands.
Distribution. — Northern Hemisphere and Tristan de Cunha; Rioja (1943) reported it from Acapulco;
this is the first record from the Gulf of California.
Family Nereidae
Ceratoncreis mirabilis Kinberg
Ceratoncreis mirabilis. Kinberg, 1866: 170; Hartman, 1948: 71-72. Nereis (Ccratonercis) mirabilis. Fauvel,
1953: 200-201. Ceratoncreis singular is. Treadwell, 1929: 1-3.
Material. — April 1962: 33 (68); October 1963: 9 (29).
Ecology. — Marked preference for sandy sediments (Table 2).
Distribution. — Cosmopolitan in warmer seas. Hartman (1948) referred C. singularis Treadwell from
Carmen Island, Baja California to this species.
1968 Reish: Bahia de los Angeles Annelids 77
Nereis procera Ehlers
Nereis procera. Ehlers, 1868: 557; Reish, 1959a: 81-82.
Material. — April 1962: 17 (65); October 1963: 6 (16).
Ecology. — Nereis procera showed a slight preference for sandy substrates (Table 2).
Distribution. — British Columbia to southern California; the distribution is extended into the Gulf of
California.
Nereis riisei Grube
Nereis riisei. Grube, 1857: 162; Hartman, 1940: 221-222; 1951: 46; Rioja, 1959: 225.
Material. — October 1963: 6 (11).
Ecology. — Nine specimens from sandy substrates; two from silty sands.
Distribution. — Known from the Gulf of California to Ecuador and Gulf of Mexico; the distribution
within the Gulf of California is extended northward.
Platynereis bicanaliculata (Baird)
Nereis bicanaliculata. Baird, 1863: 109; Platynereis bicanaliculata. Hartman, 1954: 36-39.
Material. — April 1962: 4 (32).
Ecology. — The substrate was sandy at Station 130 where 29 specimens were collected. The other stations
were composed of silty sands and sandy silts.
Distribution. — ■ Western Canada to Baja California, Gulf of California, Hawaii, Australia, and possibly
Peru (Hartmann-Schrbder, 1962a).
Family Nephtyidae
Aglaophamus dicirris Hartman
Aglaophamus dicirris. Hartman, 1950: 122-124.
Material. — April 1962: 67 (822); October 1963: 84 (509).
Ecology. — ■ Collected at 93 per cent of the stations sampled. Equally common on silty sand or sandy silt
substrates (Table 2). As widespread but less abundant in October 1963 than in April 1962; the average
number of specimens per station was only 6 (maximum 19) as compared to 12 (maximum 76). The
population, especially reduced in the inner reaches of the bay.
Distribution. — Southern California to Panama; Gulf of California; North Carolina.
Nephtys panamensis Monro
Nephtys panamensis. Monro, 1928: 81-82; Hartman, 1940: 239-240; 1950: 101.
Material. — April 1962: 24 (147); October 1963: 3 (4).
Ecology. — Commonest on sandy substrates.
Distribution. — Gulf of California to Panama.
Family Glyceridae
Glycera americana Leidy
Glycera americana Leidy, 1855: 147-148; Hartman, 1950: 73-75.
Material. — April 1962: 21 (40); October 1963: 6 (18).
Ecology. — Taken most frequently from the sandy and silty sand bottoms.
Distribution. — ■ Western Hemisphere, New Zealand, Australia.
Glycera robusta Ehlers
Glycera robusta Ehlers, 1868: 656-658; Hartman, 1950: 69-70.
Material. — April 1962: 2 (7).
Ecology. — Sandy sediments.
Distribution. — British Columbia to southern California and possibly Florida (Hartman, 1950). The dis-
tribution is extended into the Gulf of Californa.
Glycera tesselata Grube
Glycera tesselata Grube, 1863: 41-42; Hartman, 1950: 77-78; Im.ijima and Hartman, 1964: 165.
Material. — April 1962: 21 (69); Oct. 1963: 35 (658).
Ecology. — Glycera tesselata preferred sandy substrates (Table 2). Most specimens were collected in
October 1963 and these were small and presumably immature Worms.
Distribution. — ■ British Columbia south to tropical Pacific, Japan, Indo-Pacific, Caribbean Sea, and Mediter-
ranean Sea.
Family Goniadidae
Glycinde arminger Moore
Glycinde arminger Moore, 1911: 307-311; Hartman, 1950: 49-5 1.
Material. — April 1962: 57 (290); October 1963: 56 (342).
Ecology. — -Widespread, in all substrates in both survey; most specimens from sandy benthos ( Table 2).
The increase in October 1963 reflects the extreme abundance of the species (225 individuals) at station 65.
The largest sample in April 1962 was 18
Distribution. — British Columbia south to Galapagos Islands. The distribution is extended into the Gult
of California.
78 San Diego Society of Natural History Vol. 1 5
Goniada littorca Hartman
Goniada littorca Hartman, 1950: 23-26; Reish, 1963a: 425.
Material. — October 1963: 1 (1).
Ecology. — Silty sand.
Distribution. — • Southern California to San Quintin Bay; the distribution is extended into the Gulf
of California.
Family Onuphidae
Onuphis nebulosa Moore
Onuphis nebulosa Moore, 1911: 269-273; Hartman, 1944a: 75-78.
Material. — April 1962: 9 (23); Oct. 1963: 8 (44).
Ecology. — This species preferred coarser sediments.
Distribution. — Central California to Panama; this is the first report from the Gulf of California.
Family Onuphidae
Onuphis zebra Berkeley and Berkeley
Onuphis zebra Berkeley and Berkeley, 1939; 337-338; Hartman, 1944a: 71-72.
Material. — April 1962: 60 (746); Oct. 1963: 76 (566).
Ecology. — • This was one of the commonest worms in the bay. It was present in all types of substrates,
but it showed a preference for finer sediments. This species constructs a tube of silt. Onuphis zebra was
widespread throughout the bay during both surveys; a slight drop in the population was noted in the
October 1963 survey. The maximum April sample was 115; October. 23.
Distribution. — • Punta Gorda, Baja California and Guatemala; the distribution is extended northward in
the Gulf of California from near the vicinity of La Paz to Bahia de los Angeles.
Family Eunicidae
Eunice vittata (delle Chiaje)
Nereis vittata delle Chiaje, 1828: 195. Eunice vittata. Hartman, 1944a: 118.
Material. — April 1962: 1 (1); Oct. 1963: 3 (4).
Ecology. — Sands or silty sands.
Distribution. — ■ Warmer waters of the Western Hemisphere from Bermuda to West Indies and Trinidad,
from southern California to Panama, and many localities in the Gulf of California.
Nematonereis unicornis (Grube)
Lumbriconereis unicornis Grube, 1840: 80. Nematonereis unicornis. Fauvel, 1953: 249-250.
Material. — April 1962: 1 (5).
Ecology. — Sand.
Distribution. — Cosmopolitan in warmer seas; previously unknown from the Gulf of California.
Family Lumbrineridae
Lumbrineris erecta (Moore)
Lumbriconereis erecta. Moore, 1904: 490-492. Lumbrineris erecta. Hartman, 1944a: 149-150.
Material. — October 1963: 1 (1).
Ecology. — Sand.
Distribution. — Southern California to Panama, and Gulf of California.
Lumbrineris minima Hartman
Lumbrineris minima. Hartman, 1944a: 155-156; Reish, 1963a: 425-426.
Material. — April 1962: 57 (378). October 1963: 20 (53).
Ecology. — -Commonest on coarser sediments (Table 2). This is similar to the ecology off southern Cali-
fornia (Hartman, 1944a; Reish, 1959a), but unlike that at San Quintin Bay (Reish, 1963a).
Distribution. — Southern California and San Quintin Bay; the distribution is extended into the Gulf of
California.
Family Arabellidae
Drilonereis inula Moore
Drilonereis nuda. Moore, 1909a: 254-256; Hartman, 1944a: 178-179.
Material. — April 1962: 2 (2); October 1963: 3 (3).
Ecology. — Sands or sands and silts.
Distribution. — -Central California to Panama. Fauvel (1943) reported it from the Gulf of California
(locality unspecified).
Family Dorvilleidae
Protodori illea gracilis (Hartman)
Stauronereh gracilis. Hartman, 1938: 100-101. Don illea gracilis. Hartman, 1944a: 189. Protodori illea
gracilis. Hartman, 1965b: 48.
Material. — April 1962: 7 (20). October 1963: 7 (132).
Ecology. — Of 152 specimens, 141 were taken from eight stations characterized by sandy sediments.
Distribution. — California; the distribution is extended into the Gulf of California.
1968
Reish: Bahia de los Angeles Annelids
79
^T
Specimens per sample
April 1962
□ "9
i^J 10-49
ggg 100-249 (130 moiimum)
l>>ii 01 lOS i»C(il S
soc «O0 MO
Haploscoloplos e/ongofus
Figure 5. — Benthic distribution of Haploscoloplos elongatus in April 1962.
Sp*clm«n» p#f sompie
October 1963
ESS1 9
%0 10-49 (14mo„muml
■ * - i * Dt LOS IkClil)
*ao coo
Haploscoloplos e/ongofus
Figure 6. — Benthic distribution of Haploscoloplos elongatus in October 1963.
80 San Diego Society of Natural History Vol. 15
Family Orbiniidae
Hafiloscoloplos elougatus (Johnson)
Scoloplos elongata. Johnson, 1901: 412-413. Haploscohplos elongatus. Hartman, 1957: 273-275; Reish,
1963a: 426; 1965: 140; Imajima and Hartman, 1964: 274.
Material. — April 1962: 58 (1470). October 1963: 54 (252).
Ecology. — Widespread, commonest on silty sandy (Table 2). Much reduced in October 1963, especially
in inner part of bay (Figs. 5,6).
Distribution. — Japan, Beaufort Sea to San Quintin Bay; the distribution is extended into the Gulf of
California.
Orbhiia jobusoni (Moore)
Aricia jobusoni. Moore, 1909a: 260-262; Orbhiia johnsoni. Hartman, 1957: 257-260.
Material. — April 1962: 2 (4).
Ecology. — Sandy silts.
Distribution. — Central California to Costa Rica; previously unknown from the Gulf of California.
Phylo felix Kinberg
Pbylo felix. Kinberg, 1866: 25 1-252; Hartman, 1957: 262-265.
Material. — April 1962: 10 (15); October 1963: 19 (17).
Ecology. — Taken from all substrate types.
Distribution. — California, Gulf of California, Brazil, Patagonia, East Falkland Island, and Antarctic
Ocean.
Phylo nudus (Moore)
Aricia inula. Moore, 1911: 311-315. Pbylo nudus. Hartman, 1957: 268.
Material. — April 1962: 4 (7); October 1963: 6 (8).
Ecology. — Sands or a mixture of sands and silts.
Distribution. — • Southern California and Burma; the distribution is extended into the Gulf of California.
Scoloplos acmeceps Chamberlin
Scoloplos acmeceps. Chamberlin, 1919b: 15-16; Hartman, 1957: 282-283.
Material. — April 1962: 3 (7); October 1963: 2 (6).
Ecology. — Sands or sands and silts.
Distribution. — Alaska to Mazatlan; this is the first report from the Gulf of California.
Family Paraonidae
Aricidea uscbakowi Zachs
Aricidca uschakowi. Zachs, 1925: 1-3; Hartman, 1957: 321.
Material. — April 1962: 8 (176); October 1963: 24 (160).
Ecology. — Commonest on sand bottoms.
Distribution. — Russian Pacific Ocean to southern California; the distribution is extended into the Gulf
of California.
Aricidea rosea, new species
Figures 7a-c, 8
Material. — April 1962: 53 (5534); October 1963: 12 (19).
Diagnosis. — -Length to 7-8 mm; width 0.25 mm; to at least 70 setigerous segments; prostomium (Fig. 7a)
triangular in shape and rounded anteriorly, without eyespots, the median antenna extends into the second
setigerous segment; branchiae number 11 to 12 pairs, begin at setigerous segment 4 and extend to segment
15 or 16, third to ninth pair largest and may overlap with opposite member mid-dorsally; notopodia with
ciriform notopodial post-setal lobe which is well developed throughout the length of the worm; neuro-
podium poorly developed; setae of notopodium of capillary type and distally pointed (Fig. 7b); capillary
setae present in all ncuropodial lobes; beginning at about segments 20-2 5 and to the posterior end four
to six modified setae appear in the neuropodium (Fig. 7c); they are of the curved acicular type with a
secondary tooth and a pointed hood.
Type material. — -The holotypc and paratypes have been deposited in the U.S. National Museum. The type
locality is Bahia de los Angeles, Baja California, Mexico.
Remarks. — Hartman (1959b) listed nine species within the genus Aricidea (Aricidea) plus two addi-
tional species which were referred with question. Three additional species have since been described
A. (A.) lopezi ruba Hartman (1963), A. abranchiata Hartman (1965a), and A. neosuec/ca Hartman
(1965a). The different species of the genus may be conveniently separated from one another on the basis
of the number of pairs of branchiae and the nature of the modified neuroseta. Aricidea (A.) rosea can
be distinguished from the majority of the known species by the number of pairs of branchiae. There arc
11 to 12 pairs in A. (A.) rosea and 9 to 1 1 in A. (A.) lopezi ruba; they differ, however, in the
nature of their modified neuroseta.
Ecology. — This is one of the commonest polychaetes in Bahia dc los Angeles; 22 of the 5352 specimens
were collected in April 1962 (Fig. 15). The greatest concentration of A. rosea occurred in fine sediments
in the middle part of the bay.
1968
Reish: Bahia de los Angeles Annelids
81
.1 mm
•* | .006mm,
Figure 7 — Aricidea (Aricidea) rosea, n. sp.: a, anterior end, dorsal view; b, capillary-
setae from notopodium; c, modified seta from neuropodium.
Paradenitis lyra Southern
Paraonis (Paraonides) lyra. Southern, 1914: 94-95; Hartman, 1957: 334-335.
Paradoneis lyra. Hartman, 1965: 52.
Material. — October 1963: 1 (3).
Ecology. — Sand.
Distribution. — European waters and southern California; the distribution is extended into the Gulf of
California.
Paraonis gracilis oculata Hartman
Paraonis gracilis oculata. Hartman, 1957: 331-3 32.
Material. — April 1962: 13 (80); October 1963: 10 (17).
Ecology. — Commonest in sands and silty sands.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Family Spionidae
Laonicc cirrata (Sars)
Nerine cirrata. Sars, 1851: 207; Laonicc cirrata. Berkeley and Berkeley, 1952: 26; Imajima and Hartman,
1964: 281-282.
Material. — October 1963: 3 (10).
Ecology. — Silty sands.
Distribution. — -Cosmopolitan; this is the first report from the Gulf of California.
Nerinides acuta (Treadwell)
Spio acuta. Treadwell, 1914: 199-201. Nerinides acuta. Hartman, 1941: 294-296; Reish 1959a: 86.
Material. — April 1962: 5 (7); October 1963: 2 (2).
Ecology. — Sands and from sand and silts.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
San Diego Society of Natural History
Vol. 15
Figure 8 — Benthic distribution of Aricidea rosea in April 1962.
Nerinides maculata Hartman
Nerinides maculata. Hartman, 1961: 91-92; Reish, 1963a: 427.
Material. — October 1963: 1 (1).
Ecology. — Sand.
Distribution. — Southern California, San Quintin Bay, and the northern Gulf of California at San Felipe.
Nerinides pigmentata (Reish)
Spiophanes pigmentata. Reish, 1959c: 11-13. Nerinides pigmentata. Hartman, 1961: 92-93.
Material. — October 1963: 1 (2).
Ecology. — Silty sand.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Polydora heterochaeta Rioja
Polydora heterochaeta. Rioja, 1939: 308-309.
Material. — October 1963: 3 (3).
Ecology. — Sand or silty sands.
Remarks. — ■ Identified by Keith H. Woodwick.
Distribution. — Acapulco; the distribution is extended into the Gulf of California.
Polydora socialis plcn-x Berkeley and Berkeley
Polydora socialis plena. Berkeley and Berkeley, 1936: 468-469; 1952: 22.
Material. — April 1962: 27 (81); October 1963: 22 (36).
Ecology. — Occurs on all substrate; commonest on sandy silts.
Remarks. — Identified by Keith H. Woodwick.
Distribution. — Alaska and British Columbia; the distribution is extended in the Gulf of California.
Prionospio cirrifera Wiren
Prionospio cirrifera. Wiren, 1883: 409; Fauvel, 1927: 62-63; Reish, 195 9a: 87.
Material. — April 1962: 46 (623); October 1963: 62 (471).
Ecology. — This species was found at more stations, but not in greater numbers, in areas of finer sedi-
ments (Table 2). This relationship is similar to that found in Newport Bay, California (Reish, 195 9a).
Maximum count April, 1962, 48; October 1963, 34.
Distribution. — Europe, North Sea, Arctic Ocean, Bering Sea to southern California; the distribution is
extended into the Gulf of California.
Prionospio longibranchiata, new species
Fig. 9 a-e
Material. — April 1962: 2 (2); October 1963: 5 (6).
1968
Reish: Bahia de los Angeles Annelids
8 3
0.15 mm
.15 mm
i :
.014 mm
Figure 9. — Prionospio longibranchiata, n. sp.: a, anterior end, dorsal view; b, hooded
hook from neuropodium; c, anterior parapodium; d, posterior parapodium, magnification
as figure c; e, capillary seta from posterior inferior neuropodium.
84 San Diego Society of Natural History Vol. \5
Diagnosis. — • Length of holotype 5 mm; width 0.2 mm; 37 setigerous segments; incomplete specimens as
long as 12 mm, width 0.4 mm, 5 5 setigerous segments; prostomium (Fig. 9a) longer than broad, slightly
rounded and expanded anteriorly, tapers posteriorly to second setigerous segment; peristomium prolonged
anteriorly as lateral lobes to prostomium; palps broken off; 4 eyes equal in size with anterior pair farther
apart; 5 pairs of long, cirriform branchiae, some partially broken with the longest extending to segment
20; all setigerous segments biramous; anterior segments with only capillary setae; hooded hooks from
setigerous segment 17 in neuropodium, 21 in notopodium; hooded hooks (Fig. 9b) consist of one large
tooth at right angles to shaft and 3 smaller teeth, hood as indicated in Fig. b; details of anterior and mid-
body parapodial lobes shown in Fig. 9 c, d; pygidium with an unpaired longer cirrus and two shorter
ones.
Remarks. — -Thirty species are known previously from the genus Prionospio Malmgren (Hartman, 195 9b;
Hartmann-Schroder, 1962a, b; Laubier, 1962). Prionospio longibranchiata belongs to the group possessing
only cirriform branchiae, but can be distinguished by the number and nature of the branchiae.
Prionospio cirrifcra Wiren (1883), cosmopolitan in distribution, has from 6 to 12 pairs of short branchiae.
Prionospio polybranchiata Fauvel (1953), known from India, has 5 pairs of long branchiae, which resemble
those of P. longibranchiata, and at least 3 5 pairs of shorter branchiae. Prionospio hetcrobranchia Moore
(1907), from Massachusetts, P. h. tcxana Hartman (195 1a) from Texas, and P. b. navportensis, Reish
(1959c), from California, all have 5 pairs of branchiae, but the first, fourth, and fifth pairs are pinnate
in all three species.
Type material. — The holotype and paratypes have been deposited in the U.S. National Museum. Type
locality Bahia de los Angeles Baja California, Mexico.
Prionospio malmgreni Claparede
Prionospio malmgreni. Claparede, 1868: 333; Pettibone, 1954: 282-284.
Material. — April 1962: 7 (19); October 1963: 15 (59).
Ecology. — ■ Sands or sandy silts. In San Quintin Bay, P. malmgreni was more abundant in finer sediments
(Reish, 1963a).
Distribution. — ■ Northern Hemisphere and South Africa; this is the first report from the Gulf of California.
Prionospio pinnata Ehlers
Prionospio pinnata. Ehlers, 1901; 163-164; Hartman, 1960: 114-115.
Material. — April 1962: 36 (252); October 1963: 61 (755).
Ecology. — Commonest on sand and silty sands.
Distribution. — Cosmopolitan; this is the first report from the Gulf of California.
Prionospio pygmacus Hartman
Prionospio pygmacus. Hartman, 1961: 93-95; Reish, 1963a: 427.
Material. — April 1962: 51 (876); October 1963: 4 (9).
Ecology. — All substrates (Table 2).
Distribution. — Southern California, San Quintin Bay; the distribution is extended into the Gulf of
California.
PseuJopolydora rcishi Woodwick
Pseudopolydora rcishi. Woodwick, 1964: 15 2.
Material. — October 1963: 2 (4).
Ecology. — Silt and silty sands.
Remarks. — Specimens identified by Keith H. Woodwick.
Distribution. — This species was recently described from Eniwetok Atoll, Marshall Islands; the distribution
is extended into the Gulf of California.
Spiophanes bombyx (Claparede)
Spio bombyx. Claparede, 1870: 485. Spiophanes bombyx. Berkeley and Berkeley, 1952: 22-24.
Material. — April 1962: 20 (427); October 1963: 3 (8).
Ecology. — Occurred almost exclusively in sandy sediments.
Distribution. — Cosmopolitan; this is the first report of the species from the Gulf of California.
Spiophanes missionensis Hartman
Spiophanes missionensis. Hartman, 1941: 296-298; Reish, 1963a: 427.
Material. — April 1962: 9 (12); October 1963: 1 (1).
Ecology. — Taken from all substrates.
Distribution. — Southern California, San Quintin Bay; the distribution is extended into the Gulf of
California.
Family Magelonidae
Magelona calif ornica Hartman
Magclona califomica. Hartman, 1944c: 320-321; Reish, 1963a: 427.
Material. — April 1962: 14 (119); October 1963: 30 (537).
Ecology. — All specimens but one from sand or silty sands. Present at 73 per cent of the sandy stations
but only 30 per cent of the silty sand stations (Table 2).
Distribution. — Southern California. San Quintin Bay; the distribution is extended into the Gulf of
California.
1968
Reish: Bahia de los Angeles Annelids
85
Choetozone corona
M -..-■< M>0
Figure 10. — Benthic distribution of Cbaetozone corona in April 1962.
"1 TT
^3^r
Speclmervs per SQfDpl«
October 1963
m-«
12-49 (21 mo»imum)
Choetozone corona
Figure 11. — Benthic distribution of Cbaetozone corona in October 1963.
86 San Diego Society of Natural History Vol. 15
Family Disomidae
Poecilochactus johnsoni Hartman
Poecilochaetus johiisoni. Hartman, 193 9b: 164-166.
Material. — April 1962: 5 (6); October 1963: 3 (5).
Ecology. — Primarily sandy stations.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Family Chaetopteridae
Telepsavits cost arum Claparede
Telepsavus costarum. Claparede, 1868: 340; Berkeley and Berkeley, 1952: 63; Hartman, 1961: 31.
Material. — April 1962: 43 (433); October 1963: 24 (47).
Ecology. — Commonest on coarser sediments, but present at a few stations with sandy silt and silts.
Distribution. — Cosmopolitan; this is the first report from the Gulf of California.
Family Cirratulidae
Acrocirrus crassifilis Moore
Acrocirrus crassifilis. Moore, 1923: 188-190; Hartman, 1961: 31.
Material. — October 1963: 2 (5).
Ecology. — Sand or silty sands.
Distribution. — -Southern California; the distribution is extended into the Gulf of California.
Caulleriella alata (Southern)
Chaetozone alata. Southern, 1914: 112-113; Caulleriella alata. Hartman, 1961: 108.
Material. — April 1962: 10 (232); October 1963: 9 (21).
Ecology. — Preference for sand and silty sands.
Distribution. — Ireland and from British Columbia to southern California; the distribuiton is extended
into the Gulf of California.
Chaetozone corona Berkeley and Berkeley
Chaetozone spinosa. Moore, var. corona Berkeley and Berkeley, 1941: 45-46; Chaetozone corona. Reish,
1959a: 89; Hartman, 1961: 109-110.
Material. — April 1962: 49 (2112); October 1963: 3 2 (133).
Ecology. — This species was eliminated from the outer part of the bay and was reduced in the inner
reaches of the bay in October 1963 (Figs. 10, 11).
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Cirri f or mia spirabrancha (Moore)
Cirratulus spirabranchus. Moore 1904: 492-493; Cirriformia spirabrancha. Hartman, 1944b: 263.
Material. — April 1962: 2 (2); October 1963: 4 (20).
Ecology. — Sands or sand and silt mixture.
Distribution. — California; the distribution is extended into the Gulf of California.
Cossura Candida Hartman
Cossura Candida. Hartman, 1955: 44-45; Reish, 1963a: 428.
Material. — April 1962: 40 (341); October 1963: 42 (333).
Ecology. — C. Candida was found at a larger number of stations and in greater abundance with increas-
ing fineness of the sediment (Table 1). A similar ecology has been reported at San Quintin Bay
(Reish 1963a). About equal numbers were taken in both surveys.
Distribution. — Southern and Lower California; the distribution is extended into the Gulf of California.
Tharyx partus Berkeley
Tharyx multifilis parvus. Berkeley, 1929: 307, Tharyx parvus. Hartman, 1961: 113.
Material. — April 1962: 58 (1262); October 1963: 47 (520).
Ecology. — T. parvus showed a preference for the silty sands. About twice as many specimens were taken
in April 1962 than October 1963; the reduction in the population occurred largely in the outer reaches
of the bay (Figs. 12, 13).
Distribution. — British Columbia to southern California; the distribution is extended into the Gulf of
California.
Tharyx tesselata Hartman
Tharyx tesselata. Hartman, 1960: 126-127; 1961: 113.
Material. — April 1962: 14 (62); October 1963: 3 (3).
Ecology. — Sands, silty sands and sandy silts.
Distribution. — -Southern California; the distribution is extended into the Gulf of California.
Family Flabelligeridae
Pherusa neopapillata Hartman
Pherusa neopapillata. Hartman, 1961: 121-122.
Material. — April 1962: 10 (12); October 1963: 6 (10).
Ecology. — All sediment types.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
1968
Reish: Bahia de los Angeles Annelids
87
Specimens pe' somp
April 1962
HI 50-99
K§3 (00-249 (163 mo»imum)
Tharyx parvus
Figure 12. — Benthic distribution of Tharyx parvus in April 1962.
Tharyx parvus
Figure 13. — Benthic distribution of Tharyx pari its in October 1963.
88
San Diego Society of Natural History
Vol. 15
Figure 14. — Benthic distribution of Capitita ambheta in April 1962.
— Specimens per sample
October 1963
too ooo
Copitita ambiseta
Figure 15. — Benthic distribution of Capitita ambiseta in October 1963.
1968 Reish: Bahia de los Angeles Annelids 89
Pirotnis gracilis Hartman
Piromis gracilis. Hartman, 1961: 123-124.
Material. — April 1962: 3 (3).
Ecology. — -Sand and sandy silts.
Distribution. — Oaxaca, Mexico, Guatemala, and Ecuador; the distribution is extended into the Gulf of
California.
Family Opheliidae
Armandia bioculata Hartman
Armaudia bioculata. Hartman, 1938: 105; Reish, 1963a: 428.
Material. — April 1962: 3 1 (65): October 1963: 15 (264).
Ecology. — Preference for sandy substrates (Table 2).
Distribution. — California and San Quintin Bay; the distribution is extended into the Gulf of California.
Polyophthalmus pictus (Dujardin)
Nats picta. Dujardin, 1839: 293; Polyophthalmus pictus. Imajima and Hartman, 1964: 309.
Material. — April 1962: 1 (81); October 1963: 2 (2).
Ecology. — Found on sands or silty sands.
Distribution. — -Cosmopolitan; Rioja (1947) reported it from the lower part of Gulf of California.
Travisia gigas Hartman
Tunisia gigas. Hartman, 1938: 103-105; 1961: 34.
Material. — April 1962: 4 (14); October 1963: 1 (3).
Ecology. — Sandy at all stations.
Distribution. — -Central and southern California and Cabo San Lucas; the distribution within the Gulf
of California is extended northward.
Family Capitellidae
Auotomastus gordiodes (Moore)
Eunotomastus gordiodes. Moore, 1909a: 278-279; Auotomastus gordiodes. Hartman, 1947a: 442-444.
Material. — April 1962: 9 (34); October 1963: 3 (3).
Ecology. — Sand or a sand and silt mixture.
Distribution. — ■ California; the distribution is extended into the Gulf of California.
Family Capitellidae
Capitella capifata (Fabricus)
Lumbricus capitatus. Fabricus, 1780: 279; Capitella capifata. Fauvel, 1927: 154-155; Reish, 1963a: 428.
Material. — October 1963: 1 (2).
Ecology. — Silty sands.
Distribution. — Cosmopolitan; this is the first report from the Gulf of California.
Capitita ambiseta Hartman
Capitita ambiseta. Hartman, 1947a: 409-410; Reish, 1963: 429.
Material. — April 1962: 67 (18377); October 1963: 54 (215).
Ecology. — • C. ambiseta was the most abundant species collected; 18,5 5 2 specimens were taken. The popu-
lation was nearly eliminated following the warmer months with only 215 specimens collected from 5 4
stations in October 1962 (Figs. 14, 15). This species was taken more frequently, but in fewer number,
from stations composed of silty sediments. An average of 147, 255, and 182 specimens per station were
taken from sandy, silty sand, and sandy silt sediments respectively. The population of C. ambiseta in April
1962 was more extensive than that encountered at San Quintin Bay (Reish, 1963a).
Distribution. — Central California south to San Quintin Bay, Baja California; this is the first report
from the Gulf of California.
Notomastus (Clis/omastus) tenuis Moore
Nofomastus tenuis. Moore 1909a: 277-278; Notomastus (Clistomastus) t ennuis, Hartman, 1947a: 420-422;
Reish, 1959a, p. 92.
Material. — April 1962: 8 (16); October 1963: 6 (8).
Ecology. — Sands or silty sands.
Distribution. — British Columbia to southern California; this is the first report from the Gulf of California.
Family Arenicolidae
Arenicola sp.
Material. — April 1962: 1 (1).
Remarks. — -One specimen lacking the posterior end was taken; it was impossible to make a specific iden-
tification. The substrate was sandy at this station.
Family Maldanidae
Asychis disparidentata (Moore)
Maldane disparidentata. Moore, 1904: 494-496; Asycbis disparidentata. Berkeley and Berkeley, I'^l: 46-47.
Material. — April 1962: 1 (1).
Ecology. — Silt.
Distribution. — Known previously from British Columbia, California, and India; this is the first report
of the species from the Gulf of California.
90
San Diego Society of Natural History
Vol. 15
Spec
mens per somple
April
1962
EI
1-9
^
0-49
TTTTT^T
50-99
m
00-249 (110 mo.imum)
i>"i
DC t OS 1KUU5
o
=
Praxillella offinis pacifica
»oc -ooo
Figure 16. — Benthic distribution of Praxillella affinis pacifica in April 1962.
Proxillello affinis pocifica
Figure 17. — Benthic distribution of Praxillella affinis pacifica in October 1963.
1968 Reish: Bahia de los Angeles Annelids 91
Axiothella rubrocincto (Johnson)
Clymenella rubrocincta. Johnson, 1901; 418-419; Axiothella rubrocincta. Berkeley and Berkeley, ] y v 2 :
Jl; Reish, 1963a: 429.
Material. — October 1963: 1 (4).
Ecology.- — Silty sands.
Distribution. — British Columbia to San Quintin Bay; the distribution is extended into the Gulf of
California.
Heteroclymenc glabra Moore
Heteroclymene glabra. Moore, 1923: 229-230; Hartman, 1961: 37.
Material. — April 1962: 37 (382); October 1963: 1 (1).
Ecology. — Equally widespread on all types of sediments, but commonest on sandy substrates (Table 2).
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Mai da ne sarsi Malmgren
Maldane sarsi. Malmgren, 1865: 188; Fauvel, 1927: 197-199; Hartman, 1961: 37.
Material. — April 1962: 6 (28); October 1963: 5 (7).
Ecology. — Most specimens from sandy silt or a silty benthos.
Distribution. — -Cosmopolitan; this is the first report from the Gulf of California.
Praxillella affinis pacifica Berkeley
Praxillella affinis (Sars) var. pacifica. Berkeley, 1929: 313-314; Berkeley and Berkeley, 1952: 49-50;
Praxillella affinis pacifica. Hartman, 1961: 37; Reish, 1959a: 93.
Material. — April 1962: 54 (1040); October 1963: 32 (56).
Ecology. — -Occurred mainly on sand and silty sand bottoms. In October the population was nearly elimi-
nated in the outer part of the bay and was much reduced in the inner bay area. (Figs. 16, 17).
Distribution. — British Columbia to southern California; the distribution is extended into the Gulf of
California.
Family Oweniidae
Ouenia fusiformis collaris Hartman
Owenia fusiformis collaris. Hartman, 1955: 46; Reish, 1959a: 94.
Material. — April 1962: 2 (3); October 1963: 6 (8).
Ecology. — Sands.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Family Pectinariidae
Pectinaria (Pec/inaria) hartmanae, new species
Figure 18 a-e
Material. — April 1962: 6 (6).
Diagnosis. — Length 15-20 mm, width at anterior end 3-4 mm; cephalic spines brassy, number from 8 to
10 pairs, long, flattened, recurved, antennuated tips (Fig. 18a); antennular membrane with about 30 fili-
form fringes; about 30 varying sized oral tentacles; two pairs pectinate branchiae the anterior longer than
the posterior; setigerous segments 1 to 3 with only notopodial fascicle; the next 12 segments with both
components; notosetae of two types, both tapered to a point, one nearly smooth (Fig. 18b), the other
limbate, denticulated and incised (Fig. 18c); uncini in neuropodium wih 7 major teeth in two rows and
many smaller teeth medially (Fig. 18d); scaphal hcoks (Fig. 18e) number from 8 to 10, blunt, brassy in
color; anal tongue with median papillae and crenulated margin; tube, straight, length 2 5 mm; consists oi
coarse silicious grains.
Remarks. — -The subgenus Pectinaria was previously known from 18 species (Hartman, 195 9b). These
species can be grouped conveniently according to the number of segments with notosetae. Further groupings
may be made en the number of segments with uncini. P. hartmanae and P. nana Wesenberg-Lund (1949),
from the Gulf of Oman, are the only two species known with 15 pairs of notosetae. Both species have 12
uncinigerous segments. These two species arc separated from one another on the basis of 8-10 pans ,>t
cephalic spines in P. hartmanae and 11 pairs in P. nana, 8-10 pairs of scaphal hooks in the former .\ni.\
3 pairs in the latter, and antennular membrane with 30 fringes in the former and 8 in the latter.
Ecology. — The substrate consisted of silty sand at three of the stations and sand and sandy silt at the
other three.
Type material. — The holotype and three additional specimens have been deposited in the U.S. National
Museum.
Type locality. — -Bahia de los Angeles, Gulf of California, Baja California. This species is named in honor
of Dr. Olga Hartman in recognition of her contributions to the knowledge of polychaetes.
92
San Diego Society of Natural History
Vol. 15
Figure 18. — Pectinaria (Pectinar/a) hartmanae, n. sp.: a, cephalic spine; b, capillary
seta from notopodium; c, limbate seta with incised margin from notopodium; d, uncinus
from neuropodium; e, scaphal hook.
Family Ampharetidae
Amphisamytba bioculata (Moore)
Samytha bioculata. Moore, 1906: 25 3-25 5; Amphisamytba bioculata. Berkeley and Berkeley, 1952: 73.
Material. — April 1962: 6 (38); October 1963: 1 (1).
Ecology. — Preferred silty sands.
Distribution. — British Columbia, southern California, India; this is the first report from the Gulf of
California.
Amphicfcis scaphobranchiat a Moore
Ampbictcis scaphobrancbiata. Moore, 1906: 255-257; Berkeley and Berkeley, 1952: 68-69.
Material. — April 1962: 7 (23); October 1963: 7 (8).
Ecology. — Preferred silty sands.
Distribution. — British Columbia to southern California; the distribution is extended into the Gulf of
California.
Asabellides lineata (Berkeley and Berkeley)
Pseudosabellides lineata. Berkeley and Berkeley, 1943: 131-132; 1952: 71-72; 1956: 241: Asabellides lineata.
Hartman, 1961 : 39.
Material. — April 1962: 1 (1).
Ecology. — Sandy silts.
Distribution. — British Columbia, Alaska, Canadian Arctic, Hudson Bay (Berkeley and Berkeley, 1956)
and questionably southern California (Hartman, 1961). The distribution is extended into the Gulf of
California.
1968 Reish: Bahia de los Angeles Annelids 93
Family Terebclledae
Amaeana occidentals (Hartman)
Amaea occidentalis. Hartman, 1944b: 277-278; Amaeana occidentals. Hartman 1959: 495.
Material. — April, 1962: 34 (273); October 1963: 11 (15).
Ecology. — Most frequent on sandy silts but commonest on sands in April 1 962.
Distribution. — -Central and southern California; the distribution is extended into the Gulf of California.
Pista crhtata (Miiller)
Amphitrite crhtata. Miiller, 1776: 40; Pista crhtata. Fauvel, 1927: 266; Berkeley and Berkeley, 1952: 78-79.
Material. — April 1962: 3 (4); October 1963: 3 (9).
Ecology. — Sand or silty sands.
Distribution. — -Cosmopolitan; this is the first report from the Gulf of California.
Polycirris perplexus Moore
Polyeirris perplexus. Moore, 1923: 198-199; Hartman, 1961: 41.
Material. — April 1962: 31 (212).
Ecology. — Most specimens from sandy silt or silty substrates.
Distribution. — Southern California; the distribution is extended into the Gulf of California.
Family Trichobranchidae
Terebellides stroemi Sars
Terebellides stroemi. Sars, 1835: 48, Berkeley and Berkeley, 1939: 343; Reish, 1959b: 39; Imajima and
Hartman, 1964: 3 52-3 5 3.
Material. — April 1962: 7 (19); October 1963: 3 (3).
Ecology. — All substrates.
Distribution. — Cosmopolitan; Berkeley and Berkeley (1939) reported it from Puerto San Carlos (near
Guaymas) .
Family Sabellidae
Cbone mollis (Bush)
Metachone mollis. Bush, 1904: 216; Chone mollis, Hartman, 1944b: 279-280; Reish, 1963a: 430.
Material. — April 1962: 55 (553); October 1963: 14 (34).
Ecology. — Prefers coarser sediments (Table 2).
Distribution. — California and San Quintin Bay; the distribution is extended into the Gulf of California.
Euchone barnardi, new species
Figure 19 a-d
Material. — April 1962: 35 (663); October 1963: 2 (2).
Diagnosis. — -Maximum length of 2.0mm including branchial length of 0.5mm; 8 thoracic and 9 abdominal
setigerous segments, the last 3 comprise the anal depression; branchiae with 3 radioles per side, with num-
erous filaments which extend within 0.1mm of the tip, united for one-half of their length; collar little
developed dorsally but produced into 2 lobes ventrally; thoracic notopodium with 2-3 superior double-
winged capillary setae (Fig. 19a) and 3-4 inferior subspatulate setae (Fig. 19b); 3-4 long handled uncini
in thoracic neuropodium, each with a large tooth and 4-5 smaller teeth (Fig. 19c); abdominal notopodium
with 5-6 avicular hooks, each with larger tooth and 15-20 teeth (Fig. 19d); abdominal neuropodium in-
cluding and depression with 4-5 simple capillary setae.
Remarks: — Hartman (1959b) lists 13 and possibly 14 species in the genus Euchone Malmgren. Addi-
tional species described include E. limnicola Reish (1960), E. incolot Hartman (1965a), and E. tri
men tat a Reish (1965). Euchone barnardi comes closest to E. trisegtnentata, E. rosea Langerhans, and
E. incolot. All are small and the three former have only 17 setigerous segments; E. incolot possesses from
16-19 setigerous segments. The anal depression consists of 4 segments in E. rosea: it consists <>t 3 segments
in E. barnardi, E. trisegtnentata and E. incolor. The margins of the anal depression are flared m / . incolot
but not in E. barnardi or E. trisegtnentata. Euchone barnardi and E. trisegtnentata have i pairs ol radioles;
E. incolor has 3 to 4 pairs of radioles. These species all have similar setae but differ in details: superior
thoracic notopodial setae double-winged in £. barnardi and £. incolor, single in E. trisi %mentata; thoracic
neuropodia long-handled hooks with 5-6 teeth in E. barnardi. 4-5 in E. incolor, and 7 in E. trisegtnentata,
abdominal avicular hooks with 6 rows of teeth in E. barnardi, 7 in E. incolor, and 10 in £. trisegtnentata.
Ecology. — All but three of specimens were taken in the April 1962 survey. This species did not exhibit
any preference for a particular substrate type.
Type material. — The hclotype and paratypes have been deposited in the U.S. Xation.il Museum.
Type locality. — Bahia de los Angeles, Gulf of California, Baja California.
This species is named in honor of Dr. J. Laurens Barnard, in appreciation for making these collections
available for study.
94
San Diego Society of Natural History
Vol. 15
.014 mm
Figure 19. — Euchone barnardi, n. sp.: a, double winged capillary seta from superior
thoracic notopodium; b, subspatulate seta from inferior thoracic notopodium; c, long
handle uncinus from thoracic neuropodium, magnification as figure b; d, avicular hook
from abdominal notopodium.
Euchone cortezi, new species
Figure 20 a-e
Material. — April 1962: 2 (2). Both stations located near Isla Ventana in Bahia de los Angeles.
Diagnosis. — Length 2 to 6mm including the branchial length of 0.5 to 1.5mm; 8 thoracic and 12 to 13
abdominal setigerous segments, the last 5 in two and last 6 in one specimen comprise the anal depression
(Fig. 20a) ; branchiae with 7 radioles per side, united with a membrane for one-half of their length, and
the filaments extending to near the tip of the radiole; collar well developed and divided into two lobes
with incised margins along the mid-dorsal and mid-ventral line; thoracic notopodium with superior doubled-
winged capillary setae (Fig. 20b) and inferior spatulate setae (Fig. 20c); thoracic neuropodium with long
handled uncini (Fig. 20d) each with a large tooth and 3 smaller teeth; abdominal notopodium (Fig. 20e),
provided with avicular hooks each with a large tooth and 2 5 to 30 smaller teeth in 6 rows; abdominal
neuropodium including the anal depression with simple capillary setae.
Remarks. — Of the 16 to 17 known species in the genus Euchone (see under E. barnardi above), E. cortezi
comes closest to E. papillosa (Sars) (Malmgren, 1865) on the basis of the number of segments in the anal
depression. Euchone cortezi has 5 to 6 segments in the region and E. papilloma has approximately 6; how-
ever, the two species can be readily separated on the basis of the number of segments; there are 20 to 21
in the former and 33 to 34 in the latter. The inferior spatulate setae also differ.
The specific name refers to the Sea of Cortez, a name used for the Gulf of California in the past.
Type material. — The holotype and one paratype have been deposited in the U.S. National Museum.
Type locality. — Near Isla Ventana in Bahia de los Angeles, Gulf of California.
1968
Reish: Bahia de los Angeles Annelids
9S
\
.014 mm
.003 mm
Figure 20. — Euchone cortezi, n. sp.: a, posterior end showing anal depression; b, double
winged capillary seta from superior thoracic notopodium; c, subspatulate seta from
inferior thoracic notopodium, magnification as figure b; d, long handle uncinus from
thoracic neuropodium; e, avicular hook from abdominal notopodium, magnification as
figure d.
Fabricia limnicola Hartman
Fabricia limnicola. Hartman, 195 1b: 384-385; Reish, 1963a: 430.
Material. — October 1963: 2 (44).
Ecology. — Silty sands and sands.
Distribution. — Southern California and San Quintin Bay; the distribution is extended into the Cult <>t
California.
Megalomiiiii pigmentum Reish
Mcgalomma pigment ion. Reish, 1963a: 430-433.
Material. — April 1962: 22 (68); October 1963: 13 (23).
Ecology. — -This species showed a preference for sands and silty sands (Table 2); at San Quintin I5.i\
M. pigmentum was more commonly taken from finer sediments (Reish, 1963a).
Distribution. — Southern California and San Quintin Bay; the distribution is extended into the Gull ol
California.
Family Serpulidae
Hydroides recurvispina Rioja
Hydroides recurvispina. Rioja, 1941: 169-172.
Material. — April 1962: 3 (5); October 1963: 1 (1).
Ecology. — Majority from sands.
Distribution. — • Mazatlan and Acapulco; the distribution is extended northward into the Gulf of California.
96 San Diego Society of Natural History Vol. 1 5
Literature Cited
Alikunhi, K. H
1943. On an undescribed hermaphrodite species of Microphthalmia (Hesionidae) occurring on the
sandy beach at Madras. Indian Sci. Congr., Proc. 29: 149-150.
Baird, W.
1863. Descriptions of several new species of worms belonging to the Annelida Errantia and Sedentaria
or Tubicola of Milne Edwards. Zool. Soc. London, Proc. 1863, p. 106-110.
Barnard, J. L., and W. E. Grady
1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. I. General Account.
San Diego Soc. Nat. Hist., Trans. 15 (6): 51-66.
Berkeley, E.
1929. Polychaetous annelids from the Nanaimo district. 4. Chaetoptcridae to Maldanidae. Contr. Can.
Biol. Fish., new ser., 4: 305-316.
Berkeley, E., and C. Berkeley
193 6. Notes on Polychaeta from the coast of western Canada. I. Spionidae. Ann. Mag. Natur. Hist.,
ser. 10, 18: 468-477.
193 8. Notes on Polychaeta from the coast of western Canada. II. Syllidae. Ann. Mag. Natur. Hist.,
ser. 11,1: 3 3-49.
193 9. On a collection of Polychaeta, chiefly from the west coast of Mexico. Ann. Mag. Natur. Hist.,
ser. 11, 3: 321-346.
1941. On a collection of Polychaeta from Southern California. So. Calif. Acad. Sci., Bull. 40: 16-60.
1943. Biological and oceanographical conditions in Hudson Bay. 11. Polychaeta from Hudson Bay.
J. Fish. Res. Bd. Can. 6: 129-132.
1952. Annelida, Polychaeta sedentaria. In Canadian Pacific Fauna. J. Fish. Res. Bd. Can. no. 9b:
1-139.
Busch, W.
185 1. Beobachtungen iiber Anatomie und Entwicklung einiger wirbellosen Seethiere. Berlin, A.
Hirschwald. 143 p.
Bush, K. J.
1904. Tubicolous annelids of the tribes Sabellides and Serpulides from the Pacific Ocean. Harriman
Alaska Exped., N. Y., 12: 169-3 5 5.
Chamberlin, R. V.
1919a. The Annelida Polychaeta. Mem. Mus. Comp. Zool. Harvard, 48: 1-5 14.
1919b. New polychaetous annelids from Laguna Beach, California. J. Entomol. Zool. Pomona, 11: 1-23.
Chiaje, S. delle
1828. Memorie sulla storie e notomia degli animali senza vertebre del Regno di Napoli, 3: 1-232.
Claparede, E.
1863. Beobachtungen iiber Anatomie und Entwicklungsgeschichte wirbelloser thiere an der Kiiste
von Normandie angestellt. p. 1-120.
1868. Les annelides, chetopodes du Golfe de Naples. Mem. Soc. Phys. Geneve, 19: 313-584.
1870. Les annelides chetopodes du Golfe de Naples. Supplement, Mem. Soc. Phys. Geneve, 20:
365-452.
Day, J. H.
1954. The Polychaeta of Tristan da Cunha. In Results of the Norwegian Scientific Expedition to
Tristan da Cunha 1937-1938.
DUJARDIN, F.
1839. Observations sur quelques annelides marins. Ann. Sci. Natur. Paris, sec 2, 11: 281-294.
Ehlers, E.
1868. Die Bostenwurmer (Annelida Chaetopoda) nach systematischcn und anatomischen Unter-
suchungen dargestellt. Leipzig, Wilhelm Englemann. Vol. 2, p. 269-748.
1901. Die Polychaeten des magellanischen und chilenischen Strandes. Ein faunistischer Versuch.
Festschrift zur Feier des Hundertf iinfzigjahrigen Bestehous der koniglichen Gesellschaft der
wissenschaften zu Gottingen. (Abl. Math. -Phys.) . Berlin, Wiedmannsche Buchlandlung.
232 p.
Fabricius, O.
1780. Fauna groenlandica .... 452 p.
Fauvel, P.
1927. Polychetes sedentaires. Faune de France, 16: 1-494.
1943. Annelides polychetes de California recueillies par L. Diguet. Mem. Mus. Hist. Natur. Paris,
n. s. 18: 1-32.
195 3. Annelida Polychaeta. The Fauna of India including Pakistan, Ceylon, Burma and Malaya.
5 07 p.
Graviir, C.
1905. Sur un Polynoidicn {Lepid asthenia digueti nov. sp.) commensal d'un Balanoglosse de Basse-
Californie. Bull. Mus. Natur. Paris, 11: 177-181.
1968 Reish: Bahia de los Angeles Annelids 97
Grube, A.-E.
1840. Actinien, Echinodermen und Wurmen des Adratischcn und Mittelmeers. . . ., p. 61-81.
18 57. Annulata Oerstediana, pt. 2, p. 15 8-166.
1863. Beschrieburg neuer oder wenig bekanntcr Anneliden, Sechster Beitrag. Arch. Naturg. 29:
37-69.
Hartman, O.
1936. A review of the Phyllodocidae (Annelida polychaeta) of the coast of California, with
descriptions of nine new species. Univ. Calif. Publ. Zool. 41: 117-132.
193 8. Descriptions of new species and new generic records of polychaetous annelids from California
of the families Glyceridae, Eunicidae, Stauronereidae and Opheliidae. Univ. Calif. Publ. /
43: 93-1 1 1.
1939a. Polychaetous annelids, Pt. I. Aphroditidae to Pisionidae. Allan Hancock Pacific Exped. 7: 1-15 6.
1939b. New species of polychaetous annelids from southern California. Allan Hancock Pacific
Exped. 7: 157-172.
1940. Polychaetous annelids, Pt. II. Chrysopetalidae to Goniadidae. Allan Hancock Pacific Expc.l.
7: 173-287.
1941. Some contributions to the biology and life history of Spionidae from California. Allan Han-
cock Pacific Exped. 7: 289-324.
1944a. Polychaetous annelids, Pt. V. Eunicea. Allan Hancock Pacific Exped. 10: 1-238.
1944b. Polychaetous annelids from California, including the description of two new genera and nine
new species. Allan Hancock Pacific Exped. 10: 239-307.
1944c. Polychaetous annelids, Pt. VI. Paraonidae, Magelonidae, Longosomidae, Ctenodrilidae and
Sabellariidae. Allan Hancock Pacific Exped. 10: 311-389.
1947a. Polychaetous annelids, Pt. VII. Capitellidae. Allan Hancock Pacific Exped. 10: 391-481.
1947b. Polychaetous annelids, Pt. VIII. Pilargiidae. Allan Hancock Pacific Exped. 10: 482-52 3.
1948. The marine annelids erected by Kinberg with notes on some other types in the Swedish State
Museum. Ark. Zool. Svenska Vetensk. 42 A: 1-137.
1950. Goniadidae, Glyceridae and Nephtyidae. Allan Hancock Pacific Exped. 15: 1-180.
195 1a. The littoral marine annelids of the Gulf of Mexico. Publ. Inst. Marine Sci. 2: 7-124.
195 1b. Fabricinae (feather-duster polychaetous annelids) in the Pacific. Pacific Sci. 5: 379-389.
1954. Australian Nereidae, including descriptions of three new species and one new genus, together
with summaries of previous records and keys to species. Trans. Roy. Soc. So. Austr. 77: 1-41.
195 5. Endemism in the North Pacific Ocean, with emphasis on the distribution of marine annelids,
and descriptions of new or little known species, p. 3 9-60. In Essays in the Natural Sciences
in honor of Captain Allan Hancock, Los Angeles, Calif.
1957. Orbiniidae, Apistobranchidae, Paraonidae and Longosomidae. Allan Hancock Pacific Exped.
15: 211-392.
195 9. Catalogue of the polychaetous annelids of the world, parts 1-2. Allan Hancock Foundation
Publ. Occ. Pap. 23.
1960. Systematic account of some marine invertebrate animals from the deep basins off southern
California. Allan Hancock Pacific Exped. 22: 69-214.
1961. Polychaetous annelids from California. Allan Hancock Pacific Exped. 25: 1-226.
1963. Submarine canyons of southern California, Pt. III. Systematics: Polychaetes. Allan Hancock
Pacific Exped. 27: 1-93.
1965a. Deep-water benthic polychaetous annelids off New England to Bermuda and other North
Atlantic areas. Allan Hancock Foundation Publ. Occ. Pap. 28.
1965b. Supplement and index to the catalogue of the polychaetous annelids of the world, including
additions and emendations since 1959. Allan Hancock Foundation Publ. Occ. Pap. 23.
Hartmann-Schroder, G.
1962a. Zweiter Beitrag zur Polychaeten fauna von Peru. Kieler Meeresforsch. 18: 109-147.
1962b. Zur Kenntnis des Eulitorals der chilenischen Pazifikkuste und der argentinischen Kuste Sud-
patagoniens unter Ostracoden. Teil II. Die Polychaeten des Eulitorals. Mitt. Hamb. Zool. Mus.
Inst. 60: 57-167.
Imajima, M., and O. Hartman
1964. The polychaetous annelids of Japan. Allan Hanei ck Foundation Publ. Occ. Pap. 26.
Johnson, H. P.
1897. A preliminary account of the marine annelids of the Pacific coast with descriptions oi new
species. Calif. Acad. Sci. Proc, Zool. scr. 3, 1: 15 3-198.
1901. The polychaeta of the Puget Sound region. Boston Soc. Natur. Hist. Proc. 29: 381-437.
Kinberg, J. H.
1866. Annulata nova. Fbrh. Ofv. vet. Akad. Stockholm, 22: 167-179.
Laubier, L.
1962. Quelques annelides polychetes de la lagune de Venisc, description de Prionospio caspersi n. sp.
Vie et Milieu, 13:123-159.
98 San Diego Society of Natural History Vol. 15
LlIDY, J
185 5. Contributions toward a knowledge of the marine invertebrates of the coasts of Rhode Island
and New Jersey. J. Acad. Natur. Sci. Phila. 3: 135-158.
Malmgren, A. J.
1865. Nordiska Hafs-Annulater. Forh Ofv. Kongl. vet. Akad. Stockholm, 1: 51-110; 2: 181-192;
5: 355-410.
Mecznikow, E.
1865. Beitrage zur Kenntnis der Chaetopoden. Zeitschr. Wiss. Zool., 15: 328-341.
Monro, C. C. A.
1928. On the polychaeta collected by Dr. H. Montensen off the coast of Panama. Vidensk. Medd.
Natur. Foren. Kjdbenhavn. 85: 75-103.
Moore, J. P.
1904. New polychaetes from California. Acad. Natur. Sci. Phila., Proc. 56: 484-503.
1906. Additional new species of polychaeta from the north Pacific. Acad. Natur. Sci. Phila., Proc.
58: 217-260.
1907. Descriptions of new species of spioniform annelids. Acad. Natur. Sci. Phila., Proc. 59: 195-207.
1908. Some polychaetous annelids of the northern Pacific coast of North America. Acad. Natur.
Sci. Phila., Proc. 60: 321-364.
1909a. Polychaetous annelids from Monterey Bay and San Diego. Acad. Natur. Sci. Phila., Proc.
61: 225-295.
1909b. The polychaetous annelids by the U. S. S. Albatross off the coast of southern California in
1904. I. Syllidae, Sphaerodoridae, Hesionidae and Phyllodocidae. Acad. Natur. Sci. Phila.,
Proc. 61: 321-351.
1910. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of southern Cali-
fornia in 1904. II. Polynoidae, Aphroditidae and Segalionidae. Acad. Natur. Sci. Phila., Proc.
62: 328-402.
1911. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of southern Cali-
fornia in 1904. Euphrosynidae to Goniadidae. Acad. Natur. Sci. Phila., Proc. 63: 234-3 18.
1923. The polychaetous annelids dredged by the U. S. S. Albatross off the coast of southern Cali-
fornia in 1904. Spionidae to Sabellariidae. Acad. Natur. Sci. Phila., Proc. 75: 179-259.
MULLER, O. F.
1776. Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae .... 274 p.
Oersted, A. S.
1843. Groenlands Annulata dorsibranchiata. K. Danske Videns. Selsk. Natur. Math. 10: 155-216.
Parker, R. H.
1963. Zoogeography and ecology of some macroinvertebrates, particularly mollusks, in the Gulf of
California and the continental slope off Mexico. Vidensk. Medd. fra Dansk natur. Foren.
126: 1-178.
Pettibone, M. H.
1954. Marine polychaete worms from Point Barrow, Alaska, with additional records from the
North Atlantic and North Pacific. US. Nat. Mus., Proc. 103: 213-356.
1963. Marine polychaeta worms of the New England region. I. Aphroditidae through Trochochae-
tidae. U. S. Nat. Mus., Bull. 227.
Potts, F. A.
1910. Polychaeta of the Indian Ocean. Pt. II. The Palmyridae, Aphroditidae, Polynoidae, Acoetidae
and Sigalionidae. Trans. Linn. Soc. Zool., sec. 2, 16: 325-353.
Reish, D. J.
195 9a. Benthic polychaetous annelids. Pt. III. Allan Hancock Foundation Publ. Occ. Pap. 21,
p. 70-106.
1959b. An ecological study of pollution in Los Angeles-Long Beach Harbors, California. Allan Han-
cock Foundation Publ. Occ. Pap. 22.
1959c. New species of Spionidae (Annelida, Polychaeta) from southern California. So. Calif. Acad.
Sci., Bull. 58: 11-16.
1960. A new species of Sabellidae (Annelida, Polychaeta) from southern California. Ann. Mag.
Natur. Hist. London, ser. 13, 2: 717-719.
1963a. A quantitative study of the benyhic polychaetous annelids of Bahia de San Quintin,
Baja California. Pacific Natur. 3: 399-436.
1963b. Mass mortality of marine organisms attributed to the "red tide" in southern California. Calif.
Fish and Game, 49: 265-270.
1965. Benthic polychaetous annelids from Bering, Chukchi, and Beaufort Seas. U.S. Nat. Mus., Proc.
117: 13 1-158.
Reish, D. J., and J. L. Barnard
1967. The benthic polychaeta and amphipoda of Morro Bay, California. U.S. Nat Mus., Proc.
120: 1-26.
1968 Reish: Bahia de los Angeles Annelids 99
Rioja, E.
193 9. Estudios anelidologicos I. Observaciones acerca dc varias formas larvarias y postlarvarias
pelagicas de Spionidac, precedentes de Acapulco, con descripcion dc una especie neuva del genero
Polydora. Ann. Inst. Biol. Mexico. 10: 297-311.
1941. Estudios anelidologicos III. Datus par el conocimiento de la fauna de poliquetos de Lis costas
del Pacifico de Mexico. Ann. Inst. Biol. Mexico. 12: 669-746.
1943. Estudios anelicologicos VI. Aportaciones al concimiento de los I xogoninos (Anelides poliquel
de las costas del Pacifico. Ann. Inst. Biol. Mexico. 14: 207-227.
1944. Estudios anelidologicos XI. Notas sobre algunas especies de poliquetos de la costas Mexicanas
del Pacifico. Ann. Inst. Biol. Mexico. IS: 13 9-145.
1 947a. Estudios anelidologicos XVII. Contribucion as concocimiento de los anelidos poliquetos de
Baja California y mar de Cortes. Ann. Inst. Biol. Mexico. 18: 197-224.
1947b. Estudios anelidologicos XVIII. Observaciones y datos sobre algunos anelidos poliquetos de Golfo
de California y costas de Baja California. Ann. Inst. Biol. Mexico. 18: 517-526.
195 9. Estudios anelidologicos XXIII. Contribucion al conocimiento de las Islas de Revillagigeda. Ann.
Inst. Biol. Mexico. 30: 243-2 5 9.
Sars, M.
1835. Beskrivelsev og iagttagelser over nogle mockelige eller nye i havet ved den Bergcske kyst
levende dyr. ... 81 p.
185 1. Beretning om en i sommeren 1849 fortagen zoologisk reise i loften og fin marken, Nyt. Mag.
Naturv. 6: 121-211.
SCHMARDA, L. K.
1861. Neue wirbellose tliiere becbachtet und gesammelt aug in einer Reise um die Erde 1853 bis
18 57. Leipzig, vol. 1, Turbellarien, Rotatorien und Anneliden, pt. 2, p. 1-164.
Southern, R.
1914. Archiannelida and Polychaeta. /;/ Clare Island survey, pt. 47. Roy. Irish Acad. Dublin,
Proc. 31: 1-160.
Steinbeck, J., and E. F. Ricketts
1941. Sea of Cortez. A leisurely journal of travel and research. With a scientific appendix ....
Viking Press, New York. 5 98 p.
Tebble, N.
1960. The distribution of pelagic polychaetes in the South Atlantic. Discovery Repts. 30: 161-300.
Treadnxell, A. L.
1901. The polychaetous annelids of Porto Rico. U.S. Fish Comm. Bull. 20: 183-210.
1914. Polychaetous annelids of the Pacific coast in the collection of the zoological museum of the
University of California. Univ. Calif. Publ. Zool. 13: 175-234.
1923. Polychaetous annelids from Lower California with descriptions of new species. Amer. Mus.
Novitates, 74.
1929. New species of polychaetous annelids in the collections of the American Museum of Natural
History, from Porto Rico, Florida, Lower California, and British Somaliland. Amer. Mus.
Novitates, 392.
1937. The Templeton Crocker Expedition. VIII. Polychaetous annelids from the west coast of
Lower California, the Gulf of California and Clarion Island. Zoologica, 22: 139-160.
1942. Polychaetous annelids from Lower California and the Philippine Islands in the collection of
the American Museum of Natural History. Amer. Mus. Novitates. 1172.
USCHAKOV, P. V.
1955. [Polychaetes from the seas in the far East.] Akademia Nauk SSSR. Opredeliteli po Faune
USSR, no. 56, 445 p. [in Russian].
Wesenberg-Lund, E.
1949. Polychaetes of the Iranian Gulf. Danish Sci. Invest. Iran, pt. 4, p. 247-400.
Wiren, A.
1883. Chaetopoder from Sibiriska Ishafvet och Berings Haf insamlade uner Vegtf-Expeditionen
1878-79. Vega-Exped.-Vetenskapliga Iakttagelser, 2: 383-428.
Woodwick, K. H.
1961. Polydora Ticket hi, a new species of spionid polychaete from Lower California. Pacific Sci.,
15: 78-81.
1964. Polydora and related genera (Annelida Polychaeta) from Eniwetok, Majuro. and Bikini Atolls,
Marshall Islands. Pacific Sci., 18: 146-15 9.
Zachs, I.
1925. Nouvelles additions a la faune des Polychaeta du Nauman. Acad. Sci. Russ. Leningrad,
C. R., sec. A, p. 1-3.
Accepted for Publication 27 February 1968
Department of Biology, California State College at Long Beach, California.
100 San Diego Society of Natural History Vol. 15
Appendix
Table 1
Polychaetes reported from the Gulf of California. An asterisk denotes new distri-
butional records for the area. The names listed are those accepted by the author cited.
Aphroditidae
Aphrodita japonica Marenzeller. Hartman, 1939a. Isla Partida; Isla Espiritu Santo.
A. pari a Moore. Hartman, 193 9a. Bahia de San Ignacio; Sinaloa.
Pontogenia laevheta Hartman, 1939a. Isla Angel de la Guarda.
Polynoidae
Arctonoc littata (Grube). Treadwell, 1923. La Paz.
Chaetocanthus magnificus (Grube). Treadwell, 1937; Berkeley and Berkeley, 1939; Hartman, 1959.
Isla Espiritu Santo; Banco Arena.
Halosydna breihetosa Kinberg. Fauvel, 1943. Gulf.
H. fuscomarmorata (Grube). Fauvel, 1943. Gulf.
H. glabra Hartman, 1939a; Fauvel, 1943; Steinbeck and Ricketts, 1941. Bahia de la Concepcion,
Cabo Pulmo, Gulf.
H. Johnson/ (Darboux). Berkeley and Berkeley, 193 9. Georges Is.; Punta Penasco, Sonora; San Felipe.
H. latior Chantberlin. Rioja, 1947a. Topolobampo; Bahia de Navachiste.
Harmothoe hirsuta Johnson. Rioja, 1947a. Topolobampo.
Hololcpr In veleronh Hartman, 193 9a. Bahia de San Francisquito; Isla Angel de la Guarda.
Iphione muricata (Savigny). Fauvel, 1943. Gulf.
/. oiata Kinberg. Hartman, 1939a; Steinbeck and Ricketts, 1941, west side of Gulf between
24° and 26° W. Latitude.
Lepidamctria gigas (Johnson). Berkeley and Berkeley, 193 9. San Felipe.
Lepid 'asthenia digueti Gravier, 1905; Fauvel, 1943. La Paz and Gulf.
L. ornata Chamberlin. Treadwell, 1937. Banco Arena.
Lepidonotns caelorus Moore. Treadwell, 1937; Rioja, 1947a. Banco Arena and La Paz.
L. claia (Montagu). Fauvel, 1943. La Paz.
L. hupfcri Augener. Hartman, 1939a; Steinbeck and Ricketts, 1941; Rioja, 1947a. Estero de Agia-
bampo; Bahia de la Concepcion; La Paz.
L. purpureas Potts. Berkeley and Berkeley, 1939; Hartman, 1959. Isla Angel de la Guarda.
L. lersicolor Ehlers. Hartman, 1939a; Rioja, 1947a. Bahia de San Francisquito; Isla San Esteban;
Topolobampo.
Thormora johnstoni (Kinberg). Hartman, 1939a; Steinbeck and Ricketts, 1941; Rioja, 1947a. Baja
Calif, side of Gulf between 24° and 29° W\ Lattiude.
Polyodontidae
Panfhalis adumbrata Hoagland. Treadwell, 1937. Bahia de Santa Inez.
*P. pacifica Treadwell.
*Polyodonfcs from Hartman.
P. oculea (Treadwell). Hartman, 1939a; Steinbeck and Ricketts, 1941. Pichilinque Harbor, Estero de
Agiabampo.
Sigalionidae
Eupholoe philippineinis Mcintosh. Berkeley and Berkeley, 193 9. Isla Espiritu Santo.
''Sthenolepis fimhriarum Hartman; 1939a. Isla del Carmen, near Isla Espiritu Santo.
Psanimolyce myops Hartman, 1939a. Near Isla Espiritu Santo.
Sthenelah maculata Hartman, 1939a. Isla Espiritu Santo.
S. neoleanirae Hartman, 193 9a. Isla del Carmen.
*Stenelanella uniformh Moore.
Thalenessa leuisii (Berkeley and Berkeley), 1939; Hartman, 1939a; Steinbeck and Ricketts, 1941;
Fauvel 1943. Cabo San Lucas, Isla Espiritu Santo, Bahia de los Angeles.
Pareulepidae
*Pareulepu fimbriata (Treadwell).
Chrysopetalidae
Bhawania riveti (Gravier). Steinbeck and Ricketts, 1941. La Paz.
*Paleanotus bellis (Johnson). Bahia de los Angeles.
P. chrysoleph Schmarda. Rioja, 1947a. La Paz.
/'. purpurea Rioja, 1947b. La Paz.
Amphionomidae
Chloeia conspicua Horst. Rioja, 1943. Gulf.
C. entypa Chamberlin. Treadwell, 1937; Rioja, 1947a. Cabo San Lucus, La Paz, Bahia de Santa Inez,
Topolobampo.
1968 Reish: Bahia de los Angeles Annelids 10 1
C. flat a (Pallas). Treadwell, 1923. Bahia de San Francisquito.
C. pinnata Moore. Berkeley and Berkeley, 1939. Isla Espiritu Santo.
(?)C. rosea Potts [sic]. Fauvel, 1943. Gulf.
C. viridis Schmarda. Hartman, 1939; Berkeley and Berkeley. 1939; Steinbeck and Ricketts, 1941.
Common in Gulf.
Enrythoe complanata (Pallas). Hartman, 1939a; Berkeley and Berkeley, 1939; Steinbeck and Ricketts,
1941; Fauvel, 1943; Rioja, 1947a. Common in Gull.
(?)£. dubia Horst [sic]. Fauvel, 1943. Gulf.
Notopygos hispida Potts. Fauvel, 1943. Gulf.
N. ornata Grube. Hartman, 1939a; Berkeley and Berkeley, 1939; Steinbeck and Ricketts, 1941; Rioja,
1947a. Common in Gulf.
■ Pareurythoe californica (Johnson).
Euphrosinidae
Euphrosine aurantiaca Johnson. Rioja, 1947a. La Pa
£. bicirrata Moore. Hartman, 193 9a. Isla Partida.
Phyllodocidae
Anaitides lamellifera (Pallas). Fauvel, 1943. Gulf.
A. madeirensis (Langerhans). Treadwell, 1937; Steinbeck and Ricketts, 1941; Fauvel, 1943. Common
in Gulf.
* A. mucosa (Oersted).
A. panamensis (Treadwell). Fauvel, 1943. La Paz.
*A. williamsi Hartman.
*Eteone dilatae Hartman.
Eulalia myriacyclum (Schmarda). Steinbeck and Ricketts, 1941. Puerto Refugio.
Eumida sanguine a (Oersted). Rioja, 1947a. La Paz.
*Paranaitis polynoides (Moore).
Vhyllodoce digue ti Fauvel, 1943. Gulf.
Typhloscolecidae
'■'Typbloscolex mulleri Busch.
Hesionidae
Hesionc genetta Grube. Fauvel, 1943. Gulf.
H. in /err ex ta Grube. Berkeley and Berkeley, 193 9; Hartman, 1940; Rioja, 1947a. Common in Gulf.
//. pant hernia Risso. Fauvel, 1943. Isla San Jose.
Leocrates chinensis Kinberg. Hartman, 1940. Isla Espiritu Santo.
*Microphthalmus riojai n. sp.
Opbiodromus pugettensis (Johnson). Steinbeck and Ricketts, 1941; Rioja, 1947a. Cabo Pulmo,
Topolobampo.
*Oxydronms brunnea Hartman.
"'O. arenicolus glabrus Hartman.
*Syllidia liniata Hartmann-Schroder.
Pilargidae
* Ancistrosyllis bassi Hartman.
*Loandalia fanieli Berkeley and Berkeley.
"Pilargh hamatus Hartman.
Syllidae
Autolytus varius Treadwell. Rioja, 1947a. La Paz, Bahia de Navachiste.
"Brania limbata (Claparede).
"'Exogone lonrei Berkeley and Berkeley.
* Lunger hansia heterochaeta (Moore).
Odontosyllis pbosphorea Moore. Rioja, 1947a. La Paz.
*Pionosyllis gi gun tea Moore.
*Sphaerosyllis erinaceus Claparede.
Syllis elongata (Johnson). Rioja, 1947a. La Paz, Topolobampo.
Typosyllis byalina (Grube). Rioja, 1947a. La Paz.
Nereid ae
Ceratonereis mirabilis Kinberg. Treadwell, 1929; Hartman, 1940; Steinbeck and Ricketts. [941;
Fauvel, 1943; Rioja, 1947a. Common in Gulf.
C. paucidentata (Moore). Fauvel, 1943. Gulf, La Paz.
Neantbes sue cine a (Frey and Leuckart). Rioja, 1947a. Estero de Agiabampo; Topolobampo.
Nereis gritbei (Kinberg). Fauvel, 1943. Gulf.
*N. procera Ehlers.
N. rata Ehlers. Berkeley and Berkeley, 193 9. Isla Espiritu Santo.
N. riisei Grube. Hartman, 1940; Fauvel, 1943. Gulf.
N. zonata Malmgren. Hartman, 1940. Isla San Esteban.
102 San Diego Society of Natural History Vol. 15
Nicon mexicana. (Treadwell). Treadwell, 1942. Topolobampo, Isla San Jose.
N. moniloceras (Hartman). Hartman, 1940. Isla San Esteban.
Platynereis bicanaliculata (Baird). Treadwell, 1914; Hartman, 1940; Steinbeck and Ricketts, 1941.
Common in Gulf.
P. polyscalma Chamberlin. Hartman, 1940; Steinbeck and Ricketts, 1941. Between 23° and 27° W.
Latitude.
Pseud cmereis gallapagensis Kinberg. Fauvel, 1943. Gulf.
Nephtyidae
Aglaophamus dicirris Hartman. Hartman, 1940. Common in Gulf.
A. dibranchh (Grube). Treadwell, 1937; Hartman, 1940. Bahia de San Francisquito, Isla del Carmen.
Cabo San Lucas.
A. iiicrmis Ehlers. Hartman, 1940. Isla Espiritu Santo.
Nephfys caecoides Hartman. Hartman, 1940. Isla Partida.
N. magellanica Augener. Hartman, 1940. Common in Gulf.
N. panamensis Monro. Hartman, 1940. Between 24° and 29° W. Latitude.
N. singularis Hartman, 1940. Cabeza Ballona.
N. squamosa Ehlers. Hartman, 1940, 195 0. Common in Gulf.
Glyceridae
Glycera americana Leidy. Hartman, 1940. Common in Gulf.
G. capitata Oersted. Hartman, 195 0. Arroyo de San Luis.
G. capitata branchiopoda Moore. Chamberlin 1919a. 26° 48' W. Latitude, 110° 45' 20" N. Longitude.
G. dibranchiata Ehlers. Steinbeck and Ricketts 1941; Fauvel 1943. Gulf.
G. mexicana (Chamberlin). Chamberlin, 1919a. 27° 39' 40" W. Latitude, 1110 0' 30" N. Longitude.
*G. robusta Ehlers.
G. rugosa Johnson. Treadwell, 1937. Bahia de Santa Ynez.
G. tcsselata Grube. Hartman, 1940; Rioja, 1947a. Common in Gulf.
Goniadidae
*Glycinde armigera Moore.
Goniada acicula Hartman, 1940. Between 24° 22' W. Latitude and 29° 33' N. Longitude.
*G. lifforea Hartman.
Onuphidae
Diopatra chiliensis Quatrefages. Fauvel, 1943. Gulf.
D. neapolitana delle Chiaje. Fauvel, 1943. Gulf.
D. neotridens Hartman, 1944a. Isla Angel de la Guarda.
D. obliqua Hartman, 1944a. Between 27° 57' W. and 31° 19' W. Latitude.
D. ornata Moore. Berkeley and Berkeley, 193 9. Bahia de las Animas.
D. splcndissima Kinberg. Rioja, 1947a. La Paz, Topolobampo.
D. tridentata Hartman, 1944a; Rioja 1947b, Gulf.
Hyalinoecia juvenalis Moore. Treadwell, 1937; Hartman, 1944a; Rioja, 1947b. Common in Gulf.
Nothria stigmatis cirrata Hartman, 1944a. Bahia Tepoca.
Onuphis eremita Audouin and Milne Edwards. Rioja, 1947b. Topolobampo.
O. nannognathm Chamberlin, 1919a. 25° 59' W. Latitude and 108° 40' N. Longitude.
*0. nebulosa Moore.
O. vexillaria Moore, 1911. Estado de Sonora.
O. zebra Berkeley and Berkeley, 1939; Rioja, 1947a. La Paz.
Eunicidae
Eunice ajra Peters. Steinbeck and Ricketts, 1941; Hartman, 1944a. Isla Espiritu Santo.
E. a. paupera Grube. Fauvel, 1943. Gulf.
E. antennata (Savigny). Berkeley and Berkeley, 193 9; Steinbeck and Ricketts, 1941; Fauvel, 1943;
Hartman, 1944a. Common in Gulf.
E. aphroditois (Pallas). Steinbeck and Ricketts, 1941; Fauvel, 1943; Hartman, 1944a. Common
in Gulf.
E. australis Quatrefages. Fauvel, 1943. La Paz.
(?)£. coccinea Grube. [sic]. Fauvel, 1943. Gulf.
E. filamentosa Grube. Steinbeck and Ricketts, 1941; Fauvel, 1943; Hartman, 1944a. Gulf.
E. indica Kinberg. Fauvel, 1943. Isla San Jose.
E. longicirrata Webster. Berkeley and Berkeley, 1939; Hartman, 1944a. Common in Gulf.
E. mnltipec tin at a Moore. Fauvel, 1943. Gulf.
E. schemacephala Schmarda. Steinbeck and Ricketts, 1941. Isla Espiritu Santo.
E. tentaculata Quatrefages. Fauvel, 1943. Gulf, [perhaps E. apboditois, fide Hartman, 1959b].
E. vittata (delle Chiaje). Hartman, 1944a; Rioja, 1947a. La Paz to Bahia de los Angeles.
Marphysa aenea (Blanchard). Steinbeck and Ricketts, 1941; Hartman, 1944a. Isla Espiritu Santo,
El Mogote.
M. mortenseni Monroe. Berkeley and Berkeley, 1939. Isla Espiritu Santo.
1968 Reish: Bahia de los Angeles Annelids 103
M. sangumea (Montagu). Hartman, 1944a; Rioja, 1947b. Isla de Jorge. Isla Angel de la Guarda,
La Paz.
M. stylobranchiata Moore. Rioja, 1947a. La Paz. Bahia de Navachiste.
*Nematonereh unicornis (Grube).
Nicidhm cariboea (Grube). Hartman, 1944a. Isla Espiritu Santo.
Palola paloloides (Moore). Rioja, 1947a. La Paz, Topolobampo.
P. siciliensis (Grube). Hartman, 1940; Steinbeck and Ricketts, 1941, Fauvel, 1943. Common in Gulf.
Lumbrineridae
Lumbrineris bifilaris (Ehlers). Hartman, 1944a; Rioja, 1947a. La Paz, Los Frailes.
L. branchiata Fauvel, 1943. [Homonym, fide Hartman, 1959b, p. 333.]
L. brevicirra (Schmarda). Fauvel, 1943. La Paz, Isla San Jose.
L. erecta (Moore). Hartman, 1944a. Between 24° 25' and 29° 54' W. Latitude.
L. inflata Moore. Fauvel, 1943; Hartman, 1944a. Isla Partida.
L. latreilli Audouin and Milne Edwards. Hartman, 1944a. Between 25° 49' and 31° 01' \\". latitude.
L. I. japonica Marenzeller. Hartman, 1944a. Bahia de San Francisquitc.
*L. minima Hartman.
L. simplicis Hartman, 1944a, 195 9. Isla Angel de la Guarda, Bahia San Luis Gonzaga.
L. tetraura (Schmarda). Hartman, 1944a. Isla Tuburon, Isla Espiritu Santo, Bahia Tepoca.
Arabellidae
Arabella tricolor (Montagu). Hartman, 1944a; Rioja, 1947a. Common in Gulf.
A. mutans (Chamberlin) . Fauvel, 1943; Hartman, 1944a. Isla Angel de la Guarda, Bahia Agua Verde.
A. scmimacitlafa (Moore). Hartman, 1944a; Rioja, 1947b. Between 24° and 30° W. Latitude.
Drilonereis filum (Claparede). Fauvel, 1943. Gulf.
D. facata Moore. Hartman, 1944a. Isla Espiritu Santo.
D. nuda Moore. Fauvel, 1943. Gulf.
Lysaretidae
Aglaurides fulgida (Savigny). Steinbeck and Ricketts, 1941; Fauvel, 1943; Hartman, 1944a. Common
in Gulf.
Dovilleidae
Donillea articulata (Hartman). Rioja 1947a. La Paz.
D. cerasina (Ehlers). Steinbeck and Ricketts, 1941; Hartman, 1944a. Between 24° and 29° \V.
Latitude.
''Protodori illea gracilis (Hartman).
Orbiniidae
'■' Haploscoloplos elongatus (Johnson).
Naineris laevigata (Grube). Rioja, 1947b. Topolobampo.
"'Orbinia jobnsoni (Moore).
*Pbylo felix Kinberg.
*P. nudiis (Moore).
''Scoloplos acmeceps Chamberlin.
Paraonidae
* Aricidea rosea, n. sp.
*A. lischakoui Zachs.
K'Paradoneis lyra (Southern)
''Paraonis gracilis oculata Hartman.
Spionidae
Aoni.les califomiensis Rioja, 1947a. La Paz.
*Laonice cirrata (Sars).
*Nerinidis acuta (Treadwell).
*N. macula/a Hartman.
*N. pigment at a (Reish).
Polydora ciliata (Johnston). Ricja, 1943. Guaymas.
P. cirrosa Rioja, 1943. Guaymas.
*P. heterochaeta Rioja.
P. ligni Webster. Rioja, 1943, 1947a. Guaymas, Topolobampo.
P. richettsi Woodwick, 1961. Cabo San Lucus.
P. socialis (Schmarda). Rioja, 1947a. La Paz.
"'P. s. plena Berkeley and Berkeley.
*Prionospio cirrifera Wiren. Bahia de los Angeles.
::'P. longibranchiata, n. sp.
'■'P. malmgreni Claparede.
*P. pinnata Ehlers.
*?• pygmaeus Hartman.
'■' Pseinlopolydora reishi Woodwick.
104 San Diego Society of Natural History Vol. 15
*Spiophanes bombyx (Claparede).
*S. inissionensis Hartman.
Magclonidae
*M. calif or nica Hartman.
Disomidae
*Poecilochaetus johnsoni Hartman.
Chaetopteridae
Cbaetopterus variopedatus (Renier). Fauvel, 1943; Rioja 1947b. La Paz.
"'Telepsai us costaritm Claparede.
Cirratulidae
'■' Acrocirrns crassifilis Moore.
* Coulter iella alata (Southern).
*Chaetozone corona Berkeley and Berkeley.
Cirratulus exuberant Chamberlin. Treadwell, 1937. Gulf.
C. sinincolens Chamberlin. 26° 48' W. Latitude and 110° 45' 20" N. Longitude.
Cirriformia luxuriosa (Moore). Rioja, 1947a. Bahia de Navachiste.
C. spirabrancl.nl (Moore). Steinbeck and Ricketts 1941. Bahia de los Angeles, Coronados.
''Cossura Candida Hartman.
Dodecaceria pacifica (Fewkes). Rioja, 1944. Guaymas.
' Tharyx pari us Berkeley.
*T. tessclata Hartman.
Flabelligeridae
Pberusa capulata (Moore). Steinbeck and Ricketts, 1941. Isla Angel de la Guarda.
P. cruca Caparede. Rioja, 1947b. La Paz.
*P. neopapillata Hartman.
P. papilla/a (Johnson) Steinbeck and Ricketts, 1941; Rioja 1947a. La Paz.
P. plumosa (Muller). Rioja, 1947b. La Paz.
"Piromis gracilis Hartman.
Opheliidae
Ammotrypane aulogaster Rathke. Rioja, 1947a. La Paz.
'' Armandia bioculata Hartman. Bahia de los Angeles.
Polyophthalmus pictus (Dujardin) Rioja, 1947a. La Paz, Topolobampo, Bahia de los Angeles.
Traiisia gigas Hartman. Steinbeck and Ricketts, 1941. Cabo San Lucas, Bahia de los Angeles.
T. olens Ehlers. Fauvel, 1943. Gulf. [Perhaps T. chinensis Grube or T. japonica Fujiwara, fide Hart-
man, 1959b, p. 436].
Sternaspidae
Sternaspis major Chamberlin, 1919a. Between 27° 39' 40" W. Latitude and 100° 0' 30" N. Longitude.
Capitellidae
'■' Anotomastus gordiodes (Moore).
''Capitella ca pit at a (Fabricius).
*Capitita ambiseta Hartman.
Darybrancbus caditcus (Grube). Steinbeck and Ricketts, 1941; Fauvel, 1943; Rioja, 1947b. La Paz,
Isla San Jose, El Mogote.
D. glabrus Moore. Hartman, 1947a. Guaymas.
D. lumbricoides Grube. Hartman, 1947a. Isla Tiburon.
D. platyceps Hartman, 1947a. Estado de Sonora.
Notomas/us lobatus Hartman, 1947a. Consag Rock.
*N. (Clis/oinastus) tenuis Moore.
Arcnicolidae
Arenicola glassclli Berkeley and Berkeley, 1939. San Felipe.
Arenicola sp.
Maldanidae
*Asychh disparidentata (Moore).
* Axiothella rubrocincta (Johnson).
Euclymene papilla/a Berkeley and Berkeley, 1939; Rioja, 1947a. Punta Penasco.
Iletcroclywene glabra Moore.
"Maldane sarsi Malmgren.
'■'Praxillclla affiius pacifica Berkeley.
Oweniidae
Ouenia fusiformis dclle Chiaje. Steinbeck and Ricketts, 1941. EI Mogote.
*
O. /. collaris Hartman.
1968 Reish: Bahia de los Angeles Annelids 105
Sabellaridae
I dan thyrsus pennatus (Peters). Steinbeck and Ricketts, 1941. Cabo Pulmo.
Pectinariidae
Amphictene auricoma (Miiller). Rioja, 1947b. La Paz.
'' Pcrtiiitii iii hartmanae n. sp.
Ampharetidae
Ampharete boma Chamberlin, 1919a. 26° 40' W. Latitude and 110° 45' 20" N. Longitude.
*Ampbicteh scaphobranchiata Moore.
* Amphhamytha bioculata (Moore).
* Asabellides line at a (Berkeley and Berkeley).
Sabellides delus Chamberlin, 1919a. 26° 40' W. Latitude and 110° 45' 20" N. Longitude.
Terebellidae
'' Amaeana occiJeutalis (Hartman).
Artacama conifera Moore. Hartman, 195 5. Los Frailes.
Axionice mirabilis Mcintosh. Fauvel, 1943. Gulf.
Eupolymnia heterobrancbia (Johnson). Rioja, 1947a. La Paz.
Loimia medusa (Savigny). Fauvel, 1943, Gulf.
L. montagui (Grube). Rioja, 1947a. La Paz.
Neoamphitrite robust a (Johnson). Fauvel, 1943; Rioja, 1947a. La Paz.
Neoleprca spiralis (Johnson). Rioja, 1947a. Topolobampo.
Xicolca la/ens. Chamberlin, 1919a. 26° 48' W. Latitude and 110° 45' 20" N. Longitude.
Vista elongata Moore. Steinbeck and Ricketts, 1941. El Mogote.
*P. crista/a (Miiller).
Polycirrus mexicanus (Rioja), 1947a. La Paz.
''P. perplexus Moore.
Spinosphaera oculata Hartman. Rioja, 1947b. La Paz.
Terbella calif 'arnica Moore. Fauvel, 1943. Gulf.
Tbelepus comatus (Grube). Fauvel, 1943. Gulf.
T. crispus Johnson. Treadwell, 1937. Cabo San Lucas.
T. setosus (Quatrefages). Berkeley and Berkeley, 1939; Steinbeck and Ricketts, 1941. San Felipe,
Bahia de los Angeles, Punta San Marcial.
Trichobranchidae
Terbellides stroemi Sars. Berkeley and Berkeley, 193 9. Puerto San Carlos.
Sabellidae
Chone iiifundibuliformis fauieli Mcintosh. Berkeley and Berkeley, 1939. Puerto San Carlos, Bahia de
los Angeles.
C. minuta Hartman. Rioja, 1947a. La Paz.
::'C. mollis (Bush).
*Euchone barnardi, n. sp.
*£. cortczi, n. sp.
''Fabricia limnicola Hartman.
Hypsicomus pbacotacnia (Schmarda). Fauvel, 1943; Rioja, 1947a. La Paz.
Megalomma mushaensis (Gravier). Steinbeck and Ricketts, 1941; Fauvel, 1943; Rioja, 1947a. La Paz,
Isla Espiritu Santo, Coronados.
*M. pigmentum Reish.
Oriopsh armandia (Claparede). Rioja, 1947a. La Paz.
Pseudopotamilla occelata Moore. Rioja, 1947b. La Paz.
Sabella melanostigmo Schmarda. Fauvel, 1943. Gulf.
Serpulidae
Apomatus similis Marion and Bobretzky. Treadwell, 1937. Cabo San Lucas.
Eupomatu\ bumilis Bush, 1904. Guaymas.
Hydroides crucigera Morch. Rioja, 1947a. La Paz. Topolobampi .
SrH. recur i ispina Rioja, 1941. Rio Mayo.
Pomatoceros minutus Rioja, 1941, 1947a. Rio Mayo, Bahia de Navachiste.
Pomatoleios kraussi (Baird). Rioja, 1947a, Bahia de Navachiste, Topolobampo.
Pomatostegus stellatus ( Abildgaard). Fauvel, 1943. Cabo Pulmo.
Protula superb a Moore. Parker, 1963. Off mainland coast and southern limits of Gulf of California.
Protula tubularia (Montagu). Steinbeck and Ricketts, 1941. Puerto Refugio.
Sal mac in a dysteri (Huxley). Steinbeck and Ricketts, 1941; Fauvel, 1943. Bahia de los Angeles,
Bahia de San Francisquito.
Sphobranchus giganteus (Pallas), Fauvel, 1943. La Paz, Cabo Pulmo.
S. pseudoincrassatus Bush, 1904; Steinbeck and Ricketts, 1941; Ricja, 1947a. Cabo San Lucav
Topolobampo.
Spirorbis marioni Caullery and Mesnil. Bush. 1904. Guaymas, La Paz.
Vermiliopsis cornuta Rioja, 1947b. La Paz.
106
San Diego Society of Natural History
Vol. 15
Table 2
The relationship of the principle polychaetes to sediment type in Bahia de los Angeles,
Baja California. Numerals in parentheses indicate the number of stations at which the
species were collected in April 1962 and October 1963, respectively.
Sand,
40 stations
Silty Sand,
46 stations
Sandy Silt,
30 stations
Silt,
45 stations
Species
Per- Mean no. Per- Mean no. Per- Mean no. Per- Mean no.
centage specimens centage specimens centage specimens centage specimens
occur- per occur- per occur- per occur- per
station rence station rence
ence
station
ren
30
2
27
33
2
40
40
3
39
25
2
35
30
3
26
42
4
33
35
14
28
30
3
22
90
12
98
48
30
37
63
11
76
63
9
85
60
9
52
75
18
83
15
17
35
35
3
33
40
5
57
83
1 1
80
20
9
26
73
10
3 0
55
4
56
70
40
72
5
1
39
65
21
85
42
13
26
62
147
72
27
19
22
68
20
67
25
15
22
50
1 1
43
15
27
19
30
2
39
station
Pareulcpis fimbriata (13-21)
Etcone dilatae (3 5-7)
Ophiodromus pugetteiish (29-20)
Ancistrosyllh bassi (4-3 9)
Pilargis bamatus (26-9)
Langerhansia heterochaeta (21-16)
Ceratonereis mirabilh (32-9)
Nereis procera (27-6)
Aglaopbamus dicirrus (66-84)
Giycera tesselata (20-3 5)
Glycinde armigea (57-5 6)
Onuphis zebra (60-76)
Lnmbrineris minima (57-20)
Haploscoloplos elongatus (5 8-54)
Aricidea rosea
Polydora socialis plena (27-22)
Prionospio cirrifera (46-62)
P. pinnata (36-60)
P. pygmaeus (5 1-4)
Magelona californica (14-30)
Telepsaius costarum (43-23)
Cbaet ozone corona (49-32)
Cossura Candida (40-42)
Tharyx pari us (5 8-47)
Armandia bioculata (31-15)
Capitita ambiseta (67-54)
Heteroclymene glabra (37-1)
Praxillella nffinis pacifica (54-32)
Amaeana occidentalis (33-11)
Chone mollis (54-14)
Euchone barnardi (3 5-3)
Megalomma pigmentum (22-13)
2
17
2
20
3
27
2
20
2
23
2
13
2
23
3
20
9
93
5
20
4
80
6
97
5
50
22
83
76
63
1
37
12
83
16
5 3
18
60
24
3
3
4
11
33
2
67
28
67
3
27
255
83
9
27
1 1
47
6
53
9
43
10
40
3
10
2
—
3
9
2
17
2
17
1
9
3
2
1
17
3
11
10
91
1
29
4
65
13
96
3
29
11
42
87
53
4
20
12
91
7
22
18
1
5
3 8
18
6
22
6
94
9
44
7
40
182
84
6
20
14
31
4
18
9
3 3
15
24
1
2
6
1
4
10
2
4
100
1
12
1
16
1
3
12
2
2
52
5
2
1
3
23
1
MUS. COMP. ZoOL.
LIBRARY
UU 181968
HARVARD
UNlVfiftSJT*
A BIOLOGICAL SURVEY OF BAHIA DE LOS
ANGELES, GULF OF CALIFORNIA, MEXICO. III.
BENTHIC MOLLUSCA
EUGENE V. COAN
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 8, 25 SEPTEMBER 1968
.s^*>-
ssCi
rV Pata-'Bota
Isla n
Sorobado1'
(Olsla Flecha
Cerraia^l /lsla
' Ventanaj
Bahia de Los Angeles
MEXICO
206 207
Isla
Cabeza de
Caballo
Islos de
Las Gemelos
VSFS
HOUSES
104 107
•ho
NAUTICAL MILES
KILOMETERS
J 1_
113° 30
Figure 1. Map of Bahia de los Angeles. Inset, location of bay in Gulf of California.
A BIOLOGICAL SURVEY OF BAHIA DE LOS ANGEL
GULF OF CALIFORNIA, MEXICO. III. ,,k**vA*0
BENTHIC MOLLUSCA
Eugene V. Coan
Abstract. Benthic mollusks were collected by the Beaudette Foundation at Bahia de los Angeles,
Baja California, Mexico, in April 1962, and October 1963. Samples were obtained with an orange-
peel grab at 161 stations. April collections were sparse, but October samples were even sparser. The
depletion may result from seasonal predation by bottom-dwelling fishes. Fewer species and specimens
were found with increasing water depth, and with the exception of a few shallow water localities
near the head of the bay, the molluscan fauna is considered to represent a single assemblage charac-
terized by the genera Tellina, CaJulus and Nucitlana. This assemblage is thought to be typical of
silty-sand substrates in semi-protected bays in tropical and subtropical areas. The fauna of Bahia de
los Angeles is characteristic of the Panamic Province and is most similar to faunas found farther
south in the Gulf of California.
Resumen. En la Bahia de los Angeles, Baja California, Mexico, se recolectaron moluscos benticos
durante los meses de Abril 1962 y Octubre 1963. Estas colecciones fueron patrocinadas por la
Fundacion Beaudette. Las muestras corresponden a 161 estaciones, y fueron recogidas con unas tena/.is
de cuatro cucharas o sectores. Las colecciones correspondientes al mes de Abril fueron pobres, y las
de Octubre resultaron aiin mas deficientes. Esta escasez de moluscos se debe probablemente a la
predacion que efectuan los peces de fondo durante ciertas estaciones. A mayor profundidad el numero
de especies y de ejemplares por especies era menor. La fauna malacologica, excepto en las zonas mas
internas de la bahia, estaba representada por los generos Tellina, Catlitlus y Nut itlana. Este complejo
malacologico se considera tipico de los substratos de arenas salobres en bahias semi-protegidas de las
regiones tropicales y subtropicales. La fauna de la Bahia de los Angeles es caracteristica de la pro-
vincia Panamena, y es muy similar a la fauna que se presenta mas al sur en el Golfo de California.
This report deals with the benthic mollusks collected by the Beaudette Foundation
of California at Bahia de los Angeles, Gulf of California, Mexico, in April 1962 and
October 1963. A preliminary report on these collections has been presented elsewhere
(Coan, 1964). A list of mollusks obtained from shore and beach-drift collections has
been published by McLean (1961).
Methods
A detailed description of sampling methods is given by Barnard and Grady (1968).
Briefly, a grid of 161 sampling stations was established in the southern portion of the
bay (Fig. 1; see also Reish, 1968, Fig. 1). Samples were collected with an orange-peel
grab that covered approximately 0.06m- of bottom surface. In general, alternate stations
were sampled in April 1962, the remaining stations in October 1963. However, several
stations were sampled in both periods, and stations 1 through 8 were sampled onlj in
October. With the following two exceptions, one sample was made at each station: two
samples, preserved separately, were made at Station 1)8 in April; two samples, combined,
were made at Station 130 in April. These exceptions were taken into consideration in
the computations.
Several non-quantitative samples were also made by the Beaudette Foundation and
by an expedition of the Scripps Institution of Oceanography, both in April 1962. The
material is not included in the analysis, but the species are reported in the Systematic
List and in the discussion of geographic ranges. Detailed information on sampling
cedures at these stations is presented in Barnard and Grady (1968, Tables 2, 3
San Diego Soc. Nat. Hist., Trans. 15 (8): 107-132, 25
110
San Diego Society of Natural History
Vol. 15
6 A H I A DE LOS ANGELES
NAUTICAL MILE
500 1000 1500
Sediment Types
METERS
Figure 2. Sediment distribution in south portion of Bahia de los Angeles.
The material collected was sorted to phylum. Species represented only by shells were
removed from the molluscan samples to minimize possible contamination by organisms
that may have drifted from elsewhere. With small specimens this separation was difficult,
and some dead organisms may have been inadvertently included in the analyses. After
the material was air-dried, the distinction between live-collected and dead shells could
not be made as easily.
The physical features of the bay are discussed by Barnard and Grady (1968).
Figure 2 shows the major benthic sediment patterns for comparison with the distri-
butions of the species discussed.
Discussion. — Several potential sources of sampling error are not excluded by the
techniques employed. Use of an orange-peel grab virtually precluded the collection of
some deep-burrowing mollusks, and the depth of grab penetration varies with sediment
properties. Because of the large number of samples taken, some deep-burrowing species
may be represented. Future studies should utilize large grabs to supplement smaller ones.
1968 Coan: Bahia de los Angeles Moi.lusks
I I I
While the rigid grid pattern results in uniform sampling areally, it does not neces-
sarily provide sufficient data for charting precise distributional patterns. For example,
high gradient slopes were not sampled effectively because of their small areal representa-
tion. The distributional patterns of some mollusks reflect this because their depth
habitats were not sampled on the more steeply-sloping eastern side of the bay. Species
that may be especially abundant on such slopes — their distributions being more stronglj
correlated with degree of slope than with depth or sediment type — may not have been
collected.
There is a lack of quantitative samples from depths shallower than 10m. A few
shallow-water grabs were made from a skiff, but there are obvious difficulties in taking
samples with a relatively large device from a small boat. Parker (1964a, b) has men-
tioned decided differences between intertidal sandy beach assemblages and off-shore
shallow-water assemblages in the Gulf of California. At Bahia de los Angeles there
appears to be a distinct shallow-water fauna at some stations, especially numbers I
through 8, but the samples are too few to be conclusive.
Discussion
Differences between sampling periods. — Fewer mollusks than expected were col-
lected in the April survey. From 76 samples, 4060 individuals representing 126 species
were collected; in general, the specimens appeared to be juveniles. In October, only 97
species and 18 32 individuals were collected from 91 samples; a wider range of specimen
sizes seemed to occur in this period, but no quantitative measure of this was made.
Thus, there was striking depletion of the bay fauna. Several shallow-water species were
collected in October but not in April; two of these occurred in relatively large numbers.
The causes of this depletion are not known. It might result from annual or seasonal
changes in water temperature, sediment deposition patterns, behavior of the organisms,
action of bottom-feeding fishes, unsuspected changes in sampling techniques and grab
operation, or even from random variation. Too little data are available to resolve this
question. However, there are features of the depletion that may be pertinent. I noted
that the remains of dead animals were not found in greater abundance in the October
sample. If the depletion resulted from /';/ situ death, one would expect — barring other
complications such as burying, sediment shift, and decomposition — to find more residual
shells in October.
The change in the distributional pattern of species may also be instructive
(Table 1). Twenty-seven species were collected in quantities of thirty or more in-
dividuals in the quantitative sampling. Eighteen showed a decrease in numbers in the
October sample, and in ten of these the decrease seemed to be greater toward the mouth
of the bay, though not necessarily in deep water. Possibly predation is greater otl shore
than near shore, where tidal variation, turbulence, and temperature extremes may deter
predaceous fishes. The tendency for forms to decrease offshore, however, cannot account
for the virtual disappearance of some fairly common species (e.g., Nuculana acrita).
The problem is worthy of further research.
Bathymetric distribution. — The April data indicate an inverse relationship between
the numbers of species and specimens per sample and water depth. (The October data
were not analyzed in this manner.) A rather sharp break occurred at approximately 3 0m.
About 17 species and 90 specimens per station were taken from depths of less than 30m,
nine species and 30 specimens at greater depths. Keen (1964) reported that the number
of specimens at 40m at Isla Espiritu Santo, Gulf of California, was greater than at 20m,
but she included both living and non-living material.
112 San Diego Society of Natural History Vol. 15
Table 1
Changes in abundance of Mollusca between sampling periods
Species showing an apparent increase in numbers
a) due to near-shore sampling in October: Transennella humilis; Caecum sp.;
Micranellum sp. ; Elephantulum cf. E. liratocinctum; Nassariiis tiarula; Actco-
cina angustior.
b) due to greater number of unidentifiable juveniles: Tcllina sp.
Species showing little changes in numbers and
a) little change in distribution: Transennella sororcula.
b) decrease off shore (increase due to near-shore sampling) : Tcllina arenica.
Species showing little change in numbers and
a) almost complete disappearance: Nnculana acrita; Mysclla cf. M. chalcedonica;
Dentalium innumcrabilc (a deep-water form) ; Solariorbis liriope (a deep-
water form ) .
b) a uniform decrease throughout the bay: Nitcnla dcclivis; Crenclla ecuadoriana;
Mangclia cyrene; Actcocina infrequens.
c) a greater decrease off shore: Nncinclla subdola; Lucina mazatlanica; Diplodonta
sericata; Laevicardium elenense; Chione gnidia; Lyonsia gouldii; Solemya vol-
vulus; Cad nl us panamcnsis; Alabina cxcurvata; Actcocina carinata.
The Bay-wide Community. — A computer study, made under the direction of Dr.
James Rohlf, indicated that the fauna of the bay was not divisible into distinct associa-
tions or communities. Thus, with the exception of the small shallow-water fauna taken
only in October 1963, which seems to be distinct, the mollusks obtained in this study
are considered to represent a single community. Parker (1964a, b) concluded that for
the Gulf of California in general there is a sharp faunal division at 26m. No such
division was found in the present study, although the number of species declines at
depths of over 30m. There is, however, a gradational decrease in mollusk abundance
toward the mouth of the bay.
Table 2 lists the commonest mollusks collected during the survey. The three most
abundant forms, Tcllina arenica, Cadulus panamensis, and Nuculana acrita are found
throughout the bay (see Systematic List). Few of the species which Parker (1964a, b)
lists as characteristic of ll-26m or 26-65m depths were found in this study, but his
samples were from a wide area of the Gulf of California. Also, a greater variety of
depths and environments were sampled, and the sampling regions did not include bays
as protected as Bahia de los Angeles.
The Tcllina-Cadulns-N nculana assemblage does not correspond with assemblages
described by Thorson (1957) for arctic and sub-arctic regions. Parker (1960) noted
associations in the Gulf of Mexico which are somewhat similar to that at Bahia de los
Angeles. The fauna at Isla Espiritu Santo (Keen, 1964) is quite different, perhaps because
the substrate in the area is coarse coral sand. I suspect that the Tellina-Cadulus-Nuculaua
assemblage may be typical of sandy-silt substrates in semi-protected bays. Mr. Barry
Roth and I found that these three genera were the most abundant in the harbor at
Princeton, San Mateo County, California, in August 1962 and July 1964, though the
species represented all differed from those at Bahia de los Angeles.
1968 Coan: Bahia de los Angeles Mollusks 1 13
Table 2
Relative abundance of common Mollusca in Bahia de los Angeles, April and
October samples combined. A density estimate (no. specimens / m2) is given for the
commonest species.
Tellina arenica, 998 (95.5); Cadulus panamensis, 679 (65.0); Nuculana acrita,
660 (63.2); Acteocina carinata, 432 (40.5); Nassarius tiarula, 276 (26.4); Diplodonta
sericata, 215 (20.6); Solemya volvulus, 178; Lyons/a gouldii, 163; Nucinella subdola,
152; Caecum sp., 151; Mysella cf. M. chalcedonica, 150; Alabina excurvata, 143;
Nucula declivis, 117; Micranellum sp., 115; Crenella ecuadoriana, 107; Dentalium in-
numerabile, 72; Transennella humilis, 70; Elephantulum cf. E. liratocinctum, 69;
Tellina sp. (unidentifiable juvs.), 62; Laevicardium eleneme, 59; Acteocina angustior,
59; Transennella sororcula, 51; Acteocina infrequent, 5 0; Solariorbis liriope, 48; Man-
gelia cyrene, 47; Chione gnidia, 43; Lucina mazatlanica, 39.
Biogeograpbic considerations. — Of the 177 identifiable species encountered in this
study, 154 do not range into California but may extend northward on the west coast
of Baja California. Very few, however, occur north of Bahia Magdalena. Of 2 3 forms
which do range to California, 19 also extend farther south than the tip of Baja Cali-
fornia. Of 47 species encountered that occur only in the Gulf of California, 2 8 do not
occur north of Bahia de los Angeles. This is surprising because the salinity and water
temperatures at the head of the Gulf are similar to those of Bahia de los Angeles.
Systematic List
A systematic list of species collected in both the quantitative and non-quantitative
sampling is given below. The material collected is analyzed by time of collection, and
the depth range and main distribution within the area of quantitative sampling (or out-
side this area) is given for each species. The following abbreviations are used: C = central;
E = east; f.= figure(s) ; int. = intertidal; N =north; pi. = plate(s) ; s = scattered; S = south;
sub = subtidal; W = west; ::' = probably new species.
The material collected is housed in the Department of Geology, Stanford University.
California, under the collection numbers 50306-50489. For a complete Station list of
specimens order NAPS Document 008 3 from ASIS National Auxiliary Publications
Service, c/o CCM Information Sciences, Inc., 22 West 34th Street, New York, N.Y.
10001; remitting $1 for each microfiche copy and $3 for each hard-copy reproduction.
114
San Diego Society of Natural History
Vol. IS
co
U <
t 3
< <
H
CO
to
4> rt ^ u
» S £ c
C U, 4_, rt
rt rt rj -t-J
-•^ DO 3
o\ .a «
c
er 1
spec
en a
<N
-S • -*
O o «
d
c
NO
0
O)
E
a
C
rr
Ui
O
d»
o
c
rt
<N
(A
£
no
*«— »'
on
"-1
—
IH
a
^■v
<
(A
c
u
rt
s
0>
G
&
rt
</>
to
o
z
d
u
c
01
c
>"
U, C/5
z
u
z
z
o
I *
o
no
o
NO O
On
CM
O
NO
NO
o
NO
O
NO
ON
K _]
u*
<+*
^ „ «
,
„
■""■
—'
<vj >*
O
;
. <*\ , n
r a
1-1
c-
a"" —
H • *.
H-J
...
• ~H-i Ph
CO
ON '""' ON O
. c
O £ f-H 0»
V- ON
On """'
C
>- 1 O
NO NO «v" 2
^ ^ On 5
" — ^ ^ "5
^ NO NO ^ 2
^ on on £ ■>
ON ^ rt ON '^
c °
O to
uJ O ui O UJ
0 ylOO«
t^N Q.
^ CM
CM ••
u — .
^ o
K
00 rj- ON
"
NO
X °
oo ^>
On
£- ^
^ ON^-
^^
On "
ON ~
" c *
C
C
c
o
c °
a°~
a» vi
o so a.
^
0
^ 0
^ 0
UJ -i-
S "S "
S X a,
; — ' u-
<, -— OO
Q ^ ^
Z ^
o
f^
■*
ON
ON
•— '
f— «
00
.s
"3
«
c
o
c
u
u
■--
r^
w
-;
— >
u.
t^»
s
r+\
"^ f^
t3 CO
^ o
3 <s>
■5
<
Q
Z
<
D
O
D
CO
Ui
o
C/N
CO
o
ON
Q
<
Q
w
Z
u
■ft
«s
a
0»
K
id
00
c
o
<
Q
u
oi
— -
c
o
u.
oo
>^
Ui
o
c/5
<3
CO
Ui
o
C/3
nailar a multico
(Sowerby, 183
'<
l
Z
1
*
Z
£
<
^
^
^
1968 Coan: Bahia de los Angeles Mollusks 1 1 s
c
>
U
3
to ^ vs
o to S>ocn c/> c/j £> Z
— C\
so
o
o
CM m NO
s — — — « rj fN CM
n^-' — - — — * —
ON — CM — ' <n -i- <N
-t '
.* I <M .
^ 1- \0
«>* ,-r <*"
o „ —
. ~" . <^4 . rl
K NO ^ • v-
o — cm — ' m r\ ""> — i
^ 0, , — C_ — ~ — C
. ■- ** ..,.•"...
*-M O ^ \o «+h <NOv+-n3v+H^N»HONr\'+H M-. \D "^ v+-4OOv*-"r^N*Ho0
O-^ ^O _ fN •»■> ^ „ **"> „ Ol CM „ 2 . t ~ _ «- .. ^f . M5
cm — • o" ~ "~ o — ri ~ ~r ~ ~~ -1- ^ ■*.--.. o—i^i i-i— vo
"- .. 1" K .. K .. K ..
~5*i «5«;^m^ «
c* ** a " e c ~~ "** — c " c" c* - c o c "" c b
ci S a. u 8 oSKoVJoSoSSu^ ci k j u w «> <« j> « u
c» — ci — ' o — — ci ci — Cl - — — Ci ci — "T ci — o -— • ci — -
„
„
>^
-
„
in
«-^.
K
V)
c
NO
Tf"
a
**~*
—'
rt
&c
,.
"a
~E-
OO
1£
o
©
ON
"EL
ON
"a,
*-»-.*
<N
o
Mh
-o
VH
K
Mh
On
K
M-I
~
rs|
«•<-»
o
CM
-
<<t
CM
rsi
-1-'
5
\r*
*^
**
^
\^
o-
^
oo
\o
NO
oo
■ Z
vc
^O
OO
NO
ND
oo
^
^N
ON
OS
i;
On
ON
ON
«^v
ft)
ON
1—1
rH
0\
^
ON
ON
*™—
•"" '
ON
s
c\
■5 ^
5 0>
a*3»; qi-12 '5:^-^ sl Sis ;::;. i=g
J3 -
O'OO ^ioU -'U-- --- -~ - -1-.-
116
San Diego Society of Natural History
Vol. 15
c
o
.si
EUH
60
0J
</> S
u
rt
a
rt -73
**-'
w C
mC
_-
tuO ^
c
-T>
Q
OS .5J «
ug,fi
-a . -^
O o «
J3
3 .2
^
-o
w
2
(/l
60
-a
0>
«
o
CO
u
K
K
so
so
Os
a.
<
.a «
o n
c
o
so
■> — •
ON
K
—
K
JQ
so
so .
2"a
r-l a. a- ~
OS . 1-H
" "5. .
OS
as"
O . (N
as — os — !
r-( a rn ft
OS
o
OS
^- rsi ->
OS .. ..
« ^ m
" OS os
OS " "
Os
_ r-s
so
os a
so
o
-J <N
ft
uj 0
i4 0
so OS os
OS ■• ■*>
-£"" .
"* ^ _a ~ u
OS ^ os ■
c ° c
o — i tu — . <u — « rt cu ■ — >o<u— ^ ^ W ^ ^
\4 O ^O^OftUJO^^O^O^O *
-1 SO <N
OS .. OS ..
<» ~ " ^
^ "^ ^ OS
OS ■""' os — '
c a"
r-* O -" O
» ^ oo ^
OS ^ os ■"
c
a °
*! 0 ^ O
C
^ ft
3
a
w
H
<
CO
to
<
U
E
■n.
<! 8 ^
E s *
5-2 2
< ~ ~
u u
s *•
O c ^
<-* ?
^ -s ^
Z 1 «
0 5J
U U
a
z
u
p
so
a
i; £*
^: 0
2 &
a U
►3
-3
c
•2 C
— J2
<s —
■5 £
^3
«
Q
^
K^.
^
^;
oo
X
0
o
OS
*<*
M
5S
C
a>
2
cd
"3
.5
P
2 | ~S 2
T3 <U
u
u S
1968 Coan: Bahia de los Angeles Mollusks 117
Z I* Z ^ Z £* * * Y
O ' ^ fM SO "^ T\ CSI ' <*1 I — „, ("4
l -~^ — II ^^ **"* I I N-^ ' r- I V* "^ K
<M r^ONK— ' l\ "s "1" .^-l- — ~ —
IN CN SO ^t
■-< rx "* — so
— oo "s — on
- ^= -° A ei
M_l T^ ^ • 55
K . 2 . ..^ - - ... ••*
-r
<N
a
>*-!
**S
*
•>
O
*N
o
<»
£j
SO
as
as
■"•
C
Q
c
o
VI
V)
„■ « w "' i^IN SO K "> SO _
oo "3 ^OO- OO oo O _; O — ^
<N?°"^ (N *—- <N <N xs Q, «s D. .
.'.£•—'. 3 • . • ■ "a
0.%*h -3 *-w <** s»^ •-*-»_, '
<S
-f .IT NO OS =
o .. - -r-x-^lT
K tC K rj oo' *- o' £ oo' ™ a oV oo' Q On cs -* £ so ^ «
O Os O ^ — . r^ (N <N cs| ON <N ^ rj r.t *s to ,
M» K * M * « * OO OS „, oo oo ~ OO OO OO ^ OS OS **<
.^ NO «-s °^ i^ "^ i^ " >~ « « ■ SO -^ ~- »— >^ "~ _ ^ —
OS ON OS ~ ON "" OS ~ OS u ON ON R OSri
~~~c" ~ £ ~~ G "" c -£>" 2. ~ "~ "-• , „ „ C , g ^
c c c S c S e 8 e 8 i - e = , • c c c" 8 c 8 --'
l5 O s4t2^0 iiO s>o *0 *: us us u; us O * O
W_ W 3 -£ C Sf- £ a _ g . „ n S> . -2 ~
-2 aS >h ?
"7
B
q
"3 0>
>
C
u.
sU
\J
*<^
3
oo
"3
1-5
<-
Z
to
-
o
oo
g *
s
*s»* , X
a os
<
-i
^
a
>.
<
ei. g
c
O "-1
Q
1
"C
»
J3
Q
a >»
^3
q
■3 S
5
<
s
no
V.
v
^
5
s: g
<u 3
i-i ^ ,
u
X
t-l
•g
O
Z
% 'gZ
S
s
§
u
£
hj
>
:>-
S-§l 51 t^7?^^^? Sisal a H S^ -=^^1-!
c ?
^3~U^O -« ?. aa ■> to ^T-a— B VJ >-3~v_--
a<
118
San Diego Society of Natural History
Vol. 15
.S 3
^
Z
u
w
c
r!
Vt3
O
HI
60
-a
o
2
2 <y?
~ o a>
t» — *-» u
ffS 3 g
f rt rt "O
«
o
so
tx so
K
K ■
I c*s
SO i-l
so
as
2 3
o
& =
so
— ■♦ -H
3 .1
g E
<N — —
<
a c o
OS
.-h SO
so
-r
~H °°
Csl '-'
o
C
o .
so
OS
fad 0
v * i « *
■*» T, _: """ — • '
Mh' •- "-" .-
OS IT „ O
^- as « „. o
as
f
" so
SO
OS
"a,_.
OS ^
o
C
c 2 —
CO ^° CO so SO ^° OD _Q
^ OS 1^ OS SO ^ so
as"-1 as ~-< os ^asn
C " 3 _i
-~ o _- o A
w o
CLOwn U m O </) ty
^O ^Obi 0 fad
fad
^0
\o
ON
»*»\
On
J\
^H
ON
,™"
„
i-H
„
C
a
c
O
c
3
u
VI
0)
Ea
fad O fad 0 fad
0
O
"^ CO
l\
r\
CO er\
1—1
00
00
co so
^- OS
ON
CN
OS i-l
r_l
*-*
3
c
C
c °
O CA
fad
fad
fad 0
a
-
t/3
•- Q
U
•5 2
.3 2 &
sC
•3 T
■~ CO
Q.
I = I 5
U
SO
-1-
^ ^
>
3
00
ON
co
U
00
O co
u
t^
^
£
^
c
60
IS
u
0
13
w
rt
s
«
O
<a
,3,
'l-l
-0
<
Q
z
~L
c
^ as
to
^j ON
^* — .
0
•^
-*— 1—
«
2 — "
^s
09
•3 XI
«a — j r
1-1
s^ rt
3
C
03
.5
TD
» y.
j3
3
.S rt
= Q
g
u
"3 S
= Q
■_
^
5u
U
h
H
t~,
K
^
E-<
hi
OS
00
■*
,— '
OS
„
^^
60
u
C
O
0
*»«»
r
«
■r
Sw
0
O
,SJ
<a
-S
id
^:
o\
tj
c
u<
«
'"*
"a
" ■->
«
0
—
>a
7
«
*-»
0
'J
S
rt
••i
■ i;
pq
Q
C^
X
-C:
■-(
E-H
E-H
1968 Coan: Bahia de los Angeles Moi.lusks 119
3
U » C/5 <*> £ ^ C« W ^ ?
K
NO <N| NO
.IN . ** . .5
^> no ^^ I" K ^ oj
"*
_ a. • .,
u — -j
- so
ON
o
- 00
O \0
.— .
CM
fM
—
•• rt
« s
CO
oo \o
s~ o>
1^-
^ so
ON. ""
cn
ON ON
—
fH
— —
c
c S
c
c C
<U v>
o
--J ■->
QJ ■ 1
o
o --»
* o
^
*s M
"*■ £ <*
ON "^ ON
— . .. en i-H -. *N «N ^ t>< •• ■** (^
CN r~l ON
c g ts c"5 o e e e c S "2 c
N
"T3
a<
<+H
.-«
so'
CM
■-H 5
O =:
ON
C a.
r
o
NO
*
NC
■-*-^
^
o
^
rt
s
f*1!
»
o
ON
OC
X
ON
J
i^
~L
^
a
-t-I
.r.
•^_;
j.
OS
^
V-
i" 1
trf-^
NO
-r'
^
-4-
1-
f 1
.j,"
NO
\D
x
| Kl
^
ON
ON
U
ON
*""
»
c
c
~^;
0
-
1/1
U
^o ^ ^^ ^o ^^
ass w ^»> Ss" o?» Q>" 9 1- <-^r:Q=-'^-^--'-
<^y- SS^SQ ^sj *->f <^- ^^-~ r;S= as ^ y ^- =35^
120
San Diego Society of Natural History
Vol. 15
c
o
<
Q
O
Ph
o
X
<
o
bo
c
rt
4-» (J
^ 5
rt -rj
N-"'
f" c
_c
"H
nr 3
c
-Q
Q
^0
tr>
o\
—
U
rt
ft
(A
u
U
O
rt
■j
£
w
S Ji 3.2
-c c a- i-
u i; rt
(^1 trt c c
so N~'
as
a.
<
Vi
io
U
rt
c
eu
C
D
rt
-*
CO
r>
rt
Z
*"*
U
c
-3
C
2 <J
2<£
u
u
u
z
z
so
so
I I
^* OS
•— " OO
<
o
a
vo
0
~~"
Ph
K
0
■*
B<
so
H
</)
<
o
^
oo oo °°
.. _Q ■•
OO SO °°
»*s SO *"^
o o i*
O
«
«
5- K
^ 0s
<
Q
i— i
1—4
►J
<
H
Z
w
Q
S so
J- CM
</) ~ — i
c
-a
<a
w o
- so
5 oo
CM
oo
OS _
s oo a >,
£ — =: -Q
r: •*. "-■
a <*>
5 -C
o so
Sj 1^
5 OS
— a
id
? 3
c
3
■c: _o -s -o
*3 </l <3 O
:0 ^ J2
Q Q
■fc. o
8 </->
~ c
5 O
w
"t: o
? f5
w
<
Is
o
OS
oo
— 1
kJ
<
a.
H
Z
tu
i-i
w
^
(/5
-
K
«
u
■^
V.
c
■5
_D
K
O.
|
£
a
o
Ui
W
D
to
oo
3
■ '
en"
U
E
• ^2
w
Q- rt
** <
<
Q
S c
<s
u
0
J §
o -— -
ai
*~5 ^^^
Q
H
u
1968
Coan: Bahia de los Angeles Mollusks
121
UJ
Z
Z
Z Z
60
-a
o
Z
u
z
3
3
3
no
O
nc
no
(Nl
OO
ON
X Ui
NO
U
no
C
a
» • 2 ■"-
o
o\
to rx M i\ no
• S "S. J" Z JB C
~ — £3 2 c
^ on u_E " b
I 2: Ch m H
t
8^g W
* £ 5 ON <
2 on 2 - Q
o
5 - Z S
3
D
^ —
^ _
_ o
5 o
t^ I
1—1
w
u
<
&
Q
2 o
5 -J
j
j
-a
HE
-2 &
w
•5 >.
Z
<
*o u-
•3 "O
_Q
C/5
<
2 3
i£
£
■g v>
•^
Ch
h
w
On
<
•""'
oj
Q
3
C.
D.
03
60
3
O
•3
00
<
—
"^ <^J
—
</5
,^
00
w
^ O
w
Eulima or B<//
(unidentifie
'8
£
'J
0
u
<
z
w
r
i: c
<
LLl
a
3
■~
-J
DC
u
DQ
C
u
a
u
n
u
=3 <
oa ^ -e
On
q —
■c
, rd
- m
3 -2 •*
is =; <
—
<
Q
g
-
60
c
0
5 C
e u
5 =
122
San Diego Society of Natural History
Vol. 15
° « £ u
60 3 £ c
C i; <-> rt
n ™ « -n
-c -a oo 3
a c "S
o\
.Si w
<j n
B
0
°- F,
1-1
rt
i" 1
s-
. -a
(A
o
O rt
'J
Z "
O
C
c *- _
o 5 rt
Q. O p
a.
.Si w
Z
u
C
c
J
- CO
CO
CO
U
w
co
w
co
>-i no
K —■
<N —•
<N .—
_ — -f
00
c
o
u
t-t __
.5 2
s
<N
Tt"
^s
11
<N
.«
ON
i— 1
^-<
m
(N
c
o
«
ON
0
*~ '
a
J3
>>
u
u
J3
t/i
pq
eu
•4- _
_2 D.
00 „
c —
o —
u
.5 <*
X
«
~ ON
i— • " e
0
H
H
s 72 R
u! C £
s 3 P
<
Q
Z
O
H
■-, 00
3 §
~Q co
c
? ■* g «
^s 00
SS -o
5 '2
■~ 3
c
u
^ >
Z -S
o
w
Z
o
3
N '
IN
00 . .
n -—00
o Ot \o
CO —■ VH
NO
«"■+; -a "*■
c
oo
£*
" ^H " '~^
«
pq
ON 00
^h ND
ON
eq ^
r^ C
ON C ■
^- rt CM
£ 53 on
ffl pq
o
ON
00
c
o
^ ^^
K
c
o
5
3
o
u
Solitriorbis liriop
(Bartsch, 191
Tcinostoma sp.
(unidentifiabl
u
c
8
0
s
CD
c
'53
<
Q
O
CO
CO
2
~ ^J
NO ^
O ^ — .
K a S •?
~i c
C ts o
pq
U
JS u
o _c
k pq
a
t«
Cm
tn
«
^— v
<3
w
S:
•^
<
Q
-a
N
Z
^
«
0
■•^
r
o
o
c/3
cq cq
oi « «
1968
Coan: Bahia de los Angeles Mollusks
123
c
>
o
o
* -
o g
w
co
P
z
o
<u
6C
60
w
"O
-o
c
o
Ml
z
or)
3
CM
o
NO
— . r^i
S3
c;
, J
B,
K
a
a
K
...
• '
...
<N
o
no
n
NO
NO
— '
no
~^
— '
oo
o
■*■
o
o
-r
(N
rj
(N
0\
oo
ON
ON
~*
no
— '
^H
£3 ^ S3
S3 ^
OS
ON
rs"
1-
o\
no
o\
Em;
-^ MH OO
OO
oo
<5j o" ON
NO
ON
C -
ron
pi.
'en,
c
c
3
It
to
co &£
^
H
m
•-*
*■"-.
5
■C
Cv
so
SO
£
c
6.
"_ NO
ni
so
q
OO
V-
s ■=
rt
c*\
l^~
j
m
<a ^
>^\
OO
^3
*3
'— *»
oo
oc
OO
•— '
zr.
•"
«M
"^r "*^
5; ^^
w
o
rt
.— 4
_ o
jo
^^
. . — .
ki
15 ' — ■
^ - —
u
-U
—
M
—
**i v>-
5S
rt
■3 £
■*< *— <
O ri
<
Q
w
O
co
co
isoella sp.
(unidcntifiab
rt
15
_^
o
u
4-1
c
a
n
u
Bartsch, 1921
■ —
Cs
o
c
o
a.
u
rt
u
<
Q
0
w
<
u
-a
CE
*->
c
—
'2
3
•cranellum sp
(unidentified
'J
53
r
a
B
z
u
co
^ ~3
~ B
5 "3
i'i
"•— ■_<
8 3
« o
R u
-*—
Q
-->
c
u
a
L_
H
U
u
c
rt
<
S3
U
—
<
j
D
Q
0
5
rt
-o
<
<
<
E
h
—
^ -
^
•^
s
^
-^
•^
-s
u
3
§
«
S3
S3
S
s
_
U
124
San Diego Society of Natural History
Vol. 15
a
o
rt _Q
v « w c>
m 3 s c
n u s «
rt rt rt -n
■s-2 «i
C4
ON
—. u « .a
rt
a . _*
o o «
-c E
3 .2
5 «
§ E
a.
• v
o
rt
0
C
w
a.
rt
-*
«j
o
rt
Z
tJ
U
q
^3
C
2 u
< a.
- <s
>
o
o
c
>
o
c
>
It:
o
w
Z
o
>
o
c
>
Vt3
o
>
o
p*
Z
ON —
3
no
on —
^
__;
„
rsJ
_
OS
60
a
60
a. b
60
c
o
■"
O
■•
o
Cu
Lrf
rsi
Ui
no
Ih
MH
C/5
rn
co
i*n
„
CO
(N
3
rt •*"*
U
u
rt"
e*1
£
■"-'
c .
•— <
C
i^s
C M-T
CN
c >+*
ON
oo
C
*-m o\
n
^^
rt
^H
^
rt
r— 1
I as
J3
I co"
_"
CT\
I
<u — i
u
QJ 1
J
C
j* ft
i~
_* D.
l_
a>
-al
* 3
rt
rt
rt
rt
o
rt
pq
ca
PQ
PQ
^
cq
H
o
11-
PQ
pq
pa
PQ
PQ
M
60
C
o
<
Q
oo
cu
O
H
ai
w
0
to
dig
■ 2 «
5 c
•2 "°
-5 "5
.■a 3
u
■5; x
o ^
> rt
2 PQ
G
U ON
ON
60
C
■3 o
a a
C to
c
33
rt
■B,X
C PQ
0
5 - ^ «
"0 a
0> g
e u ■
PQ
u
. tn
" c
O On
vh OS
2 C
*pq
2 uT
-5 ■*
■■5-« -^pq
5 NO
S ON
■v. On
PQ «a
u
F-,
f-
K
<
Q
w
<
H
<
u
Q Q.
"3 -a
-S.PQ
u
1968
Coan: Bahia de los Angeles Mollusks
125
w
Z
z
tu
z
to
W
V5
u
C/5
w
Z
-
z
2
o
ON _Q
<N
•*
K
-r o
o —
•*
vo
a
m « —
Q » c ,J
BUI
2i a,
oo
oo
VJ
ON
t g
— * ■*> .s c
Q. s G £
S ON
■2 oo
3
4>
"5 K W
W -£ ■* <
< 5 2 Q
Q 8
o si
H o S
oo jjj
- <
Ml UJ
a
:<
pq
0
C/3
c/3
s-1
■— ' C k- "^ X "
■^ a -. a
— ^- oo -c^
-O a _ ~
rt 5 tj «*<
>■ - »
-C = —
2 ~
B?
2 5 ?>
ci ; rl
*-> -^ **l
oo s *
« m .£. >; s -„.
a ** a ja - -^
— -^ a i- Si«
■^ *- -_* *- \j
- s 5 i s .a -Hi
3 w I £ £ ~ £
* .5
"J
126
San Diego Society of Natural History
Vol. 15
c
o
.S'a
n „Q
» O
V P-t -
»3 S c
C j- <_, rt
rt PS n -*3
so
tfl
as
-.>
4_»
—
u
rt
7*
C
JS
M
s
6
PS
4->
Z
4-1
0
u
w
to
i S
o
C/5
o
0 -
>
rt
rt
£ C
u
^^
u >
2
0
en
SO
OS
o — 3 . —
n 't c £
u c
u « .2
w « «
. J* <*
►J w O
Z c
-o
c
< a.
d. to
~H CO
so
so
o
C
K
K
o
)A
•*■
■«-
so
OS
CO
os
„
^H
c
o
Wi
C
C
0)
a
csl
1-
Hi
Hi
Hi
so
so
Mh
so
so
<-H
OS
„
OS
„
^*
CO
eg
^
^
_='
JB
K
•<*■
0
p4
»*^
Ja
SO
so
O
(4
*-H
SO
l
OS
, ,
OS
<a
OS
•"■
<«
t^\
1-1
C
c
c
, J
C
rt
<+H
0>
rt
EX
0)
O
o
O
o>
u «u
uj «g
< g
g -s
u 5
sJ S
D
§Q
w
■i o
■a
■*
ON
<
a
>3 .
-
OO
<
Q
2: °°
w
*3 1~i
Z2
<-5 Ul
sU
5 2
<
5* o£
■Si c
<
2: -Q
•5 -°
2
c
z
« 3
a o
Q
>
*!i
j-
<
Z
2 £
|s
a
H
c/^
.= O
5 «;
5
D
u.
C^5 " '
o
0
-2 8
oa
^_^
W
<»
fN
<
«^"s
OO
Q
^
w
!j
5
<
c*c
u
%
2
•^1 ^*
.-
a -a
a
D
^ o
O
Cr
—
■3 cq
pi
?
0
>
<5
~5
>
:0
o
1968
Coan: Bahia de los Angeles Mollusks
127
>
o
U
Z
tc
—
Wl
~r
00
z
c
J3
3
o
•4-
1- «=
■4- ™
3
no
> —
ON
no
NO
NO
-5 K
°"4- CO
c4 no
rS °N
«i ^ "*
C _• B
§ a a
K
NO
-1-
K
K
<ni
oo
o
IN.
oo
■^
t+4
«+*
«+H
-f
o
-1-
NO
o"
■4-
Is.
■*
NO
IN.
ON
■>
ON
-f
1-
-r
M
^
X
c
C rN
"S3
*■— N
■J2 oo
<
? ON
■2 ON
&s
*■ >;
O.VE
SVC
a c
«• ^
BC —
1 5
■3«:2
■S3 c
<3 -— -
- _□
" r
-5 3
2 "w
5! n^
2 to
« — -
.2 4
~§ 1
u
H
Q
u
u
U
£
(N|
ON
u" ^
^ -, 3 -S!
O ,-. "£ — J •>■
■SI ON -
ii - = a
~ ON ~
~ On .On
?\ M **< —
<-> os y on
2f j; Sf r- 2? — "5 £
|0 ;Q S — ' C — i
§ S S k f-
^2
■e.
on fi)
~ — . ■- rs
o
oo
A
u
*
o
= on ^2 .=
3 I >: * a
(A
- i ^ ^ <■'
K
<N1
ON
J; c
"S.S
-
—
£?T ~ ~ -
— ^
<
Q
; •_
oi 2 Js
K3
^
z
-
—
—
ft.
C/5
ii
128
San Diego Society of Natural History
Vol. 15
j? 3 S c
C 1- 4_. «
rt rt rs -rj
u ^— ' <y Js
i_ c
- 13 SC 3
so
'■Si
OS
4_»
p— 1
:j
rt
4*
X
^
o
rt
4-J
u
Z
*""
o
u
r
3
_=
d
cr
o
c
o
c
rt
m
(A
E
so
•«— "
os
1-1
—
11
<
"5
rt
en
c
o
<u
C
w
&
rt
_*
(A
r>
rt
^
o
d
u
c
-a
c
< a
^ ^ ^
Z Z Z
U
u
c
rt
*-»
C
>
o
C/5 to
„, SO ^- <N ^
O
i
K
so i K os
-* SO -* —
i— I oo
-4-
■4-
so
o
as
OS
as
■4-
r-1
OS
OS
OS
OS
OS
OS
OS
OS
- - * *- 2
\4 X
c
at
r(
OS
K "
™ C
_Q 3 ^
a ^3 -* "a.
P3
0 cq
OS
"S
OS
Jh 4h ^ v+h
^J i/ * oo
**-l .—I
d _: C •
§ ^S &
tO C/5
OS
OS
o
th
(N
OS
•-1
-
OS
o
l*.
OS
V+H
4-t
—
„
I/)
-c
o
X
■4-
a
O «-. T
u
" ri on £ rsT « o
-a ^ ~ « ~ ij ^
= "5, u _* "S. S "E.
J2 s rt *?
Q *! ca Q
OS
a
OS
1-H
rt
«*■>
rC
„
U
HI
t/>
i-i
pq
,.
<N
#
»-H
, ,
_J
rt
&
Q
<
Q
Z
o
w
H
U
<
K
^^
CM
rN
rs|
(^ U1
OS
ia
■-*i
r—.
*-^
C' co
R »
U
^o
?- rt
<
■S M
■»- c
oo
£P c
g rt
Q
U
BOJS
s ^
-5
u
c
■S <
5 X
0
.^ u-
.5
•S «'
—
H
U
U
>«
>
> — '
•*-. - '
•— i
<
"5
^
^
u
^5 OS
P Q
X
to
G
O
■k. CO
OS
S °2
o
-a
OS
,*o
■^
— •
^
5 as
3 o
*— -V
•2
w
^— v 4_,
^■^ OS
~c I-
*»
<
Q
« C/3
^3 '~l
~Q rt
ij P3
(S
—3
g
ft!
5 jS
fe
—
j
UJ
2
••^ rt
§ c
rt
a,
-o
0)
c
Q
•S
u
w
c
0»
. — «
c
_jj
-a
3
<
;
a.
IhI
^^
5 —
5
5 5
B
(4
*o
rt
**> rt
i*s rt
-c
p
>o TS
«a
5
u
-2 m
-5 Q
X.
*- ^-
k.
E*h
(/I
fci
3
d
d
h
K
Ex
1968
Coan: Bahia de los Angeles Mollusks
129
u
u
Z
u
z
u
2 "5
o J2
ft. c*
3 §
2 N
1- -
a,
c -
3
C o
S 5
-r
t
• M
-i-
...
o
o
__"
c\
•
oc
a
a,
.#.
ON
0\
so
o
<^-
o
0-v
ON
<* «
\0
»"H
so
1-1 o
„
T-H
, •— i
rt .
«
K K
u
•*-,
K
u 0
I/] —
ON ^
«
Mh
as
U-
— -r
id
oo
"u
cXJ K
(A .
(JO
1-t —
i- a.
~
a
Ul
=3 a
pq
Q
pa
Q
U
Lt
pa
cc
«
«^
£
, *
—
Q.
-—>
no
^r
_-
*
NO
ON
c
on
o
NO
*->
NO
o
Si
,1
_Q
u
Is.
< >
*3 <N
—
<N
h:
ON
# ™
ca
C ^3
ft? 72
^— c
3
a 13
1<
■8 d
'5 =3
o
~S d
Vfcl ON
~-5
ft- u
-5 o
a t
51 pa
■a *J
a/;
ft- Q
<> 2
*- s
l\
"Z? ON
■a jc
On
o
0\
I <N ON
ft.
u
'CQ
£
£
ft^m
a —
a ^
a S
£ c
<s 3
^
a ,-v
~a -o
-2 rE
■s =
—
^72 ^
c
3
a =3
a
-£ d
a —
=: -
-2 ■•>
5 ~^
O o\
"ft rj
h
a
k
l<
o
^-s.
^«
-
<N
<
-
OO
<
z
■j
3
a
-^
-
■^
3
CI
3
0
_
>
"c
u
<
a
s_-<
u
-
-a
o
a
c
».2
-->
■r
-*
*
—
n
—
n
c
■j
n
n
—
-^
—
c
u
3
C
i-
3
3
£
3
_ -
130
San Diego Society of Natural History
Vol. 15
C 3
'n -a
s'5
« o £ c
C >~ w rt
rt « rt -n
" c -Q
dm M
o\
u
«
a
Ui
a
C
c>
-'.
-^
.•-
-
o
n
u
X
*""
o
c
a>
y
3
n
j:
c
O"
£
o
O
c
o
c
rt
r<
a
I/)
E
VO
C\
— '
—
—
a
«•"*»
<
w
C
c
'o
M
c
rj
(/I
(/)
6
rs
Z
"
6
c
u
c
-a
c
<
D
Z
X
a,
S
<
ui
<
Q 5
Z
o
h
I
U
o
h
Z
<
u
<
2
<S-
- £
■a on
=: co
o — •
"••4
la u
© -O
■*- . —
5 £,
« -—•
1968 Coan: Bahia de los Angeles Mollusks 13 1
Acknowledgments
Dr. Myra Keen of Stanford University is responsible for the identifications of nearly all of the
molluscan species discussed in this report and for the preliminary organization of the data. She also
offered suggestions and criticized the manuscript. Dr. James Rohlf of the University of Iowa demonstrated
the use of a computer in data analysis. Dr. Joseph Connell of the University of California at Santa
Barbara made many valuable suggestions. Dr. J. Laurens Barnard of the United States National Museum
did the field work during his association with the Beaudette Foundation, with support from the National
Science Foundation Grant 20909.
Mr. Barry Roth of San Francisco State College aided in sorting the collection; Dr. Judith Terry,
then a student at Stanford, helped in the compilation of data; Mrs. Berenice E. Coan typed lists of data
and portions of the manuscript and catalogued most of the collection.
Literature Cited
Baker, F.
1926. Mollusca of the family Triphoridae. (Expedition of the California Academy of Sciences to
the Gulf of California in 1921 — VI). Calif. Acad. Sci., Proc. (4) 15: 223-239.
Baker, F., and G D. Hanna
1927. Marine Mollusca of the order Opisthobranchiata. (Expedition of the California Academy of
Sciences to the Gulf of California in 1921— V). Calif. Acad. Sci., Proc. (4) 16: 123-13 5.
Baker, F., G D. Hanna, and A. M. Strong
1928. Some Pyramidellidae from the Gulf of California. Calif. Acad. Sci., Proc. (4) 17: 205-246.
1930. Some rissoid Mollusca from the Gulf of California. Calif. Acad. Sci., Proc. (4) 19: 23-40.
193 8. Some Mollusca of the families Cerithiopsidae, Cerithiidac and Cyclostrcmatidae from the Gulf
of California and adjacent waters. Calif. Acad. Sci., Proc. (4) 23: 217-244.
Barnard, J. L., and J. R. Grady
1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. I. General Account.
San Diego Soc. Nat. Hist., Trans. 15 (6): 5 1-66.
Bartsch, P.
1910. The West American mollusks of the genus Alaba. U.S. Nat. Mus., Proc. 39: 153-156.
1911a. Descriptions of new mollusks of the family Vitrincllidae from the west coast of America.
U.S. Nat. Mus., Proc. 39: 229-234.
1911b. The Recent and fossil mollusks of the genus Alabina from the west coast of America. U.S.
Nat. Mus., Proc. 39: 409-418.
1911c. The Recent and fossil mollusks of the genus Ccr/th/opsis from the west coast of America.
U.S. Nat. Mus., Proc. 40: 3 27-3 67.
191 Id. The Recent and fossil mollusks of the genus Ahania from the west coast of America. U.S.
Nat. Mus., Proc. 41: 333-362.
1915. The Recent and fossil mollusks of the genus Rissoitia from the west coast of America. U.S.
Nat. Mus., Proc. 49: 33-62.
1917a. Descriptions of new West American marine mollusks and notes on previously described forms.
U.S. Nat. Mus., Proc. 52: 637-681.
1917b. A monograph of West American melancllid mollusks. U.S. Nat. Mus., Proc. 53: 295-356.
1920. The West American mollusks of the families Rissoellidae and Synceratid.ie, and the rissoid
genus Barlceia. U.S. Nat. Mus., Proc. 58: 159-176.
Carpenter, P. P.
18 57. Catalogue of the collection of Mazatlan shells in the British Museum collected by Frederick
Reigen. London (Brit. Mus.). 5 52 p.
Coan, E. V.
1964. Biometrical analysis of Mollusca, Bahia de los Angeles, Baja California, [abstract]. Amcr.
Malacol. Union Rept. for 1964 (Bull. 31): 48.
Coan, E. V. and B. Roth
1966. The West American Marginellidae. Veliger 8: 276-299.
Dall, W. H.
1919a. Descriptions of new species of mollusks of the family Turridac from the west coast of
America and adjacent regions. U.S. Nat. Mus., Proc. 56: 1-86.
1919b. Descriptions of new species of Mollusca from the North Pacific Ocean in the collection ,>t
the United States National Museum. U.S. Nat. Mus., Proc. 56: 293-371.
Dael, W. H, and P. Bartsch
1909. A monograph of West American pyramidellid mollusks. U.S. Nat. Mus., Bull. 68: i-xxi -
1-258.
132 San Diego Society of Natural History Vol. 15
Hertlein, L. G., and A. M. Strong
1951. Mollusks from the west coast of Mexico and Central America. Part X (Eastern Pacific Ex-
peditions of the New York Zoological Society — LXIII). Zoologica 3 6: 67-120.
Johnson, R. I.
1964. The Recent Mollusca of Augustus Addison Gould. U.S. Nat. Mus., Bull. 239: 1-182.
Keen. A. M.
195 8. Sea shells of tropical West America. Stanford, Calif. (Stanford Univ.). 619 p.
1964. A quantitative analysis of molluscan collections from Isla Espiritu Santo, Baja California,
Mexico. Calif. Acad. Sci., Proc. (4) 30: 175-206.
1966a. West American mollusk types in the British Museum (Natural History) II. Species described
by R. B. Hinds. Veliger 8: 265-275.
1966b. Moerch's west Central American molluscan types with proposal of a new name for a species
of Semelc. Calif. Acad. Sci., Occ. Papers no. 59: 1-33.
1966c. West American mollusk types at the British Museum (Natural History) III. Alcide d'Orbigny's
South American collection. Veliger 9: 1-7.
1968. West American mollusk types at the British Museum (Natural History) IV. Carpenter's
Mazatlan Collection. Veliger 10: 389-439.
McLean, J. H.
1961. Marine mollusks from Los Angeles Bay, Gulf of California. San Diego Soc. Nat. Hist.,
Trans. 12: 449-476.
Olsson, A. A.
1961. Mollusks of the tropical Eastern Pacific, particularly from the southern half of the Panamic-
Pacific faunal province (Panama to Peru). Ithaca, New York (Paleo. Res. Inst.). 574 p.
Palmer K. v. W.
195 8. Type specimens of marine Mollusca described by P. P. Carpenter from the West Coast (San
Diego to British Columbia). Geol. Soc. Amer., Mem. 76: 1-376.
1963. Type specimens of marine Mollusca described by P. P. Carpenter from the west coast of
Mexico and Panama. Bull. Amer. Paleont. 46: 285-408.
Parker, R. H.
1960. Ecology and distributional patterns of marine macro-invertebrates, northern Gulf of Mexico.
Recent Sediments, Northwest Gulf of Mexico, 1951-1958. Tulsa, Oklahoma (Amer. Ass. Pet.
Geol.) pp. 302-381.
1964a. Zoogeography and ecology of some macro-invertebrates, particularly mollusks, in the Gulf
of California and the continental slope off Mexico. Vidensk. Medd. fra Dansk Naturhist.
Foren. 126: 1-178.
1964b. Zoogeography and ecology of macro-invertebrates of the Gulf of California and continental
slope of western Mexico. Amer. Ass. Pet. Geol., Mem. 3: 331-376.
Pilsbry, H. A., and A. A. Olsson
1945. Vitrinellidae and similar gastropods of the Panamic Province. Part I. Acad. Nat. Sci. Phila-
delphia, Proc. 97: 249-278.
1952. Vitrinellidae of the Panamic Province. Part II. Acad. Nat. Sci. Phidelaphia, Proc. 104: 35-88.
Reish, D. J.
1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. II. Benthic Poly-
chaetous Annelids. San Diego Soc. Nat. Hist., Trans. 15 (7): 67-106.
Strong, A. M., and L. G. Hertlein
1937. New species of Recent mollusks from the coast of western North America. (The Templeton
Crocker Expedition of the California Acad. Sci., 1932. — -No. 25). Calif. Acad. Sci.,
Proc. (4) 22: 159-178.
193 9. Marine mollusks from Panama collected by the Alan Hancock Expedition to the Galapagos
Islands, 1931-1932. Alan Hancock Pacific Exped. 2: 177-245.
Thorson G.
1957. Bottom communities (sublittoral or shallow shelf). Geol. Soc. Amer., Mem. 67, 1: 461-534.
Turner, R. D.
195 6. The Eastern Pacific marine Mollusca described by C. B. Adams. Mus. Comp. Zool., Harvard
Univ., Occ. Papers on Mollusks 2: 21-136.
Warmke, G. L., and R. T. Abbott
1961. Caribbean scashells: a guide to the marine mollusks of Puerto Rico and other West Indian
islands, Bermuda and the lower Florida Keys. Narberth, Pennsylvania (Livingston Publ.).
346 p.
Accepted for Publication 18 March 1968
Department of Biological Sciences, Stanford University, Stanford, California, 94305.
(*&>$. COMP. ^uOL>
J LIBRARY
0u 18 1968
HARVARD
TYPE SPECIMENS OF BIRDS IN THE
SAN DIEGO NATURAL HISTORY MUSEUM
JOSEPH R. JEHL, JR.
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 9, 25 SEPTEMBER 1968
MUS. COMP. *loOL.
LIBRARY
TYPE SPECIMENS OF BIRDS IN THE SAN DIEGO
NATURAL HISTORY MUSEUM Uoi 18 ^b3
HARVARD
Joseph R. Jehl, Jr. UNIVERSITY
The bird collections of the San Diego Society of Natural History are mainly com-
posed of specimens from the southwestern United States and Baja California, Mexico.
Among the more than 37,000 specimens are 17 holotype and 2 syntype specimens that
served as the bases for 1 8 avian subspecies. Twelve of these subspecies were proposed by
the late Laurence M. Huey, Curator of Birds and Mammals at the San Diego Natural
History Museum from 192 3 to 1961, four by the late A. J. van Rossem, one by the
late Frank Stephens, and one by J. Elton Green and Lee W. Arnold. Of the 18 forms
proposed, ten are currently considered valid by most workers.
In this list of specimens the following information is given: name as originally
published; name currently accepted by the A. O. U. Check-list, 5th ed. or, where perti-
nent, by the Check-list of the birds of Mexico, if different from the original; and type
locality. Geographic coordinates not given in the original descriptions, emendations, and
corrected spellings are given in brackets. The current taxonomic status of each race has
been reviewed. For brevity I have usually cited only the earliest published opinion on
the validity of each race, except that major revisionary studies are also cited. Unless
stated otherwise, races accepted by the first reviewer are recognized in each of the
publications cited below that appeared subsequent to the original description; con-
versely, races considered invalid by the first reviewer have been merged by all subse-
quent authors.
American Ornithologists' Union
1957. Check-list of North American birds, 5th ed.
Friedmann, H. L., L. Griscom, and R. T. Moore
1950. Distributional check-list of the Birds of Mexico, Pt. 1. Pac. Coast
Avifauna 29.
Miller, A. H., H. L. Friedmann, L. Griscom, and R. T. Moore
1957. Distributional check-list of the birds of Mexico, Pt. 2. Pac. Coast
Avifauna 3 3.
Peters, J. L., and others
1931-1962. Check-list of birds of the world. Vols. 1-7, 9, 15. Harvard
University Press.
Cory, C. B., C. E. Hellmayr, and B. Conover
1918-1949. Catalogue of birds of the Americas and adjacent islands. Pt. 1,
nos. 1-4; Pt. 2, nos. 1-2; Pts. 3-11. Field Mus. Nat. Hist. Zool. Ser.
Vol. 13.
To determine the validity of a taxon, it is often necessary to reconstruct the type
series and to independently analyze the material upon which a description is based. For
San Diego Soc. Nat. Hist., Trans 1$ (9): 133-139, : S September 1968
134 San Diego Society of Natural History Vol. 15
this reason paratypes are listed whenever possible, even though they have no nomen-
clatorial standing. No specimens formally designated as paratypes are known to be in
the collections. However, following Cochran (1961. U.S. Natl. Mus. Bull. 220: xii) I
consider paratypes to include all specimens that served as the basis for the original de-
scription regardless of whether they were formally designated.
Huey often listed catalog numbers of specimens that he referred to new taxa; such
specimens I consider unquestioned paratypes. In some instances, he gave only locality
data for referred specimens, but from these data and the dates of collection it is possible
to identify those specimens still extant in this collection. Each series of paratypes thus
identified is indicated by an asterisk. Because of the greater risk of error, I made no
attempt to identify specimens borrowed from other institutions. Stephens, Green and
Arnold, and van Rossem gave no references to specimens they examined.
Catalogue
Hydranassa tricolor occidentalis Huey
San Diego Soc. Nat. Hist., Trans. 5 (7): 83-84; 10 October 1927.
= Hydranassa tricolor ruficollis (Gosse)
Holotype. — Adult male; skin in good condition. Collected by L. M. Huey, 2 5 May
1926. Original no. 54, now SDSNH no. 10644.
Type locality. — Scammon['s] Lagoon, Baja California, Mexico; lat. 28°04'N, long.
114°05'W.
Paratypes*. — 9788, Sweetwater slough, San Diego Bay, California; 30889, La Punta,
San Diego Bay, California; 11414-11416, 1143 5, 11462, 30890, Pond Lagoon, Baja
California, Mexico; 10643, 10645-10647, Scammon's Lagoon, Baja California, Mexico;
11463, San Ignacio Lagoon, Baja California, Mexico.
Status. — Grinnell (1928. Univ. Calif. Publ. Zool. 32: 84) did not recognize this race,
presumably on the basis of a letter from van Rossem stating "material examined in
eastern museums shows great range of variation in birds of the South Atlantic portion
of North America, quite including the extremes in size recorded by Huey from Lower
California. Neither does there appear to be even an average color difference." Peters
(1931, Check-list of birds of the World, Vol. 1, p. 113) placed occidentalis in synonomy
with ruficollis. The A. O. U. Check-list (1931, 4th ed., p. 29) noted only that Huey
had proposed racial separation for the birds from Baja California. Apparently the valid-
ity of this race has never been satisfactorily determined.
Nyctanassa violacea bancrofti Huey
Condor 29: 167-168, fig. 56; 15 May 1927.
Holotype. — Adult male: skin in good condition. Collected by L. M. Huey, 2 5 May
1926. Original no. 64, now SDSNH no. 10654.
Type locality. — Scammon['s] Lagoon, Baja California, Mexico; lat. 28°04'N, long.
114°05'W.
Paratypes''-'. — 10655-10657, Scammon's Lagoon, Baja California, Mexico; 2468, San
Benito Is., Mexico.
Status. — Recognized as a valid subspecies by the A. O. U. Check-list Committee
(1931. Check-list of North American Birds, 4th ed., p. 31).
1968 Jehi: Type Specimens of Birds 135
Callipepla gambeli deserticola Stephens
Auk 12: 371-372; October 1895
= Lophortyx gambelii gambelii Gambel
Syntypes. — Adult male; skin in good condition. Collected by F. M. Stephens, 15 Jan-
uary 1890. Original no. 4969, now SDSNH no. 264. (Stephens incorrectly cited this
specimen as a female, but male sex is indicated by the original label as well as by
plumage.) Adult female; skin in good condition. Collected by F. M. Stephens, I January
1890. Original no. 4938, now SDSNH no. 265.
Type locality. — The male specimen (SDSNH no. 264) was collected at Palm Springs,
San Diego County [now Riverside County], California, elev. 100 ft. The female
(SDSNH no. 26 5) was taken at Walters (now Mecca), Colorado desert [now Riverside
County], California, elev. 2 50 ft. Since the male was listed first by Stephens, Grinnell
(1931, Univ. Calif. Publ. Zool. 38: 270) restricted the type locality to Palm Springs.
Because of this action, the male specimen must be considered a lectotype.
Status. — Considered inseparable from Callipepla gambclii by the A. O. U. Check-list
Committee (1897. Auk 14: 132).
Oreortyx picta eremophilus van Rossem
Condor 39: 22; 15 January 1937.
Holotype. — Adult male; skin in good condition. Collected by A. J. van Rossem,
27 October 193 5. SDSNH no. 17324.
Type locality. — Lang Spring, Mountain Spring Canyon, Argus Mountains, Inyo County,
California, elev. 6000 ft.
Status. — Recognized by the A. O. U. Check-list Committee (1944. Auk 61: 446).
Hellmayr and Conovcr (1948. Cat. birds of the Americas, Vol. 13, Pt. 1, No. 2, p. 229)
merged eremophilus with picta.
Endomychura hypoleuca scrippsi Green and Arnold
Condor 41: 25-29, figs. 14-16; 17 January 1939.
Holotype. — Adult male; skin in good condition. Collected by J. E. Green, 16 Ma)
193 8. Original no. 4105, now SDSNH no. 17934.
Type locality. — Anacapa Island, California.
Staiis. — Recognized by the A. O. U. Check-list Committee (1944. Auk. 61: 449).
Otus asio cardonensis Huey
Auk 43: 360-362; July 1926.
Holotype. — Adult female; skin in good condition. Collected by L. M. Huey 18 April
1923. SDSNH no. 8671.
Type locality. — Canyon San Juan de Dios (10 mi E El Rosario), Baja California,
Mexico [lat. 30° 1 l'N, long. 115°35'W].
Paratypes. — 8607, 8610-8616, 6 mi E El Rosario, Baja California, Mexico; 8671, 10 mi
E El Rosario, Baja California, Mexico.
Status. — Recognized by the A. O. U. Check-list Committee (1944. Auk 61: 450).
and by Marshall (1967. West. Found. Vert. Zool. Monograph 1, p. 13).
Dryobates villosus scrippsae Huey
San Diego Soc. Nat. Hist., Trans. 5(1): 9; 20 February 1927.
= Dendrocopos villosus byloscopus (Cabanis and Heine)
136 San Diego Society of Natural History Vol. 15
Holofypc. — Adult male; skin in good condition. Collected by L. M. Huey, 27 Sep-
tember 1926. SDSNH no. 10847.
Type locality. — La Grulla, Sierra San Pedro Martir, Baja California, Mexico [lat.
30'53'N, long. 115°26'W], elev. 7500 ft.
Paratypes*. — In his original description, Huey (op. cit.) referred 10 specimens from
Sierra San Pedro Martir to this race. Ten specimens in this collection are doubtless the
birds examined by Huey: 8761, 8762, 8786, 8814, 8815, 8842, 10829, 10848, 10853,
108 54. Huey also referred 20 specimens from the Sierra Juarez region to scrippsae. It
is virtually certain that these included the following 14 specimens: 9441, 9497, 9486,
9511, 9524, 9539, 10881, 10882, 10960, 11003, 11019, 11024, 11063, 11077.
Status. — Todd (1946. Ann. Carnegie Mus. 30: 5 12) considered D. v. scrippsae a
synonym of D. v. hyloscopus.
Dryobates nuttallii longirostrata Huey
San Diego Soc. Nat. Hist., Trans. 5(2): 17: 14 July 1927.
= Dendrocopos scalaris crcmicns (Oberholser)
Holofypc. — Adult male; skin in good condition. Collected by L. M. Huey, 1 3 April
1926. SDSNH no. 10448.
Type locality. — San Felipe, Baja California, Mexico [lat. 31°02'N, long. 11415'W].
Paratype. — 31776, San Felipe, Baja California, Mexico.
Status. — As Grinnell (1928. Univ. Calif. Publ. Zool. 32: 121) pointed out, this form
of "Nuttall's Woodpecker" was based on misidentified specimens of Ladder-backed
Woodpecker (Dendrocopos scalaris).
Aphelocoma californica cactophila Huey
San Diego Soc. Nat. Hist., Trans. 9(35): 432, 434; 1 October 1942.
= Aphelocoma coerulescens cactophila Huey
Holotype. — Adult female; skin in good condition. Collected by L. M. Huey, 24 October
1930. SDSNH no. 13 549.
Type locality. — 3 mi N Punta Prieta, Baja California, Mexico; lat. 28°56'N, long.
144°12'W.
Paratypes*. — 32230, 12041, 5 mi N Santa Rosalia Bay, Baja California, Mexico; 32229,
12,000, Mesquital, Baja California, Mexico; 11929-11932, 32225-32228, Llano de San
Bruno, Baja California, Mexico; 11988, 32224 10 mi El Canon, Baja California, Mexico;
13480, 135 52, 13 5 53, 14068, 14070, 2 5 mi N Punta Prieta, Baja California, Mexico;
11782, 9 mi N Punta Prieta, Baja California, Mexico; 13816, 14069, 3 mi N Punta
Prieta, Baja California, Mexico; 11944, Campo Los Angeles, Baja California, Mexico;
15242, 15243, Santa Teresa Bay, Baja California, Mexico; 18275, San Borjas Mission,
Baja California, Mexico.
Status. — Recognized as a valid race by the A. O. U. Check-list Committee (1945.
Auk 62: 445) and by Pitelka (1951. Univ. Calif. Publ. Zool. 5 0 (3): 2 39-243). Blake
(in Peters, 1962. Vol. 15, p. 212) considered cactophila a synonym of hypolcuca.
Auriparus flaviceps ignatius Huey
San Diego Soc. Nat. Hist., Trans. 6(13): 211-212; 30 September 1930.
-Auriparus flaviceps flaviceps (Sundervall)
1968 Jehl: Type Specimens of Birds 137
Holofype. — Adult male; skin in good condition. Collected by L. M. Huey, 8 March
1928. SDSNH no. 11886.
Type locality. — San Ignacio, Baja California, Mexico; lat. 27 17'N, [long. 1 12 5 5'W].
Paratypes*. — Huey referred 23 specimens to Ignatius, which doubtless included the
following 18 in this collection: 15 331, 15332, San Lucas, Baja California, Mexico;
32404, 32405, 10252, San Jose del Cabo, Baja California, Mexico; 32400, Calmalli,
Baja California, Mexico; 12010, 12016, 32401, Mesquital, Baja California, Mexico;
11873, 11881, 24248, 24249, 32403, San Ignacio, Baja California, Mexico; 15335-
15 338, Santa Teresa Bay, Baja California, Mexico.
Stafns. — Grinnell (1931. Condor 33: 167) considered A. f. Ignatius a synonym of
A. f. flaviceps.
Thryomanes bewickii magdalenensis Huey
San Diego Soc. Nat. Hist., Trans. 9(35): 430-431; 1 October 1942.
Holofype. — Adult male; skin in good condition. Collected by L. M. Huey, 20 Novem-
ber 1941. SDSNH no. 18453.
Type locality. — Santo Domingo, Magdalena Plain, Baja California, Mexico; lat.
25G30'N, [long. 111°55'W].
Paratypes. — 18435, 18438, 18440-18444, 18452, 18456, Santo Domingo, Baja Cali-
fornia, Mexico; 18410, Arroyo Seco, Baja California, Mexico.
Status. — Recognized by the A. O. U. Check-list Committee (1948. Auk 65: 441).
Thryomanes bewickii atricauda Huey
San Diego Soc. Nat. Hist., Trans. 9(35) 429, 431: 1 October 1942.
= Thryomanes bewickii cerroensis (Anthony)
Holofype. — Adult male; skin in good condition. Collected by L. M. Huey, 1 3 October
1941. SDSNH no. 19283.
Type locality. — San Borjas [=Borja], Baja California, Mexico; lat. 28°52'N, long.
113 5 3'W.
Paratypes. — 13817, 13556, 13861, 13862, 25 mi N Punta Prieta, Baja California,
Mexico; 11786, Punta Prieta, Baja California, Mexico; 18272, 18282, San Borjas Mission,
Baja California, Mexico.
Status. — I find no evidence that the validity of this form was reviewed by the A. O. U.
Check-list Committee. Miller, Friedmann, Griscom, and Moore (1957. Pacific Coasi
Avifauna 33: 159) synonomized T. b. atricauda with T. b. cerroensis, without stated
reasons.
Dendroica erithachoroides rhizophorae van Rossem
San Diego Soc. Nat. Hist., Trans. 8(10): 67-68; 24 August 1935.
= Dendroica petechia rhizophorae van Rossem
Holofype. — Adult male; skin in good condition. Collected by A. J. van Rossem,
30 April 1930. Original no. 13008; Dickey collection no. 30293; SDSNH no. 17090.
Type locality. — Tobari Bay, Sonora, Mexico; lat. 26 59'N, long. Ill) 56'W".
Status. — Validity of this race accepted by Miller, Friedmann, Griscom, and Moore
(1957. Pacific Coast Avifauna 33: 247).
Remarks. — Miller et al. {op. cit.) incorrectly stated that this specimen was in the
Dickey collection (T. R. Howell, pers. comm.).
138 San Diego Society of Natural History Vol. 15
Dendroica erithachorides hueyi van Rossem
San Diego Soc. Nat. Hist., Trans. 11(3): 50-51; 25 February 1947.
= Dendroica petechia casteneiceps Ridgway
Holotype. — Adult male; skin in good condition. Collected by L. M. Huey, 18 April
1927. SDSNH no. 11471.
Type locality. — San Ignacio Lagoon, Baja California, Mexico [near lat. 26°50'N, long.
113°10'W].
Status. — Considered a synonym of D. /). casteneiceps by Miller, Friedmann, Griscom, and
Moore (1957. Pacific Coast Avifauna 33: 246).
Richmondena cardinalis seftoni Huey
San Diego Soc. Nat. Hist., Trans. 9(21): 216-217; 31 July 1941.
Type. — Adult male; skin in good condition. Collected by L. M. Huey, 2 5 April 1940.
SDSNH no. 18147.
Type locality. — Santa Gertrudis Mission, Baja California, Mexico [lat. 28°03'N, long.
113°06'W].
Parafypes. — 18135, 18146, Santa Gertrudis Mission, Baja California, Mexico.
Status. — Recognized by the A. O. U. Check-list Committee (1948. Auk 65: 442).
Pipilo fuscus eremophilus van Rossem
San Diego Soc. Nat. Hist., Trans. 8(11): 70-71; 24 August 193 5.
Type. — Adult male; skin in good condition. Collected by A. J. van Rossem, 22 May
1935. SDSNH no. 17083.
Type locality. — Lang Spring, Mountain Springs Canyon, Argus Mts., Inyo County,
California, elev. 5 500 ft.
Status. — Recognized by the A. O. U. Check-list Committee (1944. Auk 61: 461)
and by Davis (1951. Univ. Calif. Publ. Zool. 52(1): 103).
Passerculus rostratus anulus Huey
San Diego Soc. Nat. Hist., Trans. 6(10): 204-206; 30 August 1930.
= Passerculus sandivichensis anulus Huey
Holotype. — Adult male; skin in good condition. Collected by L. M. Huey, 21 May 1926.
SDSNH no. 10523.
Type locality. — South side of entrance to Scammon['s] Lagoon, Baja California, Mexico;
lat. 27°54'N, long. 114°18'W.
Status. — Recognized as a valid race by the A. O. U. Check-list Committee (1944. Auk
61: 462) and by Peters and Griscom (1938. Bull. Mus. Comp. Zool. 80: 475).
Amphispiza belli xerophilus Huey
San Diego Soc. Nat. Hist., Trans. 6(16): 229-230; 24 December 1930.
= Amphispiza belli cinerea Townsend
Holotype. — Adult female; skin in good condition. Collected by L. M. Huey, 31 October
1930. SDSNH no. 13 563.
Type locality. — Santa Catarina Landing, Baja California, Mexico; lat. 29°30'N, long.
115C16'W.
Status. — This form was listed by Hellmayr (1938. Zool. Ser., Field Mus. Nat. Hist.,
Vol. 13, pt. 11, p. 543) but was apparently ignored by the A. O. U. Check-list Com-
1968 Jehl: Type Specimens of Birds 139
mittee. Miller, Friedmann, Griscom, and Moore (1957. Pacific Coast Avifauna 33: 381)
considered xerophilus "a synonym, somewhat intermediate toward A. b. cinerea."
Accepted for Publication 6 August 1968
San Diego Natural History Museum, P.O. Box /3 90, San Diego, California.
LIBRARY
OU 1
8 1968
XJl
GEOGRAPHIC VARIATION IN THE CLINGFISH,
GOBIESOX EUGRAMMUS BRIGGS
DAVID W. GREENFIELD AND JAMES W. WILEY
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 10, 25 SEPTEMBER 1968
. *"•
MU3. COM p. ^oOL
LIBRARY
GEOGRAPHIC VARIATION IN THE CLINGFISH,
GOBIESOX EUGRAMMUS BRIGGS UU lg 1959
HARVARD
David W. Greenfield and James W. Wiley
Abstract. — The clingfish, Gobiesox eugrammus Briggs, is redescribed from new material collected
at Punta Banda on the Pacific Coast of Baja California, Mexico and San Diego County, California.
The Guadalupe Island population shows some differences from the mainland population but the species
is considered monotypic. G. eugrammus was found to live near and under the sea urchin
Strongylocentrotus franciscanus. The range of Gobiesox maeandricus is extended to Mexican waters.
Resumen. — Se describe de nuevo Gobiesox eugrammus Briggs, basandose en material recogido en
Punta Banda, en la costa Pacifica de Baja California, Mexico, y en la region de San Diego, California.
Los peces que integran la poblacion que habita la zona de la Isla Guadalupe, presentan algunas
diferencias en relacion con los que se encuentran a lo largo de las costas continentales. Sin embargo,
esta especie se considera monotipica. Gobiesox eugrammus vivo junto con el erizo de mar Sfrongy-
locentrotus franciscanus, y debajo de los mismos. La region habitada por Gobiesox meandricus se
extiende hasta aguas mexicanas.
Gobiesox eugrammus Briggs was originally considered to be endemic to Guadalupe
Island. However, recent field work indicates that its range is more extensive. Briggs
(1965) first recorded the species from the mainland of Mexico at the south side of
Punta Banda. In July 1967, Wiley captured a specimen there at a depth of 30 feet and
in September 1967 he captured another individual at the same locality. In October 1967,
eleven specimens were collected by the authors, P. A. Adams, D. Hensley, and
D. Wildrick at depths of 30 to 90 feet, and in January 1968 the authors collected I J
additional specimens at the same locality. The great number of specimens captured in
these latter trips resulted from our discovery that G. eugrammus associates with the
sea urchin Strongylocentrotus franciscanus. Students from the University of California
at Los Angeles also collected two clingfish from the south side of Punta Banda in
November 1967.
These collections indicate that G. eugrammus maintains populations in the Punta
Banda region. The discovery of a specimen in the Scripps Institution of Oceanography
collections taken at Bird Rock, San Diego County, California, shows that this species
occurs at least occasionally as far north as southern California, although its status there
is not known.
Since these additional specimens substantially supplement the original description
of G. eugrammus, and since the mainland population exhibits a statistically significant
difference from the Guadalupe population, a complete description is presented. I he
uniform plan and sequence adopted by Briggs (1955) is followed, as are his methods of
taking the counts and measurements, with the exception of the dorsal and anal tin ray
counts, which do not include the two extra rays added to the counts bv Briggs. I
graphs of 31 specimens from the mainland and eight specimens from Guadalupe Island
failed to reveal the two hidden rays mentioned by Briggs. The range of body propor-
San Diigo Soc. Nat. Hist., Trans. IS (10): 141-14". 25 SEPTEMBER 1968
142 San Diego Society of Natural History Vol. 15
tions is presented first, followed by the mean. Proportional measurements were made by
arithmetically dividing one measurement into another.
MATERIAL EXAMINED
Mainland population. — United States National Museum USNM 203076 38.0 mm SL, Punta Banda,
Mexico, October 14, 1967: USNM 203077 32.8, 41.0, 41.6 mm SL, Punta Banda, Mexico, October, 1967.
Scripps Institution of Oceanography SIO5 9-307-60C 34.4 mm SL, Punta Banda, Mexico, August, 195 9:
SIO67-209-60C 43.2 mm SL, Bird Rock, San Diego Co., Calif.. October, 1967: SIO68-287-60C 20.1,
23.7, 24.2, 24.6, 26.4, 26.6, 27.6, 28.6, 35.0, 36.0, 36.7, 37.8, 42.8, 43.2, 49.0 mm SL, Punta Banda,
Mexico, January, 1968. California Academy of Sceinces CAS 24139 36.8, 39.3 mm SL, Punta Banda,
Mexico, October, 1967. Los Angeles County Museum LACM-30122-1 32.4, 31.5 mm SL, Punta Banda,
Mexico, October, 1967. Field Museum Natural History FMNH 74571 29.3, 3 1.1 mm SL, Punta Banda,
Mexico, October, 1967. Stanford University 31.4 mm SL, Punta Banda, Mexico, September, 1976. Britisli
Museum (Natural History) 1 968*8* 1 • I 40.8 mm SL, Punta Banda, Mexico, July, 1967. University of
California at Los Angeles W67-150 38.6, 43.8 mm SL, Punta Banda, Mexico, November, 1967.
Guadalupe Island population. — Scripps Institution of Oceanography SIO63-174-60A (4) — Guadalupe
Island, Mexico, April, 1963; SI063-1 67-60A (1) — Guadalupe Island, Mexico, April, 1963; SIO63-184-60A
(3) — Guadalupe Island, Mexico, April, 1963; SIOH50-42-60A (4) — paratypes — Guadalupe Island,
Mexico, February, 1950; SIOH50-32-60A (2) — Guadalupe Island, Mexico, February, 1950.
Gobiesox eugrammus Briggs
Figure 1, Table 1
Diagnosis. — A Gobiesox with poor development of papillae on head. Margin of
upper lip smooth. Dorsal origin much closer to upper pectoral base than to caudal base.
Position of anus variable, usually closer to anal fin origin than rear margin of disc, but
may be closer to rear margin of disc than anal fin origin, or equidistant, under first to
second dorsal ray. Eye 3.72 — 5.10: 4.29 in head. Eye in interorbital .89 — 1.42: 1.13.
Caudal peduncle depth in eye .92 — 1.40: 1.13. Length of disc 2.67 — 3.86: 3.03, tip
of chin to anterior margin of disc 4.77 — 6.95: 5.96, both in standard length. Each
part of disc region C consists primarily of 3 longitudinal rows of papillae; there are
7 or 8 rows across width of disc region A. A row of elongate papillae present on mem-
branes between distal half of pelvic rays. In the adult, tip of depressed anal fin extends
to or slightly beyond caudal base, as does dorsal. Dorsal rays 11 — 13 (usually 12),
anal rays 8 — 10 (usually 10), pectoral rays 21 — 24, caudal rays 12 — 16, and
vertebrae 26 — 29 (usually 27).
Description. — Standard length of specimens 12.6 to 49.0 mm. Body depressed;
depth 4.66 — 7.50: 5.87 in standard length. Caudal peduncle moderate; least depth
1.02 — 1.73: 1.39 in its length. Head well depressed; length 2.34 — 3.07: 2.72 and
width 2.29 — 3.21: 2.69 in standard length. Snout shallow with lateral outline slightly
rounded, 3.37 — 5.60: 4.49 in head. Interorbital 3.03 — 4.93: 3.84 in head. Posterior
nostril directly above anterior edge of eye; anterior nostril with a bilobed dermal flap
extending from its posterior margin. Teeth of lower jaw in two rows, outer row of 3
to 5 pairs of incisors anteriorly, rounded or slightly pointed and larger than inner row.
These followed on each side by row of enlarged canines. Teeth of upper jaw conical,
irregular in size and position, forming a deep patch, followed on each side by row of
1968
Greenfield and Wiley: Gobiesox eugrammus
14:
Figure 1. Gobiesox eugrammus (USNM 203076), Punta Banda, Mexico. Inset of
disc region.
slightly recurved canines.
All papillae on head shallow, lobe-like structures, none on upper lip margin and
none on snout above center of premaxillary groove. Upper attachment of gill membrane
opposite fourth to sixth (usually fifth) pectoral ray. Subopercular spine well developed
but covered by skin of opercle region. Fleshy pad on pectoral base conspicuous, with
free posterior margin extending from area of upper gill membrane. Attachment of axial
dermal flap opposite fifth to ninth pectoral ray. Dorsal-caudal distance extends forward
to point about midway between posterior margin of gill membrane and orbit, 2.26 —
2.90: 2.53 in standard length; postdorsal-caudal distance 2.06 — 3.87: 2.79 in dorsal
length. Depressed tip of anal fin extends slightly beyond vertical line from caudal base.
Coloration. — In alcohol, light brown dorsal and lateral sides covered with more or
less equally spaced light spots about 1 3 diameter of eye, becoming elongated into bars
posteriorly. Darker areas between light spots with numerous small or unpigmented light
spots which are visible with magnification. Body crossed by 7 dark vertical bars, fading
in intensity with preservation. First bar extends from rictus to point under anterior
TABLE 1
Counts of Mainland and Guadalupi Isi and Gobiesox eugrammus
Dorsal Rays
Anal Rays
Pecoral Rays
Principal Caudal Rays
Vertebrae
Mainland 11 12 13
8 9 10
21 22 23 24
12 13 14 15 16
26 27 28 29
3 24 5
1 8 22
4 17 7 3
2 7 17 4 1
2 16 10 1
Guadalupe Island 16 7
14
1 9 4
9 5
1 4 3
144
San Diego Society of Natural History
Vol. 15
margin of orbit. Second originates on lower portion of cheek and runs across head
directly posteriorly to eyes. Next two bands run between upper margins of pectoral
fins, close together; each often appears as a single band. Fifth band widest; extends
anteriorly to dorsal origin. Sixth band runs from anal origin to point midway along
dorsal base. Last band located between last few fin rays of dorsal and anal fins. Caudal
fin dark brown to black. Area between last dark vertical band and caudal fin appears
to have light vertical band across it. Vertical fins brownish-black, with transparent tips.
Pectorals light with peppering of melanophores on anterior two-thirds; on posterior
third of fins rays only peppered, membranes immaculate. Ventral surface of body
lighter, with only scattered melanophores.
/// life, base color of dorsal and lateral sides of body uniform translucent cream.
Ventral parts translucent; abdominal organs visible through skin. Upper parts of body
with light-orange markings, round on head, becoming oval in region of operculum and
more elongate longitudinally near caudal fin. Dorsal and caudal fins black with white
margins. Anal fin grey with white margin. Pectorals and pelvic sucker white. Eye with
red outer and yellow inner rings encircling black pupil.
When placed on a dark background, or upon death, the dorsal and lateral trans-
1.45
140
•
•
1.35
•
•
1.30
125
•
•
•
*
•
*
*
*
1.20
•
*
115
•
*
•
*
*
*
*
* *
1 10
1.05
•
*
*
*
*
*
*
*
*
*
*
*
•
•
1.00
•
•
* *
*
.95
*
*
90
,0 i2 M 16 18 20 22 24 26 28 30 3 34 38 38 40 42 44 46 48
STANDARD LENGTH mm
Figure 2. Relationship of greatest depth of caudal peduncle into eye diameter for
mainland population (stars) and Guadalupe Island population (circles) of Gobiesox
engrain nuts.
1968 Greenfield and Wiley: Gobiesox engrain nuts 145
lucent cream areas become generally darkened as melanophores expand. Concentrated
melanophores form the 7 dark vertical bars mentioned above. These bars look dark
bluish to the naked eye, but appear black under magnification.
Comparison of Mainland and Guadalupe Island Populations
Mainland Gobiesox eugrammus may be separated from the Guadalupe Island speci-
mens by a difference in the ratio of the caudal-penducle depth into the eye. The
Guadalupe population has a mean value of 1.23 with a range of 1.00 — 1.40, while the
mainland population has a mean value of 1.09 with a range of .92 — 1.30 (Fig. 2).
These differences are statistically significant at the .001 level (t = 3.929, 40 d.f.) usin<^
the "Student's" t-test. Although not as useful in separating the two populations, there are
statistically significant differences in the head width and length, tip of chin to anterior
margin of disc, all in standard length, and the width of the orbit in head length
(Table 2). Other body proportions yielding statistically significant differences, such as
length of the disc and greatest body depth, both in standard length, are a result of
TABLE 2
Comparison of morphometrics between Mainland and Guadalupe populations of G. evgrammui
Body Proportion
Guadalupe Island
Range Mean
SD
Main
Range
LAND
Mean
SD
t-VAUUE
40 d.f.
Caudal peduncle
depth in eye
1.00-1.40
1.23
.13
.92-1.30
1.09
.10
3.929 p<\001
Eye in head
3.72-5.10
4.09
.37
3.74-4.95
4.37
.3 2
2.586 p<.025
Length of disc
in SL
2.92-3.86
3.2 5
.28
2.67-3.12
2.93
.13
5.234* p<.001
Tip of chin to 4.77-6.95 5.56 .58 5.48-6.67 6.14 .40 3.885 p<\001
anterior margin
of disc in SL
Greatest depth
in SL
5.72-7.50
6.33
.52
4.66-6.65
5.66
.41
4.629:
1 p<.001
Depth of caudal
peduncle in its
length
1.06-1.73
1.44
.19
1.02-1.62
1.36
.15
1.171
not sig.
Head length
in SL
2.34-2.86
2.63
.14
2.55-3.07
2.76
.12
3.161
p<.005
Head width
in SL
2.33-3.21
2.84
.25
2.29-2.84
2.62
.13
3.880
p<.001
Snout in head
3.37-5.44
4.26
.63
3.40-5.60
4.60
.5 3
1.873
not sig.
Dorsal caudal
length in SL
2.26-2.90
2.57
.17
2.32-2.83
2.51
.12
1.344
not sig.
Postdorsal in
dorsal length
2.30-3.28
2.64
.2 5
206-3.87
2.87
.40
1.971
not sig.
Bony interorbital
width in head
3.13-4.27
3.86
.24
3.03-4.93
3.83
.44
0.222
not sig.
Eye in interorbital
.91-1.42
1.08
.13
.89-1.35
1.16
.13
1.856
nut sig.
'Significance due to allometric growth
146
San Diego Society of Natural History
Vol. 15
allometric growth. Since the sample from Guadalupe Island is composed mostly of
smaller individuals than is the mainland sample (in which allometric growth can be
demonstrated) the t-test simply was comparing large with small specimens. There are
modal differences in both the number of pectoral rays, 22 for the mainland population
and 2 3 for the Guadalupe Island population, and the number of principal caudal rays,
14 for the mainland population and 13 for the Guadalupe Island population. Since
these counts vary inversely, they probably do not result from differences in environ-
mental temperatures in the source areas.
It is possible that the mainland and Guadalupe Island populations represent distinct
subspecies; however the differences in body proportions may be just the result of
environmental conditions. Until additional information concerning the genetic status
of these populations is obtained, they will be treated simply as different at the popu-
lation level.
Ecological Relationships
G. eugrammus is a secretive form which at Punta Banda lives near and under the
sea urchin Strongylocentrotus franciscanus. After this relationship was discovered, it
was possible to locate specimens under about 5 0 per cent of the sea urchins pried off
the rocks. The clingfish generally scurry away, usually heading for the next closest
sea urchin, where they would dart under the protective cover of the spines. In the
deeper water, the orange spots made it difficult to see the clingfish against the back-
ground of the spines. Seven clingfish were placed with sea urchins in an aquarium,
where photographs of their hiding behavior were obtained (Fig. 3).
Figure 3. G. eugrammus in close association with Strongylocentrotus f rand sea mis
1968 Greenfield and Wiley: Gobiesox eugrammus 147
In captivity, clingfish readily took frozen brine shrimp which were fed to them
by hand, and also fed on the isopods that live commensally on the sea urchins. To
determine whether commensal isopods formed a major portion of the clingfish diet, we
preserved in formalin stomachs of eight specimens collected at Punta Banda. Analysis
showed that the major food items were amphipods; however, isopods, polychaetes,
crabs, snails, cowries and limpets were also present. Since the specimens were collected
at about 1000 hours, and the food items had undergone partial digestion and were well
down the digestive tract, we suggest that the clingfish may be a nocturnal feeder,
leaving the protection of the sea urchin at night.
Other species of small fishes found in close association with the clingfish include
the cottids Artedius corallinus, Artedius crcaseri and Orthonopias triads, the ronquil
Ratl.ibunella alien/, and the klipfish Gibbonsia montereyensis.
Zoogeography
Gobiesox eugrammus was apparently derived from the more northern species
G. maeandricus, which appears to be more primitive on the basis of the position of the
posterior nostril (Briggs, 19)5). Although G. maeandricus now is only found commonly
north of Point Arguello, California, with a recorded range from San Diego, California,
to British Columbia, it was probably distributed farther to the south during the Pleisto-
cene when the water temperatures were generally cooler. Individuals are occasionally
carried south by the California current, however, as evidenced by the capture of a
16.0 mm SL specimen attached to a piece of Giant Kelp, Macrocystis, between Guada-
lupe Island and the mainland at 28° 48'N, 115 5 3'W (SIO-65-524-60A) . This speci-
men extends the known range of G. maeandricus into Mexican waters.
G. eugrammus probably evolved from a population of G. maeandricus that became
isolated after the cool waters moved northward at the close of the Pleistocene. It is not
certain, however, whether the ancestral population of G. eugrammus evolved on Gua-
dalupe Island and secondarily invaded the mainland, or whether the initial isolation
occurred in areas of cold upwelling along the mainland of Baja California.
Acknowledgments
The authors are indebted to P. A. Adams, D. Hensley, and D. Wildrick for assistance in field work
and to Carl L. Hubbs, John C. Briggs, and Richard H. Rosenblatt for critically reviewing the manuscript
and making suggestions.
Literature Cited
Briggs, J. C.
1955. A monograph of the clingfishes (order Xenopterygii). Stanford Ichthyological Bull. 6: 1-224.
1965. The clingfishes (Gobiesocidae) of Guadalupe Island, Mexico. ( alif. Fish and Game 51: 123-125.
Accepted for Publication 22 August 1968
Department of Biology, California State College, Fullerton, California 92631
FIVE NEW TAXA OF HAPLOPAPPUS (COMPOSITAE
FROM BAJA CALIFORNIA, MEXICO
MUS. COMP. 2LOOL.
LIBRARY
mar 3M
REID MORAN
HARVARD
UNIVERSITY
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 11 12 FEBRUARY, 1969
^Tijuana
117'
116"
©Mexicali
BAJA CALIFORNIA
117c
I
San Quintin
• H. arborescens
peninsularis
+ H. juarezensis
■ H. odontolepis
© H. pulvinatus
a H. rosaricus
116°
El4Rosario
Figure 1. Northern Baja California, showing all known localities for the five new taxa of Haplopappus.
FIVE NEW TAXA OF HAPLOPAPPUS (COMPOSITAE)
FROM BAJA CALIFORNIA, MEXICO
Reid Moran
Abstract. — The five new plants are distinguished from their nearest relatives as follow-,.
H. (Ericameria) arborescem ssp. peninsularis, of the Sierra Juarez and Sierra San Pedro Martir,
differs from //. arborescem A. Gray ssp. arborescem, of northern California, in having 9-18 rather
than 18-2 5 florets per head; it differs from //. arborescem ssp. parishii (Greene) Moran, of southern
California, in having leaves 1 J/2-3 J/i mm rather than 3-10 mm wide. H. (Ericameria) juarczensis, of
the Sierra Juarez, differs from H. martirensis (Wiggins) Blake, of the Sierra San Pedro Martir, in
its broader leaves, smooth involucral bracts with a narrower herbaceous center, longer achenes, and
more numerous pappus bristles. H. (HazarJia?) odontolepis, of Cerro Potrero, differs from H. verru-
cosus Brandegee, of the foothills of northwest Baja California, in its hispidulous herbage, sessile
leaves, much larger heads with more numerous ray and disk florets, and apparently fertile disk florets.
//. (HazarJia) rosaricus, of the El Rosario region, differs from H. scjiiarrosus H. & A., of west
central California to northwestern Baja California, in its thinner and more branching glabrous stems,
more glutinous herbage with a lemony fragrance, smaller leaves, shorter and less exserted corollas,
and shorter style branches, achenes, and pappus. H. (Stenotus) pull inatus, of the high Sierra San
Pedro Martir, differs from others of the section in its smaller and discoid heads, with shorter corollas
and achenes, borne on peduncles shorter than the leaves; from //. acaulis (Nutt.) A. Gray, of
northwestern North America, it differs further in its smaller and thicker leaves with only the
midrib evident, sparsely glandular but otherwise glabrous herbage, fewer disk florets, and shorter
style branches.
Resumen. — Las cinco plantas nuevas se distinguen de sus parientes mas cercanos, como sigue.
H. (Ericameria) arborescem ssp. peninsularis, de la Sierra Juarez y Sierra San Pedro Martir, difiere dc
H. arborescem A. Gray ssp. arborescens, del norte de California, por tener de 9 a 1 8 flosculos por
cabezuela, en vez de 18 a 25; difiere de H. arborescens ssp. parishii (Greene) Moran, del sur de Cali-
fornia, por tener hojas de 1 J4 a 3 ]/z mm de ancho, en vez de 3 a 10 mm. H. (Ericameria)
jnarezensis, de la Sierra Juarez, difiere de H. martirensis (Wiggins) Blake, de la Sierra San Pedro
Martir, en sus hojas mas anchas, suaves bracteas involucrales con un centro herbaceo mas estrecho.
aquenios mas largos, y mayor numero de cerdas del vilano. H. (HazarJia}) odontolepis, del Cerro
Potrero, difiere de H. vernicosus Brandegee, de las vertientes al noroeste de Baja California, en sus
ramas y hojas hispidulas, hojas sesiles, cabezuelas mucho mas grandes con mas numerosos flosculos del
rayo y del disco, y flosculos del disco aparentemente fertiles. H. (Hazard ia) rosaricus, de la region
circunvecina a El Rosario, difiere de H. scjiiarrosus H. et A., del occidente del centro de California al
noroeste de Baja California, en sus tallos mas delgados, mas ramificados, y glabros, ramas y hojas mas
glutinosas, con fragancia a limon, hojas mas pequenas, corolas mas cortas y menos exsertas, y ramas
del estilo, aquenios, y vilano mas cortos. //. (Stenotus) piiliina/its, de la alta Sierra San Pedro Martir
difiere de las otras de la seccion, en sus cabezuelas mas pequenas y discoid ales, con corolas y aquenios
mas cortos, sustentadas por pedunculos mas cortos que las hojas; de //. acaulis (Nutt.) A. Gray, del
noroeste de Norteamerica, difiere, adem.is, en sus hojas m.is pequenas y grucs.is. siendo solo la
nervadura central evidente, y esparcidamente glandulares pero de otra manera glabras, menos flosculos
del disco, y ramas del estilo mas cortas.
The genus Haplopappus Cass., competently monographed by H. M Hall (1928),
is now under critical cytotaxonomic study by Dr. R. C. Jackson and associates. There
would seem to be plenty of taxonomic problems for everybody, and I would not go out
of my way to poach in another botanist's territory. However, in the course of general
collecting in the state of Baja California, which is more my territory, I have happened
San Diego Soc. Nat. Hist., Trans. 15 (11): 149-164, 12 February 1969
152
San Diego Society oe Natural History
Vol. 15
to find five new plants of this genus and have been able to study them in comparison
with related species in the same area.
In the citation of specimens, field numbers are mine unless otherwise identified.
Herbaria are cited mainly for duplicates already sent out (if not yet incorporated) ;
other duplicates are yet to be sent.
Haplopappus arborescens (A. Gray) H. M. Hall ssp. peninsularis Moran, subspec. nov.
(Fig- 2)
A subspecie typica involucri bracteis H-20 floribusque 9-14 (-IS) ct a subspecie
parishio1 fol/is angustioribus , maximis IVi-^Vi mm lath, diffcrt. Typus: Moran 13554
(SD 65769).
Figure 2. Flowering branches of Haplopappus arborescens ssp. peninsularis, X 0.75; part of the type
collection, from south of El Rayo, Sierra, Juarez, September 1966.
Rounded shrub 1-2 '/> m high, the stem to 1 dm thick at the base, the bark dark gray
or almost black. Leaves linear, acute, the largest 3-6 'j cm long, IJ4-3J4 mm wide, ca.
l/2 mm thick. Flowering shoots l/z-2l/z dm long, the cymes 1 '/j-5 cm wide, flowering
August to October. Involucres 4-5 l/z mm high, 2-3 (or, when pressed, to 4) mm wide,
of 15-20 bracts. Disk florets 9-14 (-18); rays none. Corolla 4-6 mm long. Achenes
1 J/2-2 x/z mm long, the pappus 3J4-5 mm long. Gametic chromosome number: n—9.
Type collection. — Rare in the chaparral at 1600 meters elevation, 2 miles south
of El Rayo, Sierra Juarez, Baja California, Mexico (near 31:58'N, 115°58'W), 17 Sep-
tember 1966, Moran /3 5 54 — holotype: SD 65769; isotypes: CAS, UC, US, etc.
lHaphpappus arborescens ssp. parishii (Greene) Moran, comb. nov. Bigelovia parishii Greene, Bull. Torrey
Club 9: 62. 1882.
1969 Moran: New taxa oi Haplopappus 153
Distribution. — Generally scarce, mostly in chaparral, central part of the Sierra
Juarez at 1150 to 1750 meters elevation and western slope of the Sierra San Pedro
Martir at 1200 to 2200 meters, Baja California. Collections: Sierra Juarez: 6 miles north
of Laguna Hansen, 1700 m, 13608 (SD) ; 5 miles southeast of the sawmill on the road
to La Botella, 1750 m, 1)606 (DS, ICF, KANU, SD, UC) ; 21, miles west of La
Botella, 1750 m, 13511 (DS, SD, UC, US, etc.); 1 V2 miles west of La Botella, 1750 m,
13599a (ICF, KANU, SD, UC), 13599b-k (SD); granitic wash near Mission Santa
Catarina, 3700 feet, Breedlove 4213 (DS). Sierra San Pedro Martir: 7000 feet, Brandegee
in 1893 (UC); foothills, 4000 feet, Robertson 48 (UC); north slope of Cerro Blanco,
1200 m, 14533 (KANU, SD); above Rancho San Pedro Martir on road to Corral de
Sam, 1900 m, Moran & Thome 14115 (DS, RSA, SD, UC, US, etc.); 1 mile east of
Corral de Sam, 2200 m, 15530 (KANU, SD) ; Rancho San Pedro Martir, 1700 m,
74566 (SD); south of La Corona, 2150 m, 11304 (DS, ICF, SD, UC) ; 1 '/, miles west
of Encinal, 1200 m, Moran 25 Thome 14508 (KANU, RSA, SD) ; Vallecito de la Canoa,
1900 m, Moran & Thome 14504 (DS, RSA, SD) ; west of La Grulla, 2100 m, Moran &
Thome 14502 (RSA, SD); La Encantada, 2200 m, Wiggins c5 Demaree 4952 (DS),
Moran & Thome 14)68 (CAS, RSA, SD, UC, US, etc.) ; Santa Rosa, 2050 m, Moran &
Thome 14419 (DS, RSA, SD, UC, US).
Discussion. — Haplopappus arborescens ssp. arborescens occurs mainly in northern
and central California — in the Sierra Nevada from Nevada County to Fresno County
and in the Coast Ranges from the Oregon line to Ventura County. Haplopappus
arborescens ssp. parishii* occurs in southern California — from the San Gabriel Moun-
tains of Los Angeles County and the San Bernardino Mountains of San Bernardino
County to Otay Mountain and the Cuyamaca-Laguna Mountains of San Diego Count).
Haplopappus arborescens ssp. peninsularis occurs in northern Baja California — in the
central part of the Sierra Juarez and on the west slope of the Sierra San Pedro Martir.
It grows mostly towards the upper limit of the chaparral, often in disturbed areas, and
generally is scarce. In the Sierra Juarez it seems to be local: I have found it only within
about 8 miles of Laguna Hansen, but one collection is from 26 miles to the south.
There appear to be no more northern collections, though C. R. Orcutt in particular
explored the northern part of the Sierra for plants at various seasons, making one trip
in September and another in October, when the plant would have been in flower and
very conspicuous. I found it relatively common only in one area, about 3 to 5 miles
southeast of the sawmill on the road to La Botella. In the Sierra San Pedro Martir, again
it is scarce. The cited collections represent all localities at which Dr. R. F. Thorne and
I saw this plant on an 8-day, 60-mile collecting trip through the Sierra. The most
plants we saw in one area were about 2 5, scattered along some 2 miles of trail between
Rancho San Pedro Martir and Corral de Sam.
In number of florets per head, II . a. parishii, with 9 to 12 or rarely 16, is quite
distinct from H. a. arborescens, with 18 to 25. Also, the number of involucral bracts
seems to be correspondingly smaller. In II. a. peninsularis the inflorescence tends to be
smaller than in //. a. parishii, in which the floral shoots are 1 to 5 dm long .\nd the
cymes 2 to 1 8 cm wide; but the heads are very similar. There too, the number of florets
is commonly 9 to 12 and rarely 16 or more. In one collection of //. a. peninsularis
(Z3599), besides several plants with smaller numbers, three had 11 to 14, one had 13
to 16, and one had 10 to 18.
In leaf width also, H. a. parishii, with the largest leaves 3 to 10 mm wide, is quite
distinct from H. a. arborescens, with leaves rarely more than 2 mm wide. Here //. a.
peninsularis is intermediate, with the largest leaves 1 l/2 to 3 ' 2 mm wide. Most specimens
154 San Diego Society of Natural History Vol. 15
of H. a. parisbii are quite distinct from the ssp. peninsularis, having the largest leaves
over 5 mm wide; but some, especially from San Diego County, have narrower leaves,
and a few have no leaves over 3 mm wide. It is true that some of these specimens were
collected late in the season and have only the narrower upper leaves left. But a quick
sampling of four populations (13716, 11718, 13719, 13721) in the Cuyamaca-Laguna
Mountains shows that, as might be expected, individuals vary in leaf width and that a
few individuals at flowering time may have no leaves more than 3 mm wide. Specimens
from such plants apparently cannot be distinguished from the broader-leaved specimens
of H. a. peninsularis. However, San Diego County specimens that cannot be distin-
guished by leaf width evidently are just extremes from a population of mostly wider-
leaved plants and so are best referred to H. a. parisbii on the basis of distribution. The
line between these two subspecies thus is conveniently drawn at the international boun-
dary, where there is an apparent 5 0-mile break in the distribution.
Haplopappus parisbii has always been treated as a species separate from H. arbores-
cens. Hall (1928) pointed out that it scarcely differs except in the decidedly smaller
number of florets and the broader leaves; but the two are quite distinct in these respects
as well as geographically. Clearly H. a. peninsularis, though not intermediate geographi-
cally, does connect the other two in leaf width. Since some specimens are virtually
identical, it can scarcely be treated as a species separate from H. parisbii; and if it were
treated as a subspecies of H. parisbii, the gap between H. arborescens and H. parisbii
would thereby be reduced, the main remaining difference being in the number of bracts
and florets. It therefore seems more reasonable to treat the three as subspecies of one
species.
Dr. R. C. Jackson reports a gametic chromosome number of ;/ = 9 for collection
14368.
At Rancho San Jose, this plant is known as rama hechicera [witch's bush], and
according to Mrs. Meling it is supposed to have some medicinal value.
Haplopappus juarezensis Moran, spec. nov. (Figs. 3,4)
Frutex resinosus bispidulus ad 6 dm alius. Folia oblanceolata ad oboiata, cuneata,
obtusa, integra, enriacea, glanduloso-punctafa, 5-/5 /;//// longa. Capitula solitaria vel
pauca subcorymbosaque, discnidea. Involucra turbinata, 5-/0 ////// longa, 3-5 ;;/;;/ lata,
bract eis 20-3 5, ca. 4-seriatis, lanceolatis, acutis. Flores capituli 12-26, corollis 6-9 //////
longis, ramis styloruiu 2y2-3Vi mm longis. Acbenia subcylindrica, vinacea, birsuta,
4-5 ' 2 mm longa, pappo albo, 6 - <V ;//;;/ longo. Typus: Moran 135 56 (SD 63 5/4). A
H. cuneato et H. martirensi acbeniis longioribns, pap pi aristis numerosioribus et praeterea
a priore ramulis foliiscjne bispidnlis, foliis angustioribus, bracfeis acutis, a posteriore foliis
latioribus, involucri pedunculique bracfeis leribus (in tllo granuloso-glandulosis) differt.
Stiff shrub 2-6 dm high and 5-12 dm wide, sympodially much branched, the stems
to 3 cm thick at the base, the bark gray-brown, shreddy. Herbage hispidulous with
septate conic trichomes ca. 0.1-0.2 mm long. Branchlets 2-7 (-17) cm long, the
first year ca. Yz-\ mm thick, green, hispidulous, pitted and very resinous, terete, in-
conspicuously lined downward from the midrib and margins of each leaf, each usually
terminating with a head in late summer, the second fall sometimes still leafy, to 2 mm
thick, tan, sometimes strongly angulate, with 1-3 or more ascending new shoots from
upper axils; internodes averaging 1 Yz-5 mm. Leaves sessile, oblanceolate to obovate,
cuncate, broadly acute or mostly obtuse to rounded, entire, 5-15 mm long, 2-6 mm
wide above, x/z-\ mm wide at the base, ca. Yi mm thick, sparsely hispidulous, con-
spicuously glandular-pitted, the midrib not projecting but evident dorsally as a trans-
1969
Moran: New taxa of Haplopappus
IJ5
Figure 3. Flowering plant of Haplupappns juarezensis at the type locality, southeast of Las Filipenas, Sierra
Juarez, September 1966.
lucent or glandular line, the lateral veins obscure. Heads discoid, yellow 11-14 (8-16)
mm high, with 12-26 florets, flowering July to October, often solitary and terminal but
sometimes also with 1-3 or more younger ones subcorymbose from the upper leaf axils,
the peduncles 2-18 but mostly 5-10 mm long, slender, glabrous at least above, with
small bracts transitional from the reduced upper foliage leaves to the involucral bracts.
Involucres turbinate, 5-8 (-10) mm high, 3-5 mm wide, the bracts ca. 20-35, erect, in
ca. 4 series, from triangular-lanceolate without to linear-lanceolate within, slender-acute,
1-1 '/4 mm wide, obtusely keeled, chartaceous with a narrow herbaceous or glandular
midline, the margins scarious, somewhat erose, ascending-fimbriate above. Corolla 6-8
(-9) mm long, slender, gradually ampliate upward, glabrous or slightly puberulent on
the tube, the lobes spreading, triangular-ovate, 0.8-1.2 mm long, glabrous. Style branches
long-exserted, 2V2-l)Vi mm long, slender, the appendage equalling or exceeding the
stigmatic part. Achenes subcylindric but tapering slightly to the base, faintly 5 -ribbed,
purplish red, 4-5 Vi mm long, ca. 0.6-0.9 mm thick, densely ascending-pubescent with
slender stiff hairs ca. 1-2 mm long. Pappus 6-8 mm long, of ca. 50-90 scabrous capillar)
bristles, white becoming slightly yellowish. Gametic chromosome number: n=9.
Type collection. — Occasional on granitic hills at about 1575 meters elevation, J.O
miles by road southeast of Rancho las Filipinas, Sierra Juarez, Baja California. Mexico
(near 3 1 47'N, 115 48'W), 17 September 1966, Moran I 3 5 56 -- holotypc: SD 63514;
isotypes: CAS, UC, US, etc.
Distribution. — Known only from a few scattered populations on rocky and
gravelly slopes in the southern half of the Sierra Juarez at 1150 to 1600 meters eleva-
tion. Other collections: l/2 mile southeast of Rancho Calabozo, 1600 m, 13488 (CAS,
SD, UC, US, etc.), 13593 (DS, SD, UC, US, etc.), 13594 (CAS, SD); type locality.
156 San Diego Society of Natural History Vol. 15
Figure 4. Flowering branch of Haplopappus juarezensis, X 1.3; p.irt of the type collection, from south-
east of Las Filipinas, Sierra Juarez, September 1966.
9842 (DS, ICF, SD, UC) ; y2 mile south of La Cienega, 1350 m, 13698 (DS, SD, UC,
US, etc.); Portezuelo de Jamau, 1300 m, 1 167 1 (CAS, SD, UC, US, etc.), 13889 (SD) ;
\Yz miles west of Rancho Alamito, 1150 m, 13634 (DS, SD, UC, US, etc.).
Discussion. — The new species is known only from five localities in the southern
part of the Sierra Juarez; but the area is little known botanically, and the plant very
likely is more widespread than the few collections might suggest. It is often asso-
ciated with such plants as Pinus monophylla Torr. & Frem., P. quadrifolia Pari., ///;//-
perils californica Carr., Nolina palmeri S. Wats., Quercus peninsularis Trel., Q. tur-
binella Greene, Adcnostoma fasciculatum H. & A., A. sparsifolium Torr., and Ccanothus
greggii var. pcrplcxans (Trel.) Jeps.
Haplopappus juarezensis belongs to the section Ericameria (Nutt.) A. Gray. It
most resembles H. marfirensis (Wiggins) Blake, which is similar is size and habit, with
similar pubescence and generally similar inflorescence and flowers. That differs from
H. juarezensis most conspicuously in its much narrower leaves (%-l/4 mm wide),
which give the plant a rather different appearance; the leaves also are more acute and
mucronate, with obscure midribs. Furthermore, the involucral bracts and the upper
bracts of the peduncles are granular-glandular rather than smooth; the involucral bracts
have a broader herbaceous center; the achenes are shorter (ca. 3'/2-4J/2 mm long); and
the pappus bristles are fewer (ca. 30-40). Haplopappus martirensis occurs in the Sierra
San Pedro Martir, the range immediately south of the Sierra Juarez, but at higher ele-
vations than H. juarezensis — at least from 2000 to 2750 meters. It commonly grows
in crevices of steep granitic faces.
Haplopappus juarezensis also resembles H. cuneatus A. Gray in habit and inflores-
cence, and they are the only members of the section with short entire leaves broader than
1969 Moran: New taxa of Haplopappus 157
linear. It differs from H. cuneatus in its hispidulous herbage, narrower leaves, slender-
acute involucral bracts, larger and red achenes, and more numerous pappus bristles.
Haplopappus cuneatus, which ranges from the Sierra Nevada and Arizona to central
Baja California, also occurs in the Sierra Juarez but mainly in the northern part, often
at slightly greater elevations than H. juarezensis or in more shaded spots. At Rancho
Calabozo, where the two occur together, H. juarezensis clearly favors drier and more
exposed places. No hybrids were found.
At the three southern (and lower) localities, H. juarezensis occurs with H. propin-
quus Blake, another discoid Ericameria. That is a larger shrub, glabrous, with linear
leaves, smaller heads, obtuse bracts, and smaller achenes. Again, no hybrids were found.
Dr. R. C. Jackson has made chromosome counts for collections 1355 6, 13593,
13634, 13671, and 13698. For each he reports a gametic chromosome number of n = 9.
At Santa Catarina I was told that this plant is called hierba del pasmo. I have
heard that name applied also to H. cooper/ (A. Gray) Hall; but in Baja California it
seems to be used mainly for H. propinquus Blake, a plant of widely reputed medicinal
value. The same name was reported by Blake (1926) as used in Baja California and
Sonora for H. sonoriensis Blake; but at Bahia de los Angeles that plant, which is not
used medicinally, is called romerillo and is distinguished from the medicinal hierba del
pasmo (H. propinquus), in that region occurring only in the higher mountains. The
same name is reported as used in Mexico for various plants of other families; it pre-
sumably implies that the plants are considered as remedies for tetanus.
Haplopappus odontolepis Moran, spec. nov. (Fig. 5)
Fruficnlus hispidulus resinosus 3 dm alfus, ramulis e caule crasso dense adscen-
dentibus. Folia sessilia, elliptic a, spinoso-dentata, coriacea, gland uloso- punctata, 6-1 8 mm
longa, 2-4 mm lata. Capitula solifaria rel pauca cymosaque, radiafa, flaia. Involucra
cylindro-turbinata, 1 0-12 mm longa, 3-5 /;;;/; lata, bracteis ca. 25, exterioribus foliis
similibus, spinoso-dentatis, squarrosis. Flores radii 6-8, ligulis 6-61A mm longis. Flores
disci 12-14, corollis 7J4-8J4 mm longis. Achcnia radii discic/uc (immatura) similia,
5 -cos fata, [nibescentia. Typus: Moran 12162 (SD 61119). Species H. vernicoso affinis,
sed ille subglalh'r basique gracilior, foliis snbpetiolafis, capitulis minoribus, floribus
paucioribus minoribusque, acheniis disci sterilibiis multoque deminutis. Hae duae ab omniis
congeneribus mihi notis imolucri bracteis exterioribus dentatis valde differunt.
Glutinous shrublet ca. 3 dm high, hispidulous with trichomes 0.1-0.2 mm long,
with an erect stock 5-10 cm high and to 2]/2 cm thick bearing at the summit main
crowded erect or ascending branches; first-year branches simple or few branched above,
light tan, ca. 1 mm thick, angled from leaf margins and midrib downward, sparsely
hispidulous, leafy throughout, with internodes averaging 2 J/2-6 mm, and mostly with
small leaves fascicled in the axils; older ones branching further, to 6 mm thick, the hark
dark gray, shreddy, yellowish in the new fissures. Leaves sessile .\nc\ subclasping, elliptic
to cuneate-oblanceolate, acute, 6-12 (-18) mm long, (2-) 3-4 mm wide above, 1-2 mm
wide at the base, coriaceous, glandular-pitted and slightly to heavily resinous, hispidulous
but less so ventrally, the midrib white, prominent dorsally, the lateral veins obscure, the
apex deflexed, tipped with a white spine, the margins : : crisped, each with 2-6 irregu-
larly spaced spreading white spinose teeth ' _>-l mm long, the first few leaves of the shoot
smaller and entire. Heads solitary on the branches or on branchlets mostly more than
1 cm long, bright yellow. Involucres cylindric-turbinate, 10-12 mm long, 3-5 mm wide
excluding the squarrose tips, of ca. 25 bracts; outer 8-10 bracts graduated, leaflike or
transitional, the exposed upper part herbaceous, glandular-pitted, 2-4 mm wide, with
158
San Diego Society of Natural History
Vol. 15
Figure S. Dried flowering branch of Haplopappus odontolepis, X 1.15; part of the type collection, from
Cerro Potrero, May 1965.
marginal teeth and recurved spinose tip, the covered part stramineous, 1 1/2-2 mm wide,
ciliate; the inner ca. 1 5 bracts oblong to linear, acute, cartilaginous with margins nar-
rowly scarious and erose-ciliate, 8-10 mm long, the outer of these ca. 2 mm, the inner
ca. 1 mm wide. Ray florets 6-8; corolla tube 5-5 V? mm long, the ligule 6-6l/2 mm long,
1.6-1.9 mm wide, irregularly 3-lobed for 1 mm or less; style branches linear, obtuse,
L-l Yz mm long, stigmatic to the apex, unappendaged. Disk florets 12-14; corolla slightly
ampliate, 7/4-8 J/2 mm long, the teeth triangular-ovate, 0.6-0.9 mm long; anthers 2J4-3
mm long; style branches linear, acute, 1/4-2 mm long, the puberulent appendage 1-2
times longer than the stigmatic part. Ray and disk achenes (immature) similar, 5-ribbed,
ca. 2 J/2 mm long, sparsely ascending-pubescent. Pappus of ca. 5 0 scabrous brownish-
white bristles 4-5 J/2 mm long.
Type collection. — Rare on the north slope at 1400 meters elevation, near the sum-
mit of Cerro Potrero, Baja California, Mexico (near 29°49'N, 114C37'W), 30 May
1965, Moran 121 62 — holotype: SD 61139; isotypes: CAS, GH, ICF, K, KANU, NY,
UC, US.
Distribution. — Known only from the type collection.
Discussion. — Cerro Potrero is the rounded peak at the head of Arroyo Cataviria
and about 7 miles north-northwest of ex-mision Santa Maria. It has also been called
Cerro Ugarte, but that name is not in local use. Though scarcely higher than another
peak just south of Santa Maria, apparently it is the highest summit in the 100 miles
between Cerro Chato, at the south end of the Sierra San Pedro Martir, and the Sierra
San Luis. Dominant shrubs at the summit and on the north slope of Cerro Potrero are
Salvia chionopeplica Epling, Juniperus californica Carr., and Eriogonum fasciculation
Bcnth. Other characteristic plants are Stipa speciosa T. & R., Yucca ivhipplei ssp. eremica
Epl. & Haines, Qucrcns turbinella Greene, Kibes quercetornm Greene, Primus ilicifolia
1969 Moran: New taxa of Haplopappus
(Nutt.) Walp., Rhus ovata Nutt., Brickellia frutescem A. Gray, Eriophyllum conferti-
florum (DC.) A. Gray, Haplopappus propinquus Blake, and Senecio lemmonii A. Gr.n.
In three hours on the summit of Cerro Potrero, I found only two plants of Haplopappus
odontolepis. I did not find it on the slightly lower Cerro Santa Maria, about 5 miles to
the south.
The new species is most similar to H. vernicosus Brandegee, native in the foothills
from north of San Vicente to southeast of El Rosario at elevations of about 5 0 to 900
meters. That species was known to Hall (1928) only from two specimens of the type
collection; and Wiggins (1964) commented that it was little known and apparently rare.
In fact, it occurs at many localities in the foothill area, as shown by a score of recent
collections, and usually is abundant where it occurs.
Haplopappus odontolepis resembles H. vernicosus in its slender and angled branches
and its dentate and glandular-pitted elliptic or oblanceolate leaves, and the two are
unique in the genus in their spiny-margined leaflike outer involucral bracts. However,
H. odontolepis differs from H. vernicosus in many respects. The branches are more
crowded-ascending from a thicker base. The herbage is moderately hispidulous rather
than very sparsely hispidulous or mostly glabrous. The leaves are not so conspicuously
narrowed to a petiole-like base; and their marginal teeth are not so large, though the
white spine ending each tooth is a little longer. The heads are more commonly solitary
and are almost twice as long; and the toothed leaflike outer bracts, for which the species
is named, are more numerous, larger, and more conspicuous. Both ray and disk florets
are more numerous and somewhat larger. Immature achenes of the disk florets are like
those of the rays, not greatly reduced as in H. vernicosus: probably disk as well as ray
achenes are fertile.
Though noting that its herbage was not so clearly impressed-punctate as in other
species, Hall (1928) placed H. vernicosus in Ericameria because of similarities to H.
citneatns A. Gray, especially in habit, leaf shape, and pappus. It thus became the only-
species in the section with toothed leaves and of course the only one with toothed involu-
cral bracts. From the scanty insect-damaged material, Hall thought the disk achenes
sterile and — perhaps a related condition — the style branches not differentiated into
stigma and appendage; in these respects also the species would be unique in Ericameria.
Ample material now confirms Hall's tentative view as to disk achenes and style branches.
Haplopappus odontolepis, which clearly is most similar to H. vernicosus, likewise
would be anomalous in Ericameria because of its toothed leaves; furthermore, it has larger
heads than any species included by Hall in Ericameria. However, disk achenes are like ra)
achenes and so presumably are fertile, and style branches of disk florets are appendaged.
These facts suggest (1) that the lack of differentiation of style branches in disk florets
of H. vernicosus is indeed related to sterility of achenes and (2) that the conditions of
achenes and style branches in H. vernicosus are not germane to its sectional position since
that species must in any case be placed with H. odontolepis, in which these are normal.
It now appears that the similarities of H. vernicosus to //. citneatns arc superficial
and that H. vernicosus should be removed from Ericameria and placed with //. odon-
tolepis, possibly in a section of their own. Mainly on the basis of gross morphology, but
with support from a chromosome count for H. vernicosus, Dr. Jackson suggests that
these two belong to section Hazardia or are closely related to it.
Haplopappus rosaricus Moran, spec. nov. (Fig. 6)
Frutex glaber glutinosus fragrans 2-9 dm alius, ramis gracilibus, angulatis. Folia
spatulata, spinoso-dentata, coriacea, glanduloso-punctata, \-2x \ cm longa, 5-/2 mm lata.
160 San Diego Society of Natural History Vol. 15
Figure 6. Flowering branch of Haplopappns rosaricus, X ca. 1.0; part of the type collection, from south-
east of El Consuelo, July 1967.
Capitnla thyrsoidea, discoidea, flava. Involucra turbinata, 8-10 mm longa, 3]/2-5 mm
lata, bracteh 35-60, 5-8-seriatis, erectis, oblongis, mucronatis, ca. 1 mm lath. Florcs
capital/ plerumque 17-23, corollis glabris, 5l/2-6]/2 mm longis, ramis stylorum 1.1-1.5
mm longis, appendicibus ovatis. Achenia fusiformia, 5-costata, adpresso-pubescentia,
3-3 ' S mm longa, pappi aristis ca. 25-40, scabris, 5-6 mm longis. Typus: Moran 14020
(SD 65765). Species H. squarroso proxima sed caulibus brevioribus glabris basique
ramosioribus, foliis minoribus glutinosioribus basique attenuatis, involucri sqnamis erectis,
corollis ramis stylorum acbenibus pappoqite brevioribus divert.
Glabrous, glutinous, lemony-fragrant shrub 2-9 dm high and 2-13 dm wide, with
many slender erect or closely ascending branches from a base 1-2 cm thick. Branches
zigzag, angled from the leaf margins and midrib downward, at first tan, very glutinous
and somewhat granular, ca. 1 mm thick, becoming gray and to 3 mm thick, the inter-
nodes averaging ca. 5-15 mm, the axils sometimes with a few fascicled small leaves.
Leaves sessile or subpetiolate, obovate to spatulate, obtuse to rounded, 1-2 J/2 cm long,
5-12 mm wide above, 1-2 mm wide at the base, coriaceous, conspicuously glandular-
pitted or in youth somewhat rugose, heavily glutinous, green or sometimes whitish with
the dried exudate, spinose-tipped and dentate with 2-8 teeth on each margin, though
sometimes entire in the lower half, the teeth deltoid, l/z-\l/z mm long, each tipped with
a conic white spine, the midrib inconspicuous but in dried leaves whitish and protruding
dorsally, the lateral veins obscure. Heads discoid, yellow, 9-13 mm high, with 12-30 but
commonly 17-2 3 florets, flowering June to October, terminal and solitary or mostly
thyrsoid, with 1-2 sessile or short pedunculate in each upper axil, the youngest below.
Involucres turbinate, 8-10 mm high, 3J4-5 mm wide, of ca. 35-60 bracts closely im-
bricated in 5-8 series, the bracts erect or with the tips slightly spreading, chartaceous
1969 Moran: New taxa oi Haplopappus 161
with scarious margins and short herbaceous tips, glandular internally as well as externally
near the apex, erose and weakly spinose-mucronate at the apex, ca. 1 mm wide, the outer
oblong, the inner linear-oblanceolate. Corolla 5l/2-6l/2 mm long, glabrous, gradually
widening ca. from ]/z mm at the base to 1 mm at the apex, the lobes erect, triangular
ovate, acute, 0.6-0.9 mm long. Style branches 1.1-1.5 mm long, the appendage ovate,
acute, puberulent, half as long as to nearly equalling the stigmatic part. Achenes sub-
terete, narrowed at base and somewhat at apex, 3-3 '2 mm long, reddish black, with
5 prominent white ribs, rather sparsely appressed pubescent. Pappus 5-6 mm long, of
ca. 2 5-40 scabrous unequal bristles, white in flower but becoming sordid. Gametic
chromosome number: ;/=5.
Type collect/oil. — Locally common on north slope at 75 meters elevation, Arroyo
del Campo Viejo, 2.3 miles southeast of El Consuelo and 7.2 miles north-northwest of
El Rosario, Baja California, Mexico (near 30°09/2'N, 115:46'A'W), 2 July 1967,
Moran 14020 — holotype: SD 65765; isotypes: CAS, UC, US, etc.
Distribution. — On north slopes and higher ridgetops in the foothills within 12
miles of the sea, from northeast of El Socorro to southeast of El Rosario, at 75 to 47 5
meters elevation; often rather abundant. Other collections: 5 miles west of Rancho el
Cipres, 350 m, 14017 (DS, KANU, SD) ; 4 miles ENE of El Socorro, 150 m, 14035
(CAS, SD, UC, US, etc.) ; canyon north of El Rosario, Gander 9598 (CAS, SD) ; 3 miles
north of El Rosario, 200 m, 14021 (DS, SD, UC, US); Vi mile north of El Rosario,
75 m, 14022 (KANU, SD); 7 miles west of El Aguajito, 240 m, 15384 (CAS, SD, UC,
US, etc.) ; divide 9 miles ESE of El Rosario on road to Puerto San Carlos, 47 5 m, / 1 242
(DS, ICF, RSA, SD, UC), 14025 (CAS, SD, UC, US, etc.).
Discussion. — Haplopappus rosaricus is rather abundant at most places where it
occurs. Common associates at most of the localities are Agave shauii Engelm., Rosa
minuti folia Engelm., Euphorbia misera Benth., and Ambrosia chenopodiifolia (Benth.)
Payne. Other species of Haplopappus with which it sometimes grows are H. propinquus
Blake, H. venetus ssp. oxyphyllus (Greene) Hall, and H. verrucosus Brandegee. No
hybrids were noted.
Haplopappus rosaricus is most closely related to the polytypic H. sc/narrosus H. & A.,
whose southern subspecies grindelioides (DC.) Keck extends south at least to within 40
miles of the range of H. rosaricus. In H. squarrosus, the plant is generally larger and
less branched, the base thicker (to 9 cm), the branches taller (to l'j meters) and
thicker. Various parts, but especially the stems, are pubescent, at least in those southern
forms that otherwise more closely approach H. rosaricus. The herbage is less glutinous
and has a different odor, not so sweet and lemony. The leaves are larger (ca. 1 ' _■-
6 X 1-3 cm), and they tend to be broader and clasping at the base. The involucre is
sometimes larger, and its bracts are commonly squarrose. The corolla is longer (7-11 mm)
and more exserted; the style branches are longer (1.7-ca. 3 mm); the achenes are longer
(3-5 mm or more) and often glabrous; and the pappus is longer (6-12 mm).
Dr. R. C. Jackson reports a gametic chromosome number of n-S for collection
1402 5.
Several natives of El Rosario knew no name for this plant. However, one man said
that it was used in a remedyr for toothache.
Haplopappus pulvinatus Moran, spec. nov. (Fig. 7)
Planta cues pit osa ad 18 cm lata in rupium rimis. Folia 10-20 mbrosulata, coriacea,
linearo-oblanceolata, acuta, Integra, sparsim glandulosa, /-/': cm longa, ca. I mm la/a.
bast subamplccfcntia. Cap/tula solitaria, discoidea, flava, pedunculo foliis breviore. In-
162
San Diego Society of Natural History
Vol. 15
&f&&&^Pi^x
Figure 7. Flowering plant of Haplopappm pitliinatiis at the type locality, Cerro "2828", Sierra San Pedro
Martir, July 1968.
vohicra turbinata, 5-7 ^ mm longa, 4-5 mm lata, bracteis 20-}}, }-4-seriatis, ellipticis
ad lanceolatis, acutis, marginibus scariosis, exterior/bus ca. I mm interioribus ca. 1 Vz mm
latis. Flores capituli 14-} 6, corollis glabris, }l/z-6 mm longis, ramis stylontm linearibus,
acutis, 1.3-2.1 mm longis. Achenia ll/2-} mm longa, hirsuta, pappo albo, 2l/2-5 mm
longo. Typns: Moran 15262 (SD 68466). Ab aliis sectionis Stenoti capitulis minor/bus
discoideisque, corollis achcniisque breiioribus, pedunculisquc foliis breiioribus differt;
et a H. acauli foliis minoribus crassioribus praeterque glandes dispersas glabris, costa
solum conspicua, disci floribits paitcioribus, ramisque stylontm breiioribus praeterea
differt.
Plant forming a flat or convex cushion to 10 or rarely 18 cm wide, the stems
closely branching, the interstices commonly packed with soil, the surface green with
crowded leaf rosettes. Stems to 6 mm thick at the base, with brown bark, the branchlets
1-2 mm thick, each with ca. 10-20 crowded leaves at the apex and densely covered
below with old leaves that are at first white or tan and later dark brown. Leaves ascend-
ing and commonly outcurved, coriaceous, green, linear-oblanceolate, acute, entire,
channelled ventrally, 1-1 J/z cm long, %-l '/i mm wide above, a quarter to half as wide
just above the broadened clasping base, sparsely beset with trichomes less than 0.1 mm
long each tipped with a yellowish viscid globule ca. 0.5 mm thick, the midrib promi-
nent, protruding dorsally, a parallel nerve on each side obscure except at the base, the
apex subconic, whitish. Peduncles terminal, shorter than the leaves, ca. x/z mm thick,
angled, sparsely glandular, with a few linear, acute, decurrent bracts decreasing in size
upward, the upper ca. 3 mm long. Heads solitary, discoid, yellow, 7-9 mm high, with
14-36 florets, flowering in June and July. Involucre turbinate to narrowly campanulate,
5-7/2 mm long, 4-5 mm wide (5-9 mm wide pressed), of 20-33 erect bracts imbricated
in 3-4 series, the bracts elliptic to lanceolate, chartaceous below, green above, with
scarious margins, the outer ca. 1 mm wide, acute, often sparsely glandular-ciliate, the
inner ca. 1 ]/z mm wide, mostly more broadly acute, usually fimbrillate above. Corollas
glabrous, 3 J/2-6 mm long, 0.3-0.4 mm wide at the base and slightly ampliate above,
the lobes triangular-lanceolate, acute, 0.9-1.4 mm long. Style branches linear, acute,
1969 Moran: New taxa of Haplopappus 16^
1.3-2.1 mm long, the appendage usually exceeding and often twice exceeding the stig-
tnatic part. Achenes light brown, with 5-8 unequally spaced low whitish ribs, 1 l/2-l mm
long, ca. y2 mm thick and tapering to the base, hirsute with ascending white trichomes
Yz-\ mm long. Pappus 2I/2-5 mm long, of 25-35 unequal scabrous white bristle.
Somatic chromosome number: 2«=18.
Type collection. — Rather scarce, in crevices of east- and north-facing rocks and
cliffs at 2800 meters elevation, east slope below Cerro "2828", east rim of the Sierra
San Pedro Martir, Baja California, Mexico (near 31°02'N, 115°27'W), 5 July 1968,
Moron 1 5262 — holotype: SD 68466; isotypes: CAS, GH, K, KANU, MEXU, RSA,
uc, us.
Distribution. — Known only at the type locality.
Discussion. — The new species seems rather rare and limited as to habitat: despite
wide collecting in the area on several trips, I could find only a dozen plants, all on the
east slope of the one peak. Associated plants include Selaginella asprella Maxon, Sedum
niieuw Davids., Hcuchera leptotneria var. peninsularis Rosend., Butt., & Lak., Saxifraga
eriophora S. Wats., Potentilla ubceleri S. Wats., Tanacetum sp., and Stcpbanomeria sp.
Haplopappus puhinatus belongs in the section Stcnofus (Nutt.) A. Gray (as sug-
gested by Dr. Jackson /';/ lit.), even though it differs from previous members and hence
from Hall's (1928) diagnosis in its pulvinate and scarcely subshrubby habit and in its
discoid heads. It differs from the others also in having peduncles shorter than the leaves;
and it further differs from each of them, at least on the average if not in each case
absolutely, in having smaller heads, shorter corollas, and shorter achenes. In fact, it
represents an extreme of reduction and compaction for the section Sfenofns.
Haplopappus puhinatus appears closest to H. acaulis (Nutt.) A. Gray, native
from Saskatchewan to Colorado and east central California. The smaller-leaved and
more densely caespitose forms of that variable species approach H. puhinatus in habit,
the stems and leaves are sometimes similarly glandular, and the involucral bracts are
nearly as many and in about three series. However, the leaves are wider and often
longer, with the lateral veins more evident, and are scaberulous at least on the margins
and often throughout; the disk florets are more numerous; and the style branches are
longer.
In size and shape of leaves, H. puhinatus more nearly resembles //. stenophyllus
A. Gray, native from Washington and Idaho to Nevada and northeastern California. In
that species, however, the leaves are more scattered; the stems, leaves, and bracts are
densely glandular-scabrid; the involucral bracts are fewer and nearly equal; and the
corollas are puberulent.
Haplopappus maclcanii Brandegee, of the Yukon Territory, is similar to //. pul-
linatus in having small but thick leaves. However, the stems are more elongate; the
leaves are narrower, hispid-ciliate, and more numerous on the stems; and the involucral
bracts are fewer and more nearly equal.
On 2 5 August 1968 I collected achenes at the type locality from a plant that had
provided part of the type collection. From these Dr. R. C. Jackson reports a somatic
chromosome number of 2;/= 18.
Acknowledgements
I am grateful to Drs. R. C. Jackson, Peter Raven, and Robert F. Thorne for critically reviewing
the manuscript and to Biologist Mario Sousa S. for preparing the Spanish summary. Ur. Jackson also has
kindly provided the chromosome counts reported here. And I am indebted to the Moran Travel and
Contingency Fund for supporting the field work.
164 San Diego Society of Natural History Vol. 15
Literature Cited
Blake, S. F.
1926. Asteraceae. //; Paul C. Standley, Trees and shrubs of Mexico. Contr. U.S. Natl. Herb.
23: 1401-1641. [The vernacular names contributed chiefly by Paul C. Standley.]
Hall, Harvey M.
1928. The genus Huplopappus: a phylogenetic study in the Compositae. Carnegie Inst. Publ. 389:
i-viii, 1-391, figs. 1-114, pis. 1-16.
Wiggins, Ira L.
1964. Flora of the Sonoran Desert, p. 187-1740. /;; Forrest Shreve and Ira L. Wiggins, Vegetation
and flora of the Sonoran Desert. Stanford University Press.
Accepted for Publication IS November 1968
San Diego Natural History Museum, P. O. Box 1590, San Diego, California 92112.
FOSSIL GROUSE OF THE GENUS DENDRAGAPUS
MUS. CO MP. ZOOL
LIBRARY
JOSEPH R. JEHL, JR. MAR 3 1969
HARVARD
UNIVERSITY
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 12 12 FEBRUARY 1969
FOSSIL GROUSE OF THE GENUS DENDRAGAPUS
Joseph R. Jehl, Jr.
Abstract. — Osteological variation in the Pleistocene and Recent populations of Dendragapus is
reviewed. Two species of Dendragapus are recognized from late Pleistocene deposits of the western
United States. Dendragapus lucasi, which was smaller than the extant D. obscurus, and D. g. gilli, a
large, heavy-bodied grouse, are known only from Fossil Lake, Oregon. D. nanus, also described from
the Fossil Lake deposits, is considered a synonym of D. lucasi. Another grouse, known from cave
deposits of northern California, is tentatively described as a small, southern subspecies of gilli, Den-
dragapus gilli milleri n. subsp. Phylogenetic relationships between the Pleistocene grouse and
D. obscurus are not currently resolvable.
Grouse of the genus Dendragapus* are not common in the fossil record. Three
forms, Dendragapus lucasi (Shufeldt), D. nanus (Shufeldt), and D. gilli (Shufeldt) are
known only from late Pleistocene deposits at Fossil Lake, Lake County, Oregon. These
fossils seem to have been collected from a narrow series of beds with a maximum age
of approximately 29,000 years (Allison, 1966). Abundant material referred to the
extant Blue Grouse, D. obscurus, was reported by Miller (1911) from the Samwel and
Potter Creek cave deposits of Shasta County, California. These deposits are of Upper
Pleistocene age, but they have not been precisely dated. The locations of the Pleistocene
deposits are shown in Figure 1. Dendragapus obscurus has also been reported from an
Indian midden in the Puget Sound region, Washington (Miller, 1960), and from the
Weiss and Birch Creek Valley rock shelters in Idaho (Miller, 1963). Carbon 14 dating
indicates a maximum age of 10,000 to 12,000 years for the Idaho remains.
For an earlier study (Jehl, 1967) it was necessary to assemble most of the Fossil
Lake Dendragapus material previously studied by Howard (1946), the largely unstudied
California cave fossils, and a large series of skeletons of modern D. obscurus. Because
osteological variation in obscurus proved to be greater than had been recognized pre-
viously, I restudied the differences on which the fossil species had been based in light of
this variation. In this paper I review the fossil populations of Dendragapus and I attempt
to outline their possible relationships to each other and to D. obscurus.
Materials
I examined the following skeletal material representing seven of the eight currently
recognized races (A.O.U., 1957) of Dendragapus obscurus: D. o. richardsoni, 1 com-
plete and 2 partial females, 3 complete males; D. o. pallidas, 1 complete and 2 partial
females, 1 complete male; D. o. oreinus, 3 complete and 2 partial females, 1 complete
and 1 partial male; D. o. fuliginosus, 5 complete females, 4 partial males; D. o. sitkensis,
1 partial female; D. o. sierrae, 3 complete and 3 partial females, 5 partial males, 1
unsexed; D. o. howardi, 3 partial males. Fossil material examined is indicated below
under "Referred material" and includes all known material from Fossil Lake examined
'Short (1967) has recently proposed that Canachites and Falcipennis, genera recognized by IYters (1934),
be placed in an expanded genus Dendragapus. In this paper I use DendragapUi in its traditional sense.
However, except for a single report of Canachites canadensis from the late Pleistocene of Virginia
(Wetmore, 1962: 7-8) there is apparently no fossil record of either Canachites or Falcipennis (Brod-
korb, 1964).
San Diego Soc. Nat. Hist., Trans. 15 (12): 165-174. 12 Fi bruary 1969
166
San Diego Society of Natural History
Vol. 15
by Howard with the exception of two elements of D. lucasi ( 1 coracoid, 1 tarsome-
tarsus) in the Oregon State University collections (Howard, 1946: 179).
The following abbreviations are used: AMNH, American Museum of Natural His-
tory: UCMP, University of California Museum of Paleontology; UMMP, University of
Michigan Museum of Paleontology.
<C Fossil Lake
Samwel Cave
Potter Creek Cave
CALIFORNIA
200
X
M i les
Figure 1. Pleistocene localities of Dendragapus remains.
Identification of Dendragapus material. — Howard (1946: 179) has correctly empha-
sized that there is great variation in the bones of modern species of grouse, which may
lead to difficulty in identifying fossil material. Dendragapus bones are no exception,
and there are few characters by which individual elements of this genus can be dis-
tinguished from those of Pedioecetes, Tympanuchus, and Ccntrocercus. The humeri,
femora, tibiotarsi, and carpometacarpi of Dendragapus are more robust than those of
the other named genera. The ulnae are heavier and more strongly curved. The tarsometa-
tarsi are somewhat more distinct; the shaft averages wider, the anterior metatarsal groove
shallower and less elongated, and the trochleae heavier. The coracoids are similar to
those of the other genera, but the antero-ventral process of the furcular facet, in ven-
tral view, is thicker, blunter, and less recurved toward the shaft.
Races of modern Dendragapus obscurus are based largely on color variation (Fried-
man, 1946: 69), though there is an eight to ten per cent difference in wing lengths
1969 Jehl: Fossil Grouse of the Genus Dendragapus 167
between the largest and smallest races. Too few skeletons are available for the races to
be characterized osteologically. However, I can detect no racial differences in the
skeletons at hand. All show the robustness characteristic of the species, and there is no
demonstrable geographic variation in robustness.
Dendragapus From Fossil Lake
Shufeldt (1892) described two species of grouse from Fossil Lake, Oregon, lucasi
and nanus, which he assigned to the genus Pedioecetcs. Howard (1946) reassigned both
species to Dendragapus on the basis of tarsal characters. Other characters, namely the
stoutness of the carpometacarpi and the shape of the furcular facet, confirm her reallo-
cation. Shufeldt (1892) erected a new genus, Palaeotetrix, for a third species, gilli, but
this genus is not separable from Dendragapus (Jehl, 1967).
Dendragapus lucasi
Among the galliform birds present in the Fossil Lake deposits, this species was
second in abundance only to the Sage Grouse {Centroccrcus urophasianus; Howard,
1946). Dendragapus lucasi was somewhat smaller than modern D. obscurus. All ele-
ments, except one slightly shorter tarsometatarsus, fall within the size range and pro-
portions of female D. obscurus. There is marked sexual size dimorphism in grouse, with
males averaging larger than females. The apparent small size of lucasi probably is not an
artifact of sampling, however, as it is unlikely that a sample of 20 elements from at
least six individuals was derived entirely from female birds.
The ulna, coracoids, humerus, carpometacarpi, and femur of lucasi are indistin-
guishable from those of female obscurus. Howard (1946: 180) suggested two charac-
ters, "relatively less depth of proximal end and less development of the intermetacarpal
tuberosity," by which carpometacarpi of lucasi were thought to differ from those of
obscurus. However, she presented no quantitative data in support of the first character,
and I find that the great variation in the large series of obscurus at hand encompasses
that exhibited by elements assigned to lucasi. Furthermore, in obscurus the development
of the intermetacarpal tuberosity is perhaps the most highly variable feature of the
carpometacarpus; the development of this structure in lucasi is quite within the range
of variation exhibited in obscurus.
The tarsometatarsi of lucasi are also extremely similar to those of obscurus but
show several minor average differences: the metatarsal facet is smaller and more clearly
defined proximally; the area below the internal cotyla on the posterior face is less deeply
excavated; and the ridge bordering this depression, running between the internal cotyla
and the metatarsal facet, is less well defined. The trochleae average smaller than in
obscurus, but the symmetry of the distal end of the tarsometatarsus and the position
of the internal trochlea (cf. Howard, 1946: 180) are within the range of variation of
obscurus. Also, I detect no difference in the depth of the internal edge of the shaft, nor
in the depth and prominence of the hypotarsus (cf. Howard, 1946). The morpholep
of the hypotarsus is somewhat variable in obscurus and includes the variation exhibited
by lucasi.
In summary, Dendragapus lucasi may be characterized as a Pleistocene Dendragapus
differing from the modern D. obscurus only in over-all size and in minor characters of
the tarsometatarus.
Referral material. — Ulna-1 (AMNH no. 3476, Type). Carpometacarpal-7 [UCMP no. J1748, pre-
viously tentatively referred to D. nanus, see explanation below; AMNH nos. 347JA i-1. ;4"A
3478A (2)]. Tarsometatarsus — 8 fAMNH nos. 3475 (2), both previously referred to D. iijhih;
(4), 3476 (2)]. Coracoid — 2 [AMNH no. 3478A (2)]. Humerus -
(UCMP no. 31781). Tentatively referred. — Radius — I (UCMP no. 31776).
68 San Diego Society of Natural History Vol. 15
Dcmlragapus nanus
According to Shufeldt's (1892: 414) original description D. nanus differed from
D. lucasi "only in the matter of size," nanus being smaller. However, this distinction is
no longer valid. Howard (1946) reassigned four tarsometatarsi and two carpometacarpi
from nanus to lucasi, retaining only two tarsometatarsi (AMNH no. 3475) and, tenta-
tively, one carpometacarpus (UCMP no. 31748) in nanus. All of these elements are
within the size range of D. lucasi.
Howard (1946) considered the carpometacarpus of nanus to differ from those of
obscurus (and, by implication, from those of lucasi) in having "the external surface of
the proximal end . . . less rugose and the process of metacarpal I . . . more markedly
tilted upward." However, I could not confirm these differences when I compared the
carpometacarpus of nanus with a large series of obscurus. Tarsometatarsi of nanus were
said to resemble those of obscurus, and consequently to differ from those of lucasi, by
having a "short, prominently projecting hypotarsus." As noted above, variation in the
hypotarsus in obscurus is too great to make this character of diagnostic value. Howard
also stated (p. 180) that tarsi of nanus differed from those of obscurus by the "(1)
extreme proximal location of tubercle for tibialis anticus muscle; (2) more symmetri-
cally flared distal end; and (3) relatively more slender shaft." Again, variation in
obscurus includes the differences by which nanus was distinguished. In some obscurus
the position of the tubercle is as far proximal, and in one lucasi it is more proximal; the
symmetry of the distal end is variable and some tarsi of obscurus match those of nanus
exactly; the ratio of least lateral width of the shaft to total length of the tarsometatarsus
does not indicate that the element was slenderer in nanus than in obscurus (Table 4).
In summary, I find no characters except smaller size that allow elements previously
assigned to D. nanus to be differentiated from D. obscurus. Unfortunately, the sample
of nanus is too small to indicate whether the tarsal characters that seem to distinguish
lucasi from obscurus may also separate lucasi from nanus. However, these differences
are slight, and I believe, judging from the variation in tarsal characters both in obscurus
and in the California cave material discussed below, that the tarsal elements assigned to
nanus are within the range of variation of lucasi.
Because of the similarity of grouse elements from different species and the great
variation within species, it is impossible to disprove the occurrence of two small species
of Dcndragapus in the Fossil Lake beds. Nevertheless, the characters that have been used
to differentiate D. nanus do not allow it to be separated from either D. lucasi or females
of D. obscurus. I would, therefore, withhold specific recognition from nanus and con-
sider it synonymous with its contemporary, lucasi, with which it agrees in size and
proportions, rather than with the slightly larger and (apparently) more recent obscurus.
Dcmlragapus gilli
This large grouse, which approximated in size females of Centrocercus urophasianus,
is currently known only from two carpometacarpi (AMNH no. 3474, type; UMMP
no. 48223). D. gilli differs from obscurus and lucasi only in size and robustness, but in
these characters it is completely separable. As no carpometacarpus of lucasi or even of
obscurus males attains the great size and robustness that appears to characterize gilli,
the latter must be considered specifically distinct.
The California Caves Population
Remains of Pleistocene Dendraga[)us are abundant from the Samwel and Potter
Creek caves of Shasta County, California. Loye Miller, who first reported these fossils
(1911: 396-397), assigned most to D. obscurus, but suggested that more than one
1969
Jehl: Fossil Grouse of the Genus Dendragapus
169
species might be represented for many bones were of "unusual size" and "great robust-
ness" and differed by "slight detail of the head of the tarsometatarsus." Miller was
unable to analyze these fossils, because of a lack of modern skeletal material. (Nearly
60 years later complete skeletons of male obscurus are still in too short supply.) In
examining this material I detected only one form of Dendragapus; there were no demon-
strable differences between Samwel and Potter Creek cave specimens. However, this
form differed in body size and relative stoutness of the long bones from other Dendra-
gapus. A discussion of individual elements follows.
Carpometacarpus. — Except for greater robustness, carpometacarpi of the California
cave grouse are indistinguishable from those of lucasi or obscurus; they are as robust,
but not as long, as those of gilli (Table 1). The shape of the distal portion of meta-
carpal III is highly variable and does not, as might be inferred from Figure 2, offer a
valid means of distinguishing this form.
Referred material: UCMP nos. 5401, 5621, 8926, 8910a, 9621 (2), 9621a, 9743, 10043, 10160, 10183
(5), 27312 (2), 82977. Tentatively referred: UCMP nos. 27322, 27322a.
Table 1
Measurements of Carpometacarpus in Dendragapus (in mm)
Ratio, width metacar-
pal 11 to length
carpometacarpus
Width metacarpal II X 100
S.D. No. Range and Mean S.D. No. Range and Mean S.D.
Species
No.
Total length
Range and Mean
D. obscurus cf
7
38.8-42.5
(40.6)
1.6
5
4.1-4.3
(4.2)
5
10.1-10.7 (10.4)
.18
D. obscurus ?
20
34.3-39.4
(36.7)
1.2
20
3.3-3.9
(3.6)
.18
20
8.8-10.9 (99)
.43
Samwel and
Potter Creek
cave fossils
15
34.3-40.0
(36.2)
1.7
15
3.7-4.4
(4.0)
.20
15
10.5-11.8 (11.3)
.3 8
D. lucasi
7
34.3-39.3
(36.9)
1.5
7
3.4-4.0
(3.7)
.23
7
9.9-10.8 (10.2)
.3 0
D. gilli
2
43.8, 45.1
2
5.2, 5.2
2
1 1.5, 11.9
"D. nanus"
1
38.5
1
3.8
1
10.0
Ulna. — The ulnae average slightly longer than those of female obscurus and the
shafts average approximately five per cent wider (Table 2). Some shafts appear slightly
less curved than in obscurus.
Referred material: UCMP nos. 3922, 8910a, 8926a (2), 8789, 9528, 9554e, 9556, 9728, 9746, 273 12,
27313, 27322, 27322a, 31758. Tentatively referred: UCMP nos. 3049, 3608, 9554c (2), 9554J, 10119
(3), 27322a, 31737 (6).
Table 2
Measurements of Ulna in Dendragapus (in mm)
Proximal width
(tip of olecranon
to tip of external
Greatest distal
Species
Total length
cotyla )
u idth
No.
Range and Mean
S.D.
No.
Range and Mean S.D.
No.
9
Range and Mean
9.6-10.8 (10. 1)
S.D.
D. obscurus d 9
65.5-71.0 (67.6)
1.4
9
12.5-14.0 (13.5) .46
.39
D. obscurus ? 19
57.9-64.8 (61.6)
1.8
20
11.2-12.9 (12.2) .40
20
S.4-'>.4 (9.0)
.31
Samwel and
Potter Creek
cave fossils 9
59-67 (62.8)
2.4
1
11.8
10
8.8-10.1 (9.1)
.41
D. lucasi (type) 1
63.9
1
11.5
1
8.7
170
San Diego Society of Natural History
Vol. 15
-
s
5P
w
-I
M
<
<
<
h
w
o
o
H
z
w
w
a
a
<
* -I £
<S *■ <3
^- -*^ a/;
«^ »
^, r* ss
-J s m
55S
U
S "8
C/3
5
?
■a *a «.
o
"3 -~
Q
s
~3 <=> S
X
S-S p<
■-■2
■a.
t/3
K
ON
"o
-a
— O
oo oo
d ^
— ON
<N ON
-> d
K
K
ON K
fN •—
■*
■+
o u)
~ ON
•^ ,J- ~
-a
s u
NO
ON
O
o "J
— ON
— . ON
ON ON
•4"
1969 Jehl: Fossil Grouse of the Genus Demi raga pus 171
Humerus. — Most of the humeri are fragmentary and lengths can be obtained or
estimated for only five specimens; only one of these falls beyond the length of obscurus
females. The humeri appear to have slightly smaller heads and the shafts are slightly
straighter distally, averaging approximately eight per cent wider than in obscurm
(Table 3).
Referred material: UCMP nos. 4096, 4275, 4281, 5287, 5491, 5285, 6774, 6921, 6961, 6962, 8053, 8324,
8910, 9514, 9554a, 9646, 9712, 10147, 10160, 10183, 27312 (2), 27314, 27315 (2), 27342 (3), 27325,
31722.
Table 3
Measurements of Humerus in Dendragapus (in mm)
Greatest distal
Species
Total length
width
No.
Range anil Mean
S.D.
No.
Range and Mean
S.D.
D. obscurus cT
14
69.7-75.0 (72.4)
2.1
14
14.4-15.7 (15.8)
.59
D. obscurus ?
24
61.8-69.5 (65.1)
1.9
25
12.6-14.6 (13.6)
.48
Samwel and Potter
Creek cave fossils
5
62.8, 63.5, 64*,
68*, 72.3
7
11.4-14.0 (13.5)
D. lit cast
I
66.6
1
13.6
"estimated lengths
Tarsometatarsus. — Tarsometatarsi are similar to those of lucasi and of obscurus
females, but the shafts are noticeably wider (Table 4, Fig. 2). In all other respects
including the shape of the head (cf. Miller, 1911: 397), they are within the range of
variation of obscurus. Compared with lucasi, the trochleae are much more robust, the
metatarsal facet is less well defined, the depression below the internal cotyla on the pos-
terior face is deeper, and the ridge bordering this depression is more pronounced.
Referred material: UCMP nos. 3580, 4624, 579S, 8505, 8923a, 9554, 9554a, 9634, 9644, 9674, 9729.
9740, 9784, 9790, 9803, 9828, 10183a, 27313, 27323, 27324a, 27324b, 27325a, 31718, 31722, 37662,
37663, 37664, 37665, }7666, }7667, 37668. Tentatively referred: UCMP nos. 5501, 7430, 10183.
Tibiotarsus. — The fragmentary nature of most of the tibiotarsi makes detailed
analysis impossible. Two nearly complete elements (length estimated) are longer than
those of female D. obscurus. Shaft widths are approximately five to ten per cent greater
than in obscurus elements of comparable length; the distal width is also greater
(Table 5).
Referred material: UCMP nos. 8873, 8926a, 9554a, 27322, 27323, 27323a, 27323b, 31716, 31722, 31757.
Table 5
Measurements of Tibiotarsus in Dendragapus (in mm)
Lateral width
Species
Total length
across trochleae
No.
Range and Mean
S.D.
No.
Range and Mean
S.D.
D. obscurus cf
12
89.0-97.8 (92.8)
2.6
12
9.7-10.9 (10.0)
a:
D. obscurus ¥
19
78.4-89.2 (84.3)
2.9
17
8.7-10.0 (9.3)
.45
Samwel and Potter
Creek cave fossils
5
81.5*, 83.5*, 84*,
91.5*, 91.5*
4
9.9-1D/' ( Ki.4)
"estimated lengths
Femur. — None of the five femora (UCMP nos. 9554a, 9607, 9710, 9714, 27314)
examined are complete, but it is clear that all are within the si?e range of female
172
San Diego Society of Natural History
Vol. 15
obscurus; shafts average approximately ten per cent wider than in similarly sized
obsciirus (Table 6).
Table 6
Measurements of Femur in Dendragapns (in mm)
Greatest distal
Species
Total length
■width
No.
Range and Mean
S.D.
No.
Range and Mean
S.D.
D. obscurus <$
12
73.8-79.4 (75.6)
2.1
12
13.7-15.5 (14.2)
.65
D. obscurus +
22
65.1-71.3 (67.5)
1.9
22
11.6-13.0 (12.3)
.51
Samwel and Potter
Creek cave fossils
2
65.5*, 69.0*
1
12.0
D. lucasi
1
68*
'"estimated lengths
Coracoid. — As far as can be determined, the coracoids fall within the length
range for female obscurus. Some are wider, but there is not enough material for critical
comparison. I have found no method of distinguishing these coracoids from those of
lucasi or obscurus.
Referred material: UCMP nos. 3180, 5836, 7042, 7428, 8926a, 9674, 10160, 31713, 31722, 31759.
Table 7
Measurements of Coracoid in Dendragapns (in mm)
Length from head to
Species
tip of sternal facet
No.
Range and Mean
S.D.
D. obscurus cf
13
53.6-59.0 (56.9)
2.5
D. obscurus ?
26
47.4-54.8 (50.2)
1.9
Samwel and Potter
Creek cave fossils
4
48.5*, 49.6, 50.9*,
5 1.5*
D. lucasi
2
52.7, 52.9
"estimated lengths
Other elements: Two cervical vertebrae (UCMP nos. 9521, 10183), three frag-
ments of synsacrum (UCMP nos. 963 3, 9674, 9696) and one furculum (UCMP no.
8926a) are similar to those of obscurus and are probably referable to this population.
Discussion. — The California cave population was similar to D. lucasi in size but
differed in greater stoutness of the long bones. Smaller than D. gilli, it closely resembled
that form in robustness. Compared with the extant D. obscurus, the cave grouse was
slightly smaller, nearly all elements falling within the size range of obscurus females,
but the long bones averaged five to ten per cent more robust.
In interpreting these cave deposits the possibility of biased sampling must be con-
sidered. If predators responsible for amassing these remains were unable to carry large
grouse, the resulting deposits would be composed largely of the much smaller females
and much of the variation in the fossil population would be obscured. However, many
of the bird and mammal predators known from these deposits (Miller, 1911; Furlong,
1906) could have carried larger birds with ease. I conclude that the differences ascribed
to the cave remains are valid and, therefore, that this population is worthy of taxonomic
recognition.
The affinities of this population are not obvious. Although some individual bones
cannot be distinguished from those of obscurus, the California fossil population averaged
much stouter than obscurus. I detect no geographic variation in robustness in races of
1969
Jehl: Fossil Grouse 01 the Genus Dendragapus
173
obscurus, and in other grouse that I have examined robustness of the long bones appears
to be a conservative character. For these reasons I infer that the cave grouse probably
was most closely related to the similarly robust though much larger D. gilli. Even if
this inference is correct, the nature of the relationship between these forms can only be
surmised, owing to a lack of gilli material. The cave form could have been specifically
distinct, but it is hard to believe that three species of Dendragapus occupied so small an
area of the west only a few thousand years before the first appearance of obscurus. It
seems most reasonable to treat this population as a smaller, probably contemporaneous,
southern subspecies of gilli. I name this population Dendragapus gilli milleri in honor of
Dr. Loye Holmes Miller, who first called attention to these fossils. The Oregon popula-
tion thus becomes the nominate race, D. g. gilli.
Figure 2. Left to right, carpometacarpi of Dendragapus lucasi (AMNH no. 3478 A), Dendragapus
obscurus (UMMZ 74758), Dendragapus gilli milleri (UCMP no. 82977); tarsometatarsi cf
Dendragapus lucasi (AMNH no. 3475 A) Dendragapus obscurus (UMMZ no. 208192), Den-
dragapus gilli milleri (UCMP no. 9823). All illustrations X 1.
Dendragapus gilli milleri new subspecies
Holotype. — Left carpometacarpus, lacking metacarpal III and exhibiting slight wear
on distal end. University of California Museum of Paleontology no. 82977 (Fig. 2).
Age and locality. — Late Pleistocene deposits of Samwel Cave, Shasta County, California.
Diagnosis of holotype. — Similar in robustness to carpometacarpi of D. g. gilli but
approximately 18 per cent shorter; similar in size to carpometacarpi of D. lucasi and
D. obscurus but 10 per cent more robust.
Measurement of holotype. — Total length 38.7 mm; width of metacarpal II at mid-
point 4.3 mm; depth of proximal end through process of metacarpal I and internal
crest of trochlea 12.4 mm; ratio, width of metacarpal II at midpoint to total length
1 1.2 per cent.
Referred material. — As discussed above (pp. 169-172).
Synthesis
In the late Pleistocene two species of Dendragapus inhabited a small area of the
western United States. Dendragapus lucasi and D. g. gilli occurred sympatrically in
southern Oregon; a smaller subspecies of gilli, D. g. milleri, occurred in northern Cali-
fornia. The relationships of these Pleistocene grouse to the extant D. obscurus are not
clear. If robustness of the long bones is a conservative character, as I have suggested,
lucasi would appear to be the most likely ancestor for obscurus. D. gilli may have be-
174 San Diego Society of Natural History Vol. 15
come extinct at the end of the Pleistocene, leaving no descendants, but this is far from
certain. If "Dendragapus obscurus" can be shown to comprise two species (see Fried-
mann, 1946: 68), gilli may have been ancestral to one, lucasi to the other. In that
event, the problems associated with matching ancestral and descendant populations will
be extremely difficult. Until more information is available regarding the distribution
and variation in the fossil populations, evolutionary rates, the significance of the dif-
ferences that characterize the several populations, the temporal relationships of the
Pleistocene forms, as well as taxonomic relationships within D. obscurus, further specu-
lation on relationship in this genus is best postponed.
Acknowledgments
The late R. A. Stirton, University of California Museum of Paleontology, kindly made the Samwel
Cave and Potter Creek Cave material available to me. E. H. Colbert, American Museum of Natural His-
tory, loaned specimens of D. Iiicusi and "D. iiaiitts," and allowed me to examine type material in his
care. Skeletons of D. obscurus were obtained through: N. K. Johnson and the late A. H. Miller, University
of California; O. M. Buchanan and T. R. Howell, University of California at Los Angeles; R. F. John-
ston, the University of Kansas; K. E. Stager, Los Angeles County Museum; and H. G. Lumsden. I am
indebted to C. W. Hibbard for advice and assistance throughout this study, to H. B. Tordoff, R. W.
Storer, and R. M. Mengel for commenting on an earlier draft of the manuscript, and especially to Hilde-
garde Howard for her careful and helpful editorial assistance.
Literature Cited
Allison, I. S.
1966. Fossil Lake, Oregon, its geology and fossil faunas. Oregon State University Press. 48 p.
American Ornithologists' Union
1957. Check-list of North American Birds, 5th ed. Lord Baltimore Press, Baltimore, Md.
Brodkorb, P.
1964. Catalogue of fossil birds: Part 2 ( Anseriformes through Galliformes) . Bull. Florida State
Mus. 8(3): 195-335.
Friedmann, H.
1946. The Birds of North and Middle America. U.S. Natl. Mus. Bull. 50, Part X. U.S. Govt.
Printing Office, Washington, D.C.
Furlong, E. L.
1906. The exploration of Samwel Cave. Amer. J. Sci. 22: 235-247.
Howard, H.
1946. A review of the Pleistocene birds of Fossil Lake, Oregon. Carnegie Inst. Wash. Publ. 5 5 1.
pp. 141-195.
Jehl, J. R., Jr.
1967. Birds from Fossil Lake, Oregon. Condor 69: 24-27.
Miilllr, L. H.
1911. Avifauna of the Pleistocene cave deposits of California. Univ. Calif. Publ. Geol. 6: 385-400.
1960. Some Indian midden birds from the Puget Sound area. Wilson Bull. 72: 392-397.
1963. Birds and Indians in the west. Bull. S. Calif. Acad. Sci. 62: 178-191.
Peters, J. L.
1934. Check-list of birds of the world. Vol. 2. Cambridge, Harvard University Press, 401 p.
Short, L. L., Jr.
1967. A review of the genera of grouse (Aves, Tetraoninae) . Amer. Mus. Novitates no. 2289, 39 p.
Shufeldt, R. W.
1892. A study of the fossil avifauna of the Equus beds of the Oregon Desert. J. Acad. Nat. Sci.
Philadelphia 9: 389-425.
Wetmore, A.
1962. Notes on fossil and subfossil birds. Smiths. Misc. Coll. 145(2): 1-17.
Accepted for Publication 13 January 1969
San Diego Natural History Museum, P. O. Box 1390, San Diego, California 92112.
A BIOLOGICAL SURVEY OF BAHIA DE LOS
ANGELES GULF OF CALIFORNIA, MEXICO, IV.
BENTHIC AMPHIPODA (CRUSTACEA)
J. LAURENS BARNARD
MUS. COMP. ZOOU
LIBRARY
JUL 10 1969
HARVARD
UNIVERSITY*
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 13, 27 JUNE 1969
The series on Bahia de los Angeles has been published, in part, through the generosity
of Richard F. Dwyer. Much of the research on which it is based was conducted at the
Vermilion Sea Field Station of the San Diego Natural History Museum, which is
supported by the National Science Foundation.
A BIOLOGICAL SURVEY OF BAHIA DE LOS ANGELES,
GULF OF CALIFORNIA, MEXICO. IV.
BENTHIC AMPHIPODA (CRUSTACEA).
J. Laurens Barnard
Abstract. — The gammaridean fauna of a bay in the disjunct, warm-temperate zone of the Gulf
of California is discussed. Of the 70 species encountered, 13 represent new species and 5 new sub-
species. Two new genera are proposed. In 1962 and 1963 a sharp decline in diversity and abundance
occurred in autumn in depths between 6 and 46 m. The diversity of Gammaridea is about half
that of similar depths in southern California. Approximately 70 per cent of the amphipods have
warm-temperate affinities; at least 9 cognates of Californian species occur in BLA. Shallow, sandy
bottoms, in 2-8 m depth, are dominated by Ericthonius brasiliensis and Podocerus fulatius; mid-
depths of 7-36 m on fine sands by Ampelisca compressa, A. cristata and Platyischnopus metagracilh;
and the deepest zone, 37-46 m on brown silts, by Hetcrophoxus oculatus, Ampelisca cristata and
A. compressa. None of the dominant species is endemic, but in the Californian warm-temperate
zone Podocerus fulanus is confined to lagoons.
Resumen. — Se estudian los Anfipodos Gammaridos de una bahia situada en la zona limitrofe de la
region calido-templada del Golfo de California. Se describen setenta especies, de las cuales 13 son
especies nuevas y ademas 5 nuevas subespecies. La bahia esta bajo la influencia del conocida fenomeno
del efecto del verano en la densidad faunistica, de modo que esta es pobre en relacion con la que
presenta el mar abierto. Un 70% de los Anfipodos, aproximadamente, presentan afinidades con los
de las regiones calido-templadas; por lo menos 9 especies son cognatos a las que aparecen en regiones
calido-templadas; todo lo cual indica en cierto modo, el aislamiento de esta fauna de sus fuentes
originales.
Las regiones poco profundas (de 2 a 8 metros de profundidad) con fondos arenosos, estan
dominadas por Ericthonius brasiliensis y Podocerus fulanus; las zonas de profundidad media (de 7 a
3 6 m) sobre arena fina, aparecen pobladas por Ampelisca cristata, A. compressa y Platyiscbnopiis
metagracilh; mientras que las zonas profundas (de 36 a 46 m) con fondos de fango marron, estan
habitadas por Hetcrophoxus oculatus, Ampelisca cristata y A. compressa. Ninguna de las especies
dominantcs refleja una dislocacion de la fauna al hacerse endemicas; a excepcion de Podocerus
fulanus que puede funcionar en mar abierto como una reliquia confinada a las pequefias lagunas
litorales.
This paper describes the gammaridean fauna of a small bay, Bahia de los Angeles,
on the east shore of Baja California, in the northwestern part of the Gulf of California.
Amphipoda from the gulf have been recorded previously by Shoemaker (1925). Bahia
de los Angeles is presumably within the disjunct, warm-temperate, upper gulf region
discussed by Beltran (1960) in the "Symposium: The biogeography of Baja California
and Adjacent Seas." Collection of this fauna was undertaken as part of a survey to
assess the simple mud and sand-bottom communities and the general environment
of that bay. Barnard and Grady (1968) have outlined the thermal regime, water char-
acteristics, and benthic sedimentary structure of the bay and have reported on the
phenomenon of "summer-kill" as a dominating influence on the warm-temperate fauna
in 1962 and 1963.
The amphipodan fauna of Bahia de los Angeles is impoverished, in terms of both
species and abundance of individuals, in comparison with southern California, the only
other quantatively studied region of the eastern Pacific Ocean.
Approximately 70 per cent of the Amphipoda have warm-temperate affinities, and
at least 9 cognates of warm-temperate species occur in Bahia de los Angeles (BLA).
San Diego Soc. Nat. Hist., Trans. 15 (13): 175-228, 27 June, 1969
176 San Diego Society of Natural History Vol. 15
The fauna lacks numerous species that overlap this latitude in the open part of the
Pacific Ocean. The diversity of Amphipoda is appproximately half that of similar
depths in southern California.
The fauna, which is seriously depleted during high temperatures of late summer
(2 5 C), may also be affected by redtide (Barnard and Grady, 1968). Repopulation
of the bay during winter thermal minima (ca. 15 C) apparently occurs rapidly from
depths exceeding 5 0 m; post- winter samples contain a dense fauna of polychaetes,
Amphipoda, and small mollusks, especially clams. Windrows of dead clam shells, in
adult stages, occur on sand beaches in the southwestern part of the bay. These suggest
that mass mortalities in summer occur sporadically rather than annually.
The Amphipod Fauna
Seventy species of gammaridean Amphipoda, including 2 new genera, 13 new
species and 5 new subspecies, have been identified from Bahia de los Angeles. Nearly 20
additional species are recorded in fragmented and juvenile materials (Table 1). Twenty-
seven of the 70 identified species dwell primarily on soft bottoms, 3 3 inhabit hard
bottoms and 10 occur in both habitats.
In April, 1962, Amphipoda, as well as Mollusca, Polychaeta, and small Decapoda
were conspicuous in the samples. During the resurvey in October-November 1963 the
entire fauna was so poorly represented in the first few samples that we believed the
orange-peel grab was malfunctioning. Numerous adjustments were made; control sam-
ples taken with a Van-Veen grab confirmed the sparseness of the benthic fauna.
The diversity of soft-bottom (infaunal) species in Bahia de los Angeles is far less
than on the coastal shelf of southern California even though the diversity of sediments
is similar. In depths of 2-5 5 m in southern California 136 (infaunal) species have
been recorded (see Table 9 of Barnard, 1966, subtracting 29 deep-water species from
the total of 165). Nevertheless, the abundance of individuals (2000 per s.q m*') in April
at Bahia de los Angeles equals or slightly exceeds that found at various depths in south-
ern California, but it is very low compared with parts of Bahia de San Quintin (J. L.
Barnard, 1964b) where certain biotopes have more than 20,000 individuals /m2. In
Bahia de los Angeles the greatest abundance (45 69 individuals /m2) occurs in depths
of 2-6 m, whereas in depths between 7 and 46 m the abundance drops to between 75
and 208 individuals /m2 (Table 2).
A sharp decline in abundance of all but three of 20 dominant soft-bottom gam-
marideans occurred between April 1962 and November 1963 (Table 1, species marked
with "S" and having 104,- specimens in April samples). The decline in frequency is
generally of one order of magnitude but no appreciable decline occurred in Rudiletn-
boides stcnopropodits and a slight to moderate decline occurred in Heterophoxus oculatus
and Microdcutopus schmitti. Six species were completely absent from samples in Novem-
ber, 1963. Whether an autumnal decline (1962) and winter refurbishment (1963)
occurred are unknown. Four of the six species missing in November are eurybathic
(Barnard, 1966) and presumably could repopulate shallow water depths from deep water
sources. Autumnal impoverishment occurred in depths of 40-46 m as strongly as it did
in depths of 21-30 m as seen in the following tabulation in which 19 samples were
selected from each depth in each season, indicating total specimens of Gammaridea
collected in each category.
•computed from 8775 individuals occurring in 70 samples representing about 4.38 m~; see Table 1 for
April samples. The orange-peel grab collects an area of about 0.06 m-.
1969
Barnard: Bahia de los Angeles Amphipoda
177
w
<
i
u
V.
<
o
M-!
W
o
X
<
o
z
if
< J
<
II
O H
b
3^
D
H
H
<
J
><
0 oS
D
u w
h
£ w
w
c
2
D
u
u
0
<
w
Q
<
<
o
o
H
!j
<
o
H
W
to
<
X
c
J
W
^~
H
W
►J
W
g
H
><
W
H
H
W
w
X
H
Z
?
o
z
as
Z
5
< H
7.
0
H
H
0
H
i
H
h
O
t/i
O
H
a
w
v.
CO
H on
Of
« «
Q'S
< c
c.
<~
-a.
CO
■a.
5
<3
CO
bo a
S> 8 "W
^= f ^
Cu -.
c. _£ -C,
-all II
so
ON
o
<~
■a
CO
-a. — .
S »>
CO
CO _'
£«
H
O
03
H
O
CO
-a.
co
•5"=
CO
CM CM «\ OO m* 'J- O
**s 0\ t— i CM
M-. ~H *^ i— ■ r— OOr^, -*" rj
■* ON s-n SO (N
K "~> K
N « t N N x H
+
+ + + +
4-
+
o\ o
SO 0\ K
CM
CM
K Tt
O —
so o
7) m 71
SO ON OO
co
CM
SO o
O — I
CO CO CO
rr\ ^f »-»
o
oo
CM
CO
>-wwwiHWHHwwwhhwHl-H
_
H w w T->
~5
8
a
(3
C.
So
"*■»
<3
s
■»*
w
_2
C
o
M*
■-.
«m
-C
ms
o
<3
mi
CU
k.
•ex.
5
o
Cm
<3
-a
2
.c-
41
Cm
~c«
-5
Co
"S
-a,
-
Mi
"3
"3
0
ca
<3
"3
<3
Cm
w
■_
•m
Cg
Q
<m
cm
S
s
<-*
■v.
■^
"S,
•^~
k
^T
^3
a
■v.
k
Ml
«
^
"a
<3
«
<a
<s
a
-!
s
Cj
^*
Cm
C\)
-
<i
Sj
2
CJ
a
o
Cm
Cj
<o
c-
Co
V.
o
o
o
;
C
Cm
k.
/o
<*s
"
^t
--•
-«
-c
<^
^_
q
V
MO
-fc-
<.
•5
-a,
-V.
**
£
c-
-a.
5u
■a.
%*
«,
Cm
C«
■a.
■a.
-S.
-£.
-£.
-S_
«1
Jt
q
'm
s
Co
S
s
s
s
5
S
5
S
B
t
S
?
-—
q
-1
*
^
^
^
-d
^
^
'C
^
■<
^
"<
■<
^
cc
oq
oq
*
U
o
■a.
Cm a
■3
-a -a
■a. -a.
o o
o o
u -
< -
■a. *-
- ir
S -a
-
ua -
0]
178
San Diego Society of Natural History
Vol. 15
0
H
H
0,
CQ
Q
<
X
Co
CO
-a.
S
<s
to
6« ~
^S *» <3
v -a. «
-ail II
w
«
1—1
•a.
P
C/5
1-1
01
-Q
o
%>
— -* .*— -v
u
•a. —
O
oo
wo _r
H ^
H
O
pq
H
Po
O
CO
-a.
co
•a.0
S rs
to
5-2
4'|
+
+
0\ o
1H
f-H
^\
o
oo
fH o
o
(S
— <
CM
CM CM rg iH *->
^H i-H ^H ^ "^"
^* ,—i .—<*«"» ^-i
OO ^" C\ N i"
*-» cm K cm cm ^-> —
— <+
t+t
+ +
+ + +
+ +
r, — —
*■-» -— • CM
<S| — fS
H H H f-
CO CO
N N h
CO C/1
l-H OO
co co co co co
WHl-WHHH^-'WfHfHfHHHHl-'I-'Hf-Hf-HWl-W
^ & fr & & ^ ^ & > ^> & >
©
■3
to
5
o
_<o
~5
5 13
*3 •** <o
5 "«
© -c:
-a.
o
-a
•a,
V. S "*i
?», ©
■a, -a.
o ©
tS <3 o £
© -a, £
to
a S "3
* © b
"*! S S
■a, s »»
© £
l^ © to
^ ■a. "a
■*- -a. ?
to -tr <-,
a: a: a:
s
©
** R ■»*
'So
us
■s - s
«s
3
~5
©
5
60
t3
■a, <a,
© o
Co
■a.
? «
is
o
"3
-a
8
60
o
to
5 s
o
■a,
to
o
-a
o
-a
.Si, •£
kJ -J >-J
.^* s
t3 <3
60 60
Be
o
to
-^ -», \a ^i ^.
sssss^o
8 a a s
X X X H
© © © ©
•^ *Q *^ *^
<b, -a. <a, -a.
<S « <3 «
k. K K k.
tl t! 1 «
Oo Oh Co ft,
1969
Barnard: Bahia de los Angeles Ampiiipoda
179
>^\ f\ f>
1— ^" f*
—r <~ K ~ Tf
t(- «— • <*"»
-* ^H — r j rl
+
+
+
+
+
K
V^ r*1 V^ *">
o
i^. <N i— < o
t/5 1« l/llfl fl U)
-^- -r Ix -1-
on
i/i i/i i/i i/i tfl
"*-
OO f^i •— < •— i "4*
^ g* ^ S* ^^ ^ ^
■5 ~
<, ■*
2 -
^: -*-.
o .
■°< -ft< :H 'c:
S 2 = o
c2 cC &; ft^
■a
-ft. -ft. '
o
c
.2 >
c
3
O
t/>
9 «i 2 'S S
„ «
o 2 g -5 -5 s "«
?s ft S 5 £
6C £
ft. a, ft, ft. £
ft. k: ft<
=< CO tfl 5 P
-ft.
to
•-ft.
-.
D. <o « w
S Q. ^r ■ ■ *-<
~3 us -5,
— o ^
I =: -B T3 T3 <-
5" ^ = "S *s "S s
-. =: U - - J3 -
.i! Q.
O ...
■ -3 -3
'5 '3
^ o
2 C
CJ rs C
7 ^ ~
__--. ,, ft. ft. tO CO -— — '
180
San Diego Society of Natural History
Vol. 15
TABLE 2
Dominant Amphipoda taken in several depth classes during April, 1962, survey of mud bottoms
in Bahia de Los Angeles, indicating number of individuals per square meter.
No. samples
No. dominant species, total
Species, total
Individuals/m-, total
Acuminotleutopus periculosus
Ampclisca compressa
Ampelisca crhtata
Batea rectangnlata
Batea susurrator
Corophium baconi
Ericthonius brasiliensis
Gitanopsis pusilloides
Heterophoxus oculatus
Listriella mclanica lazaris
Microdeu/opus schmitti
Microjassa macrocoxa
Monoculodes hartmatiae
Orchomenc niagdalenensis
Photis brevipes
Pbotis californica
Platyischnopus mctagracilis
Podocerus fulanus
Kudilemboides stenopropodus
2-6
53
1390
106
Depth class, m
7-16 17-26
8
4
27-36
37-46
3
13
17
24
13
11
6
6
6
5
15
14
20
22
18
4569
75
208
109
173
106
5
7
7
17
14
36
28
21
32
5
28
58
2500
21
1
22
111
100
53
4
7
Depth
21-30 m
40-46 m
April, 1962
855
1104
November, 1963
97
128
Six species of Amphipoda dominate the benthos of BLA: Ericthonius brasiliensis,
Podocerus fulanus, Ampclisca cristafa, Platyischnopus mctagracilis, Ampclisca com-
pressa, and Heterophoxus oculatus.
The shallow sandy bottoms from about 2 to 8 m are dominated by Ericthonius
brasiliensis and Podocerus fulanus (Table 2). These species are tube-dwellers; presum-
ably they attach to hard particles and in patches of red algae occurring on those bot-
toms. Seven other species occurring in these depths (Table 2) are more abundant than
any species found in depths greater than 6 m. This density pattern differs from that
found off the coast of southern California, where Amphipoda are relatively sparse on
inshore sand bottoms but are abundant in depths exceeding 6 m (Barnard, 1963).
The 6 m depth-class in BLA and the 9 m depth-class in southern California are
quite different (Tables 2, 3; cf. Barnard, 1963: Table 16) In southern California,
Ericthonius brasiliensis is 29th in abundance. Its primary position in BLA may reflect:
1) the more abundant algae in the quieter waters that act to shelter masses of tubes
from disruption by surf action; and 2) the absence of competitors resulting from
extreme eurythermic conditions. Ericthonius brasiliensis is a eurythermic species that is
cosmopolitan in tropical, warm-temperate, and even in some cold-temperate areas (viz.
Norway; Puget Sound, Washington). Whereas Ericthonius brasiliensis is able to adjust
1969
Barnard: Bahia df los Angeles Amphipoda
1S1
to wide seasonal extremes in temperature, Podocerus cristatus of the open Pacific may
not endure short-term fluctuations even though it adjusts to wide extremes in incre-
ments by latitude. This possibility is suggested by the dominance of Podocerus fulanus
in Bahia de los Angeles and in lagoons of southern California wherein P. cristatus is
absent.
In the 16 m depth-class, the frequency of Amphipoda is lowest (7$ mL'). This
corresponds to southern California where standing-crop and density of organisms
are lowest seaward of the surf zone but increase sharply on softer bottoms of deeper
waters. The mid-depth zones, generally 7 to 36 m in BLA, are dominated by Ampelisca
cristata. Platyischnopus mctagracilis and Ampelisca compressa are also fairly common in
the 26 and 36 m depth zones, but the only other species that can be considered dominant
TABLE 3
Dominant Amphipoda taken in several depth classes between 1957 and 1961 on coastal shelf
of southern California, based on 161 samples proportionally divided according to respective
depth classes; collected by "Velero IV," University of Southern California; indicating number
of individuals per square meter. Each column includes only the first 20 dominant species
ranked by individuals per square meter in that depth class and a species is not necessarily
absent from depth classes lacking numbers. A phyletic or ecologic equivalent occurring in
Bahia de Los Angeles is noted. + marks those species also occurring in Bahia de Los Angeles.
Depth class, m
Species
4-9
10-18
19-37
Occurrence in BLA
AcuniinoJcutopHs heteruropus
Amphideutopus oculatus
Ampelisca brevisimulata
Ampelisca compressa
Ampelisca cristata
Ampelisca lobata
Am pit hoe sp.
Aoroides cohimbiae
Argissa hamatipes
Batea transversa
Eohaustorius washingtonianus
Ericthonius brasilicnsis
Eurystbeus thorn psoni
Heterophoxus oculatus
Ischyrocerus pelagops
Listriella goleta
Mandibulophoxus uncirostratus
Megaluropus longimerus
Monoculodes hartmanae
Paraphoxus abronius
Paraphoxus bicuspidatus
Paraphoxus epistomus
Paraphoxus fatigans
Paraphoxus heterocuspidatus
Paraphoxus obtusidens
Paraphoxus stenodes
Paraphoxus variatus
Photis brevipes
Photis lacia
Synchelidium shoemakeri
Synchelidium spp.
9.2
1.0
4.4
7.5
4.1
9.5
0.5
3.0
30
2.3
4.9
9.7
25
55
3.8
4.1
25
31
47
11
22
27
27
21
29
12
12
11
21
12
95
16
11
15
40
20
30
17
9.1
48
64
14
63
26
8.9
9.0
26
37
10
11
27
15
7.3
36
19
7.7
36
A. periculosm
+
A. mexicana
+
+
+
+
+
+
E. tonichi
+
Microjassa macrocoxa
L. m. lazaris
Platyischnopus mctagracilis
+
+
P. cognatus
+
P. gem mat u\
P. spinous
+
P. californica
Synchelidium ^pp.
Synchelidium spp.
182
San Diego Society of Natural History
Vol. 15
is the nestler Batea susurrator at 26 m (Table 2). The species of Ampelisca are tube
builders, but their soft tubes are anchored in the substrate and lie limply on the surface.
Like all members of the Phoxocephalidae, Platyischnopus is presumed to be a burrower
and in this capacity it represents a poor substitute for the dense populations of Para-
phoxus (16 species) found off southern California. In depths exceeding 36 meters the
species of Ampelisca and Platyischnopus maintain dominance but they are joined by
another burrower, Heterophoxus oculatus (Table 2), a eurybathic organism, which is
also prominent on coastal shelves of the open sea and on the coastal slope to depths of
almost 2000 m. Note that Platyischnopus is scarcely more abundant than Photis, but
it is more widespread and occurs in nearly every sample between 7 and 46 m, whereas
Photis occurs abundantly in only a few.
The inshore sands of California in depths of 4-9 m are dominated by Paraphoxus,
Synchelidium, Mandibulophoxus (a morphological parallel to the deeper-dwelling Platy-
ischnopus of BLA), and Eohaustorius; the abundance of these burrowing forms, rather
than of infaunal tube-dwelling species of Ampelisca, may reflect the strong water motion
and shifting substrate of the region. However, tubicolous genera, such as Photis, that
attach their tubes to structures projecting above sediment level, are as abundant as some
of the species of burrowers. There is some weak evidence (Barnard, 1958b) that tubi-
colous species inhabiting hard particles are far less obligated to remain in their tubes
than are the infaunal ampehscids and can move from place to place as their epibenthic
substrates are shifted. They appear to be able to occupy tubes of other organisms with-
out necessarily constructing their own. Even though the sampling efforts (comple-
mented by dredgings in BLA) have been comparable, the diversity of the shallow-water
BLA fauna is half that of southern California (23 vs 46 species; Table 1 and Barnard,
1963, Table 16), which suggests that the quiet-water environment' BLA is even harsher
than the shifting substrate of open-sea shallows.
Of the medium-depth fauna, three of the six common species of BLA are also
abundant in southern California: Ampelisca cristata, A. comprcssa, and Photis hrevipes.
The ecological counterparts in southern California of Platyischnopus and Monoculodes
TABLE 4
Substrate Distribution of 4 Species of Gammaridea in BLA
Species
Ampelisca compressa
Ampelisca cristata
Listriclla melanica lazaris
Platyischnopus metagracilis
Substrate
Silt
Sandy
silt
Silty sand
Sand
Silt
Sandy
silt
Silty sand
Sand
Silt
Sandy
silt
Silty sand
Sand
Silt
Sandy
silt
Silty sand
Sand
Percent
of
Specimens
28
40
23
9
22
3 8
27
13
6
12
20
62
28
29
25
18
Average
per
Sample
8.9
14.0
7.3
3.4
23.0
45.0
29.0
16.0
0.4
1.0
1.4
5.2
8.5
10.0
7.5
6.7
Number
per
Positive
Sample
19.0
15.0
12.0
4.6
24.0
45.0
34.0
15.0
1.6
2.4
2.1
6.5
8.5
10.0
9.3
8.2
1969 Barnard: Bahia de los Angeles Amphipoda is3
are probably Paraphoxus and Synchelidium but the mainland coastal shelf of California
has no obvious counterpart of Batea. Batea occurs abundantly only in the Channel
Islands of California and joins Platyischnopus in a few square miles of benthos off San
Diego. Two species of Synchelidium also occur in BLA but their occurrence is very
sparse.
Of the deep-water fauna, four of the nine BLA species are also prevalent in south-
ern California: Ampelisca cristata, A. compressa, Photis brevipes, and Heterophoxus
oculatits.
Of the 27 most abundant species of southern California in depths similar to those
sampled at BLA, only 6 species or their allopatric equivalents are absent from BLA;
most of these are members of Paraphoxus. However, many other southern California
species are also absent from BLA, especially notable being a host of ampeliscids (Table 9
of Barnard, 1966). Northern species having their southern limits in southern California
should be omitted from consideration. This may be determined in Barnard (1966,
Table 9).
The distributions of the 6 dominant species at BLA in April are presented in
Figures 1-6.
Impoverishment of the Fauna
The gammaridean fauna on soft-bottoms in BLA is far less diverse than the fauna
of the coastal shelf of southern California. The pertinent members (33) of the BLA
fauna are marked in Table 1 and the 102 members of the southern California fauna
have been extracted from Barnard (1966, Table 9) by eliminating deep water species,
rarely occurring members, and those known from the literature to inhabit hard bottoms
primarily. Gammaridean genera for the most part inhabit either soft or hard bottoms
but occasional genera such as Ampelisca and Paraphoxus have members in both habitats.
These differences have been taken into account in the two faunas under consideration.
The soft-bottom Gammaridea of BLA represent 2 3 genera whereas the Gammaridea
of warm-temperate California represent 5 8 genera.
The two most diverse benthic genera in midlatitudes are Am pel/sea and Paraphoxus.
In BLA Ampelisca has 3 species on soft bottoms but in southern California it has 8.
In BLA Paraphoxus has 4 species on soft bottoms but in southern California it has 16.
Only 5 soft-bottom species of BLA (3 of Batca and 2 of Platyischnopus) have no
counterparts in the dominant fauna of soft-bottoms of mainland southern California.
Seventy-four species of the California fauna have not been found on soft-bottoms
of BLA, although 5 have been found sparsely on hard-bottoms. The remaining 69
species of southern California are not necessarily stenothermic as might be expected
from their absence in BLA. Thirty-seven of those species have depth ranges (by arbitrary
classes) from 18 to 183 m and 12 others have ranges between 18 and 5 5 m (Barnard,
1966, Table 9). They are therefore potentially eurythermic but their absence in BLA
may reflect the inability of a deme to undergo wide thermal changes annually.
There are some striking omissions from the BLA fauna of eurylatitudinal and
tropical species, for example: (1) Paraphoxus floridanus, a tropical pan-American
species, with northern limits at the Mexico-California border; (2) Paraphoxus obtusi-
dens, known from the Kurile Islands to Colombia, South America; (3) Ampelisca vene-
tiensis, known from southern California to Ecuador; (4) Ampelisca cristoides, known
from the open Gulf of California even as close as Angel de la Guarda Island; (5)
Ampelisca pacifica, California to Panama, perhaps submerging deeply in tropical waters;
(6) Ampelisca hreiisimulata, from California to Panama, in tropical depths of 9 m;
184
San Diego Society of Natural History
Vol. 15
Ampelisca compressa
Figure 1. Distribution of Ampelisca compressa Holmes.
Ampelisca cristata
Figure 2. Distribution of Ampelisca cristata Holmes.
1969
Barnard: Bahia de los Angeles Amphipoda
185
Figure 3. Distribution of Hetcrophoxus oculatus Holmes.
Figure 4. Distribution of Lhtriella melanica liizarh Barnard.
186
San Diego Society of Natural History
Vol. 15
Monoculodes hortmanae
Figure 5. Distribution of Monoculodes bartmanae Barnard.
Platyischnopus metagrocilis
Figure 6. Distribution of Platyischnopus met a gracilis Barnard.
1969 Barnard: Bahia ni los Angeles Amimiipoda
187
(7) Ampelisca kancocki, collected by the "Velero 111" near BLA in depths slightly
exceeding those of this survey; (8) Ampelisca indentata, from California to Gulf of
California; (9) Ampelisca pugetica, Washington to Angel dc la Guarda Island, on shore
at the latter; (10) Byblis veleronis, California to Gulf of California (Espiritu Santo
Island, 38 m) ; (11) Cerapus tubularis, a curylatitudinal cosmopolite; (12,13) Cheiri-
photis magacheles and Chevalia aviculae, both tropical cosmopolites, occurring in south-
ern California; (14) Mandibuphoxus uncirostratus , apparently a tropical [ndo-Pacific
species found as far north as southern California; (15) Photis elephantis, known from
southern California and with records at hand from the Galapagos Islands. Other omis-
sions from the BLA fauna include Anamixh linsleyi, Batea lobata, Eurysthetis spinosus,
Leucothoides pacifica, Metaceradocus occidentalis, Metaphoxus frequens, M. fultoni,
Parajassa angularh, and Pontogeneia minuta, all but Metaphoxus spp. probably occur
on hard bottoms, which have not been completely explored.
Affinities of the Fauna
Nine species and subspecies of BLA Gammaridea are cognates of forms with warm-
temperate affinities; 20 species occur only in warm- and cold-temperate regions; 19
species occur in or have affinities with tropical waters; 2 species have unknown affini-
ties; and the remaining 18 have eurylatitudinal distributions (Table 1). Batea transversa
coyoa may also have direct warm-temperate affinities, although the genus Batea is mainly
tropical in distribution. Acuminodeutopus periculosus is assumed to be a warm-temper-
ate species, because to date the genus has been found only in warm-temperate regions.
Of the unidentified species only Pleustes sp. has definite warm-temperate affinities. This
is the southernmost record of the genus.
More than 70 per cent of the Gammaridea in BLA have non-tropical affinities, but
if the eurylatitudinal species with unknown affinities are subtracted the percentage drops
to 61 (30 of 49 species). Soule (1960) found that 5 7 per cent of the Bryozoa occur-
ring in the northern third of the Gulf of California have non-tropical affinities. On the
other hand, Garth (1960) found that only 8 per cent of the Brachyura have northern
affinities, but his data included the entire Gulf.
Of the dominant Gammaridea with known affinities of soft-bottoms in BLA, 72
per cent (18 25) have affinities with warm regions; three species are tropical, 15 warm-
temperate, and 7 eurylatitudinal.
Systematic Procedure
Classification follows that of Barnard (1958a). Distributional information excludes
new records in BLA. Locality data occur in Barnard and Grady (1968). Materials are
summarized in Table 1. For a complete Station list of specimens order NAPS Document
00308 from ASIS National Auxiliary Publications Service, c o CCM Information
Sciences, Inc., 22 West 34th Street, New York, N. Y., 10001; remitting S1.00 for each
microfiche copy and $3.00 for each hard-copy reproduction. Samples 130 and 2 10,
represented by aliquots, have their data expanded by factors of 100 and 10 respectively.
Diverse substrate habitats of 4 species are summarized in Table 4. Sideheads of "Rec-
ords" include summary of locality data, mainly depth; of "Relationships" include, with
few exceptions, morphological differences from the nearest relative only, and occasional
comments on classificatory problems arising from this study. Diagnoses .\nc\ descriptions
are condensed but they are widely supplemented by the figures. Decisions on categorical
rank of new species and subspecies have no relationship to a numerical standard of
differential characters but are based on the differences of known species in each s^cnus.
Body sizes, quoted in mm refer to body length. Derivations of new names are found in
the appendix.
188 San Diego Society of Natural History Vol. 15
ORDER AMPHIPODA
Suborder Gammaridea
Family Ampeliscidae
Ampclisca cotnprcssa Holmes
Figure 7 c
Ampelhca comprcssa Holmes. Barnard, 1960a: 31-32; 1964a: 213.
Antenna 1 of female reaching only to end of article 4 of antenna 2, flagellum with 5 articles, in
contrast to populations of the open sea off southern California having antenna 1 reaching almost to the
end of article 5 of the second antennal peduncle. Article 2 of peduncle of antenna 1 only 1.5 times
as long as article 1, whereas in open sea specimens the article is twice as long as article 1. The entire
first antenna is stunted in comparison to specimens from southern California. The head is more fully
excavate below, appearing more narrow in the specimens from Bahia de Los Angeles.
Catalogued specimen. — USNM No. Ill 508, female, 6.7 mm, BLA 88.
Records. — Soft-bottoms, 15-46 m; in April, 152 individuals /m2; in October with 9.0 individuals /m2.
Distribution. — Western Atlantic Ocean; Caribbean Sea; eastern Pacific Ocean from Panama to Puget
Sound, Washington, 1-266 m.
Ampelhca crhtata Holmes
Figure 7 a, b
Ampelhca crhtata Holmes. Barnard, 1954a: 26-28, pis. 17, 18: 1964a: 213.
Female adults have article 2 of antenna about 1.9 times as long as article 1, in contrast to popula-
tions of the open sea off southern California having article 2 about 1.2 times as long as article 1.
Juveniles have the open sea form of antenna 1 and subadults range from 1.4 to 1.7 in the relative length
of article 2. The population in Bahia de Los Angeles is entirely of the microdentate form, having a
minute tooth on pleonal epimeron 3. Uropod 3 has the outer ramus broader and more rounded apically
than in specimens from southern California.
Records. — Soft-bottoms, 15-46 m; in April 527 individuals /m2; in October with 43 individuals /m2.
Distribution. — Caribbean Sea; eastern Pacfic Ocean from Ecuador to Tomales Bay, California, 6-152 m.
Ampelhca lobata Holmes
Ampelhca lobata Holmes. Barnard, 1954a: 11-14, pis. 5, 6; 1964a: 214.
Records. — Rocky intertidal on outer islands and at Vermilion Sea Field Station.
Distribution. — Caribbean Sea; eastern Pacific Ocean from Ecuador to Monterey Bay, California, 0-183 m.
Ampelhca mexicana Barnard
Ampelhca mexicana Barnard, 1954a: 45-46, pis. 31, 32; 1954b; 7; 1964a: 214.
Records. — Deep water zone of the bay, 2 5-46 m, primarily on silt-bottom but also on silty sand and
fine sand.
Distribution. — Caribbean Sea at Bahia Caledonia, Panama; eastern Pacific Ocean from Peru to Punta
Canoas, Baja California, 9-73 m.
Ampelhca milleri Barnard
Ampelhca milleri Barnard, 1954a: 9-11, pis. 3, 4; 1964a; 215.
Material. — BLA SIO-X, one juvenile, 2.5 mm, fitting the description except for small differences; adults
should be collected for better comparison.
Record. — Piedras Ahogadas, 24 m, shell fragment bottom.
Distribution. — Eastern Pacific Ocean from Ecuador and the Galapagos Islands to San Francisco Bay,
California, 15-187 m; in shallow water at San Francisco.
Ampelisca scbellcnbergi Shoemaker
Ampelhca scbellcnbergi Shoemaker, 1933a: 3-5, fig. 2; 1942: 9. Barnard, 1954a: 14-16, pis. 7, 8; 1954b: 2.
Record. — Rocky intertidal of Vermilion Sea Field Station.
Distribution. — Caribbean Sea and Gulf of Mexico; eastern Pacific Ocean from Peru to Laguna Beach,
California, 0-46 m.
Family Amphilochidae
Amphilochus neapolitanus Delia Valle
Amphilochus neapolitanus Delia Valle. Chevreux and Fage, 1925: 112-113, figs. 106-108. Barnard, 1962c:
126, fig. 3; 1964a:217; 1964b:105.
Record. — Sandspit north of village in tailings of fish sample.
Distribution. — Cosmopolitan in tropical and subtropical seas, 0-80 m.
Gitauopsis pusilloides Shoemaker
Gitanopsis pusilloides Shoemaker, 1942:9-11, fig. 3.
The unusual maxillipedal palp having 2 clavate spines on article 3 at the base of the finger is shown
in the drawings.
Records. — Intertidal to 9 m on coarse substrate.
Distribution. — Bahia Magdalena, Baja California, possibly north to La Jolla, California; intertidal to
about 20 m.
1969
Barnard: Bahia de los Angeles Amphipoda
189
Figure 7. Ampelisca crhtata Holmes, female, 7.0 mm, BLA 41: a, head: b, uropod 3. Ampelisca compressa
Holmes, female, 6.7 mm, BLA 88: c, head. Batca transversa coyoa, new subspecies, female, 6.9 mm, BLA
SIO-62-236: d, e, f, pereopods 3, 4, 5; g, gnathopod 2. Gitanopsis pusilloides Shoemaker, female, ovig-
erous, 1.8 mm, BLA 213: h, i, j, maxillipcd and views of palp terminus, the latter with medial setae
removed to show fused cusp-spines. Hippomedon propinquus Sars, female. 4.5 mm, BLA 103: k, 1, gnath-
opod 2; m, gnathopod 1; n, antenna 1. Am pit line ramondi Audouin, male, 6.3 mm, BLA SIO-62-2I0:
o, p, gnathopods 1, 2.
190 San Diego Society of Natural History Vol. 15
Family Ampithoidae
Ampithoc plumuiosa Shoemaker
Ampithoc plumulosa Shoemaker, 1938: 16-19, fig. 1; 1942: 39. Barnard, 1959a: 37; 1964b: 111; 1965:
20, figs. 11, 12.
Only juveniles and subadult females have been collected in BLA.
Record. — Intertidal at Vermilion Sea Field Station.
Distribution. — - British Columbia to Ecuador, intertidal and shallow subtidal waters.
Ampithoc pollcx Kunkel
Figure 8
Ampithoe pollex Kunkel, 1910: 92-94, fig. 36. Barnard, 1954c: 29-31, pis. 27, 28; 1959a: 37; 1964b,
111; 1965: 22-25, figs. 13, 14. Mills, 1964: 15-16.
Grubia indent at a Stout, 1913: 656-657.
Three adult forms of this species occur in BLA. They may represent individuals hatched in different
seasons.
Typical form, material: BLA 305 (3), SIO-62-216 (8): male gnathopod 2 is fully developed and
has the dactyl and thumb meeting; specimens fit the descriptions given by Barnard (1965); maximum
length about 5.5 mm.
Small form with aberrant gnathopod 2, material: SIO-62-215 (8); gnathopod 2 of the male has a
shortened, thicker, more curved dactyl than in the typical form; the dactyl fails to meet the thumb;
coxa 1 has a more excavate anterior margin than in the typical form; maximum length about 5.5 mm.
Large form with retarded gnathopod 2, material: BLA 305 (144), 306 (193); male gnathopod 2 is
typical of juveniles although the body length exceeds 7.0 mm; the palmar tooth is poorly developed;
specimens preserved in alcohol are more strongly pigmented than the other forms.
Material of juveniles and females not associated witli males and thus unrecognizable as to form:
BLA 214 (1), 306A (228), SIO-62-212 (3), 62-228 (3).
Records. — Intertidal on rocky shores.
Distribution. — Bermuda; eastern Pacific Ocean from Bahia de San Quintin, Baja California to Coos Bay,
Oregon, intertidal and shallow sublittoral waters.
Ampithoc ramondi Audouin
Figure 7 o, p
Ampithoc ramondi Audouin. Shoemaker, 1942: 40. Barnard 1955a: 28-29 (with references); 1964a: 217;
1965: 25-27, figs. 15, 16.
Amphithoc vaillanti Lucas. Chevreux and Fage, 1925: 33-334, figs. 341, 242. Gurjanova, 1951: 880-881,
fig. 614.
Ampithoe divhura Shoemaker, 1933b: 255-256, fig. 8.
Amphithoe intermedia Walker, 1904; 290-291, pi. 7, fig. 46. Chevreux, 1908: 515-516, fig. 29.
The references to this species have been condensed from a total of 5 5 or more published since 1900.
Male gnathopod 1 has a strongly excavate palm like that of male gnathopod 2. This is atypical of
A. ramondi as found on the outer coast of Baja California and may indicate subspeciation.
Records. — Intertidal on rocky shores.
Distribution. — Tropicopolitan, phycophilous, intertidal; northern record in eastern Pacific Ocean at Bahia
de San Ramon, Baja California.
Ampithoc tea Barnard
Ampithoe tea Barnard, 1965: 30-34, figs. 19-21.
The second antennal flagellum in both sexes is stouter than in the type-series. It resembles antenna 2
of Ampithoe sp. of Barnard (1965). Article 2 of pereopods 1-2 is short, as in the latter species. However,
article 4 of antenna 2 is like that of the type series and is not shortened as in Ampithoc sp. Probably
the latter is a rare individual of Ampihoe tea matching the phenotype in Bahia de Los Angeles but far
removed geographically from its optimal home.
Records. — Intertidal on rocky shores; shell sand, 2 5 m.
Distribution. — Southern and Baja California from Santa Barbara and Catalina Island to Punta Eugenia
and Guadalupe Island, intertidal to 67 m.
Family Aoridae
Acuminodeutopus periculosus, new species
Diagnosis. — Male gnathopod 1 having process of article 5 produced into 2 distal teeth, article 6 extending
well beyond process of article 5; coxa 1 rather shallow, evenly extended forward; gnathopod 2 with article
2 distally expanded anteriorly, article 3 with long anterior process, longer than known for A. heteruropus
Barnard (195 9a); inner ramus of uropod 3 with 2 distal spines. Pereopods 1-2 similar to those of
A. hcternropus, including 3 stout setae of article 5, otherwise with minor setal variations; pereopods 3-5
similar but with minor variations in setal placement; uropod 2 differing from that of A. hcternropus
by possession of 2 marginal spines on outer ramus; eyes slightly larger and less elongated than those of
A. heteruropus.
Holotype. — USNM No. 111461, male, 2.3 mm.
Type-locality. — BLA 3 9, April 24, 1962, 7 m.
Relationship. — Male gnathopod 1 of Acuminodeutopus heteruropus has a single tooth on article 5, a
1969
Barnard: Baiiia he los Angeles Amphipoda
191
Figure 8. Ampithoe pollex Kunkel, male, 7.0 mm, BLA 305, large retarded form: a, head .ind antennae;
b, c, d, e, f, pereopods 1, 2, 3, 4, 5; g, h, gnathopods 1, 2; i, Celson; j, k, uropod '; I, lower lip; m,
n, uropods 1, 2. Male, 5.4 mm, BLA SIO-62-215, small aberrant form: n, p, gnathopods 2, I.
192
San Diego Society of Natural History
Vol. 15
Figure 9. Acuminodeutopus periculosus, new species, male, 3.0 mm, BLA 116: a, b, gnathopod 1;
c, d, e, gnathopod 2; f, uropod 3. Female, 3.0 mm: g, h, gnathopods 1, 2.
short article 6, and gnathopod 2 has an unexpanded article 2 and no process on article 3. All males of
A. periculosus have the double cuspidation on article 5 of gnathopod 1 but the size and shape of the
teeth vary considerably.
Records. — Scattered along the shallowest perimeter on both sides of Bahia de Los Angeles in depths of
4 to 3 8 m, mainly shallower than 1 5 m, persistent in both seasons, on bottoms of coarse sand and shell,
rarely on fine sands.
Lcmbos macromamts (Shoemaker)
Bemlos macromanus Shoemaker, 1925: 36-41, figs. 10-13.
Lembos macromanus. Barnard, 1962a: 9, fig. 3; 1964b: 110, chart 12.
The largest male, 9.0 mm long, from BLA SIO-62-235, was figured in Barnard (1962a); its first
gnathopod is similar to that shown by Shoemaker (1925) and the pereon has 2 sternal teeth. Another male
of SIO-62-212 has the first gnathopod like that of L. concarus as identified by Barnard (1962a) but lacks
anterior setae on article 6 and has 3 sternal teeth on the pereon.
Records. — Rocky intertidal to at least 9 m on coarse substrate.
Distribution. — Gulf of California north to Estero de Punta Banda (Enscnada), Baja California, shallow-
water and intertidal.
Microdcutopus schmitti Shoemaker
Microdeutopus schmitti Shoemaker, 1942; 18-21, fig. 6. Barnard, 1959a: 32-33, pi. 9; 1961: 180; 1964a:
218.
The male of SIO-62-212 has a strong, sharp tooth on the anterodistal edge of article 2 on gnathopod
2; the tooth on the fifth article of gnathopod 1 is blunt, nearly spatulate, and lacks accessory teeth.
Records. — - Rocky intertidal to 44 m, primarily on coarse substrate.
Distribution . — Monterey Bay, California, to Cabo San Lucas, Baja California, 0-43 m.
Neomegampbopus roosevelti Shoemaker
N come gam phopus roosevelti Shoemaker, 1942; 36-38, fig. 13. Barnard; 1962a: 10.
Records. — Rocky intertidal to 24 m on shell sand.
Distribution. — Border of Mexico-California (32°N) to Cabo San Lucas, Baja California, 11-42 m.
Rudilemboides stenopropodus Barnard
Rudilemboides stenopropodus Barnard, 1959a: 31-32, pi. 8; 1964b; 110, chart 14.
1969 Barnard: Bahia de los Angeles Amphipoda 193
Records. — In April from rocky intertida] to 3 8 m, with most of the records below 9 m; in October from
4 to 19 m, with most of the records from 9 m or less.
Distribution. — Pt. Conception, California to Bahia de San Quintin, Baja California, 1-68 m.
Family Argissidac
Argissa bamatipes (Norman)
Argissa bamatipes (Norman). Walker, 1904: 246. Stebbing, 1906: 277. Shoemaker. 1930: 37-40, figs. I J,
16. Stephensen, 1931: 261; 1935: 140; 1940: 41; 1944: 52. Gurjanova, 1951: 327-328, fig. 193.' Barnard'
1962c: 151.
Argissa typica Boeck. Sars, 1895: 141-142, pi. 48.
Record. — Reef between Isla Ventana and Isla Cabeza de Caballo, shore.
Distribution. — North Atlantic in Gulf of St. Lawrence, Kattegat and northern Britain to Kola Bay;
Greenland; Chukchi, Bering, Okhotsk and Japan seas; California; 4-1096 m.
Genus Batea Muller
Occasionally fully adult but more often juvenile specimens of both Batea transversa and Batea
rectangnlata have the evanescent teeth on the third plconal epimera typical of Batea lobata. These are
considered as mutants. Batea lobata is distinguished from B. catharinensis largely by this difference, the
evanescence of teeth on the third pleonal epimera, although the gnathopods are slightly different. Batea
lobata might be considered a Pacific race of the Atlantic B. catharinensis, but a second Pacific cognate of
B. catharinensis, sympatric with B. lobata, is described below.
Key to Species of Batea (Note that coxa 1 is absent in this genus)
la. Coxa 2 quadrate rectangnlata
b. Coxa 2 subtriangular 2
2a. Palm of gnathopod 2 oblique 3
b. Palm of gnathopod 2 nearly transverse transversa
3 a. Teeth of third pleonal epimeron well developed 4
b. Teeth of third pleonal epimeron obsolescent lobata
4a. Articles 5 and 6 of gnathopod 2 equal in length; lobe on article 2 of pereopod 3 poorly de-
veloped; distal half only of inner margin of mandibular palp article 2 setose; rostrum evenly
tapering susurrator, n.sp.
b. Article 5 of gnathopod 2 slightly shorter than article 6; lobe on article 2 of pereopod 3
well developed; all of inner margin of mandibular palp article 2 setose; rostrum tapering
unevenly catharinensis
Batea rectangnlata Shoemaker
Batea rectangnlata Shoemaker, 1925: 31, figs. 7-9; 1926: 9-13, figs. 5-7.
Although other species of Batea in Bahia de Los Angeles differ from their cognates or their warm-
temperate relatives, this species, originally described from Bahia de San Francisquito, Gulf of California,
does not. Gnathopods, pereopods, head, and pleonal epimera correspond to those figured or described by
Shoemaker.
Records. — In April from 7 to 40 m, primarily below 15 m, on coarse sand and shell, scarce on fine sand;
in October occurring sparsely between 2 and 9 m.
Distribution. — Bahia de San Francisquito, Gulf of California.
Batea susurrator, new species
Figure 10
Diagnosis. — Rostrum tapering acutely and almost evenly; postantennal corner of lateral cephalic lobe
sharp, quadrate; mediodistal edge only of mandibular palp article 2 setose; lower lip lacking medial lobes;
articles 5 and 6 of gnathopod 1 subequal in length, both relatively thinner than in B. catharinensis (see
Shoemaker, 1926), palm oblique, slightly shorter than posterior edge of article 6. thus palm shorter than
in B. catharinensis; coxa 1 subtriangular; posterodistal corner of article 2 on pereopod 3 nearly quadrate,
not lobate; posteroventral rounded margin of article 2 on pereopod 5 with deep slit, article 2 slightly
broader than in B. catharinensis; telson cleft slightly less than halfway, edges proximally appressed cli sely,
distally gaping; posterior edge of third pleonal epimeron strongly serrate in adult, weakly in juveniles.
Pleonal epimera, uropods and mouthparts. except for mandibular palp and lower lip resemble those of
B. catharinensis; maxilla 1 with more setae on inner plate than in B. catharbtensil despite size differences:
when outer plate of maxilliped bears 2 spine-setae distally, inner plate of maxilla 1 be.irs
B. catharinensis when outer plate of maxilliped bears 4 spine-setae, inner plate of maxilla I bears only !
setae. Antenna 1 of B. susurrator bearing a distal cusp on medial surface of article 1, not known for
B. catharinensis.
Holotypc. — USNM No. 111485, male, 4.2 mm.
Type-locality. — BLA 132, April 22, 1962, 25 m.
Relationship. — This species differs from its cognate of the Caribbean-western Atlantic. B catharinensis,
by the lack of proximal setation on the medial edge of mandibular palp article 2; the
2, its palm and posterior lobe of article 5; the lack of a prominent lobe on artiele 2 ..t pereopod 3; the
evenly tapering rostrum. Batea susurrator resembles B. transversa Shoemaker (I "26) but differs
species in the absence of a lobe on article 2 of pereopod 3, the much larger second gnathopod with its
more prominent and oblique palm, and the posterior lobe of article 5. Batea susurrator might be con-
194
San Diego Society of Natural History
Vol. 15
Figure 10. Batea susurrator, new species, male, 4.3 mm, BLA 132: a, head; b, c, d, e, f, pereopods
1, 2, 3, 4, 5; g, h, i, uropods 1, 2, 3: j, telson; k, medial surface of article 1 of antenna 1; 1, man-
dibular palp; m, n, gnathopods 1, 2; o, pleonal epimera 2-3, left to right. Juvenile, 2.4 mm: p, pleonal
epimeron 3, left side.
1969 Barnard: Bahia de los Angeles Amimiipoda
195
e
fused with adults of B. lobata Shoemaker (1926) but again, the lack of a third pereopodal lobe th
larger gnathopod 2, and the sharper lateral cephalic lobes distinguish it, although the well-developed
serrations of the adult third pleonal epimeron are significant. Juveniles of B. msurrator have poorly
developed serrations but their other characters do not match those of B. lobata.
Records. — Ranging from 9 to 37 m in depth, most abundant in 2$ m on coarse gray sand.
Batea transversa coyoa, new subspecies
Figure 7 d, e, f, g
Diagnosis. — Differing from B. f. transversa Shoemaker (1926), by absence of posteroventral lobe on
article 2 of pereopod 3, by the more slender second articles of pereopods 4 and 5, the very slightly
shorter fifth article of gnathopod 2 and by the more oblique palm.
Holotype. — USNM No. 111458, female, 6.9 mm, ovigerous.
Type-locality. — BLA SIO-62-236, south arm of Bahia de Los Angeles, April 26, 1962, ca. 30 m, shrimp
trawl on N-S line between stations 156 and 45, coll. by Dr. Carl L. Hubbs' party.
Relationship. — Batea susurrator differs from B. t. coyoa in the expanded article 6 of gnathopod 2 and
the presence of a slit on article 2 of pereopod 5. Batea t. coyoa resembles a specmen of B. t. transversa
from Catalina Harbor California (Shoemaker, 1926, figs. 9 and 11) more than it does another specimen
from Pt. Loma, California (Shoemaker, 192 6, figs. 8 and 10). Correspondence is noted in the shapes of
coxae 5-6 and articles 5-6 of gnathopod 2. Gnathopod 1 of B. t. coyoa resembles that in figure 9c of
Shoemaker, especially notable being the contrast to B. susurrator and ca/hariiiensis in number, size, shape
and arrangement of setae.
Records. — On coarse substrate from 9 to 30 m.
Distribution of B. t. transversa. — Southern California, 2-27 m.
Carinobatca Shoemaker, 1926
Carinobatca differs from Batea in the presence of dorsal cusps on posterior segments and the elongate
article 1 of the palp on maxilla 1. Heretofore the absence of inner lobes on the lower lip of Carinobatca
appeared significant as a generic character but the type-species of Batea, B. cathar'mensis (Atlantic) also
lacks inner lobes. The Pacific members of Batea have inner lobes and this should be evaluated as a
generic character.
If dorsal cusps alone were the characters distinguishing Carinobatca from Batea, there would be
good cause to synonymize the genera because this kind of difference has rarely proved significant in other
Gammaridea. Characters other than the first maxillary palp are assorted in Carinobatca much as they
are in Batea and this would suggest a polyphyletic origin of the several species of Carinobatca from those
of Batea. But, the consistency in first maxillary palp indicates a monophyletic development of
Carinobatca.
Heretofore the enlarged process on article 1 of antenna 1 in members of Carinobatca represented a
good generic character, but a new species to be described below has the small process typical of Batea.
Carinobatea conductor, new species
Figure 1 1
Diagnosis. — Rostrum of medium length and stoutness in contrast to other species of Carinobatca (having
either shorter or longer rostra) ; distoventral tooth of article 1 of antenna 1 smaller than in other
species; gnathopod 1 bearing only 2 distal setae (in contrast to 6 or more in other species) ; gnathopod 2
with short article 3 (in contrast to its elongation in C. cuspidata Shoemaker, 1926), article 5 longer
than 6, its posterior lobe shallow and not pointing distally, palm oblique, equal to posterior margin of
article 6; article 2 of pereopod 3 linear, lacking posterodistal lobe; posterodistal lobe of article 2 of
pereopod 5 broadly expanded and deeply rounded (in contrast to C. carinata Shoemaker, 1926, having
a shallow, quadrate lobe); lateral margin of peduncle of uropod 1 with about 10 spines (compared witli
24 in C. cuspidata and about 11 in C. carinata); uropod 3 with flat, broad, heavily setose rami (in
contrast to C. carinata but not C. cuspidata) ; plconites 1 and 2 each with small dorsal tooth, pereonite 7
lacking tooth (in contrast to other species).
Holotype. — USNM No. 111495, ? male, 4.0 mm.
Type-locality. — BLA SIO-62-216, reef between Isla Ventana and Isla Cabeza de Caballo, shore, debris
of fish sample, April 21, 1962, coll. Dr. Carl L. Hubbs and party.
Relationship. — Similarities and contrasts to other species are stated in the diagnosis. The diversity of
characters among species of Carinobatca is much stronger than in Batea with differences in dorsal body
cusps, length of article 3 of gnathopod 2, striking differences in uropod 3 (see C. cuspidata), peduncular
spines of uropod 1, article 2 of pereopod 5, and the cusp on article 1 of antenna 1. In their linear
second articles of pereopod 3 all species of Carinobatca resemble Batea susurrator. B transversa coyoa
and B. catharinensis.
Mouthparts of C. conductor correspond to those figured by Shoemaker for C. cuspidata, especially in
the 4.0 mm male. The larger, 5.0 female has only a single accessory seta on article
palp; both sexes have a small distal cusp concealed in the terminal setae of that article; the inner plate
of maxilla 1 has 5-6 setae; in the female the first palp article of maxilla 1 is not as elongate as in
both sexes of C. cuspidata but the distal end of the palp is more strongly armed.
196
San Diego Society of Natural History
Vol. 15
Figure 11. Carinobatea conductor, new species, male, 4.0 mm, BLA SIO-62-216: a, head; b, c, gnath-
opod 2; d, c, medial surfaces of antennae 2, 1; f, g, h, uropods 1, 2, 3; i, telson; j, k, 1, m, pereopods
1, 3, 4, 5; n, coxa 4; o, gnathopod 1. Female, 5.0 mm: p, pleonal epimera 1-3, left to right; q, lateral
surface of article 1 of antenna 1.
1969 Barnard: Bahia de i.os Am.i i is Amimiipoda 197
Family Corophiidae
Corophium baconi Shoemaker
Corophium baconi Shoemaker, 1934: 356-359, fig. 1; 1949, 82. Barnard, 1959a: 38; 1961: 182; 1964b:
111-112, chart 16; 1964a: 219.
Records. — Rocky intertidal to 9 m.
Distribution. — Generally a species of lagoons and estuaries but occurring in the open sea from Peru to
the Bering Sea; off southern California in depths of 9-5 5 m and in 1-2 m in enclosed bays and harbors.
Corophium uenoi Stephensen
Corophium uenoi Stephensen 1932: 494-498, figs. 3, 4. Barnard, 1952: 28-32, pis. 8, 9; 1959a: 39. Nagata,
1960: 178. 'Barnard, 1961: 183; 1964b: 112, chart 16.
The accessory tooth above the large ventral tooth on article 4 of male antenna 2 is slightly larger
and blunter than in specimens from southern California.
Records. — Rocky intertidal of Vermilion Sea Field Station.
Distribution. — Japan; eastern Pacific Ocean from Monterey Bay, California at 24 m, to Bahia de San
Quintin, Baja California, rarely occurring in the open sea, more often in lagoons and estuaries, such as
Morro Bay, Newport Bay, and Bahia de San Quintin, intertidal to 2 m.
Ericthonius brasiliensh (Dana)
Ericthonius brasilicnsis (Dana). Barnard, 1955a: 37-38; 1959a: 39; 1961: 183; 1964a: 219.
Records. — From 0 to 42 m, primarily on intertidal or very shallow sand bottoms.
Distribution. — -Cosmopolitan in tropical, warm-temperate and some boreal seas, 0-130 m.
Rildardanus, new genus
Diagnosis. — Antenna 2 strongly ornamented; mandibular palp 3-articulate; inner plates of maxillipeds
broad, flat; gnathopod 1 in male much larger than 2, both pairs subchelate; accessory flagellum uni-
articulate, vestigial; inner rami of both uropods 1 and 2 scale-like, vestigial; uropod 3 composed only
of a setose peduncle; teison a large subcircular. simple flap.
Type-species. — Rildardanus tros, new species.
Relationship. — The shortened, scale-like inner rami of uropods 1-2 relate this genus to Pseudiricthonius
Schellenberg (1926), but that genus has a well-developed third uropod with a long ramus. Concholestes
Giles (1888) is the only other genus in the Corophiidae lacking rami on uropod 3 but the rami of
uropod 1 of that genus are not reduced, gnathopod 2 is larger than 1 and the second antenna is not
ornamented. The unelongate first urosomal segment prevents assignment of this genus to the Podoceridae.
Gnathopods of Rildardanus resemble those of Unciola most closely and probably Rildardanus is allied to
that genus, especially in view of the similarity of pleonal epimera (Sars, 1895, pi. 222), coxae, percopods
and mouthparts. This genus differs from Unciola in the loss of the ramus of uropod 3, further enlarge-
ment and special ornamentation of male antenna 2 and almost complete loss of the accessory flagellum.
The second antennae are ornamented as much as or more than they are in Corophium but the mouthparts
and gnathopods strongly differ in Rildardanus (compare Corophium in Sars, 1895, pis. 219-221).
Rildardanus tros, new species
Figures 12, 13
Diagnosis. — With characters of the genus.
Description of male. — Rostrum small, acute; lateral cephalic lobes rounded, article 3 of antenna 2
overriding them and fitting behind a ventral accessory lobe; eyes small, composed of few ommatidia;
article 1 of antenna 1 bearing 4 ventral spines and medial ridge with 5 spines; article 2 as long as 1;
article 3 shorter than 1, accessory flagellum minute, uniarticulate; antenna 2 fitting snugly to oblique
lower edge of head, article 1 large, triangular, article 2 evanescent laterally, evidenced by its blunt
gland-cone, article 3 broadened, with 1 dorsal spine on a poorly developed process distally produced
laterally, medial face with semicircle of stout spines; article 4 more slender, mediodist.il end strongly
produced acutely and bearing apical spine, lower edge when properly turned showing 5 ventral cusps
each with stout spines; article 5 very slender, about as long as 4; lateral surfaces of pereonitcs with
bulges just dorsal to coxae; anterior coxae small and acute anteriorly, fifth coxa largest, last 3 coxae with
small, hemispherical posterior lobes; gnathopods small, first larger than second, article 5 short, with blunt
posterior lobe, article 6 long, stout, posteroproximal end of article 6 produced into blunt tooth defining
false palm, finger slightly overlapping defining process, inner edge armed with sharp cusps; gnathopod 2
slender, articles 5 and 6 equal in length, article 5 poorly setose anteriorly and lacking posterior lobe,
palm of article 6 transverse, somewhat cheliform, denned by stout spine, dactyl overlapping palm;
pereopods simple, each sixth article with small distal spine at base of dactyl, pereopods 4 and 5 equal in
length, percopod 3 very short, second articles of last 3 pereopods bearing plumose setae on both anterior
and posterior edges; pleopods well-developed, decreasing in size slightly from pleopod 1 to pleopod 3.
each pleopod armed with 1 or 2 rasp-like coupling hooks; pleonites distinct, lateral surfaces of 4 and 5
with ridge, first 2 pleonal epimera with sinuoconvex posterior edges and sharp cusps at postcrovcntr.il
corners, third pleonal epimeron with cusp produced from anterior portion of ventral edge: uropod 1
slightly exceeding uropod 2, uropod 2 very short, inner rami about .me fourth as long as outer, scale-
like, each armed with distal spine, uropod 3 a simple, asymetrically expanded, terminally setose peduncle
attached to remnant of pleonite 6 below telson; teison a large, subcircular, simple flap; mouthparts
198
San Diego Society of Natural History
Vol. 15
Figure 12. Rildardanus tros, new genus, new species, male, 4.5 mm, BLA 212: a, lateral view; b, ple-
opod 3, minus setae; c, lateral view of head showing coupling process of antenna 2; d, coupling spine of
pleopod; e, pleopod 1, plumosities of setae shortened and reduced in number for clarity; f, medial view of
antenna 2, minus setae; g, pleonal epimera 1-3, left to right; h, dorsal view of head.
1969
Barnard: Bahia de los Angeles Amphipoda
199
Figure 13. Rildardanus tros, new genus, new species, male. 4.? mm, BLA 212: a, lower lip; b. c.
antennae 1, 2, medial views; d, accessory flagellum; e. gnathopod 1; f, g. gnathopod 2; h, i, uropod 1,
lateral and medial views showing small inner ramus; j, k, uropods 2, 3; 1, dorsum of telson and third
uropods; m, n, maxillae 1, 2; o, maxilliped; p, mandible; q, upper lip.
200 San Diego Society of Natural History Vol. 15
figured, generally typical of unciola-ericthonius corophiids, outer plate of maxilla 1 with 9 spines.
Female unknown.
Holotype. — USNM No. 1115 18, male, 4.5 mm.
Type-locality. — BLA 212, main west shore of Bahia de Los Angeles, west of Isla Ventana, dredge of
sand bottom, Ulva and gracilarioids, 9-16 m, April 28, 1962.
Family Dexaminidae
Polychcria osborni Caiman
Figure 2 5 g
Polychcria osborni Caiman, 1898: 268-269, pi. 32, fig. 2. Skogsberg and Vansell, 1928: 268-282, figs. 1-26.
Polychcria antarctica (Stebbing). Alderman, 1936: 63. Barnard, 1954c: 21 (not Stebbing, 1888).
Except for a greater setosity of mouthparts, uropods and thoracic appendages, these specimens fit the
morphology depicted by Skogsberg and Vansell (1928). The pleonal cpimera are drawn here because they
were not decribed by Skogsberg and Vansell. Coxae 1 and 2 are less strongly produced and the man-
dibular lobes are better developed in BLA specimens than in those from Monterey Bay, California, de-
scribed by Skogsberg and Vansell.
So much variation has been demonstrated for P. antarctica in the southern hemisphere (especially by
Schellenberg, 1931), that through a series of changes P. osborni has become a synonym of P. antarctica
(P. osborni to P. tenuipes by Stebbing, 1906; P. tenuipes to P. antarctica by Stebbing, 1910). Skogsberg
and Vansell indicated the inadvisability of such changes. A character that seems to distinguish P. osborni
from other known species and formae is the dense setation of the ventrolateral margin of the peduncle
of uropod 1; the pleonal epimera also may be characteristic but they require description on various forms.
Record. — Rocky intertidal, in tunicates, Vermilion Sea Field Station.
Distribution. — Puget Sound, Washington to southern California (32° N), intertidal, especially in
amarouciid tunicates.
Family Eusiridae
Pontogeneia nasa, new species
Figures 14, 15
Diagnosis of male. — Rostrum decurved, scarcely tapering, extending twice as far as lateral cephalic
lobes, the latter disymmetrically rounding, anteroventral margin of head straight, then slightly attenuated
to acute corner; eyes large; mandibular palp massive, article 2 broad, strongly setose mediodistally, article
3 more than 80 percent as long as article 2, setose medially for most of its free length; inner plate of
maxilla 1 with 3 terminal-subterminal setae, lateralmost very massive, medialmost subterminal; apices of
inner maxillipedal lobes narrower than in P. longleyi Shoemaker (1933b); posterodorsal ends of pleonites
1-2 projecting above following segments as in P. longleyi and P. quins ana Barnard (1964b); third pleonal
epimeron serrate behind, first and second epimera with small tooth and notch at posteroventral corners;
second articles of all pereopods somewhat thinner and relatively longer than in P. longleyi and P. quinsana;
rami of uropod 2 shorter than in P. quinsana and telson more deeply cleft, not as deeply as in P. longleyi;
male gnathopods like those of P. quinsana, gnathopod 1 slightly larger than 2, article 5 of gnathopod 2
with narrow, sharp, somewhat distally pointing lobe, posterior margins of sixth articles of both pairs of
gnathopods with 5-6 sets of stout spines, palm with larger spine on each side, palm poorly defined from
posterior margin of article 6 if judging by length of dactyl; dactyls of all pereopods with striate medial
ridge (also shown by Shoemaker for P. longleyi) ; antennae two thirds as long as body, armed with sense
organs.
Female. — Serrations of third pleonal epimeron poorly developed; fifth articles of gnathopods similar to
male, sixth articles small, narrow, palms oblique, spines at defining corners more numerous than in most
female pontogeneias; mandibular palp with article 2 scarcely broadened.
Mouthparts (mandibular body, outer plate and palp of maxilla 1, maxilla 2, outer plate and palp
of maxillipeds) like those of P. longleyi except as stated or figured; lower lip with outer lobes narrower
than in P. longleyi.
Holotype. — USNM No. 111418, male, 5.0 mm.
Type-locality. — BLA SIO-62-235, north of BLA village sandspit. beach poisoning, shore, debris of fish
sample, April 26, 1962 Dr. Carl L. Hubbs and party coll.
Relationship. — Resembles Pontogeneia quinsana Barnard (1964b) in the unusual serrations of the third
pleonal epimera and the shape of the rostrum. The latter feature also occurs in P. longleyi Shoemaker
(1933b), the male of which is unknown. The second articles of the pereopods are narrower in P. nasa
than in the other two species and small details of serrations are different (compare figures of the various
species) ; the serrate third pleonal epimera differentiate the new species from P. longleyi as does the long
third mandibular palp article and the very broad second article differentiates P. nasa from both species.
The telsonic cleft is intermediate in size, and the spination of the male gnathopods differs from that of
P. quinsana. The distal articles of the pereopods of P. quinsana are not known but the striated ridges of
the dactyls are apparently similar to those of P. longleyi.
The female has article 5 of gnathopod 2 projecting more strongly and turned more distally than in
P. longleyi or P. quinsana.
Record. — ■ Rocky intertidal to subintertidal on coarse substrate.
1969
Barnard: Bahia de los Angeles Amphipoda
201
Figure 14. Pontogeneia nasa, new species, male, 5.0 mm, BI.A SIO-62-235: a, b, c, pereopods 2, J, fj
d, pleonal epimera 1-3, left side; e, f, antennae 1, 2; g, articles 2. }, 4 of antenna 1. medial; h. ventral
surface of uropod 2; i, head; j, k, gnathopods 1. 2; I. m. defining spines of palms ..t gnathopods 1. 2; n,
medial surface of articles 6-7 of gnathopod 2.
202
San Diego Society of Natural History
Vol. IS
Figure 15. Pontogeneia nasa, new species, male, 5.0 mm, BLA SIO-62-235: a, uropod 3; b, inner plate
of maxilla 1; c, telson; d, mandibular palp; e, articles 3-4 of maxillipedal palp; f, inner plate of maxilliped.
Female, 5.4 mm: g, mandibular palp; h, i, gnathopods 1, 2; j, dactyl of pereopod 1. Male, 5.1 mm: k,
mandibular palp. Male, young 4.6 mm: 1, m, medial surfaces of articles 6-7 of gnathopods 1, 2.
Rhachotropis luculenta, new species
Figure 16
Diagnosis of male. — Head flattened, with very long, decurrent rostrum, lateral cephalic lobes strongly
projecting, falciform, apically obtuse; eyes very large, extending onto dorsal surface of head; mandibular
palp article 2 about 80 percent as long as article 3; only pleonites 1 and 2 with posterodorsal tooth,
segment 3 with 2 small posterior cusps on each side; pleonites 1-2 with posterolateral tooth on each side,
that of segment 2 connected to a lateral carina; pleonal epimera rounded at posteroventral corners, third
with serrate posterior edge; dactyls of pereopods 1-2 nearly as long as sixth articles; second articles of
pereopods 3-4 very short, broad, posterior edges with 2-3 rather large serrations; second article of pereopod
5 intermediate in width, anterior and posterior edges parallel, posteroventral corner not lobate, bearing 2
large serrations; telson cleft about halfway.
Female. — Eyes small, about two thirds as broad as length of rostrum; antenna 1 of normal female
configuration.
Holotype. — USNM No. 111546, male, 4.6 mm.
Type-locality. — BLA 124, 42 m, April 24, 1962.
Relationship. — This species is most similar to Rhachotropis inflata (Sars, 1895, pi. 152) but differs by
(1) the absence of lateral carinae on pleonite 3 and the presence of 2 posterior cusps on the posterodorsal
margin of each side; (2) the longer and more strongly deflexed rostrum; (3) the longer dactyls of the
first 2 pairs of pereopods; (4) the very broad, short, second articles of pereopods 3 and 4 with their
larger and less numerous teeth; (5) the small teeth on the unproduced posteroventral corner of the
second article of pereopod 5; and (6) a greater disproportion between the lengths of the fourth and fifth
peduncular articles of antenna 2. It resembles R. OCtdata (Hansen) (Gurjanova, 1951) in cephalic shape
but differs by the absence of a dorsal tooth on pereonite 7 and pleonite 3, the absence of a tooth on the
posteroventral corner of the second pleonal epimeron and the relatively parallel edges of the second article
on pereopod 5.
No male accessory flagellum has been detected; a small, fused scale with setae is present on the medio-
distal edge of article 3 of antenna 1 ; the female bears a uniarticulatc accessory flagellum.
Records. — On silt bottom in 3 8-46 m; possibly represented by 1 specimen in "Velero" Sta. 5114,
southern California (Allan Hancock Foundation).
1969
Barnard: Bahia de los Angeles Amphipoda
203
Figure 16. Rhachotropis luculenta, new species, male, 4.6 mm, BI.A 124: a, head; h, pereonite 7 and
pleonites 1-5, left side; c, d, antennae 2, 1 ; e, accessory flagellum; f, article 3 i>t" antenna 1; g, article 1
or antenna 1; h, article 4 of antenna 2; i, j, gnathopods 1, 2; k, 1, m, n, pereopods I, 3, 4, J; o, p, q,
uropods 1, 2, 3; r, telson; s, mandibular palp. Female, BLA 95: t, antenna I.
204
San Diego Society of Natural History
Vol. 15
Figure 17. Megaluropus longimerus falciformis, new subspecies, female, 4.1 mm, BLA SIO-62-228: a,
maxilliped; b, c, uropods 1, 2; d, mandibular palp; e, antenna 1; f, peduncular process of uropod 1.
Male, 3.6 mm: g, head; h, i, coxae 1, 3; j, gnathopod 2. Female, 5.1 mm: k, telson; 1, pleonites 1-J,
left to right; m, head. Juvenile, 2.6 mm: n, uropod 3. Megaluropus longimerus Schellenberg, southern
California, "Vclero" station 5 564, female, 6.5 mm: o, half of telson; p, pereopod 2. Female: 3.9 mm:
q, both halves of telson.
1969 Barnard: Bahia de los Angeles Amphipoda 20 5
Elasmopus rapax Costa
Elasmopus rapax Costa. Barnard, 1955a: 10-12, fig. 5 (with references); 1962b: 94-96, figs. 16, 17.
Elasmopus rapax mutatUS Barnard, 1962b: 96-98 fig. 18.
One form of this species was reported by Barnard (1962b) from BLA but a second form occurs in
SIO collections. It resembles a form from southern California (Barnard, in press) that has article 2 of
pereopod J strongly crenellated posteriorly. A third form from Tiburon Island, near BLA, is also described
by Barnard and an open-sea subspecies, E. r. mu/a/us occurs in southern California (Barnard, 1962b).
The second form from BLA is characterized as follows: pleonal cpimeron 3 like that o£ E. antennatm
and E. hoi gurus (see Barnard, 1962b), with smooth posterior margin and small tooth at posteroventral
corner; accessory flagellum elongate, 2-articulate, article 2 minute; gnathopod 2 like that of E. r. mutahts,
bearing one distal palmar process and one process in middle of palm as large as that of E. r. rapax of
Barnard (1962b), but setae denser, invading palm halfway and hiding middle process, proximal palmar
ridge and spine absent; telson short, truncate and heavily spincd as in /!. kolgurus Barnard (1962b) and
the crenellated form (Barnard, in press); uropod 3 short but inner ramus not shortened, outer ramus with
3 sets of marginal spines.
The wide variability in assortment of characters on accessory flagella, second gnathopods, fifth
pereopods, third pleonal epimera, third uropods and telsons suggests that E. ho/gurus may be another
phenotype of E. rapax.
Records. — Rocky intertidal of Vermilion Sea Field Station and barrier islands, to 24 m on barrier island
reef.
Distribution. — Circumtropical, subtropical and in some temperate seas.
Macra inaequipes (Costa)
Macra inaequipes (Costa). Barnard, 195 9a: 2 5-26, pi. 5.
The specimens correspond morphologically with those reported by Barnard (195 9a) from Newport
Bay, California. The question of the assignment of northeastern Pacific representatives to the European
species remains open to further study.
Records. — Rocky intertidal of barrier islands.
Distribution. — Possibly circumtropical and warm-temperate in the intertidal.
Megaluropus longimerus falciformis, new subspecies
Figures 17, 18
Diagnosis. — Lateral cephalic lobe with sharp cusp anteriorly; posterior edges of pleonal epimera 1-3
strongly rounded, ending posteroventrally in small cusps, 2 and 3 sparsely serrate posteriorly; dorsal edges
of pleonites 3, 4, 5 strongly serrate; coxae 1 and 2 longer than broad, coxae 2-3 rcniform, curved
posteriorwards, coxae 3-4 poorly setose, see drawings for sexual differences; article 4 of gnathopods in
both sexes strongly produced distally and posteriorly, lobe of article 5 well-developed and directed
somewhat distally in female, less strongly distinct from article 5 in male; long falciform spines on distal
articles of all pereopods extraordinarily stout and distally blunt; second articles of pereopods 3-5 more
slender than in southern California individuals; telson short, very broad, apices broad, nearly truncated,
armed with long, stout blunt spines, telson extending only halfway along inner ramus of uropod 2.
Holotype. — USNM No. 111475 female, 4.1 mm.
Type-locality. — BLA SIO-62-228, W. shore of bay, south of village, debris of fish sample on cobble
bottom, 2 m, April 24, 1962, coll. Dr. Carl L. Hubbs and party.
Relationship. — This subspecies differs from M. I. longimerus Schellenberg (as described from California
by Barnard 1962b) in (1) the more strongly convex posterior edges of the pleonal epimera and their
smaller posterodistal teeth; (2) the presence of a distinct cusp on the lateral cephalic lobes in both sexes;
(3) the bluntness of the pereopodal spines; (4) the broader telson with longer, blunter distal spines;
and (5) the poor setation of coxae 3 and 4.
Megaluropus longimerus was originally described from western Africa and its identification from the
eastern Pacific Ocean is open to question on geographic grounds. If the taxon has subspeciated in a s
geographic area then it may also be represented by different subspecies in California and Africa. The
original description is sketchy and the material should be reevaluated in light of various characters dif-
ferentiating the two Pacific subspecies. The telson and pereopod 2 of Californian material are reillustrated
herein (fig. 17, o, p, q) for comparison of spines with specimens from BLA.
Distribution. — Lagos, Nigeria; eastern Pacific Ocean from Monterey Bay, California to Bahia de San
Ramon, Baja California, 10-108 m.
Megaluropus visendus. new species
Figures 19, 20
Diagnosis. — Rostrum long, slender, acute, reaching nearly to end of article 1 of antenna 1 lateral cephalic
lobes bulbously produced and broadly rounded; eyes large, filling lateral lobes; antennae <
article 5 of second antennal peduncle much longer than article 4, accessory flagellum urn
antenna 2 peduncular base inflated, densely setose; upper lip scarcely incised
with 1 stout distal seta and numerous, slender setules lining medial edge: article 4 of maxillinedal palp
short, armed with 1 long stout spine and several setae; coxa 1 broad, anterior e
concavity above anteroventral corner; coxa 2 much broader than I
ventral edge; coxa 3 scarcely different from coxa 1 in size, anterodistal edge beveled: coxa 4 r
206
San Diego Society of Natural History
Vol. 1J
Figure 18. Megaluropus longimerus falciformis, new subspecies, female, 4.1 mm, BLA SIO-62-228:
a, b, gnathopods 1, 2; c, d, e, f, g, pereopods 1, 2, 3, 4, 5. Male, 3.6 mm: h, coxa 4.
1969
Barnard: Bahia de los Angeles Amphipoda
207
Figure 19. Megaluropus visendus, new species, male, 2.5 mm, BLA 9: a, antenna 2, flagellum cut after
article 3; b, maxilla 2; c, mandibular palp; d, maxillipedal palp articles 3-4; e, inner plate of maxilla 1;
f, upper lip. Female, holotype, 2.7 mm: g, h, i, uropods 1, 2, 3; j, telson; k, head.
than other coxae, evenly rounded below, not tapering; coxae 5-7 unusually long, coxa 5 with long anterior
lobe projecting ventrally, coxa 6 with small anterior lobe projecting anteriorly, coxa 7 with antcrovcntral
lobe; gnathopod 1 small, stout, simple, poorly setose, articles 4 and 5 rather bulbous, article 6 comparatively
slender, dactyl armed with stout, partially fused, distal spine; gnathopod 2 of female with very slender
second article bent proximally, article 4 small and not produced behind, article 5 triangular, long, with
apex of triangle forming posterior process, article 6 linear, slender, simple, article 7 about two thirds as
long as article 6, bearing 1 distal spine and several setules; gnathopod 2 of male with article 5 less tri-
angular in shape, posterior process pointing distally, article 6 inflated but slender relative to article 5,
tapering distally, without distinct palm, dactyl long, curved, about two thirds as long as palm and distally
bearing fused spine without accessory setules; pcreopod 2 longer than 1, but both otherwise similar, slender,
article 6 shorter than 5; fifth articles of pereopods 3, 4 (? and 5), about half as long as fourth articles,
these pereopods relatively shorter than in Megaluropus agilis; posterodorsal margins of pleonitcs 1 and 2
minutely serrate, segments 4 and 5 slightly elevated as in M. lottgimerus (as shown by Barnard. 1962b)
and serrate; third pleonal epimeron evenly rounded behind, densely serrate, second pleonal cpimeron sub-
quadrate at posteroventral corner, with 2 distal serrations; first pleonal cpimeron evenly convex behind,
protuberant posteriorly, rounded-quadrate at ventral corner; telson long, reaching about to middle of rami
of uropod 3, poorly armed.
Holotype. — USNM No. 1115 11, female, 2.7 mm.
208
San Diego Society of Natural History
Vol. 15
Figure 20. Megaluropus visendus, new species, female, 2.7 mm, holotype, BLA 9: a, head; b, pleonal
epimera 1-3, left to right; c, d, e, f, g, pereopods 1, 2, 3, 4, 5; h, pereopod 2, enlarged; i, j, gnathopods
1, 2. Male, 2.5 mm: k, head; 1, gnathopod 2. Arrows in figs, c and j indicate gill attachments.
1969 Barnard: Bahia de los Angeles Amphipoda 209
Type-locality. — BLA 9, 2 m, October 31, 1963.
Relationship. — This species resembles Megaluropm agilis Hock, as figured by Chtvrcux and Fage (1925)
but has numerous small differences: (1) the rostrum is narrower, longer, and more discrete; (2) the
heads of both sexes are similar and the eyes arc of the same large size, in contrast to the small female
eyes of M. agilis; (3) article 5 of second antennal peduncle is much longer than article 4; (4) the
accessory flagellum is uniarticulate; (5) coxa 1 is broader and coxae 4-7 are longer and of different shapes
than in M. agilis; (6) gnathopod 2 of the female is somewhat similar to that of M. agilis but article 5
is broader and article 6 more evenly slender; (7) gnathopod 2 of the male has article 5 much broader
and with a distinct posterior lobe; (8) pereopods 3-5 are shorter and article 5 is much shorter than in
M. agilis.
Dorsal serrations either occur or do not occur on pleonites 1-2 in the present material.
The gnathopodal differences of these two species indicate that the specimens identified by Barnard
(1964a) as M. agilis are in reality a new species. That identification was based on a scant material with
high variability but the following differences from M. agilis as figured by Chevreux and Fage (1925)
now appear significant: (1) broader first coxa; (2) the distinctly triangular shape, posterior lobation and
greater breadth of article 5 on male gnathopod 2 and the longer dactyl; (3) the more discrete rostrum of
the male; (4) the distinctive female second gnathopod which is like the first gnathopod in the new
species; but in M. agilis the first gnathopod has a broader fifth article with a distinctly subtriangular
shape, whereas in M. agilis of Barnard the article is a linear trapezoid; (5) the uniarticulate accessory
flagellum.
There are sufficient distinctions between the descriptions of M. agilis by Chevreux and Fage (1925)
and by Pillai (1957) to suggest that the latter Indian material is also a new species.
Records. — Ranging in depth from 2 to 17 m on coarse to fine sand.
Mcli/a sulca (Stout)
Caliniphargus sulcus Stout, 1913: 641-642.
Shoemaker (1941) relegated this species to Mclita pahnata (Montagu) but unlike that species, at least
as figured by Sars (1895), the second urosomal segment of Mclita sulca bears two pairs of small cusps,
each pair embracing a seta, whereas M. pahnata has a single dorsolateral cusp on each side.
Records. — Rocky intertidal to 24 m in the offshore islands and north of the village sandspit.
Distribution. — Southern California intertidal.
Meximaera, new genus
Diagnosis. — Antenna 1 with 4-articulate accessory flagellum; article 2 of mandibular palp longer than
either articles 3 or 1, article 3 slender, linear (not falcate); lower lip bearing inner lobes; inner plate
of maxilla 1 setose only terminally; inner plate of maxilla 2 scarcely setose on medial edge, considered
primarily to be setose terminally; article 4 of maxillipedal palp not claw-shaped, short, bearing several
long, distal spine-setae; gnathopods small but distinctly subchelate; uropod 3 with subequal rami, outer
minutely biarticulate; telson cleft.
Type-species. ■ — Meximaera diffidentia, new species.
Relationship. — ■ Differing from Macra Leach by the short, non-unguiform article 4 of the maxillipedal
palp and the strongly biarticulate outer ramus of uropod 3. The general aspect of Meximaera resembles
that of Maeracunha Stephensen (1949), from Tristan da Cunha. The two genera have truncate antero-
lateral cephalic margins and biarticulate outer rami of uropod 3, but Meximaera differs from Maeracunha
in the poorly or non-setose medial edges of the inner lobes of both pairs of maxillae. Meximaera differs
from Maeropsis Chevreux (1927) and Paraccradocus Stebbing (see 1906) in the poorly setose medial edge
of maxilla 2 and the biarticulate outer ramus of uropod 3. The above-two genera may be synonymous.
Maerella Chevreux (1911) has a triarticulate maxillipedal palp.
This genus appears to be a derivative of the Macra-linc with the retention (primitive) of a second
article on the outer ramus of uropod 3 and the rare occurrence of a shortened fourth maxillipedal palp
article.
Meximaera, like Linguimaera Pirlot (1936), has the anteriorly lobate lip which also is common
to several species of Maera and caused Schellenberg (1938) to synonymize Linguimaera with Macra. The
identifications of M. othonides Walker (1904), the type-species of Linguimaera, are confused. Pirlot (1936)
discounted Chilton's (1921) and K. H. Barnard's (1935) identifications and Nayar (1959), who did not
cite Pirlot (1936), has identified material with M. othonides which fails to correspond with Pirlot's
Until valid specimens of Walker's species can be reexamined minutely for a second article on the outer
ramus of the third uropod and for its maxillipedal palp article 4, there remains the possibility that
Meximaera and Linguimaera ire synonymous. If so, Linguimaera should be revived and take precedence
over Meximaera.
Meximaera diffidentia, new species
Figures 21, 22
Diagnosis. — With the characters of the genus.
Notes. — The sides of the head are almost without lobular differentiation. The dactyls of the pereopods
are complexly armed (see figures). Article 1 of antenna 1 has
palmar surface of gnathopod 2 has not only a pair of marginal defining spines but a large submargin
spine dominating the palm. Female unknown.
210
San Diego Society of Natural History
Vol. 15
Figure 21. Meximaera diffidentia, new genus, new species, male, holotype, 5.8 mm, BLA SIO-62-216:
a, head; b, c, gnathopod 1, lateral and medial views; d, gnathopod 2; e, maxilliped; f, g, h, uropods 1, 2,
3; i, pleonal epimera 1-3, left to right.
Holotype. — USNM 111497, male, 5.8 mm.
Type-locality. — BLA SIO-62-216, reef between Isla Ventana and Isla Cabeza de Caballo, shore, tailings
of fish sample, April 21, 1962, Dr. Carl L. Hubbs and party, coll.
Records. — Rocky intertidal to 24 m in the offshore islands.
Family Haustoriidae
Platyhchnopus mctagracilis Barnard
Platyischnopus metagracilh Barnard, 1964a: 22 5-226, fig. 3.
Records. — Ranging in depth from 13 to 46 m, more abundant in 20-46 m, frequency in April in latter
depths is 170 individuals/m-, in October, 2&/m~.
Distribution. — Punta Canoas to Bahia de San Cristobal, Baja California, 30-73 m.
1969
Barnard: Bahia de los Angeles Amphipoda
211
Figure 22. Meximaera diffiidentia, new genus, new species, male, holotype, 5.8 mm, BLA SIO-62-2 16:
a, tip of outer ramus of uropod 3 showing article 2; b, c, d, pcrcopods 3. 4. *; e, maxilla 1; t. gnathopod
2, medial surface of palm; g, mandible; h, maxilla 2; i. lower lip; j, telson; k. pereopod I.
212 San Diego Society of Natural History Vol. 15
Platyischnopus viscana Barnard
Platyischnopus viscana Barnard, 1964a: 226, fig. 4.
Record. — Rocky intertidal of offshore islands.
Distribution.- — La Jolla, California to Bahia Magdalena, Baja California, 17-27 m.
Family Hyalidae
Hyale rubra frequens Stout
Allorchestes frequens Stout, 1913: 650-651.
Hyale rubra (Thomson). Hurley, 1957: 910-913, figs. 30-50.
Hyale nigra (Haswell). Barnard, 1962c: 153-156, figs. 19, 20.
Hyale rubra frequens Barnard (in press).
Specimens from BLA have the elongate antennae typical of specimens from Bahia de San Quintin
(Barnard, 1964b) and the setae on the posterior edge of the hand on male gnathopod 1 are widely spread;
they occupy 30 percent of the marginal length.
Records. — Rocky intertidal to 7 m.
Distribution. — California and Baja California, intertidal.
Family Isaeidae (=Photidae)
Ampbideutopus oculatus Barnard
Amphideutopus oculatus Barnard, 1959a: 34-35, pi. 10; 1961: 181, fig. 2; 1964a: 236; 1964b: 110,
chart 15.
Records. — -Rocky intertidal of offshore islands; 9-16 m in dredge haul; all other samples in depths of
22-41 m primarily on brown silty sands.
Distribution. — Pt. Conception, California to Bahia de San Cristobal, Baja California, 2-162 m.
Eurystheus thompsoni (Walker)
Eurystheus tenuicornis var. lobata Shoemaker, 1942: 28, fig. 10 a-c.
Eurystheus thompsoni Shoemaker, 1955: 59.
Gnathopod 1 is typical of var. lobata, having the large, posterior, setose lobe on article 2.
Record. — Rocky intertidal of offshore islands.
Distribution. — Bahia Magdalena, Baja California (lobata variety).
Eurystheus tonichi, new species
Figure 2 3
Diagnosis. — Similar to E. thompsoni (Walker) (see Shoemaker, 1931, 1942, 1955) and in male under-
going similar radical transformation of gnathopod 2 and coxa 7; following characters similar to E.
thompsoni: eyes and head, with slight anterior cusp on sharp lateral cephalic lobe; epistome strongly pro-
duced and acute, other mouthparts similar but setules on inner plate of first maxilla vrey fine; pleonal
epimera 1-3 with tooth at posteroventral corners, third with very convex, nearly subacute, posterior edge
(male holotype slightly aberrant, other specimens like Shoemaker, 1931, figure 4a); pleonites 4-5 with
pairs of dorsolateral teeth similar to E. thompsoni; uropods similar, rami of uropod 3 slightly longer than
peduncle, uniarticulate, inner slightly longer than outer; telson strongly excavate posteriorly, each lobe
armed with 1 spine and setules. Characters distinct from E. thompsoni: gnathopod 1 with article 6 stouter,
more inflated; fully adult male gnathopod 2 with palm and posterior margin of article 6 confluent, simple,
distal end of palm with single cusp, dactyl folding halfway along posterior margin of article 6; young
male gnathopod 2 similar to young male of E. thompsoni (compare fig. 2 3k herein and fig. 3a of
Shoemaker, 1931).
Most of the specimens have lost pereopods 3-5 but this does not detract from the several specific
characters distinguishing this species from E. thompsani. Pereopods 4-5 of the holotype and of a young
male are very small and may be regenerate. Article 2 of pereopod 4 on the young male is narrow and
posteriorly sinuous.
Female almost identical with that of E. thompsoni, specimens from California being compared with
those of Bahia de Los Angeles (figures 23 n, o herein for gnathopod 2).
Holotype. — USNM No. 111472, male, 5.3 mm.
Type-locality. — BLA 212, west shore of Bahia de Los Angeles, west of Isla Ventana, dredge of Viva and
gracilarioids, sand bottom, 9-16 m April 28, 1962.
Relationship. — Differs conspicuously from E. thompsoni by the simple palm of the terminal adult male,
having a single cusp; in E. thompsoni the palm is distally subtransverse, short and armed with 2 cusp-
teeth, one of them defining the palm. Pereopod 3 of E. tonichi apparently does not differ from that of
E. thompsoni except by the slightly broader article 2.
Records. — Rocky intertidal to 38 m on shell sands.
Photis ?bifurcata Barnard
?Photis bifurcata Barnard, 1962a: 30-31, fig. 10; 1964a: 240.
Specimens are subadult but presumably represent this unusual species.
Records. — Ranging in depth from 38 to 46 m.
Distribution. — Monterey Bay, California to Bahia de San Cristobal, Baja California, 11-93 m.
1969
Barnard: Bahia de los Angeles Amphipoda
213
Figure 23. Eurystheus tonichi, new species, male, 5.3 mm, BLA 212: a, pleonal cpimcra 1-3, left to
right; b, maxilla 1; c, pereopod 3; d, gnathopod 2, 2 views; e, f, pereopods 4, 5; g, coxa 7; h, gnathopod 1.
Male, 4.5 mm, BLA SIO-62-215: i, j, pereopods 4, 5; k, gnathopod 2, lateral view minus sct.it. Male, SA
mm, BLA 212: 1, m pereopods 3, 4. Female, 6.3 mm, BLA 212: n, o, gnathopod 2, 2 views. Male. 5.4 mm,
BLA SIO-62-215: p, gnathopod 2, lateral.
214 San Diego Society of Natural History Vol. 15
Pbotis breiipes Shoemaker
Pbotis brciipes Shoemaker, 1942: 25-27, fig. 9. Barnard, 1962a: 31-33, fig. 11; 1964a: 240-241.
The two adult males of BLA 2 5 (3.0 mm) have the inner protuberance on article 7 of gnathopod 2
smaller and more proximal than in Calilornian specimens.
Records. — Ranging in depth from 9 to 3 6 m.
Distribution. — Coos Bay, Oregon to Bahia Magdalena, Baja California, 0-13 5 m.
Pbotis California! Stout
Pbotis californica Stout. Barnard, 1962a: 33-36, figs. 12, 13; 1964a: 241.
The male of BLA 144 (3.0 mm) has a well-developed lobe on article 2 of gnathopod 1, and the
palm of the hand, similar to the female second gnathopod, is more excavate than in specimens from
California.
Records. — Ranging in depth from 34 to 46 m.
Distribution. — Monterey Bay, California to Bahia de San Cristobal, Baja California, 10-139 m.
Family Ischyroceridae
Jassa falcata (Montagu)
Jassa falcata (Montagu). Sexton and Reid, 1951: 30-47, pis. 4-30. Barnard, 1959a: 37; 1964b: 118.
Records. — Ranging in depth from 7 to 18 m.
Distribution. — Cosmopolitan in shallow waters and intertidal zones, especially in harbors on pilings.
Microjassa macrocoxa Shoemaker
Microjassa macrocoxa Shoemaker, 1942: 44-47, figs. 16, 17.
Records. — Rocky intertidal to 3 8 m, most abundant in shallow water dredge hauls.
Distribution. — Bahia Magdalena, Baja California, 18-27 m.
Family Leucothoidae
Leucotboe alata Barnard
Leucotboc alata Barnard, 1959a: 19-20, pi. 1; 1962c: 132, figs. 7 D, E, F; 1964a: 227.
Records. — Rocky intertidal of Vermilion Sea Field Station.
Distribution. — Monterey Bay, California to Bahia de San Ramon, Baja California, 0-24 m.
Family Liljeborgiidae
Liljeborgia marcinabrio, new species
Figure 24
Diagnosis. — Lateral cephalic lobes obtuse, strongly projecting; eyes large but poorly pigmented, ommatidia
sparse; mandibular palp article 2 not exceeding article 1 in length; coxa 1 barely expanded distally, coxa
2 strongly beveled anterodistally, coxa 4 sparsely serrate posteriorly, corner at excavation rounded, not
sharp; article 4 of both gnathopods acutely produced and slightly attenuate; dactyls of gnathopods 1 and
2 with 4 and 7 teeth on respective inner edges; serrations on second articles of pereopods 3-5 strongly
demarcated; both margins of both rami of uropod 3 spinosc; telson with apices of each lobe very deeply
incised, resulting pair of cusps asymmetrical; posterodorsal edges of pleonites 1-2 each with 3 teeth, of
pleonites 4-5 each with one large, posteriorly directed tooth, pleonite 3 lacking dorsal teeth; pleonal
epimera 1 and 2 each with posteroventral tooth, third epimeron with bifid tooth.
Holotype. — USNM No. 111504, female, 5.4 mm. Unique.
Type-locality. — BLA 13 8, 46 m, April 22, 1962, brown silt.
Relationship. — This species, having the aspect of Liljeborgia kinabani (Bate) (see Sars 1895: pi. 188,
fig. 1), differs by the larger, poorly pigmented eyes, the strongly spinose rami of uropod 3, the bifid tooth
of pleonal epimeron 3, the slightly more expanded first coxa but more strongly beveled second, by the
rounded cephalic lobe and the larger, less disproportionate teeth of pleonites 4-5.
In terms of dorsal pleonal tooth-formulas this species has the following relatives, each of which is
listed with its distinctions: L. aequabilis Stebbing (in Hurley, 1954) has a single posteroventral tooth on
the third pleonal epimeron, less strongly notched telsonic apices, a denser comb of spines on article 6
of pereopods 1-2, a smooth posterior edge of coxa 4; L. akaroica Hurley (1954) has a single tooth on the
posteroventral corner of the third pleonal epimeron, and larger teeth on pleonites 4-5; L. longicornU
(Schellenberg, 1931) has only a simple notch on the third pleonal epimeron and a broadly expanded first
coxa, its peculiar dactyl of pcreopod 5 not being compared to the new species because of the loss of
pereopods in the unique specimen; L. macrodon Schellenberg (1931) again has the simply-notched third
pleonal epimeron and obsolescent medial cusps of the telsonic apices; both L. mixta Schellenberg (1925)
and L. octotlcntata Schellenberg (1931) differ from /.. marcinabrio by their simple third epimeral tooth
and sinus.
Liljeborgia marcinabrio bears a remarkable resemblance to L. bansoni Hurley (1954), from New
Zealand, in pleonal epimera, telson, pereopods, and except for pleonite 1, the dorsal sculpture. Pleonite 1
of L. bansoni has only a single medial tooth. Its second mandibular palp article is longer than the first.
Listriella melanica lazaris, new subspecies
Figure 2 6 c-g
References to the nominate subspecies are as follows: Barnard, 1959b: 16-18, figs. 1, 2; 1964a: 229;
1964b: 108, chart 8.
1969
Barnard: Bahia de los Angeles A
MIMIIPODA
215
Figure 24. Liljeborgia marcinabrio, new species, female, 5.4 mm, BLA 138: a, maxilliped; b, mandibu-
lar palp; c, pleon; d, dorsum of pleonites 1-2 (a single example); e, head; f. end of article 3 of antenna 2,
2 views; g, end of article 4 of antenna 2; h, antenna 1; i, lower lip; j, k, j;nathopods 2, 1; 1, m, n, o,
p, pereopods 1, 2, 3, 4, 5; q, r, s, uropods 1. 2, 3; t, telson.
216
San Diego Society of Natural History
Vol. 15
Figure 25. Pachynus barnardi Hurley, female 1.8 mm, BLA 97: a, b, pereopods 1, 5; c, gnathopod 1.
Orchomcne magdalenemis (Shoemaker), male, 3.7 mm, BLA 142: d, head; e, f, gnathopod 1. Volycheria
osborni Caiman, ovigerous female, 5.6 mm, BLA 214: g, pleonal epimera 1-3, left to right.
Diagnosis. — Female differing from female of nominate subspecies by distally expanding sixth article of
gnathopod 1; this structure is similar in males of both subspecies.
Holotype. — USNM No. 111481, female, 3.2 mm.
Type-locality. — -BLA 132, 25 m, April 22, 1962, bottom of shell sand.
Records. — Ranging in depth from 7 to 44 m; generally more abundant between 13 to 40 m; frequency
of 39 individuals/m- in latter depths in April.
Distribution of L. m. tnelanica. — Pt. Conception, California to Bahia de San Cristobal, Baja California,
12-97 m in the open-sea, 2-3 m in Bahia de San Quintin.
Family Lysianassidae
HippomcJon ?propinquus Sars
Figure 7 k, 1, m, n
?Hippomcdon propinquus Sars, 1895: 57, pi. 21, fig. 1. Gurjanova, 1962: 107-111, figs. 24-26 (with
references). Barnard 1964b: 80-82 (as H. denticulatus and //. propinquus).
These subadult specimens lack the notch on pleonal epimeron 3 characteristic of the H. denticulatus
group of species but Barnard (1964b) has presented evidence questioning that character as specifically
significant. The specimens from BLA differ from //. p. propinquus in Gurjanova (1962) in the non-
chelate palm of gnathopod 2 (fig. 7 k, 1) and in this character the specimens resemble those from
southern California and Bahia de San Quintin. Gnathopod 2 of BLA specimens differs from //. p. sibiricus
in the shape of the palm and the short curved dactyl. Article 1 of the flagellum on antenna 1 is short-
ened (fig. 7n) in contrast to //. p. sibiricus Gurjanova (1962) and H. p. propinquus.
Records. — -Ranging in depth from 15 to 30 m on silt and coarse sand.
Distribution. — The species occurs throughout the subarctic and boreal regions.
Barnard: Bahia de los Angeles A
MI'ltll'ODA
Figure 26. Monoculodes hartmanae Barnard, female, 4.6 mm. BLA 5 J: a, b, gnathopods I, 2. ListrielU
melanica lazar'n Barnard, new subspecies, male, 3.6 mm, BLA 132: c, gnathopod 2; d. palm of gnathopod
2; e, gnathopod 1 with offset spines and portion of palm. Female, 3.2 mm: f, g, gnathopods 1, 2.
218 San Diego Society of Natural History Vol. 15
Lysianassa dissimilh (Stout)
Nannonyx dissimilh Stout, 1913: 638-639.
Aruga dissimilh. Shoemaker, 1942: 7, fig. 2. Barnard, 1955b: 100-103, pi. 29, figs, g, i.
Lysianopsis dissimilh. Hurley, 1963: 76-77, fig. 2 Id.
Lysianassa dissimilh. Barnard (in press).
Record. — Rocky intertidal, island reef.
Distribution. — Coast of California from Tomales Bay to Isabel Island, Mexico and the Galapagos Islands,
0-73 m.
Lysianassa macromerus (Shoemaker)
Aruga macromerus Shoemaker, 1916: 157-15 8.
Lysianopsis macromerus Hurley, 1963: 77.
Lysianassa macromerus Barnard (in press).
Record. — Rocky intertidal, Vermilion Sea Field Station.
Distribution. — Intertidal of southern California.
Orchomcne magdalenensh (Shoemaker)
Figure 2 5 e, f
Orchomcnella magdalenensis Shoemaker, 1942: 4-7, fig. 1.
Orchomcne magdalenensh. Barnard, 1964a: 231-232; 1964b: 95, fig 12.
The male has a large, projecting, lateral cephalic lobe like that of the female, in contrast to the
male described by Barnard (1964b) from Bahia de San Quintin. The first gnathopod of the male but
not the female is strongly excavate in contrast to both Barnard's and Shoemaker's drawings (see figure
herein). The female from Bahia de Los Angeles has the fourth pleonite similar to that of males, unlike
outer-coast specimens.
Records. — Ranging in depth from 6 to 46 m.
Distribution. — Bahia de San Ramon to Bahia Magdalena, Baja California, 2-11 m.
Pachynus barnardi Hurley
Figure 2 5 a-c
Vachynus barnardi Hurley, 1963: 31-25, figs. 6, 7.
Only a few small specimens, approximately 1.8 mm in length have been collected in this survey.
These differ from the original description in the stouter and shorter sixth articles of pereopods 3-5 and
the somewhat stouter and shorter fifth article of gnathopod 2.
Records. — Ranging in depth from 3 0 to 46 m.
Distribution. — Monterey Bay, California to Bahia de San Ramon, Baja California, 12-183 m.
Uristes entalladurus Barnard
Uristes enalladurus Barnard, 1963: 460-462, figs. 5, 6; 1964b: 100.
The third pleonal epimeron is somewhat more quadrate and article 5 of gnathopod 2 more bulbous
than in specimens from southern California.
Record. — • Silty sand, 3 8 m.
Distribution. — Port Hueneme, California to Bahia de San Quintin, Baja California, 2-18 m.
Family Oedicerotidae
Monoculodes hartmanae Barnard
Figure 26 a, b
Monoculodes hartmanae Barnard, 1962d: 363-365, figs. 5-7.
Although Monoculodes nyci Shoemaker (1933a and see Barnard, 1962d) was expected from Bahia
de Los Angeles, because of its occurrence at San Felipe to the north of Bahia de Los Angeles, all of the
specimens of the genus collected in the latter bay belong to M. hartmanae, as shown in the accompanying
drawings of gnathopods 1 and 2.
Records. — Rocky intertidal to 41 m but generally better represented between 13 and 38 m; frequency
in the latter depths in April is 5 1 individuals/m-.
Distribution. — Pt. Conception, California to Bahia de San Quintin, Baja California, about 2 to about
146 m but scarce in waters deeper than 37 m.
Synchelidium rec/ipalmiim Mills
Synchelidium rectipalmum Mills, 1962: 17-19, figs. 5, 6B.
Records. — Rocky intertidal to 24 m on coarse substrate.
Distribution. — British Columbia to Costa Rica (specimens at hand), intertidal to 100+ m.
Synchelidium sp. G
This apparently new species is filed in Hancock and Smithsonian collections with this label; a study
of this genus is in preparation.
Records. — Ranging in depth from 42 to 46 m.
Distribution. — Coastal shelf of southern California.
1969 Barnard: Bahia de los Angeles Amphipoda 219
Westwoodilla cornuta, new species
Figures 27, 28
Description of female. — Rostrocephalon long, not strongly vaulted above, no distinct minute rostrum
appended, anterior end of head slightly downturned and armed with 2 small setae, eyes apically located in
distal half of rostrocephalon, small, eosin in color, anterolateral cephalic lobe sharp, symmetrical; article 1
of antenna 1 stout, produced dorsodistally into long, acute cusp, article 2 equal to 1, article 3 about 60
percent as long as article 2; antenna 1 reaching nearly to end of peduncular article 5 of antenna 2, the
latter as long as head and pereon together, article 5 varying from 0 percent to 3 3 percent longer than
article 4, flagellum as long as peduncle; cpistome broadly and subacutely produced forward; upper lip
broader than in W. caccula (in Sars, 1895: pi. 115) mandibular incisor simple, blunt, both sides with
lacinia mobilis and 3 spines, molar short, very broad, blunt, flat, nearly smooth, bearing single short stout
seta either simple or irregularly inflated in animals about to molt; palp article 2 proximally expanded and
densely armed with stout setae; lower lip with rather narrow but curled mandibular lobes; palp article 2
of maxilla 1 broad, distally expanding; outer lobe of maxilla 2 narrower than inner lobe; inner plates of
maxilliped much narrower than in other species of Westwoodilla; all anterior coxae with posteromedial
spine, coxa 4 narrow for the genus, coxa 1 very broadly expanded; gnathopods small, scarcely subchclate,
sixth articles narrow, fifth articles lacking posterior lobes (slightly better developed in gnathopod 1 of
young specimens), gnathopod 2 longer and more slender than gnathopod 1; telson apically truncate, with
slight lateral notches; third pleonal epimeron strongly setose ventrally and posteriorly (poorly in juveniles
3.0 mm), epimera 1-2 ventrally setose.
Male. — Antenna 2 slightly longer than in female, article 3 of first antennal peduncle as long as article 2,
both articles 2 and 3 stouter than in female.
Holotype. — USNM No. 111442, female, 4.9 mm.
Type-locality. — BLA 105, 38 m, April 28, 1962.
Relationship. — Apart from normal specific differences occurring in other taxa of the genus, this species
has characters that might serve as criteria for a new subgenus: the produced epistome; the proximally
expanded and very strongly spinose second article of the mandibular palp; the very broad, flat, and
smooth mandibular molar; the somewhat different lower lip (fig. 27i) ; the distally expanding second
article of the first maxillary palp; the narrower inner plates of the maxilliped and the narrower outer
plates of maxilla 2.
Mandibles of Californian Westwoodilla caecula (also figured in Sars, 1895, but with differences noted
herein that may be of specific value) are shown herein with a smaller, more projecting molar having
distinct cutting cuspules and spines, plus a much larger callused area (similar to Sars, 1895; pi. 117,
figure of Bathymedon) defining the primary cutting edge than are seen in the new species. Few mouth-
parts of other species of Westwoodilla have been figured, so comparisons are in order when taxonomists
reexamine them.
Specifically, the cusp of article 1 on antenna 1 is sufficient for differentiation from other known
species of Westwoodilla. Apparently the strongly setose pleonal epimera are unique in the genus. Coxa 4
is very narrow, coxa 3 is excavate posteriorly and the outer ramus of uropod 3 is conspicuously shorter
than the inner ramus, all characters not heretofore found in Westwoodilla.
Records. — From 19 to 46 m, with a frequency in April of 6.0 individuals/m-, primarily on silt and silty
sand bottoms.
Family Phliantidae
Heterophlias seclusiis cscabrosa Barnard
Heterophlias seclusiis cscabrosa Barnard, 1962b; 79, fig. 5.
Record. — Rocky intertidal of Vermilion Sea Field Station.
Distribution. — The nominate subspecies occurs in the Dry Tortugas Islands off Florida (Shoemaker,
1933b: 250-252, figs. 4, 5). The subspecies cscabrosa occurs in the eastern Pacific Ocean in southern
California, intertidal to about 16 m.
Family Phoxocephalidae
Heterophoxus oculatus (Holmes)
Heteropboxus oculatus (Holmes). Barnard, 1960b: 320-324, pis. 59-61; 1964a: 242.
Most of the specimens from the October samples are either juveniles or the adult form heretofore
known as H. o. nitcllus Barnard (1960). H. o. nitellus Barnard (1960b) is probably a synonym of
H. o. oculatus.
Records. — Rocky intertidal to 46 m, but rare in depths of less than 30 m; frequency in April at 30-46
m is 124 individuals/m-, in October, 44 individuals/m-, on silts with rare exception.
Distribution. — Puget Sound, Washington to Panama, 13-1785 m, at 2 m in Bahia de San Quintin,
Baja California.
Paraphoxus contains Barnard
Paraphoxus coguatus Barnard, 1960b: 233-235, pi. 24.
A specimen from Dillon Beach, California, is the first female of the species to be recorded. It is like
the male except for typical dimorphic characters: medium sized eyes, short second antennae and the short
inner ramus of uropod 3 which is about two thirds as long as the outer ramus. The epistomal cusp is as
short as that figured by Barnard; this character plus the long fifth articles of the gnathopods continue
220
San Diego Society of Natural History
Vol. 15
Figure 27. Westwoodilla cornuta, new species, female, 4.9 mm, BLA 105: a, head with epistome below;
b, plconal epimera 1-3 left to right; c, antenna 1; d, telson; e, f, uropods 1, 2. male, 2.9 mm, BLA 149:
g, h, antennae 1 2. Female, 5.4 mm, BLA 122: i, lower lip; j, k, maxilla 1; 1, maxilla 2; m, mandibular
palp; n, mandibular body. Female, BLA 158: o, upper lip; p, q, diagrams of mandibles to show molars and
differentiated portions of primary cutting edges. Westwoodilla caecula (Bate), southern California: r, s,
body and portion of body of right and left mandibles showing molar and differentiated portions of pri-
mary cutting edge in contrast to W. cornuta.
1969
Barnard: Bahia de los Angeles Amphipoda
221
Figure 28. Westwoodilla cornuta, new species, female, 4.9 mm, BLA 105: a, b, c, d, pcrcopods 2, 3,
4, 5 (latter reduced in size); e, coxa 3; f, g, gnathopods 1. 2; li, maxilliped.
222
San Diego Society of Natural History
Vol. 15
Figure 29. Paraphoxus gemmatus, new species, female, 5.8 mm, holotype, BLA 102: a, b, c, d, pere-
opods 2, 3, 4, 5, setae not drawn on pereopod 4; e, gnathopod 1; f, g, gnathopod 2; h, telson; 1, epistome;
j, head, dorsal; k, 1, m, uropods 1, 2, 3.
to be the principal differences separating the species from P. similis Barnard.
Record. — ■ Coarse shell sand, 6 m.
Distribution. — Southern California, surface nekton to 32 5 m, rare. A single female of this species is in
the collections of the Pacific Marine Station, Dillon Beach, near Tomales Bay, California.
Paraphoxus epistomus (Shoemaker)
Paraphoxus epistomus (Shoemaker). Barnard. 1960b: 205-209, pis. 6-8; 1964a: 243.
Paraphoxus lucubrans Barnard, 1960b: 212-218, pis. 12-14.
Record. — Gray sand, 9 m (lucubrans variety)
Distribution. — Western Atlantic Ocean, New Hampshire to South Carolina; eastern Pacific Ocean from
Mendocino County, California to Panama, 0-182 m.
Paraphoxus gemmatus, new species
Figure 29
Diagnosis of female. — Head with broad rostrum but constricted anterior to eyes; eyes medium in size;
body broad and dorsally depressed; epistome acutely produced anteriorly; maxillipedal palp article 4 with
small apical spine, one subterminal seta and one mesial seta; article 5 of gnathopods 1-2 longer than
article 6, latter moderately broad and expanded distally, palm transverse; ratio of articles 5 to 6: gnathopod
1=14:11; gnathopod 2=13:11; articles 4-5 of pereopod 3 moderately expanded, narrower than article 2,
article 6 shorter than 5, ratio of widths of articles 2, 4, 5, 6=52: 40: 45: 20; article 4 of pereopod 4
slightly broader than article 4 of pereopod 3, article 6 much longer than 5, ratio of widths of articles 2,
4, 5, 6=78: 45: 32: 13; article 2 of pereopod 5 broad, ratio of width to length about 9: 11, posterior
edge of article 2 with 5 small, symmetrical teeth, sweep-point occurring near distal end of article 4,
ventral edge broad, scarcely dipping anteriorly and slightly convex; remainder of pereopod 5 moderately
stout, article 6 longer than 5; uropods stout, rami of uropod 1 shorter than peduncle, very stout, expanded
proximally, strongly notched outer ramus with 3 marginal spines, inner with 2 large spines (shining
like jewels), lateral margin of peduncle densely spinose, with 3 large proximal setae on lateral surface,
inner margin with seta, 2 marginal spines and 1 large distal spine; uropod 2 with rami subequal to
1969
Barnard: Bahia de los Angeles Amphipoda
223
Figure 30. Garosyrrhoe disjuncta, new species, female, 4.2 mm, BLA SIO-X: a, head; b, c, d, pereopods
3, 4, J; e, f, articles 6-7 of gnathopod 2; g, h, articles 6-7 of pereopods 1.2; i. j. articles 6-7 of gnath-
opod 1; k, pleonal epimera 1-3, left to right; 1, dorsal ornamentation of plconite ?; m. dorsal ornamen-
tation of pereonite 7 and pleonites 1-2. Male, 3.0 mm: n, maxilla 1; o. inner maxillipedal plate; p,
maxilliped.
224 San Diego Society of Natural History Vol. 15
peduncle, very strongly notched and stout (not fully flattened in lateral drawing), each ramus with
stout spine; inner ramus of uropod 3 as long as outer ramus, both strongly setose; telson moderately broad,
apices rounded, each with 1 spine and 1 setule; third pleonal epimeron like P. epistomus (see pi. 6 in
Barnard, 1960b), but spines much stouter, gland cone similar.
Male. — -With typical sexual difference in its large eyes, strongly setose uropod 3, long second antennae
and the more broadly rounded, asymmetrical third pleonal epimeron. Uropod 1 with only 2 spines on
each ramus, neither expanded as much as in female. Epistomal process well-developed.
Holotype. — USNM No. 111499, ovigerous female, 5.8 mm.
Type-locality. — BLA 102, 6 m, October 31, 1963.
Relationship. — This species is similar to Paraphoxus bcterocuspidatus Barnard (1960b) but differs by the
presence of an acute, anterior epistomal cusp, the even teeth of pereopod 5 (some specimens of jP. hetero-
cuspidatus also show this), the presence of a marginal spine on both rami of uropod 3 and more than
one on each ramus of uropod 1, with the rami much stouter and more strongly notched than in
P. bcterocuspidatus.
A specimen from BLA-9, provisionally referred to this species, is a male with no epistomal process
and a fifth pereopod more like that of P. lucubrans Barnard (1960b) than that of the female of
P. gemmatus; uropod 1 and all other characters except pereopod 5 are like those of P. bcterocuspidatus.
Records. — Ranging in depth from 2 to 9 m.
Paraphoxus spinosus Holmes
Paraphoxus spinosus Holmes 1905: 477-478. Kunkel, 1918: 76-78, fig. 13. Shoemaker, 1925: 26-27.
Barnard, 1959a: 18; 1960b: 243-249, pis. 29, 31; 1961: 178; 1964b: 105.
These specimens are like those from Newport Bay, California figured by Barnard (1960b).
Records. — Rocky intertidal to 24 m on coarse substrate.
Distribution. — -Western Atlantic Ocean; eastern Pacific Ocean from Puget Sound to the Gulf of California,
0-37 m, rarely to 73 m.
Paraphoxus tridentatus (Barnard)
Pontharpinia tridentata Barnard, 1954c: 4-6, pis. 4, 5.
Paraphoxus tridentatus Barnard, 1960b: 261-262.
Paraphoxus tridentatus pallidas Barnard, 1960b: 262-263, pis. 38, 39.
These specimens are like those of P. /. pallidas figured by Barnard (1960b) but the rami of uropod
2 and the outer ramus of uropod 1 each lack the single spine characteristic of P. /. pallidas. The inner
ramus of female uropod 3 is as long as article 1 of the outer ramus, whereas irv P. t. pallidas the inner
ramus is half as long as article 1 of the outer ramus. Gnathopods 1 and 2 are slightly stouter than in
P. t. pallidus, more like those of the open-sea Oregonian form figured by Barnard (1954c).
Records. — Ranging in depth from 19 to 38 m.
Distribution. — Puget Sound, Washington to just north of Pt. Conception, California, shallow water and
intertidal.
Family Pleustidae
Paraplcustes commensalis Shoemaker
Parapleustes commensalis Shoemaker, 1952: 231-232, fig. 8 3.
This specimen, 4.0 mm long, corresponds to Shoemaker's figures and description, except that it is
morphologically better-developed, even though it is 1.5 mm shorter. The inner plate of the maxilliped has
7 of the small blunt spines (not 4), the lateral margin of mandibular palp article 3 has 3 spines (not 2)
and the gnathopodal palms have 3 sets (not 2) of double spines on the defining margins. Pleonal epimeron
3 is more strongly convex posteriorly than it is in Shoemaker's material.
Record. — Dredge, west side of bay at station 40, 9 m.
Distribution. — Santa Barbara, California, from pleopods of a spiny lobster.
Family Podoceridae
Podocerus brasilicnsis (Dana)
Podocerus brasilicnsis (Dana). Barnard, 1962a: 67, fig. 30; 1964a: 245-246.
Records. ■ — Rocky intertidal to 24 m on coarse substrate.
Distribution. — Cosmopolitan in tropical and warm-temperate, shallow seas.
Podocerus fulanus Barnard
Podocerus sp., Barnard, 195 9a: 40, pi. 14.
Podocerus fulanus Barnard, 1962a: 69.
Considerable variation occurs in the height of pereonal and pleonal dorsal teeth with large adults
having these teeth projecting much more strongly than in Barnard's (195 9a) figures. Large males have
the palm of gnathopod 1 strongly concave.
Records. — Rocky intertidal to 42 m.
Distribution. — Newport Bay, California, about 0-2 in.
Family Synopiidae
Garosyrrhoe disjunct;!, new species
Figure 30
Diagnosis. — Posterodorsal edges of pereonite 7 and plconites 1-2 each with pair of blunt dorsolateral teeth,
1969 Barnard: Bahia de los Angeles Amphipoda 22 5
each with medial hook and separated by shallow excavation, these teeth less prominent on pleonite 3,
dorsolateral edges of these segments also with small hooked teeth; teeth of posterior edge of third pleonal
epimeron much stronger than in G. bigarra (Barnard, 1962b); rostrum well -defined, acute, strongly pro-
jecting ventrally; article 5 of pereopod 3 about 80 percent as long as article 4, poorly setose posteriorly,
article 2 bulging anterodistally.
Holotype. — USNM No. 111467, female-like, 3.0 mm.
Type-locality. — BLA SIO-X, Piedras AJiogadas, 0.5 mile south of Isla Ventana, SCUBA sample of shell
fragments, pebbles, 24 m, April 22, 1962, Dr. Carl L. Hubbs and party, coll.
Relationship. — This species differs from Garosyrrhoe bigarra (Barnard, 1962b) by the longer, more acute
rostrum, the double dorsal teeth of the body segments, the stronger teeth of the third pleonal epimeron
and the shorter fifth article of pereopod 3. Pereopods 4 and 5 of G. bigarra are not known. Spination of
articles 6-7 of pereopods 1-2 differs between the two appendages (see figures). The dorsal surface of the
head bears a crest and the ventral surface of the inner maxillipedal plate bears a hooked spine. All the
specimens are either females or lack brood-plates and have short second antennae and are therefore
female-like.
Records. — Rocky intertidal to 24 m, usually on coarse substrate.
Acknowledgements
This report was supported in part by a National Science Foundation Grant (G-20909) and by the
Beaudette Foundation of California. Mr. Richard F. Dwyer, of Los Angeles, California, has generously
assisted with funds for publication. Miss Jacqueline M. Hampton prepared most of the plates. I gratefully
acknowledge the assistance of numerous other persons listed by Barnard and Grady (1968). Mr. Scott
Gray and Dr. Joel W. Hedgpeth, then of Pacific Marine Station, Dillon Beach, California, kindly loaned
me a specimen of Parapboxus cognatus for inclusion in this study. Mrs. Carolyn B. Gast prepared the
charts.
Literature Cited
Alderman, A. L.
193 6. Some new and little known amphipods of California. Univ. California Publ. Zool., 41: 5 3-74.
Barnard, J. L.
1952. Some Amphipoda from Central California. Wasmann Jour. Biol., 10: 9-36.
1954a. Amphipoda of the family Ampeliscidae collected in the eastern Pacific Ocean by the VcleroUI
and VeleroIV. Allan Hancock Pacific Expeds., 18: 1-137.
1954b. Amphipoda of the family Ampeliscidae collected by the Velcro III in the Caribbean Sea.
Allan Hancock Atlantic Exped. Rept. 7: l-!3.
1954c. Marine Amphipoda of Oregon. Oregon State Monogs., Studies in Zool., 8: 1-103.
195 5a. Gammaridean Amphipoda (Crustacea) in the collections of Bishop Museum. Bernice P. Bishop
Mus., Bull. 215.
195 5b. Notes on the amphipod genus Aruga with the description of a new species. Bull. S. California
Acad. Sci., 54: 97-103.
1958. Index to the families, genera, and species of the gammaridean Amphipoda (Crustacea).
Allan Hancock Found. Pubis., Occ. Pap. 19: 1-145.
195 9a. Estuarine Amphipoda. In Barnard, J. L., and D. J. Reish, Ecology of Amphipoda and
Polychaeta of Newport Bay, California. Allan Hancock Found. Pubis. Occ. Pap. 21: 13-69.
195 9b. Liljeborgiid amphipods of southern California coastal bottoms with a revision of the family.
Pacific Nat. 1 (4): 12-28.
1960a. New bathyal and sublittoral ampeliscid amphipods from California, with an illustrated key to
Ampelisca. Pacific Nat. 1(16): 1-36.
1960b. The amphipod family Phoxocephalidae in the eastern Pacific Ocean, with analyses of other
species and notes for a revision of the family. Allan Hancock Pacific Expeds. 18: 17'
1961. Relationship of Californian amphipod faunas in Newport Bay and in the open sea. Pacific
Nat. 2: 166-186.
1962a. Benthic marine Amphipoda of southern California: families Aoridae, Photidae, [schyrocendae,
Corophiidae, Podoceridae. Pacific Nat. 3: 1-72.
1962b. Benthic marine Amphipoda of southern California: families Tironidae to Gammandae. Pacific
Nat. 3: 73-115.
1962c. Benthic marine Amphipoda of southern California: families Amphiloch.dae, Leucothoidac,
Stenothoidae, Argissidae, Hyalidae. Pacific Nat. 3: 116-163.
1962d. Benthic marine Amphipoda of southern California: family Oedicerotidac. Pacific Nat. 3:
349-371. .
1962e. Benthic marine exploration of Bahia dc San Quintin, Baja California, 1960-61. General.
Pacific Nat. 3: 249-274.
226 San Diego Society of Natural History Vol. 15
1963. Relationship of benthic Amphipoda to invertebrate communities of inshore sublittoral sands of
southern California. Pacific Nat. 3: 437-467.
1964a. Los anfipodos bentonicos marinos de la costa occidental de Baja California. Rev. Soc. Mexicana
Hist. Nat. 24: 205-274.
1964b. Marine Amphipoda of Bahia de San Quintin, Baja California. Pacific Nat. 4: 55-139.
1965. Marine Amphipoda of the family Ampithoidae from southern California. Proc. U. S. Natl.
Mus. 118: 1-46.
1966. Benthic Amphipoda of Monterey Bay, California. Proc. U. S. Natl. Mus. 119 (3541): 1-41.
In press. Intertidal gammaridean Amphipoda (Crustacea) of California: Monterey to La Jolla. Bull.
U. S. Natl. Mus.
Barnard, J. L., and J. R. Grady
1968. A biological survey of Bahia de los Angeles, Gulf of California, Mexico. I. General account.
San Diego Soc. Nat. Hist., Trans. 15: 5 1-66.
Barnard, K. H.
1935. Report on some Amphipoda, Isopoda, and Tanaidacea in the collections of the Indian Museum.
Rec. Indian Mus. 37: 279-319.
Beltran, E., (Gen. Chm.)
1960. Symposium: The biogeography of Baja California and adjacent seas. Syst. Zool. 9: 47-91.
Calman, W. T.
1898. On a collection of Crustacea from Puget Sound. Ann. New York Acad. Sci. 11: 259-292.
Chevreux, E.
1908. Amphipodes recueillis dans les possessions francaises de l'oceanie par M. le Dr. Seurat, direc-
teur du laboratorie de recherches biologiques de Rikitea (iles Gambier). 1902-1904. Mem. Soc.
Zool. France. 20: 470-5 27.
1911. Campagnes de la Melita. Les amphipodes d'Algerie et de Tunisie. Mem. Soc. Zool. France.
23: 145-285.
1927. Crustaces amphipodes. Exped. Sci. "Travailleur" et du "Talisman" pendant les annes 1880,
1881, 1882, 1883. Malacostraces (suite). 9: 41-152.
Chevreux, E., and L. Face
1925. Amphipodes. Faune de France. 9: 1-488.
Chilton, C.
1921. Report on the Amphipoda obtained by the F.I.S. "Endeavour" in Australian seas. Biol. Res.
Fish. Exper. F.I.S. "Endeavour," 1909-14. 5: 33-92.
Garth, J. S.
1960. Distribution and affinities of the Brachyuran Crustacea. Syst. Zool. 9: 105-123.
Giles, G. M.
1888. No. 9. Further notes on the Amphipoda of Indian waters. Natural history notes from H. M.'s
Indian marine survey steamer 'Investigator,' Commander Alfred Carpenter, R.N., D.S.O.,
commanding. J. Asiatic Soc. Bengal. 57: 220-225.
Gurjanova, E.
1951. Bokoplavy morei SSSR i sopredel'nyx vod ( Amphipoda-Gammaridea) . Opred. po Faune SSSR,
Izd. Zool. Inst. Akad. Nauk. 41: 1-1031 (in Russian).
1962. Bokoplavy severnoi chasti Tixogo Okeana (Amphipoda-Gammaridea) chast' 1. Opred po Faune
SSSR, Izd. Zool. Inst. Akad. Nauk. 74: 1-440 (in Russian).
Holmes, S. J.
1905. The Amphipoda of southern New England. Bull. U. S. Bur. Fish. 24: 459-529.
Hurley, D E.
1954. Studies on the New Zealand amphipodan fauna No. 9. The families Acanthonotozomatidae,
Pardaliscidae and Liljeborjiidae. Roy. Soc. New Zealand. Trans. 82: 763-802.
1957. Studies on the New Zealand amphipodan fauna. No. 14 — The genera Hyalc and Allor-
chcstes (family Talitridac). Roy. Soc. New Zealand. Trans. 84: 903-933.
1963. Amphipoda of the family Lysianassidae from the west coast of north and central America.
Allan Hancock Found. Pubis. Occ. Pap. 25: 1-165.
Kunkfl, B. W.
1910. The Amphipoda of Bermuda. Connecticut Acad. Arts Sci. Trans. 16: 1-116.
1910. The Arthrostraca of Connecticut. Connecticut Geol. Nat. Hist. Sur., Amphipoda, Bull. 6 (1):
15-181.
Mills, E. L.
1962. Amphipod crustaceans of the Pacific coast of Canada. II. Family Ocdicerotidae. Nat. Hist.
Paps. Nat. Mus. Canada. 15: 1-21.
1964. Noteworthy Amphipoda (Crustacea) in the collection of the Yale Peabody Museum. Postilla,
Yale Peabody Mus. Nat. Hist. 79: 1-41.
1969 Barnard: Bahia de los Angeles Amphipoda 227
Nagata, K.
1960. Preliminary notes on benthic gammaridean Amphipoda from the Zostera region of Mihara Bay,
Seto Inland Sea, Japan. Publ. Seto Mar. Biol. Lab. 8: 163-182.
Nayar, K. N.
1959. The Amphipoda of the Madras Coast. Bull. Madras Govt. Mus., n.s. Nat. Hist. Sect. 6: 1-59.
Pillai, N. K.
1957. Pelagic Crustacea of Travancore. 111. Amphipoda. Bull. Cent. Res. Inst. Univ. Travancore
5: 29-68.
PlRLOT, J. M.
193 6. Les amphipodes de l'expedition du Siboga. Deuxieme partie. Les amphipodes gammarides. II. —
Les amphipodes de la mer profonde. 3. Addendum et partie generalc. III. Les amphipodes
littoraux. 1. Lysianassidae, Ampeliscidae . . . Siboga-Exped., Monog. 33e: 237-328.
Sars, G. O.
1895. Amphipoda. J;; An account of the Crustacea of Norway with short descriptions and figures
of all the species. Christiania and Copenhagen. 1: i-viii, 1-711.
Schellenberg, A.
1925. Crustacea VIII: Amphipoda. In Michaelsen, W., Beitrage zur Kenntnis der Meeresfauna West-
afrikas, 3(4): 11-204.
1926. Die Gammariden der deutschen Sudpolar-Expedition 1901-1903. Deutsch Siidpolar-Exped. 18:
235-414.
1931. Gammariden und Caprelliden des Magellangebietes, Siidgeorgiens und der Westantarktis.
Further Zool. Res. Swedish Antarctic Exped. 1901-1903. 2 (6): 1-290.
1938. Litorale Amphipoden des tropischen Pazifiks. Kungl. Svenska Vetenskapakad. Handl., ser. 3.
16(6): 1-105.
Sexton, E. W., and D. M. Reid
195 1. The life-history of the multiform species fassa falcata (Montagu) (Crustacea Amphipoda)
with a review of the bibliography of the species. J. Linnean Soc. London. 42: 29-91.
Shoemaker, C. R.
1916. Description of three new species of amphipods from southern California. Proc. Biol. Soc.
Wash. 29: 157-160.
1925. The Amphipoda collected by the United States Fisheries steamer 'Albatross' in 1911, chiefly
in the Gulf of California. Bull. Amer. Mus. Nat. Hist. 52: 21-61.
1926. Amphipods of the family Bateidae in the collection of the United States Museum. Proc. U. S.
Natl. Mus. 68(2626): 1-26.
1930. The Amphipoda of the Cheticamp Expedition of 1917. Contr. Can. Biol. Fish. 5: 221-359.
1931. A new species of amphipod crustacean ( Acanthonotozomatidae) from California, and notes
on Eurystheus tenuicornh. Proc. U. S. Natl. Mus. 78 (2861): 1-8.
193 3 a. Amphipoda from Florida and the West Indies. Amer. Mus. Novitates. 5 98: 1-24.
193 3b. Two new genera and six new species of Amphipoda from Tortugas. Papers Tortugas Lab.,
Carnegie Inst. Washington. 28(435): 245-2 5 6.
1934. Two new species of Corophium from the west coast of America. J. Wash. Acad. Sci. 24:
356-360.
1938. Three new species of the amphipod genus Ampithoe from the west coast of America. J. Wash.
Acad. Sci. 28: 15-25.
1941. On the names of certain California amphipods. Proc. Biol. Soc. Washington 54: 187-188.
1942. Amphipod crustaceans collected on the presidential cruise of 193 8. Smiths. Misc. Colls. 101
(11): 1-52.
1949. The amphipod genus Corophium on the west coast of America. J. Wash. Acad. Sci. 39: 66-82.
1952. A new species of commensal amphipod from a spiny lobster. Proc. U. S. Natl. Mus. 102:
231-233.
195 5. Notes on the amphipod crustacean Maeroides thompsoni Walker. J. Wash. Acad. Sci. 45: 59.
Skogsberg, T., and G. H. Vansell
1928. Structure and behavior of the amphipod, Polycheria osborni. Proc. Calif. Acad. Sci. ser. 4.
17: 267-295.
Soule, J. D.
1960. The distribution and affinities of the littoral marine Bryozoa (Ectoprocta) . Syst. Zool. 9:
100-104.
Stebbing, T. R. R.
1906. Amphipoda I. Gammaridea. Das Tierreich. 21: 1-806.
1910. Crustacea. Part 5. Amphipoda. Sci. Res. Trawling Exped. H.M.C.S. "Thetis." Australian Mus.
Mem. 4 (2): 565-658.
Stephensen, K.
1931. Crustacea Malacostraca. VII. (Amphipoda. III.). Danish Ingolf-Exped. 3:179-290.
1932. Some new amphipods from Japan. Annot. Zool. Japon 13: 487-501.
228 San Diego Society of Natural History Vol. 15
1935. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Tromso Mus. Skr. 3(1):
1-140.
1940. Marine Amphipoda. Zool. Iceland. 3(26): 1-111.
1944. Amphipoda. The Zool. of East Greenland. Medd. Gr^nland, 121 (14): 1-165.
1949. The Amphipoda of Tristan da Cunha. Res. Norwegian Sci. Exped. to Tristan da Cunha 1937-
1938. 19: 1-61.
Stout, V. R.
1913. Studies in Laguna Amphipoda. Zool. Jahrb. Syst. 34: 633-659.
Walker, A. O.
1904. Report on the Amphipoda collected by Professor Herdman, at Ceylon, in 1902. Suppl. Rept.
Ceylon Pearl Oyster Fisheries . 1904. 17: 229-300.
Revised Manuscript Accepted for Publication 22 October 1968
Division of Crustacea, Smithsonian Institution, United States National Museum,
Washington, D.C. 20560.
APPENDIX
Derivation of New Systematic Names
periculosus, L., perilous
susurrator, L., mutterer
coyoa, Sp., coyote
conductor, L., employer
Rildardanus tros; Ril, anagram from Lirondo, Sp., pure; Dardanus, mythical ancestor of royal family of
Troy; Tros, grandson of Dardanus and son of Ericthonius.
nasa, L., fish-trap
luculenta, L., bright
falciformis, L., sickle-shaped, likeness
visendus L., gerundive of viso, contemplate
Meximaera diffidentia; Meximaera, a Mexican Maera; difHdcntia, L., diffidence
tonichi, Mexicanization of L., tono, thunder; nidus, nest
marcinabrio, L., marcidus, perishable; abrir, Sp., to open
lazaris, Sp., lazar, to catch with lasso
cornuta, L., horn
gemmatus, L., bearing jewels
disjuncta, L., separation
AA
A RECENT MOLLUSCAN FAUNA FROM
THE CARIBBEAN COAST OF
SOUTHEASTERN PANAMA
GEORGE E. RADWIN
MUS. COMP. ZOOL.
LIBRARY
JUL 10 wo
HARVARD
UNIVERSITY
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 14, 27 JUNE 1969
A RECENT MOLLUSCAN FAUNA FROM THE
CARIBBEAN COAST OF SOUTHEASTERN PANAMA
George E. Radwin
Abstract. — This paper lists a Recent marine molluscan fauna from Payardi Island, Panama.
Twenty-nine per cent of these species have close relatives on the Pacific side of the isthmus. It is
suggested that the term "cognate" be used to replace the presently used term "analogue" in referring
to these closely related forms.
Ri sumen. — En este articulo se incluye una lista de los moluscos recientes de la is 1 a Payardi,
Panama. El veintinueve por cien de estas especies tienen parientes proximos en la parte del istmo
correspondiente al Pacifico. Se sugiere que el termino "cognado" reemplace al termino "analogo" que
luego esta usado en referencia a estas formas intimamente relacionadas.
The molluscan fauna of the Caribbean coast of Panama is of particular interest
because, except for the paper of Olsson & McGinty (1958), it has been virtually un-
reported. The building of an oil refinery at Payardi Island, Republic of Panama, six
Caribbean Sea
MILES
Figure I. Location of the studied area in reference ,,!l
San Diego Soc. Nat. Hist., Trans. [5
230
San Diego Society of Natural History
Vol. 15
miles east of the city of Colon, (Fig. 1) from May, 195 8 to August, 1959 provided
an excellent opportunity to collect and record this fauna. During construction of the
refinery material was dredged from the shoreward edge of a submerged coral reef and
pumped into a marshy near-shore area to form a spoil bank. A representative series of
mollusks was collected from this spoil by Messrs. Robert H. Stewart, Supervising
Geologist, Panama Canal Co., and Anselmo Mena, and deposited in the U. S. National
Museum. Most of this material is apparently of non-fossil origin and the Recent com-
ponent forms the basis of this report.
Systematic List of Mollusks from Payardi Island, Panama
An asterisk signifies the existence of a cognate in the East Pacific (see Table 1).
GASTROPODA
PROSOBRANCHIATA
SCISSURELLIDAE
Woodtt ardia sp.
FlSSURELLIDAE
Emarginula pttmila (A. Adams)
*Hemitoma octoradiata (Gmelin)
''Diodora cayenensis (Lamarck)
Diodora Inter] (Orbigny)
Litcapina suffusa (Reeve)
Fissurella fascicitlaris (Lamarck)
Fissitrclla nodosa (Born)
Trochidae
Calliostoma euglyptum (A. Adams)
Calliostoma jatanicnm (Lamarck)
Cittarium pica (Linne)
Tegula fasciatits (Born)
Solariorbis schumoi (Vanatta)
Solariorbis corylus Olsson & McGinty
Solariorbis hondurasensis (Vanatta)
Solariorbis infracarinata (Gabb)
Solariorbis shimeri (Clapp)
'''Anticlimax schumoi (Vanatta)
Cyclostrematidae
Cyclostremiscus sehrammi (Fischer)
Cyclostremiscus jeannae
(Pilsbry & McGinty)
Pacbystrcmiscns pule bell its
(Olsson & McGinty)
TURBINIDAE
'■'Turbo castancus Gmelin
Turbo filosus Wood
Astraea americana (Gmelin)
* Astraea caelata (Gmelin)
Astraea pboebia (Roding)
Liotia tricarinata Stearns
Phasianellidae
Tricolia pulchella (C. B. Adams)
Tricolia affinis cruenta Robertson
Neritidae
"'Neri/ina virginea Linne
Smaragtlia viridh t iridemaris Maury
Architectonicidae
* Architectonic a nobilis Roding
*'Hcliacn\ pcrricri (Rochebrune)
Vermetidae
Vermctus varians Orbigny
Aletes megintyi Olsson & Harbison
Aletes nebulosus (Dillwyn)
Aletes floridanus Olsscn & Harbison
"Stephopoma myrakeenac Olsson & McGinty
Spiroglyphus annulatus Daudin
Thylacodes sp.
Modulidae
''Modulus modulus (Linne)
'•'Modulus carchedonius (Lamarck)
Cerithiidae
Cerithium algicola C. B. Adams
"'Cerithium littcratum Born
Cerithium mocnense Gabb
''Cerithium variabile C. B. Adams
Bittium varium (Pfeifrer)
Alabina cerithioides Dall
''Litiopa melanostoma Rang
Turritellidae
Turrit 'ell a exoleta Linne
Littorinidae
Littorina sp.
RlSSOIDAE
Rissoina canccllata Philippi
*Rissoina fischeri Desjardin
Rissoina decussa/a (Montagu)
Rissoa lipeus Dall
Zebina brouuiana (Orbigny)
Alrania aberrans (C. B. Adams)
'■' Alvania auberiana (Orbigny)
Caecidae
Caecum imbricatum Carpenter
Caecum hcladum Olsson & Harbison
Caecum ryssotitum Folin
'Caecum pule helium Stimpson
Caecum jucundum Folin
Caecum datum Folin
Meioceras nit id urn Stimpson
Vitrinellidae
Cochliolepis striata (Dall)
Episcynia inornata (Orbigny)
Sansonia tuberculata (Watson)
Parciturboides interruptus (C. B. Adams)
1969
Radwin: Payardi Island Mollusks
2 3 1
Triphoridae
Tripbora turristhomae Holten
Triphora nigrocincta C. B. Adams
Clrithiopsidae
Ccritbiopsis emersoni (C. B. Adams)
Ccritbiopsis greeni (C. B. Adams)
Seila adamsi H. C. Lea
Epitoniidae
Epitonhim foliaceicostum Orbigny
Opalia crenata (Linne)
Depressiscala nautlae (Morch)
EULIMIDAE
Enlima bifasciata (Orbigny)
Balcis intermedia (Cantraine)
FOSSARIDAE
Megalomphalus pilsbryi Olsson & McGinty
Calyptraeidae
Calyptraea centralis (Conrad)
"'Crucibuliim auricula m (Gmelin)
Crepidula plana Say
Crepidula maculosa Conrad
Crepidula con vex a Say
Xenophoridae
'■'Xenophora concbyliopbora (Born)
Strombidae
''Strombus pugilis Linne
Strombus raninus Gmelin
Eratoidae
Trivia pediculus (Linne)
Trivia quadripunctata (Gray)
Erato maugeriae Gray
Cypraeidae
Cypraea cinerea Linne
''Cypraea zebra Linne
Ovulidae
Cyphoma gibbosa (Linne)
Naticidae
"'Natica canrena (Linne)
Tectonatica Pusilla (Say)
!tPolinices lacteus (Guilding)
*Sinum macula/um (Say)
Glyphepithema floridana Render
Cassididae
Cassis flammea Linne
''Morum oniscus (Linne)
Pbaliuin granulatum (Born)
''Cypraecassis testiculus (Linne)
Cymatiidae
Charonia variegata (Lamarck)
Cyniatium caribbaen m Clench & Turner
Cymatium labiosum (Wood)
*Cymatium femorale (Linne)
''Cymatium muricinuin (Roding)
'Cymatium pile are (Linne)
Cymatium nicobaricum (Roding)
Cymatium ve space um (Lamarck)
Bursidae
Bursa corrugata (Perry)
Bursa cubamana (Orbigny)
TONNIDAE
Tonna maculosa Dillwyn
Murk idai
Murex woodringi Clench & Farfante
Murex rubidm V. C. Baker
Chicoreus florifer (Reeve)
Murii opsu oxytatm \\. Smith
Muricopsis pbilippiana | Dall)
Khomurex scbrammi (Crosse)
Risomurex muricoides (C. B. Adams)
Morula nodulosa (C. B. Adams)
Morula didyma (Schwengel)
Thais deltoidea (Lamarck)
Magilidai
Coralliophila deformh (Lamarck)
Coralliophila abbreviata (Lamarck)
COLUMBI I I IDAI
"Anachis (Parvanacbis) obesa (C. B. Adams)
Anachis (Suturoglypta) pre/ri (Duclos)
'■'Astyris lunata (Say)
Coin mbella mercatoria (Linne)
Conella in uloides (C. B. Adams)
Cosmioconcha nitens (C. B. Adams)
Nitidella nit id a (Lamarck)
*Steironepion monilifera (Sowerby)
Zafrona pulcbella (Blainville)
PlSANIIDAE
Engina turbinclla Kiener
''Bailya intricata (Dall)
"Cadncifer" adelus (Schwengel)
Cantharus auritulus (Link)
Pisania pusio (Linne)
Colubrariidae
"'Colubraria testacea (Morch)
'Colubraria lanceolata (Menke)
Melongenidae
''Melongena melon gena (Linne)
Nassariidae
Nassarius cinisculns (Reeve)
Fasciolariidae
Easciolaria tulipa (Linne)
Fusilatirus cayobuesonicm (Sowerby)
Latirus infundibulum (Gmelin)
Latirus carinifera (Lamarck)
*Leucozonia nassa (Gmelin)
Leucozonia ocellata (Gmelin)
TURBINEI I.IDA I
Turbinclla angulatus Lightfoot
*Vasiiin miiricatum (Born)
Olividae
Oliia caribaeenh V>.\\\ & Simpson
Olii ella mi ea Gmelin
Oliiella cbiriquiemis Olsson & McGinty
Minioliva myrmecoon (Dall)
MlTRlDAI
Mitra barbadensh (Gmelin)
Wi/ra nodulosa (Gmelin)
Pusia albocincta (Dall)
Pusia histrio | Rei
232
San Diego Society of Natural History
Vol. 15
VOLUTIDAE
Valuta virescens Lightfoot
Marginellidae
Vrunum carneum (Storer)
Volvarina avena (Valenciennes)
Gibberula bocaseiisis Olsson & McGinty
Gibberulina ovuliformh (Orbigny)
CONIDAE
Conns spit rilis spurius Linne
Conns regius Gmelin
Conns daucus Linne
Conns granulatus Linne
Conns largillierti Kiener
Conns pygmaea Reeve
■'Conns inns Hwass
Conns mind anus Hwass
Conns jaspidea Gmelin
Terebridae
''Terebra taurinum (Lightfoot)
Terebra spei Brown & Pilsbry
Turridae
Polystira albida (Perry)
"Drillia" albinodata Reeve
"Drillia" albomacnlata C. B. Adams
"Drillia" accstra Dall
Synfoinodrillia lissotropis (Dall)
Crassispira anberti Lamy
Crassispira barfordiana (Reeve)
Crassispira cbazaliei Dautzenberg
Nannodicllit oxytata (Bush)
Bracbycytbara biconica (C. B. Adams)
Itbycythara psila (Bush)
"Mangelia" filosa Rehder
Thelecythara floridana Fargo
Acmaturris sp.
Daphnella lyninaeiformis (Kiener)
OPISTHOBRANCHIATA
BULLIDAE
Bulla occidentals A. Adams
Retusidae
Volvulella recta (Morch)
Volt nlclla sp.
Cylichna bidentata (Orbigny)
Cylichna krcbsi (Morch)
Acteocina candei (Orbigny)
Atyidae
Atys guildingi (Sowerby)
Atys riisiana (Morch)
Pyramidellidae
Pyramidella Candida Morch
Triptycbus nil ens (Morch)
Odostomia gemmulosa (C. B. Adams)
Odostomia terryi Olsson & McGinty
Odostomia (Spiroclimax) sp.
Turbonilla pupoides (Orbigny)
Tnrbonilla sp. A.
Turbonilla sp. B.
Cavolinidae
"Carolina longirostris LeSueur
SCAPHOPODA
Dentaliidae
'■'Dental in m sowcrbyi Guilding
Dentalinin antillarum Orbigny
Dentaliuin tcxasiannm Philippi
■'Cadulns tctraschistus Watson
BIVALVIA
PROTOBRANCHIA
NUCULANIDAE
''Nucnlana acuta (Say)
Nuculana Sp.
^■Adrana nencombi (Angas)
NUCULIDAE
Nucula acgeenis Jeffreys
FILIBRANCHIA
Glycymeridae
Glycymcris pcctinata (Gmelin)
Arcidae
"'Area zebra Swainson
Area imbricata Bruguiere
Anadara notabilis (Rbding)
'' Anadara chemnitzi (Philippi)
Barbatia cancellaria (Lamarck)
"'Barbatia Candida (Helbling)
Arcopsis adamsi Dall
Pinnidae
Pinna earned Gmelin
Plicatulidae
■■Plicatula gibbosa (Lamarck)
Pectinidae
"Pecten" laurenti Gmelin
*"Pecten" muscosus Wood
"Pecten" fuscopurpureus Conrad
"Pecten" antillarum Recluz
'■''Nodipecten nodosus (Linne)
Chlamys sentis (Reeve)
Cblamys imbricata (Gmelin)
■' Argopecten nucleus (Born)
Ostreidae
"'Ostrca frons Linne
Mytilidae
'''Modiolus amcricanns (Leach)
Litbopbaga antillarum (Orbigny)
Botula fnsca (Gmelin)
IsOGNOMONIDAE
''hognoinon alatus (Gmelin)
Isognomon radiatus (Anton)
Spondylidae
■'Spondylus americanus Hermann
Limidae
Lima scabra Born
Lima tenera Sowerby
Anomiidae
''Pododesmus rudis (Broderip)
1(=Adrana perproiracta (Dall). See: van Regteren Altena. 1968. Basteria *2(l-3).
1969
Radwin: Pavardi Island Mollusks
233
EULAMELLIBRANCHIA
Crassatellidae
*Crassinella adamsi Olsson
CONDYLOCARDIIDAE
*Condylocardia bernardi Dall
Lucinidae
Lucina pensylvanica (Linne)
Lucina nassula Conrad
Lucina pectinatus (Gmclin)
''■'Lucina niuricatus (Spengler)
'■Codakia orbicularis (Linne)
''Codakia orbiculata (Montforc)
''Codakia pec find I a (C. B. Adams)
Chamidae
Chama congregata Conrad
Chama macerophylla Gmelin
Cardiidae
Lac vicar. I 'iniu mortoni (Conrad)
* Laei icardinin laevigatum (Linne)
Laeiicardium multilincata Dall & Simpson
'■'Trachycardium isocardia (Linne)
*Trachycardium muricatum (Linne)
Trigoniocardia antillarum (Orbigny)
* Americardia media (Linne)
Papyridea soleniformis (Bruguiere)
Veneridae
Cbione cancellata (Linne)
"'Chione paphia (Linne)
"'Macrocall'nta maculata (Linne)
Callocardia albida (Gmelin)
*Antigona listeri (Gray)
'■'Gouldia cerina (C. B. Adams)
*Dosinia discus (Reeve)
1*1 IKK .il [DAI
: ■ Kit pcllar u typica \ I
I II I I MDAE
' Tcllina listeri Roding
Tellina punicea Born
Una angulosa Gmelin
''Tellina martinicensh (Orbij
Scissiila exilis (Lamarck)
Macoma tenia (Say)
Arcopagia fausta (Pulteney)
Sanguinolariidai
Asaphis deflorata (Linne)
Solenidae
Solecurtm cumin gianui (Dunker)
Myidae
Sphenia antillensh Dall & Simpson
Semelidae
'■'Seinelc purpurascem (Gmelin)
'■'Semele proficua (Pulteney)
Abra lioica Dall
Corbulidai
Corbula rwiftiana C. B. Adams
Corbula dietziana C. B. Adams
Corbula cubaniana Orbigny
Corbula caribaea Orbigny
Notocorbula operculata (Philippi)
Montacutidae
Mont acuta percompressa Dall
SEPTIBRANCHIA
CUSPIDARIIDAE
Cardiomya costellata (Dcshayes)
Cardiomya perrostrata (Dall)
Discussion
The geological history of southern Central America and northern South America
is not completely understood. There is good evidence, however, that several major trans-
American seaways existed in this area between the Middle Cretaceous and late Pliocene
(Simpson, 1950: 3 63). The Atrato Strait (also known as the Bolivar Trough), in
northwestern Colombia, was probably the widest and most enduring of these (Woodrin-.
1966); it may have been the last major channel (Whitmore and Stewart, 1965: 185).
The time of closure of this strait, and concomitantly the completion of the land bridge,
is somewhat in dispute. Woodring (/';/ //'//.) indicates that it could not have occurred
before Pleistocene. Some paleontologists, however, have suggested an earlier, Miocene
time of closure (Olsson, 1932: 39, 42; Durham and Allison, 1960: 67). Based on
mammal evidence, most vertebrate paleontologists accept a late Pliocene date for this
event (Whitmore and Stewart, 1965: 185).
A comparison of the Payardi Island fauna with the most complete listings <>t
Panamic-Pacific species (Keen, 1958; Olsson, 1961) shows that at least S5 of the 2'>
Payardi Island species (29^y ) either occur in both the Caribbean and the eastern Pacific
or are represented in the eastern Pacific by species here considered cognate. | fable 1 ).
-Closely related and allopatric species have often been termed analogues, particularly in malacol
literature. However, because the term "analogy" has a widespread use in biology and implies similarity due
to convergence, not relationship, the term "analogous" tor closely related species is a potential soura
confusion. Webster's New World Dictionary defines "cognate" as "related through the same origin:
derived from a common original form." As this meaning is essentially the same
"analogue", I recommend that the latter term be dropped in favor oi ite".
234
San Diego Society of Natural History
Vol. IS
Many of the species considered as cognates in the present compilation have been pre-
viously suggested by Keen (195 8) and/or Olsson (1961). Other cognates were deter-
mined by comparing morphological features of Pacific and Caribbean species; in some
cases, figures in Olsson and Keen were utilized in the absence of specimens. In addition,
some eastern Pacific cognates have almost certainly been overlooked, especially in
families such as the Turridae and Pyramidellidae. Thus, the proportion of Payardi Island
species with eastern Pacific cognates is probably greater than the 29c/( indicated. This
percentage may be moderate to high if the idea of a Miocene or early Pliocene closure of
the Atrato Strait is accepted, or as quite low in terms of a late Pliocene or early Pleis-
tocene closure. Regardless of which view is taken, no zoogeographic or evolutionary
conclusions can be drawn from this percentage of cognates.
No similar comparison may be made between the only other southwestern Carib-
bean fauna studied (Olsson and McGinty, 195 8) and compilations of the Panamic-
Pacific faunas (Keen, 195 8; Olsson, 1961). Olsson and McGinty's paper deals solely
with shallow-water mollusks collected from a variety of habitats in a relatively broad
geographical area, on several occasions spanning more than thirty years. The present
list, in contrast, includes only species collected from a relatively restricted area by an
inherently more selective method. Thus, it is hardly surprising that these two Caribbean
lists, although dealing with faunas within 200 miles of each other, show many dis-
similarities.
Table 1
Closely Related Recent Species Occurring at Payardi Island
and in Tropical East Pacific Faunas
Payardi Is. Species
Hemitonia octoradiata (Gmelin)
Diodora cayenensis (Lamarck)
Anticlimax schumoi (Vanatta)
Astraea caelata (Gmelin)
Turbo castanea Gmelin
Neritina Virginia Linne
Ah ania auberiana (Orbigny)
Caecum pulchcllum Stimpson
Rissoina fischeri Desjardin
Architectonica nobilis Roding
Heliacus perrieri (Rochebrune)
S/epbopoma myrakeenae Olsson & McGinty
Modulus modulus Linne
Modulus carchedonius Lamarck
Cerithium lariabile C. B. Adams
Cerithium lilteratum Born
Liiiopa melanostoma Rang
Crucibulum auricula (Gmelin)
Xenophora conchyiiophora (Born)
Strombus pngilis Linne
Cypraea zebra Linne
Cyphoma gibbosa (Linne)
Natica canrena (Linne)
Volinices lacteus (Guilding)
Siniim macnla/um (Say)
Phalium grauiilatum (Born)
Morum oniscus (Linne)
Cypraecassh testiculm (Linne)
Eastern Pacific Cognate
H. bermosa (Lowe)
D. inequalis (Sowerby)
A. uilletti Hertlein & Strong
A. turbanica (Dall)
T. squamiger Reeve
N. luteofasciata Miller
A. inconspicua (C. B. Adams)
C. diminuta (C. B. Adams)
R. effusa Mbrch
A. nobilis Roding
H. infundibuliforinis (Gmelin)
S. pen nat ii in Mbrch
M. disculus Philippi
M. catenulatiis Philippi
C. nicaraguense Pilsbry & Lowe
C. maculosa m Kiener
L. melanostoma Rang
C. personal ii in Keen
X. robusta Verrill
S. gracilior Sowerby
C. ceri inetta Kierner
C. emarginata (Sowerby)
N. grayi Philippi
P. nber Valenciennes
S. debile Gould
P. centriqitadrata Valenciennes
M. tuberculosum Reeve
C. tenuis Wood
1969
Radwin: Payardi Island \K>i i usks
23 5
Table 1 (cont.)
Payardi Is. Species
Cymatium femorale (Linne)
Cymatium pileare (Linne)
"Cymatium muricinum (Roding)
Muricopsis oxytatus M. Smith
. \\h vis huiata (Say)
Anachh obesa (C. B. Adams)
Steironepion monilifera (Sowerby)
Bailya intricata (Dall)
Colubraria testacea (Morch)
Colubraria lanceolata (Menke)
Melongena melon genu (Linne)
Leucozonia nassa (Gmelin)
Vasum muricatum (Born)
Mitra nodulosa Gmelin
Terebra ttmrinum (Lightfoot)
Conns inns Hwass
Cat olina longirostris LeSueur
Dentalium soucrbyi Guilding
Cadidus tetraschistns Watson
Nuculana acuta (Say)
Area zebra Swainson
Anailara chemnitzi (Philippi)
Arcopsis adamsi Dall
Barbatia Candida (Helbling)
Plica/nla gibbosa Lamarck
"Pec/en" muscosits Wood
Nodi pec ten nodosns (Linne)
Argopecten nucleus (Born)
Ostrea frons Linne
hogrtomon alatus (Gmelin)
Spondylus americanus Hermann
Modiolus americanus (Leach)
Pododesmus ritdis (Broderip)
Crassinella adamsi Olsson
Condylocardia bernardi Dall
Lucina muricatus Spengler
Codakia pectinella (C. B. Adams)
Codakia orbicularis (Linne)
Codakia orbiculata (Montfort)
Trachycardium isocardia (Linne)
Trachycardium muricatum (Linne)
Americardia media (Linne)
Laei icardium laevigatum (Linne)
Papyridea soleniformis (Bruguiere)
Dosinia discus (Reeve)
Chione paphia (Linne)
Macrocallista maculata (Linne)
Antigona listcri (Gray)
Gouldia cerina (C. B. Adams)
Rupellaria typica (Jonas)
Tcllina listcri Roding
Tellina punicea Born
Tcllina angulosa Gmelin
Tellina martinicensis Orbigny
Scmele purpiirascens (Gmelin)
Semele proficua (Pulteney)
I \SI I UN PAI II [< ( U(.\ \ I I
C. tigrinum (Broderip)
C. pileare (Linnc)
-iC. muricinum (Roding)
Al. armatm (A. Adams)
A. barfordi Strung tc Hertlein
A. diminuta (C. B. Adams)
S. melanosticta Pilsbry & Lowe
B. anomala (Hinds)
C. lucasensh Strong &; Hertlein
C. siphonata (Reeve)
M. patula (Broderip & Sowerby)
L. ritdis (Reeve)
V. caestus Broderip
M. marshalli Bartsch
T. ornata Gray
C. mus Hwass
C longirostris LeSueur
D. innumerabile Pilsbry & Sharp
C. quadrifissatus (Carpenter)
N. hindsii (Hanley)
A. pacifica (Sowerby)
A. mix (Sowerby)
A. solida (Sowerby)
B. reeceana (Orbigny)
P. spondylopsis Rochebrune
"P." relcro Hertlein
N. subnodosiis (Sowerby)
A. circularis (Sowerby)
O. serra Dall
/. chemnitzianus (Orbigny)
S. princeps Sowerby
M. americanus (Leach)
P. cepio (Gray)
C. adamsi Olsson
C. pauainensis Olsson
L. liana Pilsbry
C. cancellaris Philippi
C. pinchoti Pilsbry & Lowe
C. gal a pagan a Dall
T. consors (Sowerby)
T. senticosum (Sowerby)
A. guanacastensis (Hertein & Strong)
L. clarionense (Hertlein & Strong)
P. aspersa (Sowerby)
D. ponderosa (Gray |
C. mariae (Orbigny)
M. scftialida (Sowerby I
A. multicast at a (Sowerby)
G californica Dall
K. robusta (Sowerby)
T . cumingi ( 1 [anl
T. simulaus C. B. Adams
T. eburna Hanley
T. proclivh Hertlein & Strong
S. sparsilittca/a Dall
s. lentiatlare (Sowerby)
'^Cymatium muricinum (Roding) has been collected in the Galapagos (Emerson, pers. cci
236 San Diego Society of Natural History Vol. 15
Acknowledgments
In addition to my obvious debt to Messrs. Stewart and Mena, I wish especially to thank Dr. Joseph
Rosewater, Division of Mollusks, U. S. National Museum, for providing facilities to study this fauna.
Most of the present research was undertaken while I was a Smithsonian Predoctoral Intern. Dr. Donald
R. Moore, Institute of Marine Sciences, University of Miami, kindly assisted in identifying the caecids,
vitrinellids, cyclostremids, and other difficult groups.
Thanks are also due the following individuals for reading preliminary drafts of this paper and pro-
viding constructive citicism: Drs. Joseph Rosewater, Harald A. Rehder, and Wendell P. Woodring, U. S.
National Museum; Dr. William K. Emerson, American Museum of Natural History; Dr. Reid Moran,
Messrs. Arnold Ross and Allan J. Sloan and Mrs. Fay H. Wolfson, San Diego Natural History Museum.
Figure 1 was prepared by Miss Gail Culver and Miss Anne Acevedo, San Diego Natural History
Museum.
Literature Cited
Durham, J. W. and E. C. Allison
1960. The geologic history of Baja California and its marine faunas. Syst. Zool. 9: 47-91.
Keen, A. M.
195 8. Sea Shells of Tropical West America. Stanford University Press, 624 p.
Olsson, A. A.
1932. Contributions to the Tertiary paleontology of northern Peru. Pt. 5, The Peruvian Miocene.
Bull. Amer. Paleontol. 9: 1-272.
1961. Mollusks of the Tropical Eastern Pacific; Panamic-Pacific Pelecypoda. Paleont. Res. Inst.,
Ithaca, New York, 5 74 p.
Olsson, A. A. and T. L. McGinty
195 8. Recent marine mollusks from the Caribbean coast of Panama with description of some new
genera and species. Bull. Amer. Paleontol. 39: 5-5 8.
Simpson, G. G.
1950. History of the fauna of Latin America. American Scientist 38: 361-389.
Whitmore, F. C, Jr. and R. H. Stewart
1965. Miocene Mammals and Central American Seaways. Science, 148: 180-185.
Woodring, W. P.
1966. The Panama land bridge as a sea barrier. Proc. Amer. Philos. Soc. 110: 425-433,
Accepted for Publication 10 May 1969
Department of Marine Invertebrates, Natural History Museum, P. O. Box 1390,
San Diego. California 92112.
MUS. COMP. ZOOL.
LIBRARY
f
OCT 30 1969
1
HARVARD
UNIVERSITY.
STUDIES ON THE TETRACLITIDAE
(CIRRIPEDIA: THORACICA):
REVISION OF TETRACLITA
ARNOLD ROSS
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 15 22 SEPTEMBER 1969
STUDIES ON THE TETRACLITIDAE
(CIRRIPEDIA: THORACICA):
REVISION OF TETRACLITA
ARNOLD ROSS
ABSTRACT. — Tesseropora and Tetraclitella. subgenera of Telraclita, are elevated to full generic status ir
the absence of intergrading major characters. Newmanella gen. nov. is proposed for Telraclita (Telraclita
radiata (Bruguiere, 1789) and Tesseroplax gen. nov. for Telraclita ( Tesseropora ) unisemita Zullo. 1968. On th<
basis of shell characters the five genera here recognized fall into two groups. Monometric growth, allometry 01
the carina, and non-tubiferous radii characterize Tesseropora, Tesseroplax, and Telraclita; diametric growth
isometry of the wall plates, and tubiferous radii characterize Tetraclitella and Newmanella. Both Tesseropla.\
and Tetraclita are considered later derivatives from the Tesseropora stock; but Tetraclitella and Newmanellc
from a tesseroporan precursor. New distributional records for Newmanella radiata include the Bahamas
Puerto Rico, Dominica, and Venezuela. Apparently the antenniform ramus of cirrus III in N. radiata play;
some role during the reproductive cycle, but its exact function is unknown.
RESUMEN. — Tesseropora y Tetraclitella, subgenero de Tetraclita. estan elevados al estado generico en a
ausencia de intergradacion caracteres mayores. Newmanella gen. nov. esta propuesto para Tetraclitc
(Tetraclita I radiata (Bruguiere, 1789) y Tesseroplax gen. nov. para Tetraclita I Tesseropora ) unisemita Zullo
1968. En el base de caracteres de concha los cincogeneros reconocidos aquT, caen en dos grupos. Crecimientc
monometrico, alometria de la carena, y radios no-tubiferoso caracterizan Tesseropora. Tesseroplax. \
Tetraclita; crecimiento diametrico, isometrfa de la laminas de las paredes, y radios tubiferoso caracterizar
Tetraclitella y Newmanella. Los dos Tesseroplax y Tetraclita son considerados derivatives mas tarde de lo;
progenitores de Tesseropora, pero Tetraclitella y Newmanella de un progenitor de Tesseropora. Registro;
distribucionales nuevos por Newmanella radiata se incluyen las Bahamas, Puerto Rico, Dominica, >
Venezuela. Aparentemente, la ramal antenniforma de cirrus III en N. radiata hace un papel durante lo*
reproducti vos, pero su funcion exacto es desconocido.
The Tetraclitidae comprise a group of intertidal balanomorph barnacles occurring in
tropical and warm-temperature waters between latitudes 38° north and 52° south. Although!
these barnacles are a major component of intertidal faunas they have not received adequate
attention either taxonomically or biologically. In this, the first in a series of papers covering
the taxonomy and general biology of the tetraclitids, I re-evaluate the status of Tetraclita
and its subgenera, and propose a new classification in the light of recent studies (Ross, 1968),
and the work of Darwin ( 1 854), Nilsson-Cantell ( 1 92 1 ) and Hiro ( 1 939).
There are more than two dozen named taxa distributed presently between Tetraclita
Schumacher, 1817 (type species: T. (Tetraclita) squamosa (Bruguiere), 1789), and its
subgenera, Tesseropora Pi lsbry, 1916 (type species: T (Tesseropora) rosea (Krauss), 1848),
and Tetraclitella Hiro, 1939 (type species: T (Tetraclitella) purpurascens (Wood). 1815).
These subgenera are distinct, and their elevation to generic status clarifies relationships that
are obscured when all of the species are considered monogeneric.
Both Darwin (1854: 344) and Hiro (1939: 270) cited numerous morphological
characters that support the generic distinctness of the Caribbean western Atlantic species. /
radiata. These characters, coupled with the absence of intergrading major morphological
structures, necessitate the proposal of Newmanella gen. nov. for this species. Attempts to
assign the recently described Pliocene species from the Gulf of California, Mexico. T
(Tesseropora) unisemita (Zullo, 1968: 273), within the proposed classification posed
innumerable problems, and consequently I have placed it in a new genus, Tesseroplax.
SAN DIEGO SOC. NAT. HIST. TRANS 15(15): 237-251, 22 SEPTI MM R 1969
238 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
FAMILY TETRACLITIDAE Gruvel
Tetracaines Gruvel, 1903: 160; Nilsson-Cantell, 1921: 357, nom. transl.; Ross, 1968: 6,
nom. transl.
Definition. - Balanomorpha with shell of 4 plates; rostrum compound; compartments
demarcated along sutures on interior surface of wall when fused or corroded externally;
parietes with one or more rows of separate or confluent tubes containing living tissue or
secondarily filled with calcareous and chitinous material; overlapping plates with radii. Basis
membranous or calcareous and not forming complex interdigitations with wall. Inferior
margin of mandible pectinate or serrate, never molariform. Labrum not bullate; crest with
or without shallow notch, never incised. Cirrus II and III commonly armed with bipinnate
and other complex setae distributed along anterior curvature; cirrus III resembling II more
than IV; inner or outer ramus commonly antenniform during reproductive cycle. Basidorsal
point on intromittent organ, and caudal appendages absent. Occurring in intertidal zone,
generally on inanimate objects. Type genus: Tetraclita Schumacher, 1817.
Remarks. — Authorship of this family must be accorded Gruvel ( 1 903: 1 60) rather than
Nilsson-Cantell (1921: 357) as earlier noted (Ross, 1968: 6), and the date of authorship
accepted as 1903, because "A family group name of which the suffix is incorrect is available
with its original date and authorship, but in properly emended form" (Article 1 1 (e) (ii),
ICZN).
INTRAFAMILIAL RELATIONSHIPS
In the Tetraclitidae shell growth proceeds either diametrically or monometrically
(Darwin, 1854: 324). In monometric growth there is a direct correlation between
obsolescence of the radii, which are nontubiferous, secondary apical filling of the parietal
tubes, and enlargement of the orifice by attrition or corrosion. In diametric growth
enlargement of the orifice results directly from growth of the tubiferous radii normal to the
parietes without wearing away the peritreme, and the parietal tubes are not secondarily
filled.
These modes of shell growth allow separation of the tetraclitids into two groups (Fig. 1).
Monometric growth characterizes Tesseropora, Tesseroplax and Tetraclita, diametric
growth Tetraclitella and Newmanella. The latter method is the phylogenetically more
primitive based upon the fossil record.
Equally important in segregating the tesseroporan line from the tetraclitellan is the
allometry of the carina that occurs commonly in the former group but not the latter. In the
tetraclitellans all of the plates develop essentially isometrically. Allometry of the carina and
adjacent laterals is a somewhat recent departure from the isometry characteristic of early
balanomorphans and is always directly associated with monometric shell growth. Con-
versely, isometry of the wall plates occurs concomitantly with diametric growth.
Enlargement of the orifice in the tesseroporans, which have monometric shell growth,
results from attrition or corrosion of the peritreme, and this is facilitated by their occupancy
of a poorly protected high intertidal habitat. In the tetraclitellans enlargement of the orifice
is by diametric growth, and this group occurs in relatively well protected habitats low in the
intertidal zone. In both groups, the parallel development of a multilayered wall probably
confers a selective advantage in that it provides protection against rapacious gastropods and
other boring predators.
Tetraclita was probably derived from Tesseropora, which retains a single row of
parietal tubes. Henry (1957: 36) showed that secondary longitudinal tubules in some
individuals of Tesseropora pacifica (Pilsbry) is "the first step in the formation of a multilayer
wall." She assigned this species to Tetraclita (sensu stricto) because of the presence of
1 %l)
ROSS: STUDIES ON THE TETRACLI ill) \l
239
Figure 1 . Inferred phylogeny and generic affinities in the Tetraclitidae. Peripheral illustrations portray aspects of
shell growth and morphology, and key characters used in distinguishing the two groups in this family.
240 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
secondary tubules. However, this assignment is contrary to the original definition of the
genus; the secondary tubules are not comparable to those of Tetraclita, because there is but
one complete row in this species and in certain crowded specimens of T. rosea (see Zullo,
1968:272).
Tesserop lax also has but one row of parietal tubes, and therefore is more closely related
to Tesseropora than to Tetraclita. I regard the absence of depressor muscle crests on the
scutum and removal of the tergal spur from the basiscutal angle to be primitive characters in
Tesseroplax as well as in Tetraclitella. The septate parietal tubes in Tesseroplax represent a
unique specialization for this family, and their function remains unknown, although they
may strengthen the wall. Tesseroplax probably represents an early off-shoot from the
tesseroporan line, as Zullo (1968: 274) suggested, and it is evidently an evolutionary
terminus, since none of the presently recognized species can be shown to have evolved from
T. unisemita. The morphological evidence in support of Zullo's alternate hypothesis that T.
unisemita may be the result of "convergence on the tetraclitan shell plan from an unrelated
ancestor," is weak. The narrow opercular plates in Tesseroplax, especially the tergum, may
be a direct consequence of monometric growth and allometry in this and other tesseropo-
rans. These attributes strengthen the inference that Tesseroplax is in the direct line of descent
from Tesseropora.
Tetraclitella and Newmanella appear to have been derived independently from a
tesseroporan precursor, the shell of which had a single row of parietal tubes and developed
diametrically. The presence in these genera of two or more rows of parietal tubes, as in
Tetraclita, I consider a later development which may function as defense against boring
predators.
Since each of these tetraclitellans possesses several primitive morphological characters,
it is not possible to deduce which genus is more primitive phylogenetically. The scutum in
Tetraclitella, although elongated, is still basically triangular, and it lacks crests for the
insertion of the lateral and rostral depressor muscles. The same is true in Tesseroplax. The
wall plates in Tetraclitella are weakly articulated and do not develop the complex occlusial
surface found in Newmanella. On the other hand, the tubes in the radii of Tetraclitella, to
judge from other balanomorphs (e.g., Megabalanus, Emersonius, Platylepas) reflect a
highly advanced and complex mode of formation.
KEY TO GENERA OF TETRACLITIDAE
1 . Growth of shell monometric; radii solid 2
1 . Growth of shell diametric; radii tubiferous 4
2. Parietes with one row of tubes 3
2. Parietes with two or more rows of tubes Tetraclita
3. Parietal tubes lacking transverse septa; scutum bearing depressor
muscle crests Tesseropora
3. Parietal tubes bearing transverse septa; scutum lacking depressor
muscle crests Tesseroplax gen. nov.
4. Radii with horizontal summits, articular margins lacking teeth;
scutum transversely elongated, lacking depressor
muscle crests Tetraclitella
4. Radii with oblique summits, articular margins bearing prominent
teeth; scutum triangular, bearing depressor
muscle crests Newmanella gen. nov.
1969
ROSS: STUDIES ON THE TETRACLITIDAE
241
Tesseroplax gen. nov.
Definition. --Shell moderately large, smooth; compartments discrete; parietes with
one row of rectangular tubes secondarily filled apically, septate basally; radii narrow, non-
tubiferous; basis calcareous, with transverse septate tubes; scutum triangular, lacking crests
for depressor muscles; tergum narrow, with spur separated from basi-scutal angle.
Type species. — Tetraclita ( Tesseropora ) unisemita Zullo, 1968.
Etymology. Derived from the Greek, tesseres, four, and plax, plate, in reference to
the number of parietal plates.
Remarks. - - Tesseroplax unisemita is known only from two specimens collected from
Pliocene sediments on Angel de la Guarda Island in the Gulf of California, Mexico. Many
supposedly unique characters were attributed to this extinct species; but the absence of
depressor muscle crests on the scutum is typical of Tetraclitella; internal parietal ribs also
occur in Newmanella; and the apical filling of the parietal tubes is a characteristic of
Tetraclita as well as Tesseropora. However, Tesseroplax is the only tesseroporan possessing
septate parietal tubes and a calcareous basis also with septate tubes. These structures as well
as internal parietal ribs probably reflect a unique method of shell deposition in this lineage.
Tesseroplax is also the only tesseroporan in which the scutum lacks depressor muscle crests,
and the spur of the tergum is infolded as well as being extremely narrow and elongate.
Figure 2. Newmanella radiata, from test panel set at 1 5 m. Vieques Sound, Puerto Rico. a. contiguous specimens
viewed from left side, actual height of upper specimen 23.1 mm; b. c. external and internal views, respective:
articulated opercular plates, actual height of scutum 1 3.9 mm. tergum 9.5 mm.
242 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
Newmanella gen. nov.
Definition. - Shell relatively large, conic, ribbed; compartments discrete; parietes with
2 or more rows of irregularly shaped tubes; radii broad, summits oblique, tubes opening on
articular surface; basis calcareous, solid, scutum triangular, bearing crests for depressor
muscles; tergum narrow with spur separated from basiscutal angle; mandible with 4 teeth,
basal comb, and spine-like lower extremity; maxilla I with more than 12 spines below
subapical notch.
Type species. — Balanus radiata Bruguiere, 1789.
Etymology. -- Named in honor of Dr. William A. Newman, Scripps Institution of
Oceanography, longtime friend, and student of the Cirripedia.
Remarks. - - The presence of more than one row of parietal tubes and well developed
tubiferous radii serve to distinguish Newmanella from the tesseroporan complex. The radial
tubes in Newmanella differ from those of Tetraclitella by being irregular in outline and of
varying size and by their mode of formation, which results from development of prominent
irregular ridges on the sutural surface that branch and coalesce. The radial tubes in
Tetraclitella are uniform in size and outline and develop in a manner comparable to that of
the parietal tubes. Initially growth is normal to that of the parietes, but with subsequent
growth and sequential development of additional tubes along the outer lamina of the radius,
the earlier formed radial tubes curve and may extend downward to the base.
Newmanella radiata (Bruguiere)
[Lepas Indiae orientalis ex violaceo radiata] Chemnitz, in Martini and Chemnitz, 1785: 319, pi. 99, fig. 842.
Balanus radiata Bruguiere, 1 789: 1 68; Bruguiere, 1 79 1 , pi. 1 64, figs. 5, 5a.
Lepas purpurea Spengler, 1790: 172.
Lepas violacea Gmeiin, 1791: 3213, not Balanus violaceous Gruvel, 1903 [= Balanus abeli Lamy and Andre, 1932:
218, footnote].
Balanus radiatus: Ranzani, 1818: 75; Lamarck, 1818: 393; Ranzani, 1820: 39; Jay, 1839: 7; Lamy and Andre, 1932:
218.
Conia radiata: Blainville, 1824: 378; Blainville, 1825: 598; Blainville, 1827, pi. 85, figs. 5, 5a; Deshayes, 1831:357.
T[etraclita](C[onia}) radiata: Gray, 1825: 104.
Conia lyonsiil ex Leach, MSjSowerby, 1823, no pagination.
Tetraclita radiata: Darwin, 1854:343, pi. 1 1, figs. 5a-5d; Weltner, 1897: 258; Gruvel, 1903: 161; Gruvel, 1905:291;
Schmalz, 1906: 65, pi. 6, fig. 4; Hoek, 1907: xvi; Pilsbry, 1916: 259, pi. 61, figs. 3-3c, 4; Pilsbry, 1927: 38;
Nilsson-Cantell, 1939: 5; Pope, 1943: 244; Pope, 1945: 368; Pilsbry, 1953: 27; Southward, 1962: 163; Ross,
1968: 18.
Material. Turtle Rocks, south of Bimini, Bahamas, B.W.I., approximately
25°40'N., 79°20'W.; intertidal on exposed hull of wrecked ship; E. Kirsteuer coll., August
1967; 5 specimens.
Vieques Sound, Puerto Rico, approximately 18°12'N., 65°25'W.; on fouling test panel
set at 1 5m below surface; U.S. Naval Oceanographic Office, October 1 966; 4 specimens.
Vieques Island, Puerto Rico; intertidal; J. A. Rivera and C. B. Rivera coll., November
17, 1951; 10 specimens.
Scotts Head Bay, Dominica, approximately 15°21'40"N., 61 °22'40"W.; intertidal on
Tetraclita stalactifera (Lamarck); E. Kirsteuer and K. Rutzler coll., May 17-28, 1966; 1
specimen.
Isla de Margarita, Venezuela, approximately 10°45'N., 64°52'W.; intertidal; P. Glynn
coll., May 28, 1968; 7 specimens.
Gulfof Paria, Trinidad, approximately 10°12'N., 61°52'W.; on marine structures in the
Shell Trinidad Ltd. oilfield; R. Bacon coll., May 30, 1968; 10 specimens.
Supplementary Description. Shell white; low-conic; ribs on parietes numerous,
1969
ROSS: STUDIES ON THE TETRACLITID A I
243
approximate, narrow, prominent, generally branched basally (Fig. 2a). Orifice large,
trigonal to pentagonal; peritreme slightly toothed. Radii broad; horizontally striated:
summits oblique, at 45° or less; ridges on articular margin irregular, with interspaces
forming tubes opening on margin. Alae with summits less oblique than radii; articular
margin toothed horizontally. Wall of body cavity with regular longitudinal ribs. Fancies
with more than 1 row of irregularly spaced large and small tubes, without definite pattern;
longitudinal septa thick, crenated basally. Shell measurements are given in Table 3 o\' the
Appendix.
Basis calcareous; translucent; thin centrally, thickening peripherally.
Scutum triangular; articular ridge high, effectively erect, evenly rounded terminally.
cde
abm
h if - PUERTO RICO
degjklm- Bahamas
abc - TRINIDAD
Figure 3. Newmanella radiata. trophi. a, b, mandible: c-e, maxilla I: f, maxilla II: g-k. crest of labrum: 1 . enlarged
view of crest of labrum in m; m, labrum and palps.
244 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
about 2/3 length of margin; articular furrow extremely broad and deep; adductor muscle
ridge thin, relatively short, apically confluent with articular ridge; adductor muscle
depression shallow, poorly defined, crests for lateral and rostral depressor muscles few in
number, low, short.
Tergum triangular; longitudinal furrow broad, shallow; spur fasciole open; spur
obliquely rounded, separated from basiscutal angle by less than its own width; articular ridge
high, projecting and covering apical 1/3 of scutum when articulated (Fig. 2c); articular
furrow broad and deep; crests for depressor muscles numerous, low, regularly spaced.
Crest of labrum with broad V-shaped medial notch either smooth or laterally toothed;
teeth on crest either simple, M-shaped or comb-like, numbering commonly 3-4, ranging 2-7
(Figs. 3g-l); superolateral margins of labrum finely serrate or toothed; interspersed among
teeth, in notch, and extending laterally along crest of labrum are short, soft bristles. Palps
elongate, rectangular; basal margins free of setae; superior margin clothed with short,
broad, bipectinate setae; inner lateral face clothed completely with ctenae; distal extremity
bearing long, slender, bipinnate setae. Cutting edge of mandible armed with 5 teeth including
inferior angle; teeth 2-4 support subsidiary cusps; superior slope of tooth 4 serrate; inferior
angle bears 3-4 acicular teeth; comb between tooth 4 and inferior angle contains 17-20
acicular teeth (Figs. 3a-b). Maxilla I with deep, U-shaped notch, rarely without notch;
spination along cutting edge in three functional clusters; 2 long, stout spines above notch and
6-9 short, slender spines above or running into notch; 13-20 long, slender spines medially,
grading into basal zone of 7-13 short, slender spines. Maxilla II bilobate, taller than broad;
setae of apical lobe long, bipinnate, but shorter and bipectinate on lower lobe; region of
juncture of two lobes covered with short, narrow spines and ctenae.
Posterior ramus of cirrus I about 2/3 or less length of anterior ramus; intermediate
articles of both rami broader than high; segments of anterior ramus normal, posterior
protuberant (Fig. 4a). Rami of cirrus II either essentially equal in length or posterior ramus
antenniform (Fig. 4b); terminal segments of anterior ramus armed with few bipinnate setae;
proximal and intermediate articles of both rami bullate when cirrus normal. Rami of cirrus
III either equal in length or posterior ramus antenniform; when rami are equal, lesser
curvature of anterior ramus not covered with spines, and setae are bipectinate; when rami are
unequal in length lesser curvature of intermediate and basal segments of both rami covered
with short triangular or hook-like spines, and setae bipectinate only on anterior ramus. Cirri
IV-VI essentially equal in length with equal rami. Anterior face of posterior ramus of cirrus
IV covered with short, triangular spines; proximal segments of both rami along posterior
face also spinose. Proximal segments of cirri V-VI covered with short spines along greater
curvature only. Chaetotaxis of intermediate articles of cirri IV-VI ctenopod along anterior
curvature, with 4 pairs; between or at bases of apical pair are 1-2 short slender setae. Cirral
counts are summarized in Table 1 of the Appendix.
Intromittent organ distinctly annulated throughout its length; sparsely hirsute except
for 2 distinct clusters surrounding terminal orifice (Fig. 4e).
Remarks. - Newmanella radiata is apparently limited to the Caribbean, having been
reported from Florida (Pilsbry, 1957: 27), St. Thomas, Virgin Islands (Pilsbry, 1916: 259),
and Monos Island, Trinidad (Southward, 1962: 163). New records, which considerably fill
gaps in this species distribution include the Bahamas, Puerto Rico, Dominica, and
Venezuela.
FUNCTIONAL SIGNIFICANCE OF ANTENNIFORMY
In some individuals both rami of cirrus III are normal, but in others one is normal and
one antenniform (setae arranged in whorls at each articulation and the segments articulated
1969
ROSS: STUDIES ON THE TETRACLITIDAE
245
Figure 4. Newmanella radiata, cirral appendages, a, left cirrus I; b, left cirrus II with antenniform posterior
ramus; c, left cirrus III with antenniform posterior ramus; d, proximal segments of right cirrus III; e, distal end of
intromittent organ; f, normal left cirrus II; g, basal segments of left cirrus VI. pedicel, and proximal portion of
intromittent organ: h, normal left cirrus III.
246 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
so that the ramus can be rotated through 360°). Antenniformy is apparently not widespread
in the Chthamalidae, and is unknown in the Balanidae. Of the tetraclitid genera recognized
herein only Tetraclitella, to my knowledge, does not exhibit antenniformy, and it is
apparently rare in Tesseropora.
Darwin (1854: 83) noted variability in the occurrence of an antenniform ramus in
chthamalids, and stated that the ramus "acts as an organ of touch." In the deep sea
chthamalid Hexelasma hirsutum (Hoek), Southward and Southward (1958: 642) found the
antenniform ramus to be less sensitive to touch than rami of the other cirri. They noted also
that the antenniform rami do not form part of the food gathering cirral net, which would be
precluded by the nature of its armament, and speculated that the rami might function as
direction indicators, permitting alignment of the net with prevailing food-laden currents.
This seems unlikely because individuals with antenniform rami also occur in intertidal
habitats, where there is no obvious adaptive value for such an organ.
Pope (1965: 59) detected no correlation between the presence or absence of an
antenniform ramus and age, seasonal or environmental variables. However, seventy per cent
of the eastern Australian population of Chthamalus antennatus Darwin that she studied, had
antenniform rami, and 80 per cent of the total population were brooding. Specimens
collected three weeks later from the same area had normal rami, but only 20-35 per cent still
contained developing nauplii. At no time did all individuals in the population have
antenniform rami.
In Newmanella, the May specimens from Venezuela had antenniform rami and were
brooding nauplii, but the opposite was true of specimens collected from Dominica and
Trinidad, except for one individual from Trinidad that had antenniform rami. Specimens
taken in August from the Bahamas and in October and November from two localities off
Puerto Rico also lacked antenniform rami and eggs or nauplii in the mantle cavity (Fig. 5).
Development of the antenniform ramus in N. radiata probably begins in February or March,
and the appendage is shed in April or May only to be replaced by a normal ramus.
Specimens of Tetrachthamalus oblitteratus from Mauritius were found to have normal
rami, and one individual was brooding; those from Elat, and all but one from Seychelles, had
antenniform rami, and both populations were brooding eggs or nauplii (Newman, 1 967: 430;
pers. comm.). Newman also observed that individuals with antenniform rami either lack or
have a diminutive intromittent organ, whereas individuals with normal rami have a normally
developed intromittent organ. In Tesseropora pacifica from Guam I have observed the
simultaneous presence of an antenniform ramus, reduced intromittent organ, and eggs in the
mantle cavity.
According to Crisp and Patel (1958: 1078) copulation and fertilization commonly
follow within a few days of ecdysis. However, ecdysis early in the reproductive cycle
normally results in loss of the recently oviposited eggs, because they lie in the mantle cavity,
the lining of which is molted with exuviae of the appendages, intromittent organ, prosoma,
esophagus, and rectum (Darwin, 1854: 157). Consequently, newly fertilized individuals
normally cease molting during the brooding period, which lasts for several weeks. But, if the
molting process is resumed after fertilization but prior to naupliar release, the mantle lining
is retained and complete exuviation either accompanies or follows liberation of the nauplii
(Patel and Crisp, 1961: 103). In Balanus balanoides Linnaeus the intromittent organ is lost
during exuviation after the time of fertilization, whether the individuals have been fertilized
or not (Crisp and Patel, 1960: 33). A new one gradually develops during the summer,
reaching a normal length prior to the onset of the breeding season, which in B. balanoides
occurs annually in November or December; development of the intromittent organ in
Tetrachthamalus and Tesseropora pacifica probably follows a similar pattern as does the
1969
ROSS: STUDIES ON THE TETRACLITIDAE
247
32
30
28
26
c/)24
^22
LU
^20
CD
LU -in
if) IO
. 16
O
214
212
<
LiJ 10
8
6
ANTERIOR RAMUS
24
22
20
18
16
14
12
10
8
6
4
2
• tw"^
— /
IV
V
VI
\
32
30
28
26
24
22
20
18
16
14
12
10
8
6
POSTERIOR RAMUS
CARINOROSTRAL
DIAMETER
OF SHELL
HEIGHT OF
SHELL
IV
V VI
24
22
20
18
16
14
12
10
"!"
■ ■
8
6
4
j.
2
PUERTO RICO
IIIIIIIIHIIIIHIIIIIIIIIIIIIIIIIIIIItllUltllllllllMIIIIIIIIHIIIII
BAHAMAS
TRINIDAD
VENEZUELA
OCT. 66
AUG. 67
MAY 68
MAY 68
Figure 5. Graph of mean values from Table 1 for cirral counts of anterior ramus (top left) and posterior ram"s
(top right), and from Table 3 for carinorostral diameter of shell (bottom left) and he.ght of shell (bottom right) 0
Newmanella radiala. Vertical line = range; horizontal bar = mean.
248 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
antenniform ramus.
Thus, there is a strong correlation between the reproductive period and the presence of
antenniform rami. In all cases, antenniform rami are noted first either immediately before,
during, or immediately after the reproductive period, which may occur more than once a
year in the chthamalids and tetraclitids. The presence of an antenniform ramus has not been
reported in individuals long after termination of the reproductive cycle.
In cross-fertilizing intertidal balanomorphs copulation takes place while the animals
are submerged. Some means of indicating copulatory availability would be advantageous
during this phase of the reproductive cycle, because these animals are monoecious and every
individual in the population is potentially capable of being fertilized by another nearby
individual. Since the periods of submergence are usually brief, some adaptation would be
effective in insuring rapid selection of a responsive individual. The antenniform ramus may
elicit or detect tactile or chemical stimuli from an individual acting the role of the opposite
sex.
The correlation between brooding, antenniformy, and penis development is com-
plicated. However, it appears that individuals brooding eggs or nauplii, having antenniform
rami, and lacking or having diminutive penes assume the morphology and act the role of a
female; those individuals without eggs or nauplii in mantle cavity, and with normal rami and
penes constitute the post-reproductive segment of the population, having reverted to the
normal hermaphroditic state. Resumption of the hermaphroditic state is necessary because
these animals will cross-fertilize during the next breeding season. Loss or diminution of the
intromittent organ after fertilization is not comparable to the loss of the antenniform ramus
after release of the nauplii, because the ramus is replaced by a functional, feeding appendage,
whereas the intromittent organ remains non-functional until the onset of the following
breeding season.
ACKNOWLEDGMENTS
I am indebted to Dr. William A. Newman, Scripps Institution of Oceanography, for his continued
encouragement, loan of specimens, and valuable comments on an earlier draft of the manuscript. I thank Ernst
Kirsteuer, American Museum of Natural History, John R. DePalma, U.S. Naval Oceanographic Office, Elizabeth
Pope, the Australian Museum, Huzio Utinomi, Seto Marine Biological Laboratory, and Peter Bacon, University
of the West Indies, Trinidad, for their assistance in providing me with specimens. Miss Gail Culver prepared figures
1 and 2, and Miss Anne Acevedo figures 3 through 5.
LITERATURE CITED
Blainville, H. M. de
1824. Nematopoda. Diet. Sci. Nat. 32 (Mollusques- Morfil). Strasbourg, F.G. Levrault. 567 p.
1825-1827. Manual de malacologie et de conchyliologie. Paris, F. G. Levrault. 664 p. (vol. 1, text, 1825), 87
pis. (vol. 2, plates, 1827).
Bruguiere, M.
1789-1791. BALANITE, lepas; Linn. Encyclopediemethodique: Histoirenaturelledes Vers: 1 (1): 158-173
(text, 1789; plates, 1791).
Crisp, J. D., and B. S. Patel
1958. Relation between breeding and ecdysis in cirripedes. Nature 181: 1078-1079.
1960. The moulting cycle in Balanus balanoides L.Biol. Bull. 1 18(1): 31-47.
Darwin, C. R.
1854. A monograph on the sub-class Cirripedia. The Balanidae, the Verrucidae, etc. London, Ray Society.
684 p.
Deshayes, G. P.
1831. CONIE. Conia. Encyclopediemethodique: Histoire naturelledes Vers 2 (2): 257-258.
1969 ROSS: STUDIES ON THE TETRACLITIDAE
249
Gmelin, J. F.
1 79 1 . Systema naturae per regna tri naturae . . . editio decima tertia. aucta. reformata. Leipzig I (6): 302 I -
3909 (Vermes).
Gray, J. E.
1825. A synopsis of the genera of cirripedes arranged in natural families, with a description of some new
species. Ann. Philos.. n.s. 10(2): 97-107.
Gruvel, A.
1903. Revision des cirrhipedes appartenant a la collection du Museum D'Histoire Naturelle. Nouv. Arch.
Mus. D'Hist. Nat., ser. 4(5): 95- 1 70.
1905. Monographic des cirrhipedes ou thecostraces. Paris, MassonetCie. 472 p.
Henry, D. P.
1957. Some littoral barnacles from the Tuamotu, Marshall, and Caroline Islands. Proc. U.S. Natl. Mus. 107
(3381): 25-38.
Hiro, F.
1939. Studies on thecirripedian fauna of Japan. III. Supplementary notes on the cirripeds found in the vicinity
of Seto. Mem. Coll. Sci., Kyoto Imp. Univ. ser. B. 1 5 (2): 237-284.
Hoek, P. P. C.
1907. TheCirripedia of the Siboga- Expedition. A. Cirripedia pedunculata. Siboga-Exped. 31: 1-127.
Jay, J. C.
1 839. A catalogue of the shells, arranged according to the Lamarckian System, together with descriptions of
new or rare species, contained in the collection of John C. Jay, M. D. New York, Wiley and Putnam. 125
p. 10 pis.
Krauss, F.
1 848. Die SudaFrikanischen Mollusken. Stuttgart.
Lamarck, J. B. P. A. de M. de
1818. Histoire naturelle des animaux san vertebres 5: 375-410. Paris, Deterville.
Lamy, E. and M. Andre
1932. Notes sur les especes Lamarckiennes de Cirripedes. Compt. Rendu Cong. Soc. Sav. Paris Depart.. Sec.
Sci. 65: 212-228.
Martini, F. H. W., and J. H. Chemnitz
.1 785. Neues systematisches Conchylien-cabinet 8: 294-347, pi. 96-100.
Newman, W. A.
1967. A new genus of Chthamalidae (Cirripedia, Balanomorpha) from the Red Sea and Indian Ocean. J.
Zool. London 153: 423-435.
Nilsson-Cantell, C. A.
1921. Cirripeden-Studien. Zur Kenntnis der Biologic Anatomie und Systematik dieser Gruppe. Zool. Bidr.
Uppsala 7: 75-395.
1 939. Recent and fossil balanids from the north coast of South America. Cap. Zool. 8 (4): 1 -7.
Patel, B., and D. J. Crisp
1 96 1 . Relation between the breeding and moulting cycles in cirripedes. Crustaceana 2(2): 89-107.
Pilsbry, H. A.
1916. The sessile barnacles contained in the collection of the U.S. National Museum: including a monograph
of the American species. Bull. U.S.Natl. Mus. 93: 1-366.
1927. Cirripedia of Curacao. Bijdr. Dierk. 25: 37-28.
1953. Noteson Floridan barnacles (Cirripedia). Proc. Acad. Nat. Sci. Philadelphia 105: 13-28.
Pope, E. C.
1943. Animal and plant communities of the coastal rock platform at Long Reel. New South Wales. Proc.
Linnean Soc. New South Wales 68 (5-6): 22 1 -254.
1945. A simplified key to the sessile barnacles found on the rocks, boats, wharf piles and other installations in
Port Jackson and adjacent waters. Rec. Australian Mus. 2 1 (6): 35 1 -372.
1965. A review of Australian and some Indomalayan Chthamalidae (Crustacea: Cirripedia). Proc. Linnean
Soc. New South Wales 90(1): 10-77.
Ranzani, C.
1818. Osservazioni su i Balanidi. Opuscoli Scientifici 2: 63-93.
1820. Osservazioni su i Balanidi. Mem. Storia Nat., deca prima: 13-57.
250 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL.15
Ross. A.
1968. Bredin-Archbold-Smithsonian Biological survey of Dominica 8. The intertidal balanomorph Cirri-
pedia. Proc. U.S. Natl. Mus. 125 (3663): 1-22.
Schmalz, C.
1906. Die ordnung der Cirripedien Systematische Conchylien-cabinet von Martini und Chemnitz II (27-28):
1-82. (published 1911).
Schumacher, C F.
1817. Essai d'un nouveau systeme des habitations des vers testaces. Copenhagen. 287p.
Spengler. L.
1 790. Beskrivelse og Oplysning over den hidindtil lidet udarbeijdede Slaegt af mangeskallede Konchylier, som
Linnaeus har daldet Lepas, med tilf^iede nye og ubeskrevne Arter. (Om Conchylie-Slaegten Lepas).
Skrift. Naturhist. Selsk. 1 ( 1 ): 1 58-2 12.
Southward, A. J.
1962. On the behaviour of barnacles. IV. The influence of temperature on cirral activity and survival of some
warm-water species. J. Marine Biol. Assoc. U.K. 42: 163-177.
Southward, A. J., and E. Southward
1958. On the occurrence and behaviour of two little-known barnacles, Hexelasma hirsutum and Verruca
recta, from the continental slope. J. Marine Biol. Assoc. U.K. 37: 633-647.
Sowerby, G. B.
1823 [1821-1834]. The genera of Recent and Fossil shells. London. No pagination.
Weltner, W.
1 897. Verzeichnis der bisher beschriebenen recenten Cirripedienarten. Arch. f. Naturgesch. 1 (3): 227-280.
Wood, W.
1815. General conchology; or a description of shells arranged according to the Linnean System. London. 246
P-
Zullo, V. A.
1968. Tesseropora Pilsbry (Cirripedia, Thoracica) from the Pliocene of the Gulf of California. Crustaceana 1 5
(3): 272-274.
Department of Invertebrate Paleontology, Natural History Museum, P.O. Box 1390,
San Diego, California 921 12
APPENDIX
MEASUREMENTS FOR NEWMANELLA RADIATA
Table 1. Summary of data on cirral counts: range (R) and mean (X)
values for the number of segments in anterior (a) and posterior (p) rami.
I
II
III
IV
V
V,
a
P
a
P
a
P
a
P
a
P
a
-
N
10
10
10
10
10
10
10
10
10
10
8
9
Bahamas
R
18-23
10-15
7-9
12-19
8-12
11-24
20-24
20-24
19-25
20-25
24-26
22-26
X
18.20
12.20
8.40
15.30
9.10
19.3
21.30
21.70
22.50
23.20
24.88
24.67
N
8
8
8
7
8
7
6
6
6
6
4
6
Vieques Sound
R
19-21
11-14
7-13
11-14
8-14
12-17
20-28
23-30
24-33
25-34
26-35
26-34
Puerto Rico
X
19.75
12.00
10.37
12.71
11.62
14.57
24.00
27.00
28.00
29.66
29.75
30.16
N
5
7
8
8
8
7
8
7
7
7
7
8
Venezuela
R
14-18
9-11
6-13
8-21
6-8
14-21
12-15
13-16
13-18
14-17
14-18
14-18
X
16.00
10.00
8.00
13.25
7.25
18.42
13.38
14.43
15.14
15.14
16.14
16.25
N
8
8
8
8
8
5
8
9
8
7
7
9
Trinidad
R
16-25
8-17
7-10
16-22
7-11
12-25
13-19
16-23
16-20
17-24
17-24
16-25
X
21.00
13.62
8.75
18.50
9.00
19.00
16.38
19.00
18.50
19.85
19.85
19.88
1969
ROSS: STUDIES ON THE TETRACLITIDAE
251
Table 2. Cirral counts lor individual specimens
Dominica
ntzht
left
II
111
IV
\ I
Anterior
19
9
11
14
17
15
Posterior
13
17
21
13
15
14
Anterior
18
9
9
12
14
14
Posterior
12
16
11
12
14
Vieques Island, Puerto Rico
right
left
Anterior
22
12
15
27
37
37
Posterior
14
20
23
34
37
36
Anterior
22
13
15
26
33
27
Posterior
15
19
18
31
38
39
Table 3. Summary of data on shell and opercular valve parameters
(in mm): range (R) and mean (X) values.
Car-Ros
SHELL
OPERCULAR PLATES
Scutum Tergum
Diam.
Lat. Diam.
Height
Height
Width
Height
Width
Bahamas
(n = 5)
R
X
9.3-17.2
13.4
9.5-16.5
12.7
5.9-13.3
11.4
2.5-4.8
4.2
2.5-4.9
4.3
3.4-6.1
5.3
2.3-4.5
3.8
Vieques Sound,
(n=4)
PR.
R
X
12.9-23.1
17.6
13.3-24.6
17.8
7.5-13.9
10.4
4.6-6.6
5.5
4.1-7.0
5.2
6.1-9.5
7.5
4.3-7.6
5.7
Venezuela
(n = 6)
R
X
5.4-7.2
5.9
5.1-7.1
6.0
2.6-4.5
3.0
Trinidad
(n=5)
R
X
11.1-14.9
12.1
9.4-14.5
9.8
6.1-14.7
9.3
3.0-6.8
4.3
2.7-6.1
4.0
3.5-7.4
4.S
2.6-5.0
3.3
Dominica
(n=l)
10.0
8.1
7.8
3.1
2.9
3.5
2.6
Vieques Island,
(n=l)
PR.
31.3
29.3
28.1
10.1
10.5
13.1
13.9
MUS. COMP. ZOOL.
LIBRARY
OCT 30 1969
HARVARD
UNIVERSITY,
TYPE SPECIMENS OF MAMMALS IN THE
SAN DIEGO NATURAL HISTORY MUSEUM
SUZANNE I. BOND
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15, NO. 16 3 OCTOBER 1969
TYPE SPECIMENS OE MAMMALS IN THE SAN DIEGO
NATURAL HISTORY MUSEUM
Suzanne I. Bond
The mammal collections of the San Diego Society of Natural History comprise more
than 21,000 specimens, of which the majority are from the southwestern United States and
Baja California, Mexico. Included are holotypes of two species and 87 subspecies. In this
catalog Of type specimens, the following information is given: name as originally published,
original citation, currently accepted name if different from the original, collector, date of
collection, original number, museum number, and type locality. Unless otherwise specified,
all taxa listed here were considered valid by Hall and Kelson (Mammals of North America.
Ronald Press, New York, 1959). Geographic coordinates not given in the original
description and corrected spelling are given in brackets. All holotype specimens are in good
condition; skulls and skins are present for all.
The length of the list precluded the inclusion of paratypes. For a complete list, order
NAPS Document 00310 from ASIS National Auxiliary Publications Service, c/o CCM
Information Sciences, Inc., 22 West 34th Street, New York, New York 10001; remitting $1
for microfiche or $3 for photocopies.
FAMILY LEPORIDAE
Sylvilagus bachmani howelli Huey
San Diego Soc. Nat. Hist., Trans. 5 (5): 67-68, 6 July 1927.
Holotype. - Adult female. Collected by L. M. Huey, 10 December 1926. SDSNH no. 5939.
Type locality. - 10 mi SE Alamo, Baja California, Mexico; lat. 31 °35'N, long. 116°03'W.
Sylvilagus bachmani rosaphagus Huey
San Diego Soc. Nat. Hist., Trans. 9 (23): 22 1 -223, 3 1 July 1940.
Holotype. - Adult male. Collected by L. M. Huey, 2 June 1925. SDSNH no. 4947.
Type locality — 2 mi W Santo Domingo Mission, Baja California, Mexico; lat. 30°45'N,
long. 115°58'W.
FAMILY SCIURIDAE
Ammospermophilus harrisii kinoensis Huey
San Diego Soc. Nat. Hist., Trans. 8 (25): 352-353, 1 5 June 1937.
= Ammospermophilus harrisii saxicola (Mearns)
Holotype. -- Adult female. Collected by L. M. Huey, 22 February 1935. SDSNH no.
11284.
Type locality. Bahia Kino, Sonora, Mexico [lat. 28°40'N, long. lll°58'Wj.
Remarks. A. H. Howell (N. Amer. Fauna 56: 169, 1938) considered A. h. kinoensis a
synonym of A. h. saxicola.
Ammospermophilus leucurus canfieldae Huey
San Diego Soc. Nat. Hist., Trans. 5(15): 243-244, 27 February 1 929.
Holotype. -- Adult male. Collected by L. M. Huey, 14 February 1928. SDSNH no. 6873.
Type locality. Punta Prieta, Baja California, Mexico; lat. 28°56'N. long. 1I4°I2'W.
SAN D1LGO SOC. NAT HIST.. TRANS 15 (16): 252-263, 3 OCTOBER 1969
253 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
Citellus beecheyi nudipes Huey
San Diego Soc. Nat. Hist., Trans. 7 (2): 1 8-20, 6 October 1 93 1 .
= Spermophilus beecheyi nudipes (Huey)
Holotype. - Adult female. Collected by F. Stephens, 1 3 October 1 926. SDSNH no. 201 5.
Type locality. Laguna Hanson, Sierra Juarez, Baja California, Mexico; lat. 31°58'N,
long. 115°53'W.
Remarks. The Committee on Nomenclature, American Society of Mammalogists
recommended the use of Spermophilus to replace Citellus {J. Mamm.49(3): 605, 1968).
Citellus beecheyi rupinarum Huey
San Diego Soc. Nat. Hist., Trans. 7 (2): 17-18, 6 October 1931
= Spermophilus beecheyi rupinarum (Huey)
Holotype. Sub-adult female. Collected by L. M. Huey, 9 October 1930. SDSNH no.
8251.
Type locality. Catavina, Baja California, Mexico; lat. 29°54'N, long. 1 14°57'W.
Citellus tereticaudus apricus Huey
San Diego Soc. Nat. Hist. Trans. 5 (7): 85-86, 10 October 1927.
= Spermophilus tereticaudus apricus (Huey)
Holotype. -Adult male. Collected by L. M. Huey, 13 July 1927. SDSNH no. 6308.
Type locality. - Valle de la Trinidad, Baja California, Mexico; lat. 31°20'N, long.
li5°40'W.
Citellus tereticaudus vociferans H uey
Proc. Biol. Soc. Wash. 39: 29-30, 30 July 1926.
= Spermophilus tereticaudus tereticaudus Baird
Holotype. Adult female. Collected by L. M. Huev, 25 March 1926. SDSNH no. 5127.
Type locality. - San Felipe, Baja California, Mexico [ lat. 3 1 °02'N, long. 1 14°50'W ].
Remarks. -A. H. Howell (N. Amer. Fauna 56: 185, 1938) considered C. t. yociferans a
synonym of C. t. tereticaudus.
FAMILY GEOMYIDAE
Thomomys bottae abbotti Huey
San Diego Soc. Nat. Hist.. Trans. 5 (8): 89-90, 18 January 1928.
= Thomomys umbrinus abbotti Huey
Holotype. Adult male. Collected by L. M. Huey, 10 May 1925. SDSNH no. 4781 .
Type locality. 1 mi E El Rosario, Baja California, Mexico; lat. 30°03'N, long. 1 15°48'W.
Remarks. Hall & Kelson (Mamm. N. Amer. 1:416. 1959) considered bottae conspecific
w\ih umbrinus. See also Patton & Dingman, J. Mamm.. 49(1): 1-13. 1968.
Thomomys bottae affinis Huey
San Diego Soc. Nat. Hist., Trans. 10 (14): 254-255, 31 August 1945.
= Thomomys umbrinus affinis Huey
Holotype. Adult male. Collected by L. M. Huey, 9 May 1940. SDSNH no. 14083.
Type locality. Jacumba, San Diego County, California.
Thomomys argusensis Huey
San Diego Soc. Nat. Hist., Trans. (5): 43-44. 19 December 1931 .
= Thomomys umbrinus argusensis Huey
Holotype. Adult male. Collected by S. G. Harter. 10 August 1931. SDSNH no. 9545.
Type locality. Junction Ranch, Argus Mts., Inyo County, California.
1969 BOND: TYPE SPECIMENS Of MAMMALS 254
Thomomvs bottae aridicola Huey
San'Diego Soc. Nat. Hist., Trans. 8 (25): 354-355. 15 June 1937.
= Thomomvs umbrimts aridicola Huey
Holotype. Adult female. Collected by L. M. Huey, 1 February 1936. SDSNH no.
1 1 424.
Type locality. 10 mi S Gila Bend. Maricopa County, Arizona, about 2 mi N o[' Black
Gap.
Thomomvs bottae boregoensis Huey
San'Diego Soc. Nat. Hist., Trans. 9(15): 70-71, pi. 5, Figure D, 8 December 1939.
= Thomomvs umbrinus boregoensis Huey
Holotype. Adult female. Collected by F. M. Huey. 29 March 1930. SDSNH no. 8034.
Type locality. Beatty Ranch. Borego \ Borrego] Valley, San Diego County, Califor-
nia.
Thomomvs bottae aderrans Huey
San'Diego Soc. Nat. Hist.. Trans. 9 (15): 71-72. pi. 5. Figure E. 8 December 1939.
= Thomomys umbrimts boregoensis Huey
Holotype. Adult female. Collected by F.Stephens . 21 January 1924. SDSNH no.
1305.
Type locality. Carrizo Creek. San Diego County, California.
Remarks. J. F. Chattin (San Diego Soc. Nat. Hist., Trans. 9 (27): 275-277, 1941)
considered T. b. aderrans a synonym of T. u. boregoensis.
Thomomvs bottae borjasensis Huey
San'Diego Soc. Nat. Hist., Trans. 10 (14): 262-263, 31 August 1945.
= Thomomys umbrinus borjasensis Huey
Holotype. Adult female. Collected by F. M. Huey. 14 October 1941. SDSNH no.
14491.
Type locality. San Borjas [= Borja] Mission, Baja California, Mexico; lat. 28°52'n.
long. 113°53'W.
Thomomvs umbrinus brazierhowelli Huey
San'Diego Soc. Nat. Hist.. Trans. 12 (23): 407-408. 1 February 1960.
Holotype. ^Adult male. Collected by A. B. Howell. 26 February 1958. SDSNH no.
18725".
Type locality. -San Fernando Mission. Baja California. Mexico |lat. 29°58'N, long.
I15°15'W].
Thomomvs bottae cactophilus Huev
San'Diego Soc. Nat. Hist., Trans. 5 (15): 241-242. 27 Februan 1929.
= Thomomys umbrinus cactophilus Huey
Holotype. Adult male. Collected by F. M. Huey. 17 Februar> 1928. SDSNH no.
6808.
Type locality. Punta Prieta. Baja California. Mexico: lat. 28°56'N, long. 1 14°I2'\V.
Thomomys bottae cata\inensis Huc\
San'Diego Soc. Nat. Hist.. Trans. 7 (5): 45. 19 December 1937.
= Thomomys umbrinus catavinensis Huey
Holotype. Adult female. Collected by F. M. Hue). 10 October 1930. SDSNH no.
8256.'
Type locality. Catavina, Baja California. Mexico; lat. 29^54'N. long. I 14°57'W.
255 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
Thomomvs bottae cedrinus Huey
San Diego Soc. Nat. Hist., Trans. 12 (6): 100-101, 10 February 1955.
= Thomomvs umbrinus cedrinus Huey
Holotype. Adu\t female. Collected by L. M. Huey, 27 April 1938. SDSNH no. 13161.
Type locality. --From summit of Crossman Peak, Chemehuevis Mts.; Mohave County,
Arizona.
Thomomvs bottae comobabiensis Huey
San Diego Soc. Nat. Hist., Trans. 8 (25): 354, 15 June 1937.
= Thomomys umbrinus comobabiensis Huey
Holotype. Adult female. Collected by L. M. Huey, 22 March 1937. SDSNH no.
1 2460".
Type locality. 5 mi N W Sells, Pima County, Arizona.
Thomomys bottae cunicularius Huey
San'Diego Soc. Nat. Hist., Trans. 10 (14): 252-253, 31 August 1945.
= Thomomys umbrinus cunicularius Huey
Holotype. -Adult male. Collected by L. M. Huey, 29 November 1936. SDSNH no.
12182.
Type locality. - Los Palmitos, SE base of Sierra Juarez, Baja California, Mexico; lat.
31°44'N, long. 115°36'W.
Thomomys bottae growlerensis Huey
San Diego Soc. Nat. Hist., Trans. 8 (25): 353-354, 15 June 1937.
= Thomomys umbrinus growlerensis Huey
Holotype. -Adult male. Collected by L. M. Fluey, 16 March 1937. SDSNH no. 12387.
Type locality. - 7 mi E Papago Well, Pima County, Arizona.
Thomomys bottae homorus Huey
San Diego Soc. Nat. Hist., Trans. 1 1 (4): 55, 31 January 1949.
= Thomomys umbrinus homorus Huey
Holotype. Adult male. Collected by L. M. Huey, 6 April 1947. SDSNH no. 15689.
Type locality. - 1 mi E Rancho Lagunita, Baja California, Mexico; lat. 28°20'N, long.
113°15'W.
Thomomvs bottae hueyi Goldman
Wash. Acad. Sci. 28: 340-341, 15 July 1938.
= Thomomys umbrinus hueyi Goldman
Holotype. Adult male. Collected by L. M. Huey, 17 June 1932. SDSNH no. 10088.
Type locality. Spud Rock Ranger Station, 7400 ft., Rincon Mts., Pima County,
Arizona.
Thomomys bottae jojobae Huey
San Diego Soc. Nat. Hist., Trans. 10 (14): 256-257, 31 August 1945.
= Thomomys umbrinus jojobae Huey
Holotype. Adult male. Collected by L. M. Huey, 20 June 1927. SDSNH no. 61 16.
Type locality. Sangre de Cristo, Baja California, Mexico; lat. 31°52'N, long.
116°06'W.
Thomomys bottae juarezensis Huey
San'Diego Soc. Nat. Hist., Trans. 10 (14): 255-256, 31 August 1945.
= Thomomys umbrinus juarezensis Huey
Holotype. Adult male. Collected by L. M. Huev, 6 November 1936. SDSNH no.
5849. '
1969 BOND: TYPE SPECIMENS OE MAMMAI S 256
Type locality. Laguna Hanson, Sierra Juarez, Baja California, Mexico llat. 31°58'N,
long. 115°53'W].
Thomomys bottae lorenzi Huey
San Diego Soc. Nat. Hist., Trans. 9 (22): 219-220, 31 July 1940.
= Thomomys umbrinus lorenzi Huey
Holotvpe. Adult male. Collected by P. Covel, 17 April 1933. Original no. 175, now
SDSNH no. 10608.
Type locality. 7 mi N Boulder Creek. Santa Cruz County, California.
Thomomys bottae proximarinus Huey
San Diego Soc. Nat. Hist., Trans. 10(14): 261, 31 August 1945.
= Thomomys umbrinus proximarinus Huey
Holotvpe. Adult male. Collected by L. M. Huey. 15 August 1940. SDSNH no. 14182.
Type locality. Boca la Playa, 16 mi W Santo Tomas, Baja California, Mexico; lat.
31°32'N, long. I16°38'W.
Thomomys bottae rhizophagus Huey
San Diego Soc. Nat. Hist., Trans. 1 1 (4): 54-55, 31 January 1949.
= Thomomys umbrinus rhizophagus Huey
Holotvpe. Adult male. Collected by L. M. Huey. 1 1 April 1947. SDSNH no. 15710.
Type locality. Las Flores, 7 mi S of Bahia de Los Angeles, Baja California, Mexico;
lat. 28°50'N, long. 113°32'W.
Thomomys bottae ruricola Huey
San'Diego Soc. Nat. Hist., Trans. 1 1 (4): 53-54, 31 January 1949.
= Thomomys umbrinus ruricola Huey
Holotvpe. Adult male. Collected by L. M. Huey, 27 June 1947. SDSNH no. 15944.
Type locality. -4 mi N Santa Catarina Landing, Baja California, Mexico: lat. 29°35'N,
long. 115°17'W.
Thomomys bottae sanctidiegi Huey
San Diego Soc. Nat. Hist., Trans. 10 (14): 258-259, 31 August 1945.
= Thomomys umbrinus sanctidiegi Huey
Holotvpe. -Adult male. Collected by L. M. Huev, 18 December 1941. SDSNH no.
14886.
Type locality. Balboa Park, San Diego [San Diego County], California.
Thomomys bottae vanrossemi Huey
San Diego Soc. Nat. Hist., Trans. 8(1): 1-2, 10 August 1934.
= Thomomys umbrinus xanrossemi Huey
Holotvpe. -Adult male. Collected by L. M. Huey, 15 February 1934. SDSNH no.
10922.
Type locality. Punta Penascosa, Sonora, Mexico I lat. 31°25'N, long. 1 I3°35'W I.
Thomomys bottae xerophilus Huey
San Diego Soc. Nat. Hist., Trans. 10 (14): 257, 31 August 1945.
= Thomomys umbrinus .xerophilus Huey
Holotvpe. - Adult male. Collected by L. Mi Huey. 29 March 1936. SDSNH no. I 1827.
Type locality. Near Diablito Spring, summit of San Matias Pass (between Sierra
Juarez & Sierra San Pedro Martir), Baja California, Mexico [near lat. 31°20'N, long.
115°30'W].
Thomomys quadratus monoensis Huey
San Diego Soc. Nat. Hist., Trans. 7 (35): 373-374, 31 May 1934.
257 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
= Thomomys talpoides monoensis Huey
Holotype. -Adult male. Collected by L. M. Huey, 7 August 1932. SDSNH no. 10414.
Type locality. - - Dexter Creek Meadow, at confluence of Dexter and Wet Creeks, Mono
County, California.
Remarks. Goldman (J. Mamm. 20: 234, 1939) considered T. q. monoensis a synonym
of T. t. monoensis.
FAMILY HETEROMYIDAE
Perognathus longimembris aestivus Huey
San Diego Soc. Nat. Hist., Trans. 5 (8): 87-89, 18 January 1928.
Holotype. Adult male. Collected by L. M. Huey, 20 June 1937. SDSNH no. 61 10.
Type locality. - Sangre de Cristo in Valle San Rafael on W base of Sierra Juarez, Baja
California, Mexico; lat. 31°52'N, long. 116°06'W.
Perognathus longimembris internationalis Huey
San Diego Soc. Nat. Hist., Trans. 9(11): 47-50, 31 August 1939.
Holotype. - Adult male. Collected by L. M. Huey, 24 April 1936. SDSNH no. 1 1972.
Type locality. Baja California side of the International Boundary at Jacumba, San
Diego County, California.
Perognathus longimembris kinoensis Huey
San Diego Soc. Nat. Hist., Trans. 8 (12): 73-74, 24 August 1935.
Holotype. -Adult male. Collected by L. M. Huey, 26 February 1935. SDSNH no.
1 1 300.
Type locality. Bahia Kino, Sonora, Mexico [lat. 28°40'N, long. 1 1 1°58'W].
Perognathus longimembris pimensis Huey
San Diego Soc. Nat. Hist., Trans. 8 (25): 355-357, 15 June 1937.
Holotype. -Adult male. Collected by L. M. Huey, 22 May 1937. SDSNH no. 12579.
Type locality. 1 1 mi W Casa Grande, Pinal County, Arizona.
Perognathus longimembris venustus Huey
San Diego Soc. Nat. Hist., Trans. 6(17): 233, 24 December 1930.
Holotype. Adult female. Collected by L. M. Huey, 4 October 1930. SDSNH no. 8196.
Type locality. San Agustin, Baja California, Mexico; lat. 30°N, long. 1 15°W.
Perognathus longimembris virginis Huey
San Diego Soc. Nat. Hist., Trans. 9(12): 55-56, 31 August 1939.
Holotype. Adult male. Collected by L. M. Huey, 12 August 1937. SDSNH no. 12919.
Type locality. - St. George, 2950 ft., Washington County, Utah.
Perognathus alticola inexpectatus Huey
Proc. Biol. Soc. Wash. 39: 121-122, 27 December 1926.
Holotype. Adult male. Collected by G. Cantwell, 28 August 1936. Original no. 216,
now SDSNH no. 5724.
Type locality. 14 mi W Lebec, Kern County, California.
Perognathus formosus infolatus Huey
San Diego Soc. Nat. Hist., Trans. 12(1): 1-2, 1 March 1954.
Holotype. Adult male. Collected by L. M. Huey, 3 April 1947. SDSNH no. 15664.
Type locality. 7 mi W San Francisquito Bay, Gulf of California, Baja California,
Mexico; lat.'28°30'N, [long. I13°W].
Perognathus formosus mohavensis Huey
San Dieeo Soc. Nat. Hist., Trans. 9 (8): 35-36, 21 November 1938.
1969 BOND: TYPE SPECIMENS OK MAMMALS 258
Holotype. Adult male. Collected by L. M. Huey. 14 April 1935. SDSNH no. 1 1317.
Type locality. Bonanza King Mine, Providence Mts., San Bernardino County, Califor-
nia.
Perognathus bailey i hueyi Nelson & Goldman
Proc. Bio. Soc. Wash. 42: 106-107,25 March 1929.
Holotype. Adult female. Collected by L. M. Huey. 17 April 1926. SDSNH no. 5220.
Type locality. San Felipe, Baja California, Mexico [lat. 31°02'N, long. 1 14°50'W|.
Perognathus baileyi mesidios Huey
San Diego Soc. Nat. Hist., Trans. 13 (7): 112, 15 January 1964.
Holotvpe. ^Adult male. Collected by L. M. Huey, 13 October 1941. SDSNH no.
14470".
Type locality. -San Borja Mission, Baja California, Mexico; lat. 28°45'N I long.
113°50'W].
Perognathus arenarius albescens Huey
Proc. Biol. Soc. Wash. 39: 67-69, 30 July 1926.
Holotype. Adult male. Collected by L. M. Huey. 23 March 1926. SDSNH no. 5103.
Type locality. San Felipe. Baja California, Mexico I lat. 31°02'N, long. 1 14°50'W].
Perognathus arenarius mexicalis Huey
San Diego Soc. Nat. Hist., Trans. 9(13): 57-58, 31 August 1939.
Holotype. -Adult female. Collected by L. M. Huey. 23 November 1936. SDSNH no.
12127".
Type locality. - Los Muertos Canyon fan. GaskilPs Tank, near Laguna Salada, Baja
California, Mexico: lat. 32°27'N, long. 115°53'W.
Perognathus arenarius paralios Huey
San Diego Soc. Nat. Hist., Trans. 13 (7): 113. 15 January 1964.
Holotype. Adult male. Collected by L. M. Huey. 23 March 1926. SDSNH no. 15542.
Type locality. Barril. Baja California, Mexico; lat. 28°20'N [long. 1 12°50'W].
Perognathus arenarius sabulosus Huey
San Diego Soc. Nat. Hist., Trans. 13 (7): 1 14, 15 January 1964.
Holotype. -Adult male. Collected by L. M. Huey, 22 May 1926. SDSNH no. 5300.
Type locality. - From mainland on the S side of Scammon's Lagoon. Baja California,
Mexico [near lat. 27°40'N, long. 1 14°05'W].
Perognathus intermedius lithophilus Huey
San Diego Soc. Nat. Hist.. Trans. 8 (25): 355. 15 June 1937.
Holotype. Adult male. Collected by L. M. Huey. 5 February 1935. SDSNH no.
11211".
Type locality. Porto [= Puerto] Libertad. Sonora, Mexico [lat. 29°57'N, long.
112°40'W|.
Perognathus fallax majusculus Huey
San Diego Soc. Nat. Hist., Trans. 12 (25): 418, 1 February 1960.
Holotype. Adult male. Collected by L. M. Huey, 29 June 1947. SDSNH no. 15952.
Type locality. San Quintin, Baja California, Mexico | lat. 30°30'N, long. 1 1 5°59'W | .
Perognathus fallax xerotrophicus Huey
San Diego Soc. Nat. Hist., Trans. 12 (25): 419, I February 1960.
Holotype. -Adult male. Collected by L. M. Huey, 15 October 1930. SDSNH no. 8310.
Type' locality. 2 mi NW Chapala. Baja California. Mexico flat. 29°25'N, long.
114°20'W|.
259 SAN DIEGO SOCILTY OF NATURAL HISTORY VOL. 15
Perognathus spinatus broccus Huey
San Diego Soc. Nat. Hist., Trans. 12 (24): 410, 1 February 1960.
Holotype. -Adult male. Collected by L. M. Huey, 18 March 1928. SDSNH no. 6891.
Type locality. San Ignacio, Baja California, Mexico; lat. 27°17'N [long. 1 I2°55'W].
Perognathus spinatus lorenzi Banks
Pro. Biol. Soc. Wash. 80: 101-103, 28 July 1967.
Holotvpe. Adult male. Collected by R. C. Banks, 22 October 1964. Original no. RCB-
2366, now SDSNH no. 19901.
Type locality. South San Lorenzo Island, Gulf of California, Baja California, Mexico;
lat. 23°36'N, long. 112°51'W.
Perognathus spinatus oribates Huey
San Diego Soc. Nat. Hist., Trans. 12 (24): 409-410, 1 February 1960.
Holotype. -- Adult male. Collected by L. M. Huey, 27 February 1958. SDSNH no
18742.
Tvpe locality. - San Fernando Mission, Baja California, Mexico; lat. 30°N [long.
115°15'W].
Perognathus spinatus prietae Huey
San Diego Soc. Nat. Hist., Trans. 6(17): 232-233, 24 December 1930.
Holotype. -Adult male. Collected by L. M. Huey, 26 October 1930. SDSNH no. 8450.
Tvpe locality. -25 mi N Punta Prieta, Baja California, Mexico; lat. 29°24'N, long.
114°24'W.
Perognathus spinatus rufescens Huey
San Diego Soc. Nat. Hist., Trans. 6(17): 231-232, 24 December 1930.
Holotype. -Adult male. Collected by L. M. Huey, 10 November 1929. SDSNH no.
7446.
Type locality. - Mouth of Palm Canyon, Borego [= Borrego] Valley, San Diego
County, California.
Dipodomys mohavensis argusensis Huey
San Diego Soc. Nat. Hist., Trans. 10 (10): 131-132, 9 March 1945.
= Dipodomys panamintinus argusensis Huey
Holotvpe. - Adult male. Collected by S. G. Harter, 13 August 1931. Original no. 140,
now SDSNH no. 9552.
Type locality. Junction Ranch, 5725 ft., Argus Mts., Inyo County, California.
Remarks. ' Miller and Kellog (Bull. U.S. Nat. Mus. 205: 387, 3 March 1955) consid-
ered D. m. argusensis a synonym of D. p. argusensis.
Dipodomys cascus Huey
San Diego Soc. Nat. Hist., Trans. 12 (29): 479-480, 30 August 1962.
= Dipodomys stephensi (Merriam)
Holotype. Adult male. Collected by L. M. Huey, 17 August 1961. SDSNH no. 18961.
Type locality. 1 mi E Bonsall, San Diego County, California.
Remarks. Lackey (San Diego Soc. Nat. Hist., Trans. 14 (22): 328, 1967) considered
D. cascus a synonym of D. stephensi.
Dipodomys agilis martirensis Huey
San Diego Soc. Nat. Hist., Trans. 5(1): 7-8, 20 February 1927.
Holotype. Adult male. Collected by L. M. Huey, 28 September 1926. SDSNH no.
5762.
Type locality. La Grulla (east side of the valley). Sierra San Pedro Martir, Baja
1969 BOND: TYPE SPECIMENS OK MAMMAI S 260
California, Mexico [lat. 30°50'N, long. I 15°20'W].
Dipodomys agilis plectilis Huey
San Diego Soc. Nat. Hist"., Trans. II (10): 240, 30 April 1951.
Holotype. Adult female. Collected by L. M. Huey, 1 May 1925. SDSNH no. 4721.
Type locality. Mouth of Canyon San Juan de Dios, Baja California, Mexico: lat.
30°7'N, [long. 115°10'WJ.
Dipodomys agilis latimaxillaris Huey
Pro.*Biol. Soc. Wash. 38: 84, 26 May 1925.
= Dipodomys agilis simulans (Merriam)
Holotype. Adult male. Collected by L. M. Huey, 27 February 1925. SDSNH no.
4696.
Type locality. 2 mi W Santo Domingo Mission, Baja California, Mexico; lat. 30°45'N,
long. 115°58'W.
Remarks. Huey (San Diego Soc. Nat. Hist., Trans. 11 (10): 234, 1951) later consid-
ered D. a. latimaxillaris a synonym of D. a. simulans.
Dipodomys parlius Huey
San Diego Soc. Nat. Hist., Trans. 11 (10): 241, 30 April 1951.
Holotype. Adult male. Collected by L. M. Huey, 13 April 1923. SDSNH no. 4250.
Type locality. Santa Catarina Landing, Baja California. Mexico; lat. 29°31'N [long.
115°20'W].
Dipodomys peninsularis australis Huey
San Diego Soc. Nat. Hist., Trans. 11 (1): 249, 30 April 1951.
Holotype. Adult male. Collected by L. M. Huey, 21 November 1941. SDSNH no.
14734.
Type locality. - - Santo Domingo, Magdalena Plain, Baja California, Mexico: lat.
25°30'N [long. 112°07'W].
Dipodomys peninsularis eremoecus Huey
San Diego Soc. Nat. Hist., Trans. 1 1 (10): 248-249, 30 April 1951.
Holotype. Adult male. Collected by L. M. Huey, 31 March 1947. SDSNH no. 15619.
Type locality. 7 mi W San Francisquito Bay, Gulf of California, Baja California,
Mexico; lat. 28 °30'N Hong. 112°58'W].
Dipodomys peninsularis pedionomus Huey
San Diego Soc. Nat. Hist., Trans. 1 1 (10): 247-248, 30 April 1951.
Holotype. Adult male. Collected by L. M. Huey, 17 October 1930. SDSNH no. 8363.
Type locality. 2 mi N I label says NW] Chapala Dry Lake, on Llano Santa Ana. Baja
California, Mexico; lat. 29°30'N, long. 1 14°35'W.
Dipodomys gravipes Huey
Pro. "Biol. Soc. Wash. 38:83-84, 26 May 1925.
Holotype. Adult male. Collected by L. M. Huey, 28 February 1925. SDSNH no.
4703."
Type locality. 2 mi W Santo Domingo Mission, Baja California. Mexico: lat. 30°45'N,
long. 115°58'W.
Dipodomys antiquarius Huev
San Diego Soc. Nat. Hist., Trans. 12 (29): 477-479. 30 August 1962.
Holotype. -Adult female. Collected by G. Ives, 30 March 1960. SDSNH no. 18901.
Type locality. - - San Juan Mine, Sierra San Borja. Baja California. Mexico: lat.
28°50'N, long. 113°37'W.
261 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
Dipodomys merriami annulus Huey
San Diego Soc. Nat. Hist., Trans. 1 1 (10): 224-225, 30 April 1951.
Holotvpe. Adult female. Collected by L. M. Huey, 22 March 1947. SDSNH no.
15522.
Type locality. Barril, Gulf of California, Baja California, Mexico; lat. 28°20'N, long.
112°50'W.
Dipodomys merriami brunensis Huey
San Diego Soc. Nat. Hist., Trans. 11 (10): 225-226, 30 April 1951.
Holotvpe. -Adult male. Collected by L. M. Huey, 28 March 1928. SDSNH no. 6904.
Type locality. Llano de San Bruno, Baja California, Mexico i lat. 26°15'N, long.
111°35'W]. '
Dipodomys merriami llanoensis Huey
San Diego Soc. Nat. Hist., Trans. 11 (10): 226-227, 30 April 1951.
Holotvpe. Adult male. Collected by L. M. Huey, 15 November 1941. SDSNH no.
14676'.
Type locality. Buena Vista, Magdalena Plain, Baja California, Mexico: lat. 24°51'N,
long. 111°50'W.
Dipodomys merriami quintinensis Huey
San Diego Soc. Nat. Hist., Trans. 1 1 (10): 222-223, 30 April 1951.
Holotvpe. -Adult male. Collected by L. M. Huey, 9 April 1923. SDSNH no. 4205.
Type locality. 5 mi E San Quintin, Baja California, Mexico [lat. 30°30'N, long.
H5°50'W|.
Dipodomys merriami semipallidus Huey
San Diego Soc. Nat. Hist., Trans. 5 (5): 65-66, 6 July 1927.
Holotvpe. Adult female. Collected by L. M. Huey, 15 April 1923. SDSNH no. 4271.
Type locality. 7 mi N Santa Catarina, Baja California, Mexico; lat. 29°45'N, long.
115°10'W.
Dipodomys merriami trinidadensis Huey
San Diego Soc. Nat. Hist., Trans. 11 (10): 220-221, 30 April 1951.
Holotvpe. Adult male. Collected by L. M. Huey, 13 March 1936. SDSNH no. 11531.
Type locality. Aguajito Spring, El Valle de la Trinidad, Baja California, Mexico [near
lat. 31°28'N, long. 115°40'W].
Dipodomys deserti arizonae Huey
San Diego Soc. Nat. Hist., Trans. 12 (6): 99-100, 10 February 1955.
Holotvpe. - Adult male. Collected by L. M. Huey, 14 May 1937. SDSNH no. 12532.
Type locality. 3 mi SE Picacho, Pinal County, Arizona.
FAMILY CRICETIDAE
Peromyscus crinitus pallidissimus Huey
San Diego Soc. Nat. Hist., Trans. 6 (26): 389-390, 28 August 1931.
Holotvpe. Adult male. Collected by A. W. Anthony, 21 November 1930. Original no.
I 175, now SDSNH no. 8864.
Type locality. Small island in Gonzaga Bay, Baja California, Mexico; lat. 29°50'N,
long. 114°20'W.
Peromyscus guardia harbisoni Banks
J. Mamm. 48 (2): 215, 20 May 1967.
Holotvpe. Adult male. Collected by C. F. Harbison, 23 March 1963. Original no.
RCB-1607, now SDSNH no. 19194.
1969 BOND: TVI'i: SIM ( IMI NS ()l MAMMALS 2(0
Type locality. Isla Granite, Gulf of California, Baja California, Mexico; lat. 29°33'N,
long. 113°34'W.
Peromvscus interparietalis lorenzi Banks
J. Mamm. 48 (2): 216, 20 May 1967.
Holotype. - Adult male. Collected by R. C. Banks, 23 October 1964. Original no. RCB-
2282, now SDSNH no. 19912.
Type locality. Isla San Lorenzo Norte, Gulf of California, Baja California, Mexico;
lat. 28°42'N, long. 112°57'W.
Peromvscus interparietalis ryckmani Banks
J. Mamm. 48 (2): 216, 20 May 1967.
Holotype. Adult female. Collected by R. C. Banks, 23 October 1964. Original no.
RCB-2335, now SDSNH no. 19944.
Type locality. Isla Salsipuedes, Gulf of California, Baja California, Mexico; lat.
28°42'N, long. 112°59'W.
Peromvscus maniculatus hueyi Nelson and Goldman
San Diego Soc. Nat. Hist., Trans. 7 (7): 51-52, 15 April 1932.
Holotype. Adult male. Collected by A. W. Anthony, 21 November 1930. Original no.
1172, now SDSNH no. 8861.
Type locality. Small island in Gonzaga Bay, Baja California, Mexico; lat. 20°15'N
Hong. 114°20'W|.
Neotoma lepida aridicola Huey
San Diego Soc. Nat. Hist., Trans. 12 (15): 287-288, 25 September 1957.
Holotype. Adult male. Collected by L. M. Huey, 27 March 1947. SDSNH no. 15595.
Type locality. El Barril, Gulf of California, Baja California, Mexico; lat. 28°20'N
[long. I12°50'W].
Neotoma lepida aureotunicata Huey
San Diego Soc. Nat. Hist., Trans. 8 (25): 349-350, 15 June 1937.
Holotype. Adult male. Collected by L. M. Huey, 14 February 1934. SDSNH no.
10907'.
Type locality. Punta Penascosa, Sonora, Mexico [lat. 31°20'N, long. 1 13°40'W|.
Neotoma lepida harteri Huev
San Diego Soc. Nat. Hist., Trans. 8 (25): 351-352, 15 June 1937.
Holotype. Adult male. Collected by L. M. Huey, 8 February 1936. SDSNH no.
11462.
Type locality. 10 mi S Gila Bend (2 mi N Black Gap), Maricopa County, Arizona.
Neotoma lepida molagrandis Huey
San Dieao Soc. Nat. Hist., Trans. 10 (16): 307-310, 31 August 1945.
Holotype. Adult male. Collected by L. M. Huey, 28 April 1940. SDSNH no. 14065.
Type locality. Santo Domingo Landing, some 3 mi inland from the beach landing.
Baja California, Mexico; lat. 28°15'N [long. 1 14°02'W].
Phenacomvs silvicolus A. B. Howell
J. Mamm. 2: 98-100. pi. 7. 2 May 1921.
= Phenacomys silxicola A. B. Howell
Holotype. Young adult female. Collected by P. Walker, 25 October 1916. Originally
of the S. G. Jewett Coll., no. 1214. now SDSNH no. 16437.
Type locality. 5 mi SF Tillamook. Tillamook County. Oregon.
263 SAN DIEGO SOCIETY OE NATURAL HISTORY VOL. 15
Microtus californicus grinnelli Huey
San Diego Soc. Nat. Hist., Trans. 7 (6): 47-49, 19 December 1931.
Holotype. ^Adult male. Collected by L. M. Huey, 26 June 1927. SDSNH no. 6165.
Type locality. Sangre de Cristo in Valle San Rafael on the W base of the Sierra
Juarez, Baja California, Mexico; lat. 31°52'N, long. 1 16°06'W.
Ondatra zibethicus goldmani Huey
San Diego Soc. Nat. Hist.. Trans. 8 (32): 409-410, 18 January 1938.
Holo type. -\du It female. Collected by L. M. Huey, 11 August 1937. SDSNH no.
12915.
Type locality. - Saint George, Washington County, Utah.
FAMILY CANIDAE
Urocyon cinereoargenteus peninsularis Huey
San Diego Soc. Nat. Hist., Trans. 5 (13): 205-207, pi. 25, Figure 1, pi. 26, Figure 3,
1 September 1928.
Holotype. -Adult male. Collected by L. M. Huey, 10 March 1928. SDSNH no. 6845.
Type locality. -San Ignacio, Baja California, Mexico; lat. 27°24'N, long. 1 12°59'W.
FAMILY PROCYONIDAE
Bassariscus astitus yumanensis Huey
San Diego Soc. Nat. Hist., Trans. 8 (25): 357-359, pi. 23, 15 June 1937.
Holotype. -Adult male. Collected by L. M. Huey, 6 March 1937. SDSNH no. 12272.
Type locality. Tinajas Atlas, Gila Mts., Yuma County, Arizona.
Department of Birds and Mammals, San Diego Natural History Museum, P.O. Box
1390, San Diego, California 92112
'D" _ MUS. COMP. ZOOL.
L1BRARY
OCT 30 1969
HARVARD
UNIVERSITY
TWELVE NEW DICOTS
FROM BAJA CALIFORNIA, MEXICO
REID MORAN
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
VOL. 15. NO. 17 15 OCTOBER 1969
TWELVE NEW DICOTS
FROM BAJA CALIFORNIA, MEXICO
REID MORAN
ABSTRACT. — Described are Hedeuma martirense, Heterotheca martirensis, Stephanomeria mon-
ocephala, and Tanaceium bajacalifornicum, from the high east face of the Sierra San Pedro Martir:
Eriogonum zapatuense and Castilleja fruticosa, from Isla Guadalupe; Eriogonum angelense. from lsla Angel
de la Guarda; Dudleva nubigena (Brandegee) Britt. & Rose ssp. cerralvensis, from Isla Cerralvo; Hemizonia
greeneana Rose ssp. peninsularis, from Islas de Todos Santos and the nearby coast: Monardella lagunensis M.
E. Jones ssp. mediopeninsularis, from four peaks in central Baja California; Rhus kearneyi Bark ley ssp.
borjaensis. from the Sierra San Borja: and Rhus kearneyi ssp. virginum, from the mountains of south central
Baja California.
RESUMEN. — Se describen doce taxa nuevos de Dicotiledoneas procedentes de Baja California, Mexico:
Hedeuma martirense, Heterotheca martirensis, Stephanomeria monocephala, y Tanaceium bajacaliforni-
cum, de la alta escarpadura oriental de la Sierra de San Pedro Martir; Eriogonum zapaloense y Castilleja
fruticosa. de la Isla Guadalupe: Eriogonum angelense. de la Isla Angel de la Guarda: Dudleva nubigena
(Brandegee) Britt. y Rose ssp. cerralvensis, de la Isla Cerralvo; Hemizonia greeneana Rose ssp. peninsularis,
de las Islas de Todos Santos y costas cercanas; Monardella lagunensis ML E. Jones ssp. mediopeninsularis, de
cuatro cerros de la parte central de Baja California: Rhus kearneyi Barklev ssp. borjaensis. de la Sierra San
Borja; y Rhus kearneyi ssp. virginum, de las sierras centro-meridionalesde Baja California.
In the course of general collecting in Baja California, I have found several
previously unknown plants, twelve of which are described here. In the citation of
collections, field numbers are mine unless otherwise identified. Each of my recent
collections (above number 5000) is represented in the herbarium of the San Diego
Society of Natural History. Duplicates now being distributed are mostly not cited; they
will go to the herbaria of the Smithsonian Institution, in Washington, the Universidad
Nacional Autonoma de Mexico, in Mexico City, the University of California at
Berkeley, and various other institutions.
Eriogonum angelense Moran, spec. nov. (Fig. 1)
Planta perennis , caudice pauciramoso, erecto, 1-6 dm alto, rosula densa 2-3 ': dm
lata coronato. Folia hirtella, Vi-2 dm longa, lamina ovata, cordata. apice rotundata,
petiolo subaequilonga. Pedunculus nudus, 2 Vi-7 '/: dm altus, 2-3-chotomus, ramis 3-12
dm altis, racemos 2-13 cm longos racemose ferentibus. Involucra solitaria, brevipe-
dunculata, turbinata, 5-lobata, 2-2 Vi mm alta, extus glabra, bracteolas multas lineari-
oblanceolatas glandulosas continentia. Flores ca. 50-120, glabri, flavi, ca. 1 .6 mm longi,
segmentis oblongo-ovatis, interioribus angustioribus. Typus: Moran 10454 (SD 541 861.
Ab aliis subgeneris Ganysmae magnitudine, infiorescentiae forma, copiaque florum
involucri differ t.
Plant perennial, thecaudex erect, simple or few branched above, 1-2 (-6) dm tall, 8-
20 mm thick, covered above with old dried leaves, reddish brown below with persistent
leaf bases, in sterile condition crowned with a rosette 2-3 V: dm wide of ca. 30 close-set
leaves. Leaves green, hirtellous, the trichomes colorless, 0.3- 1.2 mm long; blades ovate,
rounded at apex, mostly cordate at base, 5-1 1 ': cm long. 3-9 cm wide, with 4-6 rather
SAN DIEGO SOC. NAT HIST.. TRANS. 15(17): 265-295. 15 OCTOBER 1969
266
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL.15
V- ■
f
Figure 1. Herbarium specimens of Eriogonum angelense (10421), from Isla Angel de la Guarda, showing sterile
rosette and old floral stem, x 0.23.
prominent ascending veins on each side, the margins irregular and slightly crisped;
petioles 4-11 cm long, 2-4 mm wide at apex, widening to 1-2 cm at the clasping base,
distinctly ca.-9-nerved below. Flowering rosettes elongating to Vi-\ dm, the floral stem
terminal, erect, glabrous, 5-18 dm tall, once dichotomous or often also with a smaller
third branch, the first internode 2 Vi -1 Vi dm tall, 6- 1 1 mm thick, each branch with its first
internode Vi-2 dm long and succeeding ones gradually shorter, each node with one
branchlet, the branchlets ascending, mostly simple, 2-13 cm long (the upper shorter),
with 4-12 nodes. Bracts in threes, shortly connate, oblong, subacute, pubescent
especially ventrally and on margins, the lowest to 1 5 mm long, those of branchlets ca. 1
mm long. Involucres solitary, on peduncles 1-2 (-4) mm long, turbinate, glabrous
without, pubescent within, 2-2 Vi mm high, 5-lobed for one-third their length with
rounded lobes, containing numerous linear-oblanceolate glandular-puberulent bractlets
1 Vi-2 mm long. Pedicels glabrous, 1 Vi-lVi mm long. Flowers ca. 50-120, glabrous
throughout. Perianth yellow, 1 .6- 1 .7 mm long, the segments oblong-ovate, subtruncate,
ca. 1 .2 mm long, the outer 0.7-0.8 mm wide, the inner 0.5-0.6 mm wide. Filaments white,
ca. 0.5 mm long; anthers light yellow, oblong, ca. 0.4 mm long. Ovary yellowish, ca. 0.5
mm long. Achenes not seen.
Type collection. -- Occasional on south side of canyon at about 550 meters
elevation, west slope of peak about 4 miles southeast of Puerto Refugio, Isla Angel de la
Guarda, Baja California, Mexico (near 29°29 '/2'N, 1 13°33'W), 22 March 1963, Moran
1969 MORAN: TWELVE NEW DICOTS
267
W454 — holotype: SD 541 86; isotypes: DS, MEXU.
Distribution. - Known only from Isla Angel de la Guarda at 350 to 800 meters:
rather scarce, in small colonies on ridges and canyonsides or less often in arroyo beds.
Other collections: a dozen plants on N and W slopes on insular divide ca. 2 miles SE of
Puerto Refugio, 500 m, 10421 (SD, UC, US); few in arroyo bed NW of Cerro Angel, 500
m, 12935 (SD); scarce on rocky slopes and in arroyo bed NW of Cerro Diablo, 350 m,
12456 (CAS, SD); half dozen plants on W slope SW of Cerro Diablo, 630 m, 12446 (SD).
Discussion. --At the type locality in March 1963, a few plants were preparing to
flower, but only the very first flowers on one plant were open. At the other places where I saw
the plant on that trip and in early March and late April 1966, none were flowering or
preparing to flower. At each of the five places, however, old floral stems with involucres still
remained.
Eriogonum angelense belongs to the subgenus Ganysma. most species of which are
annuals and much smaller. It appears to differ from all previously known species in its
generally larger size, its open inflorescence whose two or three main branches are essentially
racemes of racemes, and its more numerous flowers per involucre. It is perhaps most closely
related to E. inflatum Torr. & Frem., native from Utah to central Baja California; but it
differs further in its almost palmlike habit, its somewhat larger and non-glandular involucres
on shorter peduncles, and its smaller and glabrous perianth. Eriogonum inflatum var.
deflatum I. M. Jtn. occurs on Isla Angel de la Guarda but apparently only near the shore and
not with E. angelense.
Exploration of the peninsula of Baja California has reduced the list - never long — of
plants known only from islands off its eastern shore, and these islands appear to have few
endemics. Isla Angel de la Guarda, with four still listed, apparently has more than any other.
Besides Eriogonum angelense, these are Ferocactus johnstonianus Britt. & Rose, Penste-
mon clevelandii ssp. angelicus (I. M. Jtn.) Keck, and Hofmeisteria filifolia I. M. Jtn.
Lyrocarpa linearifolia Rollins is known only from Islas Angel de la Guarda and San
Esteban.
Eriogonum zapatoense Moran, spec. nov. (Figs. 2, 8)
Frutex decumbens 1-4 (-7) dm altus, ad 2 Vi m latus, trunco basi ad 1 dm diam. Folia
dense tomentosa, lamina crassa, oblonga, rotundata, 3-6 cm longa, l-2Vi cm lata, in
petiolum 1-3 cm longum et 1 Vi-2 mm latum gradatim angustata. Pedunculi 3-13 cm alti,
inflorescentia cymosa, V2 -2 V2 dm lata, axe 4-8-plo trifurcato. Involucra solitaria, subsessilia
infernave pedunculata, campanulata, 2Vi-4 mm longa, 1 Vi-3 mm lata, ca. 10-40-florata,
extus tomentosa, intus subglabra, dentibus 5, deltoideis, obtusis, '/: mm longis. Pedicelli
glabri. Perianthium flavum, 2Vi-3 mm longum, intus glabrum, extus basi hirtellum,
segmentis obovatis, interioribus exteriora excedentibus. Typus: Moran 15114 (SD 67852).
Ab E. giganteo atque E. molli habitu decumbenti staturaque minor e, foliis crassioribus,
pedicellisglabris, etperianthio flavo recedit.
Shrub commonly prostrate, 1-4 or rarely 7 dm high, V2 -2 V2 m wide, the herbage densely
tomentose, the tomentum at first white and with a few spreading hairs to V2 mm long, at
maturity gray and close. Trunk rarely 10 cm thick at base; branches commonly decumbent,
to 2 m long and 3 cm thick, zigzag, often bare except at summit, reddish brown beneath
shaggy brown outer bark. Branchlets 3-6 mm thick, covered and concealed by persistent
sheathing-decurrent leafbases, each with 5- 1 5 leaves in the upper 2-3 cm. Leaves tomentose,
the blades thick, oblong, obtuse to mostly rounded at apex, 3-6 cm long, 1-21/: cm wide,
conspicuously reticulate dorsally with projecting veins and venules, glabrate ventrally at post
maturity and exposing a surface which in the dried leaf is often orange-brown to dark red.
268
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
*».i
Ik
dOttfiUi mm iff "9
-
It
M. ■.:■,,.
■
Figure 2. Type specimen of Eriogonum zapatoense (\5\ 14), from IsloteZapato, Isla Guadalupe, x 0.25.
gradually narrowed below into a petiole ca. 1-3 cm long and 1 Vi-2 mm wide, the base an
obconic sheath oblique at top. Floral stem plus inflorescence Vi -2 Vi dm high, the peduncle of
one internode 3-13 cm tall, 2-3 mm thick, terminal, growth later continuing sympodially.
Inflorescence cymose, Vi-lVi dm wide, imperfectly 4-8 times trichotomous, some lower
trichotomies about equal but the upper with branches unequal or some suppressed, each fork
with a terminal involucre, or the lower sometimes with a terminal axis bearing 1-2 whorls of
branches and a terminal involucre. Bracts connate at base, the lowest unequal, oblong to
spatulate, 5-25 mm long, 1 '/2-6mm wide, the uppermost equal, triangular-ovate, ca. 1 Vi mm
long. Involucres solitary, sessile or the lower on peduncles to 5 mm long, campanulate, 2 xh -4
mm long, 1 lA-3 mm wide, glabrous within, tomentose without, with 5 triangular-obtuse
teeth ca. Vi mm long, each involucre with ca. 10-40 flowers. Pedicels 2-3 mm long, glabrous.
Perianth yellow, 2'/2-3 mm long, glabrous within, densely hirtellous without at base, the
lobes obovate, rounded at apex, glabrous on the broad thin margins, the outer 1.2-1.5 mm
long, 1.1-1.3 mm wide, the inner 1.5-1.8 mm long, 0.9-1.0 mm wide. Filaments inserted at
top of tube, ca. 1 .3-2.0 mm long, bearded at base; anthers 0.5-0.6 mm long. Achenes brown
ca. 2.2 mm long and 0.8 mm thick; styles ca. 1 .2 mm long; seed dark brown, ca. 1.7 mm long.
Gametic chromosome number: n = 20.
Type collection. Rather common on south and southwest inner slopes of the crater at
50 meters elevation, Islote Zapato (Outer Islet), 2 miles south of Isla Guadalupe, Baja
California, Mexico (near 28°51'N, 118°17'W), 21 June 1968, Moran 15114 holotype:
SD 67852; isotypes: K, MEXU, UC, US, etc.
1969 MORAN: TWELVE NEW DICOTS 269
Distribution. - Known only from Islote Zapato at elevations of 50 to 200 meters.
Other collections: Remple in July 1937 (DS); Moran 2927, March 1948 (CAS, DS, SD);
Kuijt, Miller, & Lindsay 1042, June 1955 (SD, UC); Lindsay 2623, June 1955 (DS).
Discussion. - Islote Zapato, two miles south of Isla Guadalupe, is little more than a
seabound volcanic crater, whose inner slopes rise uniformly at about 45 degrees to meet the
sheer outer seacliffs on three sides in a jagged knife-edge rim of greatly varying height: only
to the east does this sharp rim give place to a narrow and almost flat hanging valley some 200
meters above the sea. Eriogonum zapatoense is common on the inner slopes of the crater,
especially the south slope, and is occasional in the upper valley. In protected places, plants
may grow more or less erect, but most are prostrate. Commonly the trunk or the few main
branches run up slope, so that the plant is strongly asymmetrical. In many larger plants, the
lower branches are dead and the foliage is clustered at the top. The March collection is
sterile, the June collections are in flower, and the July collection is in bud. In June 1968 the
inflorescences were generally smaller than the many remaining from the year before.
Measurements cover the old as well as the new.
Eriogonum zapatoense is most similar to E. molle Greene, of Isla Cedros, Baja
California, and to E. giganteum S. Wats., of Santa Catalina, San Clemente, and Santa
Barbara Islands, California. In leaf shape it especially resembles E. giganteum ssp.
formosum (K. Brandegee) Raven. Both E. molle and E. giganteum grow erect (and E.
giganteum often much taller); and both have somewhat thinner leaves, commonly taller and
stouter peduncles, and much larger and, especially in E. molle, denser inflorescences, with
pubescent pedicels, and with white or pink flowers. My sterile collection of March 1948 was
referred to E. molle ( Moran, 1951).
From buds of the type collection. Dr. James L. Reveal reports a gametic chromosome
number of n = 20.
Although the new species has been found only on Islote Zapato, for which it is named, it
very likely occurs also on precipitous nearby Islote Toro (Inner Islet), where no botanist has
been able to collect. It is one of five species of Islote Zapato never found on the main island of
Guadalupe, the others being Erysimum insulare Greene, Dudleya guadalupensis Moran,
Rhus integrifolia (Nutt.) Rothr., and Lavatera lindsayi Moran. Coreopsis gigantea (Kell.)
Hall, a sixth species of Islote Zapato, was found on the main island in 1 875 but has not been
seen there since. Presumably all six species have been nearly or quite exterminated on the
main island by the goats that have so drastically reduced the native vegetation.
Dudleya nubigena (Brandegee) Britton & Rose ssp. cerralvensis Moran, subsp. nov.
(Fig. 3)
A subspecie typica foliis viridibus modiceve glaucis nee farinosis atque corollis flavis
nee aurantiis usque rubris sat differt. Typus: Moran 36 1 8 (SD 69717).
Caudex unbranched, erect, rarely 6 cm high, \-2Vi cm thick. Rosettes '/2-1 (-2?) dm
wide, of 15-30 leaves. Rosette leaves green or slightly glaucous, triangular-lanceolate to
elliptic-oblong, acute, apiculate, 3-10 (-15) cm long, 12-20 (-35) mm wide, 2-4 (-5'/:) mm
thick, flattish or slightly channeled ventrally, convex dorsally, the margins acute near base,
obtuse above. Floral stems 1-3 dm tall, 2-4 (-5) mm thick, naked in lower 2 Vz-\ 1 cm, with 8-
20 leaves above, the leaves ascending, narrowly triangular-ovate, obtusish, '/2-1 Vi (-3) cm
long. Inflorescence erect from the first, 5-8 cm wide, of 2 or 3 branches which bifurcate 0-2
times but usually once, the cincinni 2-5 cm long, each with 5-14 flowers. Pedicels erect (but
often perpendicular to floral stem when that has declined), ca. 1 mm thick below and thicker
upwards, the lowermost (6-) 10-15 mm long. Calyx 4-6 mm long, 4-6 (-8) mm wide,
subtruncate to rounded below in anthesis, deeply divided, the disk sometimes narrower than
270
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
Figure 3. Dudleya nubigena ssp. cerralvensis. Left: freshly collected plants of the type collection (3618), 4 April
1952, x 0.2. Right: topotype (9542) flowering in San Diego 28 October 1962, x 0.6.
corolla, the segments erect or mostly ascending with tips ca. 1 mm from corolla, triangular-
ovate, acute, 2 l/2-5 mm long, 2 '/2-3 Vi mm wide. Corolla clear yellow, pentagonal, 7 '/2-IO
mm long, 4-5 mm thick at base, 3-4 mm wide at apex, the tube 3-5 V2 mm long, the segments
oblong, 1 Vi-2 mm wide, rounded to broadly acute. Filaments light yellow, subulate, the
epipetalous 6-7 '/: mm long, adnate 2-4 mm, the antesepalous 6 V2-S mm long, adnate 2 V2-
3 V2 mm; anthers light yellow, 1 Vi-2 mm long. Nectar glands whitish, truncate, ca. lA mm
high and 1 mm wide. Gynoecium 6-7 Vi mm high, ca. 3 mm thick, white below, yellowish
above, the pistils erect, appressed, the styles ca. 1 V2 mm long. Ovules 50-80, ca. 0.5 x 0.2
mm. Gametic chromosome number: n = 17.
Type collection. - Locally common on north-facing granitic cliffs at 120 meters
elevation, in the arroyo about 1 mile east of the abandoned Ruffo Ranch, southwest side of
IslaCerralvo, Territoriode Baja California Sur, Mexico (near 24°1 l'N, 109°51'W), 4 April
1952, Moran 3618 holotype: SD 69717; isotypes: BH, DS, UC, US. Some other
duplicates may have been distributed by the Bailey Hortorium as Dudleya sp.
Distribution. - Known only from the type locality. Other collection: 9542.
Discussion. - -To judge from exploration in several parts of the island, this plant is not
common, very likely being confined to relatively cool and shaded habitats, which are scarce.
In April 1952, most plants were flowering (fig. 3, left); in April 1962, a drier season, I saw no
plants flowering and only two with old floral stems apparently of that year.
Plants of the type collection flowered in the greenhouse at Ithaca in November 1952;
some from the second collection flowered in San Diego in October 1962 and again, at what
1969 MORAN: TWELVE NEW DICOTS 271
seemed a more normal season, in the springs of 1964 and 1965. A plant of Dudleva brought
into cultivation may look more different from its former self in the wild than plants of two
different species, grown together, sometimes look from each other. The difference between
wild and cultivated plants here was striking (fig. 3). The description is based on the freshly
collected material and on the cultivated plants. The parenthetic extreme measurements are
mostly from the cultivated plants.
Brandegee ( 1 89 1 ) based Cotyledon nubigena on plants from "the summits of the Sierra
de la Laguna"; but in the herbarium he later identified with it plants from San Jose del Cabo,
near sea level. Rose (Britton & Rose, 1903) separated the lowland plants as D. xantii, the
type specimen from Cabo San Lucas. It appears that his distinctions do not hold but that
there may be other slight differences. Until further living material can be studied, however, I
would include D. xantii in D. n. nubigena, to which it is at least very similar. This subspecies
then is known from a dozen localities scattered through the Cape region of Baja California,
from Isla Espfritu Santo to Cabo San Lucas and from sea level to 1800 meters. It is
characterized by farinose leaves and orange to coral red corollas. Dudleva n. cerralvensis, in
other ways very similar, differs in having green or slightly glaucous leaves and clear yellow
corollas.
From cultivated plants of both collections, Dr. Charles H. Uhl reports a gametic
chromosome number of n= 17 for D. n. cerralvensis. He has found the same number in one
montane and several lowland collections of D. n. nubigena.
Rhus kearneyi Barkley (Fig. 4)
Rhus kearneyi has been known only from the Tinajas Altas Mountains of southwestern
Arizona, where according to Kearney and Peebles (1951) it occurs at 1000 to 1500 feet
elevation. It was at first confused with R. integrifolia (Nutt.) Rothr., of coastal southern
California and northern Baja California. In typical R. kearneyi, the puberulence of twigs
and bracts is sparser than in R. integrifolia and not closely appressed, the leaves are cordate
or subcordate at the base and perhaps a little thinner, the cilia of the sepals are non-
glandular, and the petals are non-ciliate. The twigs, bracts, and leaves, especially the under
surfaces, bear short thick glands about 0.05 mm long, which turn orange-red and may persist
as detached or easily detachable granules, resembling frass. The leaves, described as
glabrous except for the glandular hairs, in fact are sparsely puberulent; and the tiny
trichomes, if not always erect, at least are not closely appressed like the few sometimes found
on the leaf blades of/?, integrifolia.
My collections from Baja California include one from the east base of the Sierra San
Pedro Martir that seems referrable to typical R. kearneyi and several from the mountains
farther south that seem to represent two new subspecies of/?, kearneyi.
For reasons that he did not state, Barkley (1937) considered R. kearneyi most closely
related to R. standlevi Barkley, of southeastern Mexico, and to R. ovata S. Wats., of
Arizona, southern California, and northern and central Baja California. In Arizona,
according to Kearney and Peebles, R. ovata occurs at 3000 to 5000 feet and thus much higher
than R. kearneyi. Both the new subspecies of R. kearneyi occur at higher elevations than the
typical subspecies, and both occur with R. ovata.
Rhus kearneyi ssp. kearneyi
Teledo Canon, E side of Sierra San Pedro Martir, 700 m, 7550 (DS, ICF, SD, UC).
Collected in October, the specimen has only young inflorescences; but the petals in bud are
seen to be non-ciliate. This was distributed as/?, integrifolia.
272
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
CALIFORNIA
^TIJUANA
A R. kearneyi kearneyi
+ R. kearneyi borjaensis
• R. kearneyi virginum
116°
i
114
I
Figure 4. Distribution of Rhus kearneyi.
1969
MORAN: TWELVE NEW DICOTS
273
Rhus kearneyi ssp. borjaensis Moran. subsp. nov. (Fig. 5)
A subspecie typica ramulis dense foliisque sparsim hirtellis fioribusque aliquantum
mqjoribus differ t. Typus: Moran 7999 (SD 6405 1 j.
Shrub Vi-4 m high, the trunk to 1 dm thick. Herbage hirtellous with colorless trichomes
to 0.5 mm or on twigs to 0.8 mm long and also bearing tiny sessile or granular glands that
become orange-red. Branchlets 1 !/2-2'/2 mm thick, reddish brown, glandular and densely
hirtellous. Leaves simple, the petioles 3-6 mm long, 1-2 mm thick, glandular and hirtellous,
the blades coriaceous, ovate to oblong-ovate, rounded to broadly acute at apex but
commonly obtuse, cordate to broadly obtuse at base but commonly subcordate, entire or
rarely glandular-serrulate at ends of main veins, 2 '/2-4 Vi (-6 Vi)cm long, 2-3 (-4 Vi) cm wide,
with thickened and somewhat revolute margins and with about 6-12 conspicuous pale
ascending veins on each side of midrib, dark green and commonly drying brownish above,
paler and often densely glandular beneath, sparsely hirtellous on both surfaces and more
densely so on margins and midrib beneath. Inflorescence terminal, a compact panicle 1 Vi-3
cm long, the branches spicate, densely hirtellous and glandular, flowering in March and
April. Bracts persistent, ovate to rhombic-reniform, acute or obtuse, 2 Vi-1 mm long, 2-3 '/:
mm wide, densely pubescent on both surfaces. Bracteoles ovate, conduplicate, with broadly
scarious margins, 2 '/:-3 mm long, ciliate and somewhat pubescent. Sepals ovate, rounded at
apex, cupped, 2 1/2-3 mm long, 2-21/2 mm wide, ciliate with non-glandular hairs, otherwise
glabrous or sparingly pubescent. Petals white or pink, rhombic-obovate, 4-5 Vi mm long, 2-3
mm wide, slightly puberulent on midrib within, not ciliate. Filaments 1 Vi-2 mm long,
glabrous; anthers ca. 1.5 mm long and 1 .2 mm wide. Ovary pubescent. Fruit 8-11 mm long
and wide, 4-5 mm thick, red, densely glandular, densely pilose with trichomes to 1 mm long.
■ ii wilt.
Figure 5. Left: type specimen of Rhus kearneyi ssp. borjaensis (7999), from the Sierra San Borja. x 0.23. Right:
herbarium specimen of Rhus kearneyi ssp. virginum (1 1780). from the Sierra Santa Lucfa. x 0.23. This collection
has the longest leaves of any.
274 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL.15
Type collection. -Occasional at about 1400 meters elevation near the summit of
Cerro de la Mina de San Juan, Sierra San Borja, Baja California, Mexico (near 28°43'N,
113°38'W), 23 March 1960, Moran 7999 — holotype: SD 64051; isotypes: K, MEXU, UC,
US, etc.
Distribution. --Sierra San Borja, mostly at 1100 to 1650 meters elevation but in
canyons occasionally down to 800 meters. Other collections: Arroyo de la Mina de Santa
Marta, 1100 m (seen to 1450 m), 9764; W rim of Tigre Canon, 1200 m, 9927; Canon el
Terminal, 800 m, 8398; Cerro Quemazon, 1450 m, 8071; Cerro la Sandfa, 1 650 m, 11527.
Discussion. - This plant appears to differ from R. k. kearneyi chiefly in its denser and
much longer pubescence and its slightly larger flowers.
Of the other five species included by Barkley (1937) in his section Styphonia, only R.
standleyi Barkley, from southeast Mexico, and R. muelleri Standley & Barkley, from
northeast Mexico, have spreading pubescence. Rhus standleyi differs from the present plant
in its larger, subsessile, subrugose leaves, which are more closely covered with softer
pubescence and have more strongly revolute margins. Rhus muelleri has broader and more
rotund leaves, smaller sepals, and ciliate petals.
This subspecies seems well isolated from its closest relatives (fig. 4 ). At least in brief
explorations I did not find it in the Sierra San Luis to the northwest or in the Sierra San Juan
or the Sierra San Francisco to the southeast. It is named for the mountain range to which it is
apparently confined. The name "Sierra San Borja" for this range is selected perhaps
somewhat arbitrarily from those used on maps for various parts of the range, the other
names being Sierra de Calmalli, Sierra San Lino, and Sierra San Juan; still other names not
appearing on maps are used locally.
Rhus kearneyi ssp. virginum Moran, subsp. nov. (Fig. 5)
A subspecie tvpica foliis majoribus, acutis subacuminatisve differt. Tvpus: Moran
1 1 664 (SD 59520 ).'
Shrub or small tree 2-5 m high, the trunk to 2 dm thick, the bark rough, brown. Herbage
puberulent with more or less spreading often curved colorless trichomes mostly less than 0.2
mm long and also bearing tiny sessile or granular glands that become orange-red. Branchlets
1 -2 Vi mm thick, reddish brown, at first glandular and densely puberulent. Leaves simple, the
petioles 3-14 mm long, 1-2 mm thick, the blades coriaceous, oblong-ovate, rounded to
acuminate at apex but mostly acute, mostly rounded or subcordate at base, entire or
glandular-serrulate with tooth at end of each main lateral vein, 3-6 (-8 '/2)cm long, 1 '/2-3 (-4)
cm wide, with thickened and revolute margins and with 7-12 conspicuous pale ascending
veins on each side of midrib, dark green above, paler and often densely glandular beneath,
moderately puberulent to subglabrate on veins and sparsely puberulent to glabrate between.
Inflorescence a terminal spike or compact panicle of spikes, flowering in September and
October and rarely in February. Bracts persistent, rhombic-ovate, obtuse to subacute,
cupped, strongly keeled, 2-3 mm long, 2Vi-3 mm wide, appressed pubescent within and
without, pink. Sepals ovate, rounded at apex, cupped, 2-2 Vi mm long, 1 Vi -2 mm wide, pink,
ciliate with non-glandular hairs, otherwise glabrous. Petals white, narrowly obovate,
rounded at apex, 3-3 Vi mm long, 1.2-1.8 mm wide, non-ciliate, puberulent at base within,
otherwise glabrous. Filaments 1-1 Vi mm long; anthers 0.8-0.9 mm long, 0.7-0.8 mm wide.
Ovary pubescent. Fruit not seen.
Type collection. Occasional in arroyo at 1 500 meters elevation, east slope of Volcan
las Tres Virgenes, Territorio de Baja California Sur, Mexico (near 27°28'N, 1 12°35'W), 1 1
February 1964, Moran 1 1664 holotype: SD 59520; isotypes: MEXU, UC, US.
Distribution. -- Upper north slopes and downward along arroyos, Sierra de las Tres
1969 MORAN: TWELVE NEW DICOTS 275
Vfrgenes, Sierra Santa LucTa, and Sierra de la Giganta, Baja California, at elevations of
1000 to 1750 meters. Other collections: N slope near the summit of Cerro Azufre, 1680 m,
1 1640; arroyo on E slope of Cerro Barranco, Sierra Santa Lucfa, 1400 m, 1 1780; Canada de
Quemado, S side of Cerro Giganta, 1000 m, Carter & Moran 5270 (SD, UC); N slope near
summit, Cerro del Barreno, Sierra de la Giganta, 1 350 m. Carter & Moran 5358 (SD, UC).
Discussion. - - The abundance and the associates of R. k. virginum vary from one area
to another. It is fairly common at from 1200 to 1750 meters in the Sierra de las Tres
Vfrgenes, where associates include Quercus ajoensis C. H. Muller, Bernardia incana
Morton, Rhus laurina Nutt., Rhus ovata S. Wats., Ceanothus oliganthus Nutt., Rhamnus
insula Kell., and Aralia scopulorum Brandegee. In the Sierra Santa Lucfa, where I saw it
only locally, associates include Prunus ilicifolia cf. ssp. lyonii (Eastwd.) Raven, Aralia
scopulorum, and Randia megacarpa Brandegee. In the Sierra de la Giganta, associates
include Celtis reticulata Torr. and Acacia goldmanii (Britt. & Rose) Wiggins at the lower
locality, where again I saw it only locally, and Aralia scopulorum at the upper, where I saw
only one tree.
Unlike the other two subspecies, R. k. virginum appears to flower mainly in fall. In the
Sierra de las Tres Vfrgenes in February and the Sierra Santa Lucfa in March, 1964, I found
only one flowering plant, and that with very few flowers; the others lacked even young
inflorescences. In the Sierra de la Giganta in September 1967, the plants were just beginning
to flower.
Rhus k. virginum differs sharply from R. k. borjaensis and resembles R. k. kearneyi in
the sparseness of its pubescence. It differs from both in its leaves, which are mostly acute or
even acuminate and often somewhat larger. Also, the leaves usually dry light green, whereas
in the other two they are darker and often brownish. In R. k. virginum, many though not all
leaves have glandular teeth, the percentage varying in different collections. Such teeth are
poorly developed in R. k. kearneyi and in most collections of R. k. borjaensis, though on one
vigorous shoot of the latter {9764 SD) the unusually large leaves have well developed teeth.
Lack of material prevents a thoroughgoing comparison of floral details, but at least the
anthers of R. k. virginum seem smaller than in the other two, and the sepals are glabrous.
Hedeoma martirense Moran, spec. nov. (Fig. 6)
Herba glandulosa hispidula perennis, rhizomata gracilia elongata emittens, caulibus
gracilibus Vi-l Vi dm altis. Folia brevipetiolata, ovata adelliptica. obtusa, 3-9 mm longa, 2-6
mm lata, supra glabra, margine revoluta. Cymulae paucae, unifiorae. Calyx subcylindricus,
7-9 mm longus, fauce dense annulatus, dentibus duobus inferioribus 2-3 mm longis, erectis,
tribus superioribus tertia parte connatis, I Vi-2 mm longis, aetate excurvatis. Corolla alba,
19-24 mm longa, tuba gracillima, cum fauce 17-20 mm longa. Typus; Moran 15069 (SD
67322). Inter congeneros corollae tubo perlongo distinguitur, rhizomatibusque gracilibus
elongatis insignis. H. tenuifloro affinis sed foliorum petiolis brevioribus laminisque supra
glabris inflorescentiaque reductapraeterea differ t.
Low rhizomatous perennial herb with wiry stems, the herbage with glands secreting tiny
yellow globules at the surface, hispidulous with often curved and often unequal tapering
pericellular white trichomes to 0.6 mm long, in sunny places stems and undersurfaces of
leaves often purplish red. Rhizomes woody, to 5 mm thick, with dark brown bark; aerial
shoots often branching below and often crowded, flexuous, '/2-1 Vi dm tall, Vi-V* mm thick,
glandular and hispidulous with more or less decurved trichomes, quadrangular, the two faces
below each leaf pair slightly convex, the other two channeled, the upper internodes ca. 3-5
mm long; young rhizomes like the aerial shoots but to 3 dm long, stramineous, often scarcely
or not at all hispidulous, glandular-puberulent or (only after the first season?) glabrous, the
276
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
Figure 6. Pressed specimen of" Hedeoma martirense ( 15275), from the high Sierra San Pedro Martir, xO.75.
internodes !/2-l V2 cm long, the scale leaves sessile, ovate, blunt, ca. 1 mm long. Leaves
petiolate, the blade coriaceous, from ovate in the lower to elliptic in the upper, obtuse to
rounded at apex, cuneate to rounded at base, entire, 3-5 (-9) mm long, 2-3 (-6) mm wide,
dorsally glandular-pitted or subrugose and sparsely antrorse-hispidulous, ventrally eglandu-
lar and glabrous or scabrous with a few stout conic trichomes ca. 0.05 mm long, the margins
revolute, the midrib slightly projecting dorsally, the 2-3 pairs of ascending lateral veins
obscure, the petiole 1-2 mm long, channeled ventrally. Flowers May to August, the cymules
at 1-4 upper nodes, one-flowered, the peduncle Vi-2 mm long, with an apical pair of linear
bracts 1-2 mm long, the pedicel 2-3 mm long, spreading. Calyx 7-9 mm long, commonly
purplish, the tube subcylindric but slightly ampliate upward and in fruit slightly swollen in
lower third, 4 '/2-6 Vi mm long, ca. 1 mm wide (pressed to 1 V2 mm wide), with 13 prominent
ribs, sparsely hispidulous on ribs towards the base with trichomes ca. 0.3 mm long and
glandular puberulent, glabrous within except for a dense ring at the throat of upwardly
appressed and partially exserted stiff white trichomes ca. 0.6 mm long, the teeth triangular-
lanceolate, sparsely hispidulous-ciliate, the lower two erect, 2-3 mm long, subulate above,
the upper three connate one-third, 1 lA-2 mm long, slender-acute, in age somewhat
outcurved. Corolla white or slightly tinged with pink, 19-24 mm long, glabrous in lower 3-4
mm, hirtellous above with trichomes ca. 0.2 mm long, the tube 14-15 mm long, ca. Vi mm
wide below and scarcely wider above, sparsely hirtellous within over more or less of upper
half, the throat 3-5 mm long, 1 Vz-2 mm wide above, glabrous within, the upper lip 2-3 mm
long, 1 '/2-2 mm wide, emarginate to a depth of 0.2-0.6 mm, minutely papillose-puberulent
within, the lower lip 3-41/2 mm long, 3-4 mm wide (spread out), minutely papillose-
puberulent, the middle lobe 1 V2 -2 mm long, 1 xk mm wide, slightly emarginate, upcurved, the
lateral lobes ca. 1 mm long and wide. Filaments ca. 3 mm long, exserted ca. 1 mm, glabrous;
anthers divaricate, purple, ca. V2 mm long; vestiges of other stamens ca. 0.3 mm long. Style
18-23 mm long, glabrous, the posterior lobe about half the anterior or subobsolete. Nutlets
reddish brown, elliptic, 1 .4- 1 .6 mm long, 0.6-0.7 mm wide, 0.5 mm thick.
1969 MORAN: TWELVE NEW DICOTS 277
Type collection. - Rock crevices on east slope at 2700 meters elevation, east rim of the
Sierra San Pedro Martir north of Yerba Buena, Baja California, Mexico (near 31°01'N,
115°26'W), 1 June 1968, Moran 15069 holotype: SD 67322; isotypes: K, MEXU, UC,
US, etc.
Distribution. Abundant locally on north and east slopes at 2300 to 2800 meters
elevation, especially in shade but also in the open, north central Sierra San Pedro Martir.
Other collections: E slope at head of Arroyo Copal, 2300 m, 15463; N slope of Cerro "2828".
2575 m, 15446; E slope of Cerro "2828", 2800 m,15266, 2750 m, 15275.
Discussion. - This plant grows most luxuriantly on north slopes in open pine-fir forest,
where it may spread among rocks to form a rather uniform though scarcely dense
groundcover. On the steep eastern escarpment of the Sierra, it grows also on drier and more
open slopes and, at its lower limit, into the border of the montane chaparral.
Hedeoma martirense is noteworthy for its slender rhizomes and somewhat matting
habit. The long slender corolla is also remarkable, the tube plus the throat being 1 7 to 20 mm
long and the tube proper less than 1 mm wide. To judge from the descriptions of Epling and
Stewart (1939), the length is approached only in H. costaium A. Gray, where it is 6 to 18 but
commonly 1 1 to 15 mm, and in H. quercetorum Epling, where it is 14 to 16 mm (less in a
recent collection, annotated by Robert S. Irving). The variable H. costatum, ranging from
Arizona to Texas and northern Mexico, is a somewhat coarser plant, with longer internodes
and larger leaves; the leaves are serrate and more acute, pubescent on both sides, and thinner,
with more prominent veins; and each cymule has three to several flowers. In H. quercetorum,
of northeastern Mexico, the stems are less branching and less flexuous, the pubescence of
stems and leaves is shorter and finer, each cymule has one to several flowers, the calyx tube is
curved, and the upper leaves are longer and linear-elliptic, with prominent lateral veins more
or less parallel to the midrib. However, the lower leaves are similar to those of//, martirense,
being ovate with revolute margins, rather thick with the veins often obscure, purplish red
beneath, and nearly as small.
Epling and Stewart divided the genus into twelve unnamed but numbered sections.
Though not exactly fitting their sectional definition, H. martirense seems closest to section
9, in which they placed H. costatum, H. quercetorum, and five other species. It is perhaps
most closely related to the little-known H. tenuiflorum Brandegee, of the Sierra San Borja,
some 175 miles to the south-southeast in Baja California. That is a bushier plant without
rhizomes, with similar but more elliptic and sometimes toothed leaves hispidulous on both
sides and with longer petioles; the cymules commonly are more numerous and commonly
have three to several flowers each; and the corolla tube is about 13 mm long. That species
occurs in a more arid habitat, not only farther south but also at lower elevations - at about
1475 meters and probably well below; associates include Quercus turbinella Greene,
Anemone tuberosa Rydb., Ribes quercetorum Greene, Vauquelinia calif ornica (Torr.)
Sarg., Dodonaea viscosa Jacq., Rhamnus insula Kell., Idria columnaris Kell., and
Echinocereus engelmannii (Parry) R'umpl.
Monardella lagunensis M. E. Jones ssp. mediopeninsularis Moran, subsp. nov.
(Figs. 7, 12)
Suffrutex fragrans glandulosus hirtellus, trichomatibus plerumque 0.1-0.2 mm longis,
caulibus gracilibus 3-8 dm altis e basi 1-4 cm crassa. Folia triangulo-ovata. apice rotundata.
8-20 mmlonga, 4-12 mm lata, petiolis 1-8 mm longis. Capitula Vi-2 Vi cm lata ca. 20-125-
flora, bracteis adpressis submembranaceis, exterioribus ovatis 5-8 mm longis, interionbus
angustioribus. Calyx 6-7 Vi mm longus, 13-nervatus. Corolla 9-10]: mm longa. Typus:
Moran 12147 (SD 70685). A subspecie typica foliis parvioribus capitulis parvioribus ex
278
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL.15
Figure 7. Pressed specimen of Monardella lagunensis ssp. mediopeninsularis, from Cerro Potrero, x 0.5; part of
the type collection (12147).
floribus paucioribus parvioribusque constantibus trichomatibusque brevioribus differt.
Fragrant subshrub 3-8 dm high, with many slender branches from a woody base 1-2 or
rarely 4 cm thick. Herbage hirtellous with slightly curved pluricellular white trichomes 0.1-
0.2 mm long or on vigorous shoots rarely (10460) to 0.5 mm long; also glandular, the surface
shallowly pitted, each pit supporting a viscid yellowish globule ca. 0.05-0.1 mm thick.
Branches often taking two seasons to flower, often with lateral branches below by flowering
time, at first ca. 1 mm thick and closely hirtellous with slightly deflexed trichomes, dying
back about to middle after flowering, becoming woody below but rarely over 4 mm thick
except at very base, the bark tan in new fissures, becoming gray, the lower internodes ca. Vi-
1 Vi cm long, the upper longer, sometimes to 6 cm. Leaves petiolate, the blade triangular-
ovate, rounded at apex, rounded to cuneate or commonly somewhat attenuate at base, with
2-3 weak crenations on each side or commonly entire, 7-20 mm long, 4-10 (-12) mm wide,
equally hirtellous above and beneath but more conspicuously pitted beneath, the midvein
prominent beneath, with 2-4 arching laterals on each side, the margins somewhat revolute,
the petiole 1-3 (-8) mm long. Heads in May and June or a few as early as February, 1 '/2-2 ( Vi-
1969 MORAN: TWELVE NEW DICOTS 279
2 Vi) cm wide, each with ca. 20-125 flowers, the bracts appressed, thinner than leaves and
somewhat membranous, with prominent veins, often purplish, glabrous ventrally except
toward apex, hirtellous dorsally and closely ciliate with longer trichomes, the outer ovate,
acute to narrowly rounded, 5-8 mm long, 2-5 mm wide, with lateral veins arching and nearly
parallel to midrib, the inner oblanceolate or narrower, acute. Calyx 6-7 Vi mm long, ca. 13-
nerved, often purplish, hirtellous, the teeth erect, narrow-triangular, acute, nearly equal,
1 '/4-2 mm long, hirtellous within. Corolla white or slightly tinged with pink or lavender, 9-
10 Vi mm long, the tube 6-7 mm long, gradually ampliate above, sparsely retrorsely
hispidulous inside and out in upper half, the lobes oblong, rounded at apex, 2 Vi-1 Vi mm
long, 0.6-0.9 mm wide, sparsely hispidulous outside and glandular, with 3-7 sessile yellow
globules ca. 0.1 mm thick crowded at tip, the upper two lobes united ca. Vi- Vi, the lower
three free. Filaments glabrous or with a few hairs at base, apparently elongating markedly
during anthesis, the lower two inserted near sinuses, 3-5 mm long, the others inserted on base
of upper lip, 2-3 Vi mm long; anther sacs divergent. Style glabrous, apparently elongating
markedly during anthesis, becoming ca. 1 1 mm long, the stigma lobes ca. 0.3 mm long.
Nutlets oblong, tan, smooth, 1 . 1- 1 .4 mm long, 0.6-0.8 mm wide.
Type collection. -Common among rocks, mostly on the north slope, summit of Cerro
Potrero at 1400 meters elevation, Baja California, Mexico (near 29°49'N, 114°37"W), 30
May 1965, Moran 12147— holotype: SD 70685; isotypes: K, MEXU, UC, US, etc.
Distribution. - - Known only from four peaks in central Baja California (fig. 12), at 750
to 1900 meters elevation. Other collections: N slope of Cerro Santa Maria, 1200 m, 1 1481
[sterile] ; steep north-flowing arroyo on W side of peak ca. 4 miles SE of Puerto Refugio, Isla
Angel de la Guarda, 900 m, 10460; arroyo on E slope of Volcan las Tres Virgenes, 1 500 m
[seen to 1900 m],/ 1666.
Discussion. - The four known localities for this plant are scattered over some 200
miles (fig. 12). I have partially explored various peaks in the intervening Sierras San Luis,
San Borja, San Juan, and San Francisco, without finding it. At the three northern localities,
it was found mainly or entirely on north slopes and at the southernmost, so far as I
remember, mainly so.
The new subspecies differs from M. I. lagunensis in its smaller leaves, its smaller heads
with fewer and smaller flowers, and its shorter trichomes. The leaf blades in M. I. lagunensis
are mostly 12 to 22 mm long and in M. I. mediopeninsularis mostly 8 to 15. However, in the
northernmost (and type) collection, with abundant material found in better condition than
the other collections, a few vigorous shoots have leaf blades to 20 mm long. The heads in M.
I. lagunensis are mostly 2 to 3 xh but sometimes as little as 1 Vi cm wide, in M. I.
mediopeninsularis mostly 1 Vi to 2 but occasionally 2 xh cm wide. The flowers of M. I.
mediopeninsularis are similar to those of M. I. lagunensis but proportionally smaller, the
calyx, for example, 6 to 7 '/2 mm long as compared to 7 to 9 an J the corolla 9 to 101/: mm long
as compared to 1 1 to 12. In M. I. lagunensis the longer trichomes are about 0.5 to 0.6 mm
long. In the southernmost collection of M. I. mediopeninsularis the longer trichomes are
about 0.2 mm long; in the next most southern collection (actually from closer to the two
northern localities) they are mostly about 0.2 mm but on one vigorous shoot 0.4 to 0.5 mm
long; and in the two northern collections they are about 0. 1 mm long. Monardella linoides A.
Gray, of the Sierra San Pedro Martir and northward, might be said to continue the trend in
decreasing trichome length, having one-celled trichomes about 0.05 mm long; but that is a
distinct species, with narrower leaves and longer corollas.
Smaller-leaved specimens of M. 1. mediopeninsularis resemble M. thymifolia Greene,
of Isla Cedros; but in that species the leaf blades are 5 to 10 mm long, the trichomes about 0.5
mm long, and the corollas 12 to 14 mm long, with the tube conspicuously exserted from the
280
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL.15
Figure 8. Old plant of Eriogonum zapatoense, on Islote Zapato, Isla Guadalupe, showing crooked decumbent
stem; part of the type collection (151 14), 2 1 June 1968.
Figure 9. Herbaceous young plant of Castilleja fruticosa (6687), south end of Isla Guadalupe, 27 March 1958.
Note thickness of leaves.
1969
MORAN: TWELVE NEW DICOTS
281
Figure 10. Flowering plant of Heterotheca martirensis, in the high Sierra San Pedro Martir, 14 September 1968:
part of the type collection (15612).
Figure 1 1. Flowering plant of Stephanomeria monocephala. in the high Sierra San Pedro Martir. 5 July
part of the type collection (15261 ).
1968;
282
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL.15
Figure 12. Distribution of Monardella lagunensis and related species.
1969 MORAN: TWELVE NEW DICOTS 283
calyx. In his monograph, Epling( 1925) based his description of M. thymifolia on about eight
sheets of four collections, mostly past flower. He commented that the few available
specimens were uniform in appearance, and his description bespeaks uniformity (e.g. leaves
5-8 mm long, calyx 6-7 mm long, corolla 12-13 mm long). It is noteworthy that my two
collections of the species, from stations well separated on Isla Cedros, agree rather closely
with his description and so reinforce the appearance of uniformity. Monardella lagunensis
seems quite distinct from M. thymifolia.
Castilleja fruticosa Moran, spec. nov. (Figs. 9, 13)
Planta perennis, glanduloso-puberula villosaque, 2-9 dm aha, caule lignoso, ad 4 cm
diametro. Folia crassa, linearia, Vi-6 cm longa, 1-5 mm lata, basi articulata, integra
supremave trilobata. Spica densa, 2 Vi-7 cm longa, 5-20-flora, miniata, bracteis 1-2 Vi cm
longis, plerumque trilobatis. Calyx 14-21 mm longus, antice posticeque 8 Vi-14 mm latere 0-
4 Vi mm fissus, lobis subaequalibus, triangularibus usque oblongis. Corolla 1 7-26 mm longa,
infra glabra, galea 9-13 mm longa, postice lutea dense glanduloso-puberula, marginibus
hyalinis, anguste revolutis, labio inferiore 2-3 mm longo. Capsula 7-10 mm longa, lignosa.
antice concava usque transverse sulcata, sero dehiscens. Typus: Moran 15733 (SD 70310).
Species caulibus duro-lignosis atque capsulis lignosis subindehiscentibus insignita. A C.
guadalupensi pubescentiae forma (ilia tomentosa) et calyce antice posticeque aequaliter
fisso valde differt.
Woody perennial 2-5 or rarely 9 dm high and to 5 dm or more wide, but flowering while
still herbaceous, perhaps in first season. Herbage glandular-puberulent, with trichomes
mostly 0.05-0.3 mm long, and in part also villous, the longer trichomes pericellular,
tapering, unbranched, 1-2 mm long. Stem at first straight and erect, with several regular
horizontal to ascending branches, with sympodial growth after flowering becoming crooked
and irregularly close-branching, to 2 or rarely 4 cm thick, the wood hard and brittle, the bark
mostly dark brown, the few longitudinal fissures lighter brown; branches villous. Leaves
light green, rather succulent and the smaller ones subterete, linear-oblong to linear-
oblanceolate, rounded to broadly acute, entire or the uppermost 3-lobed, articulated to a
decurrent base and readily detached, Vi-\ Vi mm wide at constriction, glandular-puberulent
and the upper commonly somewhat villous near base, commonly Vi-2 Vi cm long, 1 Vi-3 mm
wide, and 1-2 mm thick, the upper ones in favorable seasons sometimes larger, even 6 cm
long and 5 mm wide, relatively thinner, and less clearly articulated at base; midvein
translucent, often impressed ventrally and in thicker leaves sometimes also dorsally, the
lateral veins not evident. Inflorescence terminal, a compact spike 2'/2-7 cm long, of 5-20
flowers, often densely villous, the bracts and calyx tipped with Indian orange, the pigment
especially concentrated in the glandular trichomes; dead axis commonly persisting one or
more seasons. Bracts 1-2 Vi cm long, Vi-2 cm wide, generally glandular-puberulent and
mostly somewhat villous, especially on lower margins and midribs, the lower ones
sometimes entire and the uppermost sometimes 5-lobed but most 3-lobed, the lobes oblong
to mostly spatulate, obtuse to rounded, the midlobe 3-9 mm wide, the lateral lobes ascending
from near the middle, 1-3 Vi mm wide. Calyx glandular-puberulent, villous below, especially
on nerves, 14-21 mm long, about equally cleft before and behind for 8'/2-12 mm, the
segments 2 Vi -4 mm wide, entire or mostly cleft 1 -4 Vi mm into subequal triangular to oblong
acutish lobes 1-2 mm wide, the anterior often slightly wider than posterior. Corolla 17-26
mm long, with 19 veins, glabrous below, sparsely glandular-puberulent towards middle;
lower lip yellowish green, glandular-puberulent especially above, projecting abruptly
anteriorly for 1 Vi-2 mm and cupped upward for 2-3 mm, the apex 9-14 mm above corolla
base, the lobes ca. 1 mm long and wide, acutish to rounded, the palate of two finlike yellowish
284
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL.15
glandular evaginations ca. 1-1 Vi mm long and 3/4-l mm high, each continuing upward as a
low ridge for ca. 1-1 Vi mm to the sinus; galea 9-13 mm long, when flattened 4 V2-8 mm wide
at base and ca. 3 mm wide just below apex, exserted ca. 2-6 mm beyond calyx, slightly curved
anteriorly, densely glandular-puberulent dorsally above, dorsally yellowish green becoming
yellow and sometimes later reddish, the hyaline margins ca. 1 mm wide above and wider
below, sharply differentiated above from the herbaceous center, closely rolled outward
below, the apex rounded. Stamens glabrous, the anterior filaments 7 Vi - 1 6 mm long, inserted
7 '/2-9 mm above corolla base, the posterior 7-15 mm long, inserted 6 V2-8 mm above corolla
base; anthers 2-2 Vi mm long, ca. Va mm wide. Ovary glabrous, greenish, laterally com-
pressed above and obliquely obovate, 2 V2 -4 mm long, 1-1 Vi mm wide, 1 Vi-2 mm thick,
the rounded apex sometimes extending beyond the style base; style a continuation of the
posterior keel, yellowish green, 17-25 mm long, exserted before anthesis and finally by 2-4
mm. Capsule woody, blackish, triangular-ovoid, tapering in width apically to a sharp
sagittal keel continuous with low anterior and posterior keels, 7-10 mm long, 3 V2S mm wide
and thick, the anterior surface concave to transversely and sometimes tightly infolded, with a
furrow to 2 mm deep; persisting on dead inflorescence for one or more seasons mostly
unopened even though containing well-formed seeds, the two cells occasionally separating
slightly at apex and less often also the valves splitting lengthwise in upper fourth. Seeds dark
brown, irregularly obovoid, 1 .6-2. 1 mm long and 1 .0- 1 .2 mm wide including loose alveolate
coat.
Type collection. - Occasional on barren mesa at 100 meters elevation, just east of the
weather station, southwest corner of Isla Guadalupe, Baja California, Mexico (near
28°53'N, 1 18°17 '/4'W), 23 February 1969, Moran 15733- holotype: SD 70310; isotypes: K,
MEXU,UC,US,etc.
Figure 1 3. Pressed specimen ofCastillejafruticosa (7850), from the south end of Isla Guadalupe, showing flowers
and persistent capsules, x 0.75.
1969 MORAN: TWELVE NEW DICOTS 285
Distribution. - - Local at the south end of Isla Guadalupe and on offlying Islote Zapato
(Outer Islet). Other collections: S inner slope of red crater 1 mile N of Morro Sur, 320 m,
6479; 1 Vi miles NE of weather station, 220 m, 7850; type locality, 6142, 6687, 12068, 13768,
Wiggins & Ernst 136 (DS, US), Carlquist 482 (RSA), Copp 175 (DS); Islote Zapato,
Remple in 1937 (DS), Moran 2922 (CAS, DS, SD), 5657, 15113, Lindsav 1805 (DS, SD),
2624 (SD), in 1956 (SD), Kuijt, Miller, & Lindsay 1050 (UC).
Discussion. This plant occurs on a barren volcanic mesa and in two craters near the
south end of Isla Guadalupe, in an area of very low rainfall but of frequent fogs. Though
fairly common locally, it is thus very narrowly restricted. In 1948 I collected it on Islote
Zapato but failed to find it on the main island, even though I collected at the very spot on the
mesa where its bright flowers are now conspicuous. In 1957 I did find it on the mesa but saw
only herbaceous plants looking almost like annuals. In 1960 it was a little more widespread
there than I have seen it before or since, and in 1 967 and later, some plants were small shrubs
with woody stems to 2 cm thick. These facts suggest that in 1957 C. fruticosa might recently
have colonized the main island from one of the nearby islets. Although the goats that have
ravaged the northern part of the island may never have been common at the desertic south
end, comparison of the vegetation with that of the islets suggests that enough goats came to
keep certain plants near the point of extinction. Since the establishment of the weather
station in 1946, however, goats have visited the south end at their peril, and the vegetation
thus has had some protection.
Among associated species, several are similarly restricted on the island. I have seen
Euphorbia pondii Millsp. only in an even smaller colony in the same part of the mesa. The
endemic Hemizonia palmeri Rose is confined to a slightly larger area within about one mile
of the south end. The endemic H. greeneana Rose ssp. greeneana, one of the most abundant
and conspicuous plants at the south end, extends north only about two miles; it occurs also on
Islote Zapato and Islote Negro. The endemic Baeriopsis guadalupensis J. T. Howell has
been found only at the south end and on Islote Zapato and Islote Negro, though possibly it
occurs farther north on unexplored seacliffs. Hutchinsia procumbens (L.) Desv. has been
found only at the south end and on Islote Zapato. Euphorbia misera Benth., common on
Islote Zapato and very rare on cliffs in the northern canyons, also occurs rarely on the south-
end mesa. Lomatium insulare (Eastw.) Munz likewise occurs on Islote Zapato and on the
south-end mesa, and it occurs also on one cliff high at the north end of the island. These last
two species, like Castilleja fruticosa, may possibly be recent colonists at the south end. A
recent unsuccessful colonist there was Lavatera occidentalis S. Wats., which is common on
Islote Zapato and is represented by two or three old shrubs on northern cliffs: a single robust
seedling 1 meter high grew in an arroyo 3 miles from the south end in 1965, when the goat
population was at a low ebb, but has since disappeared.
Castilleja fruticosa, though still herbaceous at first flowering, is remarkable for the
thick and hard woody stems of older plants. Apparently it is one of the woodiest of the genus.
The foliage leaves are quite thick and somewhat succulent, as is often true of maritime
plants. The capsules are unusually hard and woody and are distinguished by a depression on
the under side, which sometimes becomes a deep transverse groove, as if the capsule were
caved in. On most specimens, capsules from previous years persist on dead inflorescence
axes, mostly still unopened and still holding seeds. Seeds from closed capsules have proven
viable.
Castilleja fruticosa falls in the section Perichroma Pennell (Pennell, 1935). The only
recent treatment of any major part of the genus is that of Pennell ( 1 95 1 ) for the 7 1 species of
Washington, Oregon, and California — out of some 200 species all told. The genus is poorly
represented in Baja California, with apparently only four species, including two Guadalupe
286 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL.15
Island endemics, that do not occur also in Alta California; and the flora of Guadalupe Island
is more closely related to that of Alta California than to that of peninsular Baja California.
Nevertheless, the new species does not fit clearly into any of the 19 groups (apparently
subsections) informally proposed by Pennell (1951). However, it is close to groups x through
xvi, perhaps differing from all their species in having woodier stems and woodier and often
tardily dehiscent transversely grooved capsules but otherwise differing from each mainly in
the kind and distribution of pubescence.
This plant was first reported (Moran, 1 95 1 ), and has since been identified in herbaria, as C
guadalupensis Brandegee, another Guadalupe Island endemic with hard woody stems. That
plant occurs (or occurred) in a different habitat, on the higher and much moister northern
part of the island. It has not been collected since 1898 and evidently is now very rare if not
extinct. Herbarium material is meager.
The little-known C. guadalupensis is a woody plant 3 to 6 dm tall, whose stems, leaves,
and calyx are at first densely and closely tomentose with dendritic hairs. The leaves are
narrowly spatulate, rather thin, to 6 cm long and 1 cm wide. Brandegee (1903) described the
calyx as "cleft equally before and behind about one-half its length", as in the presumably
related C.foliolosa H. & A.; but in three fairly well exposed flowers of one Palmer specimen
(GH) it clearly is cut about twice as deeply behind as before — as in no other species known
to me. The corolla is slightly longer to slightly shorter than the calyx; the galea, about
equalling the tube, is densely puberulent the length of the back with pericellular but
unbranched non-glandular trichomes about 0. 1 to 0.2 mm long. The style is exserted about 3
to 5 Vi mm. Clearly this is a very distinct species.
Castilleja fruticosa thus is amply different from C. guadalupensis, notably in the cutting
of the calyx but also in the type of pubescence and in the form and size of the leaves.
Hemizonia greeneana Rose ssp. peninsularis Moran, subsp. nov. (Fig. 14)
Frutex glanduloso-viscidus prostratus vel erectus turn quoque ad 12 dm alius ramis
virgaiis. Folia inferiora subremota filiformia usque linearo-oblanceolata Integra vel 2-8-
dentata hirsuta 2-7 cm longa, superiora conferta linearia plerumque hirtella Vi-1 cm longa
V2-I mm lata. Capitula thyrsoidea plerumque 8-radiata, primis usque 14-radiatis. Involucra
4-7 mm alta, 4-6 mm diametro, bracteis receptaculi 8-13 uniseriatis ad medium infirme
connatis. Disci flores 8-21 steriles, pappo 1.9-3.3 mm longo, paleis 9-15. Typus: Moran
13437 (SD 70684). A ssp. greeneana ramis plantarum altiorum virgatis, foliis infimis saepe
longioribus remotioreque dentatis, superis plerumque hirtellis, pappo plerumque longiore
paleisque numerosioribus differ t.
Prostrate or erect shrub 2-8 or reportedly to 12 dm high and Vi-\ Vi m or more wide,
with herbage glandular-viscid throughout. Stems to 4 cm thick at base, the bark gray-brown.
Branches commonly virgate, hirsute with multicellular white trichomes to 3 mm long, rather
sparsely leafy below, the internodes ca. Vi-\ V2 cm long, the axils sometimes later floccose,
only the lowermost leaves opposite, the middle axils often with fascicled leaves, the upper
mostly with leafy branchlets; ultimate branchlets or peduncles ca. V2 mm thick, hirtellous,
closely leafy, with shorter internodes and fascicled leaves. Lower leaves filiform to linear-
oblanceolate, acute, 2-7 cm long, V2S mm wide, with subrevolute or thickened margins,
entire or often with 2-8 ascending acute teeth or lobes to 5 mm long and 1 Vi mm wide, hirsute
with trichomes to 1 mm long, somewhat succulent and to 2 mm thick in plants nearest the
shore; upper leaves gradually smaller, entire, those of the peduncles linear-oblong, obtuse,
ca. '/2-l cm long and Vi-\ mm wide, sparsely hirtellous or subglabrous except for glands.
Heads borne sometimes at least from March to November near the shore but mostly in
summer above, often numerous and subcorymbose at ends of main branches, solitary or few
1969
MORAN: TWELVE NEW DICOTS
287
i
Figure 14. Type specimen of Hernizonia greeneana ssp. peninsularis ( 13437), from Punta Banda. xO.25.
and cymose on the branchlets, yellow, 1 1-1 8 mm wide, mostly 8-rayed, but some early ones
with more rays and the first, terminating the main branches, with as many as 14. Involucre
campanulate, 4-7 mm high, 4-6 mm wide, sparsely hirsute, beset with peglike glands ca. 0.05
mm high and each tipped with a sticky yellowish globule ca. 0.1 mm thick, the bracts
lanceolate, acute, cymbiform and strongly keeled in lower three-fifths and ca. 2 Vi mm wide
(flattened), with hyaline margins, narrowed to a flatter thickish apex 2-3 mm long. Ray
florets 8-14, the tube, 1 V2-2V2 mm long, stipitate-glandular, the ray oblong to obovate,
truncate and 2-3-crenate at apex, 4 V1-6V2 mm long, 2 '/2-3 Vi mm wide, subglabrous, the style
branches slender, 2-3 mm long. Receptacular bracts 8- 1 3 (about as many as the rays), in one
series, weakly united to middle, oblanceolate, narrowly acute (ca. 30°), 5-6 mm high, to 1 Vi
mm wide, herbaceous with hyaline margins, glandular, the apex sparsely long-ciliate. Disk
florets 8-14 (-2 1 ), the corolla 4-5 mm long, the tube slightly ampliate above, subglabrous. the
lobes triangular-ovate, 3/4-l mm long, thickened and densely puberulent ventrally on
margins, the anthers yellow, ca. 2 Vi mm long, the stigma lobes ca. 2 mm long, the achenes
sterile, 2-2 Vi mm long, sparsely glandular-puberulent, the pappus 1.9-3.3 mm long, the
paleae 9- 1 5, free or united at base, unequal, irregularly lanceolate or oblong, stiff, white. Ray
achenes 2-3 mm long, triquetrous, black, transversely rugose, acute and stipitate at base,
rounded at apex, with an upcurved lateral beak 1 -2 or rarely 3 times as long as thick.
Type collection. - Occasional on east, north, and west slopes at 380 meters elevation.
SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
summit of Banda Peak, Punta Banda, Baja California, Mexico (near 31°441/2,N,
116°433/4'W), 27 August 1966, Moran 13437 — holotype: SD 70684; isotypes: K, MEXU,
UC, US, etc.
Distribution. - - Local on the northwest coast of Baja California southwest of Ensenada
and midway between there and Tijuana. Other collections: northfacing cliff at mouth of Rfo
San Miguel (or Guadalupe), 5 m, Higgins in 1946 (SD), Howe in 1964 (SD), Moran 13155,
13431, 14633, 15376, 16000; rocky cliffs near ocean 37 miles S of border [probably the same
locality], Campbell in 1937 (POM); Isla Sur, Islas de Todos Santos, Brandegee in 1897
(UC?), Moran 2803 (SBBG), 16210, Blakley 6590 (SBBG, SD), 6598 (SBBG, SD); N side of
Punta Banda near tip, 5 m, 14638; type locality, 15914.
Discussion. - The new subspecies apparently is local on cooler parts of the coast,
occurring chiefly on Punta Banda (Banda Promontory) and on the southern one of the Islas
de Todos Santos, a disconnected part of that promontory. Otherwise it has been found only
on one north-facing beach cliff 20 miles to the north. The annual rainfall on the Islas de
Todos Santos is about 10 inches (Hastings, 1964), and the coastal area is subject to frequent
fogs. In contrast, H. g. greeneana is endemic to the south end of Isla Guadalupe and the
offlying islets, a hot dry area, again with frequent fogs but with an annual rainfall of perhaps
less than 2 inches.
This plant was first collected in 1897 on the Islas de Todos Santos by T. S. Brandegee
(1899, 1900), though his specimens cannot now be found. He first thought it seemed "the
same as specimens named H. frutescens Gray, collected by Dr. Palmer on Guadalupe Island,
although neither of them exactly agrees with Dr. Gray's description of that species". But
then, evidently from the same collection, he reported H. greeneana as "growing plentifully
over the larger [southernmost] of the Todos Santos Islands." The Palmer specimens he
mentioned must have been part of the type collection of//, greeneana, but it is not clear how
Brandegee saw specimens labeled as H. frutescens.
As noted by Moran (1950), Dr. D. D. Keck considered the Todos Santos plant a new
species; but he has never published it. His conclusion was first based only on Brandegree's
collection, though apparently later supported by mine (2803). Material now available shows
that both the original H. greeneana, of Isla Guadalupe, and the Todos Santos plant are more
variable than was then evident, and their variation is roughly parallel. The differences noted
below are mostly not clear-cut but subject to exception. Thus the two plants do not appear as
different or as distinct as they might have before, and they are perhaps best treated as
subspecies of one species.
Plants of H. g. greeneana may be prostrate in exposed places; but though commonly
taller, they are still moundlike, with arching branches, even in the most sheltered places.
They are generally compact, the inflorescence being densely leafy and often many flowered
and even the lower internodes being short. On steep north slopes near the shore, plants of//.
g. peninsularis also grow prostrate, and elsewhere they may have somewhat arching
branches; but commonly they are more erect and more open, often with virgate branches and
mostly with longer internodes below.
In H. g. greeneana the leaves often are glabrous except for the glands but sometimes are
sparsely hirsute or villous-hirsute. The lower leaves are about 2 to 4 Vi cm long, seldom entire
but mostly with 4 to 12 teeth or lobes. In H. g. peninsularis the leaves, especially the
uppermost, may be glabrous except for the glands; but commonly they are hirsute or
hirtellous, the trichomes on middle and upper leaves being generally shorter and stiffer than
in H. g. greeneana. The lower leaves are commonly longer (to 7 cm) and relatively if not
actually narrower, with fewer teeth or lobes, which are thus more remote.
In H.g. greeneana the pappus of the disk achenes is about 1.3 to 2.2 mm long, of about 6
1969 MORAN: TWELVE NEW DICOTS 289
to 11 paleae, whereas in H. g. peninsularis it is about 1.9 to 3.3 mm long, of about 9 to 15
paleae. The reduction in H. g. greeneana might seem another example of decrease or loss of
dispersability in island plants (cf. Carlquist, 1966). However, the disk achenes are sterile in
all species of the section Zonamra, to which this species belongs.
The beak of the achene in H. g. greeneana is mostly 2 to 3 times as long as thick, in H. g.
peninsularis mostly 1 to 2 times as long as thick; but there are exceptions in both directions.
Hemizonia palmeri Rose, another member of the section Zonamra, is endemic with //.
g. greeneana at the south end of Isla Guadalupe, but in an even smaller area. It is a smaller,
prostrate, rather woody plant, with silky-strigose mostly entire leaves, the heads apparently
rather constantly with 8 rays and with about 10 to 12 disk florets. Apparently H. palmeri
flowers mainly in winter and H. g. greeneana more or less the year around. Hybrids are rare,
and the two species remain quite distinct.
Hemizonia greeneana is more similar to H. Clementina Brandegee and to H . frutescens
A. Gray. Hemizonia Clementina is restricted to five of the islands off southern California,
varying somewhat from island to island (Carlquist, 1965: 1 15 ff.). Hemizonia frutescens is
endemic to the high northern part of Isla Guadalupe, a cooler and moister area than the low
south end where H. g. greeneana grows. Since the discovery of H. frutescens in 1875, when
goats were already abundant, shrubby plants have been rare and confined to cliffs, though
Greene ( 1 885) reported it "common on level ground and hillsides and, in such places, strictly
annual". It is now known only on one north-facing cliff at 800 meters elevation, where about
20 plants are visible. There it is a shrub about 6 dm high, with virgate branches. Since it is
poorly represented in herbaria, detailed comparison with H. greeneana is difficult. In habit
and leaf form, both H. Clementina and H. frutescens are more similar to H. g. peninsularis
than to H. g. greeneana. which is divergent perhaps in connection with the more desertic
conditions under which it grows.
The heads of//, greeneana were described as having 8 rays, as at most seasons they do.
In both subspecies, however, early heads, terminating the main branches, may have 12 to 14
rays, a corresponding number of involucral and of receptacular bracts, and a correspond-
ingly large number of disk florets, sometimes as many as 21. Thus in head structure //.
greeneana is intermediate between the more primitive H. Clementina and the more advanced
H. frutescens. Except for having only a single series of receptacular bracts, the early heads of
H. greeneana rather closely resemble the heads of//. Clementina, which regularly have 1 2 to
14 rays and 1 5 to 30 disk florets. As the leaves of the two also are similar, H. g. peninsularis in
early flower can be confused with H. Clementina. On the other hand, the later heads of H.
greeneana resemble the heads of H. palmeri and H. frutescens. In H. frutescens. to judge
from scanty (and possibly inadequate) material, the number of rays is mostly 8 and rarely 9,
even in terminal heads of the main stems. Hemizonia frutescens differs from H. greeneana
further in its paler and perhaps thinner leaves and its longer and softer pubescence. In form,
color, and vesture of leaves, though not in habit, it bears a suggestive resemblance to putative
natural hybrids of//, palmeri with H. g. greeneana.
Heterotheca martirensis Moran, spec. nov. (Fig. 10)
Planta humilis perennis. rhizomata gracilia ad 1 '/: dm longa foliis squamiformibus
ample ctentibus ovatis obtusis ciliatis 2-4 mm longis instructa emittens. Caules graciles, 1-10
cm longi. Folia spatulata. mucronata. 1-2 Vi cm longa. 3-7 mm lata, vulgo utrinque hirsuta.
Capitulasolitaria. discoidea, fiava. plerumque 20-42-flora, pedunculis gracilibus 1-6 cm altis
0-2-bracteatis. Involucra campanulata. 6-9 mm aha. 4-6 mm lata, bracteis 24-48. in 3-4
seriebus imbricatis, lanceolatis. acutis, margine scar iosis pur pur ascentibusque. exterioribus
ca. 2 x Vi mm. interioribus ad 8 x 1-1 Vimm. Corollae 5-7 mm longae. Achenia compressa.
290 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
cuneata, 3-4 mm longa, 0.8-1.1 mm lata, 10-costata, villosa. Pappi setae sordidae,
interioribus 35-55, 4-7 mm longis, exterioribus 20-30, V2-I Vi mm longis. Typus: Moran
15612 (SD 69534). Ob habitum humilum rhizomatosum atque capitula solitaria discoidea
insignis. A liter in totogenere tan turn H. breweri et H. oregona capitula discoidea ferunt; hae
herbae erectae elatioresque sine rhizomatibus et cum capitulis plus minusve corymbosis
sunt.
Mat-forming rhizomatous perennial, from a rootcrown to 1 cm thick. Rhizomes
purplish at first, to 1 Vi dm long, ca. 1 mm thick, the internodes 1-3 mm long, the scale-leaves
clasping, triangular-ovate, obtuse, purple becoming brown, 2-4 mm long, hirsute-ciliate, the
upper passing into foliage leaves. Herbage hirsute with spreading to ascending pericellular
white trichomes Vi-2 mm long and glandular with truncate-conic white trichomes to 0.15
mm long, each tipped with a viscid yellow globule. Stems slender, 1-10 cm long above
ground, erect or decumbent, mostly leafy throughout though on longer stems the lower
leaves more scattered. Leaves thick, spatulate, 1 -2 V2 cm long, 3-7 mm wide above, Vi - 1 mm
wide above the broadened clasping base, mucronate with the whitish conic mucro commonly
deflexed and inconspicuous, glandular throughout, closely hirsute-ciliate at base, at least
sparsely hirsute on midrib dorsally but more often evenly hirsute on both surfaces, the midrib
impressed ventrally and projecting dorsally, the lateral veins ca. 3 on each side, looping,
mostly obscure. Heads discoid, yellow, solitary, flowering May to September, the peduncles
slender, 1-6 cm tall, glandular, hirsute or not, naked or with 1 or 2 linear bracts 2-3 mm long.
Involucre campanulate, 6-9 mm high, (2-) 4-6 mm wide (to 16 mm wide pressed), of 24-48
bracts well imbricated in 3 or 4 series, the bracts erect, lanceolate to linear-lanceolate, acute,
herbaceous and (at least the outer) glandular, purplish-scarious at margins, fimbrillate
above, with a prominent colorless or purplish midrib, the outer ca. 2 mm long and V2 mm
wide, the inner to 8 mm long, 1-1 Vi mm wide. Receptacle convex, alveolate. Florets (4-) 20-
42; corolla slender-funnelform, 5-7 mm long, yellow or becoming saffron red, the segments
erect, triangular-ovate, acute, 0.5-0.8 mm long; style branches flattened, linear, blunt, 1.5-
1 .9 mm long, the appendage puberulent, about equalling to twice exceeding stigmatic part.
Achenes tan, cuneate, compressed, 3-4 mm long, 0.8- 1 . 1 mm wide, 0.6-0.7 mm thick, with 10
low ribs, villous with ascending trichomes Vi - 1 mm long. Pappus sordid, double, the inner of
35-55 unequal slender barbellate bristles 4-7 mm long, the outer of ca. 20-30 similar bristles
'/2-1 Vi mm long or some of them slightly broadened and scalelike. Somatic chromosome
number: 2n= 18.
Type collection. -Common in crevices on flat granitic surfaces at 2800 meters
elevation, north slope of Cerro "2828", east rim of the Sierra San Pedro Martir, Baja
California, Mexico (near 31°02'N, 115°27'W), 14 September 1968, Moran 15612
holotype:SD 69534; isotypes:K, MEXU,UC, US, etc.
Distribution. - - Rather common, mostly on rocks, in full sun or partial shade, at 2050
to 2800 meters elevation in the north central Sierra San Pedro Martir. Other collections:
Cerro Venado Blanco, 2750 m, 15634; E slope at head of Arroyo Copal, 2500 m [ seen to
2300 m] , 15435; 2 miles E of Corral de Sam, 2050 m, 16531; type locality, 15053; SW slope
above Yerba Buena, 2500 m, Moran & Thome 14196, Moran 15067; Los Llanitos, 2500 m,
Moran & Thome 14257.
Discussion. The new species is placed in Heterotheca Cass, as emmended by
Shinners ( 1 95 1 ) to include Chrysopsis (Nutt.) Ell. The only other species with discoid heads
are H. breweri (A. Gray) Shinners and H. oregona (Nutt.) Shinners, both erect herbs with
branching stems bearing several heads. From both these species, H. martirensis differs
conspicuously in its low and rhizomatous habit, its solitary pedunculate heads, and its better
developed outer pappus. Very likely it is more closely related to some of the radiate species of
the Rocky Mountain area. In particular, it is similar in habit, leaf shape (including the
1969 MORAN: TWtLVE NEW DICOTS 291
mucronate tip), and pubescence, to Chrysopsis jonesii Blake, of southern Utah, though that
plant is more compact, with smaller leaves and smaller sessile heads.
Brandegee (1893) listed Chrysopsis sp. from the Sierra San Pedro Martir with no
comment. I have seen no collection of his.
For the type collection of H. martirensis. Dr. R. C. Jackson reports a somatic
chromosome number of 2n = 18.
Stephanomeria monocephala Moran, spec. nov. (Fig. 11)
Planta pulvinata, ad 3 dm lata. Folia rosulata 5-15 oblanceolata, acuta, V2SV2 cm
longa, 1-5 mm lata, sparsim glanduloso-puberulenta, superne utraque margine inaequaliter
0-3-dentata. Pedunculi V2-8 cm alti, superne 0-2-bracteati. Capitula solitaria, 14-21 mm
diametro, 5-8- flora. Involucra cylindracea, 7-9 mm longa, 2-3 mm lata, bracteis lanceolatis,
interioribus 5-9 aequalibus, exterioribus 3-5 imbricatis. Corollae tubus 2Vi-4 mm longus,
ligula rosea, 4 Vi -9 mm longa, 2 Vi -4 Vi mm lata, elliptico-oblonga, truncata, apice 5-lobata.
Achaenia prismatica, 2 Vi-3 mm longa, lateribus anguste sulcata, costis minute scaberulis.
Pappi setae persistentes, albae, biseriatae, interioribus 20, fere ad basin plumosis, 4-6 mm
longis. exterioribus alternantibus, minutis. Typus: Moran 15261 (SD 68877). Species habitu
pulvinato et capitulis solitariis distinctissima, pappi setis exterioribus minutis etiam
insignita.
Plant cespitose, forming dense cushions to 3 dm wide and 1 dm high, the interstices
packed solid with soil. Stems 2-5 mm thick, each branch with a rosette of ca. 5-15 leaves and
covered below with persistent dead leaves. Leaves oblanceolate to linear-oblanceolate or
occasionally spatulate, mostly acute, entire or commonly 1 -3-dentate on each margin above,
'/2-3'/2 cm long, 1-5 mm wide above, Vi-\ '/: mm wide above the broadened base,
subglabrous or (in the same plant) commonly glandular-puberulent with pericellular
trichomes ca. 0.1 mm long, the teeth spreading or slightly refkxed, triangular, the upper
mostly longer, to 1 mm long, the midrib prominent, the lateral veins obscure. Peduncle
terminal '/2-5 (-8) cm tall, slender, striate or angled, glandular-puberulent, often with 1 or 2
small lanceolate bracts above. Heads solitary, 10-12 mm high, 14-21 mm wide, with 5-8 but
commonly 6 florets, flowering May to July. Involucre cylindric, 7-9 mm long, 2-3 mm wide,
of 5-9 but mostly 6 equal bracts and 3-5 graduated shorter ones, the bracts lanceolate, acute
to narrowly rounded, 1-2 mm wide, green or purplish with scarious margins, minutely
granular-glandular and often also sparingly glandular puberulent, the longer ones sparsely
villous at apex. Receptacle shallowly pitted, glabrous. Corolla 7-13 mm long, the tube
whitish, 2 V2-A mm long, with a few scattered trichomes ventrally above, the ligule light to
deep pink or rarely white, elliptic-oblong, truncate and 5-lobed, 4 V2 -9 mm long, 2 V2AV2 mm
wide, the lobes 1-1 '/: mm long and a third as wide, triangular-ovate, obtusish. Anthers 3 V2S
mm long, sagittate at base; pollen white. Style 7-12 mm long, lavender above, ascending-
puberulent, the style branches ca. 1 mm long. Achenes light tan, pentagono-prismatic, only
slightly narrowed towards base and apex, 2 lA-3 mm long, 0.8- 1 .0 mm thick, the sides slightly
channeled or nearly flat except for a straight shallow longitudinal groove ca. 0.1 mm wide,
the angles sparsely and minutely ascending scaberulous. Pappus persistent, white, 4-6 mm
long, double, the inner bristles ca. 20, stiff, slightly widened and connate at base, plumose
nearly to base with pinnae 0.3-0.5 mm long, the outer bristles inconspicuous, alternating,
very slender, smooth or scaberulous, mostly less than 1 mm long.
Type collection. Common in crevices of north- and east-facing rocks and cliffs at
2800 meters elevation, Cerro "2828", east rim of the Sierra San Pedro Martir, Baja
California, Mexico (near 31°02'N, 1 15°27'W), 5 July 1968, Moran 15261 holotype: SD
68877; isotypes: K, MEXU, UC, US, etc.
292 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL.15
Distribution. - - Known only from the type locality. Other collections: 15064, 15332,
15403, 16533.
Discussion. - This plant appears to be quite limited as to habitat: despite wide
collecting in the general area on several trips, I found it only in the one place. However, there
are several similar cliffs on the east rim where it probably can be expected. Associated
species include Selaginella asprella Maxon, Sedum niveum Davids., Heuchera leptomeria
var. peninsularis Rosend., Butt., & Lak., Saxifraga eriophora S. Wats., Potentilla wheeleri
S. Wats., Haplopappus pulvinatus Moran, and Tanacetum bajacalifornicum Moran.
Stephanomeria monocephala differs from others of the genus in its polster habit and
solitary heads. The regular outer series of minute pappus bristles also is noteworthy; but
although the pappus of Stephanomeria is always described as uniseriate, a variable number
of similar tiny bristles occurs in several other species.
In other floral and fruiting characters, S. monocephala appears to be an average
member of the genus. However, it differs from every other species in some combination of
characters of involucre, flowers, achenes, and pappus. Mr. Leslie D. Gottlieb, a student of
the genus, suggests that it is closest to S. lactucina A. Gray, native from eastern Oregon to
Nevada and the Sierra Nevada. That is a larger plant, with stems arising singly from slender
rootstocks and usually bearing several heads; the heads are larger, with more and larger
florets; and the achenes are larger.
Tanacetum bajacalifornicum Moran, spec. nov. (Fig. 15)
Herbaaromatica caespitosa perennis, trichomatibus biramosis sparsim instructa. Folia
basalia 2-5 cm longa, 1-2 cm lata, plerumque biternate divisa, segmentis linearibus, obtusis,
1-1 Vi mm latis,petiolo laminam excedenti. Caulis florifer 1-2 dm alius, capitulum unum vel
pluria ferens, foliis superioribus integris. Involucra hemisphaerica, 4-6 mm aha, 4-8 mm
lata, bracteis 13-25, 3-seriatis, ellipticis, 1 Vi-3 mm latis, marginibus scariosis purpurascen-
tibus. Receptaculum nudum. Flores marginales pistillati 6-15, corollis 2Vi mm longis, 4-
denlatis. Flores disci perfecti 40-1 50 , corollis 3 mm longis. Pappus nullus. Achenia non visa.
Typus: Moran 15613 (SD 69075). A ceteris speciebus caespitosis turmae Sphaeromeriae
foliis caulibusque sparsim pubescentibus nee argenteis differt. A T. capitato, cui foliorum
forma similis est, capitulis majoribus minusque confertis, bracteis numerosioribus, et
floribus numerosioribus majoribusque praeterea differt.
Aromatic cespitose perennial, to 2 dm wide. Caudex to 2 dm long and to 1 cm thick at
base, the branches 2-5 mm thick, brown below with persistent shingled leaf bases. Herbage at
first thinly canescent with appressed crinkly biramous hairs, at maturity mostly subglabrate.
Leaves alternate, thick, glandular pitted, the basal onesca. 10-15, crowded, 2-5 cm long, the
base thin, clasping, ca. 4 mm wide, the petiole exceeding the blade, strap-shaped, ca. 1 mm
wide, the blade 1-2 cm wide, biternately divided or some pedately or pinnately and then
ternately divided, the segments linear or oblong, obtuse, 1-1 Vi mm wide. Floral stems
terminal, nodding at first, slender, 1 -2 dm tall, low ridged, with 10-15 linear leaves ca. 1 mm
wide, the lower 1 -2 cm long, ternately divided near apex, the upper shorter and entire. Heads
whitish, solitary or with 1 -2 smaller ones crowded below or with a few additional small ones
scattered in upper axils. Involucre hemispheric, 4-6 mm high, 4-8 mm wide (to 10 mm wide
pressed), the bracts 13-25, in about 3 nearly equal series, elliptic, acute to rounded, 4-6 mm
long, 1 Vz-3 mm wide, the midpart stiff coriaceous below, in all but the innermost thick and
green and somewhat glandular above, the broad margins scarious, purplish, erose, pilose-
ciliate in lower half, the inner narrower. Receptacle conoidal, naked, 1 '/2-3 mm high and
thick. Pistillate marginal florets 6- 1 5; corolla ca. 2 Vi mm long, tubular, nearly regular, with
4 triangular teeth; style branches ca. 0.6 mm long, well exserted. Perfect disk florets ca. 40-
1969
MORAN: TWELVE NEW DICOTS
293
Figure 15. Type specimen of Tanacetum bajacalifornicum (15613), from the high Sierra San Pedro Martir,
x 0.25.
150; corolla ca. 3 mm long, the tube equalling the conspicuously widened throat plus the 5
triangular teeth; style branches 0.6 mm long. Pappus none. Mature achenes not seen.
Type collection. Common, mostly in rock crevices, north slope of Cerro "2828"' at
2800 meters elevation. Sierra San Pedro Martir, Baja California, Mexico (near 31°02'N,
115°27'W), 14 September 1968, Moran 15613 - holotype: SD 69075; isotypes: K, MEXU,
UC, US, etc.
Distribution. - On rocks and cliffs near the east rim of the Sierra San Pedro Martir, at
2500 to 2800 meters elevation. Other collections: rock crevices of S summit ridge, Cerro
Venado Blanco, 2750 m, 15637; on rocks under pines and firs, E slope above Arroyo Copal,
2500 m, 15521; rock crevices on E slope of Cerro"2828", 2800 m, 15411.
Discussion. Tanacetum bajacalifornicum belongs to the Sphaeromeria group,
characterized in part by the lack of rays in the pistillate marginal flowers and by the presence
of malpighiaceous hairs. This group is centered in the Great Basin, and T. bajacalifornicum
is isolated some 375 miles south of the nearest other member. The other cespitose members
of the group have silvery canescent rather than sparsely pubescent herbage; and they differ
from T. bajacalifornicum further as follows (cf. Rydberg, 1916). In T. capitatum (Nutt.) T.
& G., ranging from Montana to Utah, which is most similar in leaf form, the heads are
smaller and more crowded, with fewer involucral bracts and with far fewer and slightly
294 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL.15
smaller florets. In T. compactum Hall, of the Charleston Mountains, Nevada, the plant is
smaller, the lower leaves are flabellately 5-parted into 3-lobed divisions and the upper
pinatisect, and a pappus is fairly well developed. In T. potentilloides A. Gray, ranging from
Oregon and Idaho to Nevada and eastern California, the leaves are pinnately to tripinnately
divided, the involucral bracts fewer, the receptacles white-hairy, and the florets smaller. In T.
nuttallii T. & G., ranging from Montana to Nevada, the leaves are cuneate and entire or 3-5-
lobed at the summit, the involucral bracts shorter and fewer, and the florets smaller. In T.
simplex Nels., of Wyoming, the leaves are entire or bifid or trifid at the apex, the floral stems
shorter, and the heads solitary.
ACKNOWLEDGEMENTS
I am grateful to Drs. Lincoln Constance, Peter Raven, and Robert F. Thorne, for critically reviewing all or part
of the manuscript. Drs. R. C. Jackson, James L. Reveal, and Charles H. Uhl have kindly made chromosome counts.
The maps are by Miss Gayle Culver.
Plants were collected under a series of permits kindly issued by the Direccion General de Aprovechamientos
Forestales of the Secretarfa de Agricultura y Ganaderfa of the Mexican Government; and in accordance with the
terms of the permits, specimens are being deposited in the Forestry Herbarium in Coyoacan.
LITERATURE CITED
Barkley, F. A.
1937. A monographic study of Rhus and its immediate allies in North and Central America, including the
West Indies. Ann. Missouri Bot. Gard. 24: 265-498.
Brandegee, T. S.
1 89 1 . Flora of the Cape Region of Baja California. Proc. Calif. Acad. ser. 2, 3: 1 08- 1 82.
1893. Southern extension of California flora. Zoe 4: 199-210.
1899. Island flora notes. Erythea 7: 70-71.
1900. Voyage of the Wahlberg. Zoe 5: 19-28.
1903. Notes and new species of Lower California plants. Zoe 5: 155-174.
Britton, N. L., and J. N. Rose
1903. New or noteworthy North American Crassulaceae. Bull. N.Y. Bot. Gard. 3: 1-45.
Carlquist, S.
1965. Island life: a natural history of the islands of the world. Natural Historv Press, Garden City, New York.
451 p.
1966. The biota of long-distance dispersal. II. Loss of dispersibility in Pacific Compositae. Evolution 20: 30-
48.
Epling, C. C.
1925. Monograph of the genus Monardella. Ann. Missouri Bot. Gard. 12: 1-106.
Epling, C. C, and W. S. Stewart
1939. A revision of Hedeoma. with a review of allied genera. Repert. Spec. Nov. Beih. 1 15: 1-49.
Greene, E. L.
1885. Studies in the botany of California and parts adjacent, II. Bull. Calif. Acad. 1: 179-228.
Hastings, J. R.
1964. Climatological data for Baja California. Techn. Rep. Meteor. Climat. Arid Reg. 14: 1-132. Univ.
Arizona.
Kearney, T. H., and R. H. Peebles
1951. Arizona flora. University of California Press, Berkeley. 1032 p.
Moran, R.
1950. Plants of the Todos Santos Islands, Baja California. Leafl. West. Bot. 6: 53-56.
1951. Notes on the flora of Guadalupe Island, Mexico. Madrono 1 1: 153-160.
Pennell, F. W.
1 935. The Scrophulariaceae of eastern temperate North America. Monogr. Acad. Sci. Philadelphia 1 : 1 -650.
1969 MORAN:TWFLVENEW DICOTS 295
1951. Scrophulariaceae. In L. R. Abrams. Illustrated flora of the Pacific States, vol. 3. Stanford University
Press. 866 p.
Rydberg, P. A.
1916. Carduaceae: Tageteae, Anthemideae. N. Amer. Flora 34(3): 181-288.
Shinners, L. H.
1951 . The north Texas species of Heterolheca. including Oirv5op.m(Cornpositae). Field & Lab. 19: 66-71 .
Department of Botany, Natural History Museum, P.O. Box 1390, San Diego,
California, 921 12
MUS. CO MP. ZOCL
LIBRARY
DEC 1 1 1969
HARVARD
UNIVERSITY
OBSERVATIONS ON A YOUNG PYGMY KILLER WHALE
(FERES A ATTENUATA GRAY)
FROM THE EASTERN TROPICAL PACIFIC OCEAN
WILLIAM F. PERRIN AND CARL L. HUBBS
TRANSACTIONS
OF THE SAN DIEGO
SOCIETY OF
NATURAL HISTORY
OBSERVATIONS ON A YOUNG PYGMY KILLER WHALE
(FERESA ATTENUATA GRAY)
FROM THE EASTERN TROPICAL PACIFIC OCEAN
WILLIAM F. PERRIN AND CARL L. HUBBS
ABSTRACT. An early juvenile specimen of Feresa attenuaia was captured in May 1967 during
commercial tuna fishing operations off Costa Rica. This is the first record of this species from the eastern
Pacific and from North American waters. Discrepancies in external morphology and in osteology were noted
between the specimen and earlier described adult specimens from Hawaii, Japan, and Senegal, but full
anah sis of the differences must await the capture of adult animals from the eastern Pacific or juveniles from
the other parts of the range.
RESUMEN. — En mayo 1967, durante las operaciones comercialesde pesca atunera frente a Costa Rica, se
capture unespecimenjuvenilde Feresa attenuaia en su primer ciclo vital. Estaespeciees la primera registrada
en el Pacffico oriental y en aguas norteamericanas. Se observaron diferencias en la morfologia externa y en la
osteologfa entreesteespecimen y especimenes adultos descritos anteriormente, provenientes delHauai.Japon
\ Senegal, peroel analisiscompletode lasdiferencias no se obtendra hasta que se capturen animales adultos del
Pacffico oriental o juveniles deotros lugaresen los ifmitesdedistribucion.
The pygmy killer whale, Feresa attenuaia Gray (1875), is one of the least known of the
delphinids. Gray (1871) based the genus Feresa on a skull first described as Delphinus
intermedins Gray, from an unknown locality. Gray (1875) later proposed the species
Feresa attenuata on the basis of another skull, from the "South Seas." The synonymy and
nomenclature of the species have been elucidated by Fraser ( 1 960).
The external appearance of the animal remained unknown and its geographical
distribution remained almost undetermined for three-fourths of a century, until Yamada
(1954) described a specimen from Taiji, Honshu, Japan, and gave it the vernacular name
used here. Thereafter the known range of the species was rapidly expanded, and its
characters were made more fully known. Cadenat (1958) and Fraser (1960) reported on a
specimen from Senegal, Africa. Nishiwaki et al. (1965) dealt with further material from
Japan. Pryor, Pryor. and Norris (1965) described a specimen from Hawaii. There is no
evidence that this species has ever occurred in Australia (Dawbin, pers. comm.; cf. Slijper.
1962:345).
There has been no previous evidence that Feresa attenuata occurs in the eastern Pacific,
although Hall and Kelson (1959: 830) stated that it "may occur in Pacific waters of North
America." Possibly it was observed long ago in the tropical eastern Pacific, but Scam-
mon's (1874: 104) description of the "Panama grampus" in Bahfa de Panama is not
sufficiently detailed to allow specific identification.
The first definite indication of Feresa attenuata in the eastern Pacific Ocean is furnished
by a juvenile male specimen taken in early May, 1967, approximately 300 to 400 nautical
miles off Costa Rica. It was collected by Anthony Dutra, a commercial fisherman of San
Diego. California, during tuna-seining operations, such as have recently been described
(Perrin, 1969). The net had been set on a school of "whitebelly porpoise." presumably
Delphinus sp. Mr. Dutra recognized the specimen as representing a species that he had
never seen before and saved it in the ship's freezer. It is almost certain that this juvenile
porpoise was accompanied by its mother, which apparently escaped during the purse-
seining operation. On return to San Diego. Mr. Dutra had a trophy cast made by a
SAN DIEGO SOC. NAT. HIST . TRANS. 15(18): 297-308. 24 NOVEMBER 1969
298 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
taxidermist, who brought the carcass (Fig. 1) to Scripps Institution of Oceanography for
identification. The specimen was identified as a juvenile male of Feresa attenuata on the
basis of form, size, coloration, and tooth number. The specimen was photographed,
measured, and skeletonized. The flippers and flukes were reconstructed from measure-
ments and tracings of the cast. The skeleton, a cast, and drawings of the flippers and flukes
are deposited in the Natural History Museum of the San Diego Society of Natural History
(No. 21561).
Whether this delphinid regularly inhabits eastern Pacific waters is unknown. That an
experienced and interested tuna fisherman recognized the specimen as of a species he had
never seen before, and that the species has not previously been observed by us or by
colleagues during extensive operations in the area of the eastern Pacific where tuna are
commonly caught, suggests that the juvenile (and the adult female) may have been a
straggler to the eastern Pacific from the Indo-Pacific area, in which respect this species
would be far from unique among marine organisms of the tropical Pacific (Ekman, 1953;
Briggs, 1961; Hubbs and Rosenblatt, 1961; Rosenblatt and Walker, 1963).
The general offshore area wherein the specimen was caught is about the warmest in
the eastern Pacific, with mean surface temperatures averaging about 28° C. during
May (LaViolette and Seim, 1969). This might be taken to indicate that the species is of
high-seas, tropical distribution, but other records of the species hardly confirm this.
Whereas the first specimen from Japan (Yamada, 1954) and the specimen from Hawaii
(Pryor et al., 1965) were captured in areas and during months when the surface tempera-
tures average approximately 26.5 and 23.5° C, respectively (LaViolette and Seim, 1969),
the later specimens from Japan (Nishiwaki et al., 1965) were taken from water at 13.5° C,
in January, 1963. This wide range of occurrence relative to water temperature leaves us
with little basis for predicting the probable distribution of the species.
EXTERNAL MORPHOLOGY
The specimen from off Costa Rica was a juvenile male 822 mm. long, from tip of
upper jaw to the notch in the flukes. Unerupted teeth and the small size indicate that the
animal may have been newly born. As received by the authors, minus flippers, fluke tips,
and viscera, and probably somewhat dehydrated by freezing, it weighed 9.37 kg. It does not
seem to differ from previously described specimens except in respects that reflect its youth.
External measurements were taken of the young specimen for comparison with those
recorded by Pryor etal. (1965) for an adult male from Hawaii and by Nishiwaki etal. (1965)
for 14 adult specimens and 1 fetus from Japan. Our measurements were made point-to-point
on the left side, with a large pair of precision calipers. Pryor et al. and Nishiwaki et al. made
their measurements according to the methods proposed by the Committee on Marine
Mammals of the American Society of Mammalogists (1961), which for several applicable
dimensions specify axial projections rather than point-to-point measurements. We feel that
the point-to-point method is the more precise, is readily applicable for smaller cetaceans,
and yields values that are convertible into axial projections, provided that good photographs
in several aspects have been made. Our measurements in millimeters,with thousandths of
total length from the tip of upper jaw to notch in flukes in parentheses (computed from axial
projections where appropriate), were: total length to notch of flukes, 822; tip of upper jaw to
center of eye, 82(100); tip of upper jaw to end of gape, 69(84); tip of upper jaw to blowhole,
76(92); tip of upper jaw to insertion of flipper, 186(226); tip of upper jaw to origin of dorsal
fin, 389(473); tip of upper jaw to tip of dorsal fin, 521(634); tip of upper jaw to midpoint of
umbilicus, 393(478); tip of upper jaw to midpoint of genital aperture, 485(590);tip of upper
1969
PERRIN AND HUBBS: Feresa attenuate
299
Figure I. Feresa attenuate. Top to bottom: right lateral view; dorsal view; ventral view; close-up of head in right
lateral view.
300 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
jaw to center of anus, 555(675); tip of upper jaw to origin of flukes, 747(909); tip of upper jaw
to plane of maximum girth, 379(461); maximum girth, 554(674); girth at axilla, 540(657);
girth below origin of dorsal fin, 540(657); girth above anus, 302(367); center of eye to end of
gape, 31(38); eye length, 17(21); eye height, 9(1 1); blowhole length, 6(7); blowhole width,
1 8(22); length of slit from umbilicus to anus, 1 55(189); length of mammary slit, 2(2); length
of anal slit, 13(16); insertion to tip of flipper, 160(195); maximum width of flipper, 58(71);
height of dorsal fin, 77(94); length of dorsal fin base, 1 30( 1 58); tip to tip of flukes, 210(255);
notch to nearest margin of fluke, 73(89); thickness of blubber at origin of dorsal fin, 1 3( 1 6).
Some of the proportions for the eastern Pacific specimen, despite the discrepancies in
age, fall within the ranges given for other specimens. A notable exception is the girth, which
probably was affected by the stuffing of our specimen during the casting process. In a number
of the proportions, those involving the dimensions of the anterior parts, the eastern Pacific
specimen has high or extreme values. This can be taken to reflect the youth of our specimen.
For these measurements, the fetus from Japan was even more extreme.
The long ventral groove extended from the umbilicus to the anus, as in the previously
described adult specimens. Four bristle follicles were visible on the right side of the upper jaw
and three on the left side.
Repeated freezing and thawing of our specimen no doubt obscured some of the more
labile features of the pigmentary pattern, such as the pale lateral area described by
Nishiwakietal. (1965). In the discernible pattern, however, our specimen resembled the one
from Hawaii (Pryor et al., 1965) more closely than those from Japan (Yamada, 1954;
Nishiwaki et al., 1 965), in that the ventral white patch extended from around the umbilicus in
a narrow wedge past the anus almost to the caudal end of the tail stock, rather than ceasing at
the anus. A prominent light patch between the flippers corresponded roughly with the gray
area described by Nishiwaki et al.
White markings edged each jaw, the upper the more conspicuously and the more
extensively. On the upper jaw the whitish areas broadened anteriorly, but these were
separated by a blackish area on the midline and tended to grade above into the blackish
snout, whereas posteriorly the coal-black and pure-white areas contrasted fully along the
line of contact. The white margin on the lower jaw was much narrower, and was largely
confined to the front of the mouth. No trace was evident of the irregular white blotches that
Pryor et al. ( 1 965) observed, and regarded as scars, in the adult male from Hawaii.
SKELETON
The entire skeleton was preserved to augment the very limited material available of this
species. Because the specimen was a juvenile, perhaps newly born, the skull was very
incompletely ankylosed, and some of the postcranial elements were incompletely ossified
(Figs. 2-8).
Our measurements of the skull in millimeters (with thousandths of condylobasal length
in parentheses) were: condylobasal length, 198.5; length of rostrum, 76.5(385); width of
rostrum at base, 50.3(253); width of rostrum at mid length, 37.9(191 ); width of premaxillae
at midlength, 21.9(1 10); greatest width of premaxillae, 46.9(236); tip of rostrum to anterior
margin of superior nares, 105.3(530); preorbital width, 95.5(481); postorbital width,
109.8(553); midorbital width, 101.2(510); maximum width of blowhole, 30.3(153); zygo-
matic width, 109.5(552); length of left mandible, 144.3(727); length of right mandible,
143.5(723); left coronoid height, 39.3(201); right coronoid height, 40.5(204); length of
symphysis, 11.1(56); length of left upper tooth row, 51.5(259); length of right upper tooth
row, 52.4(264); end of left upper tooth row to tip of premaxillae, 61 .4(309); end of right upper
1969
PERRIN AND HUBBS: Feresa attenuate
301
Figure 2. Skull of Feresa attenuate. Top to bottom: dorsal view; ventral view; right lateral view.
302
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
Figure 3. Feresa atienuata. Top left, posterior view of skull; top right, mandible, dorsal view; bottom (a) right
ramusof mandible, medial view; (b) leftramusof mandible, lateral view.
toothrowtotipofpremaxillae, 62.6(315); length of left lower tooth row, 70.1(353); length of
right lower tooth row, 71.4(360); end of left lower tooth row to tip of mandible, 73.2(369);
end of right lower tooth row to tip of mandible, 74.4(375); length of bulla of left tympano-
periotic complex, 33.7(170); width of bulla of left tympano-periotic complex, 21.4(108);
internal length of the braincase, 9 1 .2(459). The tooth counts were 9/9 in the upper jaw and
1 2/ 1 2 in the lower jaw for a total of 42 teeth.
1969
PERRIN AND HUBBS: Feresa altenuata
303
Figure4. Left, skull of type of Feresa attenuata B.M.N. H. no. 1874.11.25.1 (after Fraser, 1960). Right, skull of
juvenile specimen from Costa Rica, S.D.S.N.H. no. 21561.
Measurements of the skull were compared with those for the type specimen of Feresa
attenuata (British Museum No. 1672a); for the type specimen of F. intermedia (British
Museum No. 362a), synonymized with F. attenuata by Fraser (1960); and for recently
collected adult specimens from Hawaii, Japan, and Senegal. Differences in proportions
between the Costa Rica juvenile and the adults from other areas presumably are due to great
ontogenetic changes in skull structures. However, as the skull of the juvenile became
disarticulated during maceration, it is possible that proportions calculated from measure-
ments made after the cranium had been reassembled are slightly distorted.
The juvenile from off Costa Rica differs strikingly from the adults in the small
proportionate size of the rostrum (Fig. 4 ); this is consonant with the general pattern of
development in mammals, especially those with large brains: the areas concerned with
feeding grow more slowly than the brain case. The dentigerous portions of the jaws are also
Figure 5. Feresa attenuata. (a) left tympano-perioticum. medial view; (b) right tvmpano-perioticum. medial view;
(c) left tympano-perioticum, lateral view; (d) right tympano-perioticum disarticulated to show stapes (left), incus
and malleus.
304
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
Figure 6. Left and right vertebral ribs of Feresaattenuata. Anteriormost ribs at top.
1969
PERRIN AND HUBBS: Feresa attenuata
305
& • ••• & m
iWWWWW^AA
& §b 4& & ^ 4fc
AAAAAAAAA
A A A AAA A A A
AAA A A A AAA
A #% #%
#• #%
*# ••••
Figure 7. Vertebrae of Feresa attenuata. cranial view. Second cervical vertebra fused to first. Halves of neural
processes on vertebrae 1-14 and 52-55 not ankvlosed. shown in anteromedial view. Intervertebral discs present on
some vertebrae. The vertebral column became partialk disarticulated during maceration, consequent!) some-
elements may beoutol'order in this photograph.
306
SAN DIEGO SOCIETY OF NATURAL HISTORY
VOL. 15
Figure 8. Feresa atlenuata. Top, hyoid bones; bottom left, sternum and sternal ribs; bottom right, (a) proximal
ends of humeri; distal ends severed and missing; (b) scapulae; (c) pelvic bones.
1969 PERRIN AND HUBBS: Feresa attenuata 307
proportionately shorter than in the adults. The mandible (Fig. 3), however, is
proportionately nearly as large as in the adult. The premaxillae are narrow near the middle,
but the greatest width is not very different. The whole skull is narrower than in the adult (Fig.
4 ), especially in the region of the orbits and across the zygomatic arches. The condylobasal
length is 1 .90 times that of the parietal width in the juvenile, as compared with 1 .42 to 1 .73
times in the adults described by Nishiwaki et al. (1965). The skull has not become telescoped
as far as in the adults: large expanses of the frontals and parietals remain visible in dorsal
view.
Fraser (1960) stated that in the adult skulls of Feresa that he examined, the dorsal
surface of the rostral portion of the premaxillae is flat and on the same level as the adjacent
maxillae, a condition similar to that in Orcinus and Lagenorhynchus. In our specimen the
premaxillae form a considerable convexity above the level of the maxillae (Fig. 2 ) in the
distal two-thirds of the rostrum, a condition approaching that in adults ofStenella, Tursiops,
and Delphinus.
The tympano-periotic complex (Fig. 5 ) is very large proportionately in comparison to
that of the adults, viz., 170 thousandths of condylobasal length versus a range of 102 to 1 14
thousandths for the Japan specimens, in length; and 108 thousandths versus a range of 62 to
90 thousandths, in width of the bulla. The strong mastoid process figured by Yamada (1954)
is not evident. In this respect the juvenile specimen is more like adults of Grampus,
Delphinus, Stenella and related genera figured by Yamada ( 1 953). The overall appearance of
the tympanic bone very closely resembles that of adults of Globicephala, particularly in the
rounded anterior end. The incus differs from those figured by Yamada (1954) in that the
major crus is slenderer and the minor crus longer proportionately, and the facet at the end of
the minor crus is lacking.
The vertebral formula is C 7 T, : L + C 46 , for a total of 65 vertebrae, at least two fewer
than the previously recorded minimum for the species. However, the very small terminal
vertebral remnant noted by Yamada was not recovered in our preparation. The vertebral
column (Fig. 7 ) was only partially ossified. Incompletely ossified neural processes of
varying shape crested the first 55 centra, but transverse processes and chevron bones were
still completely cartilaginous. The uncinate processes noted on the thoracic vertebrae of all
previous specimens are not apparent. The first two cervical vertebral centra are fused. In the
adult specimens from Japan, the first three to six vertebrae were fused.
The first six of the 1 2 pairs of vertebral ribs are two-headed, and there are five pairs of
ossified sternal ribs (Figs. 6, 8). The ossified sternum (Fig. 8) consists of two unfused
elements, of which the anterior one is expanded and bilobed anteriorly and the second has
convex sides.
The scapulae (Fig. 8 ) are uneroded. The short acromion process turns anteriorly rather
than posteriorly as in previously described specimens (Yamada, 1 954; Pryor et al., 1 965).
ACKNOWLEDGEMENTS
We thank Mr. Anthony Dutra. the fisherman who collected the specimen, provided information, and allowed
us to photograph the cast which hangs in his home: and Mr. Hugh Lyons, who brought the animal to our attention
and cooperated otherwise. Kenneth S. Raymond prepared figure 4. Lawrence D. Ford made the photographs ol the
carcass used for figure 1 . Dr. Kenneth S. Norris has read the manuscript. Support was provided by the U.S. Bureau
of Commercial Fisheries. The researches by Hubbs on marine vertebrates are supported by the National Science
Foundation (grant GB4672).
308 SAN DIEGO SOCIETY OF NATURAL HISTORY VOL. 15
LITERATURE CITED
Briggs, J. C.
1961. The East Pacific Barrier and the distribution of marine shore fishes. Evolution 15: 545-554.
Cadenat, J.
1958. Notes sur les Delphinides Ouest-africains II — Un specimen du genre Feresa capture sur les cotes du
Senegal. Bull. Inst. Franc. Afrique Noire 20: 1486-1491.
Committee on Marine Mammals, American Society of Mammalogists
1961 . Standardized methods for measuring and recording data on the smaller cetaceans, K. S. Norris (Ed.)
J. Mamm. 42:471-476.
Ekman, S.
1953. Zoogeography of the sea. Sidgwick and Jackson, London, 417 p.
Fraser, F. C.
1960. A specimen of the genus Feresa from Senegal. Bull. Inst. Franc. Afrique Noire 22: 699-707.
Gray, J. E.
1871. Supplement to the catalogue of seals and whales in the British Museum. (Not seen.)
1875. Feresa attenuata. J. Mus Godeffroy, Hamburg, Heft 8: 184 + pi. 6.
Hall, E. R., and K. R. Kelson
1959. The mammalsof North America. Ronald Press, N.Y. 2 vols.
Hubbs, C. L., and R. H. Rosenblatt
1961 . Effects of the equatorial currents of the Pacific on the distribution of fishes and other marine animals.
Tenth PacificSci. Congr., Abst. Sym. Papers. Honolulu, Hawaii, p. 340-341.
LaViolette, P. E., and S. E. Seim
1969. Monthly charts of mean, minimum and maximum sea surface temperature of the North Pacific Ocean.
Spec. Publ. 1 23, Naval Oceanogr. Office, Washington, D.C., 58 p.
Nishiwaki, M., T. Kasuya, T. Kamiya, T. Tobayama, and M. Nakajima
1965. Feresa attenuata captured at the Pacific coast of Japan in 1963. Sci. Rep. Whales Res. Inst. Tokyo 19:
65-95.
Perrin, W. F.
1 969. Using porpoise to catch tuna. World Fishing 1 8: 42-45.
Pryor, T., K. Pryor, and K. S. Norris
1 965. Observations on a pygmy killer whale (Feresa attenuata Gray) from Hawaii. J. Mamm. 46: 450-46 1 .
Rosenblatt, R. H, and B. W. Walker
1963. The marine shore-fishes of the Galapagos Islands. Calif. Acad. Sci.Occ. Papers 44: 97-106.
Scammon, C. M.
1 874. The marine mammals of the northwestern coast of North America. Described and illustrated. Together
with an account of the American whale fishery. G. P. Putnam's Sons, N.Y. 319 p. + v. Facsimile
editions by Dover Pub., N.Y. (1968); and Manessier Pub. Co., Riverside, Calif. (1969).
Slijper, E. J.
1962. Whales. Basic Books, N.Y. 475 p.
Yamada, M.
1953. Contribution to the anatomy of the organ of hearing of whales. Sci. Rep. Whales Res. Inst. Tokyo 8: I-
80.
1954. An account of a rare porpoise, Feresa Gray from Japan. Sci. Rep. Whales Res. Inst. Tokyo 9: 59-88.
U.S. Bureau of Commercial Fisheries Fishery-Oceanography Center, La Jolla,
California, and Scripps Institution of Oceanography, University of California, San Diego
(La Jolla, Calif or n ia ).
BOUND 197071
3 2044 093 361 301
Date Due
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11